Cape stinkweeds: Taxonomy of Oncosiphon (Anthemideae, Asteraceae)

South African Journal of Botany 117 (2018) 57–70

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Cape stinkweeds: Taxonomy of Oncosiphon (Anthemideae, Asteraceae) R. Kolokoto a, A.R. Magee a,b,⁎ a b

Compton Herbarium, South African National Biodiversity Institute, Private Bag X7, Claremont, 7735 Cape Town, South Africa Department of Botany and Plant Biotechnology, University of Johannesburg, PO Box 524, Auckland Park, 2006 Johannesburg, South Africa

a r t i c l e

i n f o

Article history: Received 14 August 2017 Received in revised form 5 April 2018 Accepted 26 April 2018 Available online xxxx Edited by GV Goodman-Cron Keywords: Aromatic Ethnobotany Khoi-San Medicinal plants Natural hybrid Southern Africa Stinkruid

a b s t r a c t A taxonomic revision of the southern African endemic genus Oncosiphon Källersjö (Asteraceae: Anthemidae) is presented. The genus includes annuals or short-lived perennials with discoid or rarely radiate capitula, a conspicuously swollen corolla tube, and 4 ribbed, non-myxogenic cypselas. The Afrikaans vernacular name stinkruid (stinkweed) is commonly applied to species of this genus in reference to their strong, unpleasant odour. Three of the species are recognised as important herbal remedies in Khoi-San and Cape Dutch enthnobotany. We recognise seven species of Oncosiphon, and report the occurrence of occasional hybrids between O. suffruticosus and O. grandiflorus. The species are distinguished by their vestiture, leaf division, inflorescence structure, floral characters and the indumentum of the involucral bracts. A key to the species, the complete nomenclature, species descriptions, diagnostic characters, and distribution and ecological details are provided. © 2018 SAAB. Published by Elsevier B.V. All rights reserved.

1. Introduction Oncosiphon Kӓllersjö (Asteraceae: Anthemideae) is a genus of aromatic herbs of which all except two species, O. piluliferus (L.f.) Kӓllersjö and O. suffruticosus (L.) Kӓllersjö, are endemic to the Greater Cape Floristic Region of southern Africa (Manning & Goldblatt, 2012; Herman & Manning, 2013). These two species have also been recorded as weeds in south-west Western Australia under the names Calomba Daisy (O. suffruticosus) and Globe Chamomile (O. piluliferus) (Atlas of Living Australia, n.d., 31 July 2017, http://bie.ala.org.au/species). Oncosiphon includes both discoid and radiate species, with the former previously included in Pentzia Thunb. and the latter in Matricaria L. (Källersjö, 1988). The genus was segregated from the polymorphic Pentzia by Källersjö (1988) to accommodate those annual species with four-lobed corollas, a swollen corolla tube, and four-ribbed nonmyxogenic fruit. The genus is closely related to Foveolina Kӓllersjö, also a southern African endemic (Magee, 2011; Magee et al., 2015). Oncosiphon and Foveolina s.str. share a variously swollen floral tube but differ in their fruit morphology (four-ribbed, not myxogenic fruits with a sclerenchymous epidermis in Oncosiphon vs three-ribbed, myxogenic fruit with short-stalked glands and prominent spongy pappus in Foveolina s. str.) (Källersjö, 1988; Magee et al., 2015). Recent ⁎ Corresponding author at: Compton Herbarium, South African National Biodiversity Institute, Private Bag X7, Claremont, 7735 Cape Town, South Africa. E-mail address: [email protected] (A.R. Magee).

https://doi.org/10.1016/j.sajb.2018.04.020 0254-6299/© 2018 SAAB. Published by Elsevier B.V. All rights reserved.

phylogenetic analyses of subtribe Pentziinae by Magee et al. (2015) suggested that Foveolina is polyphyletic, with the type group (Foveolina s. str., including only two species F. dichotoma (DC.) Källersjö and F. tenella (DC.) Källersjö closely allied to and possibly embedded within Oncosiphon. Surprisingly, the two species of Foveolina s. str. were not recovered as sister taxa in the phylogenetic trees of Magee et al. (2015), despite differing morphologically only by the presence or absence of ray florets. Magee et al. (2015) recommended that further study was required to resolve these relationships before any taxonomic changes could be made. Species of Oncosiphon are commonly refered to under the Afrikaans vernacular name stinkruid in reference to their strong unpleasant odour (Watt and Breyer-Brandw, 1962). The radiate species O. africanus (P.J. Bergius) Kӓllersjö has been called wildekamomille, and O. piluliferus is known as wurmbos or miskruid (Van Wyk, 2008). Three of the species, O suffruticosus, O. piluliferus and O. africanus (as O. glabratus (Thunb.) Kӓllersjö) were recognised as important herbal remedies in Khoi-San and Cape Dutch ethnobotany by Van Wyk (2008), and have been used to treat a variety of ailments, including typhoid fever, rheumatic fever and influenza, and as a general tonic, anthelmintic and diuretic as well as a poultice for scorpion stings. The species of Oncosiphon were last treated under either Pentzia, Matricaria L. or Tanacetum L. by De Candolle (1838) and Harvey (1865). When circumscribing the genus Oncosiphon, Källersjö (1988) provided a key to the eight species that were recognised at the time, along with some taxonomic notes. Other than the subsequent reduction

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of O. glabratus into synonymy with O. africanus by Manning and Goldblatt (2005), no taxonomic review of the genus has been pursued. As a result, the species are often confused with one another (further complicated by the presence of intermediate specimens) as well as with species from other similar genera (viz. Cotula L., Foveolina and Myxopappus Källersjö). In addition, the taxonomic status of Oncosiphon schlechteri (Bolus) Källersjö has been highlighted as problematic, with some authors suggesting that it may be synonymous with O. sabulosus (Wolley-Dod) Källersjö (Herman and Manning, 2013). We provide here a comprehensive taxonomic treatment of Oncosiphon in which we include an updated key to the species, complete nomenclature, species descriptions, diagnostic characters, phenology, and distribution and ecological details. 2. Materials and methods Species descriptions and distribution data were derived from herbarium material, comprising 410 specimens, from the following herbaria: BOL, NBG, PRE, SAM and WIND (acronyms from Holmgren et al., 1990). Selected images from LINN, THUNB-UPS and JSTOR Global Plants (https://plants.jstor.org) were also studied. In addition, six of the

seven species recognised were studied in the wild. Details of capitula and flowers were photographed on an Olympus SZ61 Stereomicroscope with Olympus SC30 camera. The species are arranged in the taxonomic treatment according to presumed relationships. 3. Results and discussion Most Oncosiphon species are annual herbs, apart from O. sabulosus and O. schlechteri, which are short-lived perennials to 0.5 m tall (Fig. 1B & C). The leaves are usually bipinnatisect or sometimes tripinnatisect with mucronate lobes (Fig. 2A), and are sometimes slightly succulent in O. suffruticosus. The leaves of O. sabulosus and O. schlechteri are diagnostically different from those found in the rest of the genus: in O. sabulosus they are pinnatifid to pinnatisect and narrowly oblong to obovate Fig. 2B), while in O. schlechteri they are simple or apically 3- to 5-fid and spatulate to linear (Fig. 2C). In both species, the leaves are somewhat succulent with blunt (rather than mucronulate) lobes (Fig. 2B & C). The synflorescence is usually a lax corymb with ≤ 6 capitula (Fig. 3A & B) but O. suffruticosus is characterised by a dense compound corymb with ≥ 10 capitula (Fig. 3C), and in O. piluliferus the capitula are usually solitary. With

Fig. 1. The species of Oncosiphon. A & B, O. sabulosus; C, O. schlechteri; D, O. africanus; E, O. intermedius; F–H, O. grandiflorus; I, O. piluliferus; J, O. suffruticosus.

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Fig. 2. Leaf diversity in Oncosiphon. A, O. grandiflorus; B, O. sabulosus; C, O. schlechteri. Vouchers: A, Rösch 469 (NBG); B1, Kolokoto & Magee 4 (NBG); B2, Boucher 4059 (NBG); C1, Desmet 40 (NBG); C2, Boucher 4059 (NBG). Scale: 10 mm.

the notable exception of O. africanus, which has radiate, heterogamous capitula (Fig. 1D), the capitula are discoid and homogamous. The vestiture and orientation of the involucral bracts are important characters to distinguish several of the species. The bracts in O. grandiflorus and O. intermedius are prominently tomentose but are glabrous in the other species. At maturity the involucral bracts are usually suberect to slightly deflexed (Fig. 4H & I) so that the base of the involucre is rounded to slightly cordate. In contrast, the involucral bracts of O. piluliferus are strongly deflexed to below the level of the receptacle, rendering the involucral base deeply cordate (Fig. 4J). The disc florets are bisexual, four-lobed and yellow or fading to burnt yellow (Fig. 1A–D; F–J) in most of the species but are distinctly orange in O. intermedius (Fig. 1E). The corolla tube is usually conspicuously swollen (Fig. 4B–E), although only slightly so in O. africanus (Fig. 4A2). The latter species is also the only one with a prominently obliquely cup-shaped pappus (Fig. 4F); in the other species the pappus is small or inconspicuous (Fig. 4G). The fruits are uniformly homomorphic, four-ribbed and lack myxogenic cells (Fig. 4F & G).

4. Taxonomic treatment Oncosiphon Källersjö in Bot. J. Linn. Soc. 96(4): 310 (1988); Bremer and Humphries in Bull. Nat. His.Mus. 23 (2): 152 (1993); Goldblatt and Manning, Cape Plants: 345 (2000); Manning and Goldblatt, Plants of the Greater Cape Floristic Region: 401 (2012); Snijman, Plants of the Greater Cape Floristic Region: 307 (2013). Type: O. piluliferus (L.f.) Källersjö. Erect annual or short-lived perennial, aromatic herbs, 0.3 to 0.5 m tall; stems glabrous to pilose or cobwbebbed. Leaves alternate, apically 3- or 5-fid to pinnatifid or bipinnatisect, rarely simple, linear to oblanceolate or obovate, sparsely pilose to pilose; lobes oblong to lanceolate or oblanceolate, mucronulate or obtuse. Capitula either discoid and homogamous or rarely radiate and heterogamous (O. africanus), terminal, pedunculate, usually coymbose, corymbs lax and simple or dense and compound, sometimes solitary; peduncles sometimes thickened apically, glabrous to pilose. Involucre hemispherical or campanulate, base attenuate, round or cuneate to deeply cordate in mature capitula; involucral bracts 3-seriate, suberect to slightly or strongly deflexed at

Fig. 3. Inflorescence structure in Oncosiphon. A, simple lax corymb of O. piluliferus; B, solitary capitula of O. africanus; C, dense compound corymb of O. suffruticosus. Vouchers: (A) Van Slangeren et al. MSBYJT384 (NBG); (B) Compton 9815 (NBG); (C) Fellingham 1519 (NBG). Scale: 2.6 mm.

