Phymaspermum athanasioides, (S. Moore) Kallersjo

12. PHYMASPERMUM ATHANASIOIDES (S.Moore) 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); Retief and Herman, Pl. N. Prov. S. Afr.: 330 (1997); Germish. and N.L. Meyer, Pl. S. Afr.: 274 (2003); Klopper et al. Checklist of the flowering plants of Sub- Saharan Africa: 147 (2006); Germish. et al. Checklist S. African Pl.: 243

Multistemmed, densely leafy shrub, 0.2–2.0 m high. Stems much-branched; branches erect, grey-pubescent; fascicles never present in leaf axils. Leaves alternate, spreading, 6.0– 40.0 × 0.8–2.0 mm, linear to oblong, acute to mucronate, entire to trifid, involute, narrowed into petiole-like base, without secondary basal lobes, glabrous on both surfaces, leathery; basal swelling present, continuous with stem ribs; lobes 2–3, 0.5–10.0 × 0.5–1.0 mm, linear to oblong, acute to mucronate. Capitula disciform, heterogamous, terminal, many-headed ± 30, simple to compound corymbs, pedunculate; peduncles 4.0–35.0 mm long, glabrous to white tomentose. Involucre 4.0–8.0 × 3.5–8.0 mm, hemispherical, cyathiform to urceolate, sometimes tapering at the base; involucral bracts 4- or 6-seriate, with the peduncular bracts usually being the first two series, tightly arranged, margins and apices scarious, membranous apices of all bracts 0.3–1.2 mm long, glabrous to tomentose especially along the edges, brown margins absent, median resin canals present; outer bracts lanceolate, 2.0–3.0 mm long, acute; middle bracts deltoid to lanceolate, 2.2–4.2 mm long, acute; inner bracts deltoid to lanceolate, 2.2–4.3 mm long, acute to rounded; second inner bracts deltoid to lanceolate, 3.8– 4.6 mm long, acuminate to rounded. Receptacle convex; paleae in marginal series, oblong to linear, 3.2–3.7 mm long, acute to attenuating, scarious. Filiform florets ± 20; corolla 2.3–3.0 mm long, with glandular trichomes, zygomorphic; adaxial lobes 2, 0.7–1.0 mm long, triangular, spreading, abaxial lamina 0.3– 0.6 mm long, linear, 3-dentate. Disc florets ±100, without resin canals; corolla 2.2–3.0 mm long, with only glandular trichomes, yellow; tube 1.0–2.0 mm long; limb narrowly campanulate, 0.8–1.3 mm long (excluding lobes); lobes erect to spreading,

0.4–0.6 mm long, triangular. Anthers ± 3.0 mm long (including apical appendage); apical appendage rounded to oblong. Style 2.5–3.1 mm long (excluding branches); branches 0.4–0.7 mm long. Cypselas 2.1–2.2 × 0.5 mm, obovate, 10- to12-ribbed, apical rim thickened to membranous, entire to dentate, glandular trichomes present, sparse, scattered, not mucilaginous when soaked, discontinuous resin canals in ribs absent. Figure 15 View FIG .

Diagnostic Characters — Phymaspermum athanasioides was previously considered to be a part of a complex with P. bolusii and P. montanum , distinguished by apparent differences in their involucral bracts. However, upon further study, these characters were found to be highly variable and no clearly recognisable characters could be found to distinguish between them. As a result only a single variable species, P. athanasioides , is recognised here. It is readily distinguished from all other species in the genus by the marginal filiform florets and from the other species with corymbosely arranged capitula by the absence of fascicles in the leaf axils and by the mostly entire leaves, which rarely become 2- or 3-fid.

Distribution and Ecology — Phymaspermum athanasioides is widely distributed from around Harrismith in KwaZulu Natal to Musina in Limpopo and extending into Swaziland and further north into Zimbabwe ( Fig. 15G View FIG ). It grows in mixed bushveld and grassland, especially on quartzite or sandstone, at an altitude of 1,000 m to 1,750 m. Flowering time is throughout the year but especially in August, hence its vernacular name August Gold.

