Phymaspermum oppositifolium Magee and Ruiters, 2016

8. Phymaspermum oppositifolium Magee and Ruiters View in CoL sp. nov. —TYPE: SOUTH AFRICA. Eastern Cape, Steytlerville (3324): Kouga Mountain, Riverside (–CD), 20 January 2000, Eusten-Brown 02 (holotype: NBG!)

Single-stemmed, densely leafy shrub, 1.5 m high. Stem much-branched; branches erect to spreading, glabrous; fascicles present in leaf axils. Leaves opposite, closely imbricate, appressed to sometimes erect, 2.0–5.0 × 1.0–2.0 mm, lanceolate, acute to acuminate, sometimes mucronate, entire, without petiole-like base, without secondary basal lobes, glabrous on both surfaces, bony; basal swelling present, continuous with stem ribs. Capitula radiate, heterogamous, terminal on short axillary shoots, solitary, with 0–3 additional capitula from leaf axils below, prominent peduncles absent. Involucre 5.0–6.0 × 4.0–5.0 mm, funnel-shaped, tapering at the base; involucral bracts 4-seriate, tightly arranged, margins and apices scarious, membranous apices of all bracts 2.4–2.5 mm long, scarious texture, brown margins absent, median resin canals present; outer bracts lanceolate, 3.4 mm long, acute; middle bracts lanceolate, 4.0– 4.5 mm long, acute; inner bracts lanceolate, 5.5 mm long, rounded. Receptacle convex; paleae in marginal series, linear, ± 4.0 mm long, rounded, scarious. Ray florets ± 8; tube 1.5 mm long; limb ovate to obovate, 4.8 × 2.3 mm, apically 3-dentate, white to purple. Disc florets ± 20, without resin canals; corolla 2.0 mm long, with only glandular trichomes, yellow to purple; tube 0.8 mm long; limb narrowly campanulate, 1.0 mm long (excluding lobes); lobes erect, ± 0.4 mm long, triangular. Anthers 2.4 mm long (including apical appendage); apical appendage ovate. Style ± 2.0 mm long (excluding branches); branches ± 0.3 mm long. Cypselas 1.9 × 0.6 mm, narrowly obovate, ribbed, apical rim thickened, entire, brown, glandular trichomes present, distribution unknown, discontinuous resin canals in ribs absent. Figure 11 View FIG .

Diagnostic Characters — Phymaspermum oppositifolium is the only opposite leafed species within the genus. It shares the appressed leaves, entirely scarious involucral bracts and purplish ray florets with the closely related P. appressum from which it can be distinguished further by the funnel-shaped involucres and lanceolate involucral bracts.

Distribution and Ecology — This species is known only from the type collection in the Kouga mountains of the Eastern Cape ( Fig. 11G View FIG ), where it was collected at an altitude of 250 m. It was found flowering in January and the collector noticed seeing very few plants (Eusten-Brown pers. comm.).

9. PHYMASPERMUM ACICULARE (E. Mey. ex. DC.) Benth. and Hook. f. ex. B.D. Jacks. in Gen. Pl. 2(1): 422 (1873); B.D. Jacks., Index Kew. 11: 339 (1895); Källersjö 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 P.P.J. Herman, Pl. N. Prov. S. Afr.: 330 (1997); Germish. and N.L. Meyer, Pl. S. Afr.: 273 (2003); Klopper et al. Checklist of the flowering plants of Sub- Saharan Africa: 147 (2006); Germish. et al. Checklist S. African Pl.: 243 (2006). Oligoglossa acicularis E. Mey. ex. DC., Prodr. 6:76 (1838). Iocaste acicularis (E. Mey ex. DC.) Harv. in Harv. and Sond., Fl. Cap. 3: 160 (1864). —TYPE: SOUTH AFRICA. Northern Cape: ‘Bei Gaatjie’, December 1835, Drège 749 (lectotype: G–DC sheet G00455513–image!, designated here; isolectotypes: G–DC sheet G00386479– image!, P–image!). [Note: de Candolle cited having seen both an Ecklon as well as a Drège collection in the protologue. There are specimens of each in G–DC but the Drège collection (from locality 1,a,36) is selected here as it is rich in vegetative and reproductive material.]

