Nyctaginaceae: a Taxonomic Treatment for the Flora of Namibia

http://dx.doi.org/10.11646/phytotaxa.238.2.1

Nyctaginaceae: A taxonomic treatment for the Flora of Namibia

MADELEEN STRUWIG1*, ESMERIALDA S. Klaassen2 & EZEKIEL G. Kwembey a3 1Department of Botany, University of Zululand, Private Bag X1001, KwaDlangezwa 3880, South Africa; email: [email protected] 2National Botanical Research Institute, Private Bag X13184, Windhoek, Namibia; email: [email protected] 3University of Namibia, Private Bag 13301, Pionierspark, Windhoek, Namibia; email: [email protected] * Author for correspondence

Abstract

A taxonomic treatment of the Nyctaginaceae for the Flora of Namibia is presented, including the description of the family, keys to the genera and species, nomenclature, descriptions of all generic and infrageneric taxa, as well as data about the habitat, elevation, flowering time and distribution. Namibia is the centre of diversity for the family in southern Africa and is represented by four genera and 17 species distributed throughout the country. The names Mirabilis divaricata, Boerhavia atomaria, B. viscosa, B . coccinea f. parcehirsuta and B. stellata are typified on specimens respectively preserved at BM, PH, G, B and an illustration.

Key words: Africa, Boerhavia, Commicarpus, Mirabilis, Phaeoptilum, typification

Introduction

The family Nyctaginaceae Juss. includes about 30 genera and 300–400 species (see e.g., Bittrich & Kühn 1993, Douglas & Manos 2007) that are distributed throughout the tropical and subtropical regions of the world (Bittrich & Kühn 1993, Jordaan 2000), especially in the Neotropics and the arid territories of western North America (Douglas & Spellenberg 2010). Namibia is the centre of diversity for this family in southern Africa with 4 genera and 17 species (Makwarela 2003).

Materials and Methods

Living plants and specimens from the herbaria BLFU, BOL, J, NGB, NU, PRE, PRU, PUC, SAM and WIND (acronyms according to Thiers 2015+) were examined. The following morphological characters were measured: length and width of the leaves, length of the petiole, length of the peduncle, pedicel, upper and lower portions of the perianth, stamens and ovary, and mean length and width of anthocarps (ten specimens per species and three anthocarps per specimen). Fresh floral material was collected in situ and preserved in 4% paraformaldehyde and flowers taken from herbarium specimens were rehydrated for 10 minutes in boiling water. The terminology of the morphological characters follows Hickey & King (2000). Distribution and habitat data were obtained from herbarium labels as well as observations in the field. Distribution maps were generated using ArcView GIS 3.2a.

Taxonomic treatment

Nyctaginaceae Jussieu, nom. cons.

Type (designated by Larsen 1989: 108):—Mirabilis jalapa Linnaeus

Accepted by Duilio Iamonico: 14 Nov. 2015; published: 11 Dec. 2015 101 Description:—Annual or perennial herbs, sub-shrubs or shrubs. Exstipulate. Leaves simple, opposite, alternate or fascicled, petiolate or sessile to subsessile, entire. Inflorescence pedunculate, compound cyme or umbel, umbel sometimes verticillate, terminal or axillary. Flowers pedicillate, unisexual or bisexual. Perianth greenish yellow to pale yellow, white, red or variegated, or divided into a distinct lower and upper part; lower part green, coriaceous, constricted above the ovary, 3–5-winged or 3–10-ribbed, persistent; upper part petaloid, campanulate or infundibuliform, sometimes with a distinct basal tube, pink, purple, maroon or white, caducous after anthesis. Stamens 1–9, exserted; filaments filiform, connate into a short sheath around the ovary. Ovary sessile, subsessile or shortly stipitate, ellipsoid, one ovule; style filiform, exserted; stigma capitate, truncate or penicillate. Fruit an achene enclosed in hardened base of perianth (anthocarp). Anthocarp 3–5-winged or 5–10-ribbed, smooth or tuberculate, glabrous or covered in glandular hairs, sometimes with a hygroscopic mucoidal secretion. Seed endosperm scanty, embryo hooked or curved. Distribution:—The family Nyctaginaceae is distributed throughout the tropical and subtropical regions of the world (Bittrich & Kühn 1993, Jordaan 2000, Douglas & Spellenberg 2010).

Key to genera

1. Woody shrub plants up to 3 m high; branches terminating in spines; leaves fascicled or alternate...... Phaeoptilum - Herbaceous or subshrub-like, annual or perennial plants up to 2 m high; branches spineless; leaves opposite...... 2 2. Flowers salverform, perianth not divided into a distinct upper and lower part; anthocarp obovoid, subglobose or ellipsoid...... Mirabilis - Flowers campanulate or infundibiliform, perianth divided into a distinct upper and lower part; anthocarp ellipsoid-clavate, clavate or fusiform...... 3 3. Perianth campanulate, 1–5 mm long; lower coriaceous part of the perianth 3–5-winged or 5-ribbed, glabrous or covered in glandular hairs; anthocarp 3–5 mm long, 5-ribbed or 3–5-winged, glabrous or covered in glandular hairs...... Boerhavia - Perianth infundibiliform, 5–14 mm long; lower coriaceous part of the flower 10-ribbed with wart-like glands around the apex and surface below the apex; anthocarp 5–10 mm long, 10-ribbed with stalked or sessile glands around the apex and wart-like glands on the surface below...... Commicarpus

Phaeoptilum Radlkofer (1884: 435).

Type:—Phaeoptilum spinosum Radlkofer = Amphoranthus S.Moore (1902: 305). Type species: Amphoranthus spinosus S.Moore.

Description:—Shrub, much branched, branches ending in spines. Leaves in fascicles or alternate, sessile or subsessile, entire. Flowers solitary or in fascicles, bisexual or unisexual, usually on leafless branches. Perianth tube campanulate, 4- or 5-lobed. Stamens 8(9); of two different lengths; filaments connate into a short sheath around the ovary; reduced in female flowers. Gynoecium reduced in male flowers. Ovary stipitate, one ovule; style asymmetrically inserted at the apex of the ovary, stout. Anthocarp broadly 4- or 5-winged. Notes:—Phaeoptilum is a monotypic genus of southern Africa (Jordaan 2000).

Phaeoptilum spinosum Radlkofer (1884: 435).

Type:—South Africa. Northern Cape, Hantam region, Meyer s.n. (holotype B-100550160!). = Phaeoptilum heimerli Engler (1895: 133). Type:—Namibia . Kamelneck. Gürich 42 (holotype B-100550159!, isotype B-100550158!). = Amphoranthus spinosus S.Moore (1902: 305, fig. 441A). Type:—Namibia . Damaraland. Een s.n. (holotype BM-000839850!).

Description:—Shrub up to 3 m high; bark greyish yellow or greyish brown. Leaves linear, (9–)10–30(–40) × 1–4 mm long, apex rounded or emarginate, base attenuate; somewhat fleshy, coriaceous, yellow green to greygreen; glabrous to pubescent. Flowers scented; bracteate. Perianth 6–8 mm long; 4(–5)-lobed, lobes spreading, ovate, rounded to triangular at the tips, tomentose, greenish yellow to pale yellow. Stamens 8(9), of different lengths, longer stamens 6–12.5 mm long, slightly exserted; shorter stamens 4–10 mm long; filaments filiform; anthers oblong-elliptic, 0.5–2 mm long. Ovary fusiform-elliptic; style 1.5–3.5 mm long; stigma truncate or penicillate. Fruit 7–8 × 2.5–3 mm,

102 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. oblong-elliptic. Anthocarp 15–25 × 12–20 mm; central portion fusiform; broadly 4(5)-winged, wings semi-circular; pubescent to glabrescent, pink, red or purple maturing brown (Fig. 1). Habitat:—Grows in arid areas on rocky, gravelly and sandy soil, often along dry river courses, depressions and on plains.

Figure 1. Phaeoptilum spinosum. A. Fruiting branch. Scale bar 10 mm. B. Flower. Scale bar 2.5 mm. Drawings prepared by Gillian Condy.

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 103 elevation:— about 1500 m. Flowering time:—August to May. Distribution:—Namibia (Fig. 2), Botswana and South Africa.

Figure 2. Known distribution of Phaeoptilum spinosum in Namibia.

Specimens examined:—Namibia . Near old borehole, Swartskamp area, on Hobatere, 1914AD, 27 August 1998, Loutit 489 (WIND); Windhoek, road from Dordabis to Uhlenhorst, 2217DC, 18 December 1999, Hanke HAN2/1 (WIND); Main road, closest to Maltahöhe, 2416CD, 16 December 2002, Bridgeford BRI1/18 (WIND); West of Maltahöhe on road to Solitaire, 2416DB, 11 March 1995, Germishuizen 7811 (PRE; WIND). Bethanie district, Aruab farm 23, on top edge of Great Escarpment, 2516DA, 10 January 1998, Miller MIL 1/037 (WIND).

Mirabilis Linnaeus (1753: 177).

Type (designated by Larsen 1989: 108):—Mirabilis jalapa Linneaus = Jalapa Miller (1754: unpaged). Type specie:—Jalapa dichotoma Crantz. = Nyctago Jussieu (1789: 90). Type species:—Nyctago jalapa Candolle. = Trimista Rafinesque (1840: 12). Type species:—Trimista levigata Rafinesque. = Admirabilis Nieuwland (1914: 280). Type species:—Admirabilis peruana Nieuwland.