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Fig. 4. Floret (A–E), cypsela (F–G) and involucre (H–J) diversity in Oncosiphon. A1, O. africanus ray floret; A2, O. africanus with inconspicuous swollen corolla tube; B, O. intermedius with narrowly lanceolate corolla lobes; C, O. grandiflorus with ovate corolla lobes; D, O. sabulosus; E, O. schlechteri showing conspicuous swollen corolla tube; F, O. africanus with conspicuous pappus; G, O. piluliferus with inconspicuous pappus; H, O. grandiflorus involucral bracts, partially obscuring the receptacle; I, O. suffruticosus involucral bracts slightly deflexed, obscuring the receptacle; J, O. piluliferus involucral bracts strongly deflexed, not obscuring the receptacle. Vouchers: (A1, A2 & F) Craven 26, NBG; (B) Low 7778, NBG; (C & H) Kruger 1683, NBG; (D & I) Perry & Snijman 2452, NBG; (E) Boucher 4073, NBG; (G) Taylor 2748, NBG; (J) Hugo 1905, NBG. Scale: 2 mm (A1, H, I & J); 1 mm (A2, B, C, D & E); 0.5 mm (F & G).

anthesis, sometimes decurrent on the peduncle, glabrous or tomentose, midvein inconspicuous or conspicous, margins and apices narrowly or broadly scarious. Receptacle flat-convex to conical-globose, obscured or unobscured by involucral bracts, epaleate. Ray florets present in one species, female-fertile; tube glandular, trichomes sessile; limb oblong, white. Disc florets bisexual, corolla yellow, rarely orange (O. intermedius); tube slightly to conspicuously swollen, limb campanulate; 4-lobed, lobes ovate or narrowly lanceolate, cucullate. Anthers sagittate at base; apical appendages lanceolate to ovate. Style terete with thickened base; branches truncate, with papillate dorsally. Pappus inconspicuous to prominent, irregularly lobed or obliquely cup-shaped with adaxial lobes longer, membraneous, white. Cypselas homomorphic, subterete, tapering at base, yellowish brown, 4-ribbed, nonmyxogenic, sessile glandular trichomes present. Diagnostic characters Species of Oncosiphon can sometimes be confused with the closely related Foveolina and Myxopappus, with which they share the annual habit and four-lobed disc florets. Oncosiphon is distinguished by the 4ribbed, non-myxogenic cypselae with a sclerenchymous epidermis (3ribbed, myxogenic cypselae without sclerenchymous epidermis in Foveolina and Myxopappus), the usually inconspicuous pappus (prominent and spongy in Foveolina and Myxopappus), and the florets with the tube prominently swollen along most of its length (not swollen in Myxopappus and only slightly swollen near the base in Foveolina). In Myxopappus and Foveolina burchellii (DC.) Magee the capitula are disciform (discoid or radiate in Oncosiphon).

Distribution and ecology The species of Oncosiphon are endemic to southern Africa, with six of the seven species restricted to the Greater Cape Floristic Region. Only O. piluliferus extends more widely into the summer rainfall regions. 4.1. Key to the species of Oncosiphon 1a. Short-lived perennials; leaf lobes obtuse: 2a. Leaves pinnatifid to pinnatisect, oblong to obovate in outline; peduncles and stems glabrous; mature capitula ≥5 mm diam. with base of involucre round to cordate ……………………………… 1. O. sabulosus 2b. Leaves apically 3- to 5-fid or simple, spatulate to linear in outline; peduncles and young stems prominently cobwebbed; mature capitula ≤ 5 mm diam. with base of involucre cuneate ……… ………………………………………………………… 2. O. schlechteri 1b. Annuals; leaf lobes mucronulate: 3a. Capitula radiate; pappus well-developed, 0.5–0.7 mm long, almost half as long as corolla tube; corolla tube not conspicuously swollen ……………………………………………………………… 7. O. africanus 3b. Capitula discoid; pappus inconspicuous or small, less than 0.3 mm long, almost one quarter the length of the corolla tube; corolla tube conspicuously swollen:

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4a. Involucral bracts tomentose; receptacle flat-convex; peduncles prominently thickened below capitula: 5a. Florets orange, lobes narrowly lanceolate, 0.7–1 mm long; restricted to the Bidouw and Wuppertal valleys ………………… 4. O. intermedius 5b. Florets yellow (fading amber with age), lobes ovate, ± 0.5 mm long; widespread in Northern Cape and Western Cape ……………… ………………………………………………………… 3. O. grandiflorus 4b. Involucral bracts glabrous; receptacle globose to conical; peduncles not thickened below capitula: 6a. Capitula in dense, compound corymbs (≥ 10 capitula); involucre campanulate, 3–5 mm diam.; involucral base rounded to shallowly cordate in mature capitula with involucral bracts suberect to slightly deflexed, obscuring receptacle …………………… 6. O. suffruticosus

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Decumbent perennial herb to 0.5 m tall; stems glabrous. Leaves pinnatifid to pinnatisect, narrowly oblong to obovate, 5–20(−30) × 4–8 (−10) mm, succulent, glabrous; lobes, narrowly oblong to oblanceolate, 1–3 × ± 2 mm, apex obtuse. Capitula discoid, homogamous, in simple, lax comrybs; peduncles slender, 5–16 mm long, glabrous. Involucre hemispherical, 5–9 mm diam., base round to cordate in mature capitula; involucral bracts suberect to slightly deflexed at anthesis, some decurrent on the peduncle, apices ± level with receptacle apex, glabrous, midvein slightly prominent, yellow, margins and apices narrowly scarious; outer series narrowly lanceolate, ± 3 mm long; median series narrowly oblong, ± 3–4 mm long; inner series lanceolate to narrowly oblong, 2–3 mm long. Receptacle globose-convex, obscured by involucral bracts. Disc florets bisexual, corolla 2–3 mm long, yellow; tube conspicuously swollen from apex to base, oblong to elliptic, ± 1.3 mm long; limb campanulate, ± 1 mm long; lobes ovate, ± 0.5 mm long. Pappus inconspicuous to small, irregularly lobed, ± 0.2–0.3 mm long. Diagnostic characters

6b. Capitula solitary or in simple corymbs (≤ 6 capitula); involucre hemispherical, 5–10 mm diam.; involucral base deeply cordate in mature capitula with involucral bracts strongly deflexed, not obscuring receptacle …………………………………… 5. O. piluliferus 1. Oncosiphon sabulosus (Wolley-Dod) Källersjö in Bot. J. Linn. Soc. 96 (4): 312 (1988); Manning and Goldblatt, Plants of the Greater Cape Floristic Region: 401 (2012); Snijman, Plants of the Greater Cape Floristic Region: 307 (2013). Matricaria sabulosa Wolley-Dod in J. Bot. 39: 399 (1901); Levyns in Adamson and Salter, Fl. Cap. Penins.: 804 (1950). Pentzia sabulosa (Wolley Dod.) Hutch. in Bull. Misc. Inform. Kew: 253 (1917). Type: South Africa. Western Cape, Simonstown (3418): shore of Klein Kalk Bay (–AB), 17 Oct 1897, Wolley-Dod 3421 (K, lecto.–image!, designated by Källersjö, 1988; BOL!, isolecto.).

Oncosiphon sabulosus shares the perennial habit, blunt leaf lobes and decurrent involucral bracts with O. schlechteri. It is distinguished by the pinnatifid to pinnisect leaves (Fig. 2B) which are oblong to obovate in outline (leaves apically 3- to 5-fid or sometimes simple and spatulate to linear in outline in O. schlechteri), glabrous peduncles and stems (cobwebbed in O. schlechteri) and round to cordate involucre base with only some involucral bract decurrent on the peduncle (involucre base cuneate with several involucral bracts decurrent in O. schlechteri). Distribution and ecology Oncosiphon sabulosus is restricted to the beach and open coastal dunes from Lamberts Bay to Cape Agulhas in Western Cape (Fig. 5). Flowering is from spring to early autumn (September to March).

Fig. 5. Distribution of Oncosiphon sabulosus (stars), O. schlechteri (dark circles) and O. intermedius (open circles).

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Additional specimens examined South Africa. Western Cape. 3218 (Clanwilliam): Lamberts Bay (–AB), Kolokoto and Magee 4 (NBG); Velddrif, Rocher Pan Nature Reserve, N section of the reserve, sea dunes (–CB), Rooyen and Ramsey 351 (NBG); Rocher Pan Nature Reserve, (–CB), Boucher 4073; Le Roux and Ramsey 358 (NBG). 3318 (Cape Town): Klein Eiland in Postberg Nature Reserve (–AA), Graven 2 (NBG); Oude Post Private Nature Reserve, Langebaan Peninsula (–AA), Boucher 2975 (NBG); Yzerfontein, (–AC), Levyns 10,031 (BOL); Melkbosstrand (–CB), Heistner 2 (NBG); Dahlstrand 1048 (PRE); Cassidy 160 (NBG, BOL); Boucher 3987 (NBG); Robben Island, slope directly above high-tide mark ± widway along western edge of the island (–CD), Lloyd 492a (NBG); Robben Island (–CD), Van Jaarsveld and De Lange 9527 (NBG). 3418 (Simonstown): coast at Stony Point, near Hangklip (–BD), Pillans 8506 (BOL); shore of Klein Kalk Bay (–AB), Wolley Dod 3421 (BOL). 3419 (Caledon): Onrust, Vermont (–AC), Jaarsveld 4635 (NBG); Pearly Beach, littoral zone just above the water mark, rocky beach (–CB), Raitt 448 (NBG); Buffeljags, Bredasdorp Forest Reserve (– DA), Hugo 1636 (NBG); Agulhas, 1.3 km beyond lighthouse on dirt road westward (–DD), Taylor 10154 (NBG). 3420 (Bredasdrop): Cape Agulhas, (–CC), Taylor 4881 (NBG); Pillans 8144 (BOL); Salter 4126 (BOL); Taylor 1289 (PRE); Walsh s.n (NBG); Lekkerwater, first dune, 300 m E of strand house (–BC), Taylor 9906 (NBG). 2. Oncosiphon schlechteri (Bolus) Källersjö in Bot. J. Linn. Soc. 96(4): 312 (1988; Manning and Goldblatt, Plants of the Greater Cape Floristic Region: 401 (2012). Matricaria schlechteri Bolus, in Schltr. inBot. Jahrb. Syst. 27: 208 (1899); Hutch. in Bull. Misc. Inform. Kew: 251 (1917). Type: South Africa. Western Cape, Clanwilliam (3218): Lamberts Bay (–AB), 16 Aug 1896, Schlechter 8540 (BOL, holo.!). Erect perennial herb, to 0.3 m tall; stems cobwebbed. Leaves apically 3- to 5-fid sometimes simple, narrowly spatulate to linear, 5–15(20) × 1–3 mm, succulent, sparsely pilose; lobes lanceolate, 2 × 1 mm, apex blunt. Capitula discoid, homogamous, in simple, lax corymbs; peduncles slender, 1–15 mm long, cobwebbed. Involucre narrowly hemispherical, 3–5 mm diam., base cuneate in mature heads; involucre bracts 3-seriate, suberect at anthesis, several decurrent on the peduncle, apices ± level with receptacle apex, midvein slightly prominent, brown, glabrous, margins and apices narrowly scarious; outer series broadly ovate, ± 2.5 mm long; median series narrowly ovate, ± 3.5 mm long; inner series oblong, 2–3 mm long. Receptacle conical, slightly obscured by involucral bracts. Disc florets bisexual, corolla 2–3 mm long, yellow; tube conspicuously swollen from apex to the base, oblong, ± 1.5 mm long; limb campanulate, ± 1 mm long; lobes ovate, ± 0.5 mm long. Pappus inconspicuous to small, irregularly lobed, ± 0.3 mm long. Diagnostic characters Oncosiphon schlechteri shares the perennial habit (Fig. 1C), obtuse leaf lobes and decurrent involucral bracts with the other littoral species O. sabulosus but is distinguished by the apically 3- to 5-fid (Fig. 2C1) or sometimes simple leaves (pinnatifid to pinnisect in O. sabulosus), spatulate to linear in outline (oblong to obovate in outline in O. sabulosus), prominently cobwebbed peduncles and young stems (glabrous in O. sabulosus), and smaller involucre, ≤ 5 mm diam. (≥ 5 mm diam. in O. sabulosus), with a cuneate base and several involucral bracts decurrent on the peduncle (base round to cordate with some involucral bracts decurrent in O. sabulosus). Distribution and ecology This species is poorly collected and known from only a few localities in Namaqualand strandveld and coastal duneveld along the West Coast

