Pentzia pinnatifida var. chenoleoides Hutch.

Pentzia pinnatifida var. chenoleoides Hutch. in Kew Bull. 10: 241 (1916). —TYPE: SOUTH AFRICA. Limpopo, Tzaneen Natal (2330): Near Murchison (–DC), 1884/5, Wood 3110 (lectotype: K–image!, designated in Hilliard and Burtt (1973); isolectotype: NH–image!).

Athanasia villosa Hilliard View in CoL in Hilliard and B.L. Burtt, Notes Roy. Bot. Gard. Edinburgh 32(3): 330 (1973); Hilliard, Compositae in Natal : 342 (1977), syn. nov. Phymaspermum villosum (Hilliard) Källersjö View in CoL , in Nord. J. Bot 5(6): 535 (1986) ; K. Bremer and Humphries , in Bull. Nat. Hist. Mus. Lond. (Bot.) 23(2): 94 (1993) ; Germish. and Meyer, Pl. S. Afr.: 274 (2003) ; Klopper et al. Checklist of the flowering plants of Sub- Saharan Africa: 147 (2006) ; Germish. et al. A Checklist S. African Pl.: 243 (2006). —TYPE: SOUTH AFRICA. Eastern Cape, Kokstad (3029): Weza , above Mpetyne forest (– DA), 05 Jan 1964, Hilliard 2507 (holotype: NU!; isotypes: E–image!, K–image!, M–image!, NH–image!, PRE!, S –image!) .

Multistemmed, densely leafy shrub, 0.3–1.8 m high. Stems unbranched to few-branched; branches erect, glabrous or scabrous to silvery villous; fascicles present in leaf axils. Leaves alternate, spreading, 6.0–40.0 × 0.5–1.0 mm, linear when undissected, mucronate, sometimes involute, entire (usually on upper branches) to deeply pinnatifid, narrowed into petiole-like base when dissected, sometimes with secondary basal lobes, glabrous to silvery villous on both surfaces, leathery; basal swelling present, continuous with stem ribs; lobes 3–9, 2.0–24.0 × 0.5–1.0 mm, linear, mucronate. Capitula discoid, homogamous, terminal, few- to many-headed (5 to +100), simple or compound corymbs, pedunculate; peduncles 8.0–42.0 mm long, tomentose. Involucre 3.0–6.0 × 1.5–3.0 mm, narrowly campanulate to cylindrical, tapering at the base; involucral bracts 3- or 4-seriate, loosely to tightly arranged, margins and apices scarious, membranous apices of inner bracts ± 0.5 mm long, glabrous to villous, brown margins absent, median resin canals present; outer bracts lanceolate, 1.4–2.5 mm long, acute to acuminate; middle bracts lanceolate, 2.4–3.5 mm long, acute to acuminate; inner bracts lanceolate to oblong, 3.3–4.2 mm long, rounded. Receptacle convex; paleae in marginal series (rarely rudimentary inner paleae present), oblong to linear, 4.0– 4.8 mm long, acute, rigid with thick scarious margins and apices, scarious apices ± 0.6 mm long. Disc florets 3–18, sometimes with resin canals in the tube, limb and lobes; corolla 2.3–2.6 mm long, with only glandular trichomes, yellow; tube 1.0– 1.5 mm long; limb campanulate, 0.8–1.2 mm long (excluding lobes); lobes spreading, 0.6–0.7 mm long, triangular. Anthers 2.5–2.8 mm long (including apical appendage); apical appendage oblong. Style 2.0– 2.8 mm long (excluding branches); branches 0.5–0.6 mm long. Cypselas ca. 2.0 × 0.5 mm, oblong, 13- to14-ribbed, apical rim thickened, entire to slightly dentate, light to dark brown, glandular trichomes absent, discontinuous resin canals in ribs present. Figure 20 View FIG .