Description:—Herbaceous or suffruticose perennial, stems ascending to erect, much branched, glabrous or glandular- pubescent, nodes thickened, roots tuberous. Leaves opposite, glabrous to puberulent, ovate, oblong, lanceolate, deltoid; apex acute, acuminate or obtuse; base cordate, subtruncate or rounded, margins entire; petiolate or subsessile.

104 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. Inflorescence a 1-many flowered cyme, involucral bracts 5-lobed, connate. Flowers conspicuous. Perianth salverform or shortly funnelform; white, pink, red, yellow, magenta or variegated. Stamens 3–6, exserted. Ovary sessile or subsessile. Anthocarp obovoid, subglobose to ellipsoid; 5-ribbed or 5-angled, smooth or tuberculate, glabrous or pucescent, coffee coloured or dark-brown to black. Notes:—Mirabilis is a genus of about 60 species native to the warmer parts of America (Spellenberg 2003). It is introduced to Namibia and cultivated as a garden ornamental.

Key to species of Mirabilis:

1. Inflorescence in a many-flowered cyme; flowers 40–50 mm long; anthocarp subglobose to ellipsoid, 7–9 mm long...... 1. Mirabilis jalapa - Inflorescence in a one-flowered (rarely 2 or 3 flowered) cyme; flowers 8–13 mm long; anthocarp obovoid, 4–5 mm long...... 2. Mirabilis viscosa

1. Mirabilis jalapa Linnaeus (1753: 177) ≡ Jalapa congesta Moench (1794: 508) ≡ Nyctago versicolor Salisbury (1796: 57) ≡ Nyctago jalapa De Candolle (1805: 426) ≡ Admirabilis peruana Nieuwland (1914: 280).

Lectotype (designated by Larsen 1989: 108):—UNKOWN. Herb. Linneaus 240.2 (LINN!) Image available from http://linnean-online. org/1827/ = Mirabilis odorata Linnaeus (1755: 7) ≡ Mirabilis dichotoma Linnaeus (1762: 252) ≡ Jalapa dichotoma Crantz (1766: 266) ≡ Jalapa undulate Moench (1802: 196) ≡ Mirabilis jalapa f. odorata (L.) Heimerl (1896a: 616). Lectotype (designated by Le Duc 1995: 640):—MEXICO. Herb. Linneaus 240.1 (LINN!). Image available from http://linnean-online. org/1826/ = Nyctago mirabilis Jaumin Saint-Hilaire (1805: t.37). Lecotype (designated by Le Duc 1995: 640):—[Icon] Nyctago mirabilis in Saint-Hilaire (1805: t.37). Mirabilis pedunculata Stokes (1812: 311). = Mirabilis divaricata Lowe (1831: 17). Lectotype (designated here)—PORTUGAL. Madeira, in a garden, 1 October 1827, Lowe s.n. (BM-001122778!). = Mirabilis procera Bertoloni (1839: 15, t.1) ≡ Mirabilis jalapa var. procera (Bertol.) Choisy (1849: 428). Lectotype (designated by Le Duc 1995: 640):—[Icon] Mirabilis procera in Bertoloni (1839: t.1). = Mirabilis planiflora Trautvetter (1840: 215) ≡ Mirabilis jalapa var. planiflora (Trautv.) Choisy (1849: 428). Type:—UKRAINE. Cultivated plant, Kiev Botanical Garden (holotype KW). = Trimista levigata Rafinesque (1840: 12). Type:—Guatimala. Guatimala, Rafinesque s.n. (holotype DWC). = Mirabilis ambigua Trautvetter (1841: 97) ≡ Mirabilis jalapa var. ambigua (Trautv.) Choisy (1849: 428). Type:—UKRAINE. Cultivated plant, Kiev Botanical Garden 1840 (holotype KW) = Mirabilis jalapa subsp. ciliata Standley (1909: 368). Type:—Mexico . Oaxaca, valley of Oaxaca, 1 October 1894, Smith 791 (holotype MO, isotype US.). = Mirabilis jalapa subsp. lindheimeri Standley (1909: 368) ≡ Mirabilis jalapa var. lindheimeri (Standl.) Cory (1936: 405) ≡ Mirabilis lindheimeri (Standl.) Shinners (1951: 175). Lectotype (designated by Le Duc 1995: 640):—USA. Texas. Comal Co.: New Braunfels, June 1846, Lindheimer 158 (MO). = Mirabilis jalapa subsp. gracilis Standley (1909: 367). Type:—Mexico . Sinaloa: Culiacan, 17 September 1904, Brandegee s.n. (holotype UCR). = Mirabilis jalapa subsp. volcanica Standley (1909: 367). Type:—Mexico . Distrito Federal. Pedregal (lava bed), valley of Mexico, 19 August 1896, Pringle 6433 (holotype MO, isotype BR, GH, NDG, US).

Description:—Plants up to 1.5 m high. Leaves (38–)40–97(–105) × (22–)23–48(–50) mm; ovate, oblong, lanceolate, deltoid; apex acute or acuminate; base cordate; petiole (5–)21(–35) mm long. Inflorescence a 3–7-flowered cyme, involucral bracts 5–13 mm long, lobes acuminate, green. Flowers 40–50 mm long, open in the late afternoon, fragrant. Perianth salverform; white, pink, red, yellow or variegated. Stamens 5 or 6; exserted, filaments filiform, connate into a fleshy cup at the base; anther oblong-ellipsoid. Ovary ellipsoid or ovoid; style filiform; stigma with stalked papillae. Anthocarp subglobose to ellipsoid, 7–9 × 4–5 mm long, coriaceous, dark-brown to black.

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 105 Habitat:—This species is cultivated as a garden ornamental from where it is often escape. It can be found in disturbed places as a naturalized species. elevation:— about 650–1590 m. Flowering time:—October to April. Distribution:—Namibia (Fig. 3), Botswana, South Africa and tropical America.

Figure 3. Known distribution of Mirabilis jalapa in Namibia.

Typification:—Lowe (1831: 17) originally indicated Mirabilis divaricata to be a cultivated plant on the island of Madeira. We found one specimen at BM (barcode 001122778) bearing a plant collected in Madeira with features matching the protologue; a note by Lowe indicating it to be a new species also occurs. We consider this specimen as part of the original material and designate it here as the lectotype of the name Mirabilis divaricata. Specimens examined:—Namibia . Otjiwarongo. Acacia Park, in camping area, 2016BC, 11 April 2010, Struwig 180 (PUC, WIND).

2. Mirabilis viscosa Cavanilles (1791: 13, t.19) ≡ Oxybaphus viscosus (Cav.) L’Héritier de Brutelle ex Willdenow (1797: 185) ≡ Allionia viscosa (Cav.) Kuntze (1891: 533).

Lectotype (designated by Spellenberg 2001: 71):—[Icon] Mirabilis viscosa in Cavanilles (1791, t.19).

Description:—Suffrutescent perennial, 1.3 m long, mostly branched at the top, pubescent. Leaves become smaller to the top of the plant; leaves on the lower portion of the plant 50–160 × 50–130 mm, ovate, deltoid, apex acute or

106 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. obtuse, base cordate or subtruncate, petiole 50–80 mm long; leaves on the upper portion of the plant subsessile, 30–50 × 20–40 mm, ovate, apex acute or acuminate, base cordate or rounded; glandular-pubescent to glabrate. Inflorescence a one (rarely 2 or 3) flowered cyme; involucral bracts 9–12 mm long, pale brown to brown, with conspicuous veins, apex broadly obtuse to rounded. Flowers sessile; 8–13 mm long. Perianth salverform, magenta. Stamens 3. Anthocarp obovoid, 4–5 × 2.5 mm, tuberculate, glabrous to minutely pubescent, coffee coloured. Habitat:—Grows in stony, sandy loam or clayish soil. Flowering time:—December to June. Distribution:—Namibia (Fig. 4), northern and central Mexico and South America.

Figure 4. Known distribution of Mirabilis viscosa in Namibia.

Specimens examined:—Namibia . Otjozondjupa, Grootfontein, Eichenbach farm, 1918CA, 27 January 2006, Bartsch, Klaassen, & Hoffmann SB2180 (PUC, WIND).

Boerhavia Linnaeus (1753: 3)

Type (designated by Fawcett & Rendle 1914: 148): Boerhavia erecta Linnaeus.

Description:—Annual or perennial herbs. Stems slender, erect, decumbent, diffuse or prostrate, woody towards base, glabrate to pubescent, with glandular hairs. Leaves opposite, petiolate; lamina linear or orbicular, obtuse to apiculate, base attenuate to truncate, margins entire or sometimes undulate, glabrous to pubescent. Inflorescence a much-branched terminal or axillary compound cyme or umbel, glabrous to puberulous, bracteate. Flowers 1–5 mm long, bisexual,

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 107 shortly pedicillate. Perianth divided into a lower and upper part; lower part 1–2 mm long, green or sometimes reddish, coriaceous, constricted above the ovary, ovoid to clavate, 3–5-winged or 5-ribbed, glabrous or with glandular hairs, persistent; upper part 1–4 mm long, campanulate, sometimes with a basal tube, petaloid, pink, purple, maroon or white, caducous after anthesis. Stamens 1–5, exserted; filaments filiform, connate at base forming a short sheath around ovary; anthers dithecous. Ovary ellipsoid, stipitate, 1-ovulate; style filiform, exserted; stigma capitate. Anthocarp 3–5 × 1–3 mm, ellipsoid-clavate or clavate, 3–5-winged or 5-ribbed; acute or obtuse; glabrous or covered with glandular hairs; secretes a hygroscopic mucoidal secretion which is produced in columnar cells in the ribs, especially when wetted. Seed oblong, brown, with smooth surface; endosperm scanty; embryo hooked.