of South Africa, from Lamberts Bay in the Western Cape to Hondeklip Bay in the Northern Cape (Fig. 5). Oncosiphon schlechteri and O. sabulosus are largely allopatric, although both species have been collected around Lamberts Bay. Flowering is in late spring (October to November). Additional specimens examined South Africa. Northern Cape: 3017 (Hondeklipbaai): Groen River Mouth (–DC), Le Roux and Parsons 79 (NBG); at highwater mark, coast 4 miles (6.5 km) S of Hondeklip Bay, (–DC), Pillans 18106; Pillans 18105 (BOL). 3117 (Lepelfontein): Namaqualand sandveld, near shore shingle beach on opposite side of bay to old geologist camp (–BD), Desmet 40 (PRE). Western Cape: 3118 (Vanrhynsdrop): Doringbaai area, Zeven Puts Farm (–CD), Boucher 4059 (NBG). 3. Oncosiphon grandiflorus (Thunb.) Källersjö in Bot. J. Linn. Soc. 96(4): 312 (1988); Goldblatt and Manning, Cape Plants: 345 (2000); Manning and Goldblatt, Plants of the Greater Cape Floristic Region: 401 (2012); Snijman, Plants of the Greater Cape Floristic Region: 307 (2013). Tanacetum grandiflorum Thunb. in Prodr. Pl. Cap. 2: 147 (1800); Thunb., Fl. Cap.: 642 (1823); DC., Prodr. Sys. Nat. 17: 132 (1838). Matricaria grandiflora (Thunb.) Fenzl ex Harv. in Harv. and Sond., Fl. Cap. 3: 166 (1865), nom. illeg., non Matricaria grandiflora (Willd.) Poir (1814); Levyns in Adamson and Salter, Fl. Cap. Penins.: 805 (1950). Pentzia grandiflora (Thunb). Hutch. in Bull. Misc. Inform. Kew.: 250 (1917); Hutch, Botanist in S. Afr.: 171 (1946). Type: South Africa, Thunberg s.n. in Herb. Thunb. 18979 (UPS-THUNB, holo.—microfiche!). Cotula tripinnata Thunb. in Prodr. Pl. Cap. 2: 162 (1800); Thunb., Prodr. Fl. Cap.: 696 (1823). Type: South Africa, Thunberg s.n. in Herb. Thunb. 20281 (UPS, holo.—microfiche!). Erect, aromatic annual herb, to 0.3 (0.5) m tall; stems sparsely pilose to pilose sometimes glabrous. Leaves bipinnatisect, lanceolate to oblanceolate, 10–40 (50) × 5–20 (30) mm, pilose; lobes oblong to lanceolate, 1–5 × ± 1 mm, mucronulate. Capitula discoid, homogamous, 6–15 mm diam., in simple, lax corymbs sometimes solitary; peduncles thickened toward the base of capitula, 10–80 (130) mm long, pilose, becoming densely pilose below capitula. Involucre broadly hemispherical, 5–14 mm diam., base cordate in mature heads; involucre bracts 3-seriate, slightly deflexed at anthesis, never decurrent on the peduncle, apices ± level with receptacle apex, midvein prominent sometimes inconspicuous, dark brown, tomentose sometimes sparsely tomentose, margins and apices broadly scarious; outer series ovate, ± 3.5 mm long; median series bract broadly oblong to oval, 3–4 mm long; inner series narrowly oblong, ± 4 mm long. Receptacle flat-convex, slightly obscured by involucral bracts. Disc florets bisexual, corolla 2–3 mm long, yellow; tube conspicuously swollen from apex to the base, ± 1.5 mm long; limb campanulate, ± 1.5 mm long; lobes ovate, ± 0.5 mm long. Pappus inconspicuous to small, irregularly lobed, ± 0.2 mm long. Diagnostic characters Oncosiphon grandiflorus shares large heads, N 5 mm diam., with prominently tomentose involucral bracts (Fig. 1F) and only slightly convex receptacles with O. intermedius. It is distinguished by the yellow disc florets with ovate corolla lobes, ± 0.5 mm long (Fig. 1E; 4C; orange in O. intermedius with lanceolate, corolla lobes ± 0.7–1.0 mm long). As in several of the other species however, the yellow florets may become somewhat amber coloured as they age. Where populations of O. grandiflorus and O. suffruticosus co-occur along the West Coast, occasional hybrid individuals may be found with lax, simple corymbs with capitula of intermediate size (± 0.5 mm diam.) and sparsely pilose to pilose involucral bracts (Kolokoto & Magee 3, NBG; Boucher 4073, NBG).

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Fig. 6. Distribution of Oncosiphon grandiflorus.

Distribution and ecology This species occurs in stony or sandy soil from southern Namibia through Namaqualand and the Bokkeveld Plateau in the Northern Cape along the West Coast and near-interior mountains to the Cape Peninsula in the Western Cape, with a single record from Danger Point near Caledon (Fig. 6). It is frequent in disturbed open areas often forming large colonies. Flowering is from late winter to late spring (August to November). Additional specimens examined Namibia. 2615 (Lüderitz): Lüderitz Bay (–CA), Merxmüller and Giess 3139 (PRE); Merxmüller and Giess 28280 (PRE, WIND); 35 km N of Rosh Pinah (on road to Aus), 2–4 km W of the Strasse (–CA), Leuenberger, Raus and Schiers 3199 (WIND); 3 miles [4.8 km] E of Kaukausib Fountain (–DC), De Winter and Guess 6109 (NBG). 2616 (Aus): granite koppie on Heinrichsfelde (–CA), Mannheimer CM2495 (WIND); just before the turn-off to Aus on Keetmanshoop road (–CA), Germishuizen 10380 (PRE). 2715 (Bogenfels): Blue ridges at Pomona pumphouse (–AB), Mannheimer and Maggs-Kölling CM2645 (WIND); Pomona (–AB), Dinter 4029 (PRE, SAM); 38 km S of Grillental (–AD), Merxmüller and Giess 28382 (WIND); Klinghardt Mtns (–BC), Smook 11165 (WIND); Merxmüller and Giess 32038 (WIND); Merxmüller and Giess 32071 (PRE, WIND); Müller 839 (WIND); Klinghardt Mountains, Nomitsas (–BD), Gess and Gess 02/03/24 (WIND); Bogenfels, at the junction to

Buntfeldschuh, in sandy rocky surface (–DA), Merxmüller and Giess 28334 (PRE, WIND); Boegoeberg (–DD), Strohbach 226 (WIND). 2716 (Witpütz): dunes just S of Tsaus waterhole, (–AA), Mannheimer and Mannheimer 313 (WIND); N of Klinghardt Mountains (–AC), Klaassen and Bartsch EK502 (WIND); E side of Klinghardt Mountains (–AC), Smook 11198 (PRE; WIND); Lüderitz, Diamond Area no:1, Inselberg of Klinghardt Mountain range (–AC), Germishuizen 10101 (PRE); Sperrgebeit, Chamnaib (–CA), Gess and Gess 02/03/10 (WIND); Karas, Chamnaib, SW foothills (–CA), Mannheimer CM2043 (WIND); Skorpion Mine (–DC), Burke 97178 (WIND); Rosh Pinah, W of Vliegveld (–DD), Zietsman 1832 (WIND); Bakenberg, Witpütz-sud, Zebrafontein Lus Farm (–DD), Merxmüller and Giess 28760 (WIND); Zebrafontein, Witpütz (–DD), Wiss 2058 (WIND). 2816 (Oranjemund): between Boegoeberg turnoff and Orangemund gate (–AA), Manneheimer CM1041 (WIND); 15 km W of Rosh Pinah on the way to Obibwasser (–BA), Giess 13,821 (PRE); Diamond Area no. 1, valley between Obib Mountain Peak and Obib dunes (–BA), Van Wyk 9035 (PRE); Obib dunes (–BA), Zietsman 1868 (WIND). South Africa. Northern Cape: 2816 (Oranjemund): Oranjemund, sandy flats near Orange River (–BB), Metelerkamp 291 (BOL); Lüderitz, Oranjemund, Stinkrant Farm (–CB), Merxmüller and Giess 2323 (NBG); Groot Derm Farm, ± 30 km E of Alexander Bay (–DA), Mayer 122500 (NBG); Richtersveld, 37 km N of Lekkersing (–DB), Germishuizen 5592 (PRE); Richtersveld, Annisvlakte (–DB), Jürgens 23,130 (PRE); Witbank, (–DC), Pillans 5137 (BOL); Pillans 5138 (BOL); 40 km from Port Nolloth on road to Alexander Bay (–DD), Germishuizen 4731A (PRE, BOL);