Diagnostic Characters — Phymaspermum acerosum shares the narrowly lobed pinnatifid leaves, the absence of myxogenic trichomes on the cypselas and the presence of prominent resin canals in the cypsela ribs with P. pinnatifidum . It can be distinguished by its narrowly campanulate to oblong involucres and the absence of a prominent apical crown on the cypselas (only at most a thickened apical rim which can be entire to shortly dentate). Phymaspermum acerosum is also a much larger more robust shrub growing to more than 0.5 m tall with much thicker and woodier stems than the smaller P. pinnatifidum which tends to have much more slender and herbaceous stems.

The previously recognised species P. villosum was at first distinguished from P. acerosum by having much broader oblong and hairy involucres. Phymaspermum acerosum was considered to only have narrowly campanulate involucres however, after many specimens were examined, P. acerosum is now considered to have involucres which vary from narrowly campanulate to broad and oblong and P. villosum is now a synonym of P. acerosum .

Distribution and Ecology — Phymaspermum acerosum is widely distributed from Port Elizabeth in the Eastern Cape Province to Noodsberg in the Limpopo Province ( Fig. 20 F View FIG ) where it occurs in grassland and forest margins at an altitude

of 60–2,200 m. It flowers throughout the year, with a peak between April and July.

Additional Specimens Examined — SOUTH AFRICA. Gauteng: 2528 (City of Tshwane): Doornpoort (–CB), Mogg 2161 (PRE). Mpumalanga: 2430 (Pilgrims Rest): Ohrigstad Nature Reserve (–DC), Theron 3633 (PRE), Theron 3607 (PRE), Jacobsen 2903 (PRE); Morgenzon Nature Reserve, near Graskop (–DC), Reid 3 (PRE); Mount Sheba Nature Reserve (–DC), Kerfoot 8757 (PRE); near Graskop (–DD), Phillips 3491 (PRE); Graskop, above Kowyns Pass, along road (–DD), Koekemoer 2123 (PRE); near Mac-Mac Falls (–DD), Burtt Davy 438 (PRE, BOL). 2530 (Mashishing): 8 miles from Pilgrims Rest on Mashishing road (–AB), Story 1215 (PRE); Mashishing (–AB), Burtt Davy 1165 (BOL), Wilms 791 (BM–image), Wilms 790 (BM– image, NU); Mashishing, near turn off to Mount Anderson from Long Tom Pass (–BA), Koekemoer 2280 (PRE); Pilgrims Rest, between Mashishing and Sabie (–BA), Schyff 5465 (PRE); Between Mashishing and Sabie (–BA), Pole Evans 3995 (PRE); Hartebeesvlakte (–BA), Mohle 20 (PRE); Sabie valley (–BB), Galpin 39 (BOL); Rosehaugh (–BD), Rogers 20377 (PRE), Sim 546 (BOL); Entokozweni (–CB), Young 1476 (PRE); Emakhazeni near Entokozweni (–CA), Codd 7759 (PRE); Helopmith, Kaapse Hoop road, 12 miles out (–DB), Liebenberg 2553 (PRE). 2531 (Komatipoort): Barberton (–CC), Williams 10512 (PRE); 7 miles NW of Havelock Mine, Barberton (–CC), Codd 7846 (PRE). 2730 (Vryheid): Hoedjie, Farm Suikerhoek, Wakkerstroom (–AB), Devenish 1698 (NU). Free State: 2828 (Bethlehem): Qwa Qwa National Park Avondrust II (–BC), Zietsman 3349 (PRE). 2829 (Harrismith): Harrismith Botanic Garden (–AC) Zeyde s.n. (NBG); Rensburgskop, 7 km van Swineburne (–AC), Jacobsz 428 (PRE); Farm Gravolette, 16 miles S of Harrismith (–AC), Zeyde s.n. (NBG); Sterkfontein Dam (–AC), Blom 162 (PRE). Kwazulu Natal: 2730 (Vryheid): Mooihoek, Piet Retief district (–AC), Devenish 1638 (NU); Utrecht, on road to Wakkerstroom to Paulpietersburg, Mooifontein farm, Guquka, south hills of Pongola mountain series (–AC), Jordaan 2815 (PRE); Nauwhoek, Utrecht (–AD), Devenish 1133 (PRE two sheets), Devenish 490 (PRE); Wakkerstroom district, Guquka farm, 25 km from Wakkerstroom on the PaulPolokwane road (–AD), Meyer 218 (PRE); south side of Dumbe Mountain (–BD), Acocks 11542 (PRE); Vryheid, 12 miles from Natal Spa on Vryheid- PaulPolokwane road (–DB), Ross 1251 (NU); Bloemhof, Emnyathi (–DD), Thode 2962 (NBG); Leeuwnek Pass, Emnyathi Mountain (–DD), Shirley (NU two sheets). 