Figure 5. Boerhavia deserticola. A. Habit. Scale bar 1 mm. B. Flower. C. Anthocarp. Scale bars B–C, 1.25 mm. Drawings prepared by Gillian Condy.

Notes:—Boerhavia is a genus of about 40 species distributed in the tropical and subtropical regions of the world

108 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. and is especially diverse in south-western North America (Douglas & Spellenberg 2010). Eight species occur in southern Africa of which three are endemic to the region (B. hereroensis Heimerl, B. deserticola Codd and B. orbicularifolia Struwig) (Makwarela 2003, Struwig & Siebert 2013b) and four introduced (B. coccinea Mill., B. cordobensis Kuntze, B. diffusa L. and B. erecta L) (Makwarela 2003, Struwig & Siebert 2013b).

Figure 6. Anthocarps of the southern African Boerhavia and Commicarpus species. A. B. coccinea. B. B. cordobensis. C. B. deserticola. D. B. diffusa. E. B. erecta. F. B. hereroensis. G. B. repens subsp. repens. H. C. chinensis subsp. natalensis. I. C. decipiens. J. C. fallacissimus. K. C. helenae var. helenae. L. C. pentandrus. M. C. pilosus. N. C. plumbagineus var. plumbagineus. O. C. squarrosus. Scale bars 1 mm. Drawings prepared by Gillian Condy.

Key to the species of Boerhavia (Struwig & Siebert 2013b):

1. Plants erect; leaves mostly elliptic-lanceolate, sometimes ovate or oblong...... 2 - Plants prostrate or decumbent; leaves mostly oblong to orbicular, sometimes elliptic-lanceolate...... 3 2. Lower coriaceous part of perianth and anthocarp glabrous...... 1. B. deserticola - Lower coriaceous part of perianth and anthocarp covered in glandular hair...... 2. B. hereroensis 3. Anthocarp and lower coriaceous part of perianth clavate and glabrous...... 4 - Anthocarp and lower coriaceous part of perianth ellipsoid-clavate and covered with glandular hairs...... 5 4. Lower coriaceous part of perianth 3–4-ribbed with apices of ribs clearly expanded; anthocarps 3–4-winged, wings straight and not undulate, apices broadly truncate...... B. cordobensis

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 109 - Lower coriaceous part of perianth 5-ribbed with apices of ribs slightly expanded; anthocarps 5-winged, wings undulate, apex truncate...... 4. B. erecta 5. Anthocarp with indentation near apex, apex acute, calcium oxalate crystals conspicuous between ribs...... 5. B. diffusa - Anthocarp with rounded apex, calcium oxalate crystals not conspicuous between ribs...... 6 6. Inflorescence an umbel or flowers solitary; plant with short stems up to 100 mm long...... 6. B. orbicularifolia - Inflorescence a compound cyme; plant with long trailing stems up to 2 m long...... 7 7. Leaves > 30 mm long; anthocarp narrowly ellipsoid-clavate (≥ 4 mm long)...... 7. B. coccinea - Leaves < 30 mm long; anthocarp ellipsoid-clavate (≤ 4 mm long)...... 8. B.repens subsp. repens

1. Boerhavia deserticola Codd (1966: 119)

Type:—Namib ia. Kaokoveld Reserve, Orupembe waterhole near Sanitatas, 5 May 1957, De Winter & Leistner 5722 (holotype PRE- 0394979-0!, isotype M-0107836, S-07-13006). Image available at http://plants.jstor.org/stable/10.5555/al.ap.specimen.pre0394979-0

Figure 7. Known distribution of Boerhavia deserticola in Namibia.

Description:—Perennial herb. Stems numerous, up to 1 m long, erect from a basal rosette, pubescent with long, septate hairs. Leaves with petioles (6–)10–33(–41) mm long; lamina elliptic, lanceolate or ovate, (15–)16–48(–55) × (5–)6– 33(–34) mm; apex apiculate, acute or rounded; base shortly attenuate; margins undulate; densely pubescent with long, septate hairs. Inflorescence with primary peduncles 95–120 mm long, secondary peduncles 34–40 mm long; 2-many flowers per cluster, pedicels 2 mm long. Flowers 4–5 mm long; lower part of perianth 1–2 mm long, green tinged pink, ovoid, 5-ribbed, glabrous; upper part of perianth 3–4 mm long, with a basal tube, white or purple. Stamens 4–6; 6 mm long. Ovary ellipsoid, stipitate; style filiform, exserted. Anthocarp 3–5 × 1–2 mm, clavate, 5-ribbed, ribs conspicuous, rounded at apex, glabrous (Fig. 5 and 6C).

110 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. Habitat:—Grows in well-drained to dry stony, sandy, sandy-loam or gravelly soils in gullies, riverbeds and riverbanks, on plains, and hillsides or mountain slopes. elevation:— about 556–1029 m. Flowering time:—January to December. Distribution:—Endemic to Namibia (Fig. 7). Specimens examined:—Namibia . Kunene, on a plain in Barab river-valley, 1913DA, 06 July 2000, Gindrig & Hennig 60 (PRE, WIND); On road to Sesfontein, roadside, in dry streambank, 1913DB, 08 March 2009 , Struwig 43 (PUC, WIND); Welwitschia, Bloemhof, old 484 Farm, 2014BC, Muller & Giess 387 (PRE); Twyfelfontein, village near Twyfelfontein Lodge, in dry streambed, 2014CB, 08 February 2009, Struwig 42 (PUC, WIND); Outjo district, Spaarwater, old 711 farm, 2016AA, 30 March 1974, Merxmüller & Giess 30622 (PRE). Brandberg, Numas Gorge, 2014AA, 19 March 1977, Muller & Giess 357 (PRE); Uis, Brandberg, near entrance to Tsisab gorge, 2014AA, 06 February 2009, Struwig 38 (PUC, WIND); Uis, Damaraland, Brandberg, on road to White Lady, 2014BA, 14 December 1974, Van Wyk 692 (PUC); Uis, Brandberg, hillslope near White Lady rock painting, 2014BA, 06 February 2009, Struwig 37 (PUC, WIND); Brandberg South, 2014BC, 19 April 1990, Baker 29 (PUC, WIND); Omaruru District, Brandberg, Tsisabschlucht, 2014BC, 12 September 1967, Urschler s.n. (PRE).

Figure 8. Known distribution of Boerhavia hereroensis in Namibia.

2. Boerhavia hereroensis Heimerl (1889: 9)

Type:—Namibia . Otjimbingwe, Marloth 1403 (holotype L-0038747, isotypes GRA-0000913-0, K-000243789, E-00193548, M- 0107835 PRE-0394975-0). Image available form http://plants.jstor.org/stable/10.5555/al.ap.specimen.l0038747

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 111 Description:—Perennial herb. Stems branches upright, diffuse, densely pubescent with long, septate hairs. Leaves with petioles (7–)8–27(–38) mm long; lamina elliptic, lanceolate, linear or ovate; (18–)20–40(–41) × (5–)7–27(–34) mm; apex apiculate; base shortly attenuate or rounded; margins entire, sometimes undulate; pubescent with long septate hairs. Inflorescence with primary peduncles ± 65 mm long, secondary peduncles ± 27 mm long; pubescent; 2–4 flowers per cluster; pedicels 1 mm long. Flowers 2–4 mm long; lower part of perianth 1 mm long, ovoid, 5-ribbed, with long, septate hairs; upper part of perianth 1–3 mm long, with a basal tube; pink or purple. Stamens 2–3, 2–2.5 mm long, included. Ovary ellipsoid, stipitate; style filiform, exserted. Anthocarp 3–4 × 1–2 mm, ellipsoid-clavate, 5- ribbed, rounded at apex, pubescent with long, septate hairs (Fig. 6F). Habitat:—Grows in stony, gravelly, sandy or loamy soil in dry watercourses, riverbanks and on mountain or hill slopes. elevation:— about 500–1700 m. Flowering time:—December to May. Distribution:—Namibia (Fig. 8) and South Africa. Specimens examined:—Namibia . Twyfelfontein Lodge, hills behind lodge, south-west of main building, 2014CB, 07 February 2009, Struwig 40 (PUC, WIND); Uis, Brandberg, Hungarob valley area, 2114AB, 04 March 1978, Craven 718 (PUC); Omaruru district, Daksberge, deep shade along riverbank, 2115BD, 20 March 1965, Hardy 2025 (PRE); Usakos, Klein Spitzkuppe mountain slope amongst rocks, 2115CC, 05 February 2009, Struwig 35 (PUC, WIND); Karibib, Klippenberg Country Club, rocky hill behind restaurant, 2115DD, 04 February 2009, Struwig 34 (PUC, WIND); Omaruru District, Okasongoro farm, between Omaruru and Kalkfeld, farm dam, 2116AA, 03 March 1983, Germishuizen 2527 (PRE); Otjimbingwe, 2216AC, 01 January 1886, Marloth 1403 (BOL); Windhoek Municipality Area, 2217CA, 17 December 1962, Hanekom 354 (PRE, WIND).