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Namaqualand, Holgat River (N of Port Nolloth), near main road, about 4 km from sea (–DD), Nicholas 2654 (PRE). 2817 (Vioolsdrif): Sesfontein (–BC), Maggs N.B.G.2572/36 (NBG); dry bed of Kalkfontein River S of Brakfontein (–CC), Pillans 5563 (BOL). 2916 (Port Nolloth): 16 km from Porth Nolloth, on road to Alexander Bay (–BB), Koekemoer 899 (PRE); Germishuizen 5323 (PRE; NBG); McDougall's Bay, 2 miles [3.2 km] S of Port Nolloth (–BD), Hardy 666 (PRE); (–BD), RodriguezOubina and Cruces 2006 (PRE). 2917 (Springbok): 60 km from Port Nolloth, on road from Steinkoft to Port Nolloth on Nuwefontein-seberg (–AA) Burgoyne 1332 (PRE); Namaqualand, Klipfontein (–BA), Hutchinson 894 (BOL); Farm Zonnekwa 328, 1 km [2 km] of crossroad to Graafwater and Sonnekwa B on road from Vaalkol (–CD), Le Roux and Lloyd 504 (NBG); Spektakel (–DA), Morris 5732 (BOL); Springbok, Hester Malan Nature Reserve (–DB), Le Roux 2516 (BOL, NBG); 4 miles (6.4 km) N of O'okiep (–DB), Taylor 4004 (NBG); Mesklip (–DD), Maguire 310 (NBG); 25 km from Springbok towards Spektakelberg Pass, road side (–DA), Glen 1457 (PRE). 2918 (Gamoep): Farm Koisabes, 17 km E from Concordia-Goodhouse road on Pofadder turn-off (–AB), Le Roux 2527 (NBG); near Ratelkraal (–CA), Maguire 321 (NBG); Springbok area, Goegap National Park (–CA), Zietsman 3852 (PRE); W of Gamoep, along Hytkoras road to Kamieskroon (–CC), Koekemoer 1156 (PRE). 3017 (Hondeklibaai): Namakwa National Park, in red Kalahari sand near Goedehoop (–AB), Koekemoer 2974 (PRE); Spoeg River, Wallekraal (–AD), Pillans 18102 (BOL); on slopes and hills around Houndeklip Bay (–AD), Pillans 18103 (BOL); Farm Oubees, 9.3 km W of Springbok, Soebatsfontein road, from turn-off to Oubees Farmhouse (–BA), Le Roux and Lloyd 739 (NBG); Farm Oubees 339, 1.2 km E of Oubees/Wildepaardehoek boundary fence, 4.7 km E of SpringbokSoebatsfontein road (–BA), Le Roux and Lloyd 660 (NBG); Farm Sandkraal, 40 km W of Garies (–DA), Hilton-taylor 1372 (NBG). 3119 (Calvinia): along R355 to Calvinia, just before the road to Kaggakama (–AA), Magee et al. 156 (NBG); Nieuwoudtville, Mentjieskraal (–AC), Maketter s.n (NBG); Calvinia, Bloedzuigersfontein (–BD), Compton 9639 (NBG); De Bosch, Calvinia (–BD), Levyns 1735 (BOL); Calvinia, Botterkloof Pass (–CD), Barker 10326 (NBG). Precise locality unknown: Richtersveld, Hardy 213 (PRE). Western Cape: 3118 (Vanrhynsdorp): Zandkraal, near Gifberg (–BC), Barker 5663 (NBG, BOL); Compton 20864 (NBG); Cohen's Farm, 1 mile (1.6 km) NW of Vredendal (–DA), Merwe 158 (NBG); Vanrhynsdorp, riverbank (–DA), Kolbe 14304; Kolbe 14303 (BOL); 2 miles [3.2 km] NE of Vrendendal, (–DA), Hall 3815 (NBG); Vanrhynsdrop, Wiedouw, under the Gifberg (–DB), Zietsman 1111 (PRE); between Vredendal and Lamberts Bay, Klipheuwel (–DC), Powrie 718 (NBG); Vleikraal, E of Klawer (–DC), Walters 108 (NBG); sandy flat between Driefontein and Heerenlogement (–DC), Pearson 6808 (BOL); Brandewyn River, Clanwilliam, (–DD), Compton 6896 (NBG). 3218 (Clanwilliam): Clanwilliam (–AB), Leipoldt 286 (BOL); Rooyen, Steyn and De Villiers 162 (NBG); Maguire 1053 (NBG); Clanwilliam sports field, (–AB), Goldbatt 3035 (NBG); Nortier experimental Farm, Lamberts Bay (–AB), Boucher 2571 (NBG); ½ mile [0.8 km] from Bergiver Station, along railway line next to Kersefontein turnoff (–AD), Boucher 80 (NBG); near edge of dam below Olifantsdam Motel (–BB), Taylor 10464 (NBG); Bidouw Valley (–BB), Middlemost 1906 (NBG); Clanwilliam, Rheboksvallei picnic site (–BB), Victor 670 (PRE); Clanwilliam (–BB), in hills around Clanwilliam (–BB), Keyroles 286 (BOL); near crossing of old Citrusdal/Clanwilliam road and Algeria road (–BD), Hugo 658 (NBG); 10 km before Clanwilliam coming from Citrusdal at T-junction (–BD), Van Slegeren and Van Der Walt MSLMW399 (NBG); along the N7 between Clanwilliam and Citrusdal (–BD), Koekemer 1403 (PRE; E base of Piketberg (–DA), Pillans 9830 (BOL); along N7 between Pools and Piekenierskloof (–DB), Taylor 10,745 (NBG); Eendekuil (–DB), Low s.n (NBG); Piekenierskloof (–DB), Barnes s.n (BOL); Olifantsrivier valley, in Clanwilliam (–DB), Levyns 1208 (BOL); near Clanwilliam, along Olifantsrivier (–DB), Rodin 30467 (BOL); S of Piquetberg (−DD), Barker 5770 (NBG). 3219 (Wuppertal): Pakhuis Pass, alongside Kliphuis River, 17 km from Clanwilliam Town

on R364, along roadside (–AA), Watson and Panero 94-44 (NBG); Stormsvlei, near Wuppertal (–AA), Leipoldt 512 (NBG); Cederberg State Forest, Heuningvlei (–AA), Kruger 1683 (NBG); Tankwa Karoo National Park, 4 km W of Varsfontein (–BA), Steyn 1132 (PRE, NBG); Tankwa Karoo National Park, S of Varsfontein (–BB), Rosch 469 (NBG); Môrester Farm, 7 km N of Elandsvlei (–BC), Hilton-Taylor 1832 (NBG); Theerivier, Citrusdal (–CA), Hanekom 1156 (NBG, PRE); Citrusdal Town area, open plot on the corner of De Klerk and Park Street (–CA), Hanekom 3103 (NBG); road to Calvinia from Ceres, road to Die Mond Farm (–DA), Cowell, Bennett and Pekeur MSBP 3785 (NBG); Swartruggens, Knolfontein, 60 km NE of Ceres (DC), Jardine 1400 (NBG); Jardine and Jardine 147 (NBG); Jardine and Jardine 498 (NBG); Jardine 1653 (NBG). 3220 (Sutherland): Tankwa Karoo National Park, Paulshoek Farm, 1st jeep track S of house into mountains, near dam (–AC), Koekemoer 2838 (PRE); Koekemoer 2839 (PRE); on veld, between Klipplaat and a point about 10 miles [17 km] to NE (–BC), Pearson 3305 (BOL). 3318 (Cape Town): Darling, Platteklip (–AD), Liebenberg 8273 (PRE); roadside on R27 N of Jakkalsfontein (–AD), Goldblatt and Manning 10357 (NBG); Saron Mission Station near Porterville (–BC), Catterd E (Sheet 2) (NBG); Cape Peninsula, Kirstenbosch, new clearing (–CD), Compton 14145 (NBG); Rondebosch, Linkoping Road (–CD), M. R. L 7557 (BOL); Paardeberg, between Wellington and Malmesbury, Waterkloof, ± 1.8 km NNW of Ayama Farm house (–DB), Nicolson and Roets 661 (NBG); Kuilsrivier sand flats (–DD), Raitt s.n (NBG). 3319 (Worcester): Tanqua Karoo, Ceres (–AD), Compton 7913 (BOL, NBG); Ceres, Tanqua Karoo, near road (–BD), Levyns 11358 (BOL); Worcester, Riverside (–CB), Walters 947 (NBG). 3418 (Simonstown): Cape Peninsula, near Retreat station (–AB), Salter 7132 (BOL); at eastern base of the Muizenberg (–AB), Page 3497 (BOL); Cape of Good Hope (–AB), Moss 5730 (BOL); Salter 7113 (BOL). 3419 (Caledon): Caledon, Danger Point (–CB), Pillans 9773 (BOL). Precise locality unknown: next to high power line, Strohbach 254 (WIND). 4. Oncosiphon intermedius (Hutch.) Källersjö in Bot. J. Linn. Soc. 96(4): 312 (1988); Manning and Goldblatt, Plants of the Greater Cape Floristic Region: 401 (2012). Pentzia intermedia Hutch. in Bull. Misc. Inform. Kew. 10: 249 (1917). Type: South Africa, Anon. s.n (K, lecto.–image!, designated by Källersjö, 1988). [Matricaria grandiflora var. β in Harv. and Sond., Fl. cap. 3: 166 (1865)]. Erect, aromatic annual herb, to 0.3 m tall; stems sparsely pilose to pilose sometimes glabrous. Leaves bipinnatisect, lanceolate to oblanceolate, 10–40 (70) × 5–20 (30) mm, pilose; lobes oblong to lanceolate, 1–5 × ± 1 mm, mucronulate. Capitula discoid, homogamous, 5–18 mm diam., in simple, lax corymbs sometimes solitary; peduncles thickened toward the base of capitula, 10–80 (110) mm long, pilose, becoming densely pilose below capitula. Involucre broadly hemispherical, 5–15 mm diam., base cordate in mature heads; involucre bracts 3-seriate, slightly deflexed at anthesis, never decurrent on the peduncle, apices ± level with receptacle apex, midvein prominent, blackbrown sometimes green, tomentose, margins and apices narrowly scarious; outer bracts narrowly oblong, ± 4 mm long, middle bract lanceolate to narrowly oblong, 4–5 mm long, inner bracts narrowly lanceolate ±5 mm long. Receptacle flat-convex, slightly obscured by involucral bracts. Disc florets bisexual, corolla 3–4.5 mm long, orange; tube conspicuously swollen from apex to the base, ± 1.5 mm long; limb campanulate, ± 1.5 mm long; lobes narrowly lanceolate,± 0.7–1 mm long. Pappus inconspicuous or small, irregularly lobed ± 0.3 mm long. Diagnostic characters Oncosiphon intermedius is readily distinguished from others in the genus by the orange disc florets (Fig. 1E) and prominently lanceolate corolla lobes (± 0.7–1 mm long; Fig. 4B) which give the heads an irregular outline. All other species have yellow disc florets, sometimes fading

