Uapaca Kirkiana (Phyllanthaceae) and Its Relatives in the Northwest of the Zambezian Region

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Uapaca kirkiana (Phyllanthaceae) and its relatives in the northwest of the Zambezian region

Pierre Meerts1,2 & João Farminhão1

1 Herbarium et bibliothèque de botanique africaine, Université Libre de Bruxelles, Avenue

F.D. Roosevelt 50, CP 265, B-1050 Bruxelles, Belgium.

Link to published version:

Kew Bulletin 74 https://doi.org/10.1007/s12225-019-9826-1 2

Summary. The complex of Uapaca kirkiana is revised in the northwest of the Zambezian region (Angola, D. R. Congo, Zambia) based on herbarium taxonomy. Five taxa are recognised, i.e. Uapaca kirkiana Müll. Arg. var. kirkiana, U. kirkiana var. gossweileri

(Hutch.) Meerts, U. kirkiana var. benguelensis (Müll. Arg.) Meerts, U. teusczii Pax f. teusczii and U. teusczii f. pilosa (P. A.Duvign.) Meerts. A determination key is provided. Specimens lacking curly hairs on leaves have long been misinterpreted or overlooked. The name Uapaca teusczii Pax is resuscitated to accommodate such specimens. New diagnostic characters of U. teusczii are highlighted including a glabrous ovary, anthers with ciliate connectives and the pubescent calyx of female flowers. U. teusczii is a characteristic species of the wet miombo, widespread in Katanga and Kwango (D. R. Congo), extending locally to Angola and N

Zambia. A lectotype is selected for U. albida, U. homblei, U. kirkiana var. benguelensis, U. kirkiana var. gossweileri, and U. teusczii f. pilosa; second-step lectotypification is provided for U. munamensis.

Key Words. Angola, edible fruit, D. R. Congo, miombo, taxonomy, dry tropical woodland,

Zambia.

Introduction

The genus Uapaca Baill. (Baillon 1858) (Phyllanthaceae, Antidesmatoideae) comprises dioecious trees and shrubs, widespread in forests of tropical Africa and Madagascar. Uapaca species are amongst the most typical components of the shrub layer of dry tropical woodlands and savannahs in the Zambezian region (Werger & Coetzee 1978; White 1983). Uapaca kirkiana Müll. Arg., the most widespread species in that region, is well known for its edible, much sought fruits locally known as “masuku” (Akinnifesi et al. 2006; Ngulube et al. 1996).

Domestication of Uapaca kirkiana has been successfully attempted, and breeding programs 3

are being developed (Ngulube et al. 1996; Akinnifesi et al. 2006; Mwase et al. 2015). In this context, improving the taxonomic knowledge of the species and its relatives is essential.

Müller Argoviensis (1864a,b) described two species to accommodate specimens of

Uapaca with foliar indumentum consisting of curly, interlacing hairs i.e. Uapaca kirkiana

Müll. Arg. and U. benguelensis Müll. Arg. The former name is based on a type from Malawi and the latter, published a few days later, is based on a type from Angola. Ever since, the taxonomy and nomenclature of that complex have been confusing. Pax (1904) accepted both species, but did not indicate discriminant characters, the two species being separated based on geographical distribution. Hutchinson (1912a) pointed out that the original material on which the name U. benguelensis is based, i.e. Welwitsch 453, was heterogeneous, comprising two distinct taxa, i.e. one with petiolate leaves and the other one with sessile leaves. He described the latter as a new species, U. gossweileri Hutch., thus implicitly restricting U. benguelensis to petiolate forms. All three species (U. kirkiana, U. benguelensis, U. gossweileri) were accepted by Hutchinson (1912b) and Pax & Hoffmann (1922). The latter work distinguished

U. kirkiana from U. benguelensis based on the lower number of secondary veins and the subsessile inflorescence of the latter. De Wildeman (1936) recognised several other taxa supposedly endemic to D. R. Congo in this complex, which were later synonymised with U. kirkiana or U. benguelensis by Duvigneaud (1949). In his revision of the genus for Katanga

(D. R. Congo), Duvigneaud (1949) considerably extended the circumscription of U. benguelensis, to include forms with glabrous leaves and forms with straight hairs. Having re- examined alleged differences between U. benguelensis and U. kirkiana, Radcliffe-Smith

(1993) concluded that there was considerable overlap between them and, accordingly, he synonymised both names under U. kirkiana. However, in his subsequent treatment of the genus Uapaca in Flora Zambesiaca, Radcliffe-Smith (1996) admitted that the question was not settled yet and that there might actually be two distinct species. The African Plant 4

Database (consulted in Aug. 2018) treats U. benguelensis as a synonym of U. gossweileri

(which is certainly incorrect from a nomenclatural point of view), and U. benguelensis var. pedunculata as a synonym of U. kirkiana. The Plant List (http://www.theplantlist.org), in contrast, accepts U. benguelensis as a distinct taxon. The World Checklist of Selected Plant

Families (http://apps.kew.org/wcsp/qsearch.do?plantName=Uapaca&page=quickSearch) also recognises U. benguelensis, U. gossweileri, U. kirkiana at species level.

We here revise the taxonomy of this group, considering material from Angola, D. R.

Congo and Zambia, the region which concentrates most of the variation in the complex. Five taxa are recognized, and four new combinations are proposed.

Materials and methods

We have revised the material from D. R. Congo, Angola and Zambia kept in the following herbaria: BM, BR, BRLU, COI, K, LSHI, LISC, LISU, P, POZG, WAG. The following characters, used in previous taxonomic treatments, were recorded: twig thickness and pubescence; leaves: petiole length, lamina length and width, shape of lamina base and apex, indumentum (curly vs. straight hairs; density), number of secondary veins; inflorescence: position respective to leaves; length of male and female peduncle, length of bracts; fruits: peduncle length and diameter, fruit diameter, indumentum, length and width of pyrenes. For the sake of simplicity, male and female inflorescences are hereafter referred to as “flowers”.

For D. R. Congo, the specimens are grouped according to the phytogeographic divisions of

Robyns (1948). Within each country, provinces or regions are cited in alphabetic order.

Key to the taxa of the Uapaca kirkiana complex in NW of the Zambezian region

1. Ovary glabrous or with sparse straight hairs; anthers with connective ciliate; lower surface

of lamina of mature leaves glabrous or with straight hairs (occasionally with sparse

remains of curly hairs on veins); lamina with 7 – 12 (– 13) pairs of secondary veins; tube

of female calyx setulose, with hairs persisting at the fruiting state 2

1. Ovary densely tomentose, with curly hairs; anthers with connective usually glabrous; lower

surface of lamina pubescent, with indumentum consisting of curly hairs not restricted to

veins; lamina with (8– ) 12 – 25 pairs of secondary veins; female calyx generally glabrous

when in fruit, or with curly hairs mostly at margin 3

2. Lower surface of leaf blade glabrous Uapaca teusczii f. teusczii

2. Lower surface of leaf blade with straight hairs Uapaca teusczii f. pilosa

3. Leaves grouped in a rosette at the apex of twigs, sessile, long attenuate at base into a

winged pseudopetiole; second year’s twigs (8 –) 10 – 25 mm in diam., generally corky and

marked by prominent leaf scars Uapaca kirkiana var. gossweileri

3. Leaves not grouped in a rosette at the end of twigs, generally petiolate, occasionally

sessile, cuneate, not attenuate in a winged pseudopetiole; second year twigs generally < 10

mm in diam., not corky 4

4. Male flower head stipitate; male inflorescence globose before anthesis; inflorescences

(male and female) and fruits generally borne 5 – 30 cm below the leaves on defoliated

previous year’s twigs; male peduncles 5 – 20 mm long, mostly in fascicles of 2 – 9; lamina

with (12 –) 13 – 21 (– 25) pairs of secondary veins Uapaca kirkiana var. kirkiana

4. Male flower head sessile; male inflorescence longer than wide before anthesis;

inflorescences (male and female) and fruits mostly amongst the leaves on current year’s 6

growth; male peduncles 1 – 10 mm long, mostly solitary; lamina with 8 – 13 (– 14) pairs of

secondary veins Uapaca kirkiana var. benguelensis

Taxonomic Treatment

1. Uapaca teusczii Pax (1894: 79); Pax (1904: 371); Hutchinson (1912b: 641); Pax &

Hoffmann (1922: 301); De Wildeman (1936: 179). Type: Angola, Malange, Teuscz 423

(holotype B; isotype K!).

Uapaca angolensis Hutch. in schedis, unpublished name.

Uapaca benguelensis var. pedunculata P. A.Duvign. (Duvigneaud 1949: 890). Type: D. R.

Congo, Haut-Katanga, entre Elisabethville [Lubumbashi] et Katofio, Duvigneaud

1214U3 (holotype BRLU!; isotype FHO), synon. nov.

Uapaca benguelensis sensu White (1962: 206), Malaisse (1997: 68), Meerts (2016: 238),

Meerts & Hasson (2016: 183) non Müll. Arg.