3. Boerhavia cordobensis Kuntze (1898: 264)

Type:—Argentina. Cordoba, Lorentz 403. (holotype F-0BN003084). Image available from https://plants.jstor.org/stable/10.5555/ al.ap.specimen.f0bn003084

Description:—Stems decumbent or prostrate, puberulent, with scattered long septate hairs. Leaves with petioles (3– )5–19(–25) mm long; lamina deltoid, oblong, lanceolate, elliptic or ovate, (14–)15–42(–47) × (8–)9–32(–34) mm; apex obtuse, acute, acuminate or apiculate; base obtuse or truncate; margins entire, sometimes undulate; glabrous, with long septate hairs along veins and margins. Inflorescence with primary peduncles 60–130 mm long, secondary peduncles 20–35 mm long; 5–many flowers per cluster; pedicels 1 mm long. Flowers 3 mm long; lower part of perianth 1–1.5 mm long, clavate, 3- or 4-ribbed with apex of ribs expanded, glabrous; upper part of perianth 1.5–2 mm long, purple or pink. Stamens 2 or 3. Ovary ellipsoid, stipitate; style filiform, exserted. Anthocarp 3–4 × 2–3 mm, clavate, 3- or 4- winged, broadly truncate-apiculate, glabrous (Fig. 6B). Habitat:—Grows in stony, sandy to sandy loam soil in dry riverbeds, on floodplains, plains, hillsides or mountain slopes and ridges. elevation:— about 310–1775 m. Flowering time:—November to April. Distribution:—Namibia (Fig. 9), Botswana, South Africa and central-western Argentina. Specimens examined:—Namibia. Welwitschia, 10 km west of Khorixas on Twyfelfontein road, 2014DB, 15 April 1984, Hines 33 (WIND); Windhoek, D1463, at T-junction sign before entrance to Aris farm, 2217CC, 13 February 2009, Struwig 82 (PUC, WIND); Naukluft. At Arbeid Adelt near farm house in riverbed, 2416AA, 29 March 1995, Burke 59134 (WIND).

4. Boerhavia erecta Linnaeus (1753: 3) ≡ Boerhaavia elongata Salisbury (1796: 56)

Neotype (designated by Fawcett & Rendle 1914: 148):—Mexico. Habitat in Vera Cruz, Herb. Linneus 9.1 (LINN!). Image available from http://linnean-online.org/28/ = Boerhaavia virgata Kunth (1817: 215). Type:—Venezuela. Cumana. Bonpland & Von Humboldt 1224 (holotype P-00670034!, isotype B-W-00766010!). = Boerhaavia discolor Kunth (1817: 215). Type:—Equador. Bonpland & Von Humboldt 3845 (holotype P-00670035!). = Boerhaavia atomaria Rafinesque (1840: 40). Neotype (designated here):—Florida. Ware s.n. (PH-00007956!) Image available from https://plants.jstor.org/stable/10.5555/al.ap. specimen.ph00007956 = Valeriana latifolia Marten & Galeotti (1844: 124). Type: Mexico . Croit dans les champs de la Antigua , prés Vera-Cruz. Galeotti 2558 (holotype BR-0000005231041!). = Boerhavia paniculata Richard var. subacuta Choisy (1849: 451). Type:—BRAZIL. Gardner 2292 (holotype P-00712503!). = Boerhaavia thornberi M.E.Jones (1908: 72) ≡ Boerhavia erecta var. thornberi (M.E.Jones) Standley (1909: 381). Type:—USA. Arizona, Tucson, 20 September 1903, Thornber 10 (holotype US-661131!).

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 113 Description:—Stems decumbent or prostrate, puberulent, with scattered long septate hairs. Leaves with petioles (5–)6–34(–35) mm long; lamina oblong, lanceolate, elliptic, deltoid or ovate, (15–)20–47(–51) × (10–)12–25(–29) mm; apex apiculate, acuminate, acute or obtuse; base shortly attenuate, obtuse or truncate; margins entire, sometimes undulate; glabrous, sometimes with long septate hairs along margins and veins. Inflorescence with primary peduncles ± 115 mm long, secondary peduncles 55–65 mm long, glabrous to puberulous; 4–10 flowers per cluster; pedicels 1 mm long. Flowers 2–3 mm long; lower part of perianth 1–2 mm long, clavate, 5-ribbed, glabrous; upper part of perianth 1 mm long, pink, purple or white. Stamens 2, slightly exserted, 2–2.5 mm long. Ovary 0.5 mm; style 1–2 mm long. Anthocarp 3–4 × 1–2 mm, clavate, 5-winged, wings slightly undulate along margins, apex truncate, ending in pointed tip, glabrous (Fig. 6E). Habitat:—Grows in stony, gravelly, sandy, loamy or clay soil in riverbeds, on plains, mountain- or hill slopes and ridges, often along roadsides. elevation:— about 250–1250 m. Flowering time:—October to May. Distribution:—Namibia (Fig. 10), South Africa, Mexico, West Indies, Central America and South America. Typification:—We were not able to trace any specimen which can be considered as part of the original material used by Rafinesque to describe Boerhavia atomaria. These exsiccata were most likely amongst the specimens discarded by Elias Durand upon purchasing Rafinesque’s herbarium (Stuckey 1971). As a consequence, according to the art. 9.7 of ICN (McNeill et al. 2012), a neotypification is required. We choose a specimen collected by Ware as the neotype as it best fits the protologue.

Figure 10. Known distribution of Boerhavia erecta in Namibia.

5. Boerhavia diffusa Linnaeus (1753: 3) ≡ Boerhavia repens L. var. diffusa (L.) Hooker f. (1885: 709)

Type (conserved type proposed by Whitehouse 1998: 873):—Virgin Islands, St. Coix, Teague Bay, West Indies laboratory, 30 May 1977, Fosberg 56776 (holotype BM-000593477!, isotypes B-100346000!, BISH, GH-00460603!, K, MO, NSW). Image available from http://plants.jstor.org/stable/10.5555/al.ap.specimen.bm000593477 = Boerhavia paniculata Richard (1792: 105) ≡ Boerhavia decumbens Vahl (1804: 284). Boerhaavia laxa Persoon (1805: 36). Type:—French Guiana. Cayenne, Leblond 239 (holotype G). = Boerhaavia adscendens Willdenow (1797: 19). Type:—Guinea, Isert s.n. in Herb. Willdenow 768 (holotype BW).

Figure 11. Known distribution of Boerhavia diffusa in Namibia.

Description:—Stems decumbent, diffuse or prostrate, glabrate to puberulous. Leaves with petioles (8–)10–30(–33) mm long; lamina elliptic, lanceolate, ovate, oblong, orbicular or obovate, (18–)21–58(–61) × (11–)13–40(–46) mm; apex apiculate, acuminate, rounded or obtuse; base shortly attenuate or obtuse; margins entire, sometimes undulate; glabrous with long septate hairs along margins and scattered along veins. Inflorescence with primary peduncles 60–150 mm long, secondary peduncles 25–60 mm long, glabrate, 2–5 flowers per cluster; pedicels 1 mm long. Flowers 2–3 mm long; lower part of perianth 1 mm long, ellipsoid-clavate, 5-ribbed, glandular hairs present; upper part of perianth

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 115 1–2 mm long, pink, purple or maroon. Stamens 2 or 3, slightly exserted, 1.5–2.5 mm long. Ovary 0.5 mm long; style 1–1.5 mm long. Anthocarp 3–4 × 1–2 mm, ellipsoid-clavate, 5-ribbed, ribs indented near apex, apex ending in pointed tip, glandular hairs present on ribs. (Fig. 6D). Habitat:—Grows in moist, well-drained to dry, stony, sandy, loamy or gravelly soil on riverbanks, floodplains, plains or coastal dunes, often along roadsides in calcrete, limestone or basalt. elevation:— about 5–1370 m. Flowering time:—January to December. Distribution:—Namibia (Fig. 11), Botswana, South Africa, Mexico, West Indies, Central America, South America, Asia, Indian Ocean Islands, Pacific Islands and Austalia. Specimens examined:—Namibia . Tsumeb, main street in front of Engen, 1917BA, 10 February 2009, Struwig 74 (PUC, WIND); Near homestead on Taranaki farm, 84 km from Grootfontein on road to Rundu, 1918BA, 06 March 1995, Germishuizen 7575 (PRE, WIND); Khorixas Rest Camp, 2014BD, 08 February 2009, Struwig 70 (PUC, WIND); Omaruru, main street, in die middle of the road, 2115BD, 11 April 2010, Struwig 174 (PUC, WIND); Omaruru, Loskop farm, 2115BD, 11 April 2010, Struwig 178 (PUC, WIND).

Figure 12. Boerhavia orbicularifolia. A. Habit. Scale bar 1 mm; B. Flower. C. Anthocarp. Scale bars B–C, 1.25 mm. Drawings prepared by Gillian Condy.

Type:—Namibia . Okaukuejo. 15 km S of Okaukuejo, 6 March 1976, Giess & Loutit 14128 (holotype WIND!, isotype PRE).

Description:—Annual. Roots up to 100 mm long, swollen, turnip-shaped. Stems herbaceous, puberulent, branching directly from base, decumbent, 100–200 mm tall, not spreading. Leaves with petioles (9–)11–27(–35) mm long; lamina orbicular, oval, oblong or lanceolate, (12–)14–20(–23) × (8–)10–15(–17) mm; apex rounded or sometimes apiculate; base obtuse or rounded; margins entire, sometimes undulate; glabrous, often with septate hairs along margins. Flowers solitary or in an inflorescence. Inflorescence an umbel, peduncles 2 mm; 1–4 flowers per cluster; pedicels ± 1 mm long. Flowers 2 mm long; lower part of perianth 1 mm long, ellipsoid, 5-ribbed, puberulous; upper part of perianth 1 mm long, white. Stamens 1. Ovary ellipsoid. Anthocarp 3 × 1 mm, ellipsoid-clavate, 5-ribbed, apex rounded, glabrate with hairs on ribs and between ribs (Fig. 12) Habitat:—Grows in black limestone in vlei-like depressions that stand under water for a while after rain and in stony, sandy-loam soil on arenite in full sun. elevation:— about 1100–1348 m. Flowering time:—March. Distribution:—Namibia (Fig. 13).

