Four new species of Ursinia (Asteraceae

Phytotaxa 177 (3): 137–145 www.mapress.com/phytotaxa/ Copyright © 2014 Magnolia Press

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ISSN 1179-3155 (print edition)

PHYTOTAXA

ISSN 1179-3163 (online edition)

http://dx.doi.org/10.11646/phytotaxa.177.3.1

Four new species of Ursinia (Asteraceae, Anthemideae) from South Africa, with an updated key to the genus in Namaqualand ANTHONY R. MAGEE1,2,*, JAMES S. BOATWRIGHT3 & LADISLAV MUCINA4,5

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, P.O. Box 524, Auckland Park 2006, Johannesburg, South Africa 3 Department of Biodiversity & Conservation Biology, Faculty of Natural Sciences, University of the Western Cape, Private Bag x17, Bellville, 7535, Cape Town, South Africa 4 School of Plant Biology, University of Western Australia, 35 Stirling Hwy, Crawley WA 6009, Perth, Australia 5 Department of Geography & Environmental Studies, Stellenbosch University, Private Bag X1, Matieland 7602, Stellenbosch, South Africa * Corresponding author; e-mail: [email protected]. 1 2

Abstract Recent field and herbarium studies of the southern African genus Ursinia (Anthemideae, Asteraceae) in Namaqualand, South Africa, have revealed greater morphological variability than currently accommodated and a high percentage of misidentified specimens. In an attempt to remedy this we herein describe four new species (Ursinia arida, U. glandulosa, U. kamiesbergensis and U. laciniata) and provide a key to the species in the region, together with illustrations of their involucral bracts and paleae. The species can be distinguished by a combination of their life history, vestiture, presence or absence of appendages on the paleae, and shape of the involucral bracts and their scarious apices. Key words: Compositae, Gariep Center of Endemism, Greater Cape Floristic Region, Kamiesberg Center of Endemism

Introduction Namaqualand is a winter rainfall region within the Succulent Karoo Biome (Mucina et al. 2006) of South Africa, extending from the Orange River (ca. 28° S) southwards to the mouth of the Olifants River. Namaqualand is best known for the spectacular mass spring (July–September) flowering of annuals, largely from the Asteraceae. Ursinia Gaertner (1791: 462) (Asteraceae) is a prominent component of these spring floral displays. The genus is almost entirely southern African (one species extending to Ethiopia) and currently comprises ca. 39 species (Prassler 1967). They are easily recognized by their often large showy ray florets, paleate involucre, scarious involucral bracts and fruit crowned by a pappus of large, white spreading scales. It is the latter character which gives rise to the rather apt common name, Parachute Daisy. Ursinia is a member of tribe Anthemideae which has its origins in southern Africa (Watson et al. 2000, Oberprieler 2005, Himmelreich et al. 2008) and is placed within the early diverging subtribe Ursineae (Oberprieler et al. 2007). Ongoing field-based studies of Ursinia as well as herbarium studies revising the identity of the Namaqualand representatives (especially during the period of preparing the Ursinia treatment for Greater Cape Plants II; Magee et al. 2013), have revealed a broad spectrum of unaccommodated variability and a high percentage of misidentified specimens. Four unnamed species were discovered, two of which were recognized in Magee et al. (2013) as ‘sp. A’ and ‘sp. B’. Herein we formally describe these four species within a broader comparative context of all currently recognised Namaqualand Ursinia species and attempt to remedy the identification problems with a key to the species in Namaqualand, together with illustrations of the involucral bracts and paleae.

Materials and Methods Herbarium material of Ursinia species from Namaqualand were studied at NBG and BOL, as well as extensive field collections over several years (2004–2013). Almost all of the taxa were observed in the field, providing crucial additional insight into the species studied. The recorded distributions of the two new species were ascertained and Accepted by Alexander Sennikov: 31 Jul. 2014; published: 29 Aug. 2014

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mapped. The specimens examined are cited under each species treatment and arranged by Country, Province and then District. Within each District the specimens are organized according to geographical position, from west to east and north to south (using the Quarter Degree Reference System; Edwards & Leistner 1971, Leistner & Morris 1976).