Small tree 2 – 5 (– 7) m high. Twigs 5 – 10 mm diam. below the lowermost leaf, blackish- gray, often with a silvery shine, generally glabrous except apical bud and at insertion of peduncles, more rarely shortly rusty-beige crisped pubescent near tip. Leaves with petiole 1.5

– 4.5 (– 5.5) cm, terete or flat on upper surface, glabrous or, more rarely, thinly puberulent to floccose (short, undulate hairs), epidermis lepidote generally flaking of in thin silvery scales; leaf blade (6.6 –) 9 – 19 (– 22) × (4.5 –) 6.6 – 14 (– 18) cm, broadly elliptic, obovate, obovate-elliptic, suborbicular to almost obcordiform, with length to width ratio typically ≤

1.5, cuneate to rounded at base, broadly rounded to truncate or, more rarely, emarginate at apex; upper surface glabrous (rarely with remains of hairs on main nerve), lepidote, markedly glutinous at the beginning, green and shiny when fresh, dull greyish green to deep brown in herbarium, sparsely lepidote, lower surface very pale silvery-green when fresh, with dark 7

green reticulum, generally turning reddish-brown and rugulose in herbarium, sparsely lepidote, completely glabrous or with scattered straight pallid hairs ca. 1 – 1.5 mm long, denser on veins, occasionally with sparse remains of short curly hairs on veins; main vein slightly prominent on upper surface, with epidermis usually flaking off in small silvery scales, mostly glabrous, secondary veins 7 – 12 (– 13) pairs, slightly impressed, more or less anastomosing in arches on lower surface, tertiary veins inconspicuous on upper surface, conspicuous and slightly prominent on lower surface, irregular or more or less perpendicular to secondary veins; primary and secondary veins often with a reddish tinge in young leaves.

Male inflorescence in the axils of leaves or just below the leaves on current year’s growth; peduncle solitary, glabrous to sparsely puberulent (appressed hairs) or lepidote, 2 – 10 (– 20) mm long, ca. 1 mm thick, angular, glabrous to sparsely pilose; “flower” bud ovoid to obovoid, markedly longer than wide, bracts ovate-elliptic to oblong-elliptic, 7 – 11 × 2 – 5 mm, the outermost ones narrower, sparsely pilose outside to glabrous; male flower head sessile; calyx with simple straight hairs, filament 1 – 2 mm, glabrous or, more rarely, with 1 – 2 cilia, anther ca. 1 mm long, with connective generally ciliate at least in some stamens (cilia ca. 0.3 mm long), pistillode pubescent, ca. 1.6 mm long, distal part ca. 1.5 mm diam. Female inflorescence solitary in the axils of leaves or just below the leaves on current year’s growth; peduncle 1 – 6 mm, glabrous, lepidote or, more rarely, puberulent; bracts ca. 8 – 11 × 3.5 – 5 mm, the outermost ones pilose outside; calyx cupuliform, ca. 1 mm long, shallowly lobed, tube setulose with appressed hairs 0.5 – 1.5 mm long persisting in fruit; ovary glabrous or with sparse straight hairs, smooth and shiny or squamulose-lepidote, often blackish-green in herbarium, styles 3 (– 4), reflexed and almost hiding young ovary, multifid, adaxially pubescent. Fruit with peduncle 2 – 10 (– 15) × 2 – 3.5 (– 4) mm, typically ca. 2 – 3 times as long as thick, glabrous or sparsely puberulent; drupe globose to very shortly ellipsoidal, glabrous, ca. (15 –) 20 – 35 mm diam, with whitish lenticels, persistent calyx with a fringe of 8

cilia near base; wall thin (ca. 1.5 mm), (3 –) 4 – 5 pyrenes, (11 –) 15 – 22 (– 25) × 11 – 13 mm, ellipsoid-oblong to narrowly ovoid, rounded to pointed at base, cuneiform-apiculate at tip, with the median ridge blunt in transverse section, seed brown, smooth, ca. 12 × 8 mm.

Fig. 1. Fig. 3A, C, E, G.

DISTRIBUTION. Angola, D. R. Congo, Zambia (rare). Map 1.

HABITAT. Miombo woodland (dry tropophilous forest dominated by Brachystegia spp.) and wooded savannah with Marquesia macroura, Julbernardia paniculata, Brachystegia floribunda and Isoberlina angolensis, on deep red soil or on rocky soil; characteristic of the wet miombo of the northern part of the Zambezian region, rich in evergreen species (Berlinio-

Marquesion), in regions with rainfall > 1100 mm/y. In Kwango (SW D. R. Congo), in

“mikondo” vegetation, i.e. tropophilous forest (“Association à Julbernardia paniculata et

Brachystegia longifolia” (Devred et al. (1958)). U. teusczii usually occurs on deeper, more fertile soil than U. kirkiana. It is apparently less tolerant of fire than U. kirkiana and, unlike the latter, it does not usually persist in degraded shrub savannah; alt. 650 – 1700 m.

CONSERVATION STATUS. Least Concern (LC). The species is widespread in the northwest of the Zambezian region.

VERNACULAR NAMES. Mundengudengu musuku (Kibemba), mupupola (Kitwaba).

USES. Edible fruit.

PHENOLOGY. Flowering in the middle of the rainy season (Jan. – Feb.).

NOTES. This taxon is very distinct, in both vegetative and reproductive traits. In addition to having the lamina almost glabrous or with straight hairs only, U. teusczii is unique in the U. kirkiana complex in having a glabrous ovary. In all the material referable to U. kirkiana, the ovary is densely crisped-pubescent. Another discriminant trait that had not been reported hitherto is the ciliate stamen connective (vs. connective glabrous in U. kirkiana (rarely with a 9

single hair)). Therefore, it seems fully justified to treat this taxon at specific rank. The earliest name available at this rank is Uapaca teusczii Pax, described from Angola, of which the type specimen matches glabrous material from D. R. Congo.

Uapaca teusczii has been consistently overlooked or misinterpreted by all authors after Pax

& Hoffmann (1922). A first reason is that specimens with glabrous leaves in the U. kirkiana complex are relatively rare outside of D. R. Congo. Hutchinson (1912b) and Pax & Hoffmann

(1922) cited only one and two collections of U. teusczii from Angola, respectively. De

Wildeman (1936) described two new taxa to accommodate glabrous forms from D. R. Congo

(i.e. U. munamensis De Wild. and U. dubia De Wild.), which are in fact referable to U. teusczii var. teusczii. Duvigneaud’s (1949) treatment of glabrous specimens in Katanga was very confusing. Based on spurious differences in pyrene shape, he ascribed part of the glabrous specimens to “U. benguelensis var. pedunculata P. A.Duvign.” and part to “U. kirkiana var. dubia (De Wild.) P. A.Duvign.”. White (1962) also used the name U. benguelensis for glabrous specimens in Zambia. Glabrous specimens were overlooked by

Flora Zambesiaca (Radcliffe-Smith 1996). Recently, U. teusczii was treated as a synonym of

U. kirkiana by Lebrun & Stork (2006) and of U. benguelensis by Figueireido & Smith (2008) and by The Plant List (http://www.theplantlist.org/tpl1.1/record/kew-209563), the World

Checklist of Plant families (http://apps.kew.org/wcsp/) and the Catalogue of Life

(http://www.catalogueoflife.org/col/details/).

U. teusczii is variable for the indumentum of lower leaf surface, which can be lacking or present, consisting of straight whitish hairs. Glabrous and pubescent specimens are sympatric and often coexist in the same population; therefore, the rank of form seems the most appropriate.

1a. Uapaca teusczii Pax f. teusczii

Uapaca munamensis De Wild. (De Wildeman 1932: 206) pro parte, excl. syntype Robyns

1561; De Wildeman (1936: 153); De Wildeman & Staner (1933: 36). Type: D. R. Congo,

Haut-Katanga, Munama, vers 1250 m d'altitude, dans la savane arborée, 8 March 1926,

Robyns 1562 (Lectotype selected by Duvigneaud (1949) BR; second-step lectotype

selected here BR0000008900371!; isolectotypes BR!, K!, P!), synon. nov.

Uapaca dubia De Wild. (De Wildeman 1933b: 307); De Wildeman (1936: 109). Type: D. R.

Congo, Haut-Katanga, route de Pweto à Baudouinville [Kirungu], savane arborée vers

1200 m d'altitude, entre Miduli et Kashika, 25 April 1926, Robyns 2123 (holotype

BR0000008898456!), synon. nov.

Uapaca kirkiana var. dubia (De Wild.) P. A. Duvign. (Duvigneaud 1949: 891), synon. nov.

Uapaca benguelensis var. pedunculata P. A.Duvign. f. glabra P. A. Duvign. (Duvigneaud

1949: 890), nom. illegit.

Uapaca kirkiana sensu Radcliffe-Smith (1996) quoad Angus 2676 (FHO, K!, SRGH), non

Müll. Arg.

RECOGNITION. Leaf with lower surface glabrous (occasionally with sparse remains of short undulate hairs on main veins).

DISTRIBUTION. Angola (uncommon); D. R. Congo (widespread in Katanga); Zambia

(rare). Map 1.

SPECIMENS EXAMINED. ANGOLA. Prov. Bié. Entre Cachinges & Silva Porto, 14 Feb.

1972, Grandvaux Barbosa & Moreno 12328 (BM!). Prov. Malange. 1903, Almeida s.n.

(LISC!); District of Malange, 1903, Gossweiler 981 (BM!, BR!, K!, P!); Teuscz 423 (B, K!).

DEMOCRATIC REPUBLIC OF THE CONGO. Region IV (Kasaï). Kwango, Kahemba,

1000 m, 3 Feb. 1954, Devred 1405 (BR!); Kahemba, 6 April 1955, Devred 1801 (BR!); 11

Kwango, Ingolobo, entre Popokabaka et Munene, 27 April 1948, Duvigneaud 774U1

(BRLU!); 5 km N Kahemba, 30 May 1948, Duvigneaud 950U1 (BRLU!). Region XI

(Upper-Katanga). Kahela, forêt claire, 1 Nov. 1953, Desenfans 4656 (BRLU!); Duvigneaud

K12 (BRLU!); 50 km SE Kilembe, 26 June 1948, Duvigneaud 1086U1 (BRLU!); entre

Elisabethville [Lubumbashi] et Katofio, Duvigneaud 1214U3 (BRLU!; FHO); 12 km S

Elisabethville [Lubumbashi], route de Sakania, 26 July 1948, Duvigneaud 1269U2 & 1269U4

(BRLU!); Colline entre Elisabethville [Lubumbashi] et Jadotville [Likasi], km 30, 31 July

1948, Duvigneaud 1273U2 (BRLU!, FHO); 45 km NW Elisabethville [Lubumbashi] vers

Jadotville [Likasi], 1948, Duvigneaud 1275U1 (BRLU!); 12 km de Biano-gare, route de

Lubudi, 7 Aug. 1948, Duvigneaud 1358U1 (BRLU!); Kundelungu, par Katofio, 18 Aug.