2731 (Louwsburg): 2 miles from Vryheid on road to Louwsburg on north facing slope (–CA), Hardy 37 (PRE); 19 miles from Vryheid on Louwsberg road, Nkongolwana River (–CA), Ross 1224 (NU); Ngotshe, Itala Nature Reserve (–CB), Hilliard and Burtt 10018 (NU), Hilliard and Burtt 8546 (NU); Louwsberg (–CB), Compton 19702 (NBG); Ngotshe district, Ngome (–CD), Hilliard and Burtt 9849 (NU); Ngotshe, 7 miles W of Ngome (–CD), Codd 9580 (PRE two sheets); Ngome Forest, about 10 km from main office of forestry on road to Vryheid (–CD), Germishuizen 2103 (PRE). 2829 (Harrismith): Inkupe Hill in the Biggarsberg (–BB), Acocks 10465 (PRE); Cathedral Peak Reserve, Mikes Pass (–CC), Magee et al. 460 (NBG). 2830 (Dundee): Helpmekaar/Elandskraal road (–AD), Hilliard 1575 (NU), Shirley s.n.(NU); Nqutu (–BA), Codd 1363 (PRE); hill on east side of Zungeni Peak (–BD), J.P.H. Acocks 11553 (PRE); Pomeroy district Jobskop on summit plateau (–CA), Venter 1775 (PRE); Pangolo mountains to Kaffir drift (–CA), Thode A327 (PRE); 12 miles Muden/Weenen road (–CC), Moll 3244 (PRE two sheets, NU); Umhlumba mountain top (–CC), Acocks 13864 (PRE); Qudeni-Nqutu road (–DB), Edwards 2249 (NU). 2831 (Nkandla): Mahlabatini, Ceza forest (–AB), Hilliard and Burtt 3312 (NU); Lower part of Nkandla Forest, 14.7 miles from Nkandla Hotel on road to Eshowe (–CC), Winter 8261 (PRE). 2832 (Mtubatuba): Hlabisa, Hluhluwe Game Reserve (–AA), Ward 2620 (PRE, NU). 2929 (Underburg): 17 km E of Mooi river on Hlatikulu road, Farm Lanner Veane (–BA), Balkwill and Manning 468 (NU); Central Drakensberg, Highmoor forestry station (–BC), Breytenbach 5072 (PRE); Estcourt district side of Kamberg Nature Reserve, Rosette road (–BC), Wright 176 (NU); Impendhle district, Loteni area (–BC), Roberts s.n. (NU). 2930 (Pietermaritzburg): Karkloof (–AC), Rehman 7417 (Z–image), TL 1139 (BOL); below Bridle Road view site, Karkloof Nature Reserve (–AC), Styles 25 (NU); Hgononi Veld near Michaelhouse (–AC), Pentz and Acocks 10313 (PRE); Mount Park Dargle, Lions River (–AC), Johnson 433 (BOL); Lions River, The Dayle (–AC), Hilliard 2157 (NU); Lions River, Karkloof, near turnoff to farm Klatine (–AC), Hilliard 4857 (NU); 4 miles from Currys Post/ Nottingham road, Lions River (–AC), Moll 886 (NU); 25 km NNE Howick, Twin Falls (–AC), Grove 51 (NU); Lions River, Curry’ s Post, Mehleys Bush (–AC), Hilliard 4853 (NU); Lions River, E of Karkloof Gorge (–AC), Parry 18 (NU); Whitecliffe, Greytown (–BA), Wood 889 (SAM); De Rust, Greytown (–BA), Thode 2955 (NBG); Greytown (–BA), Wylie 34065 (PRE), Wylie 21927 (PRE); “Mowhay” Ahrens (–BB) Fisher 463 (NU); Lions River, Silverdale (–CB) Wright 1502 (NU); Swartkops Hill (–CB), Esterhuysen 20,294 (BOL); Writers Kloof (–CB), Carnegie 1216 (BOL); Chase Valley (–CB), Canham 28 (NU); World’ s View (–CB), Moll 2647 (NU); along roadside above Sweet Waters, on the way to Hilton (–CB), Warren 5 (NU); Town Bush (–CB), Randles 29 (NU); Byrne, Keerom (–CC), Strey 11268 (NU); Richmond, Byrne Valley (–CC), Schofield s.n. (NU), Galpin s.n. (BOL); Richmond (–CD), Medley Wood 9844 (NU two sheets); Richmond, Enon Estate (–CD), Hilliard 2037 (NU); Drummond (–DA) Bromhead s.n.