Figure 13. Known distribution of Boerhavia orbicularifolia in Namibia.

Specimens examined:—Namibia . 60 km N of Maltahöhe on road to Walvis Bay, 2416DA, 11 March 1995, Burgoyne 3488 (WIND).

Neotype (designated by Meikle & Hewson 1984: 318):—Jamaica. Houston s.n. (BM-000993062!). Image available from http://plants. jstor.org/stable/10.5555/al.ap.specimen.bm000993062 = Boerhavia caribaea Jacquin (1771: 5, t.84). Lectotype (designated by Whitehouse 1996: 6):—[Icon] Boerhavia caribea in Jacquin (1771: t.84). = Boerhaavia viscosa Lagasca & Rodríquez (1801: 256) ≡ Boerhaavia diffusa var. viscosa (Lag. & Rodr.) Heimerl (1897: 27) ≡ Boerhavia coccinea Miller var. viscosa (Lag. & Rodr.) Moscosa (1943: 180). Neotype (designated here):—PERU. Originally from Peru, flowered during June and July in the Royal Botanic Garden, Madrid, Cavanilles s.n. (G-00439928!). Image available at http://www.ville-ge.ch/musinfo/bd/cjb/chg/adetail.php?id=408576&base=img&lang=en = Boerhavia repens var. viscosa Choisy (1849: 453). Type:—[Egypt or Arabia] , “Thal Arbain”, May, Rueppell (holotype FR-0030534!). = Boerhavia diffusa var. hirsuta Heimerl (1889: 10). Type:—South Africa, Northern Cape, Kuruman, Marloth 1102 (holotype B, isotype PRE-0394976-0!). = Boerhaavia diffusa var. paniculata Kuntze (1891: 533). Type:—VIRGIN ISLANDS. St. Thomas, Virgin Island Kuntze 36 (holotype NY-1374969!). = Boerhavia bracteata Cooke (1909: 421). Lectotype (designated by Whitehouse 1996: 6):—South Africa. Mpumalanga: Avoca near Barberton, Galpin 1240 (K-000243774!, isolectotypes PRE-0592865-0!, GRA-0000910-0!). = Boerhaavia coccinea f. parcehirsuta Heimerl in Urban (1912: 212). Lectotype (designated here):—GRENADA. St. George, on dry ground, 20 April 1906, Broadway 3692 (B-100550649!). = Boerhavia marlothii Heimerl: (1889: 10). Type:—NAMIBIA. Hereroland, Otjimbingwe, 1886, Marloth 1372 (holotype B-100484976!, isotype PRE-0394974-0!). − Boerhavia repens L. var. diffusa sensu Baker & C.H.Wright (1909: 5), pro parte, excl. syn. − Boerhavia diffusa sensu F.W.T.A.:152 (1927), non Linneaus. − Boerhavia coccinea Miller var. α sensu Meikle (1954: 178, fig 66).

Description:—Stems decumbent or prostrate, glabrate to densely pubescent, often with long septate hairs. Leaves with petioles (3–)5–15(–25) mm long; lamina elliptic, oblong, ovate, lanceolate or oval, (11–)13–40(–42) × (9–)10–30(–33) mm; apex acute, apiculate or rounded; base shortly attenuate or rounded; margins entire; glabrous to puberulous, often with long septate hairs along veins and margins. Inflorescence with primary peduncles 54–118 mm long, secondary peduncles 32–43 mm long; 3–many flowers per cluster; pedicels 1 mm long. Flowers 1–4 mm long; lower part of perianth 1–1.5 mm long, ellipsoid, 5-ribbed, glandular hairs present; upper part of perianth 1–3 mm long, pink, purple or white. Stamens 1–5, 1.5–4.5 mm long. Ovary 0.4–0.6 mm long; style 1.3–4.0 mm. Anthocarp 3–5 × 1–2 mm, narrowly ellipsoid-clavate, 5-ribbed, with glandular hairs between ribs, ribs themselves glabrous or covered with hairs (Fig. 6A). Habitat:—Grows in stony or sandy soil on riverbanks, hills or mountains and on ridges, often along roadsides; in quartzite, dolomite or dolerite. elevation:— about 350–2000 m. Flowering time:—September to April. Distribution:—Namibia (Fig. 14), Botswana, Swaziland, South Africa, Central America, South America, Mexico, West Indies, Eurasia, Africa and Australia. Typifications:—Lagasca & Rodrígues (1801: 256) originally stated for Boerhavia viscosa, that the species originated from Peru and flowered in the Royal Botanic Garden of Madrid. One specimen at G (barcode 00439928) bears one piece of plant with morphological characteristics matching the diagnosis. A label (bottom-left of the sheet) by N. Fumeaux indicates that the plant was probably cultivated at the Botanical Garden of Madrid, while another label (on the bottom-right) by A.Heimerl report “Typ!”. Since no effective publication by Heimerl were found his typification is not effective and we here propose to designate the specimen G-00439928 as the lectotype of the name Boerhavia viscosa. Heimerl cited five syntypes for Boerhaavia coccinea f. parcehirsuta in the protologue of which Broadway 3692 is chosen here to serve as the type. Specimens examined:—Namibia . Otjiwarongo, C39, Outjo road, 1916DB, 10 February 2009, Struwig 75 (PRE, WIND); Otjiwarongo, Kudubos, road behind Camp 1, 2016DA, 12 February 2009, Struwig 77 (PRE, WIND); Kalkfeld, Otjiwarongo, at junction of D2404 and D2414, roadside, 2116AB, 12 February 2009, Struwig 81 (PRE,

118 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. WIND); Karibib, on road to Otjimbingwe, roadside, 2116CD, 03 February 2009, Struwig 67 (PUC, WIND); Okahandja, D2110, first dry streambank after entrance to farm Okatjiho, 2116DD, 12 February 2009, Struwig 55 (PUC, WIND); Karibib, on road to Otjimbingwe, 2216AA, 03 February 2009, Struwig 68 (PUC, WIND); North of Maltahöhe, south of Naukluft, Kyffhauser farm, 2416AD, 12 March 1995, Burgoyne 3499 (PRE, WIND).

Figure 14. Known distribution of Boerhavia coccinea in Namibia.

8. Boerhavia repens Linnaeus (1753: 3) subsp. repens

Lectotype (designated by Codd 1966: 121):—Egypt . Herb. Linnaeus 9.8 (LINN!). Image available from http://linnean-online.org/35/ = Boerhavia vulvarifolia Poiret (1804: 55) ≡ Boerhavia repens var. glabra Choisy (1849: 453). Type:—Egypt . Delile (holotype P-00380853!). = Boerhavia repens var. minor Delile (1813: 146) ≡ Boerhavia diffusa L. var. minor (Delile) Cufodontis (1953: 77). Type: Egypt . Delile (holotype MPU-007234!). Image available from http://plants.jstor.org/stable/10.5555/al.ap.specimen.mpu007234

Description:—Annual herb. Stems prostrate, puberulent. Leaves with petioles (3–)5–15(–20) mm long; lamina lanceolate, elliptic, ovate or oblong, (10–)14–29(–30) × (6–)7–15(–16) mm; apex apiculate, acuminate or rounded; base obtuse; margins entire, sometimes undulate; leaf pairs at nodes anisophyllous, glabrate. Inflorescence with primary peduncles 20–24 mm long, secondary peduncles 5–20 mm long; glabrate to puberulent; 2–4 flowers per cluster; pedicels 1 mm long. Flowers 2–3 mm long; lower part of perianth 1–2 mm long, ellipsoid, 5-ribbed, glandular hairs present in grooves between ribs; upper part of perianth 1 mm long, white, pink or purple. Stamens 1 or 2, included,

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 119 0.6–0.8 mm long. Ovary 0.3–0.5 mm long; style 0.6–1.2 mm long, exserted. Anthocarp 3–4 × 1–2 mm, ellipsoid- clavate, 5-ribbed, rounded at apex, glandular hairs present in area between ribs and rarely on the ribs (Fig. 6G). Habitat:—Grows in alluvial, stony, sandy, gravelly, loamy or clay soil in pans or depressions, riverbeds, plains or hill slopes, often along roadsides. elevation:— about 557–1370 m. Flowering time:—November to May. Distribution:—Namibia (Fig. 15), Africa and tropical Asia.

Figure 15. Known distribution of Boerhavia repens subsp. repens in Namibia.

Specimens examined:—Namibia . Kunene, Mahenene Research Station fields, 1714DB, 15 December 1999, Kolberg HK1006 (PRE, WIND); Twyfelfontein lodge, at visitors parking, 2014CB, 07 February 2009, Struwig 69 (PUC, WIND); Outjo district, Fransfontein, beneath dolomite mountain in rock cave, gate to the north, 2016BA, 13 March 1974, Merxmüller & Giess 30306 (PRE, WIND); M47, Capricorn Rest Camp, camp left of restrooms, 2416BA, 08 April 2010, Struwig 161 (PUC, WIND); Maltahöhe, D850, roadside, 2416DA, 09 April 2010, Struwig 170 (PUC, WIND); Maltahöhe, D850, roadside, 2416DB, 09 April 2010, Struwig 168 (PUC, WIND); Mariental District, 70 km from Mariental, Voigtsgrund farm, 2417CD, 12 March 1983, Immelmanm 588 (PRE).