1948, Duvigneaud 1388U2 (BRLU!); Mulando, 75 km W Kasenga, 19 Aug. 1948,

Duvigneaud 1394U2 (BRLU!); Kasenga, dans la vallée de la rivière Lubi, 20 Aug. 1948,

Duvigneaud 1399U2 (BRLU!); Elisabethville [Lubumbashi], Colline de Mukuen au Sud de la ville, Plaine en contrebas, 16 July 1956, Duvigneaud 2022 (BRLU!); Kasompi central, 20 July

1956, Duvigneaud 2061 (BRLU!); Atashyo, 10 km à l’ouest de Mindingi. 21 July 1956,

Duvigneaud 2072 (BRLU!); Kasompi Est, 26 July 1956, Duvigneaud 2123 (BRLU!);

Kayuka, 70 km Est de Kasaji, 13 Aug. 1956, Duvigneaud 2308B (BRLU!); Kisenge-Kapolo,

1956, Duvigneaud & Timperman 2354 (BRLU!); Tantara, April 1957, Duvigneaud 2644

(BRLU!); Mubidi, 30 km à l’est de Mitwaba, 10 Sep. 1956, Duvigneaud & Timperman 2712

(BRLU!); Luishia, 26 April 1957, Duvigneaud 2927 (BRLU!); Région de Swambo, 1 June

1957, Duvigneaud 3343 & 3346 (BRLU!); Entre Jadotville [Likasi] et Kambove, 16 June

1957, Duvigneaud 3560 (BRLU!); NE Mukoro, rivière Katendeshi, 24 June 1957,

Duvigneaud 3668U2 (BRLU!); Kasumbalesa, 3 Dec. 1959, Duvigneaud 4378 (BRLU!); Entre

Mukumbi et Swambo, 28 Dec. 1959, Duvigneaud 4797U (BRLU!); Entre Elisabethville

[Lubumbashi] et Kipushi, forêt claire sur terre rouge sur calcaire de Kakontwe, dans une 12

plaine étendue, 2 Jan. 1960, Duvigneaud 4859U1 (BRLU!); Kamwali, 19 Jan. 1960,

Duvigneaud 5117 (BRLU!); Route Albertville [Kalemie]-Kapona, 9 Aug. 1952, Galoux 246

(BRLU!); 12 km NW Lubumbashi, parcelle expérimentale de forêt claire, 9 Apr. 1958, Gathy

619 (BM!); 12 km NW Elisabethville [Lubumbashi], parcelle expérimentale de forêt claire,

1300 m, 30 April 1958, Gathy 818 (P!); 12 km NW Elisabethville [Lubumbashi], parcelle expérimentale de forêt claire, 7 May 1958, Gathy 874 (BRLU!, K!); env. de Tshisenda, forêt

Kibakabaka, 1700 m, 20 Nov. 1969, Lisowski 760 (BR!); Kundelungu, 1670 m, 4 Nov. 1969,

Lisowski, Malaisse & Symoens 7700 (BR!, LSHI); Kumanua, 1250 m, Malaisse 8626 (BR!);

Ruashi, mine 3, 9 Sep. 1958, Plancke 26/441A & Plancke 26/442 (BRLU!); Munama, 1250 m, 8 March 1926, Robyns 1562 (BR!, K!, P!); Route de Pweto à Baudouinville [Kirungu], entre Miduli et Kashika, 1200 m, 25 April 1926, Robyns 2123 (BR!); 25 km WSW Kolwezi,

1490 m, 3 Oct. 1982, Schaijes 1542 (BR!); 5 km Elisabethville [Lubumbashi], arboretum de l’Etoile, 15 April 1947, Schmitz 635 (BR!); Lubumbashi, arboretum de l’Etoile, Dec. 1947,

Schmitz 1200 (BR!); Keyberg, 29 Nov. 1948, Schmitz 2130 (BR!, BRLU!); Mbaya, 25 km S

Elisabethville [Lubumbashi], 20 Aug. 1952, Symoens 193 (BRLU!); 2 km de la station de

Keyberg, 27 Aug. 1952, Symoens 274 (BRLU!); 3 km de Kifinga, N de la Luzia, 1400 m, 29

Dec. 1958, Symoens 6271 (BRLU!); Tumbwe, entre la Kasompa et la Kilafuifui, 16 March

1961, Symoens 8328 (P!); 12 km S de Sakania, route de Ndola, 1320 m, 17 April 1962,

Symoens 9317 (BR!); 12 km NW Lubumbashi, 5 Dec. 1968, Symoens 12857 (BR!, K!);

Keyberg, INEAC, 3 Feb. 1966, Symoens 12164 (BR!). ZAMBIA. Northern Prov. Misamfu,

6.4 km north of Kasama, 5 April 1961, Angus 2676 (FHO; K; SRGH); Radcliffe-Smith, Pope

& Goyder 5646 (K!); Abercorn [Mbala], 25 May 1936, Burtt 6297 (BM!). Western Prov.

Lukanda, 25 Dec. 1907, Kassner 2141 (K!).

CONSERVATION STATUS. Least Concern (LC). This form is widespread in the northwest of the Zambezian region. 13

VERNACULAR NAMES. D. R. Congo, Kwango: mulengo; mundengudengu masuku

(Bemba), makombio (Kisela), mupopolo (Kitwaba), tokongko (Kilamba); Angola: mumbola

(Chokwe); Zambia: mupangwa.

NOTES. Specimens with only few sparse hairs, intermediate between U. teusczii f. teusczii and f. pilosa, are not rare e.g. Angola, Prov. Lunda Sul: Exell & Mendonça 388 (BM!, COI!,

LISC!); Prov. Uige: Raimundo, Matos & Maia 833 (LISC!); Zambia: Duff 48 (FHO, K!).

A few collections have unusually large leaves (e.g. leaf blade 28 × 20 cm), with up to 14 –

15 secondary veins (e.g. Desenfans 4546, 6399 (BRLU!), Duvigneaud 2920 (BRLU!)); such specimens are always sterile and certainly represent regrowth or otherwise juvenile material.

Robyns 1562, the lectotype of U. munamensis De Wild. selected by Duvigneaud (1949) is represented by two sheets. We choose the specimen with female peduncles and De

Wildeman’s handwriting on the label as the lectotype.

1b. Uapaca teusczii f. pilosa (P. A. Duvign.) Meerts, comb. nov. Type: D. R. Congo, Haut-

Katanga, Kasenga, brousse à Uapaca-Berlinia, dans la vallée de la rivière Lubi, 20 Aug.

1948, Duvigneaud 1399U1 (Lectotype selected here BRLU!).

Uapaca benguelensis var. pedunculata P. A.Duvign. f. pilosa Duvigneaud, Inst. Roy. Colon.

belge, Bull. Séances 20(4): 890 (1949).

Uapaca pilosa Hutch. var. petiolata sensu Radcliffe-Smith (1996) quoad Angus 663 (BM!,

BR!, BRLU!, FHO, K!), Milne-Redhead 2799 (BR!, K!), non P. A. Duvign.

Uapaca katentaniensis De Wild. (De Wildeman 1933a: 58); De Wildeman (1936: 132);

Duvigneaud (1949: 875); Lebrun & Stork (2006: 248). Type: D. R. Congo, Haut-

Katanga, Plateau de Biano, environs de Katentania, Nov. 1912, Homblé 819 (holotype

BR!), synon. nov.

RECOGNITION. Differs from the type form in having straight, whitish to pale yellowish stiff hairs (1 – 1.5 mm long) scattered on the lower leaf surface, often denser on veins; young twigs often more pubescent than in the typical form; petiole occasionally floccose-pubescent.

Fig. 2A.

DISTRIBUTION. Angola (uncommon); D. R. Congo (Kwango and Katanga: very widespread); Zambia (uncommon). Map 1.

SPECIMENS EXAMINED. ANGOLA. Prov. Bié. Camacupa [General Machado], entre

Cuemba e Caqueque, 1400 m, 2 Sept. 1965, Teixeira & al. 8796 (COI!, LISC!); Cuemba, a 17 km do Munhango, 13 Sept. 1965, Mendes dos Santos 1886 (LISC!); Cuemba, na estrada

Munhango-Cuemba, 13 Sept. 1965, Monteiro & Murta 1718 (LISC!); Cuemba, entre os rios

Cuíva e Coemba, 2 Sept. 1965, Mendes dos Santos 1811 (LISC!). Prov. Lunda Norte.

Muriége, 1050 m, 20 Apr. 1937, Exell & Mendonça 816 (BM!, COI!). Prov. Malange.

Cambundi-Catembo [Nova Gaia], picada pela Reserva da Palanca Preta, 1280 m, 25 Oct.