Additional Specimens Examined — SOUTH AFRICA. Limpopo: 2230 (Musina): Soutpansberg, Entabeni Forest Reserve (–CC), Codd 3038 (PRE); Venda, Musekwa location (–CC), Du Toit 298 (PRE); Hangklip, Louws Kraal, Waterberg Transvaal (–CC), Maguire 1426 (NBG). 2329 (Polokwane): Soutpansberg Mountain, Letsume at foot of cliffs (–AB), Venter 6233 (PRE), Rogers 19904 (PRE); Vivo, Farm Llewellyn 35 on Soutpansberg (–AB), Venter 10,755 (PRE); Makhado (–BB), Breyer 24364 (PRE); Blaauwberg, south slopes at plateau (–CD), Esterhuysen 21,435 (BOL); Tomazon farm (–DD), Nienaber 26 (PRE); Haenertsburg, Iron Crown Mountain (–DD), Venter 10.855 (PRE). 2330 (Tzaneen): Polokwane, Wolkberg (–CC), Van der Merwe 33 (PRE). 2429 (Zebediela): Mokopanet district, summit of highest peak, Zebedila mountains (–AA), Galpin s.n. (BOL); 40 km NE of Mokopane, Snymansdrift (–AA), Maguire 2997 (B–image). 2430 (Pilgrims rest): Shiluvane (–AB), Junod 13887 (PRE); Lekgalameetse Nature Reserve (–AB), Stalmans 83 (PRE); Lekgalameetse Nature Reserve, Balloon/mountain 1493 m, shrubland on cliff face (–AB), Stalmans 1340 (PRE). North West: 2526 (Zeerust): On road between Koster and Lichtenburg, Mabaalstat (–DC), Vahrmeyer 1438 (PRE); 13 miles from Koster on Mabaalstad road (–DC), Story 5642 (PRE). Gauteng: 2527 (Rustenburg): Rustenburg (–BA), Collins 23957 (PRE), Galpin s.n. (BOL); Magaliesberg nature area (–DC), Ortiz and Anderson 534 (PRE). 2528 (City of Tshwane): Magaliesberg, Vissershoek, upper slopes on top of range (–CA), Repton 1292 (PRE). 2627 (Potchefstroom): Witpoortjie Falls, Roodekrantz 83, Krugerdorp, 20 miles NNE of Johannesburg, Orange Grove (–BB), Mogg 21,326 (PRE); 3 km N of Krugersdorp (–BB), Codd 3149 (PRE); Paardeburg 20 miles from Johannesburg (–BD), Tufrosis 1017 (NBG). 2628 (Johannesburg): Johannesburg west koppies (–AA), Laub 3977 (SAM); Elsburg (–AA), Rogers 12143 (BOL); Jeppestown ridge (–AA), Gillfillan 6026 (K–image); Johannesburg (–AA), Moss 19206 (PRE); Witwatersrand, Hospital Hill, Orange Grove (–AA), Bryant G6 (BOL), Moss 5072 (BOL), Cowrath 405 (K–image); Kensington (–AA), Gerstner 6427 (PRE); Melville Koppies central, along path 100 metres from lapa near Kafue Road entrance (–AA), Magee 394 (NBG); Heidelberg (–AC), Schlechter 3517 (BOL). Mpumalanga: 2430 (Pilgrims Rest): Lulu mountains, farm Grootvygenboom (–CA), Mogg 16936 (PRE); The Downs, near Tzaneen (–DA), Rogers 20101 (BOL), Nel 103 (NBG); Mount Sheba estate (–DC), Boucher 1836 (NBG), Kerfoot 8755 (PRE); Morgenzon Reserve (–DC), Mohle 407 (PRE); Motlatserivierpoort Nature Reserve, Muilhuis (–DB), Zietsman 3771 (PRE). 2530 (Mashishing): Between Lydenberg and Sabie, 12 miles from Lydenberg, Long Tom Pass (–AB), Balsinhas and Kersberg 2111 (PRE), Unknown 280 (PRE); Mashishing, upper Erica knoll trail, Buffelskloof Nature Reserve (–BC), Burrows 8158 (PRE); Buffelskloof Nature Reserve, on plateau overlooking a gorge, along edge (–BC), Koekemoer 2091 (PRE); Pilgrims Rest (–BC), Rogers 14171 (BOL); Woodbush (–CC), Jenkins 13960 (PRE); Next to Berlin offramp on Kaapsehoop (–DB), Mothogoane NB0019 (PRE); Kaapse Hoop escarpment (–DB), Nel 43 (NBG), Rogers 19504 (PRE), Rogers 21414 (BOL); Near Cythna Letty Nature Reserve (–DD), Glen 1329 (PRE); 2531 (Komatipoort): Moederlief 109 JU 450m S of Mpageni beacon (–CA), Deall 3883 (PRE); Louw’ s Creek Timbers 3 km N of offices (–CB), Kluge 2439 (PRE); SE of Louws Creek. S.A.T.I.C.O forestry station, Bearded Man Peak (–CB), McMurtry 12837 (PRE); Three Sisters mountains, SE of Kaapmuiden (–CB), Meyer BP00224 (PRE); Songimvelo Game Reserve, on the farm Vooruitzicht, Malndweni, Panhandle, en route back to house (–CC), Balkwill, McCallum and Reddy 11990 (B–image); 6 miles E of Barberton on road to Havelock mine (–CC), Codd 1603 (PRE); Along firebreak from Lone Tree Hill to Saddleback Hill (–CC), Reid 1801 (PRE); Duivels Kantoor (–CC), Thode a1612 (PRE); on hill above Angle Station (–CC), Venter 4485 (PRE). 2630 (Carolina): Carolina (–AA), Rogers 19738 (PRE). Free State: 2627 (Potchefstroom): Vredefort, Dwarsberg (–CD), Du Preez 1735 (PRE); Vredefort dome, 20 km WNW of Parys, Farm Buffelskloof 511 IQ, hilly-slopes NNW of Thabela Thabeng resort (–CD), Kroon 16236 (PRE); Witkop (–DC), Louw 1881 (PRE). 2728 (Steve Vukile Tshwethe): Soutpansberg mountains, Frans Hoek Peak (–BA), Galpin s.n. (BOL). KwaZulu Natal: 2829 (Harrismith): Oliviershoek Pass, Irenesdale farm (–CA), Klapwijk 184 (PRE).