Single-stemmed, leafy shrub, 0.45–1.5 m high. Stem few-branched; branches erect, glabrous with few glands; fascicles rarely present in leaf axils. Leaves alternate, erect to spreading, 3.0–20.0 × 0.8–1.5 mm, linear, mucronate, entire, without petiole-like base, without secondary basal lobes, glabrous on both surfaces, leathery; basal swelling present, continuous with stem ribs. Capitula discoid or radiate, homogamous or heterogamous, terminal, solitary, with 3–5 additional capitula from leaf axils below, pedunculate; peduncles 7.0–48.0 mm long, glabrous. Involucre 4.0–6.0 × 3.0–6.0 mm, broadly to narrowly campanulate or funnel-shaped, usually tapering at the base; involucral bracts 3- or 4-seriate, tightly arranged, margins and apices scarious, sometimes completely scarious especially third and fourth bracts, membranous apices of inner bracts 0.5–1.0 mm long, glabrous, brown margins absent, median resin canals present; outer bracts deltoid to lanceolate, 1.2–1.8 mm long, acute, sometimes mucronate; middle bracts deltoid to lanceolate or elliptic, 1.8–2.8 mm long, rounded to acute or sometimes mucronate; inner bracts narrowly obelliptic to lanceolate or linear, 2.2–3.2 mm long, rounded; innermost bracts narrowly obelliptic to linear, 2.4–4.0 mm long, rounded. Receptacle flat to convex; paleae in marginal series, narrowly obelliptic to linear, 2.7–4.0 mm long, rounded, scarious. Ray florets (when present) 3–4; tube 1.0– 1.7 mm long; limb obelliptic, 3.0–5.0 × 1.8–2.8 mm, apically 3-dentate, white. Disc florets 13–32, without resin canals; corolla 2.0– 3.5 mm long, with only glandular trichomes, yellow; tube 1.0– 2.6 mm long; limb campanulate, 1.0– 1.7 mm long (excluding lobes); lobes spreading, ± 0.7 mm long, triangular. Anthers ± 2.8 mm long (including apical appendage); apical appendage elliptic to lanceolate. Style 1.8–2.6 mm long (excluding branches); branches 0.1–0.4 mm long. Cypselas 2.5 × 1.0 mm, obovate, 10-ribbed, apical rim thickened, entire to slightly dentate, glandular trichomes present, sparse, scattered, not mucilaginous when soaked, discontinuous resin canals in ribs absent. Figure 12 View FIG .

Diagnostic Characters — Phymaspermum aciculare is the only species in the genus which can have either discoid or radiate heads. It can be distinguished from the other radiate species by the sparsely radiate heads with no more than four rays, with limbs less than 5 mm long. Amongst the discoid species it shares the non-corymbose capitula and large cypselas (> 0.8 mm wide) with P. erubescens and P. peglerae . It can however be readily distinguished by the glabrous stem and leaves and the smaller (3.0–6.0 mm wide), obconical, sometimes few radiate capitula with 3–5 additional capitula from leaf axils below.

Distribution and Ecology — Phymaspermum aciculare is widely distributed from McGregor in the Western Cape to Bloemhof in the North West Province with a disjunction in Namibia ( Fig. 12G View FIG ). A similar Namibian disjunction has also been reported in Melolobium (Moteetee and Van Wyk 2006) as well as Dichilus ( Schutte and Van Wyk 1988) . This species prefers hard, dry soil and sometimes occurs on dolerite, at altitudes of

40 m to 1,700 m. It is reportedly grazed by livestock ( Henrici 2844, BOL; Roux 36, PRE; Liebenberg 7673, B–image, PRE). Flowering is from September to May.

Additional Specimens Examined — NAMIBIA. 2217 (Windhoek): Dobra river bank (–AC), Koenen 580 (WIND); Farm Hohenau (–CB), Walter and Walter 1177 (B–image); Lichtenstein (–CC), Dinter 3527 (B– two images, PRE, SAM).