1965, Teixeira et al. 9094 (LISC!). Prov. Uíge. Entre Quibele e Icoca, 850 m, Raimundo,

Matos & Maia 833 (LISC!). DEMOCRATIC REPUBLIC OF THE CONGO. Region III

(Bas-Congo). Kinzau, 27 Dec. 1952, Callens 3900 (BR!). Region IV (Kasaï). Kwango,

Kahemba, 1000 m, 3 Feb. 1954, Devred 1405 (BR!); 5 km N Kahemba, 30 May 1948,

Duvigneaud 950U1 (BRLU!); 115 km N de Panzi, 7 June 1948, Duvigneaud 1029U1

(BRLU!); 33 km N de Feshi, 12 June 1948, Duvigneaud 1057U1 (BRLU!). Region XI

(Upper Katanga). Matuiti, 28 Feb. 1957, Detilleux 596 (BR!); Plateau des Biano, 32 km N de Tenke, 3 Aug. 1948, Duvigneaud 1318U (BRLU!); Kasenga, dans la vallée de la rivière

Lubi, 20 Aug. 1948, Duvigneaud 1399U1 (BRLU!); Entre Mukumbi et Swambo, à 13 km à l'est de Swambo, 24 July 1956, Duvigneaud 2109 (BRLU!); Kakontwe, 24 July 1956,

Duvigneaud 2112B (BRLU!); Kasompi Est, 26 July 1956, Duvigneaud 2123 (BRLU!);

Kasompi Ouest, 30 July 1956, Duvigneaud 2173 (BRLU!); Colline de Kungumarembe, à l'est 15

de Kasompi, 30 July 1956, Duvigneaud 2181 (BRLU!); Kasompi poste, 5 Aug. 1956,

Duvigneaud 2212 (BRLU!); 24 km O. Kolwezi, 10 Aug. 1956, Duvigneaud 2288 (BRLU!);

Kapelula, 7 km à l’est de Kanzenze, 10 Aug. 1956, Duvigneaud 2294 (BRLU!); 20 km à l’est de Kasaji. Territoire Kisenge, 14 Aug. 1956, Duvigneaud 2317 (BRLU!); Dilolo, Dilungu de la Mangoa. 20 Aug. 1956, Duvigneaud 2408 (BRLU!); Katende, 50 km S Dilolo, 23 Aug.

1956, Duvigneaud 2458 (BRLU!); Territoire Kolwezi, 60 km W de Mutshatsha, 28 Aug.

1956, Duvigneaud 2535 (BRLU!); Biano, 06 Sep. 1956, Duvigneaud 2635 (BRLU!); Tantara,

April 1957, Duvigneaud 2644 (BRLU!); Route Elisabethville-Jadotville [Lubumbashi-Likasi], km 45, 24 April 1957, Duvigneaud 2912U (BRLU!); Région Kengere-Tilwizembe, 6 May

1957, Duvigneaud 3069U2 (BRLU!); Entre Kolwezi et Kengere, 7 May 1957, Duvigneaud

3071 (BRLU!); Ferme Servranckx, 11 June 1957, Duvigneaud 3468U (BRLU!); Kamwali, 4

July 1957, Duvigneaud 3928 (BRLU!); Plaine latéritique entre Kolwezi et Nzilo 1, 12 km N de Kolwezi, 14 Dec. 1959, Duvigneaud 4541 (BRLU!); 10 km N de Mitwaba, savane de dégradation au bord de l’escarpement, 16 Jan. 1960, Duvigneaud 5090U (BRLU!); Kamwali,

19 Jan. 1960, Duvigneaud 5117 (BRLU!); Kasekelesa, 4 km N du poste S Kat., 26 Jan. 1960,

Duvigneaud 5246U1 (BRLU!); Plateau de Biano, environs de Katentania, Nov. 1912, Homblé

819 (BR!); Env. Tshisenda, Lisowski 760 (BR!, K!); Musoshi, 80 km SE Lubumbashi, 1240 m, 4 Dec. 1969, Lisowski 12848, 12850 (BR!, POZG!); Kolwezi, près du lac de barrage de la

Musonoi, Symoens 386 (BRLU!). ZAMBIA. sine loco, 1932, Duff 48 (FHO, K!). Copperbelt

Prov.. Ndola, 1959, Aubréville R16 (P!). Luapula Prov. Kawambwa distr., sandy soil at foot of rocky ridge, 64 km [40 miles] S of Kawambwa on road to Ft. Rosebery [Mansa], 29 Oct.

1952, Angus 663 (BM!, BR!, BRLU!, FHO, K!). Northern Prov. On the road Samfya-

Kapalala, 22 km E of Nsonga, 28 Jan. 1963, Symoens 10038 (BR!). North-Western Prov.

Mwinilunga distr., N of Kalene hill, Zambia-Congo border, 24 Sep. 1952, Angus 537 (BM!,

BR!, BRLU!, FHO, K!); R. Matonchi, 15 Oct. 1937, Milne-Redhead 2799 (K!, LISC!); Slope 16

E of R. Matonchi, 16 Oct. 1937, Milne-Redhead 2801 (BR!, K!) & 2810 (BR!, K!). Western

Prov. 16 km [10 miles] W of Kakoma, Angus 568 (BR!, BRLU!, FHO, K!).

HABITAT. Apparently as the type form; also thicket with Uapaca-Philippia on sandy soil; wooded savannah in dambo on whitish sandy-clay soil; open Brachystegia utilis woodland on yellow compact soil; alt. 650 – 1400 m.

CONSERVATION STATUS. Least Concern (LC). This form is widesprerad in the northwest of the Zambezian region.

VERNACULAR NAMES. Angola: catetta ombula, mumbula, nbula.

NOTES. In Flora Zambesiaca Radcliffe-Smith (1996) misinterpreted U. teusczii f. pilosa as being Uapaca pilosa var. petiolata P. A. Duvign. At the fruiting state, the two taxa cannot be confused. At the vegetative state, U. teusczii f. pilosa, especially large-leaved specimens, and

U. pilosa var. petiolata can be more difficult to tell apart. U. teusczii has reddish-brown lower leaf surface in herbarium (U. pilosa var. petiolata: livid to glaucous grey-green), young leaves glutinose and virtually glabrous on upper surface (U. pilosa var. petiolata: young leaves dry, conspicuously hairy on upper surface of veins), and the epidermis of petiole and main vein in mature leaves flaking off in small rectangular silvery scales, which is not observed in U. pilosa.

Uapaca katentaniensis De Wild., a hitherto enigmatic species known from the type specimen only, belongs here. It differs from other specimens in having more pubescent twigs, and a more pubescent female calyx; very similar specimens have been collected in Zambia, e.g. Aubréville R16 (P!).

Duvigneaud (1949) did not cite a type specimen for U. benguelensis var. pedunculata f. pilosa. However, the specimen Duvigneaud 1399U1 (BRLU) bears “typus” handwritten by

Duvigneaud on the label. It is represented by three sheets. The sheet with the most complete material is chosen as the lectotype. 17

2. Uapaca kirkiana Mull. Arg. (Müller Argoviensis 1864a: 517); De Candolle (1866: 491);

Pax (1904: 370); Hutchinson (1912b: 636); Pax & Hoffmann (1922: 302); Delevoy (1929:

469); De Wildeman (1936: 135); Duvigneaud (1949: 891); Coates Palgrave et al. (1957: 180);

White (1962: 207); Radcliffe-Smith (1988: 567; 1993: 612; 1996: 100); van Wyk & van Wyk

(1997: 90); Smith & Allen (2004: 140); Coates Palgrave (2005: 485); Figueiredo & Smith

(2008: 75); Meerts (2016: 238); Meerts & Hasson (2016: 184). Type: Malawi, Southern

Province, Soche Hill, Manganja Country, 8 March 1862, Kirk s.n. (holotype K!).

Shrub or tree up to 10 m high, with a short bole usually < 2 m, evergreen to very briefly deciduous; bark dark grey, fissured, rough, wood reddish. Twigs shortly greyish to ferrugineous tomentose when young, glabrescent, with leaves spaced or more or less grouped towards the end; stipules linear, ca. 5 (– 12) mm long, pubescent, early caducous or, rarely, more or less persistent. Leaves sessile or petiolate; petiole 0 – 4.5 cm, glabrous to thinly pubescent, stiff, with epidermis often flaking off in small rectangular scales giving a silvery appearance; lamina (5 –) 9 – 20 (– 27) × (2 –) 5.5 – 15 (– 19) cm, obovate, obovate- elliptic, elliptic, long attenuate to cuneate or, more rarely, rounded at base, broadly rounded to emarginate at apex, coriaceous; upper surface green to greyish green, dull to more or less shiny, generally pubescent when young, indumentum of thin curly pale fulvous or beige hairs, glabrescent, generally with remains of hairs on veins, lepidote; lower surface lepidote, with indumentum whitish to beige or pale fulvous, browning with age, consisting of thin curly intermingled hairs, forming a thin continuous layer when very young, generally becoming sparser and patchier with age and then concentrated along veins; secondary veins 8 – 21 (–

25) pairs, anatomosing in arches, impressed on upper surface, prominent on lower surface. 18

Inflorescences borne on current year’s twigs amongst the leaves or on defoliated previous year’s twigs below the leaves. Male inflorescence globose to ovoid or obovoid before anthesis, whitish to yellow, solitary to 2 – 9 fasciculate; peduncle 1 – 20 mm, glabrous, lepidote or pubescent, with bracteoles; 4 – 7 bracts 5 – 11 × 3 – 5 mm, broadly elliptic, yellowish green, the outer pubescent without, the inner glabrous; male flowers in a globose head, calyx appressed-pubescent, 5 stamens exceeding calyx, filament 2 – 2.5 mm, anther ca.

