Genus Ekebergia
Ekebergia Sparrm. in Kongl. Vetensk. Acad. Handl. 40: 282 (1779); Sonder in Fl. Cap. 1: 247 (1860); White and Styles in F.Z. 2: 315 (1963); White in Bothalia 16 (2): 155 (1986); White and Styles in Fl. SA. 18 (3): 51 (1986); Leistner, Seed Pl. S. Afr.: 259 (2000); Coates Palgrave, Trees Southern Afr.: 451 (2002). Type species: Ekebergia capensis Sparrm.
Deciduous trees or shrubs. Leaves: imparipinate; leaflets are entire, subopposite, or opposite; leaf rachis occasionally winged. Flowers: unisexual, in axillary panicles. Calyx: short, 4–5-lobed, obtuse, imbricate in aestivation. Petals 4–5, barely elongated than the calyx, imbricate in aestivation, elliptical or oblong. Stamens 10 (occasionally 9) united in a campanulate 10-toothed antheriferous short tube; anthers sessile. Ovary: 4–5-celled, sessile, surrounded by an annular disc; ovules in pairs in each locule; style short, thick; stigma discoid, obscurely four or five-lobed. Fruit: (dry berry), globose, 4–5-celled. Seeds: mostly one per locule, exalbuminous; cotyledons thick, fleshy; radicle superior.
Key to the species
1a. Leaf petiole and rachis not winged; prismatic crystals in ray cells of bark—E. Capensis.
1b. Leaf petiole and rachis winged; prismatic crystals absent from ray cells of bark—E. pterophylla.
Ekebergia capensis
Ekebergia capensis Sparrm. in Kongl. Vetensk. Acad. Handl. 40: 282 (1779); Thunb., Fl. Cap. 542 (1823); Sonder in Fl. Cap. 1: 247 (1860); Pappe, Silv. Cap. 1: 6 (1862); C. DC. in Monogr. Phan. 1: 641 (1878); Butt-Davy, Fl. Transv. 2: 487 (1932); White and Styles in F.Z. 2: 315 (1963); Ross, Fl. Natal 216 (1972); Palmer and Pitman, Trees S. Afr. 2: 1065 (1973); White in Bothalia 16: 155 (1986); White and Styles in Fl. SA. 18 (3): 51 (1986). Type: South Africa, Western Cape, ‘ad Cap. B.-Spei’, without date, Sparrman s.n. (S-image!, holotype).
Ekebergia buchananii Harms in Engl. Bot. Jahrb. 23: 164 (1896). Type: Nyasaland, 1891, Buchanan 39 (B, holotype†; K, isotype).
Ekebergia meyeri Presl ex C. DC. in Monogr. Phan. 1: 642 (1878); Sim in For. Fl. Port. E. Afr. 26: 20 (1909); Bak. in Journ. Linn. Soc. Bot. 40: 38 (1911); Eyles in Trans. Roy. Soc. S. Afr. 5: 389 (1916); Garcia in Fl. Mocamb. 2: 141 (1954). Type: South Africa, KwaZulu-Natal, Port Natal, in Waldern und Holzungen, April 1838, Drège b (S).
Ekebergia mildbraedii Harms in Notizbl. Bot. Gart. Berlin-Dahlem 7: 229 (1917); Staner and Gilbert, F.C.B. 7: 210 (1958); White in Bothalia 16 (2): 155 (1986). Type: Cameroon, Sanga, Übergangs grenze der Hylaea südlich des Sanaga zwischen Jaunde und Dengdeng unweit der Vereinigung von Lom und Djerem. Etwa, ca 121 mi [195 km] NO Jaunde, May 1914, Mildbraed 8479 (K-image!, holotype).
Ekebergia ruppeliana (Fresen.) A. Rich. in Tent. Fl. Abyss. 1: 105 (1847); Oliv. in F.T.A. 1: 333 (1868); C. DC. in Monogr. Phan. 1: 643 (1878); Gȕrke in Engl. Pflanzenw. Ost-Afr. 231 (1895); Lebrun in Ess. For. Reg. Mont. Cong. Or. 2: 112 (1935); Brenan in T.T.C.L. 315 (1949); Eggeling and Dale, Indig. Trees Uganda Prot. 174 (1952); Staner and Gilbert, F.C.B. 7: 208 (1958); Dale and Greenway, Kenya Trees and Shrubs 267: 54 (1961). Type: Ethiopia, Abessinien, auf dem Wege von Halei nach Temben, May 1832, Ruppell s.n. (FR).
Ekebergia senegalensis Juss. in Mém. Mus. Hist. Nat. 19: 234 (1830); Exell and Mendonca, C.F.A. 2: 316 (1951); Keay in F.T.W.A. edn 2.1: 705 (1958); Staner and Gilbert, F.C.B. 7: 208 (1958); Styles in E. Afr. Agr. Forestry J. 39: 410 (1974); White in Bothalia 16 (2): 155 (1986). Type: Angola, Distr. Golungo Alto. Sobati de Quilombo, July 1856, Welwitsch 1704 (BM).
Trichilia ekebergia E.Mey. ex Sond. in Harv. and Sond. Fl. Cap. 1: 246 (1860). Type: South Africa, no date, Drège s.n. (BM).
Trichilia rueppelliana Frensen. in Mus. Senckenb. 2: 278 (1837). Type: Ethiopia, Abessinien, auf dem Wege von Halei nach Temben, May 1832, Ruppell s.n. (FR).
Large, evergreen (sometimes semi-deciduous) tree, up to 35 m tall; crown moderately dense, somewhat flattened; bole straight (sometimes bent), up to 1.2 m in diameter.
Bark: surface smooth but rough and scaly in older trees; inner bark reddish (sometimes with white bands); twigs with conspicuous whitish lenticels; cortex with ideoblast cells; dilatation of cortical tissue is by anticlinal division of the cortical parenchyma and collenchyma cells as well as tangential stretching of cells forming strands of 2–11 cells, crystals in phelloderm cells, crystals in axial parenchyma, crystals in ray cells, fibers present in secondary phloem, canal in rays.
Wood: diffuse-porous, vessels slightly angular or rounded in outline, solitary, few, and in radial multiples of 2–4, intervessel pits alternate, small to medium, 4.4–7.7 μm in diameter; fibers non-septate; axial parenchyma scanty paratracheal, up to 12 cells wide; prismatic crystals present in chambered axial parenchyma cells; rays mostly biseriate.
Leaves: imparipinnately compound with 3–7 pairs of leaflets, petiolate, lateral leaflets opposite, ovate to lanceolate, leaflets 20–120 × 20–50 mm, petioles thick at base, 30–120 mm long,
petiolules: 5–15 mm long, rachis up to 230 mm long, stipules absent, leaflet base acute, leaflet apex acuminate.
Inflorescence: a 6–10-flowered or six multi-flowered panicle or raceme.
Flowers: unisexual, regular, small, ca.6 mm in diameter, greenish-white; sepals 5, fused, ca. 2–3 mm long, densely pubescent; petals 5, free, up to 7 mm long, imbricate, tomentose; stamens 5, with filaments joined forming cup-shaped tube; anthers sessile, usually on the tube; ovary superior, sessile, with two ovules in each locule; style single, up to 10 mm long, stigma 4–5-lobed; male flowers with undeveloped ovary; female flowers with non-dehiscing anthers.
Fruit: a drupe, fleshy, up to 30 mm in diameter, globose to ellipsoid, ripens from green to pink to deep red, 2–4-seeded.
Seeds: 5–10 mm long, not arillate, dark brown (
Figure 1).
Distribution and ecology
The species grows in riverine forests, wooded grassland, and savanna woodlands from sea level to about 3000 m. It is mostly found in Limpopo, Gauteng, Mpumalanga, KwaZulu-Natal, the Western Cape, and the Eastern Cape provinces in South Africa (
Figure 2). It is also found in Eswatini, Zimbabwe, Zambia, Sudan, Ethiopia, the Congo, and Uganda.
Diagnostic characters
Ekebergia capensis is similar to E. pterophylla in that the leaves are compound with opposite leaflets ending with a single terminal leaflet. Ekebergia capensis is readily distinguished by its rough light grey to almost black bark with some buttress roots at the base and the glossy green, ovate to lanceolate leaflets, 20–120 × 20–50 mm long, rachis 200–230 mm long, not winged or only very slightly winged, and the presence of prismatic crystals in ray cells. In P. pterophylla, the leaflets are leathery, elliptic to narrowly obovate, 30–50 × 20–30 mm, and crystals are absent from the ray cells (bark tangential section).
Conservation status
Phenology
The species flowers in the summer months (September to December), and fruits ripen between November and April.
Specimens examined
Botswana. 1822 (Kangara): Gomare, fringing forest, near store (–AC), June 1946, Miller B/431 (PRE), Seronga, Okavango River (–CA), 27 September 1954, Story 4758 (PRE). 1922 (Nokaneng): Dikgathong, Thaoge River (–CD), 29 October 1975, Smith 1471 (PRE).
South Africa. LIMPOPO: 2229 (Waterpoort): Zoutpan farm, kloof behind homestead, lower slopes of berg (–CD), November 1932, Obermeyer, Schweickerdt, and Verdoorn 112 (PRE); Makuleke (–CD), 26 October 1932, Lang s.n. (NH), Soutpansberg, Wyllie’s Poort, Ingwe Motel near Wyllie’s Poort (–DD), 8 January 1963, De Winter 7816 (PRE). 2230 (Messina): Sibasa, Thohoyandou, onder kranse (–CC), 21 November 1975, Van der Waal 11 (PRE); Lake Fundudzi, between Thomason’s store and lake (–CD), December 1928, Hutchinson 2165 (PRE); Kruger National Park, NW of Punda Milia, Shipudza (–DB), November 1966, Van Wyk 46027 (PRE). 2327 (Ellisras): Langs pad vanaf Mudzinga-spruit op pad na Seville, farm Seville 250 MT (–DD), 3 November 1979, Van Wyk 2938 (PRE). 2329 (Polokwane): Zoutpansberg, foothills in ravine (–AA), 10 January 1955, Mogg 24553 (PRE); Makhado, Bluegumspoort, ca 12.4 mi [20 km] from main road to Messina on northern slopes of Zoutpansberg (–BB), 1 February 1978, Pienaar 1204 (PRE). 2330 (Tzaneen): Letaba, Duiwelskloof, Modjadjis Reserve near Duiwelskloof (–CA), 15 April 1937, Krige 39 (PRE); Polokwane, between Malati and Malopene (–DD), 22 October 1938, Acocks and Hafstrȍm 1092 (PRE). 2430 (Pilgrim’s Rest): The Downs (–AA), March 1940, Renny 64 (PRE); Letaba, Lekgalameetse, on the farm Malts 65 KT (–AA), 30 November 2017, Turpin 1660 (J); Lekgalameetse Nature Reserve, Schelem, along main road towards Malta, near Selati River, steep slope (–AB), 5 December 1984, Stalmans 304 (PRE); Pilgrims Rest, Mariepskop (–DB), 2 December 1959, Van der Schijff 4714 (PRE). 2431 (Acornhoek): Timbavati Game Reserve, Kempiana farm, on dry riverbank (–AD), 13 August 1978, Kruger 299 (PRE); Timbavati Nature Reserve, Spring Valley, ca 124.3 mi [200 km] from Mwauwe stream (–CB), 19 November 1982, Zambatis 1510 (PRE). 2531 (Komatipoort): Mbombela, Crocodile Valley Estate near Waterfall (–CA), 12 November 1973, Onderstall 129 (PRE); Barberton, Highland Creek (–CC), 29 January 1890, Galpin 786 (PRE).
MPUMALANGA: 2430 (Pilgrim’s Rest): Olifants Camp, Kruger National Park (–BD), 31 December 1984, Schijf 1220 (J); Mariepskop, rain forest near sawmill (–DB), 1 April 1959, Van der Schijff 4650 (PRE); Blyderivierspoort Nature Reserve, Swadini (–DB), 8 October 1987, Raal 1693000 (J); White River, Kruger National Park, Skukuza (–DC), 29 December 1984, Schijf 1221 (J). 2531 (Komatipoort): Kruger National Park, boskloof in Shabin (–AA), 9 October 1954, Van der Schijff 3908 (PRE); Bushbuck Ridge Nature Reserve (–CC), 25 September 2011, Balkwill and McCallum 12570 (J); Songimvelo Game Reserve, Dundar Valley (–CC), 26 September 1999, Balkwill, McCallum, and Campbell-Young 11059 (J); ca 3 mi [4.8 km] from Shia-lo-Ngubu Dam towards Louws Creek (–CD), 10 November 1973, Nel 349 (PRE).
