A synopsis of the genus Pteronia (Compositae: Astereae) in Namibia including the resurrection of Pteronia quadrifaria

Herta Kolberg

Background: The distribution of Ficus sur includes most of tropical Africa, but whilst this species was suspected to occur in Namibia, this has not been verified. Gymnanthemum coloratum is a tropical African savannah shrub or tree that has been recorded for Botswana, Swaziland and South Africa, but which has not previously been recorded for Namibia.Objectives: To formally document the first records of two plant species from Namibia and provide habitat details of the localities from which these species were recorded.Method: The data presented have resulted from botanical expeditions to the poorly known Baynes Mountains in the Koakoveld region of Namibia. Specimens of the two species in the National Herbarium, Pretoria were examined to verify the identity in the case of G. coloratum, and to document additional records in Namibia for F. sur.Results: Ficus sur was recorded from two localities, and a third locality based on a specimen in the National Herbarium, Pretoria, was verified. Gy...

Commiphora omundomba, described here as a new species, is known only from the Kaokoveld Centre of Endemism, a biogeographical region in southwestern Angola and Northwestern Namibia. It has hitherto been confused with C. dinteri from west-central Namibia, but both morphological and molecular evidence support a distinction between the two species. Phylogenetic analysis of DNA sequence data indicates that C. omundomba shares most-recent common ancestry with C. buruxa, a species limited to the Gariep Centre of Endemism in southwestern Namibia. Diagnostic morphological characters for C. omundomba include the white, viscous exudate, smooth bark and glabrous, trifoliolate leaves. Photographs of the plant and a distribution map are provided. The new species is mainly confined to near the Atlantic coast of the northern Namib Desert, and is widespread and locally common between Santa Maria in Angola and Puros in Namibia.

Commipora kaokoensis W.Swanepoel. here described as a new species, is known only from the Kaokoveld Centre of Endemism, an arid region in northwestern Namibia. Illustrations of the plant and a distribution map are provided. Diagnostic characters include the petiolate or subsessile. all-simple and relatively large leaves with the lamina obovate or elliptic. New information is provided on the leaf morphology and geographical distribution of C. namaensis Schinz and C. dinteri Engl., species with which the new species shares some similarities. When without leaves or fruit, the three species can easily be confused. A comprehensive table with diagnostic morphological features to distinguish between the three species is presented.

KEW BULLETIN (2014) 69: 9488 DOI 10.1007/S12225-014-9488-Y ISSN: 0075-5974 (print) ISSN: 1874-933X (electronic) A synopsis of the genus Pteronia (Compositae: Astereae) in Namibia including the resurrection of Pteronia quadrifaria Herta Kolberg1 & Michiel van Slageren2 Summary. An account of species of Pteronia occurring in Namibia is presented, together with an updated key. New material of P. quadrifaria Dinter conﬁrmed that this is a valid species and not a synonym of P. lucilioides DC., and is resurrected. Species descriptions, global and Namibian distributions, conservation status, habitat and phenological information are presented for the 24 species recognised from Namibia. During this study it became clear that many species names have not been properly typiﬁed. As a result arguments are presented for the choice of 10 lectotypes, eight of which concern Pteronia with ﬁve of these for names of species accepted here. Four species of Pteronia that have been described from or are at least partly occurring in Namibia but that are no longer considered part of this genus are brieﬂy discussed. Key Words. Asteraceae, key, lectotypes, southern Africa, taxonomy. Introduction During ﬁeldwork for the Millennium Seed Bank Project in the south-west of Namibia in 2009, a species of Pteronia L. (Compositae: Astereae) unknown to the ﬁrst author was found on a white quartz hill near the small town of Aus. After initial investigation and consultation with various specialists, it seemed that this plant represented a new species of the genus. Since the only published account of the Compositae for the ﬂora of Namibia (Merxmüller 1967) is outdated and needs to be revised, it was decided to combine description of this presumed new species with a synopsis and updated key of Pteronia in the country. During this process it was discovered that the new material was in fact of the little known species P. quadrifaria Dinter, which had been considered a synonym of P. lucilioides DC. by subsequent authors (Merxmüller 1955, 1967). Pteronia is a mainly southern African genus comprising about 70 species. Within the tribe Astereae it is presently placed in the subtribe Solidagininae O. Hoffm. (which consists mostly of North American genera) and in turn into the Engleria group (Bremer 1994) of that subtribe. Pteronia is considered to be isolated among the African Compositae (Bremer 1994) and most closely related to Engleria O. Hoffm., which, however, has radiate capitula compared to the discoid capitula of Pteronia. Morphologically there exists similarity with Chrysocoma L. and other African genera of the Amellus group in the subtribe Asterinae (Bremer 1994). Charac- ters that are rare in the Astereae are found in Pteronia, like a spiny habit, opposite and succulent leaves and beaked achenes (Bremer 1994). Bremer (1994) concluded that the placement of Pteronia in the subtribe Solidagininae must be considered very uncertain, a point of view discussed below. Pending a revision of the entire genus, its subtribal afﬁliation cannot be completely ascertained (Bremer 1994). The most recent comprehensive treatment (Hutchinson & Phillips 1917) is, however, over 90 years old. In Namibia, the genus was last revised by Merxmüller in 1967. Since then, one additional species (P. anisata B. Nord.) has been recorded for Namibia (Merxmüller & Roessler 1984). Hutchinson & Phillips’s (1917) revision included the 63 species accepted by de Candolle (1836) that were grouped in three sections (sect. Scepinia (Neck. ex Cass.) DC., sect. Pachyderis (Cass.) DC., sect. Pterophorus (Vaill. ex Adans.) DC.). However, they regarded the difference between Scepinia and Pachyderis as indistinct and the distinguishing character of Pterophorus (a monospeciﬁc section according to Harvey 1865) as “triﬂing”. The 61 species that they accepted were then divided into four new sections based mainly on leaf indumentum: I. Incanae — leaves with woolly, whitish indumentum II. Papillatae — entire leaf surface papillous or scabrid III. Ciliatae — leaves ciliate mostly on margins or keels but otherwise glabrous IV. Glabratae — leaves entirely glabrous and mostly ﬂeshy Accepted for publication 22 January 2014. 1 Millennium Seed Bank Partnership, National Botanical Research Institute, Windhoek, Namibia. email: hertak@nbri.org.na. 2 Millennium Seed Bank Partnership, Seed Conservation Department, Royal Botanic Gardens, Kew, Wakehurst Place, Ardingly, nr Haywards Heath, West Sussex, RH17 6TN, UK. e-mail: m.vanslageren@kew.org © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 2 of 44 Merxmüller (1952) noted that this division is not at all consistent and that, for instance, Pteronia divaricata Less. and P. lucilioides DC., which both were classiﬁed by Hutchinson & Phillips as belonging to sect. Papillatae, differ very much in the degree to which the leaves are papillate. On the other hand, P. unguiculata S. Moore and P. mucronata DC., which belong to sect. Glabratae and sect. Ciliatae respectively, are related more closely than would be expected from them being in different sections. (It should be noted that as section Ciliatae included the type of the genus, P. camphorata (L.) L., this name was not validly published and should be section Pteronia.) In view of this we have not further considered these sections. Materials and Methods Herbarium specimens at K, NBG, SAM and WIND and high resolution images of herbarium specimens at the herbaria of B, BM, BOL, G, G-DC, GH, GRA, HAL, HBG, K, KW, LD, M, NBG, NY, P, PRE, S, SAM, TUB, W (including W-Rchb.), Z were studied. Except for type specimens, these were Namibian specimens only because this study was not intended to be a full revision of the genus but merely a summary of what is known about Pteronia in Namibia. Species descriptions and the key were compiled from a combination of published information and characters observed on Namibian herbarium specimens. Global distributions were obtained from literature and online databases, mainly the African Plant Database (http://www.ville-ge.ch/musinfo/bd/cjb/ africa/index.php?langue=an). The habitat of species was derived from a summary of Namibian specimen label information only. Altitude ranges therefore are only those stated in the collection information and relate to Namibia only. Where there is only one altitude, either only one specimen contained altitude information or several specimens had exactly the same altitude. Similarly, phenology of species is that derived from Namibian specimens only. This obviously has limitations as the number of specimens with the required information KEW BULLETIN (2014) 69: 9488 may be very few. The information given should not be seen as applicable to the species in their entire distribution range. Conservation status was assessed using Guidelines for Using the IUCN Red List Categories and Criteria. Version 10 (IUCN 2013). Since mainly Namibian specimens were examined and no ﬁeldwork was done in other countries besides Namibia, the assessments were made for Namibia only (for the endemic species this is obviously a global assessment). Where available, South African assessments were obtained from literature and online databases (SANBI 2011). Pteronia L. (Linnaeus 1763b: 1176; 1764: 414); de Candolle (1836: 356); Harvey in Harvey & Sonder (1865: 95); Hutchinson & Phillips (1917: 278); Merxmüller (1967: 150); Bremer (1994: 408). Type: Pteronia camphorata (L.) L. Shrubs with opposite, alternate or fascicled leaves, these glabrous, papillate, glandular or with conspicuous bristles and entire margins. Capitula solitary or corymbose, homogamous, discoid. Receptacle convex, ﬂat or concave, often honeycombed, ﬁmbriate, setate or lacerate. Phyllaries multiseriate, gradate, imbricate; apex often mucronate; margins often membranous. Florets hermaphrodite; corollas tubular, actinomorphic, 5-lobed, yellow or white. Stamens 5. Anthers more or less obtuse, rarely acute, at base. Style with two deltoid-tipped, ﬂattened branches bearing short to long stigmatic papillae near apex. Achenes obconical, obovoid or turbinate, sometimes ﬂattened or contracted into a neck apically, glabrous, glandular or variously hairy; carpopodium a slightly asymmetrical ring with a narrow interruption. Pappus of unequal, scabrid to barbellate setae, sometimes basally connate or scale-like and broadened. Diagnostic characters of Pteronia are discoid capitula with multiseriate, gradate and densely imbricate phyllaries, bi- or multiseriate pappus consisting of bristles only (some bristles may be scale-like broadened at the base), achenes that are sometimes narrowed into an apical neck and mostly opposite leaves (Bremer 1994; Herman et al. 2000). Updated key to Pteronia in Namibia 1. Leaves with dense, white to grey, felty indumentum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Leaves glabrous, papillose, glandular, bristly, ciliate or pectinate but never felty . . . . . . . . . . . . . . . . . . . . . 4 2. Phyllaries acute, midrib broader towards apex but never mucronate; pappus yellowish-brown. . . 9. P. glauca Phyllaries mucronate; pappus maroon-purple or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Pappus maroon-purple; phyllaries reddish-purple. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7. P. eenii Pappus yellow; phyllaries yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. acuta 4. Leaves, at least those at branch tips surrounding capitula, ciliate to pectinate or bristly, papillose or glandular on surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Leaves glabrous to minutely puberulous or minutely papillate, may appear warty when dried. . . . . . . . . . .13 5. Leaf margins ciliate-pectinate or bristly, leaf surface glabrous, bristly or minutely papillose, leaves alternate, alternate to almost opposite, opposite or decussate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 9488, Page 3 of 44 Leaf margins not ciliate-pectinate or bristly, leaf surface papillose, with short bristles or glandular, leaves opposite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10 Only leaves at branch tips surrounding capitula, with ciliate margins, others glabrous to glabrescent . .17. P. pomonae All leaf margins ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Capitula 20 × 7 mm at most, narrowly cylindrical, apex of phyllaries truncate. . . . . . . . . . . . . . . . . . . . . . . 8 Capitula to 35 × 25 mm, ovoid-obconical, apex of phyllaries rounded or acute. . . . . . . . . . . . . . . . . . . . . . 9 Pappus yellowish; apex of phyllaries truncate, mucronulate, not reﬂexed; leaves opposite, connate at base, clustered along branches; achenes densely villous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. P. mucronata Pappus maroon-purple; apex of phyllaries reﬂexed, margins undulate; leaves sessile, imbricate-decussate, mostly only towards branch tips, lower leaves soon dehiscent; achenes glandular . . . . . . . . 18. P. quadrifaria Leaves alternate, margins white bristly, to 20 × 5 mm; phyllaries coriaceous, apex rounded; achenes glandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. P. onobromoides Leaves opposite, margins and keel ciliate, 8 – 10 × 4 mm; phyllaries membranous, apex acute; achenes glabrous or with a few hairs on ribs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. P. viscosa Phyllaries yellowish, without membranous margin; capitula acute in bud . . . . . . . . . . . . . . .12. P. lucilioides Phyllaries with conspicuous membranous margin; capitula not markedly acute in bud . . . . . . . . . . . . . . . .11 Capitula ovoid-campanulate, 15 – 20 × 6 – 10 mm; phyllaries ovate to orbicular; leaves sessile, oblong to narrowly ovate, apex rounded, 5 – 15 × 4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. P. inflexa Capitula cylindrical, 15 × 3 – 9 mm; phyllaries lanceolate-oblong, linear-oblong, the outer sometimes ovate; leaves narrowing into a short petiole, obovate, spathulate or orbicular, 10 – 25 × 8 – 15 mm . . . . . . . . . . .12 Capitula solitary on branch tips, 10 – 12-ﬂowered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. P. polygalifolia Capitula in terminal corymbs, each 5-ﬂowered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. P. divaricata Leaves alternate (may be almost opposite in P. acuminata). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 Leaves opposite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18 Phyllaries rounded, ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. P. ciliata Phyllaries tapering, acute, not ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15 Phyllaries with thickened, glandular midrib; apex somewhat acute, reﬂexed; margins narrow, white. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19. P. rangei Phyllaries without thickened midrib but tapering into a sharp, long tip or if a shorter tip, then margins broad membranous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16 Phyllaries broadly lanceolate; midrib narrow, brown; remainder of phyllaries transparent, membranous; apex acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20. P. scariosa Phyllaries lanceolate; broad opaque centre brownish; margins membranous, transparent . . . . . . . . . . . . . .17 Leaves to 7 × 1.5 mm, oblong to narrowly obovate, narrowed into base, apex rounded, glabrous; phyllaries glabrous, lanceolate, 2 – 3 mm long acuminate, keeled; inner phyllaries to 2.5 mm broad; margins membranous; young branches white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. P. leucoclada Leaves to 20 × 8 mm, ovate-obovate to oblanceolate, sessile or short petiolate, apex acute; phyllaries silvery-grey puberulous, linear to lanceolate, apex acutely mucronate; inner phyllaries to 2 mm broad; margins membranous; young branches white, grey or brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1. P. acuminata More than one capitulum at branch tips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15. P. paniculata Capitula solitary at branch tips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19 Opposite leaves connate at base but not sheathing, the connecting part persistent when leaves dehisce, forming rows of spinules on branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22. P. spinulosa Leaves without clear connecting part and therefore without spinules on branches (leaf bases of P. sordida may be slightly fused, but do no form spinules after dehiscence) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20 Capitula cylindrical, to 8 mm in diam.; phyllaries yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21 Capitula broadly ovoid or obconical, to 15 mm in diam.; phyllaries brownish or dark green. . . . . . . . . . . .23 Phyllaries broadest at apex, somewhat constricted below apex, mucronate, bright yellow; apex truncate, often somewhat emarginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23. P. unguiculata Phyllaries obovate-roundish, mucronulate or without mucro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22 Phyllaries obovate to linear-oblong, very shortly mucronate, pale yellow; margins lacerate . . 5. P. cylindracea Phyllaries ovate, not mucronate, green with dark brown, glandular, longitudinal lines; margins hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. P. anisata Phyllaries broadly ovate to almost circular; apex rounded; margins narrow membranous all around . . . 8. P. glabrata Phyllaries oblong-oblanceolate; apex acute to obtuse; margins membranous laterally only . . . .21. P. sordida © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 4 of 44 KEW BULLETIN (2014) 69: 9488 Synopsis of Namibian species 1. Pteronia acuminata DC. (de Candolle 1836: 361); Harvey in Harvey & Sonder (1865: 105); Hutchinson & Phillips (1917: 315); Merxmüller (1967: 154); Merxmüller & Roessler (1984: 89); Herman (2003: 276). Type: South Africa, Northern Cape Prov., between Zack and Gariep rivers, 11 Sept. 1811, Burchell 1587 (holotype G-DC!; isotypes GH-00011510!, K000273485!, PRE). Dicoma ramosissima Klatt (1896: 843); Dinter (1921: 186). Syntypes: Namibia, Aob bei Keetmanshoop, 1892, Fleck 116 (GH-00006368 (fragment), Z000003295!); Namibia, Aob bei Keetmanshoop, 1891, Fenchel 51 (GH-00006368 (fragment), Z000003294!). Pteronia carnosa Muschl. (Muschler 1911a: 97), nom. illegit., non P. carnosa Muschl. (Muschler 1911a: 95). Type: Namibia, Bezirk des Damaralandes, Berseba, Schultze 406 (lectotype K-000273487!, selected here). Pteronia feddeana Muschl. (Muschler 1911b: 384, “Feddeana”), Range (1935: 275, “Feddeana”), nom. nov. for P. carnosa Muschl. (1911a: 97). Type: as for Pteronia carnosa Muschl. (1911a: 97). Shrub, 30 – 80 cm high, to 100 cm in diam. Stems terete, glabrous, brittle, bark white, grey to pale brown. Leaves alternate to almost opposite, somewhat succulent, glaucous, glabrous, ﬂat, ovate-obovate to oblanceolate, 10 – 20 × 5 – 8 mm; apex acute, mucronate; base cuneate, sessile or short petiolate. Capitula solitary, terminal, elongate-ovoid, to 25 mm long; apex pointed in bud. Receptacle slightly convex, honeycombed, to 4 mm in diam. Phyllaries multiseriate, gradate, outer broadly lanceolate, to 4 × 2 mm, inner linear to narrowly lanceolate, to 20 × 2 mm, silvery-grey puberulous; apex acutely mucronate; margins membranous. Florets 10 – 12; corollas to 2 mm long, yellow; lobes lanceolate, subacute. Achenes compressed, densely appressed villous, to 5 mm long. Pappus to 15 mm long, golden to straw-coloured. DISTRIBUTION. Africa: Botswana, Namibia, South Africa. Map 1. 1 NAMIBIA. Hardap Region, Maltahöhe Distr.: Grootfonteiner Fläche, 29 Sept. 1959, Giess 2286 (PRE, WIND!); 3 km from Maltahöhe on road to Helmeringhausen, on road verge, 19 Oct. 1987, Kolberg & Maggs HK94 (PRE, WIND!); Farm Grootfontein (Lisbon) MAL 9, 5 Sept. 1972, Merxmüller & Giess 28243 (M, PRE, WIND!); Farm Lisbon MAL 9, 8 April 1980, Müller 1274 (WIND!); Farm Naudaus/Duwisib MAL 76/84, 20 SPECIMENS EXAMINED. 