0.7 – 1 mm, connective glabrous or, much more rarely, with a few cilia; pistillode dilated at top, pubescent. Female inflorescence solitary, peduncle 1 – 8 mm long, glabrous, pubescent or lepidote; calyx cupular, 1 – 2 mm long, 5–8-lobed, tube glabrous or, more rarely, with very few hairs, lobes glabrous or with curly hairs especially at margin, generally glabrous in fruit; ovary densely whitish to brownish crisped pubescent, styles (3–) 4, reflexed and almost hiding young ovary, multifid, adaxially pubescent. Fruit on a peduncle 2 – 10 (– 15) mm long, 1.5 –

5 mm thick, globose, (2.0 –) 2.5 – 4 cm diam., pericarp 1 – 3 mm thick, 3 – 4 (– 5) pyrenes,

15 – 20 × 11 – 14 mm ovoid to oblong-ellipsoid, narrowed at tip, cordate to tridentate at base, with the median crest exceeding or not the lateral lobes at base, often forming a beak at tip, sharp in transversal section, lateral ridges deep for most of their length, lateral lobes smooth; adaxial side straight. Seeds 1.5 × 0.8 × 0.5 cm, resembling the pyrenes in shape, brown.

NOTES. Three combinations of traits can be distinguished within U. kirkiana, which have long been recognised at species level. However, intermediate populations occur and the varietal rank seems therefore more appropriate. Inflorescence architecture is variable in U. kirkiana, with a clear geographic pattern. Collections from the western part of the range

(mostly Angola) have the inflorescence near the apex of branches, amongst the leaves; in this case, male inflorescences are solitary, not fasciculate, with a short peduncle (“benguelensis- type” inflorescence). Collections from the eastern part of the range (mostly D. R. Congo, 19

Zambia and further east) have inflorescences on previous year’s growth below the leaves; in this case, male inflorescences are 2 – 9 fasciculate and have a long peduncle (“kirkiana-type” inflorescence). The male inflorescence is globose, with a stipitate flower head in var. kirkiana and ovoid to ellipsoidal with a sessile flower head in var. benguelensis. The “benguelensis- type” inflorescence is generally correlated to a lower number of secondary veins (≤ 13 (–

14)), and the “kirkiana-type” to a larger number of secondary veins (> 12). The third morph

(“gossweileri”) is recognised by sessile leaves with long attenuate lamina, grouped in rosettes, on corky twigs. This variety is variable for inflorescence architecture.

2a. Uapaca kirkiana var. kirkiana

Uapaca homblei De Wild. (De Wildeman 1926: 309; 1936: 128); Delevoy (1929: 468). Type:

D. R. Congo, Haut-Katanga, Elisabethville [Lubumbashi], Feb. 1902, Homblé 131

(Lectotype selected here BR0000008897831!; isolectotypes BR!).

Uapaca albida De Wild. (De Wildeman 1933a: 57; 1936: 91). Type: D. R. Congo, Haut-

Katanga, Collines pierreuses, versant d’Albertville [Kalemie], 10 Sept. 1931, Delevoy

240 (Lectotype selected here BR0000008898302!; isolectotype BR!).

Uapaca neomasuku De Wild. (De Wildeman 1932: 208; 1936: 154). Type: D. R. Congo,

Haut-Katanga, sine dat. (1918?), sine rec. (holotype BR0000008895554!).

Uapaca kirkiana var. sessilifolia P. A. Duvign. (Duvigneaud 1949: 891) Type as U.

neomasuku De Wild.

RECOGNITION. Current year’s twig 4 – 8 mm in diam.; previous year’s twig 5 – 11 mm in diam., greyish, neither markedly suberised nor with many prominent leaf scars; leaves not in rosettes, more or less spaced on current year’s twigs, sessile or petiolate, petiole 0 – 35 mm; lamina (7 –) 9 – 20 (– 25) × (5 –) 6 – 19 cm, obovate, obovate-elliptic, more or less undulate- 20

sinuose, often saddle-shaped, base cuneate to attenuate, not forming a winged pseudopetiole, broadly rounded to emarginate at apex, secondary veins in (12 –) 13 – 25 pairs; inflorescences and fruits mostly below the leaves on previous year’s growth; male inflorescence globose before anthesis, 2 – 9 fasciculate, peduncle 4 – 20 mm; bracts 4 – 7 mm long, 2 – 6 mm wide, broadly ovate to almost round; male flowers in a stipitate head; fruit peduncle 5 – 15 mm long, 1.5 – 3 (– 3.5) mm thick, generally > 3 times longer than thick; pyrenes with median crest generally narrowly acute over at least half of its length. Fig. 2B; Fig. 3B, D, F, H.

DISTRIBUTION: Angola, Burundi, D. R. Congo, Malawi, Mozambique, Tanzania, Zambia,

Zimbabwe. Map 2.

SPECIMENS EXAMINED. ANGOLA. Prov. Benguela: Ganda, Babaera, a caminho da

Fazenda Vitória, 15 Jan. 1972, Menezes 4044 (LISC055227!). Prov. Bié: Andulo, picada do

Cassombo a 4 km do desvio, ca. 1400 m, 26 Nov. 1965, Teixeira et al. 9586 (LISC055370!).

Prov. Huíla: Lubango, Serra da Chella, env. Sá da Bandeira, sol rocailleux siliceux, 1850 –

2000 m, Aug. 1937, Humbert 16209 (P!).

DEMOCRATIC REPUBLIC OF THE CONGO. Region XI (Upper Katanga). Env. d’Elisabethville [Lubumbashi], July – Aug. 1919, Burtt-Davy 17884 (BM!); Territ. Mitwaba,

Luishi, 1700 m, Coget 228 (BR!); Collines pierreuses, versant d’Albertville [Kalemie], 10

Sept. 1931, Delevoy 240 (BR!); Territ. Manono, Kapona, 16 July 1957, Devred 3608 (WAG!,

BR!); Route Pweto-Baudouinville [Kirungu], en bordure des Marungu, 20 July 1957, Devred

3696 (BR!, WAG!); 64 km NE Elisabethville [Lubumbashi], 16 July 1948, Duvigneaud

1214U1 (BRLU!, FHO); 12 km S Elisabethville [Lubumbashi], route de Sakania, 26 July

1948, Duvigneaud 1269U1 (BRLU!); 35 km NW Elisabethville [Lubumbashi], brousse dense

à Marquesia-Uapaca, en contrebas d’une colline, 31 July 1948, Duvigneaud 1274U1

(BRLU!); 5 km W Jadotville [Likasi], colline gréseuse, 2 Aug. 1948, Duvigneaud 1301U1

(BRLU!); Plateau des Biano, 32 km N Tenke, 3 Aug. 1948, Duvigneaud 1318U3 (BRLU!); 21

autour de l’hôtel des Bianos, 6 Aug. 1948, Duvigneaud 1346U1 (BRLU!); N de Biano, poteau de Lubudi, 7 Aug. 1948, Duvigneaud 1353U (BRLU!); Katofio, escarpement des

Kundelungu, 18 Aug. 1948, Duvigneaud 1380U (BRLU!); Entre Kalundu et Dilolo, rivière

Luvwegi, Duvigneaud 3267U1 (BRLU!); 6 km E Swambo, 1 June 1957, Duvigneaud 3341U1

(BRLU!); Mitonte, entre Swambo et Mindingi, Duvigneaud 3397U1 (BRLU!); Chabara, 12

June 1957, Duvigneaud 3485U (BRLU!); Tantara, 23 Dec. 1959, Duvigneaud 4757U

(BRLU!); 12 km NW Elisabethville [Lubumbashi], parcelle expérimentale de forêt claire,

1300 m, 2 April 1958, Gathy 545 (BM!; P!); Elisabethville [Lubumbashi], Feb. 1902, Homblé

131 (BR!); Env. d’Elisabethville [Lubumbashi], April – May 1934, Humbert 16051 (P!) &

Quarré 4041 (BR!); Lubumbashi, Kasapa, 1230 m, 24 Dec. 1970, Lisowski 762 (BR!,

WAG!); Route Albertville [Kalemie]-Kabambare, km. 33, 8 July 1932, Luxen 148 (BR!);

Mine de Lukuni, pied du déblai, March 1939, Quarré 5320 (BR!); Plateau de la Kimilolo, 23

Feb. 1927, Ritschard 1377 (BR!); Vallée de la Lubumbashi, 9 March 1928, Ritschard 1614

(BR!); Keyberg, 1 March 1946, Schmitz 1415 (BR!); Elisabethville [Lubumbashi], arboretum de l’Etoile, April 1948, Schmitz 1693 (BR!, WAG!); Kipopo, 19 Nov. 1960, Schmitz 7238

(BR!); 2 km SW Albertville [Kalemie], 1 Oct. 1952, Symoens 881 (BRLU!); Plateau des

Kibara, 10 km Kasongami, W de Nematshila, 1520 m, 30 Dec. 1958, Symoens 6370 (BRLU!).

ZAMBIA. Copperbelt Prov. Ndola, Nov. 1959, Aubréville R21 (P!). Northern Prov. Mpika distr., Mano Camp, just outside and N of Luangwa, 1110 m, 13 May 1994, Harder, Schmidt,

Nkhoma 3125 (MO, WAG!); Abercorn [Mbala] distr., 1850 m [6000 ft], 17 Apr. 1936, Burtt

6310 (BM!, BR!). North-Western Prov. Distr. Mwinilunga, 16 km [10 miles] W of Kakoma,

28 Sept. 1952, Angus 569 (BR!, BRLU!, FHO); Slope N of Matonchi, 16 Oct. 1937, Milne-

Redhead 2805 (K!, BR!); Distr. Ndola, Mindola F.R. near Nkana, Brenan & Greenway 7960

(BR!); 24 km de Mpumba, à l’ouest de la route Mkushi–Mpika, 19 Dec. 1961, Symoens 9003

(P!); Solwezi District. Mulilanshima Hill along Solowezi-Mwinilunga road. 16 Feb. 1995, 22

Zimba 531 (MO, WAG!). Southern Prov. Mazabuka, Siamambo forest reserve, near Choma,

29 July 1952, Angus 79 (FHO, BRLU!, BR!); 43.8 km from Choma along Choma-Namwala

Road, just adjacent to Ngonga River, Simubara-Marundu Hills, 1100 m, 3 Aug. 1997, Zimba,

Harder, Schmidt & Luwiika 1088 (WAG!).