Results and Discussion Of the eleven species of Ursinia found in Namaqualand, four are from subgenus Sphenogyne (Aiton 1813: 142) Prassler (1967: 382), namely U. anthemoides (L.) Poiret (1808: 257), U. calenduliflora (Candolle 1836: 682) Brown (1887: 670), U. glandulosa and U. laciniata. They are easily distinguished by the truncate paleae, straight or slightly curved fruit and uniseriate pappus. However, U. anthemoides remains a difficult taxonomic complex in need of further study across its much broader distribution range. The remaining seven species are from subgenus Ursinia and can be easily identified by the ovate to round scale-like appendages on the paleae, as well as the geniculate fruit with a biseriate pappus composed of an outer series of broad scales and an inner series of setiform scales. The monophyly of these two subgenera was recently confirmed in a preliminary phylogenetic analysis of the genus using nrITS sequence data (Swelankomo 2008). With the exception of U. chrysanthemoides (Lessing 1832: 242) Harvey (1865: 152) (Fig. 1F), U. glandulosa (Fig. 1M) and U. laciniata, which are perennial subshrubs, all of the Namaqualand species are annual herbs. This is in contrast to the mostly perennial nature of the genus in the adjacent regions of the Greater Cape Floristic Region. As also found by Prassler (1967), the species are most easily distinguished by characters of the involucral bracts. The apices of the involucral bracts are prominently cuspidate (U. pygmaea Candolle (1836: 690), Fig 2F), acute to acuminate (U. arida, U. cakilefolia Candolle (1836: 690), U. nana Candolle (1836: 690)) or obtuse (U. kamiesbergensis, U. speciosa Candolle (1836: 690)). In the latter two species the scarious apices form broad apical appendages which are either transversely ovate, thin and transparent as in U. speciosa (Fig. 2I), or ovate, thick and opaque as in U. kamiesbergensis (Fig. 1H). Distinct plants with prominent scarious attenuate, recurved apices were found around Garies and Kamieskroon. However, upon further study these populations were found to form a continuum with more typical populations of U. cakilefolia from around Clanwilliam. The bracts may also bear diagnostic dark markings in many of the species. In U. pygmaea the entire cuspidate apex is purplish black (Fig. 2H), in U. arida (Fig. 1K) and U. cakilefolia (Fig. 1I) the margins are outlined purplish black and in U. nana (Fig. 1J) there is usually a black apical crescent-shaped mark. The paleae are also very important for distinguishing the species. As mentioned earlier, the presence or absence of rounded apical appendages is a prominent character to separate them into the two current subgenera. The shape and texture of the apical appendage is also very important. In U. cakilefolia the appendages are prominent and scale-like in texture with a lustrous, brownish-yellow to brownish-black colour. This gives the disc a prominent dark sheen where these appendages overlap over the developing florets (Fig. 1G, I). In contrast, the appendages of U. nana and U. arida are less conspicuous, membraneous in texture and somewhat undulate (Fig. 2M, N). Apical appendages are absent in U. anthemoides (Fig. 2A, B), U. calenduliflora (Fig. 2C, D), U. glandulosa (Fig. 2E) and U. laciniata (Fig. 2F). In these species, the apical margins of the paleae are typically repand to slightly erosulate, but in U. laciniata they are prominently and closely laciniate (Fig. 2F). Prassler’s (1967) separation of U. anthemoides and U. calenduliflora was based entirely on slight differences in the size of the membraneous tips of the involucral bracts. We found this character to be highly variable (Fig 2A–D) within these two species so that the separation was completely artificial. As a result different plants from the same population had often been identified as different species. Herbarium specimens of these two species appear superficially very similar as details of the ray florets are often lost. Ursinia anthemoides is a widespread and fairly variable species occurring throughout the winter rainfall region. It has whitish, yellow or apricot ray florets usually with a pinkish underside (Fig. 1C–E). On the other hand, Ursinia calenduliflora is restricted to Namaqualand occurring from the Richtersveld to Kamiesberg. The heads are quite spectacular with the ray florets orange on both surfaces (rarely yellow in the Richtersveld) and often with a purple or black spot at the base (Fig. 1A). The Namaqualand populations of Ursinia anthemoides are distinguished further by the pappus that exceeds the corolla limb in fruit and the paleae which extend far above the height of the florets soon after anthesis. An adventive population of U. calenduliflora growing along the National Road (N1) between De Doorns and Worcester was also identified (Mauve & Oliver 131, NBG; Magee & Boatwright 392, NBG—Fig. 1B), having likely been introduced from the cultivated spring flower plantings in and around Worcester. It should be noted, however, that further taxonomic examination of the more widely distributed Ursinia anthemoides, U. chrysanthemoides and U. nana, is required as these taxa, as currently recognised, are likely to represent species complexes. However, such studies are beyond the scope of this paper. 138 • Phytotaxa 177 (3) © 2014 Magnolia Press

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Figure 1. Ursinia in Namaqualand. (A) mass display of U. calenduliflora near Springbok. (B) adventive population of U. calenduliflora near De Doorns. (C–E) U. anthemoides, pinkish underside of rays. (F) U. chrysanthemoides, perennial habit. (G, H) U. kamiesbergensis, obtuse tipped and distinctly thick and opaque involucral bract appendages. (I) U. cakilefolia, black outlined involucral bracts. (J) U. nana, with involucral bracts marked apically with black crescents. (K, L) U. arida, with only outer involucral bracts outlined purplish black. (M–P) U. glandulosa, perennial habit, glaucous, slightly succulent leaves with stalked glands on the branches and leaves. Photo credits: A. le Roux (A & G), A. Magee (B–E, H–J), L. Mucina (K), J. Boatwright (F & L), P. Van Wyk (M–O).

