Plant Ecology and Evolution 146 (1): 75–94, 2013 http://dx.doi.org/10.5091/plecevo.2013.770 REGULAR PAPER Uapaca (Phyllanthaceae) in the Guineo-Congolian forest region: a synoptic revision Frans J. Breteler Herbarium Vadense, Biosystematics Group, Wageningen University, Generaal Foulkesweg 37, NL – 6703 BL Wageningen, The Netherlands. E-mail: [email protected] Background and aims – The treatment of Uapaca (Phyllanthaceae) for the Flore du Gabon revealed many, hitherto unknown, synonymies and some overlooked names that deserve a separate publication. Advantage is taken of this treatment to establish the geography of the forest species on an African scale. Methods – Normal practices of herbarium taxonomy have been applied to study all herbarium material available, mainly at BM, BR, COI, FHO, G, K, LBV, LISC, LISU, MO, OXF, P (partly), and WAG. Key results – A synoptic revision with key to the nine Uapaca species of the Guineo-Congolian forest region is presented with their synonymies, typifications, (selection of) the examined material and distribution maps. Some morphological features are illustrated to facilitate the use of the key to the species. Two specific names have been neotypified and fifteen lectotypified. Twenty-one specific names have been placed in synonymy. Key words – Uapaca, Phyllanthaceae, taxonomy, distribution, tropical Africa, Guineo-Congolian forest region. INTRODUCTION Many of these specific De Wildeman names have been placed in synonymy (e.g. in U. mole Pax). De Wildeman The forest species of Uapaca (Phyllanthaceae) are character- (1936b) used the Jordanian species concept in his treatment ised by stilt roots, whether growing in marshy areas, along of Uapaca. The application of this concept was explained by rivers, or on dry land, but stilt roots may sometimes be ab- him in detail in a previous paper (De Wildeman 1936a). sent in instances where the forest species extend into the ad- Since De Wildeman (1936b), only treatments for local or jacent savannah. This absence may be observed with Uapaca regional floras have appeared, such as Duvigneaud’s (1950) guineensis Müll.Arg. and U. togoensis Pax, but even in the about the Uapaca species of the ‘forêts claires’ of south- savannah stilt roots are often produced (Keay 1958). The ern Congo Belge, the floras of West Tropical Africa (Keay true savannah species from East and South tropical Africa as 1958), Côte d’Ivoire (Aubréville 1959), Tropical East Africa well as Angola rarely produce stilt roots. This paper provides (Smith 1988), Zambesiaca area (Radcliffe-Smith 1996), Be- a short revision of the (stilt-rooted) Guineo-Congolian spe- nin (Essou 2006), and the Woody Plants of Western African cies. Uapaca lissopyrena Radcl.-Sm., a forest species from Forests (Hawthorne & Jongkind 2006). For Madagascar the the Flora Zambesiaca area, is also stilt-rooted. This latter genus was recently reviewed by McPherson (2011), resulting species is partly treated in a supplement: a distribution map in a reduction from twelve to eight species. Breteler (2011) is provided and the examined specimens are cited. published a new species from Gabon together with a reduced key to the species that lack stipules or have rudimentary stip- After the revision by Pax & Hoffmann (1922), who ac- ules only. Radcliffe-Smith (1993) provides a historical sur- cepted 27 species, De Wildeman (1936b) was the first au- vey of the genus as regards its suprageneric classification. thor to treat the genus for continental Africa. He recognised Mennega (1987) published about the wood anatomy and 47 species, 31 of which were described by him in the period confirmed the more or less isolated position of Uapaca with- 1908–1936. Of De Wildeman’s treatment of the 47 conti- in the subfamily Phyllanthoideae of the Euphorbiaceae. In a nental African species, 29 species are completely, or nearly paper by Hoffmann et al. (2006) dealing with the phylogeny so, confined to the Guineo-Congolian forest region. Of these of the Phyllanthaceae based on molecular analysis, Uapaca forest species 21 were named by De Wildeman. is maintained in its own tribe Uapaceae Hutch. All rights reserved. © 2013 National Botanic Garden of Belgium and Royal Botanical Society of Belgium – ISSN 2032-3913 Pl. Ecol. Evol. 146 (1), 2013 Material AND METHODS Normal practices of herbarium taxonomy have been applied to study all herbarium material available, mainly at BM, BR, COI, FHO, G, K, LBV, LISC, LISU, MO, OXF, P, and WAG. Due to rebuilding activities in connection with scanning op- erations, the material of the Paris herbarium has only partly been examined. RESULTS Chorology and ecology The nine Uapaca species of the Guineo-Congolian forest re- gion (White 1979) are often wide-spread, ranging from Sé- négal (sometimes Mali) to the Democratic Republic of the Congo and Angola (fig. 1). Five species, Uapaca guineensis Müll.Arg., U. heudelotii Baill., U. mole Pax, U. pynaertii De Wild. and U. togoen- Figure 2 – Species density of the stilt-rooted