HABITAT. Shrub savannah, tree savannah, miombo woodlands, usually on shallow lateritic soil or on rocky slopes, dambo margins; more or less resistant to fire and persisting in degraded savannah; alt. 800 – 1850 m.

CONSERVATION STATUS. Least Concern (LC). This variety is widespread in most of the

Zambezian region.

VERNACULAR NAMES. Angola: o’lumbula (Umbundu). R. D. Congo: masuku, musuku.

NOTE. Petiole length varies between 0 and 35 mm; Duvigneaud (1949) recognised the sessile-leaved specimens as var. sessilifolia P. A. Duvign., but this has no taxonomic value.

Some specimens growing in harsh conditions show some of the diagnostic traits of var. gossweileri (e.g. Quarré 5320 (BR!).

The four collections from Angola are more or less intermediate with var. gossweileri. See also discussion under var. gossweileri.

2b. Uapaca kirkiana var. gossweileri (Hutch.) Meerts comb. et stat. nov. Type: Angola,

Province Lunda Sul, on the rocky hills at Gimbundo Jamaiambe [Mona Quimbundo], valley of Rio Kutato [Cutato], May 1906, Gossweiler 3802 (Lectotype selected here:

BM001209664!; isolectotypes COI00067278!, K000406461!, LISC055222!).

Uapaca gossweileri Hutchinson, Bull. Misc. Inform. Kew 1912: 101 (1912a); Hutchinson

(1912b: 635); Pax & Hoffmann (1922: 301); De Wildeman (1936: 118); Duvigneaud

(1949: 891); Devred (1957: 426); Lebrun & Stork (2006: 246); Figueiredo & Smith

(2008: 75); Meerts (2016: 238). 23

Uapaca benguelensis Müll. Arg. (Müller Argoviensis 1864: 332) ex parte; De Candolle

(1866: 491) ex parte; Hiern (1900: 963) ex parte

RECOGNITION

Crooked shrub 2 – 3 m to small tree 8 – 10 m high. Current year’s twigs 7 – 12 mm thick in the leafy portion, previous year‘s twigs much thicker, (8 –) 10 – 25 mm in diam., usually with strongly suberised cork marked with numerous prominent leaf scars; stipules filiform, hirsute, up to 12 mm long, often more or less persistent. Leaves tightly clustered in rosettes at twig apex (occasionally more widely spaced on regrowth), sessile (rarely with petiole 0 – 5 mm), lamina obovate to broadly obovate-spatulate, long attenuate in the lower half into a winged pseudopetiole with undulate margin, truncate, rounded to broadly obtuse at apex (10 –) 14 –

24 (– 27) x (5 –) 7 – 15 (– 16.5) cm, 12 – 21 pairs of secondary veins, generally anastomosing in arches; tertiary veins forming a scalariform pattern; upper surface often with copious remains of indumentum on veins. Inflorescence on previous year’s defoliated twigs or (in

Angola) amongst the leaves; male inflorescence 2 – 8 fascicled or (in Angola) solitary, with peduncle (1 –) 3 – 20 mm long; male bract length 4 – 11 mm; female flowers with calyx ca. 2 mm long, markedly lobed, with curly hairs at margin. Fruiting peduncle 2 – 5 (– 10) mm long,

2.5 – 3.5 (– 5) mm thick. Fruit typically with slightly conical apex; pericarp 2 – 3 mm thick; pyrenes 4, with median ridge typically acute in cross-section, occasionally obtuse to rounded.

DISTRIBUTION. Angola, D. R. Congo. Map. 2.

SPECIMENS EXAMINED. ANGOLA. Prov. Bié. Kuito [Silva Porto], Centro de Estudos da Ceilunga, 6 Feb. 1970, Lopes Gomes & da Silva 2803 (COI!, LISC!); Près de Silva Porto,

7 March 1974, 1640 m, Dechamps, Murta & da Silva 1381 (BR!, LISC!); Entre Cachingues e o cruzamento para Silva Porto parcela nº3, 14 Feb. 1972, Grandvaux Barbosa & Moreno

12338 (COI!, LISC!); Na estrada Santar-Luando, 14 April 1966, Monteiro & Murta 1845 24

(COI!, LISC!). Prov. Cuanza Sul. Quibala, entre Gungo e Cariango, andados 15 km, 1340 m,

17 Jan.1972 Raimundo, Matos & Figueira 980 (LISC!); Waku Kungo, a 17 km de Santa

Comba [Waku Kungo] para a Quibala, 27 Feb. 1965, Mendes dos Santos 1293 (COI!, LISC!);

Entre a estrada asfaltada Lussusso-Muquitixe e a antiga estrada Calulo-Quibala andados 4 km, ca. 1000 m, 27 Jan. 1971, Raimundo, Matos & Figueira 650 (LISC!, BR!). Prov. Huambo.

Huambo, Station IIAA Nova Lisboa, 1700 m, 14 Feb. 1974, Dechamps, Murta & da Silva

1029 (BR!); Nova Lisboa [Huambo], perímetro Florestal de Sacaála, 1700 m, 18 Feb. 1962,

Murta 114 (COI!, LISC!). Prov. Huíla. In mixed woods between Kuanxa & Fte [= Forte]

Dom Affonso at Kului [Colui], Jan 1907, Gossweiler “2937” (sphalm. for 2936)

(Lectoparatype BM!), Gossweiler 2936 (K!); Near the river Kutoto [Cutato], Cutato

Nganguela, Nov. 1906, Gossweiler 3206 (Lectoparatype BM!); Near Kaconda [Caconda],

Feb. 1906, Gossweiler 3364 (Lectoparatypes BM!, COI!, K!, LISC!); Between Lopollo and the river Monino, Feb. 1860, Welwitsch 453 (male) (Lectoparatype BM!, LISU! (2 sheets));

Near Lopollo, Welwitsch 455 (male) (Lectoparatype K!, non BM, P); Arredores da Missão

Católica de Monhino, 8 Dec. 1955, Mendes 1024 (LISC!); Kuvango [Ganguelas], Andados 15 km pelo C.F., de Vila Artur de Paiva para Cutato, 1450 m, 21 Jan. 1960, Mendes 2134

(LISC!); Kuvango [Ganguelas], Vila Artur de Paiva, margens do Cubango, 1450 m, 5 Jan.

1960, Mendes 1984 (LISC!). Prov. Kwando-Kubango. Rio Cutato [Kutato], Febr. 1906,

Gossweiler 3206 (Lectoparatype BM!). Prov. Lunda Sul. On the rocky hills at Gimbundo

Jamaiambe [Mona Quimbundo], valley of Rio Kutato [Cutato], 6 May 1906, Gossweiler 3802

(BM!, COI!, K!, LISC!); Minungo, Alto Chicapa, na Picada das Grutas do Musseque, 11 May

1963, Martins 71 (LISC!, MO, P!). Prov. Malanje. Quizanga near Pança Mencha on the road to Quepacaca, 16 June 1903, Gossweiler 980 (Lectoparatypes BM!, P!); Mucari, Caculama, entre Caculama e Caxinga, próximo da sanzala do soba Quibugo, 5 June 1970, Grandvaux

Barbosa, Moreno & Sousa 11922 (LISC!); Kela [Quela], 8 July 1927, Carrisso & Mendonça 25

85 (COI!); Reserva da Palanca Preta, Mulundo, 24 Aug. 1965, Correia 3170 (LISC!);

Planalto de Malange, Kela, Proximum flumen Luando-Lui, 1200 m, 16 Jan. 1931, male pl.,

Gossweiler 9550 (BM!, COI!, K!, LISC!); Nova Gaia [Cambundi-Catembo], picada pela

Reserva da Palanca Preta, 1280 m, 25 Oct. 1965, Teixeira et al. 9096 (LISC!).

DEMOCRATIC REPUBLIC OF THE CONGO. Region IV (Kasaï). Nto Muniungu, 14

April 1953, Callens 1152 (BR!); Kunge-Bansa, vers Dinga, savane, 19 Feb. 1950, Callens

2441 (BR!); Kunga Bansa, 15 Feb. 1930, Callens 2534 (BM!); Kasinsi, Isangi, Callens 2561

(BR!); Kambangu, 10 Feb. 1952, Callens 3216 (BR!, K!); Région de Mawanga, 30 April

1953, Callens 4060 (BR!); Muniungu, Jan. 1955, Callens 4807; Kahemba, 29 April 1955,

Devred 1914 (BR!, WAG!); Panzi, 16 June 1955, Devred 1984 (BR!); Kahemba, 1248 m, 1

Sept. 1949, Dubois 1483 (BR!, WAG!); 25 km E Popobakata, 27 April 1948, Duvigneaud

772U1 (BRLU!); Muniungu, 25 km NE Popobakata, 6 June 1948, Duvigneaud 839U2

(BRLU!); Muniungu, 6 May 1948, Duvigneaud 840U1 (BRLU!); 115 km N Panzi, 7 June

1948, Duvigneaud 1029U2 (BRLU!, FHO); Entre Feshi et la Luié, 11 June 1948, Duvigneaud

1052U1 (BRLU!); Entre Ndinga et Mpandi, 450 m, Germain 2233 (BR!); Entre Kwango-

Wamba, village Pandi, 25 km E Dinga, 700 m, 11 Aug. 1944, Germain 2755 (BR!);

Kahemba, 22 Jan. 1942, Jernander 74 (BR!). Region XI (Upper Katanga). Kayuka, 70 km à l’est de Kasaji, 13 Aug. 1956, Duvigneaud 2308A (BRLU!); Kisenge, route de Divuma,

Duvigneaud 2329 (BRLU!); Dilolo, dilungu de la Mangoa, 20 Aug. 1956, Duvigneaud 2408

(BRLU!); Dilolo, Tshisenge, Duvigneaud 3273U (BRLU!); Kasekelesa, chantier 1, 25 Jan.