Four new species of Ursinia (Asteraceae)

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Figure 2. Involucral bracts (outer to inner) and palaea (outer and inner) of Namaqualand Ursinia. (A) U. anthemoides (Gubb 83, NBG). (B) U. anthemoides (Compton 11449, NBG). (C) U calenduliflora (Goldblatt 2386, NBG). (D) U. calenduliflora (Mauve & Oliver 131, NBG). (E) U. glandulosa (Oliver et al. 297, PRE). (F) U. laciniata (Winter NGS283, NBG). (G) U. chrysanthemoides (Maguire 1000, NBG). (H) U. pygmaea (Mucina 030906/25, NBG). (I) U. speciosa (Mucina 040806/32, NBG). (J) U. kamiesbergensis (Mucina 140907/09, NBG). (K) U. cakilefolia (Mucina 020905/03, NBG). (L) U. cakilefolia (Magee & Boatwright 344, NBG). (M) U. nana (Mucina 300806/18, NBG). (N) U. arida (Schlieben 8984, NBG). Scale: 1 mm. Artist: J.S. Boatwright.

4. Key to the Namaqualand species of Ursinia 1. – 2. – 3. – 4. – 5. – 6. – 7.

Paleae truncate, without rounded scale-like appendages; fruit straight or slightly curved with a uniseriate pappus of five broad scales ................................................................................................................................................................................................. 2 Paleae with ovate to rotund scale-like appendages; fruit geniculate with a biseriate pappus composed of an outer series of broad scales and an inner series of setiform scales.......................................................................................................................................5 Woody perennials shrublets . ..............................................................................................................................................................3 Annual herbs ......................................................................................................................................................................................4 Stalked glands on leaves and stems; young leaves and involucral bracts white felty; paleae apices repand to slightly erosulate ...... ........................................................................................................................................................................................U. glandulosa Stalked glands absent; leaves and involucral bracts glabrous to sparsely pilose; paleae apices prominently laciniate........................ ........................................................................................................................................................................................... U. laciniata Pappus not exceeding corolla limb in fruit, paleae not exceeding the height of the florets after anthesis; rays orange on both surfaces, rarely yellow, often with purple or black spot at base............................................................................... U. calenduliflora Pappus exceeding corolla limb in fruit, paleae extending far above the height of the florets after anthesis; rays whitish, yellow or apricot usually with a pinkish underside ....................................................................................................................U. anthemoides Perennial subshrubs............................................................................................................................................ U. chrysanthemoides Annual herbs........................................................................................................................................................................................6 Involucral bracts cuspidate (including some of the innermost bracts) with purplish black apices; restricted to quartz patches.......... ........................................................................................................................................................................................... U. pygmaea Involucral bracts not cuspidate, margins and apices if marked then variously outlined purplish black.............................................7 Middle involucral bracts with acute to acuminate apices....................................................................................................................8

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– 8. – 9. – 10. –

Middle involucral bracts with obtuse apices.....................................................................................................................................10 Paleae appendages prominent, scale-like, firm, lustrous, brownish-yellow to brown-black, flat; margins clearly defined, somewhat thickened......................................................................................................................................................................... U. cakilefolia Paleae appendages inconspicuous, membraneous, transparent, somewhat undulate; margins not prominently thickened................9 Inner paleae oblong with small ovate appendages; involucral bracts very broadly ovate with black crescent marks apically............ ................................................................................................................................................................................................. U. nana Inner paleae lorate with elliptic appendages; outer involucral bracts outlined purplish black, inner bracts unmarked.......... U. arida Middle involucral bracts with transversely ovate, thin, transparent, yellowish-brown apical appendages....................... U. speciosa Middle involucre bracts with ovate, thick, opaque, cloudy apical appendages....................................................U. kamiesbergensis