1 With Specimens Examined we list inspected sheets from herbaria (marked with “!”) and other herbaria which are known to hold a duplicate specimen. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 Map 1. Known distribution of Pteronia acuminata in Namibia. May 1956, Volk 12557 (WIND!); Mariental Distr.: Farm Haribes GIB 18/19, 30 May 1963, Leippert 4734 (WIND!); Torro plain at station Ebenerde, 10 Sept. 1963, Merxmüller; Giess 3602 (M, PRE, WIND!); Karas Region, Bethanie Distr.: N of Helmeringhausen, 7 Sept. 2002, Burke 02074 (WIND!); Farm Goais 21 km (13.3 miles) S of Helmeringhausen, 19 May 1965, Giess 8811 (PRE, WIND!); Farm Huns, BET 106, 12 Aug. 1976, Giess 14601 (PRE, WIND!); Karasburg Distr.: Klein Karas, 7 Aug. 1923, Dinter 4862 (SAM, Z!); Farm Windkraal WAR 34, 20 May 1963, Giess, Volk & Bleissner 7120 (WIND!); Farm Vrede, S of house, 10 Sept. 2005, Kolberg & Tholkes HK1688 (K!, WIND!); Farm Mooiplaats WAR 97, 6 Oct. 1977, Merxmüller & Giess 32521 (M, PRE, WIND!); Klein Karas, April 1890, Schäfer 222 (K!); Farm Rishon, Karasberge, Graberberg. Near Post Ofﬁce tower, 23 June 1989, Van Wyk 8696 (PRE, WIND!); Keetmanshoop Distr.: 7 miles W of Aroab on road to Keetmanshoop, 3 May 1955, De Winter 3381 (PRE, WIND!); Aob bei Keetmanshoop, 1891, Fenchel 51 (Dicoma ramosissima syntype GH-00006368 (fragment), Z-000003294!); Aob bei Keetmanshoop, 1892, Fleck 116 (Dicoma ramosissima syntype GH-00006368 (fragment), Z-000003295!); Berseba Reservat KEE 170, 13 May 1963, Giess, Volk & Bleissner 6864 (WIND!); Berseba, Aug. 1905, Schultze 406 (P. feddeana lectotype K-000273487!); Farm Springboktrek South 223, on S end of pan N of homestead, 12 Feb. 1997, Strohbach, Kubirske & Sheuyange 2786 (WIND!); Farm Springboktrek South 223, on S end of pan N of homestead, 18 Feb. 1998, Strohbach 3650 (WIND!); Lüderitz Distr.: State land, Nuobrivier, Huns Mts, S of Farm Uitsig LU 82, 9 June 1976, Giess & Müller 14327 (WIND!); State land, 3 km S of border of Farm Uitsig, 25 Sept. 1976, Giess & Wendt 14702 (WIND!); Aus – Rosh Pinah road in KEW BULLETIN (2014) 69: 9488 washes near Aus Marble turnoff, 29 Sept. 2004, Mannheimer, Maggs-Kölling & Loots CM2656 (WIND!). SOUTH AFRICA. Northern Cape Prov., Carnarvon Distr.: at Karel Kriegers grave, between the Karmee Bergen [Kareeberge] and Buffels Bout, between Zack and Gariep rivers, 11 Sept. 1811, Burchell 1587 (P. acuminata holotype G-DC!; isotypes GH-00011510, K00273485!, PRE). HABITAT. Mostly on calcareous soil in dwarf shrub savanna; altitude c. 950 m; summer or summer-winter transition rainfall areas of south-central and southeastern Namibia (Map 1). CONSERVATION STATUS. In Namibia this species does not qualify for any of the threatened categories according to the ﬁve IUCN criteria. No threats to populations were identiﬁed and both the estimated Extent Of Occurrence (EOO) and Area Of Occupancy (AOO) of this fairly widespread species are well above the maxima for the threatened categories (IUCN 2001, 2013). The Namibian status therefore is Least Concern (LC). The South African threat status is cited as LC (SANBI 2011). PHENOLOGY. Flowering: April to October. Fruiting: August to February. NOTES. Muschler (1911a) caused some confusion by publishing two species under the name Pteronia carnosa, one of which (1911a: 97) he corrected in the same year to P. feddeana Muschl. (Muschler 1911b). Hutchinson & Phillips (1917) considered this taxon to be conspeciﬁc with P. acuminata, which was followed by more recent authors (Merxmüller 1952, 1967; Herman 2003), an opinion with which we agree. The second P. carnosa (Muschler 1911a: 95), an illegitimate name, is now considered a synonym of P. glabrata L. f. (q.v.). In his description of P. carnosa Muschler (1911a: 97) cited Schultze 406 and von Trotha 129. Schultze 406 was traced in K, but the von Trotha specimen was not found in any herbarium where it was likely to be housed. Muschler worked from B and the specimen was most likely preserved there and destroyed during World War II. Schultze 406 at K is therefore selected as the lectotype. 9488, Page 5 of 44 campanulate-obconic, 10 – 15 × 10 mm, surrounded at its base by short leaves; apex rounded in bud. Phyllaries multiseriate, gradate, outer linear lanceolate, to 5 mm long, inner linear-oblong, to 10 mm long, yellow, woolly pubescent on outside; apex acute, mucronate; margins very narrow, membranous. Florets c. 14; corollas to 6 mm long, pubescent on outside, yellow; lobes linear, subacute; tube ribbed at base. Achenes obconic, long sericeous, c. 3 mm long. Pappus setae barbellate, to 7 mm long, yellow to straw-coloured. DISTRIBUTION. Africa: Namibia, South Africa. Map 2. SPECIMENS EXAMINED. NAMIBIA. Hardap Region, Maltahöhe Distr.: Naukluft plateau, 6 July 1993, Bridgeford 142 (WIND!); Naukluft, Kapokvlakte, on plateau, Nov. 1995, Bridgeford 95360 (WIND!); Naukluft Plateau, Kapokvlakte, March 1994, Günster 9452 (WIND!); Farm Zaris MAL 103, at peak of pass, 4 Sept. 1972, Merxmüller & Giess 28202 (M, WIND!); Farm Rooiberg-Süd, 1935, Steyn 9960 (WIND!); Farm Wereldend 115, Helmeringhausen, 17 May 1956, Volk 12798 (WIND!); Karas Region, Bethanie Distr.: Farm Aruab 23, plains on S part, April 1998, Miller MIL1/ 065 (WIND!); Lüderitz Distr.: Farm Witpütz Nord LU 22, 1 km E of police station, 30 Sept. 1975, Giess 13779 (WIND!); State land 3 km S of border to Farm Uitsig, 26 Sept. 1976, Giess & Wendt 14706 (WIND!); Farm Plateau LU 38, not far from the farmhouse, Aus, 8 Sept. 1963, Kräusel & Wiss 2016 (WIND!); Chamis, Sept. 1905, Schultze 433 (lectotype K-00273412!). HABITAT. Rocky areas in desert-dwarf shrub savanna transition zone; c. 1600 m. CONSERVATION STATUS. In Namibia this species is evaluated LC (IUCN 2001, 2013) as no threats to populations could be identiﬁed and the estimated EOO and AOO are above those that would qualify it for the 2. Pteronia acuta Muschl. (Muschler 1911a: 99); Hutchinson & Phillips (1917: 286); Dinter (1926: 131); Range (1935: 275); Merxmüller (1967: 154); Merxmüller & Roessler (1984: 89); Herman (2003: 276). Type: Namibia, Bezirk des Damaralandes, Chamis, Sept. 1905, Schultze 433 (lectotype K-00273412!, selected here). Shrub, 15 – 50 cm high, to 30 cm in diam. Stems appressed pubescent, pale grey, Leaves opposite, usually clustered along stems, pale grey, densely felty to woolly pubescent, linear to linear-lanceolate, trigonous, 4 – 10 × 1 – 1.5 mm; apex acute to subobtuse; connate at base. Capitula solitary, terminal, Map 2. Known distribution of Pteronia acuta in Namibia. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 6 of 44 threatened categories. The South African threat status is recorded as Least Concern (LC) (SANBI 2011). PHENOLOGY. Flowering: March to September. Fruiting: July to November. NOTES. In Namibia the species occurs along the southwestern escarpment between the coastal Namib desert and the inland plateau (Map 2) and prefers altitudes above 1500 m. When capitula are immature and the yellowish pappus is not visible, this species could be mistaken for Pteronia eenii but the known distributions of these species do not overlap (compare Maps 2 and 7). In his description Muschler (1911a: 100) cited Schultze 433 and von Trotha 147a, both with exactly the same locality and date. Schultze 433 could be traced in K, but von Trotha 147a was not found in any herbarium where it was likely to be housed. As Muschler worked from B any original specimen was most likely destroyed during World War II. Schultze 433 at K is thus selected as the lectotype. 3. Pteronia anisata B. Nord. (Nordenstam 1971: 10); Roessler & Merxmüller (1982: 192); Merxmüller & Roessler (1984: 89); Herman (2003: 276). Type: South Africa, Northern Cape Prov., Namaqualand Div., KEW BULLETIN (2014) 69: 9488 Richtersveld, Cornell's Kop, N slopes, 31 Oct. 1962, Nordenstam 1694 (holotype S-G-5140!; isotypes M0104529!, PRE, SAM). Aromatic shrub, to 40 cm high. Stems brown when young, dark grey when older. Leaves opposite, clustered along stems, succulent, green with brown resin dots, surface minutely papillate, terete to trigonous to somewhat ﬂattened, to 20 × 1.5 mm, base somewhat clasping with tuft of hairs in axil. Capitula solitary, terminal, cylindrical, to 10 × 6 mm; apex rounded in bud. Receptacle with lobed and fringed scales. Phyllaries 3 – 4-seriate, ovate to oblong, outer 3 × 1.5 mm, inner 10 × 4 mm, green with dark, resinous striations; apex obtuse; margins membranous white. Florets to 8; corollas 7 – 7.5 mm long, yellow; lobes triangular with few resin glands on outside; tube puberulous at base. Achenes obovoid with constricted apical neck, densely off-white sericeous, 3 – 4 × 2 mm. Pappus of numerous setae united into an annulus at base, 5 – 6 mm long, pale red-brown. Fig. 1. DISTRIBUTION. Africa: Namibia, South Africa. Map 3. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Lüderitz Distr.: Lorelei, fountain with "waterfall" in dune area Fig. 1. Pteronia anisata showing the bead-like veins on the phyllaries. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 PHOTO: H. KOLBERG. KEW BULLETIN (2014) 69: 9488 9488, Page 7 of 44 recorded in southern Namibia in 1977, some 170 km NNW of the type locality. Only 30 years later the species was found again just north of the Namibian – South African border (Map 3 & Fig. 1). The known distribution of Pteronia anisata suggests that it is nearendemic to Namibia and rare. Map 3. Known distribution of Pteronia anisata in Namibia. (NE), 12 Oct. 2007, Kolberg & Tholkes HK2381 (K!, WIND!); Sperrgebiet. Daberas Vley, W of dunes, E of Schakalsberge, 16 Oct. 2008, Kolberg & Tholkes HK2660 (K!, WIND!); Sperrgebiet, 19 km S of Obib Fountain, track turns E and leads over hill, 24 Oct. 2008, Kolberg & Tholkes HK2695 (K!, WIND!); Diamond Area 1, Tsaus Spinnenberg, Oct. 1977, Wendt 15/3 (M, WIND!). SOUTH AFRICA. Northern Cape Prov.: Namaqualand Div., Richtersveld, Cornell's Kop (near Annisfontein), N slopes, 31 Oct. 1962, Nordenstam 1694 (holotype S (coll. S-G-5140)!; isotypes M-0104529!, PRE, SAM). HABITAT. On blue dolomite or white quartz outcrops in steppe dominated by succulent species; 200 – 450 m. CONSERVATION STATUS. Pteronia anisata occurs in the protected diamond mining area of Namibia where human access and activities are strictly controlled and limited. Mining occurs in very restricted localities and mostly along the coast, outside the distribution area of this species. Although the estimated EOO and AOO would qualify the species as VU under criterion B, the required two criteria of fragmentation, decline or ﬂuctuation of populations have not been observed. Similarly, under criterion D2 P. anisata is known from fewer than 5 localities but this is not coupled with the required plausible future threat. The species therefore is evaluated as LC in Namibia (IUCN 2001, 2013), but the scarcity of herbarium specimens and information from recent collections suggests that it is rare in Namibia. PHENOLOGY. Flowering: October. ETYMOLOGY. The speciﬁc epithet denotes that the plant smells of aniseed (Nordenstam 1971). NOTES. Described in 1971 from the Richtersveld in South Africa (Nordenstam 1971), Roessler & Merxmüller (1982) reported that this species was also 4. Pteronia ciliata Thunb. (Thunberg 1800: 144); de Candolle (1836: 359); Harvey in Harvey & Sonder (1865: 102); Hutchinson & Phillips (1917: 317); Merxmüller (1967: 155); Merxmüller & Roessler (1984: 89); Herman (2003: 277). Type: South Africa, Western Cape Prov., Cape of Good Hope, Thunberg s.n. (holotype UPS-THUNB; isotype S-07-7225!). Pteronia ciliata Thunb. var. [“β”] subtrigona DC. (de Candolle 1836: 360). Type: South Africa, Western Cape Prov., Olifantrivier, W. F., Drège s.n. [5664 — see Notes] (holotype G-DC; isotypes HBG-505132!, P-027011!). Pteronia ciliata Thunb. var. ecklonis Harv. (Harvey in Harvey & Sonder 1865: 102, “Ecklonis”). Type: South Africa, Northern Cape Prov., Namaqualand, Nov. 1837, Ecklon & Zeyher 238 (holotype PRE). — see Notes. Pteronia ciliata Thunb. var. thunbergii Harv. (Harvey in Harvey & Sonder 1865: 102, “Thunbergii”), nom. illegit. Type: as for P. ciliata. Pteronia turbinata DC. (de Candolle 1836: 362); Harvey in Harvey & Sonder (1865: 106). Type: South Africa, Northern Cape Prov., Gariep, Klein-Namaqualand, Drège s.n. [5665] (holotype G-DC; isotypes HAL-0110987, HBG-505180!, NY-00232710!, P027281!, P-027282!, P-027283, P-027284!, PRE, WRchb-1889-0278258!, W-0008809!). — see Notes. Shrub, to 100 cm high. Stems slightly puberulous, pale grey. Leaves alternate, clustered on stem tips, succulent, glabrous, linear, grooved above, keeled below, to 10 × 1 – 1.5 mm; apex obtuse, base cuneate; pleasant aroma. Capitula solitary, terminal, ovoid-cylindrical, to 15 – 20 × 5 – 7 mm; apex contracted. Receptacle concave, ﬁmbriate-honeycombed. Phyllaries 7 – 10seriate, gradate, ovate to linear, to 12 mm long, straw-coloured; midrib faint; apex obtuse; margins distinctly ciliate. Florets to 15; corollas to 10 mm long, glabrous, greenish-yellow; lobes linear-lanceolate, acute. Achenes obconic, long sericeous, to 3 mm long. Pappus multiseriate, to 5 mm long, brownish-yellow. DISTRIBUTION. Africa: Namibia, South Africa. Map 4. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Lüderitz Distr.: Diamond Area 1, Klinghardts Mts, 2 Jan. 1996, Burke 96134 (WIND!); Sperrgebiet. N Klinghardts Mts, 5 Aug. 2001, Klaassen & Bartsch EK484 (WIND!); First level section above the E foot of Rooiberg (Sperrgebiet), 4 Sept. 1992, Kubirske, Strohbach & Swart 42 (WIND!); Sperrgebiet. Klinghardts Mts, 11 Sept. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 8 of 44 Map 4. Known distribution of Pteronia ciliata in Namibia. 2005, Kwembeya EKw64 (WIND!); Granite outcrops on road to Obib, SW of Farm Spitzkop, 1 Sept. 1963, Merxmüller & Giess 3424 (M, WIND!); W edge of Obib Mts Hill 3 km N of Obibwasser, 2 Sept. 1977, Merxmüller & Giess 32382 (M, WIND!); Gabusib, Feb. 1992, Strohbach 73 (WIND!). SOUTH AFRICA. Northern Cape Prov.: Klein Namaqualand, between Kaus, Natvoet and Doornpoort [III B 6], 17 Oct. 1830, Drège s.n.; Namaqualand, Drège s.n. [5665] (P. turbinata holotype G-DC; isotypes HAL-0110987, HBG-505180!, NY-00232710!, P-027281!, P-027282!, P-027283, P027284!, PRE, W-Rchb-1889-0278258!, W-0008809!); Namaqualand, Nov. 1837, Ecklon & Zeyher 238 (P. ciliata var. ecklonis holotype PRE, possible isotype HBG-505133!); Western Cape Prov.: Ebenezar, on rocky, dry karoo hills [III E a 4], Nov. 1833, Drège s.n. (K-000273499!); Olifantrivier, W. F., Drège s.n. [5664] (P. ciliata var. subtrigona holotype G-DC; isotypes HBG-505132!, P027011!); Cape of Good Hope, Thunberg s.n. (P. ciliata holotype UPS-THUNB; isotype S-07-7225!). HABITAT. Rocky slopes in steppe dominated by succulent species; 700 – 800 m. CONSERVATION STATUS. The estimated EOO and AOO of this species falls below the threshold value for the VU category, but two of the additional criteria of population fragmentation, decline or ﬂuctuation could not be satisﬁed. Also under criterion D2, although the number of localities is ≤5, qualifying for VU, a plausible future threat could not be established. The Namibian threat status thus is LC (IUCN 2001, 2013). Present data, however, suggest that the species is rare in Namibia. The South African threat status is LC (SANBI 2011). PHENOLOGY. Flowering: February to September. Fruiting: August. VERNACULAR NAME. Biltongbos (Afrikaans, South Africa). © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 NOTES. Only a few collections have been made in Namibia (and all of these since 1963) but Pteronia ciliata is more widespread in South Africa. The locality cited with the holotype (as well as in de Candolle’s Prodromus) of Pteronia ciliata var. subtrigona is only “Olifantrivier”, a river that runs through the Western Cape Province in South Africa. Similar to P. latisquama (a synonym of the accepted P. glauca, see below) it is problematic to ascertain which collections are possibly isotypes of the varietal name. Critical evaluation includes P-027011, as it carries not only “5664” on the label, but has another small label attached, which we believe may well present locality details, date, and coding in Drège’s handwriting. It reads (in our annotated version): “/11 33 [as in: November 1833] [unreadable location name] bei Ebenezar [which is on the Olifant River], — 300 [as in: meters altitude] (III, E, a)”. (The coding system of III, E, a, etc. is explained below in the notes at P. glauca.). More locality details are provided by, for example HBG-505132 (annotated with “Pteronia ciliata β. DC.” and considered an isotype): “Ebenezar, auf steinigen, trockenen (karrooartigen) Hügeln” with the locality code “III E a 4” added. Other specimens marked similarly “5664” but in clearly different handwriting are P-027010 and P-027013. A similar problem is a possible isotype (HBG-505133) of P. ciliata var. ecklonis: this is reportedly an Ecklon & Zeyher collection, but does not show no. 238 and while considered an (iso-) type this is with a question mark. The locality cited for the type of Pteronia turbinata is the one recorded on the holotype, but more precise locations are found on some of the isotypes, such as “zwischen Kaus, Natvoet und Doornpoort [III B 6]”, Drège s.n. (HAL-0110987, HBG-505180, W-0008809); but “17 / 10 30 (= 17 Oct. 1830), Kaus” (in what looks like Drège’s handwriting) and the number 5665 are on isotype P-027281. The date mentioned on K-00273497 is 1837, which could refer to a collection or herbarium dispatch or receipt date. This specimen, as well as K00273498 does not refer to the number “5665” and both are not considered to be isotypes. It should be noted that here and elsewhere for taxa described in the Prodromus by Auguste Pyramus de Candolle, we follow the information provided to us by Dr Fernand Jacquemoud in Geneva who kindly inspected the type material. 5. Pteronia cylindracea DC. (de Candolle 1836: 363); Harvey in Harvey & Sonder (1865: 106); Hutchinson & Phillips (1917: 313); Dinter (1926: 132); Range (1935: 275); Merxmüller (1967: 155); Merxmüller & Roessler (1984: 89); Herman (2003: 277). Type: South Africa, Northern Cape Prov., at Buffels Bout, Carnarvon Div., 12 Sept. 1811, Burchell 1603 (holotype G-DC; isotypes K-00273480!, P-027025!, PRE). KEW BULLETIN (2014) 69: 9488 Shrub to 40 cm high. Stems glabrous, older branches very woody, bark grey. Leaves opposite, mostly clustered along stems, glabrous, linear, ﬂat or grooved above, to 10 × 1 mm; apex hooked; base sometimes eared and clasping stem but not connate. Capitula solitary, terminal, cylindrical, 20 – 25 × 6 mm; apex acute. Phyllaries multiseriate, gradate, obovate to linear-oblong, 5 – 15 × 4 mm; pale, matte yellow, chaffy-membranous; apex obtuse, mucronulate; margins lacerate. Florets c. 5; corollas to 15 mm long, exserted up to 8 mm from phyllaries, yellow; lobes linear lanceolate, subacute; tube widening in upper half. Achenes narrowly obconic, long sericeous and densely sessile glandular, c. 5 mm long. Pappus multiseriate, to 10 mm long, straw-coloured. DISTRIBUTION. Africa: Botswana, Namibia, South Africa. Map 5. SPECIMENS EXAMINED. NAMIBIA. Khomas Region, Windhoek Distr.: Khomas Hochland, 14 May 2000, Burke 00090 (WIND!); Farm Harris (22), 32 km (20 miles) SW of Windhoek, 2 March 1955, De Winter 2546 (WIND!); Farm Mahonda WIN 39, 12 July 1963, Giess 7628 (WIND!); Farm Bergland, 12 Aug. 1963, Merxmüller & Giess 3593 (M, PRE, WIND!); Verdwaal (REH 41), 29 Aug. 1972, Merxmüller & Giess 28087 (M, PRE, WIND!); Farm Naos, 1 Jan. 1953, Schwerdtfeger 4227 (WIND!); Avis, Windhoek, 6 May 1964, Seydel 4016 (WIND!); Avis, the mountain parallel with the Avis R., at power lines, 26 July 1966, Seydel 4432 (WIND!); Farm Haris, on S part of pan, 18 April 2000, Strohbach BS5102 (WIND!); Farm Lichtenstein Süd, dissected plain towards SW corner, 29 April 2002, Strohbach BS5586 (WIND!); Grazing plots. Neudam Experimental Farm, 23 June 1959, Van Vuuren 595 (PRE, WIND!); Omaheke Region, Gobabis Distr.: Map 5. Known distribution of Pteronia cylindracea in Namibia. 