Commicarpus Standley (1909: 373)

Type (designated by Howard 1988: 178): Commicarpus scandens (L.) Standley (as Boerhavia scandens Linnaeus).

Description:—Perennial herbs or sub-shrubs. Stems long, slender, erect, climbing, often much branched, often woody

120 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. towards the base. Leaves opposite, petiolate, glabrous to puberulous; lamina cordate, deltoid, lanceolate, orbicular, ovate; apex acute, apiculate, acuminate, rarely rounded or retuse, base cordate to subcordate, shortly attenuate, rounded, obtuse, truncate; often fleshy, margins entire, sometimes repand. Inflorescence umbellate, sometimes verticillate; glabrous to puberulous; bracteate. Flowers pedicellate; flowers (4–)5–21(–23) mm long, bisexual. Perianth divided into lower and upper part; lower part 2–3 mm long, green, sometimes reddish, coriaceous, constricted above the ovary, cylindrical, clavate or broadly clavate, glabrous or with glandular hairs, with ten narrow, longitudinal grooves, five to ten prominent glands around the apex and less prominent glands scattered over the surface further below, persistent; upper part (2–)3–15(–17) mm long, infundibuliform, petaloid, pink, purple or white, sometimes with distinct basal tube, caducous after anthesis. Stamens 2–4, exserted; filaments filiform, connate at the base into a short sheath around the ovary; anthers dithecous. Ovary ellipsoid, stipitate, one ovule; style filiform, exserted; stigma capitate. Anthocarp (3–)5–8(–9) × 2–3 mm, 10-ribbed, clavate, elliptic-clavate or fusiform, apex surrounded by ten stalked glands or five stalked glands alternating with five sessile glands, sessile wart-like glands scattered over the surface below the apex; glabrous or covered with glandular hairs; secretes a hygroscopic mucoidal secretion which is produced in columnar cells in the ribs, especially when wetted. Seed oblong, brown, with smooth surface; endosperm scanty, embryo hooked. Notes:—Commicarpus is a genus of ± 30–35 species distributed worldwide throughout the tropical and subtropical regions with most of the species occurring in Africa and Western Asia (Douglas & Spellenberg 2010). North-eastern African and southern Arabia form the centre of diversity for this genus (Thulin 1990). Eight species of Commicarpus occur in southern Africa with three species (C. decipiens Meikle, C. fallacissimus (Heimerl) Heimerl ex. Oberm., Schweick. & I.Verd. and C. squarrosus (Heimerl) Standl.) endemic to Namibia (Makwarela 2003). Commicarpus chinensis (L.) Heimerl subsp. natalensis Meikle however, does not occur in Namibia, but along the northern coast of KwaZulu-Natal, South Africa; Mozambique and the eastern seaboard of tropical Africa (Makwarela 2003, Klopper et al. 2006) (see fig. 6H for anthocarp comparison).

Key to the species of Namibian Commicarpus (adapted and expanded from Struwig et al. 2011):

1. Shape of the anthocarp is elliptic-clavate, tapering to both ends; lower coriaceous part of the flower elliptic with five prominent glands around the apex...... 1. C. pilosus - Shape of the anthocarp is fusiform or clavate, not tapering or tapers noticeably from apex to base; lower coriaceous part of the flower with usually more than 5 prominent glands (up to 10) around the apex...... 2 2. Anthocarp fusiform; lower coriaceous part of the flower cylindrical or broadly elliptic...... 3 - Anthocarp clavate; lower coriaceous part of the flower clavate or elliptic...... 5 3. Ring of five stalked glands alternating with five sessile glands around the apex, anthocarp 4–6 mm long; lower coriaceous part of the flower broadly elliptic with 5 shortly stalked glands alternating with 5 sessile less prominent glands around the apex...... 2. C. squarrosus - Ring of ten stalked glands of the same length around the apex, anthocarp 7–9 mm long; lower coriaceous part of the flower cylindrical with 10 prominent glands around the apex...... 4 4. Surface of the anthocarp below the apex covered sparsely with shortly stalked (<1 mm long) glands grouped together; leaves cordate, ovate and deltoid...... 3. C. plumbagineus var. plumbagineus - Surface of the anthocarp below the apex covered with scattered sessile glands; leaves lanceolate-ovate...... 4. C. fallacissimus 5. Anthocarp broadly clavate, surface covered with prominent glandular hairs, apex surrounded by five shortly (<1 mm) stalked glands; lower coriaceous part of the flower clavate, surface covered with prominent glandular hairs; leaves lanceolate-ovate...... 5. C. decipiens - Anthocarp clavate; surface glabrous to puberulent, apex surrounded by five stalked glands alternating with five sessile glands; lower coriaceous part of the flower elliptic or clavate with one or two rows of five prominent glands around the apex; leaves cordate, ovate and deltoid...... 6 6. Lower coriaceous part of the flower elliptic; anthocarp tapers noticeably from apex to base, apex surrounded by five thin, long stalked glands (up to 2 mm long)...... 6. C. helenae var. helenae - Lower coriaceous part of the flower clavate; anthocarp does not taper noticeably from apex to base, apex surrounded by five thickly stalked glands, stalks c. 1 mm long...... 7. C. pentandrus

1. Commicarpus pilosus (Heimerl) Meikle (1978: 249) ≡ Commicarpus fallacissimus (Heimerl) Heimerl forma pilosus Heimerl (1937: 233)

Type:—South Africa. Transvaal, near Vivo, 15 April 1934, Schweickerdt & Verdoorn 653 (holotype PRE!, isotype K-000243814). Image available from http://plants.jstor.org/specimen/pre0393215-0

Description:—Herbs to sub-shrubs of up to 1 m high, pubescent, woody toward base, older stems white, decumbent.

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 121 Leaves with petioles (7–)8–16(–20) mm long; lamina cordate, ovate, oval, orbicular or oblong, (12–)16–40(–49) × (10–)12–38(–39) mm, apex apiculate, rounded or acute, base shortly attenuate to rounded, truncate or cuneate; slightly fleshy; pubescent. Inflorescence with peduncles 20–105 mm long; six flowers per umbel; pubescent; pedicels 1–5 mm long. Flowers 5–12 mm long; lower part of perianth 2–3 mm long, elliptic, five prominent sessile glands around the apex with smaller, less prominent glands scattered over the surface below; upper part of perianth 3–9 mm long, pink or purple. Stamens 2 or 3, filaments 6–10 mm long; anthers 0.5–0.8 mm long, transversely elliptic to subcircular. Ovary 0.65–0.75 mm long; style 5–7 mm, long exserted. Anthocarp 5–7 × 1–2 mm, 10-ribbed, elliptic-clavate tapering to both ends, five shortly stalked glands around apex, sessile glands scattered over surface below apex; glabrous (Fig. 6M). Habitat:—Grows in stony soil, gravel, clay or sand on plains, hill- or mountain slopes on calcrete, quartzite, sandstone or dolerite. elevation:— about 200–1350 m. Flowering time:—October to May. Distribution:—Namibia (Fig. 16), Botswana, South Africa and Zimbabwe.

Figure 16. Known distribution of Commicarpus pilosus in Namibia.

Specimens examined:—Namibia .—Otjimbingwe, Otjozondu farm, marble ridge on the shore of Omusema, 2216AA, 01 March 1967, Seydel 4488 (PRE); Otjimbingwe, Otjozondu farm, on the mountain 2216AA, 12 April 1965, Seydel 4344 (PRE).

122 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. Figure 17. Commicarpus squarrosus. A. Flowering and fruiting branches. Scale bar 10 mm. B. Flowers. C. Anthocarp. Scale bars B–C, 1.67 mm. Drawings prepared by Gillian Condy.

Type:—Namibia. Gross-Namaland, Rehoboth, Fleck 241A (holotype Z-00016202!).

Key to the varieties of Commicarpus squarrosus (Struwig & Siebert 2013a):

1. Flowers pink; widely distributed from the Kunene to Karas regions; occurs on sandy soil...... C. squarrosus var. squarrosus - Flowers purple; restricted to Hardap region; occurs on loamy, Ca- and Mg-rich soils with high concentrations of heavy metals...... C. squarrosus var. fruticosus

Figure 18. Known distribution of Commicarpus squarrosus var. squarrosus in Namibia.

1. Commicarpus squarrosus (Heimerl) Standley var. squarrosus

Description:—Much branched subshrub, young shoots purplish, older branches whitish, up to 1 m high; pubescent to glabrescent. Leaves with petiole (8–)9–18(–23) mm long; lamina elliptic, ovate, orbicular, trullate, oval or lanceolate, (10–)11–25(–30) × (8–)10–23(–27) mm, apex apiculate, rounded, retuse or emarginate, base attenuate; fleshy, sometimes reddish, pubescent. Inflorescence with five flowers per umbel; peduncles 30–90 mm long, glabrescent; pedicels (8–)12–25(–26) mm long, tinged purple. Flowers 5–8 mm long; lower part of perianth 2 mm long, broadly elliptic, green-purplish, five shortly stalked, pink, prominent glands alternating with five sessile, less prominent glands around the apex; smaller, less prominent glands scattered over the surface below; upper part of perianth 3–6 mm long, pink or purple. Stamens 2 or 3, filaments 5 mm long. Ovary and style 6 mm long. Anthocarp 4–6 × 1–2 mm, fusiform, apex surrounded by five stalked glands alternating with five sessile glands, sessile glands scattered over surface below apex; glabrous (Fig. 6O and 17).