1960, Duvigneaud 5239U1-U6 (BRLU!); Kasekelesa, entre les chantiers 2 et 3, 25 Jan. 1960,

Duvigneaud 5242U3 (BRLU!).

HABITAT. Savannahs, shrub savannahs, miombo woodlands often at margin of dambos. In

Kwango (SW of D. R. Congo), U. kirkiana var. gossweileri occurs in Zambezian tropophilous forests (“mikondo”) (“Association à Julbernardia paniculata et Brachystegia longifolia, sous- 26

association à Cryptosepalum pseudotaxus et Uapaca gossweileri”), often forming a fringe at the margin of dambos on more or less hydromorphic soil (Devred et al. 1958). Also found forming a 5 – 10 m wide fringe at the contact zone between frequently burnt savannah vegetation and semi-evergreen woodlands (“mabwati”) dominated by Berlinia giorgii,

Brachystegia div. sp., (Duvigneaud 1949; Germain 1949) (“Association à Marquesia macroura et Uapaca nitida, sous-association à Brachystegia wangermeeana et Brachystegia spiciformis, variante à Uapaca gossweileri” (Devred et al. 1958)). In Upper Katanga, occasionally on metalliferous soil (manganese); alt. 450 – 1700 m.

CONSERVATION STATUS. Least Concern (LC). The variety has a large distribution range and is known from many collections.

VERNACULAR NAMES. Angola: ambula, bula, lombula, lumbula, mumbula, omumbula.

D. R. Congo: mbula mbula.

USES. Edible fruit.

NOTES. Var. gossweileri is the most difficult taxon to circumscribe in the U. kirkiana complex. In its most typical form, with thick corky twigs, stunted growth habit and thickened fruit wall, var. gossweileri is a striking plant. Such forms are mostly restricted to SW D. R.

Congo (Kwango) and Angola, where they have a distinctive ecology, occurring as a fringe between miombo woodland and savannah often in infertile soil. However, many collections from Angola are less extreme, for twig characters in particular.

Var. gossweileri is polymorphic for male inflorescence position and structure. While many collections have inflorescence traits similar to var. kirkiana (especially in Kwango), some specimens from Angola have inflorescence traits similar to var. benguelensis, i.e. with male inflorescences grouped near the apex of twigs, amongst the leaves, with solitary peduncle <

10 mm long (e.g. Murta 114 (LISC!), Gossweiler 3802 (LISC!) ). Such geographic pattern of variation possibly points to introgression of var. gossweileri with var. benguelensis in their 27

area of sympatry. Alternatively, var. gossweileri could represent a polyphyletic assemblage, comprising forms having evolved independently within both var. kirkiana and var. benguelensis, possibly in response to similar selective pressure (fire?). This latter hypothesis is supported by the sporadic occurrence of plants showing some of the characteristic traits of var. gossweileri in other parts of the distribution range of var. kirkiana, e.g. in Zambia,

Burundi and Tanzania (e.g. Radcliffe-Smith 1988: 568). Resolving phylogenetic relationships requires molecular markers.

Lectotypification of Uapaca kirkiana var. gossweileri

Hutchinson (1912a) cited 7 syntypes for Uapaca gossweileri, i.e. Gossweiler 980, 2937, 3206,

3364, 3802, Welwitsch 453 (pro parte, male specimen), Welwitsch 455 (K, non BM). The handwritten number “Gossweiler 2937” on the specimen BM001209666 is probably a lapsus calami for Gossweiler 2936 since Gossweiler 2937 (BM000884457!) is the type specimen of

Crossandra angolensis S.Moore (Acanthaceae). The duplicate in K bears the correct number

Gossweiler 2936. The specimens distributed to different herbaria under Welwitsch 453 and

Welwitsch 455 represent a heterogeneous assemblage, because Welwitsch lumped together under one number all the collections which he considered to be conspecific (Albuquerque et al. 2009). Therefore it is preferable not to consider that material for selecting a lectotype. The other syntypes all show the diagnostic traits of var. gossweileri. Gossweiler 3802

(BM001209664) is selected as the lectotype because it has been widely distributed.

1c. Uapaca kirkiana var. benguelensis (Müll. Arg.) Meerts comb. et stat. nov. Type:

Angola, Province Huila, inter Lopollo and flumen Monino, Welwitsch 453 (female plant only)

(Lectotype selected here G00319292!; isolectotype COI00067278!). 28

Uapaca benguelensis Müller Argoviensis, J. Bot. 1: 332 (1864b); De Candolle (1866: 491),

ex parte; Hiern (1900: 963) ex parte; Pax (1904: 371), ex parte; Engler (1910: 612);

Hutchinson (1912b: 637); Pax et Hoffmann (1922: 303); De Wildeman (1936: 95);

Duvigneaud (1949: 890); Figueiredo & Smith (2008: 75).

Uapaca kirkiana sensu Radcliffe-Smith (1996) quoad Loveridge 791 (BR!, K; LISC!; SRGH) non Müll. Arg.

RECOGNITION

Tree up to 12 m. Twigs ca. 5 – 8 mm in diam. in the leafy part, pubescent, swollen at insertion of fruiting peduncles; previous year’s twigs neither corky nor with prominent leaf scars.

Leaves more or less spaced on current year’s twigs, not forming a rosette, always petiolate, petiole 10 – 45 mm (typically 15 – 20% of leaf blade length), generally pubescent, leaf blade

5 – 17 × 2 – 11.5 cm, obovate to broadly elliptic, rounded to slightly emarginate, obtusely cuneate to rounded at base, more rarely acute, with 8 – 13 (– 14) pairs of veins, generally densely lepidote on both surfaces, lower surface greyish to fulvous crisped pubescent, upper surface often remaining more or less puberulous (more so than var. kirkiana), veins persistently pubescent. Inflorescence borne near the apex of twigs or amongst the leaves or, more rarely, just below the leaves; male peduncle solitary, 1 – 10 mm long, generally puberulous or lepidote; male inflorescence ovoid to obovoid before anthesis, markedly longer than wide, bracts elliptic 7 – 11 mm long, the outermost ones sparsely pilose without; male flower head sessile; calyx of female flower with sparse curly hairs, especially near margin.

Peduncle of fruit 2 – 5 (– 8) mm long, 3 – 5 mm diam., generally less than two times longer than thick; fruit wall thinner than in the type variety (≤ 1 mm), shrinking or tearing in herbarium; pyrenes with median crest obtuse or blunt over at least half of its length, sometimes acute in upper third. 29

DISTRIBUTION. Angola; D. R. Congo (rare); Zambia (rare). Map 2.

SPECIMENS EXAMINED. ANGOLA. Prov. Bié. Entre o cruzamento da estrada de Silva

Porto-Serpa Pinto e Chinguar, 11 April 1971, Grandvaux Barbosa 12177 (FHO, LISC!);

Andulo, estrada Nharea Lubia a 30 km de Nharea, 24 Nov. 1965, Teixeira et al. 9532 (LISC!); a 5 km de Cachingues para Chitembo, 7 April 1971, Grandvaux Barbosa 12112 (COI!, FHO,

LISC!); Chitembo, Cachingues, próximo de Cachingues, 5 Sept. 1971, Grandvaux Barbosa &

Moreno 12234 (FHO, LISC!); Noharea, 20 Oct. 1966, Monteiro & Murta 2039 (LISC!).

Prov. Huambo. Huambo, Bailundo, proximo do Bimbe, na estrada para a Fazenda Aurora andados 8 km, 1800 m, 25 Jan 1973, Raimundo, Matos & Figueira 1268 (BM!, BR!, LISC!);

Huambo [Nova Lisboa], entre Chinguar e Silva Porto a 29 km de Chinguar, 13 Nov. 1965,

Grandvaux Barbosa 11018 (LISC!). Prov. Huila. Inter Lopollo et flumen Munhino, Feb.

1860, Welwitsch 453 (GDC, COI!); Inter Lopollo et flumen Monino, March 1860, Welwitsch

454 (BM!, LISU!); Inter Lopolo et lacum magnum de Turantula, 1500 – 1650 m [5000 – 5500 ped.], March 1860, Welwitsch 455 (female plant) (BM!, LISU!; P!); sine loco, Antunes vel De

Kindt s.n. (LISC!); Lubango, Lopolo, 1790 m, April 1899, De Kindt 1011 (LISC!); Lubango,

Munhino [Monyino], 1790 m, 5 April 1900, De Kindt 3034 (K!, LISC!, P!); Lubango,

Munhino, arredores da Missão Católica, 2 Nov. 1955, na encosta oriental do planalto da

Humpata, Mendes dos Santos 24 (LISC!); [Ganguelas] Kuvango, andados 15 km pelo C.F., de

Vila Artur de Paiva para Cutato, 1450 m, 21 Jan. 1960, Mendes 2135 (LISC!); entre

Tchivinuiro e Banja, 20 Dec. 1961, Grandvaux Barbosa 9707 (BR!, LISC!). Prov. Kwando-

Kubango: Menongue, nas margens do rio Cutato, 24 Feb. 1906, Gossweiler 3205 (BM!, BR!,

COI!, K!, LISC!); Menongue, vale do rio Cutato, no caminho para Vila da Ponte, 2 May

1906, Gossweiler 3801 (BM!, BR!, COI!, K!, LISC!). Prov. Lunda Sul: Caculo, Alto chicapa, Rio Colundo, Hiemilignosa, July 1954, Barros Machado 385 (LISC!).