9488, Page 9 of 44 Farm Sturmfeld, Steinhausen, 19 June 1958, Schwerdtfeger 4081 (WIND!); Farm Sturmfeld GO 252, Steinhausen, 14 July 1962, Toelken 156 (WIND!); Otjinene Distr.: among rocks and stones on banks of Eiseb omuramba, 54 miles from Gam, on way to Windhoek, 31 Aug. 1955, Story 5340 (PRE, WIND!); Otjozondjupa Region, Okahandja Distr.: Farm Osema OK 63, 25 June 1965, Giess 9006 (WIND!). SOUTH AFRICA. Northern Cape Prov.: at Buffels Bout, Carnarvon Div., 12 Sept. 1811, Burchell 1603 (holotype G-DC; isotypes K-00273480!, P-027025!, PRE). HABITAT. Plains, slopes, dry river banks in high altitude shrubland and woodland on Kalahari sands; 1800 – 2100 m. CONSERVATION STATUS. This relatively widespread species has an estimated EOO and AOO well above the thresholds for the threatened categories. No small or restricted populations or decline in population size could be identiﬁed to satisfy any of the ﬁve criteria according to IUCN (2013). The Namibian conservation status is therefore LC (IUCN 2001). SANBI (2011) lists the South African threat status as LC. PHENOLOGY. Flowering: January to August. Fruiting: July to December. NOTES. It is very difﬁcult to separate this species from the closely related Pteronia unguiculata, and Merxmüller (1967) remarked that there is an overlap in the phyllary characters whereas it is these same characters that are used to separate these species. The specimens at WIND could mainly be separated on a phytogeographical basis: P. cylindracea occurs in Kalahari sand in the north and east of the country while P. unguiculata is found on rocky, mountainous terrain at higher altitudes in the south-west and west of Namibia. In general, capitula of P. unguiculata are smaller and obtuse in bud, while those of P. cylindracea are acute; the phyllaries of P. unguiculata are shiny, bright yellow while those of P. cylindracea are a matte, pale yellow. More material is needed from the north-east of Namibia and neighbouring areas in Botswana and South Africa to arrive at a more deﬁnite conclusion on the relationship between these two species, taking it outside the scope of this study. 6. Pteronia divaricata Less. (Lessing 1832: 196); de Candolle (1836: 357); Harvey in Harvey & Sonder (1865: 99); Hutchinson & Phillips (1917: 296); Merxmüller (1967: 155); Merxmüller & Roessler (1984: 89); Herman (2003: 277). Type: South Africa, Western Cape Prov., Cape of Good Hope, s.coll. s.n. (lectotype LINN 982.1!, selected by Lowrey in Jarvis & Turland (1998: 358) for Chrysocoma oppositifolia). Eupatorium divaricatum P. J. Bergius (1767: 229); Thunberg (1800: 142), nom. illegit. — see Notes. Chrysocoma oppositifolia L. (Linnaeus 1760: 18, 1763a: 97, 1763b: 1177), non P. oppositifolia L. Type: as for P. divaricata — see Notes. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 10 of 44 Shrub to 150 cm high. Stems puberulous, tan to grey. Leaves opposite, grey-green, surface puberulous-papillate and glandular, ﬂat, obovate to orbicular, 25 × 15 mm; apex subacute to obtuse; base narrowed into a short petiole; aromatic. Capitula in terminal corymbs, turbinate, 15 × 3 – 4 mm; apex acute in bud. Phyllaries multiseriate, outer broadly ovate, 2.5 mm long, inner linear-oblong, 8 mm long, green to yellow, glabrous; apex obtuse or subacute; margins membranous. Florets c. 5, sweetish smell; corollas 10 – 12 mm long, white to cream; lobes linear-lanceolate, acute; tube gradually widening, ribbed and pubescent below middle. Achenes obovoid, compressed, sparingly pubescent, densely glandular, c. 5 mm long. Pappus multiseriate, setae connate at base, barbellate, 8 – 10 mm long, whitish or with reddish tint. DISTRIBUTION. Africa: Namibia, South Africa. Map 6. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Lüderitz Distr.: S of Lüderitz. Diamond Area No. 1. Mountain slopes E of Aurus Peak in Aurus Mts, 11 Aug. 2001, Burgoyne 8431b (PRE, WIND!); Rooiberg, Sperrgebiet, 28 Sept. 1996, Burke 96231 (WIND!); About 34 km (21 miles) N of Lorelei Copper Mine on the road to Witputz, 15 Sept. 1958, De Winter & Giess 6408 (PRE, WIND!); Farm Namuskluft LU 88, 8 km E of Rosh Pinah, 13 Aug. 1976, Giess 14614 (PRE, WIND!); Ai-Ais conservation area, McMillan's Pass, 2 km SE of summit of pass along track, 12 Oct. 2005, Kolberg & Tholkes HK1697 (K!, WIND!); Namuskluft LU 88, 13 Sept. 1963, Kräusel & Wiss 2071 (WIND!); 35 km N of Rosh Pinah (road to Aus), 2 – 4 km W of road, 29 Oct. 1983, Leuenberger, Raus & Schiers 3248 (WIND!); Base of Rooiberg, 29 Sept. 1996, Mannheimer & Mannheimer 420 (WIND!); Aurus basin, 10 Aug. 2001, Mannheimer CM1558 (WIND!); Namuskluft, hill behind hall, 11 Map 6. Known distribution of Pteronia divaricata in Namibia. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 Sept. 2002, Mannheimer CM2256 (WIND!); Farm Witputz, 26 Aug. 1963, Merxmüller & Giess 3196 (M, PRE, WIND); Namuskluft LUS 88, 9.5 km E of Rosh Pinah, 18 Sept. 1972, Merxmüller & Giess 28533 (M, PRE, WIND!); Farm Zebrafontein LUS 87, 2 April 1972, Merxmüller & Giess 28783 (M, WIND!); Aurus Mts, quartz koppie 7 km N of the mountains, 20 Sept. 1977, Merxmüller & Giess 32189 (M, PRE, WIND!); Farm Zebrafontein, 22 Sept. 1981, Müller & Horn 1564 (WIND!); Sperrgebiet, Aurus Mts, main complex around the beacon, 6 Sept. 1992, Oliver 10148 (WIND!); Numaeis, S of Witputz, Sept. 1957, Rusch 4678 (WIND!); Sperrgebiet, N end of Aurus Mts on W side, 8 Aug. 2001, Smook 11255 (PRE, WIND!); Sperrgebiet, N end of Aurus Mts on W side, 8 Aug. 2001, Smook 11263 (PRE, WIND!); Sperrgebiet, SE side of Aurus Mts just below the highest peak, 11 Aug. 2001, Smook 11341 (PRE, WIND!); Rooiberg, 30 Sept. 1992, Strohbach 299 (WIND!); Rosh Pinah, Namuskluft 88, 31 Aug. 1989, Van Wyk 8827 (PRE, WIND!); Rosh Pinah. Namuskluft 88, 31 Aug. 1989, Van Wyk 8828 (PRE, WIND!). SOUTH AFRICA. Western Cape Prov.: Groenkloof, Drège s.n. (P-027029!, PRE); Cape of Good Hope, s.coll., s.n. (lectotype LINN 982.1!). HABITAT. Rocky slopes and sandy plains in steppe dominated by succulent species; 600 – 1000 m. CONSERVATION STATUS. The Namibian distribution of this species is restricted with an estimated EOO below the threshold for the VU category under criterion B1 but two of the additional criteria of population fragmentation, decline or ﬂuctuation could not be identiﬁed (IUCN 2013). The Namibian threat status therefore is LC (IUCN 2001). The South African threat status is also recorded as LC (SANBI 2011). PHENOLOGY. Flowering: April to November. Fruiting: August to December. VERNACULAR NAMES. Geelknopbos, spalkpenbos (Afrikaans, South Africa). NOTES. This species is restricted to the extreme southwest of Namibia (Map 6), which receives winter rainfall. It is more common in the Northern and Western Cape Provinces of South Africa. According to Merxmüller (1967) Pteronia divaricata is reported to have yellow ﬂorets in South Africa whereas in Namibia only white to cream ﬂorets have been seen. The type of Pteronia divaricata is the same as that of the Linnaean Chrysocoma oppositifolia. The latter name was ﬁrst mentioned in the thesis of Jacob Printz, Plantae rariores Africanæ, which appeared in 1760 and where the Chrysocoma is cited under no. 49 on p. 18. The thesis is also part of Linnaeus’s Amoenitates academicæ, vol. 6 (1763a: 97), and this version is cited as “Amœn. acad. 6. afr. 49” in the second edition of Species plantarum, vol. 2 (1763b: 1177). Both works by Linnaeus are cited by Bergius when he renamed the KEW BULLETIN (2014) 69: 9488 9488, Page 11 of 44 species Eupatorium divaricatum (1767: 229), but such a complete renaming is superﬂuous and illegitimate. Lessing’s recombination into Pteronia using the epithet divaricata created, however, a legitimate replacement name (as P. oppositifolia would be blocked by the earlier Linnaean name); its author citation should ascribe to Lessing only. 7. Pteronia eenii S. Moore (1902: 325); Hutchinson & Phillips (1917: 287); Merxmüller (1952: 124; 1967: 155); Merxmüller & Roessler (1984: 89); Herman (2003: 277). Type: Namibia, Damaraland, 1879, Een s.n. (holotype BM-000903813!). Dicoma seitziana Dinter (1921: 186, “Seitziana”). Type: Namibia, Otjozondjupa Region, Grootfontein Distr., Gemsbocklaagte, Aug. 1911, Dinter 3007 (lectotype P00138958!, selected here; isolectotypes four sheets in SAM: SAM-0071799-1to -4, all “!”) — see Notes. Pteronia feldtmanniana Dinter ex Merxm. (Merxmüller 1955: 79), nom. nud. — see Notes. Shrub to 50 cm high and in diam. Stems white to grey woolly. Leaves opposite, densely grey, woolly-felty, Fig. 2. Capitulum of Pteronia eenii near Grootfontein. linear, 8 – 15 × 1 – 1.5 mm; apex obtuse; base slightly eared and clasping stem. Capitula solitary, terminal, about half its length enclosed by leaves, obconic, 13 – 15 × 8 mm; apex acute in bud. Receptacle ﬂat. Phyllaries multiseriate, gradate, lanceolate to linear-lanceolate, 4 – 11 mm long, red-brown, maroon to purple, woolly outside; apex acute, sharply mucronate; margins narrowly transparent-maroon membranous. Florets c. 15; corollas 7 mm long, glabrous, yellow; lobes linearlanceolate, subacute, tubes constricted and ribbed at base. Achenes turbinate, 3.5 mm long, densely appressed white villous. Pappus multiseriate, to 11 mm long, purple to maroon. Fig. 2. DISTRIBUTION. Africa: endemic to Namibia. Map 7. SPECIMENS EXAMINED. NAMIBIA. Unknown Region and Distr.: Damaraland, 1879, Een s.n. (P. eenii holotype BM-000903813!); Khomas Region, Windhoek Distr.: Windhoek and surroundings, 9 March 1988, Bohlmann 88/ 83 (WIND!); Kunene Region, Opuwo Distr.: 9.5 km NW of Ombombo (former road D3708?), 13 June 2007, Kolberg & Tholkes HK2327 (K!, WIND!); Outjo Distr.: Farm Westland, cattle post to Elf, 21 Oct. 1998, Hobohm HOB1 28 (WIND!); Omaheke Region, PHOTO: H. KOLBERG. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 12 of 44 KEW BULLETIN (2014) 69: 9488 VERNACULAR NAME. |haurus (Khoekhoegowab, Namibia). USES. Forage. NOTES. Until recently this Namibian endemic has been Map 7. Known distribution of Pteronia eenii in Namibia. Gobabis Distr.: On S banks of Epukiro, Sturmfeld, 20 June 1953, Schwerdtfeger 4115 (WIND!); Otjozondjupa Region, Grootfontein Distr.: Gemsbocklaagte, Aug. 1911, Dinter 3007 (Dicoma seitziana lectotype P00138958!, isolectotypes SAM-0071799-1 to -4!); Grootfontein, 11 July 1934, Dinter 7691 (HBG!, WIND!); Farm Rietfontein GR 344, 26 April 1963, Giess, Volk & Bleissner 6523 (PRE, WIND!); Farm Achlam GR 583, on Elefantenberg, 12 Nov. 1976, Giess 14790 (PRE, WIND!); Rietfontein GR 344, 3 km (2 miles) towards Waterberg, 27 April 1963, Kers 356 (WIND!); 6.5 km N of Grootfontein on road to Rundu, 23 June 2008, Kolberg & Tholkes HK2633 (K!, WIND!); Near Meteor, Grootfontein, 19 July 1965, Leach & Bayliss 13013 (PRE, WIND!); Farm Nassau, 11 Jan. 1953, Walter & Walter 963 (WIND!); Otjiwarongo Distr.: Omatjenne Experimental Farm, 1940, Pfeiffer s.n. (WIND-34113!); Okosongomingo, June 1940, Volk 3062 (WIND!). HABITAT. Rocky calcrete areas in shrubland or woodland; 1400 – 1700 m. CONSERVATION STATUS. Evaluated by the 1994 IUCN criteria as Lower Risk — Least Concern (LRlc) by Craven & Loots (2002). Using more recent criteria (IUCN 2001) the evaluation is found here to be Least Concern (LC) since the estimated EOO and AOO are well above the maxima for the threatened categories and none of the other criteria A to E are met (IUCN 2013). The possible reduction in population size (see Notes) would also be below the thresholds for the threatened categories (at least 30% for VU) resulting in an evaluation of LC. Since this is a Namibian endemic species, this is a global conservation status (IUCN 2001). PHENOLOGY. Flowering: January to July. Fruiting: June to November. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 recorded only in the central and north-eastern parts of the country (Map 7). In 2007 the ﬁrst author found a small population of Pteronia eenii in north-western Namibia on calcareous soil typical for the species (Fig. 2). This disjunct distribution could be explained by either undercollection in the north-central areas of Namibia or by the species’ preference of calcrete substrates, which are not prevalent in the far central north, or over-utilisation of the species as a forage (reported on several herbarium specimens), thus reducing population sizes. The holotype of Pteronia seitziana was destroyed in Berlin, and we designate the P collection, indicated as “isotype” (unlike the collections in SAM) as the lectotype. The name P. feldtmanniana used by Dinter on his collections was listed by Merxmüller (1955: 79) as synonymous with P. eenii, but was never validly published. 8. Pteronia glabrata L. f. (Linnaeus ﬁlius 1782: 356); Thunberg (1800: 143); de Candolle (1836: 362); Harvey in Harvey & Sonder (1865: 102); Hutchinson & Phillips (1917: 321); Dinter (1926: 132, 1931: 167); Range (1935: 275); Merxmüller (1952: 125; 1967: 155); Merxmüller & Roessler (1984: 89); Herman (2003: 277). Type: South Africa, Western Cape Prov., Cape of Good Hope, Thunberg s.n. (holotype UPS-THUNB; isotype S-G-5142!). Pteronia sesuviifolia DC. (de Candolle 1836: 360). Type: South Africa, Western Cape Prov., Kaus, R III, 1835, Drège s.n. [2778] (G-DC no. G-00322348! lectotype, selected here). — see Notes. Pteronia carnosa Muschl. (Muschler 1911a: 95). Type: Namibia, Lüderitzbucht, 4 Jan. 1910, Dinter 1026 (holotype B†; isotype SAM-71804-0!). Pteronia succulenta auct. non Thunb.: Dinter (1926: 132); Range (1935: 276). Pteronia anisata Dinter ex Merxm. (Merxmüller 1952: 124 – 125), nom. nud. Pteronia glabrata L. f. var. succulenta (Thunb.) Merxm. (Merxmüller 1952: 125), quoad specim. cit., non quoad basionym. Shrub, to 60 cm tall. Stems glabrous, brittle, initially with pale grey bark, brown when older. Leaves opposite, ﬂeshy to succulent, glabrous, ﬂattish, oblong to linear, boat-shaped, adaxially grooved, to 10 – 25 × 3 – 5 mm, apex rounded, base clasping stem but not connate with opposite leaf. Capitula solitary, terminal, ovoid, to 20 × 15 mm, apex rounded in bud. Receptacle honeycombed and slightly ﬁmbriate. Phyllaries multiseriate, gradate, broadly ovate to orbiculate, 3 – 15 mm long, glabrous, papery and brown at maturity, opaque central part green with red margin; apex rounded; margins narrow, membranous all round. KEW BULLETIN (2014) 69: 9488 9488, Page 13 of 44 Florets 7 – 12; corollas c. 10 mm long, glabrous, bright yellow; lobes linear-lanceolate, subacute; tube gradually widening. Achenes turbinate, densely appressed villous, to 3.5 mm long. Pappus multiseriate, setae connate at base, to 10 mm long, brown to golden. Fig. 3. DISTRIBUTION. Africa: Namibia, South Africa. Map 8. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Lüderitz Distr.: Diamond Area No 1, Chamnaub Inselberg, NE of Boegoeberg, 28 Aug. 2002, Bartsch, Loots & Mannheimer SB930 (WIND!); Diamond Area 1, Namitsis Inselberg, about 30 km S of Klinghardts, 1 Sept. 2002, Bartsch, Loots & Mannheimer SB956 (WIND!); Diamond Area 1, at foot of Tsabiams Mtn, 4 Sept. 2002, Bartsch, Loots & Mannheimer SB1021 (WIND!); Diamond Area 1, en route from Tsabiams to Grillental, 5 Sept. 2002, Bartsch, Loots & Mannheimer SB1031 (WIND!); Diamond Area 1, Rote Kuppe – Chamais road, 6 Sept. 2002, Bartsch, Loots & Mannheimer SB1036 (WIND!); Diamond Area No 1, S of Lüderitz, about 48 km from Pomona on road to Klinghardts Mts, 4 Aug. 2001, Burgoyne 8286 (PRE, WIND!); Diamond Area 1, Klinghardts Mts, 21 Sept. Fig. 3. Flowering Pteronia glabrata. Map 8. Known distribution of Pteronia glabrata in Namibia. 1996, Burke 96115 (WIND!); Oranjemund. Gais area, E of Hohenfels on the road to Jakkalsberge, 5 Sept. 1958, De Winter & Giess 6201 (PRE, WIND!); Mile 50 PHOTO: H. KOLBERG. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 14 of 44 on road Lüderitz to Oranjemund, 6 Sept. 1958, De Winter & Giess 6224 (PRE, WIND!); Farm Spitskop LU 111, 35 km (22 miles) N of Lorelei Copper mine on road to Witputs, 15 Sept. 1958, De Winter & Giess 6411 (PRE, WIND!); Lüderitzbucht, 4 Jan. 1910, Dinter 1026 (P. carnosa isotype SAM-71804-0!); Alicetal bei Pomona, 13 Aug. 1929, Dinter 6591 (NBG-202184-0, PRE160652-0); E of Oranjemund, 34 km from the check point on the road to Sendelingsdrif, 25 Sept. 1997, Gess & Gess 97/98/64 (WIND!); 3 km (2 miles) SE of Lüderitz, 9 Aug. 1959, Giess & Van Vuuren 731 (PRE, WIND!); S of Lüderitzbucht, on the way to the lighthouse, 12 Sept. 1967, Giess 10196 (WIND!); About 40 km N of Rosh Pinah, Farm Witpütz Süd, 29 Sept. 1983, Goldblatt 7019 (PRE, WIND!); Namuskluft, 28 Oct. 1970, Jankowitz 12/190 (WIND!); Ai-Ais conservation area, McMillan's Pass, 2 km SE of summit of pass along track, 12 Oct. 2005, Kolberg & Tholkes HK1698 (K!, WIND!); Sperrgebiet. Lüderitz – Oranjemund road at N turn-off to Bogenfels, 14 Oct. 2006, Kolberg & Tholkes HK2103 (K!, WIND!); Tafelberg, SE of Klinghardts, Sperrgebiet, Aug. 1971, Logan & Jensen 899 (WIND!); Tafelberg, N of Affenrücken (coast), Sperrgebiet 1, Aug. 1971, Logan & Jensen 959 (WIND!); Just outside Lüderitz on road to Aus, 15 Aug. 2001, Loots SL114 (WIND!); Lüderitz – Oranjemund road, 27 Aug. 2002, Mannheimer CM2021 (WIND!); Namitsas, S of Klinghardt Basin, 1 Sept. 2002, Mannheimer 2110 (WIND!); Namitsas, S of Klinghardt Basin, 1 Sept. 2002, Mannheimer CM2122 (WIND!); Limestone-dolomite ridge, NE Klinghardts, 3 Sept. 2002, Mannheimer CM2175 (WIND!); Road to Grillental from Kaukausib, 5 Sept. 2002, Mannheimer CM2201 (WIND!); Blue ridges at Pomona pumphouse, 27 Sept. 2004, Mannheimer, Maggs-Kölling & Loots CM2646 (WIND!); Road between Grillental and Drachenberg, 9 Sept. 2005, Mannheimer CM2737 (WIND!); Nautilus, N of Lüderitz, 23 Aug. 1963, Merxmüller & Giess 3074 (M, WIND!); Farm Spitzkop, plain before Numais-bank, 1 Sept. 1963, Merxmüller & Giess 3412 (M, PRE, WIND!); Lüderitzbucht, 8 Sept. 1972, Merxmüller & Giess 28277 (M, PRE, WIND!); Buchuberge, 10 Sept. 1972, Merxmüller & Giess 28302 (M, PRE, WIND!); 23 km S of Grillental, 12 Sept. 1972, Merxmüller & Giess 28380 (M, PRE, WIND!); Klinghardt Mts, S part, in the region of Sargdeckel, 17 Sept. 1977, Merxmüller & Giess 32109 (M, PRE, WIND!); Klinghardt Mts, near foot of the mountain, 27 July 1977, Müller 699 (WIND!); Schakals Mts, 31 July 1977, Müller 756 (PRE, WIND!); Sperrgebiet, on main road between Lüderitz and Oranjemund, 14 Aug. 2001, Smook 11383 (PRE, WIND!); (Buchuberg) Found on NW slope, 28 Sept. 1992, Strohbach 218 (WIND!); Namuskluft, 22 Aug. 1989, Strohbach 458 (WIND!); McMillians Pass, 23 Aug. 1989, Strohbach 486 (WIND!); Rosh Pinah, Namuskluft 88, 31 Aug. 1989, Van Wyk 8822 (PRE, WIND!); Farm Zebrafontein 87, Mountains © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 near farmhouse on Spitskop 111, Feb. 1989, Van Wyk 8986 (PRE, WIND!); Diamond area no. 1, Obib Springs, where the stream exits from the mountain, 3 Sept. 1989, Van Wyk 9041 (PRE, WIND!); 30 km S of Auffenrücken, June 1993, Williamson 4617 (WIND!); 30 km S of Affenrücken, June 1993, Williamson 4618 (WIND!); Daberas gorge, 55 km NE of Oranjemund, 16 July 1993, Williamson 5035 (WIND!); 10 km S of Chameis gate, 110 km N of Oranjemund, 21 July 1993, Williamson 5075 (WIND!). SOUTH AFRICA. Western Cape Prov.: Cape of Good Hope, Thunberg s.n. (P. glabrata holotype UPS-THUNB; isotype S-G-5142)!); Kaus, R III, 1835, Drège s.n. [2778] (P. sesuviifolia lectotype G-DC no. G-00322348!); Olifant R. W (of “West”?), R I, 1835, Drège s.n. [?778 or 2778] (P. sesuviifolia syntypes G-DC no G-00322349!, P-027091!, P-27092!, P-027093 (ex hb. Hennecart)!); possible syntypes HAL-0110985!, HBG-505117!, HBG-503991!, NY-00232709!). HABITAT. In gravel or sand of plains and hill slopes in steppe dominated by succulents; mostly near the coast; 50 – 800 m. CONSERVATION STATUS. The Namibian distribution of this species falls almost entirely into the protected diamond mining area, which has now been declared a national park, limiting human access and activity. The effect of mining on Pteronia glabrata is minimal since mining is restricted to small areas and governed by a sound environmental management policy. The species is widely distributed (large estimated EOO and AOO) and no threats could be identiﬁed, thus resulting in a Namibian evaluation of LC (IUCN 2001, 2013). The South African threat status is listed as LC (SANBI 2011). PHENOLOGY. Flowering: February to October. Fruiting: June to November. NOTES. Among earlier collectors in Namibia there has been some confusion around this species and its difference (or not) to Pteronia succulenta. Material of P. succulenta, which occurs only in South Africa, must be inspected to verify if these are two separate species or not, but this falls outside the scope of this study. The syntypes of Pteronia sesuviifolia in G-DC are on one sheet: “Kaus, Drège, and Olifantrivier, Drège”, with both showing the number 2778. (On the Olifantrivier collection the label number is unclear and appears to have the 2 and the ﬁrst of the two number 7s superimposed, raising the impression of “778” only.) We have selected the Kaus material as lectotype because, though lacking (parts of) ﬂowers, the overall quality of the material is better. It should be noted that, here and elsewhere, after designating a lectotype the remainder of all original material (if any) is not to be named “paralectotype” or “lectoparatype” (whatever one prefers), but remain “syntype(s)” in accordance with Turland’s (2013: 62) interpretation — an opinion we have followed in all cases where we have chosen a lectotype among syntypes. KEW BULLETIN (2014) 69: 9488 There are syntypes of the Olifantrivier collection in P (P-027091!, P-27092!, P-027093 (ex hb. Hennecart)!)) and G-DC (G-00322349!). Potential syntypes have annotations, sometimes indicating effectively the same location but lack the number 2778 (unlike the P specimens) or lack the precise locality. We identiﬁed HAL-0110985, HBG-505117, HBG503991 and NY-00232709 in this respect. A Drège specimen at Vienna, W-0008793, has a locality "near Kaus" and what we assumed to be a collecting date, "1837-12" (Dec. 1837), but which is probably a herbarium dispatch or a receipt date. Similarly, W-0008794 shows “1837-12” and is from near the Ebenezar, Olifantrivier locality. We therefore exclude these specimens as syntypes. Also W-Rchb. 18890292322 is not exactly from any of the two locations in the protologue and should also, we believe, be excluded as original material — contrary to the annotations in W. 9488, Page 15 of 44 splitting on older stems. Leaves opposite, clustered along stems, densely silver-grey felty, linear to linearlanceolate, 4 – 10 × 2 – 2.5 mm, apex obtuse or subacute, base sessile; aromatic. Capitula numerous, densely covering plant, all along branches, narrowelongate ellipsoid to turbinate, 12 – 15 × 8 mm; apex acute in bud. Receptacle ﬁmbriate. Phyllaries multiseriate, outer broadly lanceolate, inner oblonglanceolate, yellow to straw-coloured; grey felty on outside; midrib pubescent, thickened towards apex but never tapering into a mucro; apex acute; margins narrow glabrescent, indistinctly ciliate. Florets c. 4; corollas to 10 mm long, yellow; lobes triangular, obtuse; tube widening upwards, ribbed at base. Achenes obovate, contracted into a distinct neck, villous in lower half, to 3 mm long. Pappus multiseriate, c. 10 mm long, straw-coloured. DISTRIBUTION. Africa: Botswana, Namibia, South Africa. Map 9. 