KWAZULU-NATAL: 2632 (Bela vista): Charter’s Creek, Lake St Lucia (–AD), 1 January 1967, Hitchins 245 (NH, NU); Ndumo Game Reserve, Usutu riverine woodland (–CD), 1 November 1969, Moll 4236 (NH); Sihandla riverbank (–DD), 26 November 1967, Moll and Strey 3912 (NH); Kosi Bay, NW of Lake Nhlange (–DD), 26 October 1989, Ward 2622 (NH); Kosi Bay, NW side of Lake Nhlange (–DD), 19 July 1986, Venter 11540 (NH). 2730 (Vryheid): Utrecht, Donkerhoek (–AD), 17 September 1963, Devenish 1039 (PRE); Pongola Bush Nature Reserve, above Uitvlugt farm (–BC), 9 October 1990, Glen 2358 (PRE). 2731 (Louwsburg): Ingwavuma, summit of mountains near Magistrate Court (–AA), 23 September 1943, West 2109 (NH); Ingwavuma, Ndumu Game Reserve (–AA), 14 January 1964, Tinley 853 (NH, NU); uPhongolo Municipality, ± 300 m S of Mthembu Store, eMboloba (–AC), 21 February 2007, Keswa 1757 (NH); Ngotshe, Pongola town (–BC), 25 October 2001, Meyer BP00979 (PRE); Mpondwane, eastern slopes of Lebombo Mountains (–BD), 18 November 2001, Vahrmeijer HV00273 (PRE); Craigadam farm, Itala Nature Reserve (–CB), 19 January 1978, McDonald 487 (PRE, NU); Ndumu Game Reserve, Usutu River Forest (–DC), 10 September 1970, Pooley 841 (NH, NU). 2732 (Ubombo): Mandozi, ca 10.8 mi [17.4 km] from Manzengwenya on road to Mbazwana (–BC), 26 October 1966, De Winter and Vahrmeijer 8539 (PRE); Tembe Elephant Park (–AC), Ward 1070 (NH); Lake Sibaya area (–BC), 10 April 1958, Tinley 210 (PRE, NU); Lake Sibaya area (–BC), 11 January 1964, Strey 5128 (NH, NU); Manzengwenia (–BC), 10 December 1964, Vahrmeijer and Tȍlken 248 (NH); Sodwana Bay, edge of swamp forest (–DA), 17 September 1965, Vahrmeijer 1153 (PRE); Sodwana Nature Reserve (–DA), 16 October 2001, Schuhardt DS00216 (PRE). 2830 (Dundee): Dundee, hill above reservoir (–AA), 26 November 1955, Edwards 1078 (PRE, NU); Oudeni, Gonzaga Mission (–DB), 8 September 1939, Gerstner 3550 (PRE); Krantzkop, a few miles from Jamesons Drift (–DD), 22 March 1956, Edwards 1285 (PRE, NU). 2831 (Nkandla): Hlabisa, Charter’s Creek (–BB), 14 October 1963, Barker 10018 (NH); Nkandla Forest, ca 14.7 mi [23.7 km] from Nkandla hotel on road to Eshowe (–CA), 10 April 1964, De Winter 8266 (PRE); Nkandla Forest (–CA), 18 August 1987, Cunningham 1987 (NU); Silverglen Nature Reserve (–CC), 7 March 1994, Anna Jäger 18 (NU); Silverglen Nature Reserve (–CC), May 2010, Amoo 23 (NU); ca 4 mi [6.4 km] N of Nkwalini (–DA), 9 October 1946, Acocks 12958 (PRE); Ngoye Forest (–DC), 11 October 1984, Jordaan 396 (NH); Mtunzini (–DD), 17 February 1961, Wells and Edwards 32 (NH). 2832 (Mtubatuba): Hluhluwe Game Reserve (–AA), 27 June 1967, Wearne 86 (NH, NU); Hluhluwe Game Reserve (–AA), 13 November 1953, Ward 1697 (NH, NU); St Lucia, western shores, Nyalazi Forest Station (–AB), 11 March 1982, Nicholas and Van Wyk 1254 (NH); Nyalazi State Forest, Kentron (–AB), 8 October 1986, MacDevette 1136 (NH, NU); Hlabisa, St Lucia Estuary, edge of Kubululu Lake (–AD), 28 November 1965, Moll 2808 (PRE, NU); Maphelane Nature Reserve road, S of NPB office, dune forest (–AD), 23 October 1985, Fokkens 19 (NH); Nseleni Village, Nseleni Nature Reserve, NE of reserve and staff quarters complex (–CA), 8 November 2000, Ngwenya 1835 (PRE); Mbonambi Village (–CA), 13 December 2007, Nkuna and Mabatha 2339 (PRE); Richards Bay, Kleiklipklofie, 500 m from beach (–CC), 17 October 2001, Meyer 3971 (PRE). 2930 (Pietermaritzburg): Lions River, Karkloof, Ehlatini farm (–AC), 10 February 1967, Moll 3435 (NH, NU); Silverglen Nature Reserve, E of Umlazi Nursery (–CD), 28 August 1985, Buthelezi 589 (NH); Camperdown, Umgeni Dam (–DA), 20 May 1965, Moll 1724 (PRE, NU); Umlaas Valley (–DA), 13 Nov, 1933, Forbes 1181 (NH); near Chakaskraal at the edge of the bush on top of a rocky bank (–DB), September 1916, Thode s.n. (NH); Shongweni Dam (–DC), 21 February 41966, Morris 710 (NH); Umlaas Valley (–DC), 13 November 1933, Smuts and Gillet s.n. (NH); Silverglen Nature Reserve, next to resource centre (–DD), 12 March 1996, Crouch 742 (NH); Westville North (–DD), 11 September 1966, Coleman 332 (NH); Kranzkloof Nature Reserve (–DD), 2 3 July 2001, Grace 5 (NU). 2931 (Stanger): ca 25 mi [40.2 km] from Tongaat (–AC), 8 February 1965, Moll 1525 (NH, NU); Stanger, Ballito, Rainfarm Game Reserve (–AC), 15 December 2015, Cheek 2426 (NH); Lower Tugela, New Guelderland (–AD), June 1966, Stewart 33 (PRE); New Guelderland, 5 miles from the sea (–AD), March 1966, Stewart 6 (NH); Amatigulu Nature Reserve, Trail 2 (–BA), 4 December 2015, Steyn 2184 (PRE); Inanda, Saunders Park south, Tongaat River (–CA), Ward 5848 (PRE, NU); Port Zimbali, dune forest (–CA), 5 September 1985, MacDevette 322 (NH, NU); Musgrave road, Branksome Towers (–CC), 1 December 1983, Adamson s.n. (NH). 3030 (Port Shepstone): Mapumulo (–AA), 15 May 1987, Cunningham 2695 (NU); KwaDumisa, ± 600 m NE of eMphambanyoni Sowing Centre, eMphambanyoni (–AB), 20 July 2006, Keswa 857 (NH); Dumisa, 400 m NE of Vulamehlo Community Poultry Project, Mphambanyoni (–AB), 28 July 2006, Keswa 921 (NH); Gilbraltar, outcrop of Table Mountain Sandstone, at top of Umzimkulu Gorge (–AC), 13 March 1973, White 10527 (NH); Hlutankungu, Dumisa (–AD), 25 September 1912, Rudatis 1681 (NH); South coast, Mkomazi River, Delos Estate 10342 (–BA), 6 September 1994, Ward 12666 (NH, NU); Friedennau, Alexandra City (–BC), 10 September 1910, Rudatis 862 (NH); Ngome, edge of golf course (–BD), 21 December 1960, Mauve 4106 (PRE); Scottburgh, in die bosse van die kusgordel (–BD), 26 May 1941, Goossens 1708 (PRE); Ntimbankulu/Dweshula, near foot path to Mabhaleni Village, Ntimbankulu Forest (–CB), 15 February 2001, Ngwenya 2064 (NH); Umzimkulu Gorge, Gibraltar (–CB), 13 March 1973, White 10527 (PRE); Umtamvuna Nature Reserve, below Aerodrome (–CC), 15 October 1983, Abbott 1421 (NH); South coast, Palm Beach (–CD), 5 January 1981, Schrire 566 (NH); Alfred, Uvongo Hills Forests (–CD), 17 October 1973, Mogg 38276 (PRE). 3130 (Port Edward): Mzamba Casino, floodplain of Mzamba River, on the road to the pump house (–AA), 8 January 1993, Abbott and Arkell 19 (NH).
WESTERN CAPE: 3322 (Oudtshoorn): Wilderness, The Lakes, road to Rondevlei (–DC), 26 January 2001, Meyer BP00442 (PRE). 3323 (Willowmore): near Keurbooms River (–AB), 27 February 1984, Linger 83 (PRE); Keurbooms River (–DD), 11 November 1894, Schlechter 5942 (PRE); Keurbooms Strand (–DD), November 1961, Taylor 2913 (PRE).
EASTERN CAPE: 3029 (Bizana): ca 5 mi [8 km] from Bizana on Kokstad road (–BD), 19 December 1944, Acocks 10943 (PRE). 3129 (Port St. Johns): Transkei, Umsikaba River, Ndindini (–BD), 10 November 1970, Strey 10123 (PRE); 2nd Beach (–DA), 12 May 2002, Schuhardt DS02955 (PRE); Intafufu River (–DA), 11 October 1968, Mill 425 (PRE). 3323 (Humansdorp): Tsitsikamma National Park, kampeerterrein (–DC), 7 January 1960, Brand 165 (PRE). 3325 (Uitenhage): Winterhoek Mountain, Essenwood (–CA), 12 September 1930, Fries, Norlindh, and Weimarck 869 (PRE); Van Stadensrivier Reserve (–CA), 22 April 2002, Potgieter FP00739 (PRE); Baakens River, just below Mangold Park (–DC), 6 August 1974, Olivier 1101 (PRE). 3326 (Grahamstown): Kenton-on-Sea, Bloukrans (–DA), 15 March 2002, Potgieter FP00582 (PRE); Port Alfred, Kenton-on-Sea, top of dune (–DA), 23 May 1991, Steel 770 (PRE); 3327 (Peddie): Kaffraria, East London West; wooded kloof beyond lighthouse (–BB), 22 December 1900, Galpin 5842 (PRE); East London coast (–BB), April 1897, Sim 2152 (PRE); Kleinemonde, Brink’s plot (–BB), 8 January 1984, Eales 10 (PRE).
Eswatini. 2631 (Mbabane): Manzini, Timbutini (–AD), 22 November 1963, Compton 31785 (PRE), Hlatikulu, hills to N of Grand Valley Road, at the foot of hill up to Hlatikulu (–CD), 30 July 1981, Prior 485 (PRE).
Ekebergia pterophylla
Ekebergia pterophylla (C. DC.) Hofmeyr in Journ. Bot. 63: 57 (1925); White in Bothalia 16 (2): 157 (1986); White and Styles in Fl. SA. 18 (3): 53 (1986). Type: South Africa, near ‘Gwenberg’, Medley (‘Imley’) Wood 1022 (G, holotype; K, isotype).
Trichilia alata N.E.Br., Bull. Misc. Inform. Kew 1896 (117–118): 160 (1896). White in Bothalia 16 (2): 157 (1986). Type: KwaZulu-Natal, Groenberg (not located), Wood 1043 (K, lectotype; NH! (2x), isolectotypes). [Lectotype designated by White in Bothalia 16 (2): 157 (1986)].
Trichilia pterophylla C. DC. in Bull. Herb. Boissier 2: 581 (1894); White in Bothalia 16 (2): 157 (1986). Type as above.
Small, sturdy, deciduous shrub or tree, up to 10 m tall; crown flat; bole straight in a sheltered position; young branches stout and thick. Bark surface smooth but rough and mottled in older trees; inner bark brownish (sometimes with white bands); twigs with reduced or no lenticels; pith with prismatic crystals and druses; cortex with ideoblast cells; dilatation of cortical tissue is by anticlinal division of the parenchyma and cortical collenchyma cells as well as tangential stretching of cells forming strands of 2–11 cells; phelloderm cells with crystals; axial parenchyma with crystals; fibers absent in secondary phloem ray cells without crystals; canal in rays.
Wood: diffuse-porous, vessels slightly angular or rounded in outline, solitary, few, and in radial multiples of 2–4, intervessel pits alternate, minute, 2.7–3.8 μm in diameter; fibers non-septate; axial parenchyma scanty paratracheal, up to 5 cells wide; prismatic crystals present in chambered axial parenchyma cells; rays mostly uniseriate.
Leaves: imparipinnate, leathery compound with 2–3 pairs of leaflets, petiolate, lateral leaflets opposite, elliptic to narrowly obovate, leaflets 30–50 × 20–30 mm, leaflet base acute, leaflet apex narrowly tapered to rounded and may be notched; petioles broadly winged, petiole 20–30 mm long,
petiolules: absent; rachis broadly winged; stipules absent; margin entire, extends down to rachis and rolled under.
Flowers: unisexual, in panicles, small, ca. 30 mm in diameter, white or slightly pink, sweetly scented, star-shaped in compact clusters; sepals 4–5, ca. 2–4 mm long; petals 4–5, imbricate; stamens 10, united or joined connate, anthers sessile; ovary superior, 4–5-celled, sessile, with two ovules in each locule, surrounded by an annular disc; style short and thick, up to 7 mm long, stigma obscurely 4–5-lobed, male flowers with undeveloped ovary, female flowers with non-dehiscing anthers.
Fruit: a drupe, fleshy, up to 10 mm in diameter, globose, ripens from green to dull yellow, red, or black, mostly 1-seeded, indehiscent.
Seeds: one or two, without arils. (
Figure 3).
Distribution and ecology
The species grows in montane and coastal forests, amongst rocks on outcrops or in ravines, open woodland, forest margins, and sometimes close to the escarpment below an elevation of 2000 m. It is distributed in Limpopo, Mpumalanga, KwaZulu-Natal, and Eastern Cape Provinces in South Africa. It is also found in Eswatini (
Figure 4).
Diagnostic characters
Ekebergia pterophylla is similar to E. capensis in that the leaves are compound with opposite leaflets ending with a single terminal leaflet. E. pterophylla is readily distinguished by its broadly winged rachis and petiole, lack of petiolules, leathery, elliptic to narrowly obovate leaflets, 30–50 × 20–30 mm vs. non-winged petiole, presence of petiolules, and glossy, ovate to lanceolate leaflets, 20–120 × 20–50 mm in E. capensis.
Conservation status
Phenology
The species flowers during the late winter to spring months (July to November) and fruits between December and May.
Specimens examined
South Africa. LIMPOPO: 2229 (Waterpoort): Soutpansberg, Makhado, plaas Wellington, noordekant van plaas (–DC) 25 March 1994, Rossouw 215 (PRE); Soutpansberg, Masekwaspoort road (–DD), 15 May 1994, Balkwill 9113 (J); N of the Soutpansberg, on Fife farm 790 MS, on road to “Budworth”, slopes below south-facing cliffs (–DD), 14 May 1994, Cron 293 (J). 2230 (Messina): Makhado, ca 9.3 mi [15 km] NE of Makhado on the Witvlag road, Studholme farm (–CC), 4 April 1997, Burgoyne 6177 (PRE); Thononda, naby Kruin (–CD), 5 October 1981 Van Wyk and Theron 4630 (PRE). 2329 (Polokwane): Blaauwberg, near Malebochs kraal on hill side (–AA), 27 April 1954, Codd 8705 (PRE); Llewellyn, 35 LS farm, behind homestead (–AB), 20 July 1981 Venter 6194 (PRE); Happy Rest, ca 13 mi [20.9 km] W of Makhado (–BA), 28 February 1946, Gerstner 6027 (PRE); Lajuma Research Centre (–BA), 28 October 2015, Balkwill, Reddy and Oosthuizen 13673 (J); ca 8 mi [12.9 km] N of Makhado, near Mount Summit (–BB), 27 August 1961, Van Vuuren 1250 (PRE); The Downs (–CD), December 1918, Rogers 22010 (PRE). 2430 (Pilgrim’s Rest): Lekgalameetse Nature Reserve, The Downs, The Knuckles (–AA), 17 April 1986, Stalmans 1267 (PRE).
MPUMALANGA: 2430 (Pilgrim’s Rest): Blyde River, Drie Rondawels (–DA), 9 October 1971, Jordaan 114 (PRE); Mariepskop, on Blyderivier road (–DB), 28 May 1960, Van der Schijff 5120 (PRE); Ohrigstad Dam Nature Reserve (–DC), 20 January 1976, Theron 3432 (PRE); Kowyns Pass, Graskop (–DD), 10 September 1963, Rauh and Schlieben 9723 (PRE); Fairy Land Private Nature Reserve (–DD), 14 April 1979, Dyer s.n (J). 2530 (Mashishing): Buffelskloof Nature Reserve, top of gorge, open woodland (–AD), 12 December 1996, Crouch 747 (PRE); Montrose Falls (–BC), 2 April 1944, Mogg 30429 (PRE); Starvation Creek Nature Reserve, teen rand van eskarpement (–DA), 27 October 1977, Kluge 1096 (PRE); Buffelskloof Nature Reserve, krantz edge (–DA), 12 June 2002, Crouch 945 (NH); Kaapsehoop, Berlin (–DB), 25 April 1923, Hofmeyr 8 (PRE); Witklip, Klipkop, 19 September 1973, Kluge 221 (PRE); Barberton, valley above Agnes Mine (–DD), 1 November 1986, Cadman 3627 (J). 2531 (Komatipoort): Sabie Hoek Forest (–AA), 27 January 1906, Burtt Davy 5217 (PRE); Krokodilpoort, Beacon Hill (–AC), 11 May 1972, Nel 210 (PRE); Nelspruit, on the farm Moederlief 209 JU (–CA), 30 April 2014, Buthelezi 9 (J); Havelock Mine (–CC), October 1955, Miller 3079 (PRE); Barberton, on the farm Dycedale 368 JU (–CC), 23 November 2013, Balkwill, Reddy, McCallum and Cron 12826 (J); Barberton Mountainlands Reserve, Wonder Scheur 362 JU (–CC), 10 November 2014, Balkwill, Mbatha, Oosthuizen, McCallum and Adebowale 13146 (J); SW of Shiyalongubo Dam wall (–CD), 1 February 2004, McMurtry 11660 (PRE).