Taxonomy 1. Ursinia arida Magee & Mucina, sp. nov. Type:—South Africa. Northern Cape Province, Vioolsdrif (2817): Richtersveld, Eksteenfontein, S of Rooiberg and N of Klipbokkop at road to Hellskloof Canyon (–CD), 31 August 2006, Mucina 310806/07 (holotype NBG!, isotype K!). Erect to sprawling annual herb, 0.80–0.25 m tall; single-stemmed or well branched at the base, branches spreading to decumbent. Leaves cauline, alternate, 12.0–40.0 × 5.0–15.0 mm, bipinnate, glandular-punctate, sparsely pilose; lobes lanceolate to linear, 1.0–8.0 × ± 0.5–1.5 mm, mucronate. Capitula radiate, heterogamous, ca. 20–35 mm in diam., solitary; peduncle 40–140 mm long at anthesis, sparsely pilose. Involucre hemispherical to sometimes campanulate, 4.0–5.0 × 10.0–15.0 mm, glabrous; involucre bracts 4- to 6-seriate, margins and apices somewhat scarious; outer bracts ovate, 1.0–1.5 mm long, acute, outlined purplish black; middle bracts ovate, 2.0–3.0 mm long, acute, some outlined purplish black; inner bracts ovate, 3.0–4.0 mm long, without dark markings, apical appendages ovate to broadly ovate, obtuse. Receptacle paleate. Paleae scarious, apically with elliptic appendages, heteromorphic; outer broadly oblong, appendages narrower than limb; inner lorate, appendages only slightly broader than limb; appendages shorter than or equal to florets, not enclosing developing florets. Ray florets neuter, 12 to 20; lamina 6–12 mm long, yellow. Disc florets bisexual, numerous, yellow, 5-lobed; lobes cucullate, with sessile glandular trichomes. Anthers slightly sagittate at base; apical appendages cordate. Style terete; branches linear, truncate, sweeping hairs apically-dorsally. Pappus biseriate, white, outer scales 5, transversely obovate, inner scales 5, setaceous, unequally developed, some rudimentary. Cypselas oblanceolate, geniculate, ± 3.0 × 1 mm, heteromorphic, scattered, most white, granulate, some black, rugose. Distribution and ecology:—Ursinia arida is confined to the Richtersveld and northern Bushmanland in the Northern Cape Province (Fig. 3) and should be considered an endemic of the Gariep Center of Endemism (Van Wyk & Smith 2001). In the Richtersveld it occurs on a variety of substrates in vegetation types such as Dg 6 Helskloof Canyon Desert and Dg 2 Richtersberg Mountain Desert (Jürgens et al. 2006). The species appears to be particularly common in quartz-rich patches both in the Richtersveld (SKr 11 Rooiberg Quartz Vygieveld) and in Bushmanland (SKr 18 Bushmanland Inselberg Shrubland, SKr 19 Aggeneys Gravel Vygieveld) within the Succulent Karoo Biome (Mucina et al. 2006). We expect this species to be found also in southern Namibia, especially in the Fish River Canyon and in the desert landscapes of the Gariep River valley (upstream from Vioolsdrif and Noordoewer). Diagnostic characters:—Ursinia arida shares the inconspicuous and transparent paleae appendages and acute involucral bracts with Ursinia nana but differs in that the inner paleae are lorate with elliptic appendages (Fig. 2M), rather than oblong with small ovate appendages. The species has also often been misidentified as U. cakilefolia from which it differs in that the paleae are shorter than the florets and have elliptic appendages. In U. cakilefolia the paleae are longer than the florets, and therefore conspicuous, and have prominent scale-like, ovate to transversely ovate appendages (Fig. 2K, L) that are brownish-yellow to brown-black and which enclose the developing florets giving the disc a reflective appearance (Fig. 1I). Additional specimens examined:—South Africa. Northern Cape Province: 2817 (Vioolsdrif): Vioolsdrif, Die Toon, E. of Richtersveld National Park (–AD), July 1995, Williamson & Williamson 5657 (NBG); Richtersveld, Hellskloof Canyon, lower part close to bushman’s paintings (–CA), 31 August 2009, Mucina 310806/24 (NBG); Richtersveld, E of Eksteenfontein and SW of Rooiberg Guest House (–CD), 29 August 2006, Mucina 290806/04 (NBG); Richtersveld, Eksteenfontein, S of Rooiberg and N of Klipbokkop at road to Hellskloof Canyon (–CD), 31 August 2006, Mucina 310806/07 (NBG). 2819 (Ariamsvlei): 28 miles west of Pofadder (–DD), 23 May 1961, Schlieben 8984 (NBG, NYBG). 2917 (Springbok): O’okiep (–DB), 28 September 1939, Thorus 39 (NBG). 2918 (Gamoep): 48 km E of Springbok, at the edge of Bushmanland (–AD), Goldblatt & Manning 11332 (NBG), Boatwright & Manning 641 (NBG); Naib se Berg between Springbok and Aggeneys (–AD) Mucina 080905/4 (NBG); near Ratelkraal (–CA), 7 September 1950, Barker 6621 (NBG); Areb, 30 miles NE of Springbok (–CB), 12 April 1926 (NBG), Barker 8363 (NBG); 45 km E of Springbok on Pofadder road (–CB), 14 May 2000, Manning 2247 (NBG); Bushmanland, on road Springbok to Aggeneys, Farm Kangas (–CB), 2 September 2006, Mucina 020906/02 (NBG). Four new species of Ursinia (Asteraceae)