124 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. Habitat:—Grows in stony soil, gravel or sandy loam on hill- or mountain slopes on dolomite and granite. elevation:— about 655–1200 m. Flowering time:—December to May. Distribution:—Endemic to Namibia (Fig. 18). Specimens examined:—Namibia. Khowarib Rest Camp, hills surrounding rest camp, 1913BD, 09 February 2009, Struwig 45 (PUC, WIND); Twyfelfontein Lodge, hillslope behind lodge, SW of main building, 2014CB, 07 February 2009, Struwig 41 (PUC, WIND); Numas Gorge, high up in kloof, underneath Acacia montis-usti behind big boulders, 2114BA, 06 February 2009, Struwig 39 (PUC, WIND); Brandberg Mountain, on top, Baswald Rinne Area, 2114BC, 19 May 1977, Craven 490 (PUC, WIND); Klein Spitzkoppe, mountain slope amongst rocks, 2115CC, 05 February 2009, Struwig 36 (PUC, WIND); Swakopmund district, 63 miles south-east of Walvis Bay in Namib area, 2214DA, 02 March 1965, Barnard 85 (PRE, WIND); Tsaobis Leopard Farm, south of Karibib, 2215DD, 21 February 1990, Hardy 7017 (PRE, WIND); Tsais-Maltahöhe farm, 2416AB, 16 May 1978, Müller & Tilson 894 (PRE, WIND); C19. Tsaris Mountains, 2416AB, 08 April 2010, Struwig 164 (PUC, WIND); Maltahöhe, Mooirivier MAL 160 farm, on south facing slopes, 2416CA, 11 April 1980, Müller 1362 (PRE, WIND); Kuibis, 2616DD, 01 March 1912, Range 1283 (BOL).

Figure 19. Known distribution of Commicarpus squarrosus var. fruticosus in Namibia.

Commicarpus squarrosus (Heimerl) Standley var. fruticosus (Pohnert) Struwig (2013a: 63) ≡ Commicarpus fruticosus Pohnert (1953: 337)

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 125 Type:—Namibia . Tsarisberge, Walter 2095 (holotype M-0107832, isotype WIND!). Image available from http://plants.jstor.org/ stable/10.5555/al.ap.specimen.m0107832

Description:—The var. fruticosus differs from the nominal one in having the flowers purple. Habitat:—Grows in stony soil, gravel or sandy loam on hill- or mountain slopes on dolomite and shales. elevation:— about 1008–1370 m. Flowering time:—December to May. Distribution:—Endemic to Namibia (Fig. 19). Specimens examined:—Namibia. Naukluft Mountains at Büllspoort, 2416AA, 16 December 1947, Rodin 2833 & Strey 2132 (BOL); C14, Naukluft Mountains, mountain slope behind the river, 2416AA, 13 February 2009, 07 April 2010, 08 April 2010, Struwig 59, 160, 163 (PUC, PRE, WIND).

3. Commicarpus plumbagineus (Cavanilles) Standley (1916: 101) var. plumbagineus ≡ Boerhavia plumbaginea Cavanilles (1793: 7, t.112)

Figure 20. Known distribution of Commicarpus plumbagineus var. plumbagineus in Namibia.

Lectotype (designated by Meikle 1978: 244):—Spain. Orcellitani, at foot of the mountain (Orihuela), 14 July 1791, Cavanilles s.n. (MA- 475446!) Image available from http://plants.jstor.org/stable/10.5555/al.ap.specimen.ma475446 = Boerhavia dichotoma Vahl (1804: 290). Type:—Arabia . Forsskål (holotype C). = Boerhavia verticillata Poiret (1804: 56) ≡ Commicarpus verticillatus (Poir.) Standley (1916: 101).

126 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. Type:—Senegal. Roussillon 83 (holotype P-LA-00380848!). = Boerhavia commersonii Baillon (1885: 484) ≡ Commicarpus commersonii (Baill.) Cavaco (1953: 297). Type:—Madagascar. N. Madagascar, without precise locality, Commerson (holotype FI). − Boerhavia africana sensu Merrill (1935: 150) non Loureiro. − Commicarpus africanus sensu Dandy (1950: 152), quoad descr., non C. africanus (Lour.) Dandy.

Description:—Herbs procumbent, prostrate or scrambling (2 m high), pubescent to glabrescent, stems up to 1 m long. Leaves with petiole (7–)8–29(–31) mm long, lamina cordate, elliptic, orbicular, ovate or lanceolate, (26–)28–84(–87) × (16–)17–67(–68) mm; apex apiculate, rounded or acute; base obtuse, almost truncate, subcordate, shortly attenuate or cuneate; pubescent to glabrescent. Inflorescence with 12 flowers per umbel; peduncles 38–85 mm long pubescent; pedicels 3–6 mm long. Flowers 10–20 mm long; lower part of perianth 2–7 mm long, cylindrical, apex surrounded by ten prominent, sessile glands with a few glands scattered on the surface below; upper part of perianth 7–14 mm long, white. Stamens 3–5, filaments 12–18.5 mm long, anthers 0.6–0.8 mm long. Ovary 0.75–1 mm long; style 15-18 mm. Anthocarp 8–9 × 1–2 mm, fusiform, with ring of ten stalked glands around apex and few shortly stalked glands grouped below apex; pubescent (Fig. 6N). Habitat:—Grows in well-drained to moist stony soil, sand, loam, gravel or clay on streambanks, valleys, plains and on hill- or mountain slopes on granite, quartzite or dolerite elevation:— about 50–1420 m. Flowering time:—January to December. Distribution:—Namibia (Fig. 20), Botswana, Swaziland, South Africa, Spain, Palestine, Arabia, Senegal, Nigeria, Chad, Rwanda, Burundi, Zaire, Cameroon, Sudan, Eritrea, Ethiopia, Somalia, Uganda, Kenya, Tanzania, Mozambique, Malawi, Zambia, Angola and Madagascar. Specimens examined:—Kaokoveld, on the banks of the Otjitanga river, north of the Anglo American geologist base camp, 1713AD, 10 March 2003, Bartsch SB1299 (WIND); Katima Mulilo, 15 km from Bukalo at turn-off to Muyako, along main road, 1724DA, 28 Februay 1982, Müller 1805 (WIND); Popa Falls Rest Camp all within the Popa Falls Rest Camp, 1821BA, 21 February 1988, Hines 953 (PRE; WIND).

4. Commicarpus fallacissimus (Heimerl) Heimerl ex. Obermeyer, Schweickerdt & I.Verdoorn (1937: 233) ≡ Boerhavia fallacissima Heimerl ex Schinz (1890: 223) ≡ Commicarpus fallacissimus (Heimerl) Pohnert (1953: 337), nom illeg. (art. 52 of ICN, McNeill et al. 2012)

Type:—Namibia , Hereroland, Lüderitz 171 (holotype B!, isotypes B-100354878, K-000243813!). Image available from http:// herbarium.bgbm.org/object/B100354878.

Description:—Herb to sub-shrub 1 m high, branches upright, decumbent; glabrous to pubescent. Leaves with petioles (5–)6–17(–20) mm long; lamina ovate, elliptic or cordate, (24–)25–45(–48) × (8–)10–27(–28) mm, apex acute or apiculate, base shortly attenuate, obtuse, cordate or almost truncate; glabrous to sparsely pubescent. Inflorescence with 5 or 6 flowers per umbel; peduncles 28–70 mm long, glabrate; pedicels 5–7 mm long. Flowers 8–11 mm long; lower part of perianth 3–5 mm long, cylindrical; ten shortly stalked, prominent glands around the apex and sessile, less prominent glands scattered on the surface below; upper part of perianth 5–6 mm long, pink or purple. Stamens 2. Ovary style long exserted. Anthocarp 8–9 × 1–3 mm, fusiform, apex surrounded by ten stalked glands, sessile glands scattered over surface below apex, glabrous (Fig. 6J). Habitat:—Grows in well-drained, sandy to loamy, gravelly or stony soil on hill- or mountain slopes with shale, dolomite, granite or calcrete. elevation:— about 152–1651 m. Flowering time:—November to May. Distribution:—Endemic to Namibia (Fig. 21). Specimens examined:—Namibia . Joubertpass to Opuwo, against slope in stony/rocky soil, 1813DD, 09 February 2009, Struwig 46 (PUC, WIND); Near Merwe farm, about 20 km west of Otavi on road to Outjo, mountain slope, 1916DB, 09 March 1997, Germishuizen 9553 (PRE); Windhoek, National Botanical Garden, Lily Walk, 2217CA, 03 February 2009, Struwig 33 (PUC, WIND); C24, after the crossing of the D2161 and C24, 2316CD, 13 February 2009, Struwig 58 (PUC, WIND); C14, Naukluft Mountains, mountain slope behind river, 2416AB, 08 April 2010, Struwig 165 (PUC, WIND).

5. Commicarpus decipiens Meikle (1978: 246)

Type:—Namibia . Okahandja, 7 May 1907, Dinter 167 (holotype K-000243811!, isotypes BM-000910378, E, SAM) Image available from http://specimens.kew.org/herbarium/K000243811

Description:—Herbs up to 2 m high, pubescent. Stems erect, decumbent or scrambling, woody towards base. Leaves with petiole (5–)6–10(–14) mm long; lamina ovate, elliptic, (20–)22–50(–57) × (10–)12–28(–37) mm; apex acute, acuminate or apiculate; base rounded to shortly attenuate; pubescent. Inflorescence with 10 flowers per umbel; peduncles 23–70 mm long, pubescent; pedicels 2–8 mm long. Flowers 11–21 mm long; lower part of perianth 2–6 mm long, clavate; prominent, sessile glands and trichomes scattered over surface; upper part of perianth 8–16 mm long, white. Stamens 2 or 3, filaments 20 mm long; anthers 0.8 × 1 mm, kidney-shaped. Ovary style long exerted. Anthocarp 5–8 × 2–7 mm, broadly clavate, apex surrounded by five semi-sessile glands, prominent sessile glands scattered over surface below apex, surface covered with prominent glandular hairs (Fig. 6I). Habitat:—Grows in stony soil or humus rich sandy loam on streambanks or hill- and mountain slopes, often underneath trees. elevation:— about 1449–1544 m. Flowering time:—October to June. Distribution:—Endemic to Namibia (Fig. 22).