KWAZULU-NATAL: 2730 (Vryheid): Utrecht (–AD), July 1922, Thode A219 (PRE); Grootspruit, near Grootspruit on hill (–BC), 3 February 1968, Strey 8048 (PRE). 2731 (Louwsburg): On quartzite ridge (–CA), 6 November 1964, MacDevette 760 (NH); near Louwsburg, Ngotshe, edge of krantz (–CB), 12 October 1946, Acocks 13033 PRE. 2830 (Dundee): ca 19 mi [30.6 km] from Nqutu on Qudeni road (–BA), 20 November 1957, Edwards 2204 (PRE, NU). 2831 (Nkandla): Babanango, Scheepershoek NHS, Mahlanganeni River valley (–AC), 11 June 1993, Scott-Shaw 5668 (NU); Mtunzini, Ngoye Forest Reserve, eastern section (–DC), 5 October 1992, Williams 927 (NH); Ngoye Forest Reserve (–DC), 21 November 1990, Ngwenya 3327 (NH); Ngoye Forest (–DC), September 1969, Garland 327 (NH); Ngoye Forest (–DC), 12 October 1984, Jordaan 420 (NH); Mtunzini, Ngoye Forest Reserve (–DC), 23 February 1961, Well and Edward 134 (NU); Ngoye Forest Reserve, rocky outcrop on top of spur (–DD), 21 November 1990, Hutchings 3327 (NH); Mtunzini, Ubisana Valley (–DD), 17 August 1964, Venter 1694 (NH). 2832 (Mtubatuba): Hluhluwe Game Reserve (–AA), 28 November 1959, Ward 3341 (NH, NU). 2929 (Underberg): Hlatikulu (–BA), 16 October 1959, Compton 29240 (NH). 2930 (Pietermaritzburg): KwaSangwana, edge of cliff (–BC), Ngwenya 598 (NH); Little Noodsberg (–BD), October 1989, Scott-Shaw 2127 (NU); Noodsberg, Ozwatini plateau, forest margin (–BD), 13 October 1989, Williams 515 (NH); Mapumulo, on the Ozwatini Plateau (–BD), 13 October 1989, Balkwill 4940 (J); Cato Ridge, Mkabela, near Ekuthuleni, forest margin (–DA), 10 October 1989, Williams 377 (NH); Craig Lea farm, Table Mountain Sandstone, at top of cliffs of small uninhabited tributary valley of Ngeni Valley, open bushland among rocks (–DA), 24 February 1973, White 10308 (NH); New Hanover, near Ekuthuleni, Windy Hill (–DA), 10 October 1989, Balkwill 4706 (J); Greater Durban area, Inanda Mission Reserve 4579, Armstrong Hill, Showe Forest (–DB), 16 July 1991, Ward 12083 (NH); Hammarsdale (–DC), 4 July 1928, Forbes 299 (NH); Ismont Krantz, TMS (–DC), 26 January 1969, Strey 8359 (NH); Winston Park, Ntendeni Hill (–DD), 29 August 1971, Ward 7187 (NU). 3030 (Port Shepstone): Alexandria, Dumisa Station, Umgaye (–AD), 21 July 1911, Rudatis 1431 (PRE); Oribi Gorge Hotel, forest outskirts near waterfall (–CA), 13 August 1953, Story 4147 (PRE); Oribi Gorge Natuurreservaat, op rand van afgrond naby bopunt van viewpoint wandelpad (–CA), 6 December 1975, Van Wyk and Venter 1294 (NU); Umtamvuna Tributary (–CB), 3 February 1966, Nicholson 329 (NH); Izotsha River Gorge, TMS outcrop at top of cliffs (–CB), 13 March 1973, White 10523 (NH); Beacon Hill (–CB), 1 January 1967, Strey 7231 (NH); Umtamvuna Forest Reserve (–CC), 11 October 1969, Strey 9152 (NH, NU); Beacon Hill, ca 15.5 mi [25 km] from Port Edward (–CC), 14 July 1976, Venter 1017 (PRE); Paddock, Ngotshe Falls (–CC), 21 June 1963, Nicholson 42 (NH); ca 5 mi [8 km] NW of Uvongo, kloof forest on TMS (–CD), 2 January 1970, Moll 4934 (NH); Izotsha, TMS krantzes (–CD), 13 March 1973, Strey 11118 (NH, NU); Mgongongo, kloof forest, top of cliffs (–CD), 19 January 1969, Strey 8335 (NH, NU). 3130 (Port Edward): Beacon Hill, cliff bush (–AA), 8 February 1972, Strey 10627 (PRE, NU); Umtamvuna Nature Reserve (–AA), 25 January 1985, Abbott 2501 (NH); Umtamvuna Valley, Umtamvuna Nature Reserve (–AA), 14 April 1988, Harte 216 (J).
Eswatini. 2631 (Mbabane): Dalriach, amongst rocks (–AC), 18 October 1958, Dlamini 46043 (PRE); Sibebe Hill, ca 4.3 mi [7 km] NE of Mbabane (–CB), 12 September 1979, Prior 306 (PRE); near Hlatikulu (–CD), 16 October 1959, Compton 29240 (PRE); Abens farm (–CD), 23 October 1956, Compton 26109 (NH).
Genus Nymania
Nymania Lindb. in Not. Sällsk. Fauna Fl. Fenn. Förh. 9: 290 (1868); Marloth in F.S.A. 2 (1): 112 (1925); Verdoorn in the flowering plants of Africa 37: 1454 (1965); Dyer, Gen. 1: 299 (1975); Pennington and Styles in Blumea 22: 460 (1975); White in Bothalia 16 (2): 146 (1986); White and Styles, F.S.A. 18 (3): 39 (1986); Archer in Strelitzia 10: 354 (2000); Archer in Strelitzia 14: 630 (2003). Type species: Nymania capensis (Thunb.) Lindb.
Aitonia Thunb. in Physiogr. Sälsk. Handl. 1 (3): 166 (1776); Thunb. Nova genera Plantarum 1:52 (1782); Thunb. in Fl. Cap. 1: 508 (1823); Sond in Fl. Cap. 1: 243 (1860); White in Bothalia 16 (2): 146 (1986). Type species: Aitonia capensis Thunb.
Shrub or small tree. Leaves: simple, entire, spirally arranged in fascicles on short lateral shoots. Flowers: bisexual, solitary, axillary; calyx 4-lobed near the base; petals four, free, imbricate; stamens eight, filaments distinctly curved, fused near the base, without appendages, anthers glabrous, fixed dorsally towards the base, versatile; ovary 4-lobed, with four locules, loculi with two collateral ovules; disc thin, partially joined to the base of the staminal tube; style long, slender, with simple, minute, scarcely expanded style head. Fruit: a papery, inflated, deeply lobed capsule, laterally compressed with wing-like locules. Seeds: reniform; testa thick, woody, minutely verrucose, pubescent; cotyledon flat, slender, collateral; radicle superior, short.
The species
Nymania capensis (Thunb.) Lindb. in Not. Sällsk. Fauna Fl. Fenn. Förh. 9: 290 (1868); White in Bothalia 16 (2): 146 (1986); White and Styles, F.S.A. 18 (3): 40 (1986). Type: South Africa, without specific locality, no date, Thunberg s.n. (UPS-image!, holotype).
Aitonia capensis Thunb. in Physiogr. Sälsk. Handl. 1:166 (1776). Type: As for Nymania capensis.
Woody shrub or slender (single-stemmed) tree, up to 4 m tall; bole straight; branches rigid.
Bark: surface smooth but rough with regular flakes on older trees; inner bark whitish; twigs with lenticels; pith with prismatic crystals and druses; cortex without ideoblast cells, dilatation of cortical tissue is by anticlinal division of the parenchyma and cortical collenchyma cells as well as tangential stretching of cells forming strands of 2–7 cells; phelloderm cells with crystals; axial parenchyma with crystals; fibers absent in secondary phloem; ray cells with crystals.
Wood: diffuse-porous, vessels slightly angular or rounded in outline, numerous, solitary, in radial to diagonal multiples of 4–7, very narrow, tangential to dendritic pattern, intervessel pits alternate, minute, 2.7–5.1 μm in diameter; fibers non-septate; axial parenchyma scanty paratracheal, diffuse, in solitary strands near the vessels, in strands of 2–6 cells wide; rays mostly 1–2-seriate.
Leaves: simple, small, alternate, entire, stiff, leathery, growing in tufts along the branches, oblanceolate, 5–20 × 1–4 mm, dull green, slightly pubescent, sessile, leaf base acuminate, leaf apex acuminate.
Flowers: solitary, bisexual, borne on pedicels in leaf axils, in panicles, small, ca. 25 mm in diameter, white to red to pink, papery; sepals 4-lobed near the base, ca. 1–3 mm long; petals 4, imbricate; stamens 8, inserted at the base of the disc, filaments curved, fused at the base, anthers medifixed; ovary superior, sessile, 4-lobed, four locules with each containing two collateral ovules; style longer than stamen, style head simple, minute, scarcely expanded, ca. 8 mm long, stigma simple.
Fruit: an inflated capsule, large, conspicuous, loculicidal, papery, up to 40 mm in diameter, deeply lobed with four horizontally flattened wing-like expansions, ripens from carmine to silvery gray.
Seeds: borne in a papery, inflated capsule are distinctive, pea-shaped, and dark brown (
Figure 7).
Distribution and ecology
The species grows in hot, dry, rocky habitats near dry, sandy rivers. It is endemic to South Africa and Namibia. It is mostly found in the Northern Cape, Western Cape, and Eastern Cape provinces in South Africa. It occurs in southern Namibia, Namaqualand, the Richtersveld, Bushmanland, Ceres Tanqua Karoo, Worcester Robertson Karoo, and the Little Karoo in South Africa (
Figure 8).
Conservation status
Phenology
The species flowers in the winter months (June to August) and fruits between September and November.
Diagnostic characters
Nymania capensis is the only species in the genus Nymania, and it is readily identified by the small, simple leaves and the shape and color of its bladdery fruit. The wood is distinct in having numerous and narrow 24–44 μm (in tangential diameter) vessels (wood cross section). The species is referred to as “klapper” or “klapperbos” because the capsules are sometimes popped for fun.
Specimens examined
Namibia. 2316 (Nauchas): Ohangwena, Hardap, Rehoboth to Kobos (–BD), 22 May 1995, Van Slageren and Strohbach MSMS 046 (PRE); ca 9.3 mi [15 km] W of Rehoboth (–BD), 25 April 1976, Plowes 7162 (PRE). 2615 (Luderitz): Namusberge, ca 9.3 [15 km] from Rosh Pinah, Spitskop farm (–DD), 30 October 1983, Leuenberge, Raus, and Schiers 3263 (PRE). 2618 (Keetmanshoop): Itzawisis KEE 9 farm, Gully (–AA), 13 May 1963, Giess, Volk, and Bleissner 6875 (PRE); Great Karasberg, ca 30 mi [48.3 km] S of Narubis on road to Grunau flats and low rantjies at foot of mountain (–AD), 29 April 1955, De Winter 3319 (PRE); ca 3.4 mi [5.5 km] N of Narubis along Karasburg to Keetmanshoop highway (–DC), 3 February 1974, Davidse and Loxton 6309 (PRE). 2718 (Grunau): Klein Karas (–CA), 3 August 1923, Dinter 4822 (PRE). 2818 (Warmbad): bank of streamlet ca 20 mi [32.2 km] on Good House Road (–BD), 24 July 1937, Galpin 14142 (PRE); Keimas WAR 99 farm, below Glimmerschieferkuppe (–CC), 25 May 1972, Giess and Muller 1215 (PRE).
South Africa. NORTHERN CAPE: 2816 (Oranjemund): Namakwaland, Richtersveld National Park (–BB), 3 August 1998, Brand 30 (PRE); Richtersveld, Numees Mine, SE-facing gulley, rocky dry watercourse (–BD), 27 August 1987, Pienaar 963 (PRE). 2817 (Vioolsdrif): Richtersveld National Park, NE region (–AA), 8 July 1980, Van Wyk BSA123 (PRE); Richtersveld, ca 22 mi [35.4 km] S of Kuboos (–AC), 21 July 1948, Reynolds 5436 (PRE); Richtersveld, abandoned farm, ca 18 mi [29 km] from Eksteenfontein on road to Mount Stewart, rocky slopes (–CB), 29 August 1987, Germishuizen 4586 (PRE); ca 20 mi [32.2 km] beyond Vioolsdrif on way to Stinkfontein (–CD), 12 May 1969, Werger 400 (PRE). 2820 (Kakamas): On road from Upington to Namibian border post, at Naroegas (–AA), 17 December 1992, Jordaan 2588 (PRE); Gordonia, Riemvasmaak (–AD), 8 August 1989, Balkwill and Balkwill 4526 (J); Augrabies Falls National Park (–CA), 22 August 2005, Mothogoane 592 (PRE); Augrabies Falls National Park, along entrance road (–CB), 10 October 2011, Du Plessis 31 (PRE); Besiespoort farm, ca 15.7 mi [25.3 km] N of Kakamas-Keimoes road (–DA), 1 August 1984, Steiner 664 (PRE); Keimoes, ca 2.2 mi [3 5 km] from Keimoes on road to Kakamas (–DB), 22 June 1999, Meyer 2077 (PRE); Kenhardt, Letterkop Botanical Reserve, bergkamp (–DC), 12 July 1946, Wasserfall 1059 (PRE). 2821 (Upington): 22 mi [35.4 km] N of Upington (–AA), 25 April 1928, Pole-Evans 2123 (PRE); ca 21.7 mi [35 km] from Upington to Keetmanshoop (–AC), 14 July 1985, Joffe 38 (PRE); Namaqualand, Orange River, 13 mi [20.9 km] from Upington (–CA), 19 July 1967, Van der Schijff 6922 (PRE). 2822 (Glen Lyon): Langeberg range, Dunmurray (–BC), 22 March 1920, Pole-Evans 86 (PRE); Sunnyside Hill (–BD), 1 April 1989, Saaiman 380 (PRE); Hay, Langeberg Mountains, Excelsior 0.286 on rocky slopes and summit of Langeberge (–CD), 8 July 1936, Acocks 497 (PRE). 2823 (Griekwastad): Postmasburg, 11 miles NW of Witsands (–AC), 24 July 1958, Leistner 1169 (PRE); Griqua Town, naby Griekwastad (–CC), 26 April 1961, Mostert 1310 (PRE); Herbert, Campbell Tweefontein, below escarpment edge (–DC), 14 May 1955, Van Eck 17 (PRE). 2919 (Pofadder): Boomrivier farm (–AB), 7 June 2005, Mnengwane 18 (PRE, NU); Gemsbokvlakte (–BA), 16 July 1988, Dean 550 (PRE). 2922 (Prieska): Orange River Valley, ca 13 mi [20.9 km] E of Draghoender Station (–AC), 15 May 1946, Codd 1219 (PRE); Niekerkshoop farm, Maritz Dam (–BD), 1983, Gubb KMG12814 (PRE); ca 8 mi [12.9 km] NW of Prieska (–DA), 13 May 1961, Schlieben 8782 (PRE). 2923 (Douglas): Herbert, outside Douglas (–BB), September 1946, Brueckner 749 (PRE). 3023 (Britstown): ca 63 mi [101.4 km] W of Britstown (–CA), 19 October 1928, Pole-Evans 2308 (PRE).