Figure 3. Known geographical distribution of Ursinia arida (circles), Ursinia glandulosa (diamond), U. kamiesbergensis (triangles) and U. laciniata (stars; the white star indicates a specimen where the precise locality is unknown).

2. Ursinia glandulosa Magee & Boatwr., sp. nov. Type:—South Africa. Northern Cape, Vioolsdrif (2817): Richtersveld, Rosyntiesberg (–AC), 30 August 1977, Oliver, Tölken & Venter 297 (holotype PRE!). Woody perennial shrublet, 0.30–0.40 m tall; densely branched, branches stout, with stalked glands. Leaves cauline, alternate, 8.0–12.0 × 4.0–7.0 mm, trifid to bi-trifid, glandular-punctate, white felty when young, becoming glabrescent; stalked gland present; lobes narrowly ovate to lanceolate, 1.0–5.0 × ± 1.0 mm, mucronate; mucro acute. Capitula radiate, heterogamous, ca. 15–20 mm in diam., solitary; peduncle 50–80 mm long at anthesis, sparsely pilose. Involucre hemispherical, 5.0–7.0 × 8.0–10.0 mm, white felty; involucre bracts 5- to 6-seriate, brownish along inner margins; outer bracts narrowly ovate, 1.5–2.0 mm long, margins scarious; middle bracts narrowly oblong, 4.0–4.5 mm long, margins scarious; inner bracts narrowly oblong, 5.0–7.0 mm long, apical appendages broadly ovate, obtuse. Receptacle paleate. Paleae scarious, truncate, repand to slightly erosulate. Ray florets neuter, ca. 12 to 16; lamina 4–6 mm long, yellow. Disc florets bisexual, numerous, yellow, 5-lobed; lobes cucullate, with sessile glandular trichomes. Anthers slightly sagittate at base; apical appendages cordate. Style terete; branches linear, truncate, sweeping hairs apicallydorsally. Pappus uniseriate, scales 5. Cypselas unknown. Distribution and ecology:—Ursinia glandulosa is an endemic species of the Richtersveld (Fig. 3) where it is found in two major vegetation types, i.e. SKr 8 Rosyntjiesberg Succulent Shrubland (here at high elevations, 1060 m on Rosyntjiesberg Quartzite) and SKr1 Central Richtersveld Mountain Shrurbland (Mucina et al. 2006). Diagnostic characters:—Ursinia glandulosa, as the name implies, differs from all other currently recognised Ursinia species by the conspicuous stalked glands on the leaves and branches (Fig. 1P). This species has previously been confused with U. punctata Brown (1887: 670) with which it shares the woody perennial habit, weakly pinnatisect leaves, apical appendages only on the outer involucral bracts and the truncate paleae. It differs further from the latter in the short stout branches (Fig. 1M), the densely white felty young leaves and involucral bracts (Fig. 1N, O) and the presence of apical appendages (Fig. 2E) only on the innermost involucral bracts (branches lanky, young leaves sparsely pilose, involucral bracts glabrescent to sparsely pilose and both inner two-most involucral bracts apically appendaged in U. punctata). Additional specimens examined:—South Africa. Northern Cape Province: 2817 (Vioolsdrif): Armmanshoek (–AC), September 1995, Williamson & Williamson 5777 (NBG). 142 • Phytotaxa 177 (3) © 2014 Magnolia Press