9. Pteronia glauca Thunb. (Thunberg 1800: 144); de Candolle (1836: 365); Harvey in Harvey & Sonder (1865: 107); Hutchinson & Phillips (1917: 289); Range (1935: 275); Merxmüller (1955: 79; 1967: 156); Merxmüller & Roessler (1984: 90); Herman (2003: 277). Type: South Africa, Western Cape Prov., Cape of Good Hope, Thunberg s.n. (holotype UPS-THUNB; isotype LD-1254625!). Pteronia latisquama DC. (de Candolle 1836: 363). Type: South Africa, Northern Cape Prov., Zwischen (noted on label as “Zw:”) Camisberge und OnderBokkeveld, R V, 1835, Drège s.n. [607] (holotype GDC no. G-00322350!). — see Notes. Pteronia glabrata auct. non L. f.: de Candolle (1836: 362); Hutchinson & Phillips (1917: 289, noting “DC, not of L. f.”). Pteronia candollei Harv. (Harvey in Harvey & Sonder 1865: 105, “Candollei”). Type: South Africa, Northern Cape Prov., Fraserburg Div., between Karree R. and Quaggas Fontein, Burchell 1410 (holotype K00273422!). Pteronia thymifolia Muschl. & Dinter (Muschler 1911a: 100). Type: Namibia, Bezirk des Damaralandes, Farm Hoffnung, 1900 m, 20 Aug. 1909, Dinter 967 (holotype B†; isotypes K-00273419!, K-000273424!, SAM-71826-0). Pteronia arcuata Dinter (1932: 182). Type: Namibia, Aus, in der Mesembr.-Steppe am Wege nach Gubub [Kubub], 26 Oct. 1922, Dinter 4152 (holotype B†; isotypes HBG-505144!, PRE-156640-0, SAM71805-0, Z-000052352!, Z-000052353!, Z000003814!), synon. nov. Pteronia glauca Thunb. subsp. arcuata (Dinter) Merxm. (Merxmüller 1955: 79; 1967: 156). Shrub, to 80 cm tall and 120 cm in diam. Stems sometimes arching to ground, grey pubescent, bark SPECIMENS EXAMINED. NAMIBIA. Erongo Region, Omaruru Distr.: Brandberg, upper reaches of Königstein, 5 May 1993, Craven 4010 (WIND!); Valleys of upper Brandberg Mt, June 1955, Wiss 1427 (PRE, WIND!); Hardap Region, Maltahöhe Distr.: (Naukluft Park) 17 May 1979, Cronje 2 (WIND!); No 9. NaukluftPlateau, 9 April 1971, Meyer 491 966 (WIND!); Karas Region, Bethanie Distr.: Farm Gamochas BET 31, 6 Sept. 1972, Merxmüller & Giess 28257 (M, PRE, WIND!); Keetmanshoop Distr.: Farm Witmond, on rim of W plateau, 31 March 1998, Strohbach & Dauth 3826 (WIND!); Farm Rishon, on top of Graberberg at Telecom tower, 1 April 1998, Strohbach & Dauth 3835 (WIND!); Lüderitz Distr.: Aus, Mesembr.-Steppe along road to Gubub [Kubub], 26 Oct. 1922, Dinter 4152 (P. Map 9. Known distribution of Pteronia glauca in Namibia. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 16 of 44 arcuata isotypes HBG-505144!, PRE-156640-0, SAM71805-0, Z-000052352!, Z-000052353!, Z-000003814!); Farm Klein Aus, W of Aus, 11 Aug. 1959, Giess & Van Vuuren 768 (PRE, WIND!); Aus, Kubub, 4 Oct. 1959, Giess 2387 (PRE, WIND!); Farm Zebrafontein, track from road D463; at locked gate just before homestead, 20 Oct. 2007, Kolberg & Tholkes HK2410 (K!, WIND!); Sperrgebiet, 19 km S of Obib Fountain, track turns E and leads over hill, 24 Oct. 2008, Kolberg & Tholkes HK2694 (K!, WIND!); Farm Plateau LU 38, 8 Sept. 1963, Kräusel & Wiss 2015 (WIND!); (Aus) quartz foothills, 17 Sept. 2005, Mannheimer CM2822 (WIND!); Aus, in the river on the way to Helmeringhausen, 7 Aug. 1963, Merxmüller & Giess 2918 (M, PRE, WIND!); Klinghardt Mts, near top of the mountain, 28 July 1977, Müller 703 (PRE, WIND!); Farm Plateau, Aus, 13 April 1953, Walter & Walter 2555 (WIND!); Maltahöhe Distr.: Farm Chamchawib, Helmeringhausen, 15 Aug. 1963, Merxmüller & Giess 2812 (M, PRE, WIND!); Khomas Region, Windhoek Distr.: Windhoek, 1955, Basson 137 (WIND!); Khomas Hochland, 14 May 2000, Burke 00089 (WIND!); Farm Hoffnung, 20 Aug. 1909, Dinter 967 (P. thymifolia isotypes K-00273419!, K000273424!, SAM-71826-0); Lichtenstein, 15 Oct. 1934, Dinter 7892 (Z!); Windhoek, Augeigas, Daan Viljoen road, turn-off to Matchless Mine, ± 500 m from turn-off, 6 Aug. 2000, Friedrich FRI2 43 (WIND!); Farm Keres WIN 39, on the Kereshöhe, 6 Nov. 1965, Giess 9038 (PRE, WIND!); Moltkeblick, Auasberge, 29 Aug. 1971, Giess & Von Alvensleben 11493 (PRE, WIND!); Farm Hohenau, Aug. 1963, Kräusel & Wiss 1994 (WIND!); Farm Harris, 19 July 1965, Leach, Bayliss & Giess 12938 (PRE, WIND!); Farm Regenstein, below SW mountain foot towards plain, 22 Aug. 1972, Merxmüller & Giess 28008 (M, PRE, WIND!); Farm Regenstein, near Windhoek, 12 April 1953, Schwerdtfeger 2/294 (WIND!); Farm Bellerode, 13 km (8 miles) from Windhoek, 17 March 1965, Toelken & Hardy 712 (PRE, WIND!); Farm Immental, Von Koenen s.n. (WIND-34158!); Farm Voigtland, Okamatuja, 17 Nov. 1952, Walter & Walter 118 (WIND!). SOUTH AFRICA. Western Cape Prov.: Boschjemanskarroo [III A b 1], Drège s.n. (HAL0111565!, HBG-505116!, HBG-503989!, P-027102!, P027103!, P-00179775!, PRE, W-0008795); Aasvogelberg [II e 11], Drège s.n. (HAL-0111564!, K-000273420!, P00179776!, P-027104!, P-027101!, P-027100!, W0008796!); Fraserburg Div., between Karree R. and Quaggas Fontein, Burchell 1410 (P. candollei holotype K00273422!); Cape of Good Hope, Thunberg s.n. (P. glauca holotype UPS-THUNB; isotype LD-1254625!); Northern Cape Prov.: between Camisberge and OnderBokkeveld, R V, 1835, Drège s.n. [607] (P. latisquama holotype G-DC no. G-00322350!). HABITAT. Rocky hills and mountains in high-altitude shrubland, dwarf shrub savanna, desert-dwarf shrub transition and steppe dominated by succulent species; 450 – 1900 m. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 CONSERVATION STATUS. This species is evaluated LC in Namibia with a very large estimated EOO and AOO that puts it outside the thresholds for threatened species (IUCN 2013). In addition, no threats causing population decline could be identiﬁed. The threat status for South Africa is also listed as Least Concern (LC) (SANBI 2011). PHENOLOGY. Flowering: May to November. Fruiting: September to December. VERNACULAR NAMES. Boegoekaroo, geelboegoekaroo, perdekaroo (Afrikaans, South Africa). NOTES. This species is restricted to higher altitudes or mountains in Namibia. Plants with arching branches found near Aus were described as Pteronia arcuata Dinter, later considered a subspecies of P. glauca (Merxmüller 1955: 79). The arching branches, however, are not consistently displayed. Plants with arching and non-arching branches have been found by the ﬁrst author in the same population. The stem habit seems to be determined by the plant’s environment and arching branches occur mostly on plants in hard, rocky soil, such as the specimens near Aus. For this reason subsp. arcuata is not upheld here. An enumeration of localities and species collected by J. F. Drège in southern Africa (mainly in presentday South Africa) was published in 1843 (Drège 1843) with an introduction (on pp. 3 – 43) by E. Meyer. It is structured along an elaborate site coding system, using numerals and letters such as I, II, III, A, B, a, b, 1, 2, etc., a combination of which is often found on what have been or are being designated (iso) types of, in our case, Pteronia species. For example “III.E.a” in association with P. sesuviifolia (q.v.) is listed by Drège in 1843 on p. 108. Although this works well with most species as a precise source of additional locality data it fails with P. latisquama where none of the potential isotypes is directly linked to (part of) the “Camisberge, Onder-Bokkeveld” of the holotype. One also needs to keep in mind that this enumeration was published six years after de Candolle’s 1836 treatment of the genus in the ﬁfth volume of his Prodromus — and thus that Candolle’s locality names could easily have been included in the Drège listing. That localities in the protologue by de Candolle do not match any in Drège/Meyer was already annotated in association with specimens in W (0008795 and -8796; see below) by L. Pignotti. We found two sets of specimens with the name P. latisquama, but have been unable to link any of them to the holotype locality. One set of specimens carries: (1) Boschjemanskarroo (the locality “III A b 1” in Drège 1843: 69), found in HAL-0111565!, HBG-505116!, HBG-503989!, P-027102!, P-027103!, P00179775!, PRE, and W-0008795!, sometimes with the (presumed) de Candolle number [607] added. The name is also on K-000273421!, but we exclude this as “1837” is cited, which is possibly a herbarium despatch or receipt date. Of all these specimens P-027103 seems KEW BULLETIN (2014) 69: 9488 9488, Page 17 of 44 the most authentic as we think it shows an original Drege label with his handwriting and with “III A b 1” cited. The other carries: (2) Aasvogelberg (the locality II.e.11 in Drège 1843: 65), and is present in HAL-0111564!, K-000273420!, P-00179776!, P027104!, P-027101!, P-027100!, W-0008796!. 10. Pteronia inflexa L. f. (Linnaeus ﬁlius 1782: 356); Thunberg (1800: 144); de Candolle (1836: 365); Harvey in Harvey & Sonder (1865: 99); Hutchinson & Phillips (1917: 300); Merxmüller (1967: 156); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: South Africa, Western Cape Prov., Cape of Good Hope, Thunberg s.n. (holotype UPS-THUNB; isotype LD-1254505!). Pteronia lupulina DC. (de Candolle 1836: 357). Type: South Africa, Northern Cape Prov., Fraserburg, between Zack R. and Kopjes Fontein, 4 Sept. 1811, Burchell 1495 (holotype G-DC; isotypes K-00273447!, P-027188!, PRE157943-0, W-0008801!). Pteronia lupulina DC. var. rotundifolia DC. (de Candolle 1836: 358). Type: South Africa, Western Cape Prov., Nieuweveld, R IV, 1835, Drège s.n. [767] (holotype GDC no. G-00322347!; isotypes may exist — see Notes). Shrub, to 30 cm tall. Stems sparingly pubescent, bark pale grey to grey, splitting on old growth. Leaves opposite, coriaceous, surface densely short bristly and sessile glandular, ﬂat, elliptic to narrowly ovate, 5 – 15 × 4 mm; apex rounded; base sessile; margins pectinate. Capitula solitary on short branchlets, ovoid-campanulate, 15 – 20 × 6 – 10 mm; apex rounded to truncate in bud. Receptacle honeycombed. Phyllaries multiseriate, ovate to orbicular, to 5 × 3 mm, green to brown, glabrous except for a few glands on outside; apex rounded; margins distinct, translucent, membranous. Florets c. 10; corollas to 10 mm long, glabrous, yellow; lobes lanceolate, subacute; tube gradually widening. Achenes obconical, sparsely villous, to 4 mm long. Pappus c. 10 mm long, straw-coloured. DISTRIBUTION. Africa: Namibia, South Africa. Map 10. SPECIMENS EXAMINED. NAMIBIA. Hardap Region, Maltahöhe Distr.: Farm Naudaus/Duwisib MAL 76/ 84, 10 May 1956, Volk 12771 (WIND!); Farm Naudaus/ Duwisib 76/84, 3 May 1956, Volk 12772 (WIND!); Karas Region, Bethanie Distr.: Farm Goais BET 13, 23 June 1974, Giess 13353 (PRE, WIND!); Keetmanshoop Distr.: Farm Witmond 162A, NW of farmhouse on sloping plain, 14 Feb. 1997, Strohbach, Kubirske & Sheuyange 2872 (WIND!); Lüderitz Distr.: Farm Plateau, 20 Aug. 1963, Merxmüller & Giess 3012 (M, PRE, WIND!). SOUTH AFRICA. Western Cape Prov.: Cape of Good Hope, Thunberg s.n. (P. inﬂexa holotype UPSTHUNB; isotype LD-1254505!); Fraserburg, between Map 10. Known distribution of Pteronia inﬂexa in Namibia. Zack R. and Kopjes Fontein, 4 Sept. 1811, Burchell 1495 (P. lupulina holotype G-DC; isotypes K-00273447!, P027188!, PRE-157943-0, W-0008801!); Nieuweveld, R IV, 1835, Drège s.n. [767] (P. lupulina var. rotundifolia holotype G-DC no. G-00322347!); between Waschbank and Rietpoort, Nieuweveld, 3500 (ft.), (loc.) I d, 26 Nov. 1826, Drège s.n. (P-027186!). HABITAT. Gravelly plains and slopes of dwarf shrub savanna and desert. None of the Namibian specimens examined present any altitudinal data. CONSERVATION STATUS. The uncertain identity of Namibian specimens (see Notes) together with insufﬁcient information on these led us to conclude that the status Data Deﬁcient (DD) must be assigned for Namibia (IUCN 2001, 2013). Least Concern (LC) is recorded as the South African threat status (SANBI 2011). PHENOLOGY. Flowering: February to August. NOTES. According to Hutchinson & Phillips (1917) this species belongs to what they called sect. Ciliatae with ciliate leaf margins. The leaves of Namibian collections are, however, papillate-glandular and do not have ciliate margins, which would put the species into their sect. Papillatae. In Hutchinson & Phillips’ (1917) description of Pteronia inﬂexa there is also no mention of ciliate or pectinate leaf margins. P. inﬂexa has rarely been recorded in Namibia and the distribution in south-western and south-eastern Namibia is disjunct from its distribution in the Karoo of South Africa. More Namibian material must be collected and compared with South African specimens to verify if P. inﬂexa does indeed occur in Namibia or if these specimens belong to another species. Possible isotypes of Pteronia lupulina var. rotundifolia may exist: relevant specimens in P (e.g. P-027186! and © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 18 of 44 027187!) and W (e.g. W-Rchb.-1889-0292329! and W0020460) all show the handwritten annotation of the full name, but do not show the year or associated number 767. 11. Pteronia leucoclada Turcz. (Turczaninow 1851: 65); Harvey in Harvey & Sonder (1865: 105); Hutchinson & Phillips (1917: 314); Merxmüller (1967: 156); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: South Africa, Western Cape Prov., Cape of Good Hope, Zeyher 811 (holotype KW-001000912!; isotypes K000273482!, K-000273483!, K-000273484!, P-027205!, P027206!, PRE-0159187-0!, PRE-0587284-0!, SAM-00377380!, W-Rchb.-1889-0278261!, Z-000052407!). Shrub, to 50 cm tall. Stems smooth, glabrous, white when young. Leaves alternate, clustered along stem, ﬂeshy, glabrous, oblong to narrowly obovate-clavate, to 7 × 1.5 mm; apex rounded; narrowed into base. Capitula solitary, terminal, obconic to almost spherical, to 15 × 10 mm, apex rounded in bud. Receptacle honeycombed, ﬁmbriate. Phyllaries multiseriate, lanceolate to linear, keeled, 2 – 3 mm long, inner phyllaries to 2.5 mm broad, glabrous; apex acuminate; margins membranous. Florets c. 20; corollas to 5.5 mm long, yellow; lobes unequal, triangular-lanceolate, subacute. Achenes compressed turbinate, densely villous, to 2 mm long. Pappus multiseriate, sparsely barbellate, to 6 mm long, golden. DISTRIBUTION. Africa: Namibia, South Africa. Map 11. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Karasburg Distr.: Farm Vrede, 19 July 2005, Bruyns 10107 (BOL, WIND!); Klein Karas, 6 Aug. 1923, Dinter 4849 (Z!); Farm Klein-Aub WAR 52, 17 May 1963, Map 11. Known distribution of Pteronia leucoclada in Namibia. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 Giess, Volk & Bleissner 7022 (PRE, WIND!); Farm Udabis WAR 77, 19 May 1963, Giess, Volk & Bleissner 7108 (WIND!); Near border between Huniam Ost and Uheib 84, 19 April 1997, Strohbach 3429 (WIND!); Keetmanshoop Distr.: Klein Karas Mts, 6 March 1999, Burke 99149 (WIND!). SOUTH AFRICA. Western Cape Prov.: Cape of Good Hope, Zeyher 811 (holotype KW001000912!; isotypes K-000273482!, K-000273483!, K000273484!, P-027205!, P-027206!, PRE-0159187-0!, PRE-0587284-0!, SAM-0037738-0!, W-Rchb.-18890278261!, Z-000052407!) HABITAT. Dwarf shrubland; c. 950 m. CONSERVATION STATUS. This species has a relatively restricted range in SE Namibia and present information suggests that it is rare here. The estimated EOO would qualify it for VU status but no two of the additional three criteria under criterion B can be satisﬁed (IUCN 2013). For criterion D2 the number of localities is restricted but no additionally required plausible future threat could be identiﬁed to qualify Pteronia leucoclada as VU. The Namibian conservation status is thus assessed here as LC (IUCN 2001). The South African threat status is also LC (SANBI 2011). PHENOLOGY. Flowering: March to July. Fruiting: April. V E R N A C U L A R N A M E S . Bleekbossie, witbossie, witbasbossie (Afrikaans, South Africa). NOTES. Thus far only a few collections have been made from the extreme south-east of Namibia. The greater part of the species’ range falls into adjacent areas in South Africa. Although Turczaninow cites “Zeyher coll. n. 811.” it is often impossible to separate Zeyher from “Ecklon & Zeyher” collections — indeed the isotype in Z carries a label to that extent and does not indicate who of the two actually collected no. 811. (In contrast, K, KW, SAM, P-027206, PRE and W-Rchb. all have “Zeyher 811” as the sole collector.) Ecklon & Zeyher’s collections from “S. Africa” (Lanjouw & Staﬂeu 1957: 176) are in many herbaria. They are not, however, reported from KW where Turczaninow’s types are, and thus the type of leucoclada from KW presents a rare exception. We follow Staﬂeu & Cowan (1986: 538) on the location of the type, which in this case also carries “veriﬁed by Turczaninow”. As with other Pteronia species (see above), the various isotypes provide more locality details than the holotype: the SAM and W-Rchb. collections have “Bitterfontein, Bosmansland” on the label. Isotype P027205 has the Zeyher 811 “communicated” by Drège, and with the locality described as: “Clanwilliam. A Tulbaghskloof ad Pikenierskloof. 500′ – 1000′. Martii.”