(NU), Camperdown, Drummond (–DA), Camden Smith 32 (NU); Table Mountain (–DA), Killick 628 (NU), Johnstone 195 (NU); Ndwedwe, Inanda (–DB), Wood 921 (BOL), Hilliard 2052 (NU); Inanda (–DB) Wood s.n. (Z–image); Near Summerveld Equine Hospital Assagay, Ethekwini Municipality (–DC), Wragg 671 (NU); Hammarsdale area, Hector (ESKOM) substation site (–DC), Ward 13017 (NU); Camperdown, Nagle Dam (–DC), Wells 1543 (NU); M. Stainbanks farm, Mid Illovo (–DC), Young 1087 (NU); Summerveld part of Assagay (–DC), Wragg 88 (NU); Krantzkloof (–DD), Schlechter 3182 (BOL), Clarkson 29 (NU), Galpin s.n. (NBG), Galpin 12078 (PRE); Pinetown, Everton, Eskotene (–DD), Hilliard 1501 (NU), Hilliard 4865 (NU-3); Gillitts area, Stockville Valley, Farm Stockville 1382 (–DD), Ward 11272 (NU, PRE); Pinetown, Mbilo River Valley, below Paradise Valley Nature Reserve (–DD), Ward 10956 (NU); Hillcrest (–DD) Neuwoudt 31 (NU), Ross 336 (NU); Marianhill (–DD), Strey 9193 (NU, PRE); Fields Hill (–DD), Rehman 7977 (Z–image). 3030 (Port Shepstone): Farm Fairfield, S of Dumisa on SAPPI property (–AB), Balkwill 10284 (B–image); Hlokozi, Alexandra city (–AD), Rudatis 2962 (NBG); Umgai (–AD), Strey 10974 (PRE,NU); Umbogintwini (–BB) Brooks 7 (NU); Vernon Crookes Nature Reserve (–BC), Ward 9886 (NU); Oribi Flats, top of Oribi Gorge (–CA), Mc Clean 579 (PRE); boundary between Alfred and Port Shepstone escarpment above Oherburn (–CA), Hilliard 1638 (NU); Port Shepstone (–CB), Dinock Brown 405 (BOL); Near Port Shepstone on St. Faith’ s road (–CB), Hafström and Acocks 1855 (PRE); Above Mthamvuna River, borderland Estate Farm (–CC), Nicholson 1473 (PRE); Mthamvuna River (–CC), Nicholson 1014 (PRE); Zuurberg (–CC), Tyson 2162 (BOL, SAM), Tyson 1060 (SAM); ‘Umgeni ad cattarhactam’ (–CD), Rehman 7472 (Z–image). 3130 (Port Edward): 5 miles from Port Edward on Izingolweni road (–AA), Hilliard 1130 (NU); Mthamvuna Nature Reserve 20 km NW of Port Edward on Izingolweni road (–AA), Nicholson 2249 (PRE); Mthamvuna Nature Reserve, hidden valley, Farm Clearwater, near cottages (–AA), Hilliard and Burtt 10246 (NU), Jordaan 3684 (PRE), Balkwill 382 (NU), Germishuizen 8083 (PRE). EASTERN CAPE: 3028 (Matatiele): Maclear, Little Pot River (–CC), Hilliard 3899 (BOL, NU). 3029 (Kokstad): Insizwa, Mount Ayliff (–CD), Strey 10759 (NU), Hilliard and Burtt 6531 (NU, PRE two sheets); Insizwa roadside bank (–CD), Strey 11250 (NU, PRE); Bulembu, forest marqin (–DA), Kotze 600 (PRE); Alfred district, Mpetyn forest, Weza (–DA), Hilliard 2503 (NU); Ntlenzi kloof forest (–DC), Strey 8689 (NU, PRE); 5 miles from Bizana, on Kokstad road (–DD), Acocks 10953 (PRE); Bizana division 5–7 miles E, near Umzamba River Mouth (–DD), Lewis 4652 (SAM). 3128 (Mthatha): Roma (–CA), Schmitz 9062 (PRE); Bhaziya Mountain (–CB), Baur 185 (SAM); Misty Mountain, approx 8–9 miles from Mthatha (–DB), Sole 1 (NBG). 3129 (Bizana): Pondoland (–AB), Bowker 5569 (SAM). 3226 (Fort Beaufort): Alice, Pefferskop (–DB), Acocks 8980 (PRE). 3227 (Stutterheim): Keiskammahoek (–CA), Story 3487 (PRE); Dohne Hill (–CB), Sim 78 (NU), Sim 77 (NU); Perie, Kaffraria (–CC), Sim 19736 (NU, PRE); King William Town (–CD), Tyson 5561 (SAM); near Komsha (–DB), Flanagan 57 (PRE); Elliot 99 (PRE). 3228 (Butterworth): Kei River (–CA), Flanagan 1476 (NBG), MacOwen 1476 (K–image); West of Kei Mouth (–CB), Flanagan 57 (SAM); Centane valleys (–CB), Pegler 472 (BOL). 3325 (Port Elizabeth): Sydenham (–DC), Medley-Wood 12427 (NU).