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amber with age, with ovate lobes (± 0.5 mm long). Oncosiphon intermedius is evidently related to O. grandiflorus, with which it shares large capitula (N5 mm diam.), prominently tomentose involucral bracts (Fig. 1F), and only slightly convex receptacles (Fig. C). Distribution and ecology Oncosiphon intermedius is largely restricted to the Bidouw and Wuppertal valleys in the Cederberg Mountains of the Western Cape Province. It has however also been collected slightly north on the Bokkeveld Escarpment of the Northern Cape Province (Fig. 5). It favours sandy soils with some disturbance. Flowering is in spring (September to October). Additional specimens examined South Africa. Northern Cape: 3119 (Calvinia): sandy soil near river, Lokenburg, 21 m S of Nieuwoudtville (–CA), Story 4427 (PRE). Western Cape: 3219 (Wuppertal): after turnoff from Wuppertal to Bidouw Valley, red sandy valley (–AA), Mauve and Oliver 92 (NGB); N slopes of Wuppertal (–AA), Goldblatt and Manning 13250E (NBG); Wuppertal road, near Pakhuis Pass (–AA), Levyns 11207 (BOL); Welbedacht, Bidouw Valley (–AA), Maguire 1859 (NBG); Compton 6908 (BOL); Leivis 22,178 (BOL); Clanwilliam, Mertenhof in Bidouw (–AA), Van Breda 4266 (PRE); Bidouw (–AB), Bolus s.n (BOL); van Rooyen, Steyn and De Villiers 700 (NBG); Bidouw Valley, Zandrift Farm (–AB), Welman 136 (PRE); Bidouw Valley (–AB), Maguire 1876 (NBG, BOL); Clanwilliam, Hoeck (–AB), Schlechter 8705 (BOL); Cederberg, Matjiesrivier Nature Reserve, Wildehondshoek (–AD), Lechmere-Oertel 846 (NBG); Clanwilliam, Matjiesrivier Nature Reserve, Sandleegte A (–AD) Low 7778 (NGB); Cederberg, Matjiesrivier (–AD), Wagener 274 (NBG); Zuurfotein, Ramkraal, S of hut (–AD), Low 7530 (NBG); Cederberg, Oudekraal Farm, between Ramkraal and Oukraal (–AD), Koekemoer 2458 (PRE). 5. Oncosiphon piluliferus (L.f.) Källersjö in Bot. J. Linn. Soc. 96(4): 312 (1988); Manning and Goldblatt, Plants of the Greater Cape Floristic Region: 401 (2012); Snijman, Plants of the Greater Cape Floristic Region: 307 (2013). Cotula pilulifera L.f., Suppl. Pl.: 378 (1781). Matricaria pilulifera (L.f) Druce. in Rep. Bot. Soc. Exch. Club Brit. Isles for 1916: 635 (1917). Pentzia pilulifera (L.f.) Fourc. in Trans. Roy. Soc. South Africa 21: 87 (1932). Type: South Africa. Western Cape, Cape (3321): Laingsburg-Prince Albert div., National Road over Dwyka River (–BA), 07 Jun 1962, Nordenstam 161 (S, neo.– image!, designated by Källersjö, (1988); K–image!, isoneo.). Tanacetum obtusum Thunb. in Prodr. Pl. Cap. 2: 147 (1800); Thunb., Fl. Cap.: 641 (1823). Type: South Africa, Thunberg s.n. in Herb. Thunb. 18984 (UPS, holo.–microfiche!). Cotula globifera Thunb. in Prodr. Pl. Cap. 2: 162 (1800); Thunb., Fl. Cap.: 969 (1823). Tanacetum globiferum (Thunb.) DC. in Prodr. Sys. Nat. 17: 132 (1838). Matricaria globifera (Thunb.) Fenzl ex Harv. in Harv and Sond., Fl. Cap. 3: 165 (1865). Pentzia globifera (Thunb.) Hutch. in Bull. Misc. Inform. Kew: 251 (1917). Type: South Africa. Western Cape, Cape Town (3318), Swartland: ‘Groenekloof’ [Mamre], (–CB), Thunberg s.n. in Herb. Thunb. 20220 (UPS, holo.– microfiche!). Erect, strongly aromatic annual herb, to 0.3 m tall; stems sparsely pilose to pilose sometimes glabrous. Leaves bipinnatisect sometimes tripinnatisect, lanceolate to oblanceolate, 10–25 × 5–15 mm, sparsely pilose to pilose; lobes oblong to oblanceolate, 1–3 × ± 1 mm, mucronulate. Capitula discoid, homogamous, 5–10 mm diam., in simple, lax corymbs sometimes solitary; peduncles slender, 5–50 mm long, glabrous to sparsely pilose. Involucre hemispherical, 5–10 mm diam., base deeply cordate at anthesis; involucral bracts 3-seriate, strongly deflexed at anthesis, never decurrent on the peduncle, apices level with

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receptacle base, midvein prominent to inconspicuous, dark brown or green, glabrous, margins and apices narrowly scarious; outer bracts oblong to narrowly elliptical, 2.5–3.5 mm long; middle bract oblong to broadly oblong, 2.5–3 mm long; inner bracts obovate, ± 2.7 mm long. Receptacle conical, not obscured by involucral bracts. Disc florets bisexual, corolla 2–3 mm long, yellow; tube conspicuously swollen from the apex to the base, lanceolate to oblong, 1–2 mm long; limb campanulate, ± 1 mm long; lobes ovate, ± 0.5 mm long. Pappus inconspicuous to small, irregularly lobed, ± 0.1 mm long. Diagnostic characters Oncosiphon piluliferus is recognised by the simple corymbs (rarely solitary) of moderately-sized, hemispherical capitula with glabrous involucre 5–10 mm diam (Fig. 3A, B). It is most likely to be confused with O. suffruticosus, also with a glabrous involucre that are campanulate 3–5 mm diam. with smaller, capitula in dense, congested corymbs. The two species differ also in the shape of the involucre. In O. piluliferus the base of the involucre is deeply cordate due to the strongly deflexed bracts, which expose the receptacle (Fig. 4J) whereas in O. suffruticosus the base of the involucre is round to shallowly cordate and the involucral bracts only slightly deflexed, thus obscuring the receptacle. Distribution and ecology Oncosiphon piluliferus is widely distributed in the drier parts of South Africa, occurring from Namaqualand to Worcester, eastwards to East London and extending north through the Free State and Mpumalanga to Pretoria (Fig. 7). The species favours sandy soils on roadsides or or other disturbed areas. Flowering is from late winter to early summer (August to December). Additional specimens examined South Africa. North West: 2725 (Bloemhof): Leeuwfontein, 18 km W of Wolmaransstad (–BB), Van Wyk 1779 (PRE). GAUTENG: 2528 (Pretoria): Pretoria (–CA), Mogg 16609 (PRE). FREE STATE: 2727 (Kroonstad): Kroonstad, Kroonheuwel, Johnny's dam (–CA), Siebert and Siebert 1991 (PRE). 2925 (Jagersfontein): Fauresmith railway station grounds (–CB), Smith 986 (PRE). 2926 (Bloemfontein): Bloemfontein (–AA), Van Wyk 5269 (PRE); Pott 2077 (NBG). 3324 (Ladismith): Fouriesburg, Ficksburg (–DD), Rogers 15920 (BOL). Northern Cape: 2824 (Kimberley): Kimberley (–DB), Moran 15,856 (BOL). 2918 (Gamoep): E of Springbok, Dikbek Farm (–DA), Crosby 688 (PRE); between Gamoep and Aggeneys, SE of Springbok (–DC), Koekemoer 2523 (PRE). 3018 (Kamiesberg): Platbakkies, E of store (–BC), Welman 00076 (PRE). 3019 (Loeriesfontein): mountain pass, S of Klipplaat (–CC), Pearson 3962 (BOL); Loeriesfontein road, 30 miles [48.3 km] from Calvinia (–CD), Johnson 594 (NBG). 3025 (Colesburg): Middelburg, Grootfontein, waste ground (–CD), Leistner 489 (PRE); Theron 302 (PRE); Comins 841 (PRE); Middelburg (–CB), Hauafau 1394 (PRE); Middelburg Road Railway station (–CD), Flanagan 1390 (NBG, SAM). 3118 (Vanrhynsdorp): 14 km S of Nuwerus (AB), Hugo 2904 (NBG). 3119 (Calvinia): Calvinia, Akkerendam (–BD), Taylor 2748 (NBG, BOL); Calvinia (–BD), Hanekom 2494 (PRE); Beli, Elandsvlei and Die Bos, Calvinia (–DC), Barker 8296 (NBG). 3120 (Williston): Calvinia, Sandwerf on road to Brandvlei (–AC), Germishuizen 4009 (PRE, NBG). 3121 (Fraserburg): Fraserburg, Rietvlei (–AC), Shearing 74 (PRE). 3124 (Hanover): Graaff-Reinet, Farm Blue Gum house, along top of streambank (–DD), Retief and Reid 544 (PRE) 3220 (Sutherland): Succulent Karoo around Farmhouse “Clifton”, Farm Windhoek 112 (–AD), Clark and O'Connor 165 (NBG); Klein Roggeveld, Oranjefontein (–DC), Compton 8126 (NBG); Roggeveld, Soekop, Grasvlake camp (–AA), Rösch 743 (NBG); Rösch 277 (NBG). Western Cape: 3219 (Wuppertal): Cederberg, Matjesrivier (–AD), Wagener 275 (NBG). 3218 (Clanwilliam): Piketberg, plateau on the

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Fig. 7. Distribution of Oncosiphon piluliferus.

Elandsberg (–AD), Pillans 7964 (BOL). 3221 (Merweville): Layton, river bed Rietvlei weir (–BB), Shearing 1341 (PRE). 3222 (Beaufort West): Karoo National Park, Middle Plateau (–AD), Rubin and Sachse 706 (PRE); Beaufort West (–BC), Cooper 186 (BOL); Beaufort West, Sunnyside on river bed (–BC), Esterhuysen 2744 (BOL); Beaufort West, Farm Aardoons, at windmill near farmhouse (–DD), Retief and Reid 145 (PRE). 3319 (Worcester): Bokkveld Karoo (–BB), Marloth 12532 (NBG); Ceres, Hottentots Kloof(–BC), Compton 1206 (NBG); along N1, top of Hex River Valley, Bo-Plaas around cellphone transmitter (–BD), Botha (3) 131 (NBG); Ceres, Tankwa Karoo (–BD), M.R.Levyns 5085 (BOL); Lemoenpoort Farm of Mr. P Viljoen (–CD), Walters 2186 (NBG); Worcester, open area between Federated Timbers and locomotive sheds (–CB), Walters 601 (NBG); Worcester (–CB), Olivier 159 (NBG); Bokkeveld, Ceres (–CB), Joubert 5 (NBG); Karoo Garden, Worcester (–CB), Compton 18651 (NBG, BOL); Karoo Garden veld (–CB), Dobay 50 (NBG); natural grown part of Karoo Botanic Garden outside Worcester (–CB), Van Slangeren, Townsend and Young MSBYJT384 (NBG); Worcester, Karoo Desert National Botanical Garden (–CB), Joffe 1107 (PRE); main road passing De Doorns (–BC), Mauve and Oliver 139 (NBG); Worcester, Wolvekop (–CB), Compton 12,774 (NBG); Private Nature Reserve on Farm Doringkloof, southern foothills of Voetpadsberg (–DA), Morley 351 (NBG); 20.5 km S of N1 on the road to Montague (R318), on E of highway (–DB), Bayer and Chandler SAF-01134 (PRE). 3320 (Montagu): Avondrus Farm, 30 km SE of Touwsrivier (–AC), Hilton-Taylor 1939 (NBG); Touwsrivier, Die Vlakte (–AC), Theron 7 (NBG); Laingsburg, Karoo Garden, Whitehill (–BA), Compton 7768