SOUTH AFRICA. North West: 2725 (Bloemhof): Leeuwfontein, 10 km W from Wolmaranstad (– BB), Van Wyk 1451 ( PRE). Free State: 2824 (Kimberley): Mcgregor Hills, Kamfersdam (– DB), Esterhuysen 3683 ( BOL). 2826 (Brandfort): Krugersdriftdam Nature Reserve on Deelkop (– CC), Muller 1507 ( PRE). 2925 (Jagersfontein): Bloemfontein, Bestersput near Petrusberg (– BA), Acocks 8475 ( BOL, PRE); Fauresmith (– CB), Henrici 4710 ( PRE); Fauresmith, Rotorua (– CC), Henrici 2844 ( BOL). Northern Cape: 2822 (Glen Lyon): Hay division, Floradale (– BD), Esterhuysen 2359 ( BOL two sheets). 2823 (Griekwastad): Past Sharon farm on road to Postmasburg at high powerline (– AB), Germishuizen 8586 ( PRE). 2824 (Kimberley): Rust en Vrede (– DA), Anon. H1251 ( PRE). 2924 (Hopetown): Kraanvogel Valley (– CD), Acocks 1681 ( BOL). 3022 (Carnarvon): Grootfontein S of Prieska, Leeuput (– BD), Roux 36 ( PRE). 3123 (Victoria West): 36 km from Victoria West on main road to Britstown, farm Rietpoort (– AA), Herman 1152 ( PRE). 3220 (Sutherland): Salpeterkop SE slope, between Amandelboom and De Hoek, Fraserburg (– BD), Moffet and Steensma 3964 ( PRE). Western Cape: 3222 (Beaufort West): Nieuwveld (– AB), Marloth 4753 ( PRE, BOL) Drège 389 ( SAM); north end of Molteno Pass (– BA), Acocks 14338 ( PRE); Bleak house farm (– BA), Russel, Robinson and Herman 337 ( PRE); Sunnyside (– BC), Esterhuysen 2718 ( BOL). Eastern Cape: 3026 (Aliwal North): Burghersdorp (– CD), Guthrie 4207 ( BOL); Niewejaarspruit, between Orange and Caledon rivers (– DB), Zeyher s.n. ( HBG –image), Ecklon and Zeyher s.n. ( HAL –image); Eland’ s Hoek (– DC), Bolus 235 ( BOL). 3225 (Mahlobyanini East): Cradock, Mountain Zebra Park, on plains below house of Berghof (– AD), Liebenberg 7673 ( B –image, PRE); Cradock, Bergkwagga Park (– AD), Liebenberg 7144 ( PRE); S of Cradock (– BA), Maguire 685 ( NBG); Cradock (– BA), Zeyher 225 ( SAM two sheets), Zeyher 961 ( K –image, S –image); near Mortimer in Cradock (– BC), Kensit 9292 ( BOL). 3124 (Hanover): Herbert, Eureka (– CB), Acocks 8749 ( BOL); Sneeuwbergen (– DC), Bolus 1846 ( NBG); north slopes of Sneeuwberg, Leopard’ s valley (– DC), Bolus 3431 ( BOL). 3125 (Steynsburg): 4 miles from Middelburg on Rietpoort road (– AC), Commins 703 ( BOL); Grootfontein, Middelburg (– AC), Theron 331 ( PRE). 3226 (Fort Beaufort): Lefiswane (– CA), Tyson 38747 ( SAM). 3325 (Port Elizabeth): Uitenhage, Ecklon 1142 ( G –DC!).

PRECISE LOCALITY UNKNOWN:Luaggasvlakte, MacOwan 1087 ( PRE); MacOwan 38749 ( SAM).

10. PHYMASPERMUM ERUBESCENS (Hutch.) 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 N.L. Meyer in Pl. S. Afr.: 274 (2003) ; Klopper et al. in Checklist of the flowering plants of Sub- Saharan Africa: 147 (2006); Germish. et al. Checklist S. African Pl.: 243 (2006). Brachymeris erubescens Hutch., Bull. Misc. Inform. Kew,1916: 173 View in CoL (1917).—TYPE: SOUTH AFRICA. Eastern Cape, Mthatha (3128): Tembuland: Tabase, near Bazeia (– DA), Baur 334 (lectotype: K –image!, designated by Källersjö 1986; isolectotype: SAM!) .