DEMOCRATIC REPUBLIC OF THE CONGO. Region IV (Kasaï): Kwango, Dinga, 31 30

Aug. 1955, Devred 2524 (BR!); 40 km SW Kenge vers Popobakaba, 9 May 1948,

Duvigneaud 854U1 (BRLU!); Entre Kwango-Wamba, village Duripangi, 31 July 1944,

Germain 2527 (BR!, K!). Region XI (Upper Katanga). Lualaba, 9 Aug. 1956, Duvigneaud

& Timperman 2261 (BRLU!); Kolwezi, plateau de la Manika, 10 Aug. 1956, Duvigneaud

2267 (BRLU!); Kolwezi, Plateau de la Manika, au N du plateau, 29 Aug. 1956, Duvigneaud

& Timperman 2545 (BRLU!). ZAMBIA. North-Western Province. Dist. Mwinilunga, 27 km [17 miles] S Mwinilunga on Kabompo road, 6 June 1963, Loveridge 791 (BR!, K!,

SRGH); same loc., same date, Edwards 650 (BR!, COI!, SRGH).

HABITAT. Miombo woodland, wooded savannah, shrub savannah and thickets, often at forest edge; alt. 800 – 1800 m.

CONSERVATION STATUS. Least Concern (LC). This variety is widespread in the northwest of the Zambezian region.

VERNACULAR NAMES. Angola: catembula, mombula, mumbula, nombula, omumbula;

Kwango (R.D. Congo): mubula bula, mbula (Kiyaka).

NOTES. Uapaca kirkiana var. benguelensis and U. kirkiana var. kirkiana are quite distinct in reproductive traits, even though both are variable. Due to overlap in vegetative traits sterile specimens with 12 – 14 pairs of secondary veins can be difficult to tell apart. The two varieties have distinct, though overlapping, distribution range. Var. benguelensis is widespread in Angola and also occurs, though much more sparsely, in SW D. R. Congo and

NW Zambia. Var. kirkiana is very widespread in Katanga, Zambia and the rest of the

Zambezian region being much sparser and less typical in Angola (see note under var. gossweileri).

Some specimens are more or less intermediate between U. teusczii f. teusczii and U. kirkiana var. benguelensis and could result from introgression (e.g. Angola, Prov. Bié,

Teixeira et al. 9696 (LISC!); D. R. Congo, Kwango, Duvigneaud 839U1 (BRLU!). 31

Collections from Kwango, at the northern margin of the distribution range, are somewhat untypical in having reduced indumentum and larger leaves (e.g. Devred 2524 (BR!)).

Phiri (2005: 51) erroneously cites U. benguelensis Müll. Arg. as a synonym of U. robynsii

De Wild.

Lectoypification of Uapaca benguelensis Müll. Arg.

Müller Argoviensis (1864b) indicated Welwitsch 453 as the type of Uapaca benguelensis.

Material with that collection number has been found in BM, COI, G-DC and LISU.

Hutchinson (1912a) rightly pointed out that the material distributed with the number

“Welwitsch 453” was heterogeneous, comprising a taxon with sessile leaves, that he described as a new species (U. gossweileri Hutch.), and a taxon with petiolate leaves. Hutchinson thus implicitly restricted U. benguelensis Müll. Arg. to the petiolate morph, but he did not designate a lectotype. Since U. benguelensis Müll. Arg. was published before Welwitsch’s collection was split, the lectotypification should consider all the material, not only the material in BM (Albuquerque et al. 2009). The protologue of U. benguelensis contains a description of ovary, while the sessile-leaved specimens (i.e. U. gossweileri) are all male or sterile. Therefore, it is appropriate to choose a female specimen as the lectotype. The material in BM (BM000911092) comprises one sheet with two differently looking twigs, i.e. a sterile twig on the left with petiolate leaves and a male twig on the right with sessile leaves. The two specimens are either from different trees or, if from the same tree, could be a hybrid. In any case, it is deemed inappropriate to select such spurious material as the lectotype. The material in LISU consists of two sheets [LISU223847 & LISU223850], both with large sessile leaves typical of var. gossweileri. The material in COI comprises one sheet with a sterile twig and a female twig, both with petiolate leaves, obviously corresponding to var. benguelensis. The material in G-DC (G00319292) comprises one female twig, clearly belonging in var. 32

benguelensis. We choose Welwitsch 453 [G00319292 (G-DC)] as the lectotype. Notice that this specimen is labelled as a holotype in G-DC, in error. The female specimen in COI is an isolectotype.

Unplaced

Uapaca kirkiana var. kwangoensis P. A.Duvign. (Duvigneaud 1949: 891). Type: D. R.

Congo, Kwango, Duvigneaud 822 (BRLU! (10 sheets)).

The type material of Uapaca kirkiana var. kwangoensis P. A. Duvign is unusual in combining very short, sparse curly hairs on lower surface of leaf and long attenuate leaf base; it could be interpreted as a hybrid between U. teusczii and U. kirkiana var. gossweileri. The ovary with sparse curly hairs is consistent with that hypothesis.

Acknowledgements

We are grateful to Tariq Stevart (Missouri Botanical Garden) for assistance in the production of distribution maps and to Guy Josens (Université Libre de Bruxelles) for assistance in taking microphotographs. Michel Hasson (Biodiversité au Katanga, Lubumbashi) is gratefully acknowledged for allowing us to use his photographs. We are grateful to the curators of the following herbaria for welcoming us in their collection: BM, BR, COI, K, LISC, LISU, P,

POZG. JF is a research fellow of the FRIA-FNRS (Fonds pour la Formation à la Recherche dans l'Industrie et dans l'Agriculture, Belgium).

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Table 1. Diagnostic characters separating Uapaca teusczii and the three varieties of Uapaca kirkiana.

Character U. teusczii U. kirkiana U. kirkiana U. kirkiana

var. kirkiana var. benguelensis var. gossweileri

Position of Amongst the leaves or 5 – 30 cm below the Amongst the leaves or Variable: below the

inflorescences just below the leaves leaves, on previous just below the leaves leaves or, more rarely,

year’s growth amongst the leaves

Length of male peduncle 2 – 10 (– 20) mm 4 – 20 mm 1 – 10 mm 1 – 20 mm

Number of male Solitary (1 –) 2 – 9 fascicled Solitary Variable: solitary or 2 –

peduncles 9 fascicled

Male bracts length 7 – 11 mm 4 – 7 mm 7 – 11 mm 4 – 11 mm

Anther connective Ciliate Glabrous (rarely with 1 – Glabrous (rarely with 1 – Glabrous (rarely with 1 –

2 cilia) 2 cilia) 2 cilia)

Length of fruit peduncle 2 – 10 (– 15) mm 5 – 15 mm 2 – 5 (– 8) mm 2 – 5 (– 10) mm

Diam. of fruit peduncle 2 – 3.5 (– 4) mm 1.5 – 3 (– 3.5) mm (2.5 –) 3 – 5 mm 2.5 – 3.5 mm

Ovary Glabrous or, rarely, with Tomentose (curly hairs) Tomentose (curly hairs) Tomentose (curly hairs) 38

sparse straight hairs;

smooth or lepidote

Twig diameter below the 5 – 10 mm 5 – 11 mm 5 – 8 mm (8 –) 10 – 25 mm

leaves (previous year’s

growth)

Leaves More or less spaced More or less spaced More or less spaced Grouped in rosettes

Petiole length 1.5 – 4.0 (– 4.5) cm 0 – 3.5 cm 1.0 – 4.5 cm 0 (– 0.5) cm

Number of pairs of 7 – 12 (– 13) (12 –) 13 – 25 8 – 13 (– 14) 12 – 21

secondary veins

Straight hairs on lower Lacking or present Lacking Lacking Lacking

surface of leaf blade

Curly hairs on lower Lacking; rarely: very Copious, not restricted to Copious, not restricted to Copious, not restricted to

surface of leaf blade sparse and restricted to main veins main veins main veins

main veins 39

Figures

Fig. 1. Photographs of Uapaca teusczii in the field. A vegetative twig; B tip of twig with stipules; C twig with female inflorescences in the axil of leaves; D twig with young fruits.

Scale bar = 1cm. PHOTOS: M. HASSON.

Fig. 2. Lower surface of leaf blade. A Uapaca teusczii f. pilosa with straight hairs; B Uapaca kirkiana var. kirkiana with curly hairs. PHOTOS: M. HASSON.

Fig. 3. Comparison of Uapaca teusczii (left-hand pictures) and Uapaca kirkiana var. kirkiana

(right-hand pictures). A & B ovary; C & D stamen; E & F male inflorescence; G & H female calyx at fruiting stage. Scale bar = 1 mm. PHOTOS: G. JOSENS, J. FARMINHAO, P.

MEERTS.

Maps

Map 1. Distribution of Uapaca teusczii in the northwest of the Zambezian region (Angola, D.

R. Congo, Zambia).  U. teusczii f. teusczii; U. teusczii f. pilosa.

Map 2. Distribution of Uapaca kirkiana in the northwest of the Zambezian region (Angola,

D. R. Congo, Zambia). U. kirkiana var. kirkiana; U. kirkiana var. benguelensis;

U. kirkiana var. gossweileri.

Table

Table 1. Diagnostic characters separating Uapaca teusczii and the three varieties of Uapaca kirkiana. 40

Fig 1

Fig 2

Fig 3

Map 1

Map 2