Specimens examined:—Namibia . Tsumeb, at junction of D3022 and D2863, Otavi mountains, 1917BD, 10 February 2009, Struwig 47 (PUC, WIND); Otjiwarongo District, road Waterberg-Otjiwarongo, Okosongomingo farm 149, 55km south-east of Otjiwarongo, 2017CA, 03 April 1968, Wanntorp &Wanntorp 605 (PRE); Otjiwarongo, Klein Waterberg, mountain slope amongst rocks, 2017CA, 11 February 2009, Struwig 51 (PUC, WIND); Otjiwarongo, Klein Waterberg, at foot of mountain in shade of trees, 2017CA, 11 April 2010, Struwig 181 (PUC, WIND); Omaruru, Loskop farm, 2115BD, 11 April 2010, Struwig 176 (PUC, WIND); Omaruru district, Kalkfeld, Okozongoro farm, at farm dam, 2116AA, 03 March 1983, Immelman 453 (PRE); Okahandja, D2110, first dry streambank after entrance to Okatjiho farm, 2116AB, 12 February 2009, Struwig 54 (PUC, WIND); Windhoek, 2217CA, 17 December 1962, Hanekom 351 (PRE).

6. Commicarpus helenae (Roemer & Schultes) Meikle (1971: t.3694) var. helenae ≡ Boerhavia helenae Roemer & Schultes (1822: 73)

Neotype (designated by Whitehouse 1996: 14):—St. Helena, Ladder Hill, Burchell 1 (K-000243790!) Image available from http:// apps.kew.org/herbcat/getImage.do?imageBarcode=K000243790 = Boerhavia stellata Wight (1846: 6, t.875); ≡ Commicarpus stellatus (Wight) Berhaut (1953: 51). Lectotype (designated here):—[Icon] Boerhavia stellata in Wight (1846: t.875). − Commicarpus verticillatus sensu Baker & Wright (1909: 6), sensu Hutchinson & Dalziel: (1927: 153), sensu Heimerl (1934: 117), sensu Balle (1951: 86, t.7), sensu Meikle (1954: 177).

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 129 Description:—Herbs 0.5 m high, glabrous to pubescent, branches upright, procumbent. Leaves petiole (4–)5–15(– 22) mm long; lamina deltoid, ovate, (18–)19–45(–58) × (13–)15–38(–44) mm; apex apiculate, rounded to acute or acuminate; base shortly attenuate, cordate, subcordate, obtuse or truncate; slightly fleshy; glabrous to sparsely pubescent. Inflorescence with five flowers per umbel; peduncles 35–120 mm long, pubescent; pedicels 1–2 mm. Flowers 4–7 mm long; lower part of perianth 2–4 mm long, elliptic; with five shortly stalked, prominent glands around the apex, smaller, less prominent glands scattered over the surface below; upper part of perianth 2–4 mm long, light pink to almost white. Stamens 2 or 3, filaments 2–6 mm long; anthers 0.3–0.5 mm long, transversely elliptic to round. Ovary 0.5–0.6 mm long; style 3–3.5 mm. Anthocarp 5–6 × 2 mm, clavate, tapering noticeably from apex to base, apex surrounded by five thinly, long (1–2 mm long) stalked glands alternating with five sessile glands, 1 or 2 rings of sessile glands on surface below apex, glabrous (Fig. 6K). Habitat:—Grows in well drained, stony, sandy, loamy or gravelly soil in calcrete or sandstone on riverbanks and floodplains or edges of pans and depressions. elevation:— about 200–1247 m. Flowering time:—November to May. Distribution:—Namibia (Fig. 23), Botswana, South Africa, Iran, Arabia, Egypt, Canaries, India, Cape Verde Islands, St. Helena, Tibesti, Zaire, Sudan, Eritrea, Ethiopia, Somalia, Uganda, Kenya, Tanzania and Angola.

Figure 23. Known distribution of Commicarpus helenae var. helenae in Namibia.

Typification:—The illustration of Boerhavia stellata in Wight (1846: t.875) is part of the original material and it is here designate as the lectotype. Specimens examined:—Namibia . Omitengundi, 500 m north of road junction, 1713AB, 09 April 1998, Schulte

130 • Phytotaxa 238 (2) © 2015 Magnolia Press STRUWIG ET AL. 108530 (WIND); Khowarib Rest Camp, underneath Acacia and mopane trees, 1913BD, 09 February 2009, Struwig 44 (PUC, WIND); Etosha National Park, Charitsaubplain, near pan with Acacia reficiens, 1916AA, 18 April 1973, Le Roux 520 (WIND, PRE); Spitzkoppe, 2115DC, 03 March 1985, Craven 2022 (WIND); Okomitundu farm, underneath Acacia trees in front of gate to house, 2216AB, 12 April 2010, Struwig 183 (PUC, WIND).

7. Commicarpus pentandrus (Burchell) Heimerl (1934: 117) ≡ Boerhavia pentandra Burchell (1822: 432) ≡ Boerhavia burchellii Choisy (1849: 455), nom. superfl. et illeg. (art. 52 of ICN, McNeill et al. 2012)

Type:—South Africa. Northern Cape, along the Vaal River, Griqualand West, November 1811, Burchell 1765 (holotype K- 000243804!) Image available from http://apps.kew.org/herbcat/getImage.do?imageBarcode=K000243804 = Boerhavia transvaalensis Gandoyer (1919: 221). Type: SOUTH AFRICA. Transvaal, Marabastad, Schlechter 4339 (holotype LY, isotype K-000243806!) Image available from http://apps. kew.org/herbcat/getImage.do?imageBarcode=K000243806

Figure 24. Known distribution of Commicarpus pentandrus in Namibia.

Description:—Prostrate or procumbent herbs up to 1 m, from a woody rootstock up to 30 mm thick and ± 300 mm long, pubescent. Stems sometimes tinged purple. Leaves with petiole (4–)5–11(–12) mm long; lamina ovate, elliptic, orbicular, deltoid, (13–)15–30(–39) × (13–)14–26(–29) mm; apex apiculate, rounded to acute; base cordate, truncate, cuneate, rounded, obtuse, subcordate or shortly attenuate; slightly fleshy, upper sides darker than undersides; sparsely pubescent. Inflorescence with six flowers per umbel; peduncles 30–150 mm long, pubescent; pedicels 2–18 mm.

A taxonomic treatment for the Flora of Namibia Phytotaxa 238 (2) © 2015 Magnolia Press • 131 Flowers 12–23 mm long; lower part of perianth 2–6 mm long, clavate, with one or two rows of five prominent sessile glands around the apex with smaller, less prominent glands scattered over the surface below; upper part of perianth 9–17 mm long, purple or pink. Stamens 4–6, filaments 11–17 mm long, anthers 0.7–0.8 × 1.2–1.5 mm, transversely elliptic. Ovary 0.75–1 mm long; style 14–20 mm. Anthocarp 7–9 × 2–3 mm, clavate, five thickly stalked glands alternating with five smaller, less prominent glands around the apex, sessile glands scattered over surface below apex, glabrous (Fig. 6L). Habitat:—Grows in well-drained to moist, stony or gritty soil, gravel, sand, loam, clay or turf, in bare patches, on floodplains, plains, depressions, valleys, hillslopes or outcrops; on dolerite, granite, shale, limestone and calcrete in grassland, savanna and bushveld. elevation:— about 5–1775 m. Flowering time:—October to May. Distribution:—Namibia (Fig. 24), Botswana, South Africa, Swaziland, Lesotho, Mozambique, Zimbabwe and Zambia. Specimens examined:—Namibia . Outjo district, Etosha National Park, near Homob waterhole, 1916AA, 02 November 1966, Tinley 12856 (PRE, UNIN); between Outjo and Etosha, 1916CC, 26 February 1959, Wedermann & Oberdieck 2318 (PRE); Tsumeb, at junction of D3022 and D2863, Otavi mountains, 1917BD, 10 February 2009, Struwig 48 (PUC, WIND); At Gaub farm on road to Grootfontein, 1917BD, 03 March 1995, Germishuizen 7431 (PRE); Otjiwarongo, 8 km S of Otjiwarongo, 2016BC, 14 March 1973, Giess 12612 (PRE); Otjiwarongo, Gobabis road, corner of C30 and C22, 2016DB, 11 February 2009, Struwig 52 (PUC, WIND); Windhoek district, 46 miles east of Windhoek on road to Gobabis, 2217CA, 25 February 1955, De Winter 2527 (PRE); D1463, at T-junction sign just before entrance to Aris farm, 2217CC, 13 February 2009, Struwig 57 (PUC, WIND).

Acknowledgements

The Directors and Curators of the cited herbaria are acknowledged for giving access to the study material. The Research Directorate, Unit for Environmental Sciences and Management of the North-West University and the South African Biosystematics Initiative (SABI) provided financial support. Ms. G. Condy (South African National Biodiversity Institute) for preparing the drawings. The staff of Windhoek National Herbarium for testing the keys.

References

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