Phymaspermum equisetoides Thell. View in CoL in Vierteljahrsschr. Naturf. Ges. Zurich. 68. 446 (1923); Källersjö in Nord. J. Bot 5 (6): 535 (1986) ; K. Bremer and Humphries in Bull. Nat. Hist. Mus. Lond. ( Bot. ) 23(2): 94 (1993) ; Klopper et al. Checklist of the flowering plants of Sub- Saharan Africa : 147 (2006). syn. nov.—TYPE: SOUTH AFRICA. Eastern Cape, Kokstad (3029): Pondoland , Mnceba (– CC), Dec 1913, Jaccotet and Jaccotet 58 (lectotype: Z sheet 3788–image!, designated here; isolectotype: E–image!, Z sheet 3787– image!) [Note: There are two sheets of this collection in Z. Sheet 3788 is selected here as it is rich in flowering and leaf material.]

Multistemmed, leafy shrublet, up to 0.3 m high. Stems few-branched; branches erect, sparsely hairy to silver villous or hispid; fascicles sometimes present in leaf axils. Leaves alternate, apressed to erect, 2.0–10.0 × 1.0 mm, linear to lanceolate, acute to acuminate, revolute, entire, without petiole-like base, without secondary basal lobes, glabrous to silver-villous adaxially, glabrous abaxially, fleshy to leathery; basal swelling present, continuous with stem ribs. Capitula discoid, homogamous, terminal, solitary, with 0–3 additional capitula from leaf axils below, pedunculate; peduncles 15.0–60.0 mm long, glabrous. Involucre 5.0–7.0 × 5.0–7.0 mm, hemispherical to cyathiform, rarely tapering at the base; involucral bracts 4-seriate, tightly arranged, margins and apices scarious, membranous apices of all bracts 0.8–1.3 mm long, glabrous, brown margins absent, median resin canals present; outer bracts deltoid to lanceolate, 2.7–3.5 mm long, acute to attenuating; middle bracts triangular ovate, 4.0– 4.3 mm long, acuminate; inner bracts lanceolate, 4.6–5.2 mm long, rounded. Receptacle convex; paleae in marginal series, oblanceolate to oblong, 4.6– 5.2 mm long, rounded, membranous with scarious margins and apices, scarious apices 0.6–1.2 mm long. Disc florets ±20, without resin canals; corolla 2.0–3.0 mm long, with only glandular trichomes, yellow; tube 1.0– 1.2 mm long; limb narrowly campanulate, 1.0– 1.2 mm long (excluding lobes); lobes spreading, 0.6–1.0 mm long, triangular ovate. Anthers 2.8–3.6 mm long (including apical appendage); apical appendage obovate. Style 2.4–2.8 mm long (excluding branches); branches 0.4–1.0 mm long. Cypselas 3.6 × 1.0 mm, obovate, 11-ribbed, apical rim thickened, dentate, glandular trichomes present, sparse, scattered, not mucilaginous when soaked, discontinuous resin canals in ribs absent. Figure 13 View FIG .

Diagnostic Characters — Phymaspermum erubescens could be confused with P. peglerae or discoid specimens of P. aciculare but can be readily distinguished by the sparse to dense silvery villous or hispid indumentum on the stems and leaves (densely bronze villous in P. peglerae and glabrous in P. aciculare ), the shorter leaves, 2.0–10.0 mm long and smaller cypselas (when compared to P. peglerae ) ≤ 4 mm long.

Distribution and Ecology — Phymaspermum erubescens is a relatively poorly collected species which occurs from Lady Frere to King Williams Town in the Eastern Cape ( Fig. 13F View FIG ). It favours sandstone in grasslands at altitudes of 600–1,700 m. Flowering is from October to March.

Additional Specimens Examined — SOUTH AFRICA. Eastern Cape: 3127 (Lady Frere): Farm Ebuhleni ca. 27 km N of Indwe along R 396, track up to Vodacom tower (–AD), Magee et al. 436 (NBG); Farm Ebuhleni, SW mountain slopes, N of homestead (–AD), Bester 7396 (PRE); hills near Cala (–DA), Esterhuysen 29184 (BOL); Ngcobo (–DB), Acocks 12161 (PRE). 3227 (Stutterheim): King Williams Town (–CD), Flanagan 2164 (NBG).

PRECISE LOCALITY UNKNOWN: Kaffraria, Sim 19851 (PRE).

11. PHYMASPERMUM PEGLERAE (Hutch.) 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) ; 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). Brachymeris peglerae Hutch. View in CoL in Bull. Misc. Inform. Kew, 1916: 174 (1917). —TYPE: SOUTH AFRICA. Eastern Cape, Mthatha (3128): ‘ Tembuland , Mthatha River banks’ (– BC), 31 Jan 1861, Pegler 1601 (holotype: K –image!; isotypes: BOL! two sheets, PRE!). [Note: The specimen in K is clearly marked in Hutchinson’ s hand as “Type”.]