WESTERN CAPE: 3320 (Montagu): ca 25 mi [40.2 km] NE of Montagu Dam (–CA), 21 November 1968, Van Breda 4132 (PRE); ca 24.9 mi [40 km] W of Ladismith near turn-off to Plathuis (–DB), 14 September 1988, Hilton-Taylor 2314A (PRE); Touwsberg, Zorgvliet farm (–DB), 5 October 1993, Bredenkamp 779 (PRE); between Barrydale and Ladismith (–DC), 1 May 1974, Goldblatt 1695 (PRE); Overberg, Barrydale (–DC), 27 November 1936, Phillips 2026 (J). 3321 (Ladismith): Sevenweekspoort (–AD), 31 October 1980, Van Wyk 358 (PRE); in Huisrivier Pass, between Ladismith and Calitzdorp (–BC), 25 November 2000, Van Wyk EVW0144 (PRE); Touwsberg, NW side of Touwsberg farm, Rietfontein 139 (–BC), 6 October 1993, Bredenkamp 851 (PRE). 3322 (Oudtshoorn): Cango Caves, hillsides and open flats (–AC), 13 January 1929, Mogg 11844 (PRE); Schoemanspoort, ca 0.6 mi [1 km] N of Schoemanshoek on slopes of gorge (–AD), 22 October 1989, Greuter 22149 (PRE); Turnoff to Zebra, Succulent Karoo, edge of dirt road (–AD), 28 September 2002, Mucina 280902/10 (PRE); De Rust, Aangenaam farm (–BC), 3 April 1973, Dahlstrand 2401 (PRE); Along George-Oudtshoorn Road (–CA), 27 July 1965, Kirkman 3008 (PRE); between Oudtshoorn and George, ca 5 mi [8 km] S of Oudtshoorn (–CA), 31 January 2001, Meyer BP00506 (PRE); Cango Caves (–CB), 29 June 1941, Phillips and Rensberg 2027 (J); Volmoed, ca 37.3 mi [60 km] beyond Hartenbosch on road to Calitzdorp, near Volmoed (–CC), 7 December 1976, Venter 1412 (PRE); Uniondale, Kamanassie Mountain, at Voorzorg Poort (–DA), 5 October 1989, Van Tonder 33 (PRE).
EASTERN CAPE: 3225 (Somerset East): Pearston, Buffelshoek Nek, foot of pass (–AC), 26 October 1945, Story 81 (PRE); Klein Visrivier Valle, Buffelsfontein (–CB), August 1967, Van der Walt 165 (PRE); De Toekomst (–DA), November 1930, Long 84 (PRE). 3323 (Willowmore): ca 40.4 mi [65 km] from Willowmore to Port Elizabeth, into Baviaanskloof Nature Reserve, Gruisbult farm (–DB), 8 November 1999, Nienaber EN731 (PRE). 3324 (Steytlerville): Noorspoort farm of George Craven, picnic area (–AD), 9 November 2004, Bredenkamp 3198 (PRE). 3325 (Port Elizabeth): Kommadagga (–BB), 7 October 1975, Bayliss BRI B6077 (PRE); Bottom of Suurberg Pass (–BC), 16 August 1973, Bayliss BRI B478 (PRE); Great Winterhoek mountains, Uitenhage, Coega catchment basin, Groendal Wilderness Reserve, Prentice Kraal farm (–CB), 15 September 1976, Scharf 1942 (PRE); Settlers Park Nature Reserve (–DC), 12 July 2002, Potgieter FP01324 (PRE).
Genus Trichilia
Trichilia Browne in Civ. Nat. Hist. Jamaica 278 (1756); Sonder in Fl. Cap. 1: 246 (1860); Staner and Gilbert, F.C.B. 7: 157 (1958); White and Styles in F.Z. 2: 297 (1963); De Wilde in Meded. Land. Sch. Wagen. 68 (2): 1–207 (1968); Pennington and Styles in Blumea 22: 467 (1975); White and Styles in F.S.A. 18 (3): 53 (1986); White in Bothalia 16 (2): 157 (1986); Leistner, Seed Pl. S. Afr.: 260 (2000); Coates Palgrave, Trees Southern Afr.: 453 (2002). Type species: Trichilia hirta L.
Deciduous trees, shrubs, or suffrutices. Leaves: imparipinnate, leaflets entire, alternate, rarely trifoliolate or unifoliolate. Flowers: unisexual, in cymose panicles. Calyx: short, 4–5-lobed or toothed. Petals: 4–5, much elongated than the calyx, imbricate, oblong or ovate; stamens 10, filaments connate in the lower half, free part heavily villous, with a pair of agile appendages at the apex; ovary small, occasionally adnate to disc, 2–4-celled, two collateral ovules in each cell; style short or long, continuous with the ovary; stigma capitate, 2–4-lobed; disc fused to base of staminal tube. Fruit: a loculicidal capsule, 2–4-celled, 3-valved septiferous, 1–2 seeds in each locule. Seeds: large, covered by red fleshy aril; cotyledons thick and short; radicle immersed.
Key to the species
1a Fruit with a distinct neck up to 10 mm long; leaf apex rounded; 13–16 pairs of narrowly spaced side veins on leaflets; flower densely pubescent; tree of lower-rainfall woodland and riverine thickets; ideoblast cells in the bark cortex—T. Emetica.
1b Fruit without a distinct neck; leaf apex acute to acuminate; 8–12 pairs of widely spaced side veins on leaflets; flower sparsely pubescent; tree of high-rainfall montane forest; ideoblast cells absent from bark cortex—T. dregeana.
Trichilia dregeana
Trichilia dregeana Sond. in Fl. Cap. 1: 246 (1860); White and Styles in F.Z. 2: 298 (1963); Ross, Fl. Natal 216 (1972); Styles in E. Afri. Agric. Journ. 39: 418 (1974); White in Bothalia 16: 158 (1986); White and Styles in F.S.A. 18 (3): 55 (1986); Styles and White in Fl. Eth. 3: 483 (1990). Type: South Africa, KwaZulu-Natal, Stanger (2931): Port Natal, Gueinzius s.n. (TCD-image!, lectotype; G; K; PRE!; S; SAM). [Note: lectotype designated by De Wilde, 1968].
Trichilia chirindensis Swynnerton and Bak. in Journ. Linn. Soc. Bot. 40: 39–41 (1911); Eyles in Trans. Roy. Soc. S. Afr. 5: 390 (1916); Brenan in T.T.C.L. 319 (1949). Type: Zimbabwe, Chirinda Forest, 22 September 1905, Swynnerton 1 (BM).
Trichilia dregeana Sond. var. oblonga Harv. ex Sond. in Fl. Cap. 1: 246 (1860) Type: South Africa, KwaZulu-Natal, Stanger (2931): Port Natal, no date, Sanderson s.n. (TCD-image!, holotype).
Trichilia dregei E. Mey., Zwei Pflanzengeogr. Docum. 1: 227, nomen (1843); Harms in Nat. Pflanzenf. 2nd ed. 19B1: 110. 1940. Type: South Africa, KwaZulu-Natal, Stanger (2931): Port Natal Durban, no date, Drège 39 (holotype in P, not seen).
Trichilia schliebenii Harms in Notizbl. Bot. Gart. Berl. 11: 1070 (1934). Type: Tanzania, Bezirk Morogor: Uluguru-Gebirge, Nordwestseite, Nebelwald, 11 March 1933, Schlieben 3636 (MA).
Trichilia splendida A. Chev. Bull. Soc. Bot. Fr. 58, Mémoires 8d: 147 (1911); Staner in Bull. Jard. Bot. Brux. 16: 159 (1941); Keay in F.W.T.A. 1: 705 (1958); Staner and Gilbert, F.C.B. 7: 165 (1958). Type: Guinea, Guinée Française: Cercle du Kissi, Kissidougou, February 1909, Chevalier 20708 (P).
Trichilia strigulosa Welw. ex C. DC. in Monogr. Phan. 1: 658 (1878); Exell and Mendonça in C.F.A. 2: 314–15 (1951). Type: Angola, Golungo Alto: Sporadica, Prope Undella, April 1856, Welwitsch 1311 (BM-image!, holotype).
Trichilia umbrosa Vermoes. in Rev. Zool. Bot. Afr. 10, Suppl. Bot. 53 (1922). Type: Congo, the Democratic Republic, 4 April 1919, Vermoesen 1948 (BR-image!, lectotype).
Trichilia vestita C. DC. in Bull. Herb. Boiss. 4: 428 (1896). Type: Angola, Golungo Alto. Prope, October 1855, Welwitsch 1309/10 (BM).
Large, evergreen tree, up to 40 m tall; crown largely dense; bole straight (sometimes buttressed); up to 2 m in diameter.
Bark: surface grey to grey-brown, smooth but rough and mottled with pink in older trees; inner bark cream-coloured, rapidly turning pink to reddish brown; twigs with reduced lenticels; pith without prismatic crystals and druses; cortex without ideoblast cells; dilatation of cortical tissue is by anticlinal division of the parenchyma and cortical collenchyma cells as well as tangential stretching of cells forming strands of 2–11 cells; phelloderm cells with crystals; axial parenchyma with crystals; fibers present secondary phloem; ray cells with crystals, rays without canal.
Wood: diffuse-porous, vessel rounded in outline, few, solitary, in clusters of 2–4 or in radial multiplies of 2–4, 23–192 μm in diameter, intervessel pits alternate, minute, 2.2–3.9 μm in diameter; fibers non-septate; axial parenchyma scanty paratracheal, diffuse, confluent, in narrow bands of three cells wide, axial parenchyma cells chambered without crystals; rays 1–2-seriate.
Leaves: imparipinnately compound with 2–5 pairs of leaflets ending with a single terminal leaflet, petiolate, lateral leaflets opposite or alternate, obovate to oblanceolate, leaflets 180–200 × 60–90 mm, dark glossy green above, lighter below, distinctly broadest near apex, lateral veins 8–12 pairs, leaflet base rounded to obtuse, leaflet apex acute to acuminate and sometimes notched, margin entire and slightly rolled under; petioles thick at base, pubescent, 130–160 mm long, petiolules 10–20 mm long, pubescent; rachis thick at base, pubescent; stipules absent.
Inflorescence: a multi-flowered or few-flowered panicle, sometimes many-flowered in males but always few-flowered in female inflorescences.
Flowers: unisexual, creamy-white to velvety, sweetly scented; sepals 5, cup-shaped, ca. 3.5–7.5 mm long; petals 5, up to 24 mm long, imbricate, pubescent; stamens 10, united into a tube, single pistil in female flowers; ovary superior, 3-celled, pubescent, with 1–2 ovules in each locule, disc fused to base of terminal tube; style short, up to 6 mm long, stigma capitate, 2–4-lobed, male flowers with rudimentary ovary, female flowers with non-dehiscing anthers.
Fruit: a woody capsule, up to 32 mm in diameter, globose, on a short, stout stalk without a distinct neck attaching to the stalk, 3-lobed, up to 6-seeded.
Seeds: large, black, covered by a bright red aril (
Figure 11).
Distribution and ecology
The species is a fast-growing, frost-sensitive tree found in coastal and montane evergreen forests, mainly on mountain slopes. It grows up to 1220 m above sea level. It is found in the Limpopo, Mpumalanga, KwaZulu-Natal, and Eastern Cape provinces of South Africa (
Figure 12), but also occurs in Angola and the DRC, as well as Eswatini, Mozambique, Zimbabwe, Tanzania, Kenya, and Ethiopia. It grows in high-rainfall areas and is often found outside the natural habitat because of its ornamental uses.
Diagnostic characters
Trichilia dregeana is similar to T. emetica in that the leaves are imparipinnate, compound, and spirally arranged with opposite leaflets, ending with a single terminal leaflet. Trichilia dregeana is readily distinguished by the 8–12 pairs of side veins on its leaflets, the acute to acuminate leaflets, the down-curved leaf apex, and the fruit without a distinct neck attaching to the stalk. Trichilia emetica has 13–16 pairs of side veins on the leaflets, rounded leaflet apices, and a distinct neck joining the fruit capsule to the stalk.
Conservation status
Phenology
The species flowers during the spring to early summer months (October to December) and fruits between December and May.
Specimens examined
South Africa. LIMPOPO: 2230 (Messina): Sibasa, Pepiti Falls (–CD), 19 September 1934, Pole-Evans 3757 (PRE). 2330 (Tzaneen): Duiwelskloof, Westfalia Estate office (–CA), 25 October 1963, Scheepers 1234 (PRE); Westfalia Estate, near rain gauge (–CA), 29 May 1966, Scheepers 1251 (PRE). 2331 (Phalaborwa): PWD works area, ca 8.7 mi [14 km] S of Phalaborwa (–AA), 2 October 1984, Retief 267 (PRE).
MPUMALANGA: 2530 (Mashishing): Mbombela (–BD), 17 July 1906, Legat 2831 (PRE). 2531 (Komatipoort): White River, ca 2.6 mi [4.2 km] from Kaapmuiden turnoff, Boondocks Lodge (–CA), 15 October 2014, Balkwill, Mbatha, and Oosthuizen 13078 (J).
KWAZULU-NATAL: 2632 (Bela Vista): Maputa, Nkanini (–DD), 21 September 1965, Vahrmeijer 1222 (PRE). 2831 (Nkandla): St. Lucia Estuary Forest, S of township (–BB), February 1965, Tinley 1086 (NU); Lower Nkandla Forest (–CA), 18 June 1956, Edwards 1445 (PRE, NU); Machubeni Village, NW of Reserve office complex, Nkandla Forest Reserve (–CA), 15 May 2001, Ngwenya 2210 (PRE); Entabeni, Native Trust Ground (–DC), May 1943, Loock 9498 (PRE); Mthunzini, Oyameni village c. 200 m from Chief Ranger’s residence, Ngoye Forest Reserve (–DD), 19 September 2002, Ngwenya 2695 (NH). 2832 (Mtubatuba): Hluhluwe Game Reserve, Emabomvini (–AA), 8 August 1970, Bourquin 774 (PRE); St Lucia Estuary (–AB), 28 November 1965, Moll 2801 (NH, NU); Richards Bay, Harbour (South), approach Cut Dune (–CC), 13 August 1996, Ward 13789 (NH). 2930 (Pietermaritzburg): Pinetown, Hillcrest, Tunzini road (–BB), 13 July 1964, Ross 1271 (NH, NU); City Centre, Methodist church Longmarket Street, opposite 20th Century (–CB), 30 September 1981, Nicholas 1124 (NH, NU); Camperdown, Umgeni Dam (–DA), 20 May 1965, Moll 1731 (PRE, NU); Isipingo, near Umlaas River (–DD), 26 April 1949, Ward 856 (PRE, NU). 2931 (Stanger): Durban, Pigeon Valley, left side of entrance to Pigeon Valley (–CC), 12 November 1964, Thorns 5348 (PRE, NU); Durban Botanic Gardens (–CC), 27 September 1964, Strey 5340 (NH). 3030 (Port Shepstone): Ezinqoleni, ca. 40 m south of Oribi Gorge, hutted camp reservoir (–AC), 16 September 2006, Ngwenya 1028 (NH); Dumisa, Fairfield (–AC), December 1945, Bayer 1400 (PRE, NU); Umtamvuna bridge, riverbank bush (–BC), 26 December 1966, Strey 7122 (NH, NU); Thuthwini, Ntimbankulu/Dweshula, near foot path to Mabheleni village through Ntimbankulu Forest (–CB), 12 February 2001, Ngwenya 1980 (NH); Shelly Beach hinterland (–CD), 9 September 1967, Strey 7672 (NH, NU).
EASTERN-CAPE: 3129 (Port St. Johns): Near Mbotyi Forest (–BC), 22 May 1970, Jenkins SN4 (PRE); Lusikisiki, Lupatana Forest (–BD), 12 December 1986, Van Wyk and Mathews 7916 (NH); Goss Point, riverine forest (–BD), 10 November 1970, Strey 10151 (NH); ca 10 mi [16.1 km] from Port St. Johns on road to Lusikisiki (–DA), 20 May 1970, Jenkins SN3 (PRE); Ntafufu Forest (–DA), 10 May 1969, Strey 8511 (NH). 3227 (Stutterheim): near Komga (–DB), January 1890, Flanagan 459 (PRE). 3228 (Butterworth): Kentani Hill, Nomakeye (–AA), 12 July 1966, Strey 6611 (PRE).
Eswatini. 2631 (Mbabane): Manzini, Timbutini (–AD), 28 September 1961, Compton 30756 (PRE).