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3. Ursinia kamiesbergensis Magee & Mucina, sp. nov. Type:—South Africa. Northern Cape, Kamiesberg (3018): Before turnoff, near Bovlei, to Pedroskloof on road from Kamieskroon to Leliesfontein (–AA), 2 September 2011, Magee & Boatwright 337 (holotype NBG!, isotypes BOL!, PRE!, K!). Sprawling to rarely erect annual herb, 0.15–0.30 m tall; single-stemmed or well-branched at the base, branches decumbent. Leaves cauline, alternate, 12.0–50.0 × 7.0–15.0 mm, bipinnate, glandular-punctate, sparsely pilose; lobes lanceolate to linear, 1.0–8.0 × ± 0.5– 1.0 mm, mucronate. Capitula radiate, heterogamous, ca. 20–30 mm in diam., solitary; peduncle 130–190 mm long at anthesis, sparsely pilose. Involucre hemispherical, 4.0–6.0 × 12.0–18.0 mm, glabrous; involucre bracts 4- to 6-seriate, apices with thick, opaque, cloudy scarious appendages; outer bracts ovate, 1.5–2.0 mm long, margins scarious, apical appendages lanceolate to ovate, acute, purplish black along inner margins; middle bracts ovate, 3.5–4.5 mm long, margins partly scarious, apical appendages ovate, obtuse, purplish black along inner margins; inner bracts ovate, 6.0–7.5 mm long, apical appendages ovate to broadly ovate, obtuse or emarginate. Receptacle paleate. Paleae scarious, apically with ovate to transversely ovate appendages, heteromorphic; outer broadly oblong, appendages narrower than limb; inner narrowly oblong, appendages broader than limb; appendages longer than florets, enclosing developing florets to form reflective disc. Ray florets neuter, 20 to 25; lamina 8–15 mm long, yellow. Disc florets bisexual, numerous, yellow, 5-lobed; lobes cucullate, with sessile glandular trichomes. Anthers slightly sagittate at base; apical appendages cordate. Style terete; branches linear, truncate, sweeping hairs apically-dorsally. Pappus biseriate, white, outer scales 5, transversely obovate, inner scales 5, setaceous, unequally developed. Cypselas oblanceolate, geniculate, ± 3.5 × 1 mm, heteromorphic, scattered, most white, granulate, some black, rugose. Distribution and ecology:—Ursinia kamiesbergensis is restricted to coarse sandy soils derived from the Mokolian Kamieskroon Gneiss of the top plateaus of the Kamiesberg Mountains, Northern Cape Province (Fig. 3), within the Kamiesberg Center of Endemism (Van Wyk & Smith 2001). We may hypothesise that this species originally occurred in herb-rich and lightly, locally disturbed (grazing wildlife and burrowing animals) openings in the renosterveld scrub. Today it is very common especially in disturbed flats formerly used as arable fields formed by clearing of the renosterveld scrub on deep coarse-grained sandy and loamy-sandy soils. The vegetation type is the FRg 1 Namaqualand Granite Renosterveld (Fynbos Biome; Rebelo et al. 2006). Diagnostic characters:—Ursinia kamiesbergensis has previously been confused with U. cakilefolia but differs from that species in that the apical appendages of the involucral bracts are obtuse tipped and distinctly thick and opaque (Fig. 1H, 2J). Ursinia cakilefolia usually lacks prominent apical appendages on the involucral bracts (Fig. 2K) but when prominent appendages are present, as found in populations around Kamieskroon (Fig. 2L), they are acute to acuminate, thin, transparent and yellowish-brown. Prominent, obtuse apical appendages are also found in Ursinia speciosa (Fig. 2I) but are distinctly transversely ovate and not as much ovate as in U. kamiesbergensis. According to the taxonomic concepts of Prassler (1967), U. kamiesbergensis falls within the variability of a more broadly conceived Ursinia cakilefolia. Yet it was Prassler’s remark on page 457 that brought the Kamiesberg taxon to our attention: ‘Besonders erwähnt sei die Aufsammlung SALTER 1446 A, die durch breitere Hüllschuppen abweicht und bei der es sich vielleicht um eine distinkte Sippe handelt. Um dies zu entscheiden, wären jedoch weitere Aufsammlungen, wohl auch eingehende Feldstudien erforderlich.’ (Translation: “Noteworthy is the collection of Salter 1446 A which differs in broader bracts and perhaps should be recognized as a distinct species. To establish that, more collections as well as focused field studies are needed.”). The description of U. kamiesbergensis herein confirms Prassler’s assumption. Additional specimens examined:—South Africa. Northern Cape Province: 3017 (Hondeklipbaai): Kamieskroon Mountains (–BB), 29 August 1937, Compton 6801 (NBG); 3018 (Kamiesberg): Kamieskroon, Kamiesberg Pass, top plateau (–AA), 14 September 2007, Mucina 140907/09 (NBG, 2 sheets); Farm Hoorngat 420 (–AC), 24 August 1983, Le Roux 3165 (NBG); between Leliefontein and Kamieskroon (–AC), October 1933, Mathews s.n. (BOL); Studer’s Pass, south of Farm Koorlandskloof (–AC), 6 September 2005, Mucina 060905/01 (NBG, 4 sheets), 6 September 2005, Mucina 060905/02 (NBG, 4 sheets); Farm Welkom (–AC), 19 September 1971, Oliver 3531 (NBG); Farm Wolfhok, 30°23’04.7’’S 18°06’30.8’’E (–AC), 11 September 2007, Snijman 2127 (NBG). Precise locality unknown: Valleys in the Kamiesberg, October 1931, Mathews 20049 (BOL); Kamiesberg, December 1932, Mathews s.n. (BOL); Ex hort. Pillans’ Garden, August 1912, Pearson s.n. sub BOL27326 (BOL).