(NBG); Compton 11817 (NBG); Compton 3545 (BOL); Compton 2924 (BOL); veld infront of Matjiesfontein Hotel (–BA), Magee and Boatwright 173 (NBG); Laingsburg (–BB), Bohren 9131 (NBG); Montagu, roadside (–CC), Levyns 429 (BOL); Hot baths, near Montagu (–CC), Page 15443 (BOL). 3321 (Ladismith): Fisantrivier, 40 km SE of Sutherland (–AC), Moffett 3765 (NBG); Farm Buffelsdrif, 50 m N of road 1.9 km E of rail crossing at Winkelplaas (–CA), Laidler 237 (NBG); Farm Buffeldrif, mid-slope of hill S of Winkelplaas road, 1.5 km from junction with Barrydale-Ladismith road (–CA), Laidler 234 (NBG); Muiskraal, along main road to Ladismith on open ground (–CC), Bohnen 9131 (NBG); Klein Karoo by Muiskraal (–CC), Bohnen 8403 (NBG); Muiskraal, 10 km N of Garcia's Pass near dam (–CC), Mauve, Van Wyk and Pare 6 (NBG). 3322 (Oudtshoorn): Prince Albert, Tierberg study site, on disturbed silty soil of termite mound and on alluvium near drainage line (–AB), Dean 1028 (BOL); Dean 532 (BOL); Schoeman's Poort, 13 miles [20.9 km] from Oudtshoorn (–AC), Heginbotham 224 (NBG); lower northern slopes of Swartberg, Farm Frisgewaagd (–AD), Vlok 1607 (PRE); on the Meiringspoort road, 12 km from Klaarstroom, alongside road and next to river (–BC), Watson and Panero 94-114 (NBG); Oudshoorn, Kombuis Camp (–BC), Bolus 12015 (BOL); Oudtshoorn (–CA), Hall s.n. (NBG); Rogers s.n. (BOL); on lands of Olifantsrivier, 13 miles [20.9 km] E of De Rust (–CA), Theron 2037 (PRE); 60 km past Hartenbosch to Calitzdrop near Volmoed (–CA), Venter 1411 (PRE); between George and Oudtshoorn (–CD), Rycroft 2620 (NBG); Wildernis National Park, Rondevlei towards office (–DC), Rubin 498 (PRE); 3419 (Caledon): Riviersonderend (–BB), Schlechter 5624 (NBG, BOL);

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Swellendam, E of Bushman's River (–BB), Maguire 530 (NBG); Napier (– BD), Jardaan 175 (NBG); Albany, E of Bushman's River (–DA), Maguire 530 (NBG); 50 km beyond Gansbaai on road to Bredasdrop at roadside (–DA), Venter 1318 (PRE). 3420 (Bredasdrop): Bredasdrop, 3 km N of gates of De Hoop Nature Reserve (–AD), Mauve, Reid and Wikner 64 (NBG); Malgaspont (–BC), Hugo 1950 (NBG); De Hoop-Potberg Nature Reserve, Potberg Farm, along track to coast, 1.5 km from the coast (–BC), Burgers 1549 (NBG); Cape Algulhas (–CC), Pillans 8142 (BOL). 3421 (Riversdale): Blombos Farm, near Cape Infanta (–AC), Walters 1580 (NBG); 34 km from Riversdale on road to Blombos (–AC), Germishuizen 4208 (PRE’); Stilbaai, Geelkrans Nature Reserve (–AD), Van der Merwe 101 (NBG). 3422 (Mossel Bay): Mossel Bay, Hartenbos River Mouth (–AA), Parsons 494 (NBG); Parsons 343 (NBG); Mossel Bay (–AA), Steyn s.n. (NBG). Eastern Cape: 3026 (Aliwal North): Bethulie, between the rivers park (–AC), Roberts 5542 (PRE); Aliwal North (–DA), Gerstner 141 (PRE). 3125 (Steynsburg): White Ridge, Middelburg (–AC), Duthie s.n. (NBG); Steynsburg, Farm Bulhoek, at house of Paul Kruger's birth (–BC), Retief and Germishuizen 282 (PRE). 3126 (Queenstown): Molteno, Bamboeshoek (–AC), Balsinhas 3336 (PRE); Queenstown (–DD), Sidey 3778 (PRE). 3222 (Beaufort West): Courlands Kloof, near Nelspoort (–BB), Pearson 1464 (SAM). 3223 (Rietbron): Rietbron, back street along water furrow (–CC), Retief and Reid 485 (PRE). 3224 (Graaff-Reinet): Graaff-Reinet, Karoo National Reserve, windmill valley (–AD), Linger 2115 (PRE); Graaff-Reinet (–BC), Bolus s.n. (BOL); Graaff-Reinet to Jansenville, turn left at Pearson road, left at Petersburg road for 38 km to main gate Sante Sana (–BD), Nienabar EN579 (PRE). 3225 (Somerset East): Graaff-Reinet, Sante Sana Game Farm at Petersburg settlement (–AC), Meyer 4171(2) (PRE); Cradock, National Bergkwagga Park (–AD), Zietsman 1311 (PRE); Cradock, National Bergkwagga Park, between Weltevrede and Berghof's huishekke (–AD), Liebenberg 7601 (PRE); Cradock (–AD), Van Ham 18831 (PRE). 3226 (Fort Beaufort): Tankastad-Queenstown (–AB) Pole-Evan 1760 (PRE); Fort Hare, Honeydale Farm (–DD), Russel 3334 (BOL). 3228 (Butterworth): Kei River (–CA), Flanagan 1057 (BOL, SAM). 3323 (Willowmore): road to Willowmore, 3.5 miles [5.6 km] from Uniondale (–AD), Fourcade 4674 (NBG); Uniondale (–CA), Fourcade 1162 (BOL). 3325 (Port Elizabeth): Uitenhage, Groendal Wilderness Res, Zunga catchmbasin, Farm Deysels Kraal, Great Winterhoek Mtns (–CA), Scharf 1992 (PRE); Uitenhage (–CD), Ecklon 223 (BOL); Markman Industrial Area (–DC), Dahlsstrand 3152 (PRE); Port Elizabeth, Zwartkops River estuary, 2 km below Redhouse (–DC), Glen 856 (NBG). 3326 (Grahamstown): Glen Melville, Ecca River Valley (–BA), Taylor and Edwards 8758 (PRE); Botha Hill (–BC), Schlechter 6085 (BOL); Grahamstown, Oatlands (–BC), J.R and B.R 192 (BOL); Alexandria, Bushman's River Mouth (–DA), Barker 1166 (NBG). Precise locality unknown: Kenhardt, Middlemost s.n. (NBG). 6. Oncosiphon suffruticosus (L.) Källersjö in Bot. J. Linn. Soc. 96(4): 312 (1988); Goldblatt and Manning, Cape Plants: 345 (2000); Manning and Goldblatt, Plants of the Greater Cape Floristic Region: 401 (2012); Snijman, Plants of the Greater Cape Floristic Region: 307 (2013). Tanacetum suffruticosum L. in Sp. Pl. 2: 843 (1753); Thunb., Prodr. Fl. Cap. 2: 641 (1823). Cotula tanacetifolia L. in Syst. Nat. 2: 564 (1767), nom. illeg. superfl. Matricaria suffruticosa (L.) Druce. in Rep. Bot. Exch. Cl. Brit. Isles. 3 (5): 421 (1914); Levyns in Adamson and Salter, Fl. Cap. Penins.: 804 (1950). Pentzia tanacetifolia Hutch. in Bull. Misc. Inform. Kew: 253 (1916), nom. illeg. Pentzia suffruticosa (L.) Hutch., Botanist S. Afr.: 171 (1946): Merxm. in Mitt. Bot. Staatssamml. München: 486 (1967). Type: South Africa, Anon s.n. in Herb. Linn. No. 987.11 (LINN, lecto.–image!, designated by Källersjö, 1988). Tanacetum multiflorum Thunb. in Prodr. Fl. Cap. 2: 642 (1823); DC., Prodr. Sys. Nat. 17: 132 (1838). Matricaria multiflora (Thunb.) Fenzl ex Harv. in Harv. and Sond., Fl. Cap. 3: 166 (1865). Type: South Africa, Swellendam, Thunberg in Herb. Thunb. No. 18983 (UPS, holo.– microfiche!).