Trichiliaemetica
Trichilia emetica Vahl, Symb. Bot. 1: 31 (1790); Oliver in Fl. Trop. Afr. 1: 335 (1868); C.D.C. in A. and C.CD., Mon. Phan. 1: 660 (1878); Gȕrke in P.O.A. C: 231 (1895); Harms in Engl. Nat. Pflanzenf. 3 (4): 305 (1896); Bak. in J.L.S. 37: 133 (1905); Holland in The Useful Pl. of Nigeria, Kew Bull., add. series 9 (1): 146 (1908); Engler in Engl. and Drude, Veg. der Erde, 9, Die Pflanzenw. Afr. 3 (1): 821 (1915); Vermoesen in Rev. Zool. Afr. 10, Suppl. Bot.; 34 (1922); Harms in Nat. Pflanzenf. 2 (19B1): 109 (1940); Staner in Bull. Jard. Bot. Brux. 16 (2-3): 175 (1941); Andrews in Fl. Pl. Anglo-Egyp. Sudan 2 (331): 123 (1952); Eggeling and Dale, Indig. Trees Uganda Prot. 2: 195 (1952); Dalziel in Useful Pl. W. Trop. Afr. 328 (1955); White and Styles in F.Z. 2: 299 (1963); De Wilde in Meded. Land. Sch. Wagen. 68: 50 (1968); Styles in E. Afri. Agric. Journ. 39: 418 (1974); White in Bothalia 16 (2): 158 (1986); White and Styles in F.S.A. 18 (3): 57 (1986); Styles and White in Fl. Eth. 3: 483 (1990).
Trichilia emetica Vahl subsp. emetica, Persoon in Syn. Pl. 1: 468 (1805); Oliver in Trans. Linn. Soc. of London 29 (1): 44 (1872); Schweinfurth in Bull. Herb. Boiss. 7 (2): 295 (1899); Schinz and Junod in Mém. Herb. Boiss. 10: 44 (1900); Giubs in Bot. Journ. Linn. Soc. 37: 435 (1906); Monro in Proceed. Rhod. Sei. Ass. 8 (2): 67 (1908); Sim in For. Fl. and For. Res. Port. E. Afr. 26: 15 (1909); Legat in Kew Bull. 53 (1910); Engler in Engler and Drude, Veg. der Erde, 9, Die Pflanzenw. Afr. 3 (1): 821 (1915); Eyles in Trans. Roy. Soc. S. Afr. 5: 389 (1916); De Wildeman in Contr. Fl. Kat. 105 (1921); Vermoesen in Rev. Zool. Afr. 10 (1): B34 (1922); Gossweiler in Fl. Exot. Angol. in Agronomia Angolana 1: 121 (1950); Exell and Mendonça in C.F.A. 2: 314 (1951); Miller in Journ. S. Afr. Bot. 18:40 (1952); Pardy in Rhod. Agr. Journ. 51 (6): 492 (1954); Dale and Greenway, Kenya Trees and Shrubs 272 (1961); White and Styles in F.Z. 2 (1): 299 (1963). Type: Yemen, Hadie mountain, Forsskâl 478 (C-image!, holotype; isotype in BM).
Mafureira oleifera Bertol. in Mem. Acc. Sci. Bol. 2: 269 (1850). Type: Mozambique, Fornasini (P).
Trichilia roka Chiov. in Flora Somala 2: 131 (1932); Brenan in Mem. N.Y. Bot. Gard. 3: 235 (1953); Garcia in Contr. Conhec. Fl. Moçamb. 2: 142 (1954); Williamson in Useful Pl. Nyasal. 119 (1956); Keay in F.W.T.A. 1: 705 (1958); Staner and Gilbert, F.C.B. 7: 163 (1958). Type: Same T. emetica.
Trichilia umbrifera Swynn. and Bak. in Journ. Linn. Soc., Bot. 40: 39 (1911). Type: Mozambique, Lower Umswirizwi, 300 m, 1 November 1905, Swynnerton 148 (BM; K).
Medium to large, evergreen tree, up to 30 m tall; crown largely dense, wide spreading in the open; bole straight, sometimes fluted; young branches longitudinally striped.
Bark: surface smooth to slightly rough, dark grey to brown, unevenly fissured; inner bark whitish to brownish; twigs with conspicuous lenticels; pith without prismatic crystals and druses; cortex with round-shaped scattered ideoblast cells; dilatation of cortical tissue is by anticlinal division of the parenchyma and cortical collenchyma cells as well as tangential stretching of cells forming strands of 2–10 cells; phelloderm cells without crystals; axial parenchyma with crystals; fibers present in secondary phloem; ray cells with crystals; rays without canal.
Wood: diffuse-porous, vessel rounded in outline, few, wide, 23–192 μm in diameter, solitary, in clusters of 2–4 or in radial multiples of 2–4, intervessel pits alternate, minute, 2.2–3.9 μm in diameter, helical thickenings on vessel walls; fibers non-septate; axial parenchyma scanty paratracheal, diffuse, confluent, and in narrow bands of three cells wide, prismatic crystals absent in chambered axial parenchyma cells; rays1–2-seriate.
Leaves: imparipinnately compound with 2–6 pairs of leaflets ending with a single terminal leaflet, petiolate, lateral leaflets nearly opposite to opposite, elliptic to obovate, leaflets 120–150 × 50–60 mm, leaflet base rounded, leaflet apex rounded to pointed, lateral veins 13–16 pairs; margin entire, rolled under and sometimes undulating; petioles densely pubescent, 130–180 mm long; petiolules densely pubescent, 5–6 mm long; rachis densely pubescent; stipules absent.
Inflorescence: a multi-flowered, condensed panicle.
Flowers: unisexual, green to creamy green, sweetly scented, produced in tight clusters; sepals 5, cup-shaped, ca. 2–5 mm long, pubescent; petals 5, free, creamy green to yellow, ca. 10–16 mm long, imbricate, pubescent; stamens 10, filaments united at the base with anthers attached in the middle, pubescent; ovary superior, 3–4-celled, sessile, with two ovules in each locule, densely pubescent; style short, up to 8 mm long; stigma globose, 2–4-lobed, male flowers with simple ovary, female flowers with non-dehiscing anthers.
Fruit: a dehiscent capsule, up to 26 mm in diameter, globose, 3-lobed, differentiated from a 5–10 mm long neck, up to 6-seeded.
Seeds: large, black, nearly covered by a bright red aril (
Figure 13).
Distribution and ecology
The species grows in riverine forest and at medium-to-low-elevation woodland and forest fringes, as well as in coastal forests, from sea level to elevations between 1700 and 2000 m. It is found in the Limpopo, Mpumalanga, and KwaZulu-Natal provinces in South Africa (
Figure 14), from where the distribution extends to Namibia, Botswana, Eswatini, and Mozambique, and further north to West, Central, and East Africa, as well as Yemen.
Diagnostic characters
Trichilia emetica is similar to T. dregeana in that the leaves are imparipinnate, compound, and spirally arranged with opposite leaflets, ending with a single terminal leaflet. Trichilia emetica is readily distinguished by the presence of 13–16 pairs of side veins on its leaflets, rounded leaflet apices, the presence of a distinct neck joining the fruit capsule to the stalk, and the presence of ideoblast cells in the cortex of the bark (cross section). Trichilia dregeana has 8–12 pairs of side veins on the leaflets, acute to acuminate leaflet apices, fruit on a short, stout stalk without a distinct neck attaching to the stalk, and an absence of ideoblast cells in the cortex of the bark (cross section).
Conservation status
Phenology
The species flowers during the late winter to spring months (August to November) and fruits between December and April.
Specimens examined
Namibia. 1823 (Siambisso): Caprivi Strip, E of river, Cuando River (–BA), October 1945, Curson 1184 (PRE). 1725 (Livingstone): Eastern Caprivi, Mpilila Island, Chobe River (–CC), 15 January 1959, Killick and Leistner 3388 (PRE). 1724 (Katima Mulilo): Anthill on Zambezi flood plain (–AD), 27 August 1967, Von Breitenbach 1217 (PRE).
Botswana. 1824 (Kachikau): Northern Linyanti River, Shaile hunters camp airstrip (–AA), 20 January 1979, Smith 2623 (PRE); ca 6 mi [9.7 km] S of Kachikau and ca 30 mi [48.3 km] N of Goha (–AB), 8 July 1837, Erens 360 (PRE); Shaile Camp, edge of Linyati R. floodplain (–AB), 12 July 1977, Edwards and Ward 4532 (NU). 1724 (Katima Mulilo): Chobe River, ca 50 mi [80.5 km] N of Kachikau (–DD), 12 July 1937, Erens 4201 (PRE). 1725 (Livingstone): Bechuanaland Protectorate area, Kasane Rapids, on banks of the Chobe River (–CC), 26 July 1950, Robertson and Elffers 52 (PRE).
South Africa. LIMPOPO: 2229 (Waterpoort): Soutpansberg, Franzhoek farm (–DD), 20 July 1935 Galpin 14949 (PRE). 2230 (Messina): Sibasa, Tate Vondo Forest Reserve (–CD), 3 November 1976, Hemm 11 (PRE); Thengwe, near road at Thengwe location (–DA), 15 December 1979, Netshiungani 1144 (PRE). 2231 (Pafuri): Kruger National Park, Punda Milia, near Kamp (–CA), 2 February 1962, Ihlenfeldt 2297 (PRE). 2330 (Tzaneen): Klein Letaba River, ca 5 km E of Giyani (–BC), 3 May 1977, Liengme 101 (PRE); Klein Letaba River at Giyani (–BD), 14 May 1981, Liengme 577 (PRE); Letaba, Duiwelskloof, Westfalia, petrol pumps (–CA), 5 June 1958, Scheepers 358 (PRE); Letaba River, op pad na die eiland vanaf Tzaneen (–CD), 29 November 1974, Pienaar 503 (PRE); Great Letaba River, ca 50 mi [80.5 km] N of Gravelotte STA (–DA), 28 July 1934, Galpin 13866 (PRE). 2430 (Pilgrim’s Rest): Ngwabitsi River, gentle slope (–BD), 22 August 1985, Krynauw 736 (PRE); ca 12.4 mi [20 km] SW of Marulaneng, Blydeberg Nature Reserve (–BD), 10 October 1987, Balkwill and Cadman 3791 (J). 2431 (Acornhoek): Avoca, near Barberton (–AD), 17 September 1890, Galpin 1060 (PRE); ca 12 mi [19.3 km] from Newington to Bushbuck ridge (–CC), 19 March 1972, Buitendag 915 (PRE).
MPUMALANGA: 2430 (Pilgrim’s Rest): Picnic places near bridge in Erasmus Pass (–BC), 7 January 1063, De Winter 7806 (PRE); Swadini Blydepoort Dam, below dam wall, next to Blyde River (–DB), 10 September 2002, Male JM00240 (PRE). 2431 (Acornhoek): Timbavati Game Reserve, Kempiana farm (–AD), 10 March 1970, Porter 300 (PRE); Satara rest camp (–BD), 6 April 1990, Du Toit 231 (PRE); Bushbuckridge, Township Ga-Moreku (–CB), 21 January 2019, Shai 15 (PRE). 2530 (Mashishing): Lowveld Botanical Garden, Mbombela (–BD), 17 March 2002, Meyer 1 (PRE). 2529 (Witbank): Loskop Dam, Olifants River, near Groblersdal (–AD), 25 January 1990, Van Vreden 19 (PRE). 2531 (Komatipoort): Kruger National Park, rest camp of Pretoriuskop (–AB), 1 October 1950, Van Zinderen Bakker 313 (PRE); Nelspruit, on the farm Okkernootboom (–AC), 1 September 1988, Barker 23 (J); Plaston next to Mvangatini village, ca 1.2 mi [2 km] before the village (–AC), 12 January 2010, Lukhele UPP083 (PRE); Barberton, Sheba Siding (–CA), 11 November 1931, Galpin 3066 (PRE); between Louws Creek and Adamanda mine (–CB), 8 July 1906, Burtt Davy 2821 (PRE); Barberton (–CC), November 1931, Smith 7068 (PRE); Komati River (–DB), 20 July 1935, Nel 154 (PRE).
KWAZULU-NATAL: 2632 (Bela vista): Itala Nature Reserve, Ongegund section (–AD), 20 July 1982, Porter and Ward 138 (NH); Ndumo Game Reserve (–CD), 4 October 1988, Ward 2398 (NH). 2731 (Louwsburg): Pongola, KwaBhembe, ca 0.6 mi [1 km] from the W of Mlomokazulu high school (–AC), 9 December 2006, Magubane 1430 (NH); Flats E of Pongola Poort (–BD), 21 January 1964, Huntley 785 (NH, NU); Umfolozi, between Black Umfolosi and Nongoma (–DC), 16 August 1945, Acocks 11675 (PRE). 2732 (Ubombo): 10 mi [16 km] from Ingwavuma to Ndumo, riverbank (–AA), 20 November 1967, Moll and Strey 3704 (NH); Ingwavuma, Tembe Elephant Park (–AB), 4 September 1985, Ward 1026 (NH); I. D. C. Rice project site, ca 1.2 mi [2 km] from Phelandaba turn-off on Mbazwana road (–BA), 4 December 1985, Germishuizen 3620 (NH); Manzengwenya, ca 1.8 mi [3 km] S of De Wets Bay (–BB), 24 June 1986, MacDevette 864 (NH); Lake Sibayi eastern shore (–BC), 4 December 1969, Moll 4917 (NH); Pongola River near Tobotini (–BC), 10 April 1966, Strey 6600 (NH); Ndumo Game Reserve (–BC) 16 September 1971, Pooley 1449 (NH, NU); Mkuze Game Reserve near Mkuze River (–CA), 18 September 1962, Gush 26 (PRE, NU); Mkuzi Game Reserve (–CA), 26 September 1976, Goodman 772 (NU); between Mkuze and Hluhluwe on N2, farm iNtendele (–CC), 8 October 2001, Meyer P00958 (PRE); Sodwana State Forest, forest patch near chief’s kraal (–DA) 6 October 1986, MacDevette 1208 (NH, NU). 2830 (Dundee): Nquthu, B 1060 eMondlo, Engodini Village (–BA), 11 January 2002, Madi 55 (NH); ca 11 mi [17.7 km] from Kranskop on Middeldrift road (–DD), 20 June 1964, Edwards 3326 (PRE, NU). 2831 (Nkandla): Nkandla, Middeldrift (–CC), 9 June 1956, Edwards 1413 (PRE, NU); Hamewith near Mtunzini (–DC), 1 May 1919, Mogg 4447 (PRE); Mtunzini, Umlazi Nature Reserve (–DD), 15 September 1962, Ward 4311 (PRE, NU); Babanango (–DD), 19 April 1926, Watt and Breyer-Brandwijk 1171 (PRE). 2832 (Mtubatuba): Hluhluwe Game Reserve (–AA), 25 August 1955, Ward 2689 (NU); Mansiya/Nyimane, Hluhluwe Game Reserve (–AA), 8 October 1971, Hitchins 355 (NH, NU); Ncemane (–AB), 22 January 1949, Bayer 1463 (NU); ca 49.7 mi [80 km] from Mbazwana to Hluhluwe (–AB), 8 December 2001, Schuhardt DS00807 (PRE); St Lucia Forest Station, garden (–AC), 18 July 1965, Pegel 28 (NH); Richards Bay camping sites (–AC), 10 July 1965, Venter 1941 (NH); Hlabisa, Palm Ridge farm (–AC), 22 April 1968, Harrison 518 (NH); St. Lucia Hotel (–AD), 21 October 1964, Strey 5376 (NH); Dukuduku State Forest, St. Lucia (–AD), 12 October 1983, Nicholas 1635 (NH, NU); Inkuhlu (Ixolo), St. Lucia (–AD), 22 September 1986, Cunningham 2143 (NH, NU); Dukuduku State Forest, near Futululu (–AD), 3 September 1986, MacDevette 1044 (NH, NU); On Richards Bay road, near Ensileni River (–BC), 15 July 1951, Lawn 2085 (PRE); North coast, Lake Nhlabane area (–CB), 27 October 1991, Ward and Rajh 11419 (NH, NU). 2930 (Pietermaritzburg): Umvoti Valley, SW of Mapumulo (–BA), February 1965, Moll 529 (PRE). 2931 (Stanger): ca 16 mi [25.7 km] from Stanger on Noodsberg to Waterberg road (–AC), 18 June 1964, Edwards 3317 (PRE); Sinkwazi strand, south part of urban area (–AD), December 1973, Grobbelaar 806 (PRE); Amatikulu Nature Reserve, coastal woodland on riverbank (–BA), 20 September 1987, Ward 2134 (NH); Verulam (–CA), 14 September 1965, Moll 2020 (PRE, NU).