4. Ursinia laciniata Magee & Mucina, sp. nov. Type:—South Africa. Northern Cape, Springbok (2917): Steinkopf, Klipfonteinberg (–BA), 5 October 2014, Winter NGS283 (holotype NBG!, isotypes K!, PRE!). Woody perennial shrublet, 0.30–0.40 m tall; densely branched, branches lanky, stalked glands absent. Leaves cauline, alternate, 6.0–10.0 × 3.0–7.0 mm, 3(5)-lobed, glandular-punctate, glabrescent, polished; stalked gland absent; lobes lanceolate to narrowly lanceolate, 1.0–4.0 × ± 0.5– 1.0 mm, mucronate; mucro acute. Capitula radiate, heterogamous, ca. 15–20 mm in diam., solitary; peduncle 40–60 mm long at anthesis, glabrescent. Involucre hemispherical, 4.0–6.0 × 8.0–11.0 mm, glabrescent; involucre bracts 4- to 5-seriate, brownish along inner margins; outer bracts narrowly ovate, 1.5–2.0 mm long, margins scarious; middle bracts narrowly oblong, 4.0–4.5 mm long, margins scarious; inner bracts narrowly oblong, 5.0–6.0 mm long, apical appendages ovate, obtuse. Receptacle paleate. Paleae scarious, truncate, closely laciniate. Ray florets neuter, 10–14; lamina 4–6 mm long, yellow. Disc florets bisexual, numerous, yellow, 5-lobed; lobes cucullate, with sessile glandular trichomes. Anthers slightly sagittate at base; apical appendages cordate. Style terete; branches linear, truncate, sweeping hairs apically-dorsally. Pappus uniseriate, scales 5. Cypselas unknown. Distribution and ecology:—This species is known so far from only a few localities in between the Richtersveld and the Kamiesberg (Fig. 3). Near Steinkopf it grows among large granite (Precambrian gneiss) supporting dense succulent karoo scrub, belonging to the SKr15 Anenous Plateau Shrubland (Mucina et al. 2006). Diagnostic characters:—Ursinia laciniata may be confused with the similar but allopatric U. punctata which is also a woody perennial with dissected leaves and truncate paleae. However, U. laciniata is easily distinguished from the latter by the polished, mostly trifid leaves with acute mucros, the glabrescent leaves and involucral bracts, the narrowly oblong middle involucral bracts with only the innermost bracts apically appendaged (Fig. 2F) and the paleae with prominently laciniate apical margins (Fig. 2F). In U. pungens the mature leaves are mostly 5–10 lobed with prominently long acuminate mucros, the leaves and involucral bracts are sparsely pilose, the middle involucral bracts are ovate to narrowly ovate with at least the inner two-most involucral bracts apically appendaged and the apical margins of the paleae are repand to slightly erose. Ursinia laciniata may be confused with the closely allied U. glandulosa with which it shares the acute leaf mucros and narrowly ovate middle involucral bracts but can be distinguished most prominently by the absence of stalked glands, the glabrescent leaves and involucre, and the laciniate paleae. Additional specimens examined:—South Africa. Northern Cape Province: 2817 (Vioolsdrif): Richtersveld, Kalkfontein (–CC), 23 August 1925, Marloth 12655 (PRE). Precise locality unknown: Kamiesbergen, 10 September 1925, Marloth 12685 (PRE).

Acknowledgements The curators and staff of the cited herbaria are thanked for their kind hospitality and assistance during visits. The Northern Cape Province Department of Environment and Nature Conservation are thanked for providing the relevant permits. Annelise le Roux (CapeNature) is gratefully acknowledged for providing two of the images of U. kamiesbergensis and U. calenduliflora and Pieter van Wyk for providing the images of U. glandulosa. This work is based upon research supported by the National Research Foundation and Department of Science and Technology under the SABI Programme and the University of Johannesburg (AM). LM acknowledges the field support of N. Swelankomo and logistic support of the Iluka Chair (The University of Western Australia). LM also appreciates the support of the National Geographic Society (Washington D.C.) who financed the field work explorations in Namaqualand and contributed to the serendipitous discovery of a plant specimen here serving as the type of U. laciniata.