Uapaca species in sis Pax, show this distribution pattern. Of the remaining Africa. four species Uapaca acuminata (Hutch.) Pax & K.Hoffm., U. staudtii Pax, and U. vanhouttei De Wild. are confined but one species occur over considerable ranges may be the to Lower Guinea and Congolia, whereas U. niangadoumae result of seed dispersal by birds. Although dispersal by birds Breteler is restricted to a small area in NW Gabon. One spe- like hornbills seems likely, there are no references for this cies is also found East of the Guineo-Congolian region, in dispersal so far. Another difference in the diversity pattern of the area of the Flora of Tropical East Africa (Smith 1988): Uapaca compared to other forest taxa is that Uapaca extends U. mole (as U. paludosa Aubrév. & Leandri ) is reported with several species into the savannah zone, especially that from Uganda and Tanzania. Uapaca heudelotii is also known to the North of the West African forest block. from Angola, south of the Congo river. The presence of stilt roots may suggest that Uapaca spe- The highest diversity of this group is found in Lower cies of the Guineo-Congolian forest region are to be found Guinea (fig. 2), which is a usual pattern for African forest under marshy conditions and/or in periodically inundated ar- taxa (e.g. Wieringa 1999, Versteegh & Sosef 2007, Breteler eas only. This is true for some species. Uapaca heudelotii is & Nguema Miyono 2008, Breteler & Wieringa 2008, only found along rivers and U. niangadoumae, U. staudtii Breteler 2010, 2011). While most of these taxa have a more and U. vanhouttei are species of marshy or periodically inun- or less clear area of richest diversity in Southern Cameroon dated areas. Uapaca acuminata is a dry land species, which and Gabon, Uapaca shows a wider core area, ranging from has never been found in wet conditions, but has a stilt root Nigeria to D.R.Congo and showing a high species richness system that is the most impressive of all Uapaca species of in Upper Guinea as well. This more extended pattern is the the region, with a height of up to at least six meters. Ua- result of the fairly large ranges most species occupy. That all paca mole shows a wide ecological range. It thrives well in marshy areas, but also in secondary forests on slopes. The same holds more or less for Uapaca pynaertii. Uapaca guineensis and U. togoensis are mostly found along rivers and in gallery forests, but also in adjacent dry land forests and savannahs. In Gabon, Uapaca togoensis has been ob- served in such a condition, colonising a man-made savannah. Morphology The stilt roots of the forest species of Uapaca are more or less terete, which character is helpful in identifying the genus in the field. Saplings of not more than 10 cm in diameter and less than 10 m in height are usually already provided with stilt roots, whether growing in wet or marshy sites or in dry land forests. Uapaca mole, U. staudtii and U. vanhouttei are always markedly stipulate, whereas U. acuminata, U. niangadou- Figure 1 – Distribution of the stilt-rooted Uapaca species in Africa, mae and U. pynaertii are devoid of stipules. In Uapaca heu- U. lissopyrena has been pictured in yellow squares, the other species delotii (fig. 3F) the stipules are replaced by tufts of hairs and in red circles. The forest block is indicated with green, altitude with in U. guineensis and U. togoensis the stipules are usually a grey-scale. very small, leaving small scars, or are even absent. 76 Breteler, Synopsis of Guineo-Congolian Uapaca (Phyllantaceae) Key to the Guineo-Congolian forest species of Uapaca 1. Stipules absent, at least inconspicuous; scars of stipules absent or inconspicuous….......................…2 1. Stipules present, distinct, fugacious or ± persistent, scars of stipules conspicuous….......................…7 2. Branchlets glabrous at most with some axillary hairs……………………………........................…….3 2. Branchlets pubescent or puberulous, at least around or at the base (stipular area) of the petioles (fig. 3F)...................................................................................................................................................4 3. Leaves papery to subcoriaceous, (5–)8–10(–14) × (2.5–)3–5.5 cm, generally short- ly acuminate at apex (fig. 3A), with (5–)6–9(–10) pairs of main lateral nerves; inflores- cence < 1 cm in diam.; from southern Nigeria to western D.R.Congo and northern Angola (Cabinda)……….............………………………....................................……………..1. U. acuminata 3. Leaves stiff, thickly coriaceous, (14–)16–23(–42) × 6–8(–14) cm, obtuse to retuse (rare- ly to 1.5 cm acuminate) at apex, with (7–)8–12 pairs of main lateral nerves; (fig. 8E); in- florescence (1.5–)2–4 cm in diam.; from Guinea to D.R.Congo and Central African Republic…………….…….…………......................................…………………………6. U. pynaertii 4. Branchlets with tufts of red, ± curly hairs in the leaf axils and often also lateral at the base of the pe- tioles (fig.