Eswatini. 2531 (Komatipoort): Piggs Peak, near Lomati River (–CD), 29 June 1959, Compton 28968 (PRE). 2631 (Mbabane): Manzini, Mpisi, rocky hill (–BC), 20 December 1961, Compton 31147 (PRE); Hlatikulu, Usutu Poort, riverside (–CA), 30 October 1961, Compton 30927 (PRE); ca 9 mi [14.5 km] N of Sitobela (–DC), 30 April 1957, Murdoch 159 (PRE). 2731 (Louwsburg): Nsoko, Canterbury farm (–BB), 2 July 1966, Bayliss 3474 (PRE).
Genus Turraea
Turraea L. in Mant. Pl. 2: 150 (1771); Exell and Mendoca in C.F.A. 2: 318 (1951); White and Styles in F.Z. 2: 307 (1963); Killick in Flower. Pl. Afr. 38: 1499 (1967); White and Styles in F.S.A. 18 (3): 41 (1986); Pennington and Styles in Blumea 22: 454 (1975); White in Bothalia 16 (2): 161 (1986); White in Flower. Pl. Afr. 50: 1962 (1988); Williams in Bothalia 19: 31 (1989); Leistner, Seed Pl. S. Afr.: 260 (2000). Type species: Turraea virens L.
Small to medium tree (usually suffruticose, sometimes deciduous). Leaves: simple (seldom compound), petiolate, or subsessile. Flowers: bisexual, in short axillary panicles, solitary or fasciculate; calyx 5-lobed or toothed, cup-shaped; petals 5, long, imbricate or contorted; stamens 10–20, filaments connate into a long, slender tube widening upwards, sometimes with notched lobes, anthers shortly apiculate; ovary sessile, occasionally ribbed, with (3)4–10(20) locules, with two collateral, oblique or superposed ovules in each locule; disc small, annular or absent; style exserted, filiform, stigma discoid, capitate, or globose. Fruit: a (3)4–10(20)-valved, woody or leathery, loculicidal capsule with 1–2 seeds in each locule. Seeds: red or black, fleshy, and covered by a red or orange aril.
Key to the species
1a Woody shrubs or trees; leaves simple; style-head distinctly wider than the style (section Turraea).
2a Petals short, less than 25 mm long; flower clusters at the tip of the shoot—T. nilotica.
2b Petals long, more than 35 mm long; absence of clustered flowers at the tip of the shoot.
3a Leaf less than 50 mm long; white, non-scented flowers; crystals in ray cells—T. obtusifolia.
3b Leaf 50–120 mm long; light green-scented flowers; no crystals in ray cells—T. floribunda.
1b Suffrutices; leaves simple or compound; style-head only slightly wider than the style [section Nurmonia (Harms) F.White].
4a Leaves simple; calyx lobes linear; petals oblong—T. pulchella.
4b Leaves compound, trifoliolate, or imparipinnate; calyx lobes oblong; petals broadly obovate—T. streyi.
Turraeafloribunda
Turraea floribunda Hochst. in Flora 27 (1): 297 (1844); C. DC. in Monogr. Phan. 1: 445 (1878); Bak. in Journ of Bot. 41: 12 (1903); Brenan in T.T.C.L. 320 (1949); Stapf in Curtis’s bot. Mag. 1: 8944 (1923); Eggeling and Dale, Indig. Trees Uganda Prot. 200 (1952); Staner and Gilbert, F.C.B. 7: 154 (1958); Dale and Greenway, Kenya Trees and Shrubs 274 (1961); White and Styles in F.Z. 2: 314 (1963); Killick in Flower. Pl. Afr. 38 (1967); Palmer and Pitman in Trees S. Afr. 2: 1061 (1973); White and Styles in F.S.A. 18 (3): 45 (1986); White in Bothalia 16 (2): 151 (1986). Type: South Africa, KwaZulu-Natal, Pietermaritzburg (2930): Umlaas River (–CD), 10 January 1839, Krauss 342 (TUB-image!, holotype; G, K, M, W, isotypes).
Turraea heterophylla sensu Sond. in Fl. Cap. 1: 245 (1860); sensu Medley Wood, Natal Plants 3, t. 246 (1902).
Deciduous shrub or small tree, up to 10 m tall; crown round; young branches loosely branched, pubescent, single or multi-stemmed, sometimes scrambling.
Bark: surface smooth to slightly rough with boat-shaped elongated fissures, light grey to brown; inner bark creamy to brownish; twigs with conspicuous lenticels; pith with prismatic crystals and druses; cortex with ideoblast cells; dilatation of cortical tissue is by anticlinal division of the parenchyma and cortical collenchyma cells as well as tangential stretching of cells forming strands of 2–7 cells; phelloderm cells with crystals; axial parenchyma with crystals; fibers present in secondary phloem; ray cells without crystals, rays without canal.
Wood: diffuse-porous, vessels slightly angular or rounded in outline, numerous, solitary, in clusters of 3–5, or in radial to diagonal multiples of 2–6, intervessel pits alternate, minute, 2.3–4.1 μm in diameter, with slit-like to oval apertures mostly united in grooves; fibers non-septate; axial parenchyma scanty paratracheal, diffuse, vasicentric, confluent, aliform, in bands up to 10 cells wide, prismatic crystals absent in chambered axial parenchyma cells; rays 1–2-seriate.
Leaves: simple, rarely compound, alternate, ovate to lanceolate, 25–120 × 10–70 mm, petiolate, noticeable herring-bone venation on both sides, margin entire, leaf base acute, leaf apex acuminate; petioles pubescent, 8–10 mm long; stipules absent.
Inflorescence: a (2–)3–18-flowered false raceme.
Flowers: bisexual, axillary or solitary, small, ca. 25 mm in diameter, light green, sweetly scented, in tight clusters at the end of the slim branches; sepals five, cup-shaped, small, densely pubescent; petals five, light green, ribbon-like; stamens 10, united into a thin, white tube slightly shorter than the petals; anthers form a ring at the tip of the staminal tube, forming a knob-like head; ovary sessile, 4–10-locular, with two ovules in each locule, pubescent, intensely sulcate, and transversely ribbed; style projects beyond the anthers, up to 20 mm long.
Fruit: a segmented capsule with 5–10 segments, woody, up to 26 mm in diameter, dark brown or black.
Seeds: small, glossy, orange to red, and borne on the opened woody star-shaped fruit (
Figure 15).
Distribution and ecology
The species grows in open woodland, coastal bush, riverine forest, dune forest, and secondary forest from sea level to about 750 m. It is found in the Gauteng, KwaZulu-Natal, and Eastern Cape provinces in South Africa (
Figure 16) and also in Eswatini (not shown in the map) and further north to Kenya and Uganda.
Diagnostic characters
Turraea floribunda is similar to T. obtusifolia in that the leaves are simple and mostly in fascicles. Turraea floribunda is readily distinguished by a noticeable herring-bone venation on both sides of the leaf: light green flowers with long, narrow, and sweet-scented petals vs. white flowers with wide, non-scented petals in T. obtusifolia.
Conservation status
Phenology
The species flowers during the spring and summer months (September to February) and fruits between February to July.
Specimens examined
South Africa. KWAZULU-NATAL: 2731 (Louwsburg): Ngotshe, Ngome forest margin (–CD), 27 July 1944, Gerstner 4874 (PRE). 2732 (Ubombo): Gwaliweni Forest (–AC), 19 November 1982, Abbott 532 (NH); Jozini to Hlatikulu Forest (–AC), 18 November 2016, Balkwill and McCallum 13932 (J); Ingwavuma (–AD), 15 November 1938, Gerstner 4116 (NH); Sibayi Nature Reserve; ca 4.3 [7 km] SW of Mabibi (–BC), 9 February 2018, Bester 14098 (PRE); Lake Sibiya, East Shore (–BC), 3 December 1982, Ward 286 (NH); Kosi Bay Coastal Forest Reserve, Lake Sibaya, eastern side of Sibaya along road between lake and first dune (–BC), 24 August 1994, Lubbe 250 (NH); Jozini Dam, on N side of valley (–BD), December 1964, Wearne 26 (NH, NU). 2830 (Dundee): Manzengwenya, coastal forest (–DA), 19 March 1965, Vahrmeyer 446 (NH); Mfongosi, in kloofs and fertile valleys (–DB), 4 April 1926, Watt and Breyer-Brandwijk 1095 (PRE). 2831 (Nkandla): ca 3 mi [4.8 km] E of Biyela store (–AC), 10 October 1946, Codd 1897 (PRE); Hlabisa, Hluhluwe Game Reserve, Hluhluwe River (–BB), 29 July 1983, Phelan 707 (NU); Umfolozi River (–BD), 22 October 1910, Rudatis 1213 (PRE); Mtunzini, Siyayi Lagoon, NE side, full shade (–DD), 14 February 1986, Venter 11474 (PRE); ca 3 mi [4.8 km] from Mandeni, off Tugela road (–DD), 23 November 1956, Edwards 1625 (NU). 2832 (Mtubatuba): Hluhluwe Game Reserve, Egodeni (–AA), 19 March 1972, Hitchins 790 (PRE, NU); Hluhluwe Game Reserve (–AA), 22 April 1959, Guy 107 (NH, NU); Lake St. Lucia, Charter’s Creek (–AB), 15 March 1967, De Winter 8729 (PRE); Edge of Lake Futululu (–AC), 25 November 1965, Moll 2751 (NH, NU); Mapelane (–AD), 25 February 1982, Gordon 44 (NH, NU); N coast of St. Lucia system (sensu lato), Mapelane Nature Reserve (–AD), 19 April 2008, Nkuna and Mabatha 2382 (PRE); Lake Bhangazi (south) area, eastern side of lake (–BA), 15 February 1997, Ward 13915 (PRE); Kwambonambi, Amangwe Forest (–CA), 18 August 1982, Von Breitenbach PRF 19580 (PRE); Richards Bay village area (–CC), 15 June 1969, Venter 5539 (PRE). 2930 (Pietermaritzburg): Botanic Gardens Estate, Pietermaritzburg (–CB), 22 October 1996, Stielau 247 (NH); Hela Mountains Game Valley Estates (–CC), 4 October 1982, Schrire 961 (NH); Camperdown, Nagle Dam, upper limit of bush (–DA), 14 May 1957, Wells 1278 (PRE); Nshongweni Dam (–DC), 22 February 1966, Morris 795 (PRE); Elsinore farm, Hope Valley (–DC), 27 September 1994, Sikhakhane 455 (NH); Silverglen Medicinal Plant Nursery (–DD), 17 June 2003, Sewram 10 (NH); Burman Bush Nature Reserve (–DD), 30 May 1986, Nicholas and Ngwenya 2193 (NH). 2931 (Stanger): Lower Tugela, S of Umvoti road at roadside (–AB), 24 September 1962, Johnson 1460 (NH); New Guelderland Stasie (–AD), 11 January 1967, Stewart 102 (NH); Hawaan Forest, ca 6.2 mi [10 km] N of Durban, Umhlanga Rocks (–CA), 22 May 1976, du Toit 1196 (NH); North coast, Umhlanga Rocks, Hawaan Forest (E), Umhlanga Lagoon Nature Reserve (–CA), 19 May 1990, Ward 10921 (NH, NU); Durban, Burman Bush Nature Reserve (–CC), 30 May 1986, Ngwenya 278 (NH); Scout Camp grounds, Burman Bush, Durban (–CC), 2 April 1984, Bodenstein 75 (NH). 3030 (Port Shepstone): Dumisa Stasie, Ellismere Umgayi (–AD), 22 October 1910, Rudatis 1213 (PRE); Umbogintwini (–BB), 6 January 1980, Van Wyk and Botha 3313 (PRE); Mtwalume, Koelwaters (–BC), 7 February 1975, Badenhorst 41 (PRE); Oribi Flats (–CA), 10 October 1996, Edwards 1317 (NH, NU); Oribi Gorge Nature Reserve, N of plains (–CB), 9 January 1981, Balkwill and Crankshaw 80/101 (J); Umtamvuna Nature Reserve, below Aerodrome (–CC), 15 October 1983, Abbott 1420 (NH); Uvongo, Skyline farm, rolling grassland hills with forest in depressions (–CD), 26 July 1966, Mogg 38493 (PRE).
EASTERN CAPE: 3129 (Port St. Johns): Lusikisiki, NW facing slope at Ndindindi Forest, Mkumbeni Location (–BC), 16 March 1994, Wopula 134 (NH); Mkambati Game Reserve (–BD), 15 January 1997, Sebothoma 152 (PRE); Ntafufu River, N of Port Johns (–CA), 10 May 1969, Strey 8512 (PRE); Thembuland, Coffee Bay (–CC), 5 November 1919, Tyson 10 (PRE). 3228 (Butterworth): Dwesa Nature Reserve (–BD), 17 November 1977, Jacot Guillarmod 7882 (PRE); Amathole, SE of Idutywa, Dwesa Nature Reserve (–BD), 19 October 1987, Fishwick 37 (J); Kentani, Kei Hills (–CB), November 1900, Pegler 727 (PRE); Umzimvubu District, Silaka Nature Reserve, ca 2.8 mi [4.5 km] SW of Port St. Johns (–CB), 26 January 2016, Bester 13446 (PRE); Willowvale, Qora River Mouth, northern bank of estuary above boat houses (–CC), 6 November 1991, Cloete 1261 (NH); Silaka Reserve, along road below main gate (–DA), 3 October 1992, Cloete 2075 (NH); 3326 (Grahamstown): Alexandria, NE of Woody Cape (–CB), 4 August 2007, Pienaar 0008 (PRE). 3327 (Peddie): E of East London (–BB), November 1961, Smith 46011 (PRE).
Turraea obtusifolia
Turraea obtusifolia Hochst. in Fl. Zonas Vida Ecuador 27: 296 (1844); White and Styles in F.S.A. 18 (3): 45 (1986); White in Bothalia 16 (2): 149 (1986). Type: South Africa, KwaZulu-Natal, Stanger (2931): Natal Bay (–CC), 7 January 1839, Krauss 308 (TUB-image!, holotype; G, K, isotypes).
Turraea oblancifolia Bremek. in Ann. Transvaal Mus. 15: 245 (1933). Type: South Africa, Limpopo, Waterberg (2428): between Warmbaths and Nylstroom (–AD), 15 January 1931, Bremekamp and Schweickerd 4 (PRE-image!, holotype; K, isotype).
Semi-evergreen (sometimes deciduous) shrub, scrambling shrub, or small tree, up to 5 m tall.