References Aiton, W.T. (1813) Hortus Kewensis. Longman, Hurst, Rees, Orme, and Brown, Paternoster Row, London, 568 pp. Brown, R. (1887) Plant New or Noteworthy. The Gardeners’ chronicle: a weekly illustrated journal of horticulture and allied subjects 3(1): 670. Candolle, A.P. de (1836) Prodromus systematis naturalis regni vegetabilis, vol. 5. Treuttel et Würtz, Paris, 706 pp. Edwards, D. & Leistner, O.A. (1971) A degree reference system for citing biological records in southern Africa. Mitteilungen des Botanische Staatssammlung München 10: 501–509. Gaertner, J. (1791) De fructibus et seminibus plantarum, vol. 2. Sumtibus Auctoris, Typis Academiae Carolinae, Stuttgard, 678 pp.

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Harvey, W.H. (1865) Compositae. In: Harvey, W.H. & Sonder, O.W. (Eds.) Flora Capensis, vol. 3. Hodges, Smith & Co., Dublin, pp. 44–530. Himmelreich, S., Källersjö, M., Eldenäs, P. & Oberprieler, C. (2008) Phylogeny of southern hemisphere Compositae-Anthemideae based on nrDNA ITS and cpDNA ndhF sequence information. Plant Systematics and Evolution 272: 131–153. http://dx.doi.org/10.1007/s00606-007-0634-y Jürgens, N., Desmet, P.G., Rutherford, M.C., Mucina, L. & Ward, R.A. (2006) Desert Biome. In: Mucina, L. & Rutherford, M.C. (Eds.) The vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria, pp. 300–323. Leistner, O.A. & Morris, J.W. (1976) South African place names. Annals of the Cape Provincial Museums 12: 1–565. Lessing, C.F. (1832) Synopsis generum Compositarum. Black, Young & Young, London, 473 pp. Magee, A.R., Mucina, L. & Snijman, D.S. (2013) Ursinia. In: Snijman, D.S. (Ed.) Greater Cape Plants II: Namaqualand-southern Namib and Western Karoo. Strelitzia 29. South African National Biodiversity Institute, Pretoria, pp. 331–332. Mucina, L., Jürgens, N., Le Roux, A., Rutherford, M.C., Schmiedel, U., Esler, K., Powrie, L.W., Desmet, P.G., Milton, S.J., Boucher, C., Ellis, F., Lambrechts, J.J.N., Ward, R.A., Manning, J.C. & Midgley, G.F. (2006) Succulent Karoo Biome. In: Mucina, L. & Rutherford, M.C. (Eds.) The vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria, pp. 220–299. Oberprieler, C. (2005) Temporal and spatial diversification of Circum-Mediterranean Compositae–Anthemideae. Taxon 54: 951–966. http://dx.doi.org/10.2307/25065480 Oberprieler, C., Himmelreich, S. & Vogt, R. (2007) A new subtribal classification of the tribe Anthemideae (Compositae). Willdenowia 37: 89–114. http://dx.doi.org/10.3372/wi.37.37104 Poiret, J.L.M. (1808) Encyclopédie Méthodique: Botanique VIII. Panckoucke, Plomteux, Paris, 881 pp. Prassler, M. (1967) Revision der Gattung Ursinia. Mitteilungen der Botanischen Staatssammlung München 6: 363–478. Rebelo, A.G., Boucher, C., Helme, N., Mucina, L., Rutherford, M.C., Smit, W.J., Powrie, L.W., Ellis, F., Lambrechts, J.J., Scott, L., Radloff, F.G.T., Johnson, S.D., Richardson, D.M., Ward, R.A., Procheş, S.M., Oliver, E.G.H., Manning, J.C., Jürgens, N., McDonald, D.J., Janssen, J.A.M., Walton, B.A., Le Roux, A., Skowno, A.L., Todd, S.W. & Hoare, D.B. (2006) Fynbos Biome. In: Mucina, L. & Rutherford, M.C. (Eds.) The vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria, pp. 52–219. Swelankomo, N. (2008) Molecular Phylogeny, Radiation Patterns and Evolution of Life-History Traits in Ursinia (Anthemideae, Asteraceae). MSc. dissertation. Stellenbosch University, Stellenbosch, South Africa. Van Wyk, A.E. & Smith, G.F. (2001) Regions of Floristic Endemism in Southern Africa. Umdaus Press, Hatfield, 199 pp. Watson, L., Evans, T.M. & Boluarte, T. (2000) Molecular phylogeny and biogeography of tribe Anthemideae (Asteraceae), based on chloroplast gene ndhF. Molecular Phylogenetics and Evolution 15: 59–69. http://dx.doi.org/10.1006/mpev.1999.0714