. However, this annotation is certainly not in Zeyher’s handwriting. Furthermore, P-027206 is the only isotype with a year of collecting: 1847. Though the total of these isotypes provides much more detail, they are not necessarily Zeyher’s notes. KEW BULLETIN (2014) 69: 9488 12. Pteronia lucilioides DC. (de Candolle 1836: 358); Harvey in Harvey & Sonder (1865: 100); Hutchinson & Phillips (1917: 293); Dinter (1926: 132); Range (1935: 275); Merxmüller (1952: 126; 1967: 156); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: South Africa, Northern Cape Prov., Klein Namaqualand, R. II, Drège s.n. [2779] (holotype G-DC; isotype P027210!). Pteronia lucilioides DC. var. β sparsifolia Harv. (Harvey in Harvey & Sonder 1865: 100). Type: South Africa, Namaqualand, Wyley s.n. (not traced). Pteronia gymnocline auct. non DC.: (Meyer in) Drège (1843: 94, 214); Range (1935: 275); Hutchinson & Phillips (1917: 294). — see Notes. Pteronia bromoides S. Moore (1904: 1011). Type: Namibia, Gross Namaland, Jakalskopje, Dinter 1197 (holotype BM-000903815!). Pteronia beckeoides auct. non DC.: Dinter (1926: 132); Range (1935: 275, “beckioides”); Merxmüller (1955: 80). Pteronia roesemanniana Dinter ex Merxm. (Merxmüller 1952: 128; 1955: 80), nom. nud. — see Notes. Shrub to 2.5 m tall, virgate, sparsely branched; near coast only to 30 cm high and much-branched. Stems scabrid-puberulous, glabrescent, grey. Leaves opposite, clustered on stems, matte grey-green, surface with acute conical, multicellular papillae interspersed with sessile, spherical glands, linear to obovate or oblongspathulate, boat-shaped, 5 – 8 × 1.5 mm; apex obtuse; base sessile, not connate with opposite leaf. Capitula solitary, terminal, cylindrical, 15 – 20 × 5 – 10 mm; apex sharply acute in bud. Receptacle deeply honeycombed. Phyllaries multiseriate, oblong-oblanceolate to linear-oblong, 6 –18 × 3 – 5 mm, pale yellow, glabrous except for ﬁnely puberulous outer midrib; apex rounded to subacute to cuspidate; margins not distinctly membranous. Florets c. 12; corollas to 10 mm long, yellow; lobes lanceolate, subacute; tube gradually widening. Achenes obovoidturbinate, densely glandular and sparsely setose, to 6 mm long. Pappus multiseriate, barbellate, to 10 mm long, straw-coloured. Figs 4 – 5. DISTRIBUTION. Africa: Namibia, South Africa. Map 12. SPECIMENS EXAMINED. NAMIBIA. Erongo Region, Karibib Distr.: Okongava, Kalkberge, 2 Feb. 1934, Dinter 6933 (K-000273436!, PRE, WIND!); Farm Okongawa, Marmorberg, 9 May 1958, Seydel 1553 (WIND!); Hardap Region, Maltahöhe Distr.: Losberg, path to the repeater, very close to the top and on highest plateau, 1 July 2004, Clapham & Drayer 145 (WIND!); Mountain Zebra Park, Naukluft MAL 9, 2 June 1968, Giess 10445 (PRE, WIND!); Found above the Hudup R., some 1.6 km N of Maltahöhe, 5 Sept. 1972, Merxmüller & Giess 28222 (M, PRE, WIND!); Farm Naudaus/Duwisib, Helmeringhausen, 20 May 9488, Page 19 of 44 1956, Volk 12496 (WIND!); Near Farm Maguams, slopes of Schwarzrand, 22 March 1953, Walter & Walter 2115 (WIND!); Karas Region, Bethanie Distr.: 38 km (23.5 miles) N of Helmeringhausen, 19 Oct. 1949, Acocks 15631 (PRE, WIND!); Farm Mara, kloof on E side of Konkiep near house, 11 July 1988, Craven 3310 (WIND!); Farm Mara, O'Connell's prospecting kloof, in Hunsberg Mts, above middle windpump, 24 Sept. 1989, Craven 3521 (WIND!); Farm Mara E of Hunsberg Mts, In kloof of mountain, W of farm house, 23 June 1991, Craven 3862 (WIND!); Farm Mooifontein BET 50, 19 May 1965, Giess 8817 (PRE, WIND!); Farm Heigums BET 105, 16 June 1976, Giess & Müller 14433 (WIND!); Farm Chamchawib, Helmeringhausen, 15 Aug. 1963, Merxmüller & Giess 2813 (M, PRE, WIND!); Farm Kosos, 15 Aug. 1963, Merxmüller & Giess 2819 (M, PRE, WIND!); Farm Saraus BET 16, 5 Sept. 1972, Merxmüller & Giess 28226 (M, PRE, WIND!); Farm Aruab 23, near Aris Post, 5 April 1998, Miller MIL1/ 55 (WIND!); Karasburg Distr.: Farm Kromrivier WAR 359, May 1963, Giess, Volk & Bleissner 6997 (WIND!); Farm Sperlingspüts WAR 259, 6 Aug. 1976, Giess 14516 (WIND!); Spes Bona, 31 Aug. 1970, Jankowitz 81/1493 (WIND!); Karas Mt on route to the Telecom tower on farm Rishon, 14 Feb. 1997, Strohbach 2854 (WIND!); Goodhouse Poort, 29 Aug. 1989, Van Wyk 8711 (PRE, WIND!); Keetmanshoop Distr.: 56 km (35 miles) SW of Narubis, 30 April 1955, Acocks 18045 (PRE, WIND!); Farm Kochena KEE 74, in the mts N of the farmhouse, 12 May 1972, Giess & Müller 11895 (PRE, WIND!); Farm Pieterskloof (Kraaikluft) KEE 370, Mt. N of Schroffenstein, 14 May 1972, Giess & Müller 11926 (PRE, WIND!); Lüderitz Distr.: Diamond Area 1, Drachenberg, 9 Sept. 2005, Bartsch, Mannheimer & Kwembeya SB2048 (WIND!); Rosh Pinah. Nooitgedacht, 10 Aug. 2000, Bruyns 8330 (BOL, WIND!); Dik Willem, 29 April 2000, Burke 00076 (WIND!); SE of Aurus Mts, Sperrgebiet, 24 Sept. 1999, Burke 99212 (WIND!); SE of Aurus Mts, Sperrgebiet, 24 Sept. 1999, Burke 99215 (WIND!); Skorpion turn-off D716 c. 5 km N of Rosh Pinah, Bührmann BUH1 76 (WIND!); Diamond area no. 1, Grillental, marble hills, 6 Sept. 1958, De Winter & Giess 6228 (PRE, WIND!); Farm Namuskluft LU 88, c. 29 km (18 miles) S on road to Lorelei, 13 Sept. 1958, De Winter & Giess 6337 (NBG, PRE, SAM, WIND!); Great Namaland, Jakalskopje, Dinter 1197 (P. bromoides holotype BM-000903815!); Kahanstal, 9 Dec. 1934, Dinter 8192 (K!); Hunsberge; in the Nuob R., 26 Sept. 1976, Giess & Wendt 14714 (WIND!); Farm Witputz Nord LUS 22, June 1976, Giess & Wendt 14783 (WIND!); About 40 km N of Rosh Pinah, Farm Sud Witputs. Granitic outcroppings NE of farm houses, 29 Sept. 1983, Goldblatt 7015 (PRE, WIND!); N of Witputz, 2 Oct. 1978, Hardy & Venter 4863 (PRE, WIND!); Sperrgebiet, N Klinghardt Mts, 6 Aug. 2001, Klaassen & Bartsch EK507 (WIND!); C13 on way to Rosh Pinah © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 20 of 44 Fig. 4. The typically acute budding capitula of Pteronia lucilioides. from Aus, 8 Aug. 2001, Klaassen & Bartsch EK535 (WIND!); Sperrgebiet, 50 km from Rotkop Station, powerline track, 23 Oct. 1987, Kolberg & Maggs HK210 (WIND!); Farm Klein Aus Vista, Geister Trail from cabin in Geisterschlucht, rocky hill slopes to SW, 4 Oct. 2006, Kolberg & Tholkes HK2049 (K!, WIND!); Namuskluft LU 88. Witputz, 13 Sept. 1963, Kräusel & Wiss 2074 (WIND!); White quartz outcrop, Sperrgebiet, 3 Sept. 1992, Kubirske, Strohbach & Swart 18 (WIND!); NW corner of the farm Namuskluft, 3 Feb. 1981, Lavranos, Barad & Pehlemann 19226 (WIND!); Namuskluft, July 1971, Logan 414/1493 (WIND!); Namuskluft, July 1971, Logan 414 (WIND!); Between Klinghardts and Aurus towards Heioab, 12 Aug. 2001, Loots SL108 (WIND!); S of Tsaus (Sperrgebiet), 26 Sept. 1996, Mannheimer & Mannheimer 324 (WIND!); S of Tsaus (Sperrgebiet), 26 Sept. 1996, Mannheimer & Mannheimer 325 (WIND!); Rocky area next to borehole, near Aurus Mts, 26 Sept. 1996, Mannheimer & Mannheimer 329 (WIND!); On kopje near access to Aurus basin, 28 Sept. 1996, Mannheimer & Mannheimer 383 (WIND!); Aurus fountain on W-face, 9 Aug. 2001, Mannheimer CM1545 (WIND!); N of Dik Willem, 31 July 2002, © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 PHOTO: H. KOLBERG. Mannheimer 1933 (WIND!); Boegoeberg, N end of ridge, 29 Aug. 2002, Mannheimer 2061 (WIND!); Namitsas, S of Klinghardt Basin, 1 Sept. 2002, Mannheimer CM2123 (WIND!); (Sperrgebiet), 3 Sept. 2002, Mannheimer CM2173 (WIND!); Road from Grillental to Kaukausib, 5 Sept. 2002, Mannheimer CM2206 (WIND!); Namuskluft farm, just SW of MacMillans pass, 9 Sept. 2002, Mannheimer CM2221 (WIND!); Valley adjacent to Kahanstal, 12 km SE of Rosh Pinah on road to Loreley, 21 Sept. 1972, Merxmüller & Giess 28643 (M, PRE, WIND!); 9.5 km S of Grillental, 15 Sept. 1977, Merxmüller & Giess 32021 (M, PRE, WIND!); Farm Spitzkop LUS 111, 25 Sept. 1977, Merxmüller & Giess 32310 (M, WIND!); Numaeis South, Witputz, 1957, Rusch 4720 (WIND!); Sperrgebiet, between Aurus and Klinghardt Mts, 12 Aug. 2001, Smook 11347 (PRE, WIND!); Diamond area no. 1, Obib Mt. peak, 3 Sept. 1989, Van Wyk 9014 (PRE, WIND); Huns Mts, 24 km up Letterklip R., 26 Sept. 1983, Walter 159 (WIND); Aus, Farm Plateau, on the slopes of plateau, N of the farmhouse, 13 April 1953, Walter & Walter 2575 (WIND!); Diamond Area 1, Tsaus Mt. Oct. 1977, Wendt B/1 (WIND!); Diamond area 1, Tsaus Spinnenberg, Oct. 1977, Wendt 8/2 (WIND!); KEW BULLETIN (2014) 69: 9488 9488, Page 21 of 44 Fig. 5. Pteronia lucilioides. A habit; B leaf, abaxial; C capitulum; D phyllaries; E ﬂoret, side view; F ﬂoret, opened; G achene and pappus; H achene; J pappus setae of mixed length. A – B, E Kolberg & Tholkes HK2049; C – D, F – J Dinter 8192. DRAWN BY JULIET BEENTJE. Narudous Poort (Anusi 73), between Aus and Rosh Pinah, 30 Aug. 1989, Wittneben 8771 (PRE, WIND!); Uitsig, 40 km from Witputz, 1 Sept. 1989, Zietsman 1892 (WIND!). SOUTH AFRICA. Northern Cape Prov: Klein Namaqualand, R. II, Drège s.n. [2779] (P. lucilioides holotype G-DC, isotype P-027210!). HABITAT. Slopes, gorges, dry rivers and washes in dwarf shrub savanna, dwarf shrubland, desert-dwarf shrub © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 22 of 44 KEW BULLETIN (2014) 69: 9488 in the Northern and Western Cape Provinces of South Africa. Images of South African material were seen and it seems highly likely that P. lucilioides and P. gymnocline are conspeciﬁc, but this cannot be conﬁrmed without seeing more material. The variability of leaf indumentum density in Namibia also points towards this assumption. The sharply acute immature capitula are a constant character in all Namibian and South African material seen of P. lucilioides (Figs 4 & 5). Hutchinson & Phillips (1917: 294) retained P. gymnocline DC. while they listed P. gymnocline E. Mey. in Drege ("not of DC.") as synonym of P. lucilioides together with P. bromoides S. Moore and P. lucilioides var. sparsifolia Harv. Map 12. Known distribution of Pteronia lucilioides in Namibia. transition and steppe dominated by succulent species; 300 – 2000 m. CONSERVATION STATUS. This common and very widespread species has an estimated EOO and AOO well above the threshold for any of the threatened categories and no threats causing decline, fragmentation or ﬂuctuation of populations could be identiﬁed (IUCN 2013). The Namibian status thus is LC (IUCN 2001). The South African threat status is listed as LC too (SANBI 2011). PHENOLOGY. Flowering: February to October. Fruiting: June to December. NOTES. The species is near-endemic to Namibia with the greater part of its distribution extending from the Karibib area southwards to the Orange river with a few localities south of this river in South Africa. The specimens from the Karibib district (Dinter 6933 and Seydel 1553) to the best of current knowledge must be placed with Pteronia lucilioides, although this locality is disjunct from the rest of the species’ range. The collection Dinter 6933 in K carries the original (Dinter) identiﬁcation of Pteronia roesemanniana, a Dinter manuscript name, which was re-determined by Merxmüller as P. lucilioides. The leaves of Pteronia lucilioides are densely papillose and the capitula typically sharply acute in bud. In its range in southern Namibia P. lucilioides exhibits considerable variability in growth form: nearer the coast plants are low, much-branched shrublets, while further inland they are mostly virgate, sparsely branched shrubs which may be over 2 m tall. The density of leaf indumentum also varies but leaves of Namibian material were never completely glabrous. Glabrous leaves are the main character that distinguishes this species from P. gymnocline DC. (de Candolle 1836: 359). The latter is reported to occur © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 13. Pteronia mucronata DC. (de Candolle 1836: 362); Harvey in Harvey & Sonder (1865: 106); Hutchinson & Phillips (1917: 308); Merxmüller (1955: 80; 1967: 157); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: South Africa, Eastern Cape Prov., ZwartRuggens, Drège s.n. [2158] (holotype G-DC; isotype P027224!; other isotypes may exist — see Notes). Pteronia dinteri S. Moore (1904: 1012); Dinter (1926: 132, “Dinteri”); Range (1935: 275, “Dinteri”). Type: Namibia, Hereroland, Südrand der Etosapfanne [Etoschapfanne], 23 July 1899, Dinter 739 (holotype BM-001114680!; isotype Z-000052414!). Pteronia mucronata DC. subsp. dinteri (S. Moore) Merxm. (Merxmüller 1955: 80). Shrub to 70 cm tall. Stems much-branched, short, glabrous, grey to black, striate. Leaves opposite, clustered along stems, dark green, surface glabrous or scattered white bristly, linear, boat-shaped or somewhat keeled, to 8 × 1 mm; apex obtuse; base connate; margins white ciliate; aromatic. Capitula solitary, terminal, narrowly cylindrical, 12 – 20 × 5 – 7 mm; apex rounded in bud; base acute. Receptacle honeycombed. Phyllaries multiseriate, ovate to oblong-linear, 3 – 10 mm long, bright yellow, shiny, glabrous; midrib thickened towards apex and forming a small, recurved mucro; apex truncate, slightly emarginate; margins jagged, ciliate. Florets c. 12 – 15; corollas to 10 mm long, sparsely glandular, yellow; lobes lanceolate, subobtuse. Achenes obconical, densely villous, to 4 mm long. Pappus c. 10 mm long, yellow. DISTRIBUTION. Africa: Namibia, South Africa. Map 13. SPECIMENS EXAMINED. NAMIBIA. Hardap Region, Maltahöhe Distr.: Naukluft. Kapokvlakte on plateau, 1 Oct. 1995, Bridgeford 95359 (WIND!); Bergzebrapark Naukluft MAL 9, 2 June 1968, Giess 10441 (WIND!); Bergzebrapark Naukluft MAL 9, NW gorge, 3 Sept. 1972, Merxmüller & Giess 28191 (M, WIND!); Farm Rooiberg-Süd, 1935, Steyn 9960 (WIND!); Rehoboth Distr.: 23 km W along road D1261 from junction on road C24 (NE of Nauchas), 28 Aug. 2009, Kolberg & Tholkes HK2823 (K!, WIND!); Farm Narais North 245, KEW BULLETIN (2014) 69: 9488 Map 13. Known distribution of Pteronia mucronata in Namibia. 14 June 2004, Wittneben W04 248 (WIND!); Karas Region, Bethanie Distr.: Farm Soutkuil, stock camp before turning off to road to Bobbejaankranz, 22 Sept. 1989, Craven 3472 (WIND!); Fish R. Canyon, 10 Aug. 1976, Giess 14581 (PRE, WIND!); Fish R. Canyon, 10 Aug. 1976, Giess 14583 (WIND!); Farm Aruab 23, area around Aris Post, 5 April 1998, Miller MIL1/ 49 (WIND!); Farm Aruab 23, S part of farm, April 1998, Miller MIL1/ 088 (WIND!); Karasburg Distr.: Farm Mooirivier, 1 Sept. 1997, Strohbach, Kubirske & Sheuyange 2972 (WIND!); Keetmanshoop Distr.: Farm Dassiefontein 87, E camp, 29 March 1998, Strohbach & Dauth 3780 (WIND!); Lüderitz Distr.: Namuskluft, Rosh Pinah, Aug. 2003, Bruyns 9847 (BOL, WIND!); Farm Kolke LU 84, 4 Oct. 1975, Giess 13831 (PRE, WIND!); Farm Piet-se-Puts LUS 77. Black limestone gorge at the gate of farm Kanies LUS 71, Sept. 1976, Giess & Wendt 14748 (WIND!); Farm Namuskluft 88, 20 Sept. 2003, Klaassen, Bartsch & Loots EK1204 (WIND!); Pockenbank, 15 – 20 km E of main road against N boundary to Kokerboomkloof, 30 Oct. 2000, Loots SL20 (WIND!); Farm Witpütz South, Aug. 1963, Merxmüller & Giess 3213 (M, PRE, WIND!); Diamond Area 1, centre of Tsausberg, Oct. 1977, Wendt 3/4 (WIND!); Diamond Area 1, Tsaus Spinnenberg, Oct. 1977, Wendt 15/2 (WIND!); Khomas Region, Windhoek Distr.: Weener farm, Gamsberg, on E side, Craven 5111 (WIND!); Gurumanas REH 241, 29 Aug. 1972, Merxmüller & Giess 28073 (M, WIND!); Weissenfels, 30 Aug. 1972, Merxmüller & Giess 28110 (M, PRE, WIND!); Farm Naos, 15 July 1953, Schwerdtfeger 4350 (WIND!); Kunene Region, Outjo Distr.: S edge of Etosha Pan, 23 July 1899, Dinter 739 (P. dinteri holotype BM-001114680!; isotype Z000052414!). SOUTH AFRICA. Eastern Cape Prov.: Jansenville Div., Zwart-Ruggens, Aug. 1827, Drège s.n. 9488, Page 23 of 44 [2158] (P. mucronata holotype G-DC; isotype P027224!; potential isotypes HAL-0110982!, HBG505099!, HBG-503990!, K-000273457!, P-027225!, P027226!, P-027227!, PRE, SAM-0016168-0!, TUB005029!, W-0020461!, W-0008803!, W-Rchb.-18990219753!). HABITAT. Mountains, hills and rocky outcrops of dolomite or calcrete in dwarf shrub savanna, desertdwarf shrub transition, desert and steppe dominated by succulent species; 1200 – 1800 m. CONSERVATION STATUS. Estimated EOO and AOO for Pteronia mucronata in Namibia are above the maxima that would qualify it for a threatened category (IUCN 2013). In addition, no threats that would cause qualiﬁcation under any of the ﬁve IUCN criteria were established (IUCN 2013). The conservation status in Namibia is therefore LC (IUCN 2001). The South African threat status is recorded as LC (SANBI 2011). PHENOLOGY. Flowering: June to October. Fruiting: October to March. VERNACULAR NAMES. Kersbossie (Afrikaans, South Africa). NOTES. The consistently ciliate leaf margins or keels best distinguish Pteronia mucronata from P. unguiculata, in addition to the fewer and smaller ﬂorets, achenes and shorter pappus in the latter. Specimens with sparsely ciliate margins can be mis-identiﬁed as P. unguiculata also because the distribution areas overlap. In Namibia P. mucronata is conﬁned to higher altitudes of the south-west and the central highlands. However, the type specimen of P. dinteri, Dinter 739, even though its locality on the “southern edge of the Etosha pan” covers a large area (and was therefore not indicated on Map 13), does not conform to this. This area in the central north of Namibia is only around 1100 m above sea level without any high mountains (1200 – 1800 m high mountains is more typical of the mucronata distribution) although it has calcareous and/or saline soils. The vegetation on the fringes of the Etosha pan consists of dwarf shrubs, which is, together with the soils, comparable to the habitat of P. mucronata. The closest records of P. mucronata (Map 13) are in the central highlands, approximately 500 km due south of the Etosha pan; on the other hand a disjunct locality of P. unguiculata (see distribution on Map 23) at the Brandberg is only about 250 km SW thereof. Dinter 739 has clearly pectinate leaves though, which place it into P. mucronata even though P. unguiculata would make more sense from a phytogeographical perspective. Because of differences in number of ﬂorets (12 – 15 in P. mucronata; c. 5 in P. dinteri), capitula length (P. mucronata: 15 – 20 mm; P. dinteri: 10 – 14 mm) and pappus length (P. mucronata: 10 mm; P. dinteri: 7 mm), Merxmüller (1955) did not agree with the sinking of P. dinteri into P. mucronata by Hutchinson & Phillips (1917: 308) and kept it as a subspecies, which would support the odd distribution of this specimen. He later © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 24 of 44 (Merxmüller 1967), however, included both P. dinteri and P. mucronata subsp. dinteri under P. mucronata. No other species of Pteronia besides the clearly different P. eenii has been found in the area directly south of the Etosha pan. A thorough investigation of this extensive area would be needed to conﬁrm the presence of any other species of Pteronia, especially of P. mucronata, to better assess the anomaly of the Dinter 739 location. At his locality II c 4 “Zwart-Ruggens, Karrooﬂäche, 2000 – 3000 Fuss, August” Drège (1843: 61) lists Pteronia mucronata. Parts, or all, of this locality data is found in association with many potential isotypes that all lack a reference to the de Candolle number “[2158]” — for example HAL-0110982, HBG-503990, HBG-505099, K000273457, P-027225, P-027226, P-027227, PRE, SAM0016168-0, TUB-005029, W-0020461, W-Rchb.-18990219753. Similar to other cases, those presented with the year “1837” are also excluded (for example K000273458 and W-0008803, the latter actually indicated as an isotype by W) as it is uncertain if this is a collection date or a herbarium dispatch or receipt date. 14. Pteronia onobromoides DC. (de Candolle 1836: 364); Harvey in Harvey & Sonder (1865: 109); Hutchinson & Phillips (1917: 299); Dinter (1931: 167); Range (1935: 276); Merxmüller (1967: 157); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: South Africa, Western Cape Prov., Olifantrivier W (of “West”?), Drège s.n. [5660] (lectotype G-DC, no. G-00322353!, selected here; isolectotype P-027233! (sub nom. P. mucronata but with “Ebenezar” and [5660] added); other isolectotypes may exist — see Notes). Shrub, to 50 cm tall. Stems glabrous, smooth, pale greywhite. Leaves alternate, glabrous, ﬂat, linear-oblong, distinctly 1- veined beneath, to 20 – 40 × 3 – 5 mm; apex obtuse or subacute; base truncate; margins stifﬂy white ciliate. Capitula solitary, terminal, obconical, 35 × 15 – 25 mm. Receptacle honeycombed and ﬁmbriate. Phyllaries multiseriate, gradate, orbicular to ovate to linear-oblong (inner), to 8 mm long, greenish, glabrous, coriaceous; without midrib; apex truncate or obtuse; margins coriaceous. Florets c. 15; corollas to 20 mm long, glabrous, yellow; lobes linear-lanceolate, acute; tube slender, ribbed. Achenes compressed oblanceolate, densely golden sessile glandular, not hairy, to 7 mm long. Pappus multiseriate, barbellate, to 15 mm long, golden-brown. DISTRIBUTION. Africa: Namibia, South Africa. Map 14. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Lüderitz Distr.: Oranjemund, slopes towards diamond mine, 24 March 1958, Merxmüller & Giess 2326 (M, PRE, WIND!). SOUTH AFRICA. Western Cape Prov.: Olifantrivier W (of “West”?), Drège s.n. [5660] (P. onobromoides lectotype G-DC, no. G-00322353!; isolectotype P-027233!; possible isolectotypes: HAL© The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 Map 14. Known distribution of Pteronia onobromoides in Namibia. 0110983!, HBG-505195!, K000273445!, TUB-005030!, W-Rchb.-1889-0219717!, W-Rchb.