Bark: surface smooth, grey, pubescent; inner bark brownish; twigs with brown lenticels; branches glabrous; pith without prismatic crystals and druses; cortex without ideoblast cells; dilatation of cortical tissue is by anticlinal division of the parenchyma and cortical collenchyma cells as well as tangential stretching of cells forming strands of 2–5 cells; phelloderm cells with crystals; axial parenchyma with crystals; fibers absent in secondary phloem; ray cells with crystals; rays without canal.
Wood: diffuse-porous, vessel rounded to slightly angular in outline, numerous, solitary, in clusters of 3–5, or in radial to diagonal multiples of 2–6, intervessel pits alternate, minute, 2.3–4.1 μm in diameter, with slit-like to oval apertures commonly united in grooves; axial parenchyma scanty paratracheal in solitary strands near the vessel; prismatic crystals absent in chambered axial parenchyma cells; fibers non-septate; rays 1–2-seriate.
Leaves: simple, alternate, mostly in fascicles, variable in size and shape, oblanceolate to obovate, 20–50 × 8–25 mm, sometimes with two lobes at the tip, lower surface glabrous, glossy green, leaf base decurrent, leaf apex acute to acuminate; petiole very short or subsessile, 2–30 mm long, pubescent; stipules absent.
Inflorescence: a 1–3-flowered axillary raceme.
Flowers: bisexual, large, ca. 38 mm long, white; sepals 5-lobed, long, ca. 3–5 mm long, pubescent; petals 5, white, ligulate, narrow, 30–35 × 6 mm, glabrous, longer than staminal tube; stamens 10, united into a thin, white tube slightly shorter than the petals; anthers oblong, apiculate; ovary 5-locular, sessile, glabrous; style short, exserted, up to 5 mm long.
Fruit: a segmented capsule with 5–10 segments, woody, up to 15 mm in diameter, green.
Seeds: small, orange to red, kidney-shaped, covered by a fleshy, white aril (
Figure 17).
Distribution and ecology
The species grows in coastal forest, coastal dunes, woodland, wooded grassland, and bushland among rocks, from sea level to elevations of 1400 m. It can be found in Limpopo, North-West, Gauteng, Mpumalanga, KwaZulu-Natal, and Eastern Cape provinces in South Africa and Eswatini (
Figure 18), as well as in Mozambique and Zimbabwe.
Diagnostic characters
Turraea obtusifolia is similar to T. floribunda in that the leaves are simple and mostly in fascicles. Turraea obtusifolia is readily distinguished by its glossy, glabrous, oblanceolate to obovate leaves (20–50 × 8–25 mm, which might be lobed), white non-scented flowers, and presence of crystals in ray cells (bark tangential section) vs. ovate to lanceolate 25–120 × 10–70 mm leaves, light green scented flowers, and lack of crystals in ray cells (bark tangential section) in T. floribunda.
Conservation status
Phenology
The species flowers during the summer months (December to February) and fruits between January and June.
Specimens examined
Botswana. 2227 (Palapye): Selebi (–BB), December 1977, Kerfoot and Falconer 39 (PRE). 2324 (Lephepe): Kweneng, between Molepolole and Botlapatlou in river where rock engravings are found (–BC), 10 December 1986, Barnard 211 (PRE). 2425 (Gaborone): SE Botswana, Mokolodi Village, Mokolodi Reserve, ca 9.3 mi [15 km] S of Gaborone (–DD), 21 November 1996, Cole 1176 (PRE). 2525 (Mafeking): Kenye, Kenye Hospital (–AB), 3 June 1990, Hartley 1031 (J); Ramathlabama on Ghanzi road (–DA), 24 March 1976, Du Preez and Steenkamp 58 (PRE).
South Africa. LIMPOPO: 2229 (Waterpoort): Little Leigh farm, ca 12.4 mi [20 km] NW of town, Soutpansberg (–DD), 1 May 2000, Meyer 2819 (PRE), N of Soutpansberg, farm Davenham 740 MS (–DD), 8 May 1991, Balkwill 6334 (J). 2230 (Messina): Beacons Field, 212 MT, S of Mauluma, suid van randjie (–CC), 3 November 1979, Van Wyk 3001 (PRE). 2327 (Ellisras): Mogol Nature Reserve (–DD), February 1981, Fourie 2627 (PRE). 2330 (Tzaneen): Gravelotte, Eden 757 LT (–DC), 15 February 1987, De Villiers 50 (PRE). 2331 (Phalaborwa): Skiettocht Military Base (–CC), 11 March 1991, Manning 714 (PRE). 2427 (Thabazimbi): On Kareehoek farm, at the S foot of the Kranzberg between quartzite boulders (–BC), 9 May 1977, Venter 1955 (PRE); ca 18.6 mi [30 km] W of Vaalwater, Welgevonden Estate (–BD), 6 March 1993, Balkwill and Sebola 8023 (J); W of Vaalwater, on the farm Onvermoeid 246 KQ (–BD), 11 March 1990, Balkwill, Gesell and Williamson 5489 (J); ca 18.6 mi [30 km] W of Vaalwater, farm Van der Waltsdrift (–BD), 27 March 1989, Balkwill 4358 (J); ca 11 mi [17.7 km] NW of Rooiberg Tin Mine on road to Thabazimbi (–DC), 5 March 1948, Codd 3756 (PRE). 2428 (Nylstroom): N of Naboomspruit, Heinrichan LePhalala farm, central part of farm (–AB), 24 June 1993, Burgoyne 1610 (PRE); Geelhoutkop (–AD), January 1918, Breijer TRV 17826 (PRE); Mr Wilsons farm, ca 1.2 mi [2 km] along Warmwater Bronne turnoff, ca 7.5 mi [12 km] from Nylstroom (–CB), 6 February 1972, Stirton 5 (PRE). 2430 (Pilgrim’s Rest): Near Penge Mine, Sakarora waterfall (–AD), Burrows 3575 (J); Hoedspruit, Lissataba Private Nature Reserve, tussen klippe (–BC), 1 January 1987, Van Heerden 749 (PRE); Sekukuni, Driekop farm (–CA), 26 February 1936, Barnard 489 (PRE); Lydenburg, Steelpoort (–CB), 24 April 1999, Dehning 45 (J). 2431 (Acornhoek): Klaserie Private Nature Reserve, Dover 33 KU farm (–AA), 30 January 1980, Zambatis 922 (PRE).
NORTH WEST: 2426 (Mochudi): Ga-suping, Logaga 124 KP (–CC), 20 February 1985, Krynauw 204 (PRE). 2525 (Mafeking): Bophuthatswana, Motswedi Kloof (–BD), 1 January 1978, Peeters, Gericke and Burelli 545 (PRE). 2526 (Zeerust): Pilanesberg National Park, GA-Malau picnic site (–BB), 6 January 1991, Glen 2485 (J).
GAUTENG: 2527 (Rustenburg): De Wildt, near Pretoria (–DB), December 1928, Murray 638 (PRE); Hekpoort, in valley (–DC), 1 February 1959, Repton 5233 (PRE). 2528 (Pretoria): Hebron (–AA), April 1933, Dehnke TRV 31994 (PRE); Magaliesberg (–CA), 22 July 1976, Van Son 52174 (PRE); Pretoria National Botanical Garden, W of restrooms on N side of ridge (–CB), 21 September 2018, Bester 14245 (PRE). 2529 (Witbank): Dennilton, in vicinity of Bundu Inn Caravan Park in light shade of larger trees (–AC), 15 November 1977, Pienaar 1193 (PRE).
MPUMALANGA: 2429 (Zebediela): Farm Uitvlugt 887 KS, foot of Thaba Ya Sekhukhune Mountains, Steelpoort River Valley near future dam wall (–DD), 13 March 2007, Sachse 437 (PRE). 2431 (Acornhoek): Kruger National Park. Nkhuhlu exclosures, Skukuza, Sabie River (–DD), 22 March 2011, Van Coller, Siebert and Siebert 4425 (PRE). 2529 (Witbank): SE of Loskopdam Nature Reserve (–AD), 13 December 2001, Potgieter FP00190 (PRE). 2530 (Mashishing): Driekop farm, Sekukuniland (–AB), 4 January 1939, Mogg and Barnard 605 (PRE); Mbombela, Lowveld Botanical Garden side (–BD), 6 December 1969, Buitendag 381 (PRE). 2531 (Komatipoort): ca 4 mi [6.4 km] N of Brits on quartzite koppie (–AC), 6 December 1934, Mogg 14601 (PRE); Near Impala Siding, between Barberton and Komati Poort (–BC), 8 January 1929, Hutchinson 2493 (PRE); Malelane (–CB), December 1924, Murphy TRV26125 (PRE); Barberton between Malelane and Hector Spruit (–CC), 8 January 1929, Pole-Evans 24 (PRE).
KWAZULU-NATAL: 2632 (Bela vista): Ingwavuma, Ndumo Game Reserve (–CC), 11 February 1960, Tinley 574 (PRE, NU); Ingwavuma, Ndumu Hill, Ndumu Game Reserve (–CD), 15 February 1969, Pooley 384 (NH, NU). 2731 (Louwsburg): uPhongolo, ± 400 m W of Ivy’s Tuck Shop, eNcotshane (–BC), 16 February 2007, Keswa 1707 (NH); Jozini, Kwaliweni Forest, semi-shade, slope 5 deg (–BD), 21 February 1986, Venter 11580 (PRE); Itala Nature Reserve, Mtunzini contour path (–CB), 10 December 1987, MacDevette 2225 (NH, NU); Nongoma, ca 3.1 mi [5 km] W of Nongoma at disused coal mine near Vuna River (–CD), 28 March 1985, Reid 1074 (PRE). 2732 (Ubombo): Lebombo Mountains (–AA), 26 January 1963, Strey 4804 (PRE); Lebombo bushveld alongside road from Josini to Hlatikulu (–AC), 6 January 1986, Ward 1298 (NH); Lebombo Mountains, ca 11.8 mi [19.0 km] N of Josini (–AC), 28 November 1978, Tȍlken and Germishuizen 5791 (NH); Kosi Bay, N of Banga Nek (–BB), 24 June 1986, Buthelezi 667 (NH); Near Manzengwenya Inspection Quarters (–BB), 29 November 1969, Moll 4808 (NH); Lebombo Mountain, along road between Jozini and Hlatikula (–BC), 13 March 1993, Van Wyk 737 (NH); Mkuze Game Reserve (–BC), 29 January 1982, Bamps 7071 (PRE); Ingwavuma, Manzengwenya, Sibayi Coastal Woodland (–BD), 16 December 1984, Ward 896 (NH); Josini Mountain slopes (–CA), 14 May 1970, Strey 9795 (NH); Top of Jozini Pass opposite trading store (–CA), 18 February 1982, Reid 496 (PRE); Makowe Hills (–CC), 24 November 1960, Wells 2127 (NH); Sodwana Nature Reserve (–DA), 17 October 2001, Schuhardt DS00229 (PRE). 2830 (Dundee): Mfongosi (–DB), 22 June 1956, Edwards 1495 (PRE, NU). 2831 (Nkandla): Hlabisa, Hluhluwe Game Reserve, Vumbe stream bed (–BB), 1 November 1983, Phelan 734 (NU); Umfolozi Game Reserve, Thoboti camp area (–BD), 14 March 1966, Mthonti 13 (NH, NU); Umfolosi Game Reserve, Rustic Camp N aspect, open park-like vegetation, on rocky slope (–CC), 17 January 1981, Nichols 416 (NH); Eshowe (–CD), 28 January 1951, Lawn 1909 (NH); Hlabisa, E. shores, St. Lucia dune forest, N of Bangazi (–CD), 12 January 1978, Pooley 2071 (NU); Ngoye Forest, seaward side of forest (–DC), 19 February 1961, Wells and Edwards 77 (PRE); Ngoye Forest (–DC), 5 January 1986, Cunningham 2335 (NH); Ngoye Forest, outside northern boundary (–DC), 12 October 1984, Buthelezi 472 (NH); Mtunzini, Ngoye Forest Reserve (–DC), 8 February 1963, Huntley 267 (NU); Ngoye Forest Reserve (–DC), 30 March 1984, Balkwill, Cadman, Edwards, Getliffe and Meyer 1338 (J); Ngoye Forest Reserve, eastern section (–DD), 8 December 1993, Williams 1110 (NH). 2832 (Mtubatuba): Hluhluwe Game Reserve (–AA), 20 July 1961, Hitchins 56 (PRE); St Lucia Estuary, Sugarloaf campsite (–AD), 16 February 1986, Venter 11482 (PRE); Maphelane Nature Reserve (–AD), 2 July 1986, MacDevette 932 (NU); Maphelane Nature Reserve (–AD), 15 January 1985, Gordon 159 (NH); Cape Vidal, dune forest (–BA), 23 February 1982, Gordon 20 (NH); Hlabisa, Cape Vidal, dune forest (–BA), 3 June 1982, MacDevette 326 (NH, NU). 2930 (Pietermaritzburg): Nursery in Silverglen Nature Reserve, Chatsworth, Durban (–DD), 17 January 2005, Styles 2262 (NH). 2931 (Stanger): Ballitoville coastal forest, N of stream (–CA), 17 April 1986, MacDevette 758 (NH, NU); Hawaan Forest, S bank of Umhlanga River (–CA), 18 February 1970, Ross and Moll 2309 (NH); Balitoville, Zimbali Dune Forest (–CA), 1 July 1985, MacDevette 280 (NH, NU); Umgeni, Durban (–CC), 9 February 1914, Medley-Wood I2573 (PRE, NU). 3030 (Port Shepstone): Sea Park (–CB), 25 November 1968, Strey 8065 (NH); Uvongo Nature Reserve, ca 190 mi [305.8 km] from Nicholson Wandelpad aan regterkant (–CD), 14 July 1976, Venter 1035 (PRE); Hibberdene, behind first dune, coastal forest (–DA), 26 March 1976, Mogg 36964 (PRE).
EASTERN CAPE: 3129 (Port St. Johns): Mgwelyana (Kambati), dune bush (–BD), 20 May 1969, Strey 8660 (NH); Coffee Bay, near coast (–CC), November 1919, Tyson 30 (PRE); Mtata mouth, first kloof on S bank (–CC), 8 January 1985, Abbott 2396 (NH); Transkei, farm Hluleka near Ngqeleni (–CD), 5 January 1968, De Winter 8861 (PRE). 3227 (Stutterheim): Amalinda (–DD), 1 January 1954, Nanni 97 (PRE). 3228 (Butterworth): Willowvale, Qora River Mouth, N bank of estuary (–AD), 21 December 1991, Cloete 1531 (NH); Quora Mouth, dune bush (–BC), 25 March 1973, Strey 11186 (NU); Elliotdale, Bashee River Mouth, forest edge, N of estuary (–BD), 19 December 1991, Cloete 1484 (NH); Kentani coast (–CB), December 1913, Pegler 337 (PRE); Qora River Mouth, dunes at river mouth (–DD), 13 September 2001, Rourke 2210 (PRE). 3326 (Grahamstown): Alexander Fontein near chicken farm (–CB), 13 October 1954, Johnson 1063 (PRE); Kenton-on-Sea, Bushmans River mouth (–DA), December 1949, Leighton 3136 (PRE); Port Alfred, West slopes (–DB), October 1916, Tyson TRV 17229 (PRE). 3327 (Peddie): Kidds Beach (–BA), December 1943, Giffen FH1793 (PRE); East London, ca 13.8 mi [22.2 km] from East London on Kidds Beach/Peddie Road (–BA), 21 July 1955, Comins 1265 (PRE); East London, Bulura (–BB), 5 May 1972, Von Gadow 59 (PRE).
Eswatini. 2631 (Mbabane): Lubombo Mountains, ca 10.6 mi [17 km] N of Siteki on Siteki/Mhlume road between Groenpan and Cyrildene farms (–BD), 2 May 1976, Culverwell 16 (PRE); Hlatikulu, Ingwavuma Port hillside (–CD), 13 November 1959, Compton 29436 (PRE). 2531 (Komatipoort): Tshaneni, Lowveld near homestead (–DC), 20 December 1969, Barrett 401 (PRE).