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Tanacetum leptolobum DC. in Prodr. Sys. Nat. 17: 133 (1838). Matricaria multiflora var. leptoloba (DC.) Harv. in Harv. and Sond., Fl. Cap. 3: 166 (1865). Type: South Africa, on the banks of Gariep, 1835, Drège 2855 (G-DC, holo.–image!; HAL–image!, P (3 sheets)–image!; TUB– image!, iso.). Erect, strongly aromatic annual herb, to 0.5 m tall; stems sparsely pilose to pilose or sometimes glabrous. Leaves bipinnatisect to tripinnatisect, lanceolate to oblanceolate, 10–40 × 5–15 mm, sometimes succulent, sparsely pilose to pilose; lobes oblong to lanceolate, 1 × ± 1 mm, mucronulate. Capitula discoid, homogamous, congested, compound corymbs; peduncles slender, 5–10 mm long, glabrous to sparsely pilose. Involucre campanulate, 3–5 mm diam., base rounded or sometimes shallowly cordate in mature heads; involucral bracts 3-seriate, slightly deflexed at anthesis, never decurrent on the peduncle, extending above receptacle apex, midvein inconspicuous, glabrous, margins and apices broadly scarious; outer series ovate to broadly ovate, ± 1.8 mm long; median series oblong to broadly elliptical, 2–3 mm long; inner series bracts elliptical, ± 2 mm long. Receptacle conical, obscured by involucral bracts. Disc florets bisexual, corolla 1.0–2.5 mm long, yellow; tube conspicuously swollen from apex to base, lanceolate to oblong, ± 1.4 mm long; limb campanulate, ± 1 mm long; lobes ovate, ± 0.5 mm long. Pappus inconspicuous to small, irregularly lobed, ± 0.1–0.2 mm long. Diagnostic characters Oncosiphon suffruticosus is readily confused with Oncosiphon piluliferus, with which it shares bipinnatisect leaves (Fig. 2A) and glabrous involucral bracts, but is distinguished by the dense, compound corymbs (Fig. 3C) with ≥10 heads (solitary or in simple corymbs with ≤6 heads in O. piluliferus), smaller campanulate heads, 3–5 mm diam. (larger, hemispherical head, 5–10 mm in diameter, in O. piluliferus), and involucral bracts that are only slightly deflexed and so obscure the receptacle (Fig. 4I; involucral bracts strongly deflexed, not obscuring the base of the receptacle in O. piluliferus). Occasional hybrids between O. suffruticosus and O. grandiflorus were observed throughout the West Coast where the two species commonly co-occur. The hybrid individuals have intermediately sized (± 6 mm diam.), sparsely pilose to pilose heads, in lax simple corymbs (Kolokoto & Magee 3, NBG; Boucher 4073, NBG). Distribution and ecology This species is widely distributed along the coast in the winterrainfall region of southern Africa, occurring from southern Namibia through Namaqualand, in the Northern Cape, to Gansbaai, in the Western Cape (Fig. 8). The species is very common, growing in sandy soils often right up to the beach where it receives regular salt spray. It also favours disturbance, where it occurs in large numbers. Flowering is in spring to early summer (September to December). Additional specimens examined Namibia. 2716 (Witputz): Rosh Pinah, Namuskluft Farm (–DD), Van wyk 8837 (WIND); Mittendorf 37 (WIND); Mittendorf 115 (PRE). 2816 (Oranjemund): Numees experimental site, (–BD) Jurgens 10046 (NBG). South Africa. Northern Cape: 2816 (Oranjemund): Richstersveld National Park, top of Helshoogte Pass, in road cutting approximatly half way up pass (–BD), Bergh 2109 (NBG); Numees copper mine, plants growing near mine shaft entrance (–BD), Nicholas 2491 (PRE); Numees (–BD), Koekemoer 922 (PRE); head of Helskloof, Hottentotsparadyskloof (–BD), Thompson and Le Roux 95 (NBG); Swartwater, Richtersveld (–DD), Herre s.n. (NBG). 2817 (Vioolsdrif): Kouamsrivier, nearby Oranjerivier (–AA), Dreyer 136 (PRE); SW of Kodasprek (–AA), Thompson and Le Roux 253 (NBG); N foot of Rosyntjieberg (–AC), Thompson

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Fig. 8. Distribution of Oncosiphon suffruticosus.

175 (NBG); Ritchtersveld National Park, Die Koei (–AC), Zietsman 3834 (PRE); large kloof on N side N of Lelieshoek (–AC), Oliver, Tölken and Venter 369 (NBG); Zebrakloof, NE of Rosyntjieberg (–AC), Germishuizen 5561 (PRE); Richtersveld, kloof by Doomport (–AC), Herre s.n (NBG); Die Toon, E of Ritchtersveld National Park, (–AD) Williamson 5791 (NBG); Ploegwater, S portion of Ploegberg (–CA), Germishuizen 5482A (PRE); among Arundo in pool at Brakwater (–CC), Pearson 6072 (NBG). 2916 (Port Nolloth): Port Nolloth (BD), Venter 9124 (PRE). 2917 (Springbok): Klipfontein, Steinkopt (–BA), Herre s.n. (NBG); Hutchinson 890 (BOL); Kosies (–BA), Williamson 3940 (NBG); Eenriet Mountain, about 6 miles [7 km] N of Steinkopf (–BA), Wisura 2269 (NBG); 10 km S of Kleinsee, on way to Hondeklipbaai (–CA), Hugo 2861 (NBG); Spektakelpas, Farm Eselsfontein (–CA), Zietsman 867 (PRE); to foot of Wilderperdehoekpas and Soebatsfotein side (–CA), Zietsman 968 (PRE); Okiep (–DB), Morris 5733 (BOL); 3 km E of Springbok, disturbed area along road (–DB), Zietsman 668 (PRE); 7 km S of Springbok, on the Springbok camping ground (–DB) Greuter 21671 (PRE); Springbok, road border (–DB), Schlieben 11520 (NBG); 5 miles [8 km] N of Komaggas (–DB), Johnson 209 (NBG); Deurdrif Farm 219, 8 km S of Springbok (–DD), Bredenkamp 2144 (PRE); 6 miles [9.6 km] from Springbok, on Leliefontein road, ½ mile [1.6 km] off road, in hills near dry river bed (–DD), Thompson 463 (NBG); Hester Malan Nature Reserve, Springbok (–DB), Le Roux 2519 (NBG, BOL). 2918 (Gamoep): Namaqualand, near Ratelkraal (–CA), Maguire 316 (NBG). 2919 (Pofadder): Pofadder (–AB), Conradie 49 (NBG). 3017 (Hondeklipbaai): Namakwa National Park, between Riethuis and Taaibosduin, on granite

hill, foot of the hill (–AB), Koekemoer 3324 (PRE); Hondeklipbaai, sands near to the sea (–AD), Magee and Boatwright 333 (NBG); Kookfontein Farm, 5 km N of Soebatsfontein (–BA), Hilton-Taylor 1257 (NBG); Oubees Farm, 9.3 km W of Springbok to Soebatsfontein road, from turn-off to Oubees Farmhouse (–BA), Le Roux and Lloyd 740 (NBG); Kamiesberg, Skilpad Wildflower Reserve, front of the house (–BB), Cruz 71 (NBG); between Bowesdorp and Springbok (–BB), M.R.L 6951 (BOL); near Garies (–DB), Rodin 1378 (PRE, BOL); Garies, 1 mile [1.6 km] S of town (–DB), Hardy 493 (PRE); near road, 3 km S of Island Point, 200 m from coast (–DC), O'Callaghan,Van Wyk and Morley 157 (NBG); 9 km W of Nariep, Farm Roodeheuwel (–DC), Perold 1648 (PRE); Farm Ou Dam, 12,5 km NE of Kotzerus (–DD), Hilton-Taylor 1477 (NBG). 3018 (Kamiesberg): Kamiesberg (–AC), M.R.L 4080 (BOL); 5.6 miles [9 km] from Bitterfontein to Poffader, 1 mile [1.6 km] off road in kloof near wheat fields, broken Namaqualand veld dry hills (–CC), Thompson 396 (NBG). 3120 (Williston): near Williston, roadside (–BC), Bayliss and Bri 558 (PRE). 3220 (Sutherland): Tanqua Karoo, Tankwa National Park, Maansdam (–AA), Rösch 782 (NBG). Western Cape: 3118 (Vanrhynsdrop): Meerhofkasteel Farm, 15 km W of Nuwerus (–AA), Hilton-Taylor 1230 (NBG); 10 km from Koekenaap to Nuwerus (–AD), Stirton 10,135 (NBG); Vanrhynsdrop, Knersvlakte (–BC), Compton 20866 (NBG); Schlechter 8159 (BOL); Vredendal, Geelwal Farm (–CA), Le Roux and Ramsey 101 (NBG); Koekenaap, coast between Olifants River and Brand se Baai (–CA), Desmet 3601 (NBG); Vredendal (–CB), Krohn 6 (NBG); 13 km from Van Rhynsdrop to Nieuwoudtville (–DB), Magee and Boatwright 355 (NBG); National

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Road near Klawer, disturbed road verge (–DC), Thompson 2314 (NBG). 3119 (Calvinia): Meulsteenvlei, near Nieuwoudtville (–AC), Taylor 3928 (NBG); Nieuwoudtville Reserve, on slopes in red clay amongst boulders (–AC), Perry and Snijman 2399 (NBG); Nieuwoudtville (–AC), Makotter s.n. (NBG); Nieuwoudtville Reserve, on Dwyka tillite flats (–AC), Perry and Snijman 2452 (NBG); Calvinia, Akkerendam (–BD), Taylor 2751 (NBG); near Calvinia on Clanwilliam road (–BD), M.R.L 5077 (BOL). 3217 (Vredenburg): Vredenburg (–DD), Botha and Coetzee 1701 (PRE). 3218 (Clanwilliam): Wadrifsoutpan, W of railway (–AB), O'Callaghan, Van Wyk and Morley 102 (NBG); coast, inland dunes, S of Elandsbaai (–AD), Goldblatt and Porter 13,678 (NBG); Clanwilliam, Haas Drift, N facing slope of small kloof (–BB), Levyns 1285 (BOL); 3.6 miles [5.7 km] from Clanwilliam, W dry slopes of Pakhuis Pass (–BB), Marsh 368 (NBG); Velddrif, beside dirt road, E of pan, N of house (–CB), Brandt 11 (NBG); 2 km S of Dwarskersbos, on allotment Soverby (–CB), Boucher 4080 (NBG); Velddrif, Rocher Pan Nature Reserve, left and right of W track, (–CB), Rooyen and Ramsey 145 (NBG); Velddrif, Rocher Pan Nature Reserve (–CB) Le Roux and Ramsey 340 (NBG); near Velddrif (–CC), Mostert U35 (NBG); S bank of Berg River, ± 3 km of rail bridge (–CC), O'Callaghan 1255 (NBG); Eendekuil, by railway line (–DB), Howes 185 (PRE); Piketberg (–DC), Guthrie s.n. (NBG); along W coast road, near Pools (–DD), Magee and Boatwright 162 (NBG); Piketberg, N side of road, top of Versveld's Pass (–DD), Pillans 7165 (BOL). 3219 (Wuppertal): Bidouw Valley (–AB), Middlemost 1894 (NBG); lower slopes of Bavianskop above Olifantsdam Motel (–BB), Taylor 10461 (NBG). 3317 (Saldanha): Langebaan, 1 km S of Perlemoenpunt (–BB), O'Callaghan and Steensma 1717 (NBG); Saldanha Bay, rocks by shore (–BB), Compton 17885 (NBG); granite koppie just outside Saldanha (–BB), Mauve 5341 (PRE). 3318 (Cape Town): Langebaan, hill S of village (–AA), Manning 2926 (NBG); Langebaan (–AA), Taylor 3764 (NBG); Hopefield (–AB), Schlechter 5293 BOL); Montagu s.n. (SAM); Malmesbury (–BC), Werdernam and Oberdieck 436 (PRE);, Riebeek Kasteel, N slope of Botma's Pass (–BD), Pillans