Multistemmed, densely leafy shrublet, 0.6–0.7 m high. Stems few-branched; branches erect, silvery villous; fascicles sometimes present in leaf axils. Leaves alternate, erect, 9.0–18.0 × 1.0 mm, linear, acuminate to mucronate, entire, without petiole-like base, without secondary basal lobes, silvery-villous on both surfaces, fleshy; basal swelling present, continuous with stem ribs. Capitula discoid, homogamous, terminal, solitary, pedunculate; peduncles 25.0–80.0 mm long, densely silvery-tomentose. Involucre 8.0–12.0 × 7.0–10.0 mm, hemispherical to cyathiform, not tapering at the base; involucral bracts 3-seriate, tightly arranged, margins and apices scarious, membranous apices of all bracts 0.5–1.0 mm long, densely villous, brown margins absent, median resin canals present; outer bracts lanceolate, 2.7 mm long, attenuating; middle bracts lanceolate, 3.9 mm long, acute to rounded; inner bracts oblong linear, 5.5 mm long, rounded. Receptacle convex; paleae in marginal series, oblong, ± 6.3 mm long, rounded, membranous with scarious margins and apices, scarious apices ± 1.3 mm long. Disc florets ± 20, without resin canals; corolla 2.3 mm long, with only glandular trichomes, yellow; tube 1.0 mm long; limb campanulate, 1.0 mm long (excluding lobes); lobes spreading, ± 0.5 mm long, triangular-ovate. Anthers ±3.0 mm long (including apical appendage); apical appendage obovate, obtuse. Style 4.0 mm long (excluding branches); branches 0.5 mm long. Cypselas 4.6 × 1.0 mm, obovate, 12-ribbed, apical rim thickened, dentate, glandular trichomes present, sparse, scattered,

(2006). Pentzia athanasioides S. Moore View in CoL in J. Bot. 41: 133 (1903). Brachymeris athanasioides (S. Moore) Hutch. View in CoL in Bull. Misc. Inform. Kew 1916: 173 (1917). —TYPE: SOUTH AFRICA. Gauteng, Johannesburg (2628): Near Hospital Hill (–AA), Rand 758 (lectotype: BM–image!, designated here). [Note: This is the only specimen housed in BM and corresponds closely to the original description.] Brachymeris montana Hutch. View in CoL in Bull. Misc. Inform. Kew 1916: 174 (1917), syn. nov. Phymaspermum montanum (Hutch.) 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 (2006). — TYPE: SOUTH AFRICA. Limpopo, Pilgrims Rest (2430): Mt. Marovougne (–AB), April, Junod 1278 (holotype: K–image!). [Note: The specimen in K is clearly annotated in Hutchinson’ s hand as “Type specimen”.]

Brachymeris bolusii Hutch. View in CoL in Bull. Misc. Inform. Kew 1916: 174 (1917), syn. nov. Phymaspermum bolusii (Hutch.) 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 (2006). —TYPE: SOUTH AFRICA. Mpumalanga, Komatipoort (2531): Drakensberg , Devils Kantoor Mt. (– CC), Sept 1886, Bolus 7786 (holotype: K – image!; isotype: BOL!) [Note: The specimen in K is clearly annotated in Hutchinson’ s hand as “Type.”]

not mucilaginous when soaked, discontinuous resin canals in ribs absent. Figure 14 View FIG .

Diagnostic Characters — Phymaspermum peglerae shares the discoid non-corymbose heads and large fruit with P. erubescens and P. aciculare but can be distinguished by the densely bronze villous indumentum (sparse to dense silvery villous or hispid indumentum in P. erubescens and glabrous in P. aciculare ), the usually larger leaves 9.0–18.0 mm long and the largest fruit in the genus> 4 mm.

Distribution and Ecology — Phymaspermum peglerae is known from only two very old collections along the banks of the Mthatha River in the Eastern Cape ( Fig. 14F View FIG ). This species flowers in midsummer, around January.

Additional Specimens Examined — SOUTH AFRICA. Eastern Cape: 3128 (Mthatha): Mthatha River bank (–BC), Bolus 10133 (BOL).