SWAZILAND. 2631 (Mbabane): Komati Bridge (– AA), Bayliss 1541 ( PRE); Ukutula (– AC), Compton 25113 ( PRE); Mbuluzi falls (– AC), Compton 25134 ( PRE); hill west of Mbabane (– AC), Compton 25844 ( PRE); Black Mbuluzi valley (– AC), Compton 25933 ( NBG); Fonteyn (– AC), Compton 25975 ( NBG, PRE); Forbes Reef (– AC), Compton 31675 ( NBG); Mdimba Mountain, Manzini (– AC), Dlamini s.n. ( PRE); Kangwane mountain grassland (– AC), Loffler 1182 ( NBG); Manzini district tea road, 6 km from junction (– BC), LaCroix 4924 ( PRE).

ZIMBABWE. 1832 (Mutare): Rhodes Inyanga experiment station, Inyanga National Park (– BD), Burrows 757 ( NBG). 1932 (Melsetter): road from Vumba to Umtali, Rhodesia (– BB), Morris 346 ( BOL).

13. PHYMASPERMUM WOODII (Thell.) Källersjö View in CoL in Nordic J. Bot. 5(6): 535 (1986); K. Bremer and Humphries in Bull. Nat. Hist. Mus. Lond. (Bot.) 23(2): 94 (1993); Retief and Herman, Pl. N. Prov. S. Afr.: 330 (1997); Germish. and N.L. Meyer, Pl. S. Afr.: 274 (2003); Klopper et al. Checklist of the flowering plants of Sub- Saharan Africa: 147 (2006); Germish. et al. Checklist S. African Pl.: 243 (2006). Pentzia woodii Thell. in Vierteljahrsschr. naturf. Ges. Zürich. 61: 454 (1916). Athanasia woodii (Thell.) Hilliard in Hilliard and B.L. Burtt, Notes Roy. Bot. Gard. Edinburgh 32 (3): 331 (1973); Hilliard, Compositae in Natal: 339 (1977). —TYPE: SOUTH AFRICA. KwaZulu Natal, Pietermaritzburg (2930): Curries Post (– AC), 24 Apr. 1891, Wood 4457 (lectotype: Z –image!, designated here; isolectotype: BOL!, NH –image!) [Note: The specimen in Z, where Thellung was based is selected here.]

median resin canals present; outer bracts deltoid to lanceolate, 2.2–3.3 mm long, acute to acuminate; middle bracts deltoid to ovate, 2.7–3.4 mm long, acuminate; inner bracts deltoid to lanceolate, 2.7–4.0 mm long, acuminate to rounded; second inner bracts ovate, 3.0–5.0 mm long, rounded; innermost bracts ovate, 4.3–5.8 mm long, rounded. Receptacle convex to flat; paleae in two marginal series, linear to oblong, 4.3–6.0 mm long, rounded, rigid with scarious margins and apices, scarious apices 0.8–1.1 mm long. Disc florets 37–55, without resin canals; corolla 2.3–3.2 mm long, with only glandular trichomes, yellow; tube 1.4–1.8 mm long; limb narrowly campanulate, 1.0– 1.4 mm long (excluding lobes); lobes spreading, 0.6–1.0 mm long, narrowly triangular. Anthers 2.7–3.8 mm long (including apical appendage); apical appendage oblong. Style 2.0– 3.3 mm long (excluding branches); branches 0.5 mm long. Cypselas 1.8–2.0 × 0.5 mm, oblong to obovate, 11-ribbed, apical rim thickened, entire to slightly dentate, glandular trichomes present, sparse, scattered, not mucilaginous when soaked, discontinuous resin canals in ribs absent. Figure 16 View FIG .

Diagnostic Characters — Phymaspermum woodii shares the narrowly campanulate to urceolate involucres and flat, deltoid to ovate, closely appressed involucral bracts with P. argenteum