PRECISE LOCALITY UNKNOWN: Natal, Badenhuizen and party J28934 View Materials (NU), Wood 268 (BOL, SAM), Drège 16936 (SAM), Cooper 1139 (K–image, BM–image); Prope Macamae, Leu 157 (BOL); Entumani Zululand, Medley Wood 3969 (SAM); Zululand, Gerrard 1038 (BM–image); Plant 100 (TCD–image, S–image).

SWAZILAND. 2631 (Mbabane): Mucuceni hills (– AA), Compton 31524 ( PRE, NBG). 2531 (Komatipoort): Devils Bridge, Piggs Peak (– CD), Compton 30044 ( NBG).

LESOTHO. 2828 (Bethlehem): Lelingvana Basutoland, Leribe, Mealavaneng (– CC), Dieterlin 822 ( BOL, PRE two sheets, SAM), Dieterlin 881 ( PRE, SAM); Witzieshoek (– DB), Thode 5621 ( NBG).

ACKNOWLEDGMENTS. The curators and staff of the cited herbaria are thanked for their kind hospitality and assistance during visits and for making specimens available on loan. Mr. Nick Helme is thanked for bringing his collection of what turned out to be a new species from around Robertson to our attention and the landowner, Mr. Ernst Bruwer, for allowing us to collect type material and study the plant in situ. Dr. AnneLise Scutte-Vlok and Mr. Jan Vlok are thanked for suppling us with material and images of P. aphyllum . Dr. Laurent Gautier and herbarium staff from Conservatoire Botanique de la Ville de Genève are thanked for supplying electronic images of type material. Cape Nature, the Eastern Cape Department of Economic Development, Enviromental Affairs and Tourism, and Ezemvelo KZN Wildlife are thanked for providing the relevant permits. This work is based upon research supported by the National Research Foundation and Department of Science and Technology under the SABI Programme and the University of Johannesburg.