Turraea nilotica
Turraea nilotica Kotschy and Peyr. in Plantae Tinneanae 12: 6 (1867); Oliv. in F.T.A. 1: 331 (1868); C. DC. in Mon. Phan. 1: 445 (1878); Gȕrke in Engl. Pflanzenw. Ost-Afr. 231 (1895); Monro in Proc. Rhod. Sci. Ass. 7: 68 (1908); Sim, For. Fl. Port. E. Afr. 25 (1909); Eyles in Trans. Roy. Soc. S. Afr. 5: 388 (1916); Brenan in T.T.C.L. 321 (1949); Suesseng. and Merxm. in Proc. and Trans. Rhod. Sci. Ass. 43: 109 (1951); Garcia in Contr. Conhec. Fl. Moçamb. 2: 140 (1954); Pardy in Rhod. Agr. Journ. 52: 38 (1955); Williamson in Useful Pl. Nyasal. 120 (1956); Palgrave in Trees of Central Afr. 231 (1957); Dale and Greenway, Kenya Trees and Shrubs 275 (1961); White, F.F.N.R. 182 (1962); White in Bothalia 16 (2): 149 (1986); White and Styles in F.S.A. 18 (3): 43 (1986). Type: Somalia, Rive del Giuba a Ferdadale, 21 October 1913, Paoli 981 (FT-image! syntype).
Turraea randii Baker f. in Journ. Bot. 37: 427 (1899); Gibbs in Journ. Linn. Soc., Bot. 37: 435 (1906). Type: Zimbabwe, Salisbury, July 1898, Rand 562 (BM).
Turraea tubulifera C. DC. in Ann. Conserv. Jard. Bot. Genève, 10: 133 (1907). Type: Tanzania, Tanganyika, Zanguebar Bagamoyo, no date, Sacleux 471 (H).
Shrub or small deciduous tree, up to 10 m tall; young branches pubescent and smooth in older trees.
Bark: surface pubescent in younger stem, smooth to cork-like and flaking with age; twigs with conspicuous whitish lenticels.
Leaves: simple, elliptic to obovate, 100–160 × 80–120 mm, alternate, pubescent with curved edges, margin entire and rolled below, venation conspicuous below, petiolate, leaf base cuneate, leaf apex tapering to rounded; petioles pubescent, 15–18 mm long, stipules absent.
Inflorescence: a 5–18-flowered, sessile or subsessile fascicle in the axils of fallen leaves.
Flowers: in axillary clusters, small, ca. 25 mm in diameter, greenish white to yellow with age, sweetly scented; sepals 5, ca. 3 mm long, denticulate, puberulous; petals 5, 13–24 × 3–5 mm, greenish white turning yellow with age, sparsely puberulous; stamens 10, united into a long, thin white staminal tube, staminal tube pubescent, as long as anthers; ovary sessile, 4–10 locules, with two ovules in each locule; style exserted, projects beyond the staminal tube.
Fruit: a segmented capsule with 5–10 segments, glabrous, thinly woody, and up to 13 mm in diameter.
Seeds: black, 5 × 3 mm, kidney-shaped, and covered by orange to red arils (
Figure 19).
Distribution and ecology
The species grows in bushland and scrub woodland below an elevation of 800 m. It is found in the Limpopo and Mpumalanga provinces in South Africa and in Botswana (
Figure 20). It also occurs in Mozambique, Zimbabwe, Malawi, Uganda, Kenya, Ethiopia, and Sudan.
Diagnostic characters
Turraea nilotica can be distinguished from T. obtusifolia and T. floribunda by the shorter petals (up to 22 mm long) and the staminal tube that is distally expanded (petals more than 30 mm long and the staminal tube not expanded at the tip in the other two species above). It is also superficially similar to Vangueria infausta in that the leaves are simple, pubescent, and elliptic to obovate in shape. Turraea nilotica Is readily distinguished by its alternatively arranged leaves, slender petals, and exerted style, which is approximately twice the length of the staminal tube vs. opposite leaves, wide petals, and a short style in V. infausta.
Conservation status
Phenology
The species flowers during the winter and spring months (June to October) and fruits between September and February.
Specimens examined
Botswana. 1724 (Katima Mulilo): Kabulabula, Chobe River, Bechuanaland North (–DD), July 1930, Van Son TRV 28944 (PRE).
South Africa. LIMPOPO: 2229 (Waterpoort): Soutpansberg, Free State Mine (–DD), July 1917, Breijer TRV 17571 (PRE). 2230 (Messina): Musunda near road to Musunda Village (–DE), 2 October 1979, Khorommbi 1005 (PRE); Vuvha (–CD), 7 October 1981, Van Wyk and Theron 4943 (PRE); Musunela (–DB), 10 July 1981, Netshiungani 1448 (J). 2231 (Pafuri): Kruger National Park, Punda Milia, Sjantangalane area (–CA), 19 October 1966, Van Wyk 4748 (PRE). 2328 (Baltimore): Senwabarwana, Blouberg Nature Reserve (–BB), 28 July 1973, Davidson s.n. (J). 2329 (Polokwane): Buffelshill (–CD), November 1918, Rogers TRV 18935 (PRE); Munnik (–DE), 18 January 1946, Gerstner 5807 (PRE). 2330 (Tzaneen): Elim (–AA), June 1930, Obermeyer 507 (PRE); Koedoes River Valley, on ridge (–CA), September 1923, Keet 1188 (PRE); Moeketsi, 9 min NW of Mooketsi on Soekmekaar road (–CA), 21 August 1968, Leistner, Thom and Gillham 3286 (PRE); E of Tzaneen on road to Gravelotte (–CD), 15 September 1972, Van Vuuren 1823 (PRE); Hans Merensky Nature Reserve, ex Transvaal Nature Conservation Herbarium (–DA), 16 September 1969, Oates 32 (PRE); Kruger National Park, Rubbervale (–DC), 3 October 1932, Lang TRV 32116 (PRE). 2331 (Phalaborwa): Silonque plot 2, ca 6.2 mi [10 km] N of Phalaborwa at foot of Masakoleng outcrop (–CC), 21 October 1987, Retief 643 (PRE). 2431 (Acornhoek): Klaserie, Guernsey farm (–AC), 13 June 1971, Burger 43 (PRE); Ba-Phalaborwa, E of Klaserie, Umbabat Bushveld (–CA), 2 October 1990, Shackleton 715 (J).
MPUMALANGA: 2430 (Pilgrim’s Rest): ca 12.4 mi [20 km] from Hoedspruit farm called Trekkers, on slope of Blyde River Canyon Mountain, ca 9.3 mi [15 km] from Swadini (–BD), 3 October 2002, Male JM00274 (PRE); Escarpment joining N. section of Drakensberg, Chester farm (–BD), 16 September 1984, Rushworth 34 (NU); Klaserie, on road to Mariepskop (–DB), 17 November 1987, Venter 12688 (NU). 2431 (Acornhoek): Pilgrims Rest near Bushbuck Ridge P.O. (–CC), 18 August 1946, Acocks 12892 (PRE); Lothian (–CC), August 1923, Keet 1258 (PRE). 2530 (Mashishing): Mbombela, Lowveld Botanical Garden, garden side (–BD), 6 October 1969, Buitendag 144 (PRE); Mashishing, Lowveld Botanical Garden, garden side central area (–BD), 24 September 1970, Buitendag 680 (PRE). 2531 (Komatipoort): Shabin, Kruger National Park (–AA), 20 August 1952, Van der Schijff 710 (PRE); Kruger National Park near Pretoriuskop gate (–AA), 28 August 1950, Codd 6117 (PRE); Kruger National Park (–AA), 14 January 1952, Van der Schijff 176 (PRE); Kruger National Park, Pretoriuskop, Pretoriuskop and neighbourhood (–AB), 26 October 1950, Story 3908 (PRE); Hazyview township, about 50 m W of the Hazyview residential area, turn-off on the Hazyview-Skukuza Road (–BA), 4 October 2000, Schmidt 2631 (PRE).
Turraea pulchella
Turraea pulchella (Harms) T.D.Penn. in Blumea 22 (3): 454 (1975); White and Styles in F.S.A. 18 (3): 47 (1986). Type: South Africa, Eastern Cape, Butterworth (3228): Transkei, near Kentani (–CB), March 1903, Pegler 730 (B, holotype†; Kentani District, 305 m, fl.fr. 15 Oct. 1904, Pegler 730 (K-image!, neotype, designated by White (1986); BOL, GRA, PRE!, SAM, isoneotypes).
Nurmonia pulchella Harms. in Ber. Deutsch. Bot. Ges. 35: 81 (1917). Type: Same as for Turraea pulchella.
[Note: Pegler 370 in the Berlin herbarium was previously designated as type, but it was destroyed during WWII, so White (1986) chose Pegler 370 in the Kew Herbarium as neotype, with the explanation that there is no definite evidence that it is a duplicate of the Berlin specimen, even though it bears the same collector’s number].
Shrublet, up to 0.3 m tall; rootstock strong, woody; branchlet strong, woody.
Leaves: simple, setulose, ovate with the broadest part close to the leaf apex, shallowly lobed in the upper half, 25–30 × 10–17 mm, alternate, petiolate, pubescent, margin entire, leaf base acute, leaf apex obtuse; petioles 5–8 mm long; stipules absent.
Inflorescence: a (2–)3–18-flowered false raceme.
Flowers: bisexual, axillary or solitary, white, spicy fragrance, produced in 2–3 cymules; sepals free, small, hispidulous; petals 5, oblong, white, pubescent; stamens 10, united into a staminal tube which splits at the tip, as long as anthers; anthers apiculate; ovary sessile, 5-locular, with two ovules in each locule, pubescent; style up to 20 mm long, head capitate with 5 stigmatic lobes at the tip.
Fruit: a segmented capsule with 5 segments, thinly woody, up to 10 mm in diameter (
Figure 21).
Distribution and ecology
The species grows on the boundaries between grassland and thicket (Valley Bushveld) above river gorges or on top of the sandstone plateau at elevations between 300 and 650 m. It is found in KwaZulu Natal and the Eastern Cape Province in South Africa (
Figure 22).
Diagnostic characters
Turraea pulchella is similar to T. streyi in that the leaf surfaces are setulose and flowers occur in cymules. Turraea pulchella is readily distinguished by its simple ovate leaves, larger flowers, and style head, which is differentiated into 5 stigmatic lobes and oblong petals vs. compound, mostly trifoliolate and lobed leaves, smaller flowers, a less differentiated style head, and obovate petals in T. streyi.
Conservation status
Phenology
The species flowers during the late spring to summer months (September to March).
Specimens examined
South Africa. KWAZULU-NATAL: 2930 (Pietermaritzburg): Krantzkloof Nature Reserve (–DA), 2000, Scott-Shaw 10363 (NU); Matabetule Plateau (–DB), 22 Oct, 1987, Williams 45 (NH); Inanda, Matabetule Plateau (–DB), 27 November 1987, Williams 84 (NH); Matabetule Plateau, c. 20 km NW of Durban (–DB), 18 September 1987, Williams 36 (NH); Matabetule Plateau (–DB), 15 February 1988, Williams 217 (NH); Inanda, Greater Durban Metropolitan area, Hammarsdale area, Hector (ESKOM) (–DC), 9 December 1995, Ward 13427 (NH); eThekwini, Tabankulu Hill (W of Shongweni exit from N3, with cane watch tower) (–DC), 3 January 2006, Wragg 1469 (NU); Pinetown, Marianhill (–DD), 29 September 1994, Ngwenya 1290 (N, PRE). 3030 (Port Shepstone): Mzamba, E of Greenville Mission on slopes above Umtamvuna, on level ground above Umtamvuna River (–CC), 10 January 1995, Abbott 6659 (NH).
EASTERN CAPE: 3228 (Butterworth): Kentani Valley (–CB), October 1911, Pegler 730 (NH).
Turraea streyi
Turraea streyi F. White and Styles in Bothalia 16 (2): 153 (1986); White and Styles in F.S.A. 18 (3): 47 (1986). TYPE: South Africa, KwaZulu-Natal, Port Shepstone (3030): St Michael’s-on-Sea, Deppe’s farm, a heavily browsed ‘bush’ in bushveld (–CD), 19 Sept 1966, Strey 6876 (PRE!, holotype; NH!, NU, isotypes).
A multi-stemmed rhizomatous suffutex, up to 0.75 m tall, branchlets setulose.
Leaves: imparipinnately compound, mostly trifoliolate, alternate, with 2–5 divisions, the leaflets lobed, with rounded to acute apices, 50–70 × 8–20 mm, surface setulose, green, petiole very short, sessile, 2–20 mm long, stipules absent.
Flowers: in 2–3-flowered cymules, sepals 3–5 mm long, hispidulous; petals 5, broadly obovate, white, glabrous, 90–110 × 20–40 mm; staminal tube, up to 9 mm long, pubescent; ovary 5-locular, with two collateral ovules in each locule, pubescent; style short, slightly exserted, up to 10 mm long, pubescent.
Fruit: a segmented capsule, with 5–10 segments, woody, up to 13 mm in diameter. (
Figure 23).
Distribution and ecology
The species grows at the margins of scrub forest, bushland, and coastal grassland at elevations between 30 and 100 m. It is found between Port Shepstone and Stanger in KwaZulu-Natal (
Figure 24). It is now also known from a large population of ca. 60 to 80 individuals in a new locality, ca. 20 km north of Amandawe (ca. 15 km south of the Umkomaas River towards Amandawe). A few plants have also been found in Amandawe Village (see below). Amandawe is situated ca. 7 km east of Scottburgh and ca. 60 km south of Durban.
Conservation status
The current status of the species is given as Critically Endangered (Possibly Extinct,
http://redlist.sanbi.org/species.php?species=957-9, (accessed on 28 March 2019)). This assessment was based on the fact that the two known wild subpopulations have completely disappeared due to the encroachment of invasive alien plants and that only a small subpopulation of three introduced (planted) individuals exists near the type locality. The assessment concluded that there is not much hope of finding the plant elsewhere in its former range [
55,
56]. Recently, however, the two new populations referred to above were discovered by Mr. Lloyd Mhlongo, a PhD botany student at the University of Johannesburg [
57]. This means that the conservation status needs to be reassessed. According to the criteria of the IUCN [
53,
54], the species remains critically endangered.
Diagnostic characters
Turraea streyi is similar to T. pulchella—both species are small suffrutices with setulose leaf surfaces and white flowers borne in 1–3-flowered cymules. Turraea streyi is readily distinguished by its compound leaves (usually trifoliate but sometimes with five leaflets), the smaller size of the flowers, and broad, obovate petals (simple leaves, larger flowers, and oblong petals are diagnostic for T. pulchella).
Specimens examined
South Africa. KWAZULU-NATAL: 2931 (Stanger): King Hamlyn’s farm (–AD), 19 September 1971, Moll 5502 (NH). 3030 (Port Shepstone): Ca. 20 km north of Amandawe (–BA), Grieve KWG2973 (NH); Uvongo Road (–CD), 1 October 1987, Williams 44 (NH); Uvongo Road (–CD), 27 November 1969, Strey 9288 (NH); St. Michael’s-on-Sea, Deppe’s farm next to “Skyline” (–CD), 1 October 1987, Williams 43 (NH); Deppe’s farm, next to “Skyline”, St. Michael’s-on-Sea (–CD), 10 December 1968, Ross 1853 (NH); Port Shepstone, St Michael-on-Sea. Deppe’s farm (–CD), 19 September 1966, Strey 6876 (NH, NU).