-1889-0278273!). Northern Cape Prov.: Namaqualand, Ecklon 31 (P. onobromoides syntype G-DC no. G-00322351!); Ecklon 73 (P. onobromoides syntype G-DC no. G-00322352!). HABITAT. Sandy slopes, coastal steppe dominated by succulent species; no altitudinal data are present on the examined collections. CONSERVATION STATUS. The South African threat status is Least Concern (LC) (SANBI 2011). There are very few collections from Namibia, suggesting that it is rare but the limited information available does not allow an evaluation at this stage, and it must be assigned Data Deﬁcient (DD) status in Namibia (IUCN 2001). PHENOLOGY. Fruiting: March. NOTES. This coastal species has been collected only twice in Namibia (only one of which was seen by us), likely caused by being in the access-restricted diamond mining area in the south-west of the country. Elsewhere it is common along the coast of the Northern and Western Cape Provinces of South Africa. De Candolle (1836: 364) lists two locations with collectors: “in Africa Capensi ad Olifant-rivier (Dreg.!) et in Namaqualand (Eckl.!).”. Inspection in G-DC revealed that there are three collections involved (mounted together on a single sheet), one by Drège with the associated number [5660], and two by Ecklon, nos 31 and 73. These are all of comparable quality, but our research showed that only of the Drège one there is the (distinct) possibility that it has been distributed to other herbaria. Only for this reason we have selected the Drège collection as the lectotype. The enumeration of 1843 shows that Drège collected Pteronia onobromoides at locality III E a 4, which is Ebenezar and located on the Olifant River, KEW BULLETIN (2014) 69: 9488 9488, Page 25 of 44 and it seems reasonable to assume a copy of this gathering was seen by de Candolle. Some possible isolectotypes are the Drège specimens in HAL, HBG, K, TUB and W-Rchb., which are annotated with the locality "Ebenezar" and the code III E a 4, though none carries [5660], which would have demonstrated a more direct link with the G-DC lectotype. Ecklon 31 and 73 in G-DC are now the remaining syntypes. 15. Pteronia paniculata Thunb. (Thunberg 1800: 143); de Candolle (1836: 365); Harvey in Harvey & Sonder (1865: 102); Hutchinson & Phillips (1917: 312); Dinter (1931: 168); Merxmüller (1967: 157); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: South Africa, Western Cape Prov., Cape of Good Hope, Thunberg s.n. (holotype UPS-THUNB). Pteronia ﬂexicaulis auct. non L. f.: de Candolle (1836: 360). Shrub, to 50 cm tall. Stems glabrous, grey. Leaves opposite, clustered along stems, ﬂeshy, lacquered, viscid, bright green, glabrous, linear, terete to boat-shaped, to 15 – 25 × Fig. 6. Fruiting Pteronia paniculata. 1 – 2 mm; apex recurved, subobtuse; base connate, forming a c. 5 mm long sheath. Capitula in dense terminal cymes, cylindrical to narrowly turbinate, 10 × 3 – 4 mm; apex obtuse in bud. Receptacle honeycombed, remains long after seed is shed, conspicuous. Phyllaries multiseriate, ovate-oblong to oblanceolate, 3 – 8 × 1 – 2 mm, lime-yellow, glabrous, lacquered; apex obtuse; without conspicuous membranous margins. Florets to 4; corollas to 5 mm long, glabrous, yellow; lobes lanceolate, subacute; tube ribbed. Achenes compressed obovoid, contracted terminally, long villous, to 2.5 mm long. Pappus bi-seriate, bristles broadened at base, scale-like, c. 4 mm long, white to golden. Fig. 6. DISTRIBUTION. Africa: Namibia, South Africa. Map 15. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Lüderitz Distr.: Diamond Area 1, Skorpion, 27 Aug. 1997, Burke 97208 (WIND!); Kovis Mts, 32 km (20 miles) E of Lüderitz, on the Lüderitz – Aus road, 6 March 1963, De Winter & Hardy 7912 (PRE, WIND!); Aus, 21 Oct. 1922, Dinter 4147 (Z!); Farm Klein Aus, W of Aus, 9 Aug. 1959, Giess & Van Vuuren 932 (PRE, WIND!); Rocky gorge just as you exit Klinghardt Mts, travelling PHOTO: H. KOLBERG. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 26 of 44 KEW BULLETIN (2014) 69: 9488 Merxmüller (1955: 80, 1967: 157); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: Namibia, Gross Namaland, Gubub [Kubub], südöstlich von Aus, Schenck 147 (holotype Z-000003817!). Pteronia quinquecostata Dinter (1932: 183). Type: Namibia, Jakkalskuppe, 1 Nov. 1922, Dinter 4184 (holotype B†; isotype SAM-0071810-0!). Pteronia kingesii Merxm. (Merxmüller 1952: 125). Type: Namibia, Klein Aus, 27 June 1949, Kinges 2280 (holotype M-0104532!; isotype PRE). Map 15. Known distribution of Pteronia paniculata in Namibia. E, 11 Sept. 2005, Mannheimer CM2759 (WIND!); (Aus) quartz foothills, 17 Sept. 2005, Mannheimer CM2821 (WIND!); Kovis Mts, E side, mostly at foot of mountain, 13 Sept. 1972, Merxmüller & Giess 28433 (M, PRE, WIND!); Klinghardt Mts, S part near the Sargdeckel, 19 Sept. 1977, Merxmüller & Giess 32120 (M, WIND!); Gabusib R., 18 Feb. 1992, Strohbach 72 (WIND!); Boegoeberg, 28 Sept. 1992, Strohbach 219 (WIND!); S Namib, Diamond Area no. 1, Aurus Mts (N), 21 April 1988, Ward & Seely 10238 (WIND!); Diamond Area no. 1, Rooiberg, Oct. 1977, Wendt 23/1 (WIND!). HABITAT. In windblown sand on slopes, steppe dominated by succulents; 700 – 1400 m. CONSERVATION STATUS. Although this species' distribution in Namibia is fairly restricted, the estimated EOO and AOO still exceed the threshold for threatened categories (IUCN 2013). In addition no threats are known that would cause population fragmentation, decline or ﬂuctuation to the extent that the species would fall into any threatened category according to criteria A to E of the IUCN system (IUCN 2001). The Namibian conservation status is thus evaluated as LC. The South African threat status is also listed as LC (SANBI 2011). PHENOLOGY. Flowering: August to October. Fruiting: February to April (the exceptional four – six-month period rather than the two – three months as would normally be expected could be ascribed to the limited Namibian data on which this assessment is based). VERNACULAR NAMES. Gombossie (Afrikaans, South Africa). NOTES. In Namibia this species is restricted to the extreme south-west, which receives winter rainfall. 16. Pteronia polygalifolia O. Hoffm. (Hoffmann 1893: 73); Dinter (1926: 132); Range (1935: 276); © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 Shrub, much branched, 80 cm tall. Stems scabrid, brittle, pale grey, striate. Leaves opposite, soft, surface rough papillate, glandular, ﬂat, obovate, spathulate to orbicular, to 10 × 8 mm; apex rounded to subacute; base narrowed into a short petiole. Capitula solitary, terminal, cylindrical, 15 × 3 – 9 mm; apex acute in bud. Receptacle slightly concave. Phyllaries multiseriate, gradate, lanceolate to oblong, to 5 × 2 mm, pale yellow, glabrous; apex subacute; margins membranous. Florets 10 – 12; corollas c. 7 mm long, well exserted above phyllaries, pale yellow; lobes linear-lanceolate, acute. Achenes compressed obovoid, glandular, long white villous, to 3 mm long. Pappus setae basally connate, at most 8 mm long, white to pale golden. Fig. 7. DISTRIBUTION. Africa: endemic to Namibia. Map 16. SPECIMENS EXAMINED. NAMIBIA. Hardap Region, Maltahöhe Distr.: Satanskop off farm Springbokvlakte 166, 23 July 1983, Craven 4218 (WIND!); Namibrand Nature Reserve, 19 Sept. 1997, De Winter 10227 (WIND!); Farm Vreemdelingspoort 141, July 1971, Logan 311 (WIND!); Karas Region, precise district unknown: Gross Namaland, Oranje, 1880, Steingröver 1 (Z!); Bethanie Distr.: Mara farm 114, 11 July 1986, Craven 2666 (WIND!); Lüderitz Distr.: Aus. Dikwillem, 31 July 2002, Bartsch SB873 (WIND!); Granite hills 3 km (1 mile) W of Aus, 11 Sept. 1958, De Winter & Giess 6261 (WIND!, PRE); Jakkalskuppe, 1 Nov. 1922, Dinter 4184 (P. quinquecostata isotype SAM-0071810-0!); Huns Mts, S of farm Uitsig LU 82, state land, 9 June 1976, Giess & Müller 14287 (WIND!, PRE); Aus (?), May 2005, Greyling & Wolf CW 05-08 (WIND!); Klein Aus farm; on slope of Zipfel hill, 27 June 1949, Kinges 2280 (P. kingesii holotype M-0104532!; isotype PRE); Aus Municipal campsite, 7 Aug. 2001, Klaassen & Bartsch EK523 (WIND!); Farm Klein Aus Vista, Geister Trail from cabin in Geisterschlucht, rocky hill slopes to SW, 4 Oct. 2006, Kolberg & Tholkes HK2051 (K!, WIND!); Farm Klein Aus Vista, Geisterschlucht. Rocky gorge E of cabin, 20 Sept. 2007, Kolberg, Van Slageren, Tholkes & Whaley HK2347 (K!, WIND!); Municipal campsite at Aus, 22 Sept. 2003, Mannheimer CM2455 (WIND!); Koppie W of Aus, Sept. 2005, Mannheimer CM2832 (WIND!); Heinrichsfelde, near koppie W of chalets, 29 July 2006, Mannheimer CM3030 (WIND!); Aus, mts on Farm Klein Aus, 18 Aug. 1963, Merxmüller & Giess 2961 (M, WIND!); Kubub Mts S KEW BULLETIN (2014) 69: 9488 Fig. 7. Fruiting Pteronia polygalifolia. 9488, Page 27 of 44 PHOTO: A. MCROBB, RBG KEW. of Aus, 21 Aug. 1963, Merxmüller & Giess 3028 (M, WIND!); Farm Eureka 49, 14 Sept. 1972, Merxmüller & Giess 28449 (M, WIND!); Aus townlands, 10 Oct. 1979, Owen Smith 1274 (WIND!); Great Namaland, Gubub [Kubub], SE of Aus, 14 July 1885, Schenck 147 (P. polygalifolia holotype Z-000003817!); Kubub, March 1885, Schinz 697 (Z!); Namib Desert Park. SW side of Map 16. Known distribution of Pteronia polygalifolia in Namibia. Hauchab Mts, approx. 100 m above base of outcrop, 19 Aug. 1980, Seely & Ward 26 (WIND!); Farm Neisip 34. Aus, 10 June 1972, Wiss 2554 (WIND!). HABITAT. Rocky slopes (mostly granite) in desert and desert-dwarf shrub transition zone; 750 – 1650 m. CONSERVATION STATUS. Using the 1994 IUCN criteria, Craven & Loots (2002) evaluated this species as Lower Risk – Least Concern (LRlc). Using more recent criteria (IUCN 2001) Pteronia polygalifolia is re-evaluated as LC here because its estimated EOO and AOO are larger than the maxima qualifying it for a threatened category. No threats to the species are known that would cause qualiﬁcation under any of the other criteria in the IUCN system (IUCN 2013). Since this is a Namibian endemic, this is a global conservation status. PHENOLOGY. Flowering: May to October. Fruiting: September to December. NOTES. This species has been recorded only in southwestern Namibia (Map 16). 17. Pteronia pomonae Merxm. (Merxmüller 1952: 127; 1967: 157); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: Namibia, Alicetal bei Pomona, 6 June 1929, Dinter 6412 (holotype M-0104533!; isotypes B-10-0097214!, B-10-0097215!, HBG505135!, HBG-505187!, HBG-505188!, HBG© The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 28 of 44 505189!, HBG-505191!, K-000273528!, PRE-01616310!, S-07-7260!, Z-000052428!). Pteronia cancellata Dinter (1932: 184), nom. nud. — see Notes. Pteronia villosa auct. non L. f.: Dinter (1931: 169, author as “L.”;1932: 184); Range (1935: 276, author as “L.”). Shrub, much-branched, to 50 cm tall. Stems glabrous, pale grey when young, dark grey and ﬁssured when older. Leaves alternate to almost opposite, clustered on dwarf shoots, terminally with long, white marginal bristles forming cage around capitula, lower leaves without bristles, glabrous, linear, terete, 10 × 1 mm; apex obtuse; base slightly eared, sessile, not connate. Capitula solitary, terminal, cylindrical to ovoid, 8 – 10 × 4 – 5 mm; apex rounded in bud. Receptacle ﬁmbriate. Phyllaries multiseriate, narrowly oblong to elliptic, 2 – 3 × 5 – 10 mm, green to yellow, glabrous; apex rounded; margins dark brown or red on outer and narrow hyaline on inner phyllaries. Florets 9 – 12; corollas to 6 mm long, pilose, bright yellow; lobes linear-lanceolate, acute; tube gradually widening. Achenes turbinate compressed, densely sericeous, glandular, to 2 mm long. Pappus setae connate at base, c. 4 mm long, golden. Fig. 8. DISTRIBUTION. Africa: endemic to Namibia. Map 17. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Lüderitz Distr.: Diamond Area No. 1, Chamnaub Inselberg, NE of Boegoeberg, 28 Aug. 2002, Bartsch SB929 (WIND!); Diamond Area 1, Namitsis Inselberg, about 30 km S of Klinghardts, 1 Sept. 2002, Bartsch, Loots & Mannheimer SB947 (WIND!); Rote Kuppe – Chamais road, 6 Sept. 2002, Bartsch, Loots & Mannheimer SB1035 (WIND!); Diamond Area 1, Klinghardt's Mts, 21 Sept. 1996, Burke 96156 (WIND!); W side of Münzen Mts, 30 Aug. 1958, De Winter & Giess 6120 (PRE, WIND!); Gais area, E of Hohenfels, on the road to Jakkalsberge, Oranjemund, 4 Sept. 1958, De Winter & Giess 6199 (PRE, WIND!); Diamond area no 1, Oranjemund, Gais area, E of Hohenfels, on the road to Jakkalsberge, 5 Sept. 1958, De Winter & Giess 6202 (WIND!); Coastal desert on gneiss hills at Halenberg, 15 Oct. 1922, Dinter 4086 (K!, PRE); Alicetal near Pomona, 6 June 1929, Dinter 6412 (P. pomonae holotype M-0104533!; isotypes B-10-0097214!, B-10-0097215!, HBG-505135!, HBG-505187!, HBG-505188!, HBG-505189!, HBG505191!, K-000273528!, PRE-0161631-0!, S-07-7260!, Z000052428!; P. cancellata vouchers B-10-0097214!, B-100097215!, E-00413376!, HBG-505135!, HBG-505187!, HBG-505188!, K-000273528!, S-07-7260!, SAM0071825-0!, Z-000052428!); Klinghardt Mts, 31 Aug. 2002, Gess & Gess 02/03/28 (WIND!); Farm Witputz Nord LU 22. 20.5 km SW of the police station, 30 Sept. 1975, Giess 13766 (PRE, WIND!); Farm Spitzkop LU 111, 14 Aug. 1976, Giess 14630 (PRE, WIND!); Sperrgebiet, turn-off at N entrance to Klinghardt Mts, 4 Aug. 2001, Klaassen & Bartsch EK466 (WIND!); 39 km from Rotkop on powerline track, low koppie, 22 © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 Oct. 1987, Kolberg & Maggs HK194 (PRE, WIND!); Kovis Mts, campsite, at base of mountain, SW of radio mast, 8 Oct. 2006, Kolberg & Tholkes HK2079 (K!, WIND!); Sperrgebiet, Lüderitz – Oranjemund road at N turn-off to Bogenfels, 14 Oct. 2006, Kolberg & Tholkes HK2098 (K!, WIND!); Close to the mainroad, 9 Sept. 2005, Kwembeya EKw57 (WIND!); Tafelberg, SE of Klinghardt Mts, Sperrgebiet 1, Aug. 1971, Logan & Jensen 902 (WIND!); Schwarzkop, NE of Bogenfels, Sperrgebiet 1, Aug. 1971, Logan & Jensen 1093 (WIND!); Pietab 1, Klinghardt Mts, Sperrgebiet, Aug. 1971, Logan & Jensen 1137 (WIND!); Between Chamnaib and Bogenfels, 29 Aug. 2002, Mannheimer CM2046 (WIND!); Namitsas, S of Klinghardt Basin, Sept. 2002, Mannheimer CM2095 (WIND!); E Sperrgebiet, 2 Sept. 2002, Mannheimer CM2148 (WIND!); Road to Grillental from Kaukausib, 5 Sept. 2002, Mannheimer CM2202 (WIND!); 23 km S of Grillental, 12 Sept. 1972, Merxmüller & Giess 28381 (M, PRE, WIND!); 9.5 km S of Grillental, 15 Sept. 1977, Merxmüller & Giess 32019 (M, PRE, WIND!); Klinghardt Mts, in the S part, in the region of Sargdeckel, 17 Sept. 1977, Merxmüller & Giess 32110 (M, WIND!); Klinghardt Mts, near the foot of the mountain, 28 July 1977, Müller 696 (WIND!); Sperrgebiet, hills on N side of Klinghardt Mts, 13 Aug. 2001, Smook 11374 (PRE, WIND!); Diamond area no. 1, c. 18 km W of Rosh Pinah, towards Obib Mt, 1 Sept. 1989, Van Wyk 8890 (PRE, WIND!); S Namib, Diamond area no. 1, S of Klinghardt Mt, 21 April 1988, Ward & Seely 10233 (WIND!); Diamond Area 1, Tsaus Spinnenberg, Oct. 1977, Wendt 8/1 (WIND!). HABITAT. Plains and slopes in steppe characterised by succulents; 200 – 700 m. CONSERVATION STATUS. Present information leads to an evaluation of LC (IUCN 2001) for this relatively abundant species. Both estimated EOO and AOO are well above the thresholds for threatened categories and no population decline, fragmentation or ﬂuctuation could be identiﬁed that would qualify this species as threatened (IUCN 2013). Pteronia pomonae is endemic to Namibia and is LC thus the global status. PHENOLOGY. Flowering: July to October. Fruiting: October to January. NOTES. The characteristic long ciliate leaves at the base of capitula make this Namibian endemic species easily recognisable amongst Pteronia species in the country (Fig. 8). Pteronia cancellata is a manuscript name (Dinter 1932: 184) associated with Dinter 6412, now the type of P. pomonae, and found on vouchers in (at least) B, E, HBG, K, S, SAM, and Z, as follows: B-100097214!, B-10-0097215!, E-00413376!, HBG505135!, HBG-505187!, HBG-505188!, K000273528!, S-07-7260!, SAM-0071825-0!, and Z000052428!. Of these vouchers E and SAM do not present a reference to P. pomonae on the sheet, KEW BULLETIN (2014) 69: 9488 9488, Page 29 of 44 Fig. 8. Pteronia pomonae. A habit; B (upper) leaf, abaxial; C (upper) leaf, adaxial; D capitulum, side view; E (1 – 5) phyllaries, left to right: outer to inner; F ﬂoret, side view; G corolla, opened and stamens; H style; J (young) achene with pappus; K pappus setae detail. A – K Kolberg & Tholkes HK2098. DRAWN BY JULIET BEENTJE. while all others do. Rather than describing and adopting the name cancellata, Merxmüller (1952) chose to ignore the epithet and replace it with pomonae. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 30 of 44 Map 17. Known distribution of Pteronia pomonae in Namibia. 18. Pteronia quadrifaria Dinter (1932: 182). Type: Namibia, Gross Namaland, Zwartaus, 17 March 1929, Dinter 6190 (holotype B†; lectotype HBG-505185!, selected here; isolectotype HBG-505186!). Erect, intricately branched shrub, to 25 cm tall. Stems pale tan or pale grey, lower layers white, farinose, bark KEW BULLETIN (2014) 69: 9488 on older stems dark grey, longitudinally ﬁssured, leaf scars conspicuous, spiny remains of parts connecting opposite leaves most conspicuous near branch tips. Leaves decussate-imbricate, densely arranged at branch tips, less imbricate on ﬂowering branches, dehiscent at base of branches, somewhat ﬂeshy, surface minutely papillose, margins ciliate-pectinate, more so towards leaf apex, sometimes abaxial keel pectinate at apex, narrowly oblong to linear-lanceolate, apex obtuse, boat-shaped, convex beneath, midrib indented on slightly concave adaxial surface, 4 – 5 × 1 – 1.5 mm; apex obtuse; base sessile, somewhat connate, with spinules between bases of opposite leaves. Capitula solitary, terminal, obconical to ovoid, 10 – 12 × 4 – 6 mm; apex constricted, truncate in bud. Receptacle epaleate. Phyllaries multiseriate, gradate, spirally arranged, c. 20 per capitulum, oblong, 3 × 1.5 mm to 10 × 1 mm, pale yellow-green, shiny, papillose towards apex on abaxial surface; apex of outer acute, of inner truncate; margins undulate, membranous and reﬂexed at right angles. Florets 5 – 6; corollas 6.5 – 7 × 1 mm, short papillate at area where translucent part meets opaque part, pale yellow; lobes narrowly acute, short bristly on outer surface; tube gradually widening towards apex, basal third translucent, rest opaque. Stamens 5, ﬁlaments fused to base of corolla. Anthers fused into a tube, about half as long as stamens, bases shortly sagittate, tips acute. Style biﬁd, branches acute, half the length of style; stigmatic papillae short. Ovary obconical to obtrigonous, densely cov- Fig. 9. Pteronia quadrifaria showing the reﬂexed, undulate phyllaries, maroon pappus and densely imbricate, decussate leaves. PHOTO: H. KOLBERG. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 ered with pale yellow, spherical, sessile glands, densely hirsute. Achenes compressed obconical, 9488, Page 31 of 44 densely long, white hirsute and densely covered with golden, spherical, sessile glands, 4.5 – 5 × 2 – 2.5 mm. Fig. 10. Pteronia quadrifaria. A habit; B detailed positions of leaves; C leaf; D capitulum; E (1 – 7) sample of phyllaries; F side view of ﬂoret and (young) achene; G ﬂoret, side view; H corolla tube, opened; J stamen; K style; L style branches; M young achene; N pappus setae (three lengths). A – N Kolberg & Tholkes HK2893. DRAWN BY JULIET BEENTJE. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 32 of 44 Pappus setae fused at base into a ring, 4 – 8 mm long, bristles barbellate, of different lengths, from half the length of ﬂoret to longer than ﬂoret, maroon-purple. Figs 9 – 10. DISTRIBUTION. Africa: endemic to Namibia. Map 18. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Lüderitz Distr.: Zwartaus, 17 March 1929, Dinter 6190 (P. quadrifaria lectotype HBG-505185!, isolectotype HBG-505186!); Aus, hill N of Aus with NBC mast, below quartz ridge, 12 Oct. 2009, Kolberg & Tholkes HK2893 (K!, WIND!); Aus townlands, white quartz hill with NBC mast, N of town, 13 Oct. 2010, Kolberg & Tholkes HK2994 (K!, WIND!). HABITAT. Quartz or granite hills in steppe characterised by a diversity of succulent species; c. 1600 m. CONSERVATION STATUS. A preliminary global conservation status of EN D1 can be assigned to this Namibian endemic species because of the small number of individuals found at the only site known (IUCN 2013). The population found recently was the ﬁrst one in Namibia since Dinter’s 1929 collection, and consists of about 50 – 70 old plants on white quartz with an estimated EOO well below 100 km2 and an AOO smaller than 10 km2. No young plants were seen. The species was not found on other hills with similar substrate in the Aus vicinity. At the same time no visible or increasing threat to the population could be observed. Dinter (1932), however, reported having found this species on granite of which there are many hills in the Aus area that still need to be investigated. Until the possible distribution area is searched more closely for further populations of this species, no ﬁnal assessment of the conservation status can be made. KEW BULLETIN (2014) 69: 9488 PHENOLOGY. Flowering: October. Fruiting: December. NOTES. Dinter (1932: 182) described Pteronia quadrifaria from granite hills near Aus. The collected material at the time was, by Dinter’s account, sterile (Dinter 3580) or “with sterile heads” (Dinter 6190). Indeed, both specimens at HBG have no intact ﬂowering parts. Dinter’s original collections were at Berlin (Lanjouw & Staﬂeu 1954: 163) and are now considered lost. Of the two syntypes Dinter 3580 was not found anywhere else, but two specimens of 6190 were located at HBG, each with annotations in Dinter’s handwriting; the collection HBG-505185, being a bit more copious, is selected as the lectotype. Pteronia quadrifaria was treated by Merxmüller (1955) as a synonym of Pteronia lucilioides DC., based mainly on the papillate leaves of the Dinter 6190 specimen. He argued that the growth form of P. lucilioides is very variable in Namibia and that Dinter 6190 represents a densely branched, small, shrubby form. Around Aus, however, P. lucilioides is a tall, virgate shrub and the stunted dwarf shrub forms are found mostly nearer to the coast. Based on recent ﬂowering and fruiting collections the differences between the two species were conﬁrmed, such as the apex of the budding capitula that is constricted to truncate in P. quadrifaria and sharply acute in P. lucilioides (compare Figs 4 and 9). Furthermore the densely imbricate, decussate leaf arrangement of P. quadrifaria clearly differs from the opposite leaves of P. lucilioides, and the undulate, reﬂexed phyllaries of the former differ from the smooth, straight phyllaries of the latter. P. quadrifaria can also be clearly distinguished from P. lucilioides by the maroon-purple pappus (straw-coloured in the latter). Reviewing all presented evidence we conclude that this merits the resurrection of P. quadrifaria as a distinct entity. Pteronia quadrifaria is similar to P. mucronata in its leaves with ciliate margins and truncate phyllaries, but differs in its decussate-imbricate leaf arrangement, its phyllary apex not being mucronate but undulate-reﬂexed, and the conspicuous maroon-purple pappus (Fig. 9). 19. Pteronia rangei Muschl. (Muschler 1911a: 96); Hutchinson & Phillips (1917: 319); Dinter (1926: 132); Range (1935: 276); Merxmüller (1952: 128; 1967: 158); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: Namibia, Bezirk von Gross Namaqualand, Aus, Tafelberg, 1600 m, Oct. 1906, Range A 26 (holotype B†; lectotype BOL-138777!, selected here; isolectotype K-000273503!). Map 18. Known distribution of Pteronia quadrifaria in Namibia. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 Shrub, to 60 cm tall. Stems glabrous, cream to pale grey, leaf scars distinct. Leaves alternate, clustered along stems, somewhat succulent, greyish-green, aromatic, surface glabrous and tuberculate, linear, subterete, curved towards stem, to 25 × 1 mm; apex obtuse; base sessile, with tuft of hairs in axil. Capitula solitary, KEW BULLETIN (2014) 69: 9488 9488, Page 33 of 44 Fig. 11. Pteronia rangei. A habit of branches; B branchlet with leaves; C capitulum, side view; D (left to right) phyllaries in outer, mid, and inner position in capitulum, and an inner one viewed abaxially; E (left and right) ﬂoret side view with stamens (detail, left), and overview from the side with stamens protruding (right); F ﬂoret side view with style branches; G corolla opened with stamens and style (separate); H young achene with pappus; J achene detail; K pappus setae detail. A – K Kolberg & Tholkes HK2892. DRAWN BY JULIET BEENTJE. terminal, cylindrical to campanulate, elongate, to 15 × 10 mm; apex truncate in bud. Phyllaries multiseriate, gradate, oblong-lanceolate to linearlanceolate, to 10 × 2 mm, green to yellow, glabrous; © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 34 of 44 midribs thickened, glandular, forming a recurved, small mucro; apex subacute; margins membranous. Florets c. 12; corollas to 8 mm long, yellow; lobes linear lanceolate, subacute; tube contracted at apex, strongly ribbed in lower half. Achenes oblong-cylindrical, appressed villous, to 3 mm long. Pappus multiseriate, barbellate, c. 12 mm long, pale yellow to golden brown. Fig. 11. DISTRIBUTION. Africa: endemic to Namibia. Map 19. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Bethanie Distr.: Farm Kosos 11, 15 Aug. 1963, Merxmüller & Giess 2817 (M, WIND!); Farm Saraus BET 16, 5 Sept. 1972, Merxmüller & Giess 28227 (M, WIND!); Keetmanshoop Distr.: Farm Lourensia, N of homestead, 28 March 1998, Strohbach & Dauth 3764 (WIND!); Lüderitz Distr.: Kuibis, 2 Nov. 1922, Dinter 4181 (Z!); Farm Kolke LU 84, not far from the farmhouse, 11 June 1976, Giess & Müller 14332 (WIND!); 70 km from Rotkop Station on power-line track, W slope of mountains, 23 Oct. 1987, Kolberg & Maggs HK208 (WIND!); Farm Zebrafontein, track from road D463; at locked gate just before homestead, 20 Oct. 2007, Kolberg & Tholkes HK2411 (K!, WIND!); 21 km E along road D463 from Witpütz junction on Aus – Rosh Pinah tar road, 12 Oct. 2009, Kolberg & Tholkes HK2892 (K!, WIND!); Aus, Tafelberg, 1600 m, Oct. 1906, Range A 26 (P. rangei lectotype BOL138777!, isolectotype K-000273503!); Farm Plateau 38, on the slope of the ridges N of the farmhouse, 13 April 1953, Walter & Walter 2577 (WIND!). HABITAT. Rocky slopes and plains of dwarf shrub savanna, desert-dwarf shrub transition and steppe dominated by succulent species; 1300 – 1400 m. CONSERVATION STATUS. Data Deﬁcient (DD) according to Craven & Loots (2002) and Loots (2005). During recent ﬁeldwork the ﬁrst author could not identify any threats to populations and the species is more abundant and widespread than the small number of herbarium specimens would suggest. Reevaluation using IUCN (2001) criteria resulted in a global LC status (Pteronia rangei is endemic to Namibia) also because estimated EOO and AOO are well above the thresholds for threatened status (IUCN 2013). PHENOLOGY. Flowering: June to October. Fruiting: October to December. NOTES. The tuberculate leaf surface is unique among the Namibian species of Pteronia (Fig. 11). This Namibian endemic has been recorded only in the central south and south-west of the country (Map 19). We have chosen a lectotype since the original holotype in B is assumed lost. Isotypes were found only in BOL and K. As the BOL isotype is of altogether better quality, it is selected here as the lectotype. The isolectotype in K (fragments only) notes (strung together here): “ﬂowers yellow, shrub, ½ m, Tafelberg 1600 m, © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 Oct. 06, Deutsch – Südwest Afrika, Dr. Range”, while BOL has the altitude in feet (4800'). It should be noted that none of the lectotype collections actually carries “Aus” as the locality, only “Tafelberg” — of which there is one near Aus where the type was collected at a place called Kubub. This is in accordance with Paul Range’s handwritten list of localities, seen by the ﬁrst author. 20. Pteronia scariosa L. f. (Linnaeus ﬁlius 1782: 356); Thunberg (1800: 144); de Candolle (1836: 361); Harvey in Harvey & Sonder (1865: 104); Hutchinson & Phillips (1917: 315); Dinter (1926: 132); Range (1935: 276); Merxmüller (1952: 128; 1967: 158); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: South Africa, Western Cape Prov., Cape of Good Hope, Thunberg s.n. (holotype UPS-THUNB; isotype LD-1254325!). Pteronia lycioides Muschl. ex Dinter (Dinter 1926: 132); Range (1935: 275). Type: Namibia, Büllsporter Fläche, Dinter 2146 (not traced, likely B†). Shrub, to 50 cm tall. Stems glabrous, tan to greyish-brown, tips somewhat spinescent. Leaves alternate, thickish, coriaceous, glabrous, obovate to oblong-lanceolate, to 10 × 3 mm; apex rounded to subacute; base cuneate. Capitula solitary, terminal, subcampanulate, to 20 × 18 mm; apex acute in bud. Receptacle ﬂat, surface slightly jagged. Phyllaries multiseriate, gradate, outer much smaller than inner, broadly lanceolate, keeled, 3.5 × 2 mm to 15 × 4 mm, pale reddish-brown, membranous, translucent, glabrous; midrib narrow, brown, extending into a sharp point; apex mucronate, acute; margins ﬁnely lacerate. Florets c. 14; corollas to 2.5 mm long, yellow; lobes linear-lanceolate, subacute, margins reddish-brown when dry; tube cylindric, 5ribbed near base. Achenes obovoid, contracted into a short, glabrous neck, long villous, to 4 mm long. Pappus setae fused into a basal ring, to 13 mm long, brown to reddish-brown. DISTRIBUTION. Africa: Namibia, South Africa. Map 20. SPECIMENS EXAMINED. NAMIBIA. Hardap Region, Maltahöhe Distr.: Büllsporter Fläche, 20 Sept. 1947, Strey 2130 (NBG!, PRE); Khomas Region, Windhoek Distr.: Farm Göllschau, 19 Nov. 1934, Dinter 7980 (P!, WIND!). SOUTH AFRICA. Western Cape Prov.: Zilverfontein, 25 Oct. 1830, Drège s.n. (HBG-505184!, K-000273490!, P-027261!); Cape of Good Hope, Thunberg s.n. (P. scariosa holotype UPS-THUNB, isotype LD-1254325!). HABITAT. Seasonally ﬂooded plain in dwarf shrub savanna. CONSERVATION STATUS. This species has restricted distribution (small estimated EOO and AOO) and small number of populations in Namibia but no KEW BULLETIN (2014) 69: 9488 9488, Page 35 of 44 Map 19. Known distribution of Pteronia rangei in Namibia. Map 20. Known distribution of Pteronia scariosa in Namibia. speciﬁc threats are known from the area of distribution. However, Pteronia scariosa has not been collected in Namibia in the past 65 years and no speciﬁc efforts were made to ﬁnd it. Because of the lack of sufﬁcient recent data, the Namibian status is DD (IUCN 2001, 2013). Existing herbarium collections suggest that it is rare in Namibia. The South African threat status has been recorded as LC (SANBI 2011). PHENOLOGY. Flowering: September to November. NOTES. Only three specimens are known from Namibia, of which Dinter 2146 could not be traced (it was undoubtedly in B and is now considered lost), and the species has not been collected since 1947. The Namibian population is disjunct from the distribution of the species in South Africa’s Northern and Western Cape Provinces. More material needs to be collected in Namibia. The chances of plants still being at the recorded localities are good, since these are in relatively protected commercial farmland that has been utilised sustainably. opposite, glabrous or minutely puberulous, linear to linear-lanceolate, boat-shaped, to 6 × 2 mm; apex obtuse; bases somewhat fused. Capitula solitary, terminal, obconical, base narrow, 15 × 10 – 12 mm; apex subacute to truncate in bud. Receptacle ﬁmbriate. Phyllaries multiseriate, oblong-oblanceolate, closely imbricate, to 10 × 3 mm, greenish-yellow, glabrous; midrib dark greyish-green; apex acute to obtuse; lateral margins membranous. Florets 5 – 9; corollas to 12 mm long, yellow; lobes lanceolate, subacute; tube contracted near base. Achenes oblong, densely villous, to 3.5 mm long. Pappus multiseriate, 8 – 9 mm long, golden to straw-coloured. 21. Pteronia sordida N. E. Br. (Brown 1906: 108); Hutchinson & Phillips (1917: 325); Merxmüller (1955: 80, 1967: 158); Merxmüller & Roessler (1984: 90); Herman (2003: 278). Type: South Africa, Eastern Cape Prov., Conway Farm, Middelburg Div., 1100 m, Aug. 1899, Gilﬁllan in herb. Galpin 5527 (holotype K000273517!; isotype PRE-161726-0). Pteronia chlorolepis Dinter (1932: 181). Type: Namibia, Gross Namaland, Aus, 26 Oct. 1922, Dinter 4153 (lectotype HBG-505115!, selected here; isolectotypes SAM-0071816-0!, Z-000003813!). — see Notes. Pteronia glomerata auct. non L. f.: Range (1935: 275). Shrub to 35 cm tall. Stems mostly glabrous, sometimes whitish pubescent, older bark grey, ﬁssured. Leaves DISTRIBUTION. Africa: Namibia, South Africa. Map 21. SPECIMENS EXAMINED. NAMIBIA. Hardap Region, Rehoboth Distr.: 23 km W along road D1261 from junction with road C24 (NE of Nauchas), 28 Aug. 2009, Kolberg & Tholkes HK2824 (K!, WIND!); Karas Region, Bethanie Distr.: 28 km S of Helmeringhausen on road to Aus, 3 Oct. 2006, Kolberg & Tholkes HK2042 (K!, WIND!); Tirasberge, 16 Aug. 1963, Merxmüller & Giess 2866 (M, WIND!); Lüderitz Distr.: Aus, 26 Oct. 1922, Dinter 4153 (P. chlorolepis lectotype HBG-505115!; isolectotypes SAM-0071816-0!, Z-000003813!); Aus townlands, S of town, 3 Oct. 2006, Kolberg & Tholkes HK2045 (K!, WIND!); Farm Witpütz Süd, about 5 km NE of house, calcrete ridges E of track to road D463, 25 Oct. 2007, Kolberg & Tholkes HK2443 (K!, WIND!); Aus, at river on road to Helmeringhausen, 7 Aug. 1963, Merxmüller & Giess 2917 (M, PRE, WIND!); Farm Plateau, 20 Aug. 1963, Merxmüller & Giess 3014 (M, WIND!); Gross Namaland, Farm Plateau at Schakalskuppe station, 13 April 1953, Walter & Walter 2558 (WIND!); Farm Plateau near Aus, 9 Sept. 1963, Wiss 2021 (WIND!). SOUTH AFRICA. Eastern Cape Prov.: Conway Farm, Middelburg Div., Aug. 1899, Gilﬁllan in © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 36 of 44 herb. Galpin 5527 (P. sordida holotype K-000273517!; isotype PRE-161726-0). HABITAT. Gravelly and rocky hills and plains in desert, desert-dwarf shrub transition and dwarf shrub savanna; 1200 – 1650 m. CONSERVATION STATUS. In Namibia the estimated EOO and AOO (22,500 km2 and 2,500 km2 respectively) just exceed the maximum for the VU category under criterion B (IUCN 2013). Pteronia sordida is known from only four localities in Namibia, which would qualify it for a VU status under criterion D2, but the necessary additional criteria for threatened status (plausible threat, population decline, fragmentation or ﬂuctuation) are not established (IUCN 2001, 2013). This results in a status of LC in Namibia, the same as in South Africa (SANBI 2011). PHENOLOGY. Flowering: April to September. Fruiting: August to November. NOTES. The distribution is disjunct in Namibia with populations along the south-western escarpment and one record from central Namibia. The Namibian populations are in turn disjunct from the South African ones in the Northern, Western and Eastern Cape Provinces. The Namibian distribution can be explained by the species’ preference for higher altitudes but more material is needed to conﬁrm this. The HBG sheet of Dinter 4153 is chosen as the lectotype of Pteronia chlorolepis since it is marginally more informative than the other two extant duplicates seen. The other syntype associated with the name (Gross Namaland, Aus, 7 April 1929, Dinter 6259), was probably only present in B and is now considered lost. 22. Pteronia spinulosa E. Phillips in Hutchinson & Phillips (1917: 320); Merxmüller (1967: 158); Merxmüller & Roessler (1984: 90); Herman (2003: 279). Type: Namibia, Map 21. Known distribution of Pteronia sordida in Namibia. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 Angra Pequena [Lüderitzbucht], 18 January 1907, Galpin & Pearson 7645 (lectotype SAM-0001648-0!, selected here; isolectotypes BOL-138778!, K-000273506!, PRE-0160582-0!). Shrub, to 40 cm tall. Stems with rows of spinules where leaf connections remain, glabrous, glaucous-grey. Leaves opposite, clustered along stems, slightly succulent, glabrous but densely warty when dry, linear, boat-shaped to trigonous, keeled, to 12 × 4 mm; apex obtuse; base connate. Capitula solitary, terminal, oblong-ovoid, 12 – 15 × 6 – 7 mm; apex subacute in bud. Receptacle deeply honeycombed, ﬁmbriate. Phyllaries multiseriate, oblong, closely imbricate, 2 – 10 mm long, bright yellow, glabrous; midrib with oblong thickening; apex rounded; margins not obviously membranous. Florets to 10; corollas to 6.5 mm long, glabrous, bright yellow; lobes narrowly triangular, subobtuse. Achenes obovoid, long appressed villous, 3 – 4 mm long. Pappus c. 5 mm long, straw-coloured. Figs 12 – 13. DISTRIBUTION. Africa: endemic to Namibia. Map 22. SPECIMENS EXAMINED. NAMIBIA. Karas Region, Lüderitz Distr.: Diamond Area 1, Chameis, 22 Aug. 1997, Burke 97169 (WIND!); Bogenfels, 5 Sept. 1958, De Winter & Giess 6216 (NBG, PRE, WIND!); Lüderitzbucht, Jan. 1929, Dinter 5994 (Z!); Lüderitzbucht, 18 Jan. 1907, Galpin & Pearson 7645 (P. spinulosa lectotype SAM0001648-0!; isolectotypes BOL-138778!, K-000273506!, PRE-0160582-0!); Spencer Bay – Nordhuk, 12 Jan. 1974, Giess & Robinson 13192 (PRE, WIND!); Sperrgebiet, Pomona, road to house, 3 Aug. 2001, Klaassen & Bartsch EK460 (WIND!); Lüderitz peninsula, Mesemb Bay, 10 Oct. 2006, Kolberg & Tholkes HK2091 (K!, WIND!); Sperrgebiet, Granitberg Station, N of Bogenfels, N of old mine dumps, 17 Feb. 2007, Kolberg & Tholkes HK2199 (K!, WIND!); Lüderitzbucht, Nov. 1908, Marloth 4621 (GRA-0002940-0!, PRE0160583-0!); Lüderitzbucht, on rocky terrain towards Sturmvogelbucht, 21 March 1958, Merxmüller & Giess 2246 (M, PRE, WIND!); Nautilus, N Lüderitz, 23 Aug. 1963, Merxmüller & Giess 3075 (M, WIND!); Bogenfels, 9 Sept. 1972, Merxmüller & Giess 28327 (M, PRE, WIND!); Valley W of Elizabeth Bay, June 1993, Williamson 4543 (WIND!). HABITAT. Gravelly or rocky plains and hill slopes with windblown sand in desert and coastal steppe characterised by a diversity of succulent species; 50 – 950 m. CONSERVATION STATUS. This species is restricted almost entirely to the protected diamond mining area, now a national park. Although Pteronia spinulosa is found mostly close to the coast where mining activities are also concentrated, a signiﬁcant threat that would cause population reduction has not been identiﬁed. Mining here also underlies a sound environmental management policy that would control future threats. The estimated EOO and AOO and population sizes and number of locations are well above KEW BULLETIN (2014) 69: 9488 the threshold for the threatened categories (IUCN 2001, 2013). The conservation status was assessed by Loots (2005) as LC using IUCN (2001) criteria. This is a Namibian endemic and the status thus global. PHENOLOGY. Flowering: August to October. Fruiting: December to March. NOTES. Edwin Phillips (in Hutchinson & Phillips 1917) listed Galpin & Pearson 7645 and Marloth 4621, these together being the syntypes of Pteronia spinulosa. The collections are equal in terms of quality and documentation, preventing an obvious choice between them for a lectotype. It appears, however, that the Galpin & Pearson specimen is more widely distributed than the Marloth one, and is therefore chosen as the lectotype. Since Phillips worked at the South African Museum at the time of publishing this species, the Galpin & Pearson specimen at SAM is selected as lectotype among the four existing specimens. Marloth 4621 from Angra Pequena [Lüderitzbucht] remains the other syntype. This coastal species is endemic to Namibia (Map 22). The spinules that remain after leaf dehiscence can best be seen on younger twigs because they wear off on older branches (Figs 12 & 13). 23. Pteronia unguiculata S. Moore (1904: 1012); Hutchinson & Phillips (1917: 326); Dinter (1926: 133); Range (1935: 276); Merxmüller (1967: 158); Merxmüller & Roessler (1984: 90); Herman (2003: 279). Type: Namibia, Gross Namaland, Gubub 9488, Page 37 of 44 [Kubub], July 1897, Dinter 1233 (holotype BM001114681!, isotype Z-000003820!). Shrub, to 100 cm tall, much-branched. Stems glabrous, pale grey to grey-brown. Leaves opposite, clustered along stems, glabrous, linear to clavate, terete, to 10 × 1 mm; apex obtuse; base slightly eared, sessile. Capitula solitary, terminal, elongate-cylindrical, 15 × 8 mm; apex obtuse in bud. Phyllaries multiseriate, broadly obovate to elliptic, broadest at apex, slightly contracted below apex, 7 – 9 × 5 mm, bright yellow, glabrous; apex truncate, mucronulate, often emarginate; margins entire, at most indistinctly lacerate-ciliate. Florets c. 10; corollas to 5 mm long, glabrous, bright yellow; lobes narrlowly lanceolate. Achenes turbinate, long sericeous, with few sessile glands, to 3 mm long. Pappus c. 5 mm long, pale yellow. Fig. 14. DISTRIBUTION. Africa: Namibia, South Africa. Map 23. SPECIMENS EXAMINED. NAMIBIA. Erongo Region, Omaruru Distr.: Brandberg, Amis Kloof, Sept. 1979, Craven 1097 (WIND!); Brandberg, near Horn on the higher plains on NW Brandberg, 19 July 1984, Craven 1883 (WIND!); Brandberg, upper reaches of Königstein, 6 May 1993, Craven 4009 (WIND!); Hardap Region, Maltahöhe Distr.: Namib Naukluft Park, Naukluft Plateau, 6 July 1993, Bridgeford 141 (WIND!); E facing path up Losberg to “repeater” (a telecoms mast) in the Namibrand Nature Reserve, 1 July 2004, Clapham & Drayer 149 (WIND!); Farm Mooirivier 169, 2 Sept. 1978, Horn 31 (WIND!); (Naukluft park) Fig. 12. Horizontal-spreading branches of Pteronia spinulosa with fruiting capitula just prior to releasing achenes. PHOTO: H. KOLBERG. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 38 of 44 KEW BULLETIN (2014) 69: 9488 Fig. 13. Pteronia spinulosa. A habit; B leaf, side view; C leaf showing ﬂeshy adaxial surface; D detail of “spinules” (remnants after pair of leaves drop off); E capitulum; F capitulum with most phyllaries removed; G range of phyllaries; H budding ﬂoret, side view; J ﬂoret, side view; K style branches, detail; L young achene with pappus; M pappus setae, detail. A – M Kolberg & Tholkes HK2199. DRAWN BY JULIET BEENTJE. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 9488, Page 39 of 44 Map 22. Known distribution of Pteronia spinulosa in Namibia. Tweelingpan No. 9, 9 April 1971, Meyer 500 987 (WIND!); Losberg on the NRNR. (Namibrand Nature Reserve), 6 Oct. 2001, Rahn 36 (WIND!); Bergzebrapark-Naukluft, 20 June 1968, Van der Fig. 14. Flowering Pteronia unguiculata branches. Map 23. Known distribution of Pteronia unguiculata in Namibia. Westhuizen 3 (WIND!); Farm Naudaus/Duwisib MAL 76/84, 20 May 1956, Volk 12756 (WIND!); Farm Maguams-Krähwinkel, Helmeringhausen, 23 March PHOTO: I. DINTER. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 40 of 44 1953, Walter & Walter 2141 (WIND!); Rehoboth Distr.: Springbokvlakte (Farm Nauzerus), 1 Nov. 1997, Bührmann & Bührmann BUH1/13 (WIND!); Quartz dyke, Bastardland, Nov. 1934, Dinter 8045 (Z!); Karas Region, Bethanie Distr.: 38 km (23.5 miles) N of Helmeringhausen, 19 Oct. 1949, Acocks 15634 (PRE, WIND!); Farm Kosos, Helmeringhausen, Aug. 1963, Merxmüller & Giess 2818 (M, WIND!); Farm Saraus (BET 16), 5 Sept. 1972, Merxmüller & Giess 28228 (M, PRE, WIND!); Rolling hills on Farm Aruab 23, July 1998, Miller MIL1/097 (WIND!); Karasburg Distr.: 20 km (12.25 miles) NNE of Grünau, 13 Oct. 1949, Acocks 15562 (PRE, WIND!); Farm Vrede, 19 July 2005, Bruyns 10106 (BOL, WIND!); Farm Witpütz WAR 258, 15 May 1963, Giess, Volk & Bleissner 6951 (WIND!); Farm Genadendal, WAR 264, 20 May 1972, Giess & Müller 12087 (PRE, WIND!); Farm Sandfontein, WAR 148, 5 Aug. 1976, Giess 14505 (PRE, WIND!); 16 km SE of gate to Farm Pioneer homestead along WarmbadOnseepkans road, 8 Sept. 2005, Kolberg & Tholkes HK1663 (K!, WIND!); Farm Aussenkehr, 13 km along German Outpost Trail (NE) from Noordoewer road, 18 Oct. 2005, Kolberg & Tholkes HK1728 (K!, WIND!); Farm Udabis, 15 Oct. 1979, Lind 482 (WIND!); Farm Sperlingspütz. No. 259, 21 Sept. 1979, Owen Smith 1209 (WIND!); Lüderitz Distr.: Aus – Rosh Pinah Road, ± 7 km from Aus, on road verge, 2 Aug. 2002, Bartsch SB890a (WIND!); Rosh Pinah. Nooitgedacht, 10 Aug. 2000, Bruyns 8332 (BOL, WIND!); Sperrgebiet, SW corner of Letterkuppe Mts, 9 Sept. 2003, Burke 03182 (WIND!); Gubub [Kubub], July 1897, Dinter 1233 (P. unguiculata holotype BM-001114681!, isotype Z000003820!); Kubub, Aus, 4 Oct. 1959, Giess 2388 (PRE, WIND!); N of Aus, quartz ridge, 28 Sept. 1975, Giess 13738 (PRE, WIND!); Farm Weissenborn LU 45, slopes of Rietrivier, 10 July 1940, Kinges 2468 (PRE, WIND!); Farm Plateau, found in the saddle in front of the farmhouse. Aus, Sept. 1963, Kräusel & Wiss 2012 (WIND!); Klein Aus Vista campsite koppies, Oct. 2002, Mannheimer CM2285 (WIND!); Paddaput, E of house, 21 Sept. 2004, Mannheimer, Maggs-Kölling & Loots CM2616 (WIND!); (Aus) Quartz foothills, 17 Sept. 2005, Mannheimer CM2820 (WIND!); Aus, found on the mountains on farm Klein-Aus, 18 Aug. 1963, Merxmüller & Giess 2968 (M, PRE, WIND!); Gross Namaland, Aus, 17 Nov. 1884, Schenck 203 (Z!); Khomas Region, Windhoek Distr.: Farm Areb-Nord REH 202, Nauchas, 30 Aug. 1972, Merxmüller & Giess 28113 (M, PRE, WIND!). HABITAT. Rocky mountains in desert, desert-dwarf shrub transition, dwarf shrub savanna, dwarf shrubland, escarpment and steppe characterised by succulent species; 500 – 1700 m. CONSERVATION STATUS. This is the most widespread species of Pteronia in Namibia with an estimated EOO at least 40-fold the maximum for a status of VU under © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 criterion B1 (IUCN 2013). Populations are fairly large and the estimated AOO also much larger than required for classiﬁcation in a threatened category. No threats are identiﬁed that negatively affect population size, further disqualifying P. unguiculata as a threatened species under the IUCN (2001) system. The Namibian conservation status therefore is LC (IUCN 2001, 2013). The South African threat status is also LC (SANBI 2011). PHENOLOGY. Flowering: March to November. Fruiting: July to January. USES. Browsed by animals. NOTES. The species seems to be closely related to Pteronia mucronata (Merxmüller 1952) but the absence of bristles on the leaves in P. unguiculata distinguishes the two species (Fig. 14); see also the discussion under P. mucronata. Specimens from the Brandberg show most similarities with P. mucronata but the leaves are not at all ciliate or pectinate. From a phytogeographical point of view these specimens would also better ﬁt into P. mucronata, another high altitude species found to the south of the Brandberg (see Map 13). In his description of the species Moore (1904) discussed the similarity of P. unguiculata and P. cylindracea (q.v.), claiming there is a clear distinction in the shape of the capitula (short, narrowly ovoid in P. unguiculata (Fig. 14), long, strictly cylindrical in P. cylindracea) and apices of phyllaries (entire or very nearly so in P. unguiculata, ciliate-lacerate in P. cylindracea). This distinction is, unfortunately, not always that clear in Namibian material. 24. Pteronia viscosa Thunb. (Thunberg 1800: 144); de Candolle (1836: 364); Harvey in Harvey & Sonder (1865: 108); Hutchinson & Phillips (1917: 306); Merxmüller (1967: 159); Merxmüller & Roessler (1984: 90); Herman (2003: 279). Type: South Africa, Western Cape Prov., Cape of Good Hope, Thunberg s.n. (holotype UPS-THUNB). Shrub, to 60 cm tall. Stems glabrous, greyish-white. Leaves opposite, thick, ﬂeshy, glabrous except for keel and sometimes lower surface that may be setulose-ciliate, lanceolate to oblong-lanceolate, keeled, 8 – 10 × 4 mm; apex slightly mucronate, base slightly connate; margins and keel bristly-ciliate. Capitula solitary, terminal, obconic, 20 – 25 × 15 mm. Receptacle honeycombed. Phyllaries multiseriate, outer broadly ovate, scaly, apex subacute, inner linear-lanceolate, membranous, apex acute; 8 – 15 mm long, rugulose distally on outside. Florets c. 15; corollas to 12 mm long, glabrous, yellow; lobes ovate-lanceolate, subacute; tube slender, slightly angular at base. Achenes compressed, obovoid, ribbed, glabrous or with a few hairs on ribs, to 7 × 4 mm. Pappus c. 12 mm long, straw-coloured. DISTRIBUTION. Africa: Namibia (uncertain — see Notes), South Africa. Map 24. KEW BULLETIN (2014) 69: 9488 SPECIMENS EXAMINED. SOUTH AFRICA. Eastern Cape Prov.: Uitenhage, Zeyher 98 (P!); Western Cape Prov.: Matjiesvalei, Drège s.n. (P-027299!); Zwart Ruggens, Drège s.n.[5661] (P-027299! to -301!); Cap de Bonne Espérance, Ecklon s.n. (P-027302!). HABITAT. Steppe dominated by a diversity of succulent species. CONSERVATION STATUS. There is only one specimen reported to be from Namibia (Pearson 3677); however, as this could not be found and occurrence in Namibia is not entirely certain, no assessment can be made of the species’ status in Namibia, which therefore has to be Not Evaluated (NE) (IUCN 2001). The South African threat status is recorded as LC (SANBI 2011). VERNACULAR NAMES. Gombossie (Afrikaans, South Africa). NOTES. According to the specimens cited by Hutchinson & Phillips (1917: 307), this species occurs in Namibia. Their cited specimen (Pearson 3677, Karas Region, Lüderitz District: Great Namaqualand, 18 km W of Aus) could, however, not be traced in NBG, K, PRE, SAM or WIND, and its occurrence in Namibia could therefore not be veriﬁed. In South Africa the species occurs in the Western and Eastern Cape Provinces, disjunct from the Namibian locality of Pearson 3677. According to Pearson (1911) he spent time at this locality on 20 February 1909. Other Pearson specimens from that time have collector’s numbers far removed from 3677 (in the 4- or 5-thousands). Specimens with numbers close to 3677 were collected far from the Aus locality or during different years. It is therefore possible that there is some error and that this species does not occur in Namibia. The Pearson locality (18 km W of Aus) needs to be searched to clarify this issue. 9488, Page 41 of 44 Pteronia species reported from Namibia but no longer belonging to the genus Pteronia aizoides Muschl. (Muschler 1911a: 102). Type: Bezirk [Distr.] of Great Namaqualand, Dinter 1388 (holotype B†; isotype K!, located in unsorted Pteronia collections and rather poor material, consisting of loose leaves and seeds with pappus only). Currently a synonym of Eremothamnus marlothianus O. Hoffm. Pteronia engleriana Muschl. (Muschler 1911a: 98, “Engleriana”); Dinter (1926: 132, “Engleriana”); Range (1935: 275, “Engleri”). Type: Bezirk [Distr.] of Great Namaqualand, Aus, in 1400 m Meereshöhe, Dinte (the -r is missing) 1107 (holotype B†; isotypes SAM-071501, K-000415078!). Currently a synonym of Amphiglossa tomentosa (Thunb.) Harv. Pteronia geigerioides Muschl. ex Dinter (1926: 132); Range (1935: 275), nom. nud. Bremer (1983: 195) considered it a synonym of Asaemia minuta (L. f.) K. Bremer, while Källersjö (1991: 39) placed it under the now accepted Athanasia minuta (L. f.) Källersjö subsp. minuta. Pteronia marlothiana (O. Hoffm.) Dinter (1926: 132); Range (1935: 275). Type: [Namibia] Namaland, Angra Pequena [Lüderitz and environs], in saxosis desertis, alt. 10 m, ﬂorif. m. Aprili 1886, Marloth 1154 (lectotype (chosen for Eremothamnus marlothianus) PRE0206236-0!, selected here; isolectotypes NBG-02002550!, SAM-0039695-0!, PRE-0594888-0!). Marloth collections from SW Africa are reported from L, M, OXF, and PRE (Vegter 1976: 504). In this particular case they were not found in (K or) L, M, or OXF, but NBG, PRE and SAM did have what are effectively isotypes. The PRE collections are considered the original set of Marloth’s material (Glen & Germishuizen 2009: 286) and hence the PRE copy of 1154 is chosen as lectotype. Currently its accepted identity is that of the basionym Eremothamnus marlothianus O. Hoffm. Pteronia minuta L. f. (Linnaeus ﬁlius 1782: 357). Type: (South Africa) Cap. bonæ spei” [Cape of Good Hope], Thunberg 18977 (holotype UPS-THUNB, selected by Bremer 1983: 195). Distribution maps presented by Bremer (1983) and Källersjö (1991) show presence in Namibia. Currently the basionym of Athanasia minuta (L. f.) Källersjö subsp. minuta. Map 24. Reported but unconﬁrmed distribution of Pteronia viscosa in Namibia. Conclusions Some uncertainties still exist on the status of some of the Namibian species of Pteronia and more specimens from outside Namibia need to be studied to clarify these issues. A revision of this largely African genus over its entire range is needed. For this synopsis many specimens were seen only as high-resolution photographs and © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 42 of 44 although these are a tremendous help, they cannot completely replace looking at the real thing. Acknowledgements The Millennium Seed Bank Partnership is acknowledged for funding of ﬁeldtrips. In Namibia the Ministry of Environment and Tourism is thanked for the plant collecting permit and the National Botanical Research Institute (NBRI), Ministry of Agriculture, Water and Forestry for use of their facilities. Patricia Craven was very helpful in obtaining additional information, literature and specimen photographs and getting some of the herbarium specimen data onto a BRAHMS database, while Tyrone Tholkes is gratefully acknowledged for assistance during ﬁeldwork. At Kew we are grateful to Juliet Beentje for her wonderful line drawings of ﬁve Pteronia species, to Dr Nicholas Hind for advice on Compositae terminology and for help with identifying suitable herbarium material for some of the line drawings, to Rafaël Govaerts for his help with some nomenclatural tangles, and to Dr Wolfgang Stuppy for his improvement of the images and maps. Furthermore, we thank Dr Dee Snijman (NBG, Cape Town) for her search for Pteronia, Amphiglossa, Athanasia and Eremothamnus specimens at SAM and NBG, Mr Paul Herman (PRE) for advice on Eremothamnus, Mr John Hunnex for ﬁnding two Pteronia holotypes in BM, and Mr Thomas Hillmann for providing a photograph of Dinter 6190 while Herbarium Hamburgense was off-line. At the herbarium G in Geneva Drs Fernand Jacquemoud and Matthieu Perret, as well as Sylviane Mura are thanked for their invaluable help in checking the many Candolle types in G-DC involved in this study and making them available through high-resolution images. HK thanks the curators of K, NBG, SAM and WIND for making their collections available for study. We thank the three reviewers for useful suggestions and improvements. References Bergius, P. J. (1767). Descriptiones Plantarum ex Capite Bonæ Spei. L. Salvius, Stockholm. Bremer, K. (1983). Taxonomy of Asaemia with notes on Stilnophyton (Compositae-Anthemideae). Nord. J. 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Boissier 4: 843. Lanjouw, J. & Staﬂeu, F. A. (1954). Index Herbariorum, Part II(1), Collectors, First instalment A – D. Reg. KEW BULLETIN (2014) 69: 9488 Veget. 2: 163. Utrecht – Netherlands: International Bureau for Plant Taxonomy and Nomenclature of the International Association for Plant Taxonomy. ____ & ____ (1957). Index Herbariorum, Part II(2), Collectors, E – H. Reg. Veget. 9: 176. Lessing, C. F. (1832). Synopsis Generum Compositarum, Berolini, Sumtibus Dunckeri et Humblotii, Paris & London. [Pteronia on pp. 195 – 196] Linnaeus, C. (1760). Plantae Rariores Africanæ. L. Salvius, Stockholm. [Thesis of Jacob Printz, presided over by Linnaeus to whom new names have to be attributed] ____ (1763a). Amoenitates academicæ, vol. 6. L. Salvius, Stockholm. [The thesis of Jacob Printz, cited as Plantae africanæ rariores, as no. 106 on pp. 77 – 115] ____ (1763b). Species plantarum (ed. 2) Vol. 2. L. Salvius, Stockholm. ____ (1764). Genera Plantarum. L. Salvius, Stockholm. Linnaeus, C. (ﬁlius) (1782). Supplementum Plantarum. Impensis Orphanotrophei, Brunsvigæ. [Title page “1781” but published April 1782] Loots, S. (2005). Red Data Book of Namibian Plants. Southern African Botanical Diversity Network Report 38: 37, 122. Lowrey, T. K. (1998). Pteronia. In: C. E. Jarvis & N. J. Turland (eds), Typiﬁcation of Linnaean speciﬁc and varietal names in the Compositae (Asteraceae). Taxon 47: 365. Merxmüller, H. (1952). Compositen Studien II. Mitt. Bot. Staatssamml. München 1: 120 – 129. ____ (1955). Pteronia. In: K. Suessenguth & H. Merxmüller (eds), Taxa praecipue africana. Mitt. Bot. Staatssamml. München 2: 67 – 83. ____ (1967). 139: Asteraceae. In: H. Merxmüller (ed.), Prodromus einer Flora von Südwestafrika. J. Cramer, Lehre. ____ & Roessler, H. (1984). Compositen-Studien XI. Neue Übersicht der Compositen Südwestafrikas. Mitt. Bot. Staatssamml. München 20: 61 – 96. Moore, S. (1902). A contribution to the Composite ﬂora of Africa. J. Linn. Soc., Bot. 35: 325 – 326. ____ (1904). Compositae. Bull. Herb. Boissier Ser. II. 4: 1011 – 1013. Muschler, R. (1911a). Compositae africanae novae. I. Bot. Jahrb. Syst. 46: 51 – 124. [Pteronia on pp. 95 – 103] ____ (1911b). LXXXVII. Vernonia De Wildemaniana Muschler, nom. nov. Repert. Spec. Nov. Regni Veg. 9: 384. Nordenstam, B. (1971). New South African Compositae. Bot. Not. 124: 10 – 14. Pearson, H. H. W. (1911). 1. On the collections of dried plants obtained in South-West Africa by the Percy Sladen Memorial Expeditions, 1908 – 1911. Ann. S. African Mus. 9: 1 – 20. Range, P. (1935). Die Flora des Namalandes. VIII. Repert. Spec. Nov. Regni Veg. 38: 256 – 280. 9488, Page 43 of 44 Roessler, H. & Merxmüller, H. (1982). Weitere Neufunde aus Südwestafrika. Mitt. Bot. Staatssamml. München 18: 187 – 200. South African National Biodiversity Institute (SANBI). (2011). SANBI Integrated Biodiversity Information System (SIBIS). Website http:// sibis.sanbi.org/faces/Home.jsp?1=1. Accessed 2 December 2012. Staﬂeu, F. A. & Cowan, R. S. (1986). Taxonomic literature. Volume VI: Sti – Vuy, p. 538. Bohn, Scheltema & Holkema, Utrecht/Antwerpen; Dr. W. Junk b.v., Publishers, The Hague/Boston. Thunberg, C. P. (1800). Prodromus Plantarum Capensium, quas, in Promontorio Bonæ Spei Africes, annis 1772 – 1775. Pars posterior. Joh. Fr. Edman, Uppsala. Turczaninow, N. S. (1851). Synanthereae quaedam hucusque indescriptae. Bull. Soc. Imp. Naturalistes Moscou 24: 59 – 95. Turland, N. J. (2013). The Code decoded. A user’s guide to the International Code of Nomenclature for algae, fungi, and plants. Koeltz Scientiﬁc Books, Königstein. Vegter, I. H. (1976). Index Herbariorum, Part II(4), Collectors, M. Regnum Veg. 93: 504. Bohn, Scheltema & Holkema, Utrecht. Index of plant names Accepted names in bold; synonyms in italics. Chrysocoma oppositifolia L. = Pteronia divaricata Less. Dicoma ramosissima Klatt = Pteronia acuminata DC. Dicoma seitziana Dinter = Pteronia eenii S. Moore Eupatorium divaricatum P. J. Bergius = Pteronia divaricata Less. Pteronia acuminata DC. Pteronia acuta Muschl. Pteronia aizoides Muschl. = Eremothamnus marlothianus O. Hoffm. Pteronia anisata B. Nord. Pteronia anisata Dinter ex Merxm., nom. nud. = Pteronia glabrata L. f. Pteronia arcuata Dinter = Pteronia glauca Thunb. Pteronia beckeoides auct. non DC. = Pteronia lucilioides DC. Pteronia bromoides S. Moore = Pteronia lucilioides DC. Pteronia cancellata Dinter, nom. nud. = Pteronia pomonae Merxm. Pteronia candollei Harv. = Pteronia glauca Thunb. Pteronia carnosa Muschl. (Muschler 1911a: 95) non P. carnosa Muschl. (Muschler 1911a: 97) = Pteronia glabrata L. f. Pteronia carnosa Muschl. (Muschler 1911a: 97) non P. carnosa Muschl. (Muschler 1911a: 95). = Pteronia acuminata DC. Pteronia chlorolepis Dinter = Pteronia sordida N. E. Br. Pteronia ciliata Thunb. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 9488, Page 44 of 44 Pteronia ciliata Thunb. var. ecklonis Harv. = Pteronia ciliata Thunb. Pteronia ciliata Thunb. var. subtrigona DC. = Pteronia ciliata Thunb. Pteronia ciliata Thunb. var. thunbergii Harv. = Pteronia ciliata Thunb. Pteronia cylindracea DC. Pteronia dinteri S. Moore = Pteronia mucronata DC. Pteronia divaricata Less. Pteronia eenii S. Moore Pteronia engleriana Muschl. = Amphiglossa tomentosa (Thunb.) Harv. Pteronia feddeana Muschl. = Pteronia acuminata DC. Pteronia feldtmanniana Dinter ex Merxm., nom. nud. = Pteronia eenii S. Moore Pteronia ﬂexicaulis auct. non L. f. = Pteronia paniculata Thunb. Pteronia geigerioides Muschl. ex Dinter, nom. nud. = Athanasia minuta (L. f.) Källersjö Pteronia glabrata L. f. Pteronia glabrata auct. non L. f. = Pteronia glauca Thunb. Pteronia glabrata L. f. var. succulenta (Thunb.) Merxm. = Pteronia glabrata L. f. Pteronia glauca Thunb. Pteronia glauca Thunb. subsp. arcuata (Dinter) Merxm. = Pteronia glauca Thunb. Pteronia glomerata, auct. non L. f. = Pteronia sordida N. E. Br. Pteronia gymnocline auct. non DC. = Pteronia lucilioides DC. Pteronia inflexa L. f. Pteronia kingesii Merxm. = Pteronia polygalifolia O. Hoffm. Pteronia latisquama DC. = Pteronia glauca Thunb. Pteronia leucoclada Turcz. Pteronia lucilioides DC. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2014 KEW BULLETIN (2014) 69: 9488 Pteronia lucilioides DC. var. sparsifolia Harv. = Pteronia lucilioides DC. Pteronia lupulina DC. = Pteronia inflexa L. f. Pteronia lupulina DC. var. rotundifolia DC. = Pteronia inflexa L. f. Pteronia lycioides Muschl. ex Dinter = Pteronia scariosa L. f. Pteronia marlothiana (O. Hoffm.) Dinter = Eremothamnus marlothianus O. Hoffm. Pteronia minuta L. f. = Athanasia minuta (L. f.) Källersjö subsp. minuta Pteronia mucronata DC. Pteronia mucronata DC. subsp. dinteri (S. Moore) Merxm. = Pteronia mucronata DC. Pteronia onobromoides DC. Pteronia paniculata Thunb. Pteronia polygalifolia O. Hoffm. Pteronia pomonae Merxm. Pteronia quadrifaria Dinter Pteronia quinquecostata Dinter = Pteronia polygalifolia O. Hoffm. Pteronia rangei Muschl. Pteronia roesemanniana Dinter ex Merxm., nom. nud. = Pteronia lucilioides DC. Pteronia scariosa L. f. Pteronia sesuviifolia DC. = Pteronia glabrata L. f. Pteronia sordida N. E. Br. Pteronia spinulosa E. Phillips Pteronia succulenta auct. non Thunb. = Pteronia glabrata L. f. Pteronia thymifolia Muschl. & Dinter = Pteronia glauca Thunb. Pteronia turbinata DC. = Pteronia ciliata Thunb. Pteronia unguiculata S. Moore Pteronia villosa auct. non L. f. = Pteronia pomonae Merxm. Pteronia viscosa Thunb.

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