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9530 (BOL); Melkbosstrand (–CB), Jordaan s.n. (NBG); Milnerton, Marine Drive, adjacent to the road and Diep River (–CC), Tywalana and Van Wyk 28 (NBG); Robben Island, slope directly above high-tide mark, midway along W edge of the island (–CD), Lloyd 492b (NBG); Walgate 619 (NBG); Van Jaarsveld and De Lange 9539 (NBG) Kirstenbosch, near old entrance (–CD), Henderson 1395 (NBG); Cape Peninsula, Newlands, waste ground (–CD), Compton 10029 (NBG); Cape Peninsula, sea shore E of Mouille Point (–CD), Pillans s.n. (BOL); Paarden Eiland, Klein Zoar Vlei (–CD), Linder 2172 (BOL); Cape Town, Mowbray, Durban Road station (–CD), Bolus 3853 (BOL); Cape Town, Sea Point(–CD), Tyson s.n. (NBG); Wolley Dod 1884 (BOL); Cape Town, Lion's Head, (–CD), Breori s.n. (PRE); railway enclosure, near Sea Point (–CD), Stokoe s.n. (PRE); near Sea Point (–CD), Stokoe s.n. (PRE); Newton s.n. (BOL); Adamson s.n. (BOL); Walgait s.n. (BOL); MacDowan s.n. (SAM); frequently along shore between Cape Town and Sea Point (–CD), Gulthrie s.n. (BOL); Paardeberg, between Wellington and Malmesbury, Fynbos estate, 750 m S of Farm house (–DB), Nicolson and Roets 649 (NBG); Bellville, University of Western Cape, Cape Flats Nature Reserve (–DC), Low 790 (NBG); Milnerton (–DC), Andreae 383 (PRE); Maitland Railway (–DC), Guthrie 739 (BOL). 3322 (Oudtshoorn): George, Blanco (–CD), Meyer s.n. (NBG). 3418 (Simonstown): Wildevoëlvlei S of mouth (–AB), O′ Callaghan 672 (NBG); Bergvliet Farm (–AB), Purcell s.n. (SAM); Strandfontein (–BA); Rycroft 2369 (NBG); Swartklip, ½ mile (1 km) from sea, hot N slope of dunes (–BA), Taylor 8044 (NBG); Strand, near sea in sandy soil (–BB), Parker 3883 (NBG, BOL); Gordon's Bay, roadside (–BB), Duttie 316 (NBG); Macassar Sewerage Works area, at Eerste River Mouth, Zandoliet Farm (–BB), Boucher 3469 (NBG). 3419 (Caledon): De Kelders, near Klipgat Pump Station, next to the road to the sea (–AB), Fellingham 1519 (NBG); between Stanford and The Kelders (–CB), Pillans 8130 (BOL); Gansbaai, Langbosch Farm (–CB), Walters 1052 (NBG); Danger Point (–CB), Compton 10192 (NBG); N.S.Pillans 8130 (BOL); Gansbaai (–CB), Walters 56 (NBG). Precise locality unknown: Calvinia, Schmidt 526 (PRE).

Fig. 9. Distribution of Oncosiphon africanus.

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7. Oncosiphon africanus (P.J.Bergius). Källersjö in Bot. J. Linn. Soc. 96 (4): 312 (1988); Manning and Goldblatt in Bothalia 35: 55 (2005); Goldblatt and Manning, Cape Plants: 345 (2000); Manning and Goldblatt, Plants of the Greater Cape Floristic Region: 401 (2012). Matricaria africana P.J.Bergius. in Descr. Pl. Cap: 296 (1767); Levyns in Adamson and Salter, Fl. Cap. Penins.: 804 (1950). Type: South Africa. Western Cape, Cape Town (3318): Paarden Eiland, Klein Zoar vlei near the Brooklyn Chest Hospital (–CD), in damp sand near the water, 2 Mar 1980, Linder 2208 [BOL, neo.!, designated by Källersjö, 1988]. Matricaria capensis L. in Mant. Pl.: 115 (1767); Thunb., Prodr. Pl. Cap. 2: 161 (1800); Thunb., Fl. Cap.: 693 (1823); DC., Prodr. Sys. Nat. 17: 50 (1838); Harv. in Harv. and Sond., Fl. Cap. 3: 164 (1865). Type: South Africa, Anon s.n. in Herb. Linn. No. 1013.9 (LINN–image!, lecto., designated by Källersjö, 1988). Chrysanthemum hirtum Thunb. in Prodr. Pl. Cap. 2: 161 (1800); Thunb., Fl. Cap.: 693 (1823). Matricaria hirta (Thunb.) DC. in Prodr. Sys. Nat. 17: 51 (1838); Harv. in Harv. and Sond., Fl. Cap. 3: 165 (1865). Type: South Africa, Thunberg s.n. in Herb. Thunb. 20,133 (UPS-THUNB holo.–microfiche!). Chrysanthemum glabratum Thunb. in Prodr. Pl. Cap. 2: 161 (1800). Matricaria glabrata (Thunb.) DC., Prodr. Sys. Nat. 6: 51 (1838); Harv. and Sond., Fl. Cap. 3: 165 (1865); Goldblatt and Manning, Cape Plants: 345 (2000). Oncosiphon glabratum (Thunb.) Källersjö in Bot. J. Linn. Soc. 96 (4): 312 (1988). Type: South Africa, Thunberg s.n. in Herb. Thunb. No. 20132 (UPS, holo.–microfiche!). Sprawling or sometimes erect, annual herb, to 0.3 m tall; stems glabrous to sparsely pilose. Leaves 1- or 2-pinnatisect, lanceolate to oblanceolate, 10–30 (50) × 5–20 mm, sparsely pilose; lobes oblong to lanceolate, 3–9 mm × ± 1 mm, mucronulate. Capitula radiate, heterogamous, 15–26 mm diam., in simple, lax corymbs sometimes solitary; peduncles thickened toward the base of capitula, 10–50 (75) mm long, sparsely pilose near the apex. Involucre broadly hemispherical, 7–13 mm in diam., base shortly attenuate to slightly cordate in mature heads; involucral bracts 3-seriate, suberect to slightly deflexed at anthesis, never decurrent on the peduncle, apices above the receptacle apex, midvein inconspicuous, glabrous, margins and apices broadly scarious; outer series ovate, ± 2 mm long; median series ovate to broadly ovate, 3–4 mm long; inner series broadly ovate, 2–4 mm long. Receptacle conical-globose, obscured by involucral bracts. Ray florets female-fertile; tube 2–3 mm long, glandular trichomes sessile; limb 4–5 mm long, oblong, white. Disc florets bisexual, corolla 2–3 mm long, yellow; tube not conspicuously swollen, ca. 1.6 mm long; limb campanulate, ± 1.2 mm long; lobes ovate, ± 0.5 mm long. Pappus prominent, obliquely cup-shaped, adaxially longer, irregularly lobed, ± 0.7 mm long. Diagnostic characters Oncosiphon africanus is the only radiate species (Fig. 1D & 4A1) in the genus, furthermore the cypselas have a very prominent pappus (Fig. 4F), almost half the length of the corolla tube. All other species have discoid capitula with the pappus inconspicuous to small and less than a quarter the length of the corolla tube. The species can be confused with another white, radiate annual occurring in sandy areas along the West Coast of southern Africa, Foveolina tenella (DC.) Källersjö, but that species has a spongy pappus, fewer ray florets, and distinctive cypselas with 3-ribs and two lateral rows of cavities on the adaxial surface.

Distribution and ecology The species is restricted to saline, seasonally waterlogged soils along the West Coast of South Africa, from Milnerton to Leipoldtville (Fig. 9). It is locally common. Flowering is in spring (September to October). Additional specimens examined South Africa. Western Cape. 3218 (Clanwilliam): outside Leipoldtville, on road to Sandberg (–BC), Hugo 667 (NBG); Clanwilliam, Bergvlei (–BC), Compton 5086 (NGB); 5 km N of Velddrif, just S of salt works, Farm Velddrif 110 (–CA), Helme 6161 (NBG); Farm Modderfontein 28, just N of Rocher Pan (–CB), Helme 6228 (NBG); Farm St Helenafontein, 29.5 km E of Rocher Pan, 0.7 km E of Farmhouse (–CB), Helme 6232 (NBG); salt marsh opposite Berg River Mouth (-CC), O'Callaghan 1159 (NBG); Salt pans, Berg River Station (–CD), Compton 15129 (NGB, BOL); Berg River (–CD), Compton 9469 (NGB). 3318 (Cape Town): West Coast National Park, N of Yzerfontein, found in sea sand (–AA), Craven 26 (NBG); 2 miles [3 km] from Koperfontein, on road to Hopefield (–AB), Nordenstam 3283 (PRE); Yzerfontein (–AB), Bolus s.n. (BOL); halfway between Yzerfontein and turnoff to Langebaan, near Salt Pan (–AC), Kӓllersjö 29 (BOL); Mamre Road (–BD), Compton 9815 (NBG); Milnerton, dunes near Foreshore (–CD), Andreae 380 (PRE); Rugby near Milnerton, near salt marsh (–CD), Levyns s.n. (BOL); Levyns 7016 (BOL); Paarden Island (–CD), Pole-Evans 16870 (BOL); Rietvlei (–CD), Zeyher s.n. (SAM).

Acknowledgements The curators and staff of the cited herbaria are thanked for their hospitality and assistance during visits and for making specimens available on loan. The Northern Cape Department of Environment and Nature Conservation and CapeNature are thanked for providing the relevant permits to conduct field visits and collect specimens. This work is based upon research supported by the National Research Foundation (grant number 90646), University of Johannesburg and the South African National Biodiversity Institute. In particular we would like to thank one of the reviewers, Prof. J.C. Manning, whose careful review and valuable comments improved the manuscript greatly. References Atlas of Living Australia, d. http://bie.ala.org.au/species, Accessed date: 31 July 2017. Herman, P.P.J., Manning, J.C., 2013. Asteraceae. In: Snijman, D.A. (Ed.), Plants of the Greater Floristic Region 2: The Extra Cape Flora. Strelitzia 30: 307. South African National Biodiversity Institute, Pretoria. Holmgren, P.K., Holmgren, N.H., Barnett, L.C., 1990. Index Herbariorum, Part 1: The Herbaria of the World. New York Botanical Garden, New York. Källersjö, M., 1988. A generic reclassification of Pentzia Thunb. (Compositae– Anthemideae) from South Africa. Botanical Journal of the Linnean Society 96, 299–322. Magee, A.R., 2011. The identity of the type of Tanacetum burchellii DC (Asteraceae, Anthemideae), and the correct author citation for Pentzia punctata Harv. ex Hutch. South African Journal of Botany 77, 251–252. Magee, A.R., Nicholas, A.N., Tilney, P.M., Plunkett, G.M., 2015. Phylogenetic relationships and generic realignments in the early diverging subtribe Pentziinae (Asteraceae, Anthemideae). Botanical Journal of the Linnean Society 178, 633–647. Manning, J.C., Goldblatt, P., 2005. Two new species of Asteraceae from Northern and Western Cape, South Africa and a new synonym. Bothalia 35, 58. Manning, J.C., Goldblatt, P., 2012. Plants of the Greater Cape Floristic Region 1: The Core Cape Flora. Strelitzia 29. South African National Biodiversity Institute, Pretoria. Van Wyk, B.-E., 2008. A review of Khoi-San and Cape Dutch medical ethnobotany. Journal of Ethnopharmacology 119, 331–341. Watt, J.M., Breyer-Brandw, M.G., 1962. Medicinal and Poisonous Plants of Southern and Eastern Africa. 2nd ed. E. & S. Livingstone, Edinburgh, London.