bs_bs_banner Botanical Journal of the Linnean Society, 2013, 173, 407–441. With 5 figures Taxonomy and phylogenetics of Cuviera (Rubiaceae–Vanguerieae) and reinstatement of Globulostylis with the description of three new species BRECHT VERSTRAETE1*, OLIVIER LACHENAUD2,3, ERIK SMETS STEVEN DESSEIN3 and BONAVENTURE SONKÉ2,5,6 FMLS1,4, 1 Plant Conservation and Population Biology, KU Leuven, Kasteelpark Arenberg 31, PO Box 2437, 3001 Leuven, Belgium 2 Service d’Evolution Biologique et Ecologie, Université Libre de Bruxelles, CP 160/12, 50 Avenue F. Roosevelt, 1050 Bruxelles, Belgium 3 National Botanic Garden of Belgium, Domein van Bouchout, 1860 Meise, Belgium 4 Naturalis Biodiversity Center, Leiden University, PO Box 9517, 2300 RA Leiden, The Netherlands 5 Plant Systematic and Ecology Laboratory, Higher Teachers’ Training College, University of Yaounde I, PO Box 047, Yaounde, Cameroon 6 Africa and Madagascar Department, Missouri Botanical Garden, PO Box 299, St Louis, MO 63166-0299, USA Received 19 May 2012; revised 12 November 2012; accepted for publication 19 April 2013 Generic circumscriptions in tribe Vanguerieae (Rubiaceae) have been under discussion for a long time. Recent molecular studies, while providing new insights, have not yet solved all the problems. In this study, the taxonomy and phylogenetics of the genus Cuviera s.l. are investigated. On the basis of molecular and morphological evidence, Cuviera is restricted to a group of ten West and Central African species. Globulostylis, previously included in Cuviera, is reinstated as a distinct genus, with eight species from Central Africa. Both genera are revised; the latter includes three new species (G. dewildeana, G. rammelooana and G. robbrechtiana) and two new combinations (G. leniochlamys and G. uncinula). The close relationship of Cuviera s.s. and Globulostylis to Vangueriella is shown. Six aberrant species (most from East Africa) are excluded from Cuviera, but further work is needed before they can be confidently assigned to other genera. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441. ADDITIONAL KEYWORDS: Cameroon – Equatorial Guinea – Gabon – Tropical Africa. INTRODUCTION Tribe Vanguerieae are palaeotropical in distribution and among the larger tribes in Rubiaceae, comprising approximately 600 species in 25 currently accepted genera (Govaerts et al., 2012). The tribe is delineated by a set of morphological characteristics, in particular the specialized hood-like structure of the stigma (Lantz & Bremer, 2004), the valvate corolla lobes, the *Corresponding author. E-mail: brecht.verstraete@bio.kuleuven.be pendulous ovules and the drupaceous fruits with oneseeded pyrenes in each locule. Generic delimitation in the tribe is still problematic as several genera lack good diagnostic characters. Not surprisingly, recent molecular studies are partially in conflict with morphological classifications (Lantz et al., 2002; Lantz & Bremer, 2004, 2005). At present, molecular sampling and tree resolution are insufficient to answer all questions related to the generic delimitations in the tribe. Detailed molecular studies in particular groups of the tribe are rare (e.g. Razafimandimbison et al., 2009) and many taxonomic problems are still to be solved. Furthermore, many © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 407 408 B. VERSTRAETE ET AL. species remain to be described from Central and West Africa, but also from Madagascar and Asia. The main aim of this study is to elucidate the phylogenetic position of the African genus Cuviera DC., especially in relation to Vangueriella Verdc. Cuviera was described by de Candolle (1807) with one species, C. acutiflora DC. It was regarded as closely related to Vangueria Juss. because of the solitary pendulous ovules in each locule. Further species were added by various authors (Bentham in Hooker, 1849; Hiern, 1877; Schumann, 1888, 1899, 1903; Krause, 1912; Wernham, 1914, 1918; Mildbraed, 1924; Verdcourt, 1957, 1981; Hallé, 1960). Wernham (1913) described the genus Globulostylis Wernham with two species, G. minor Wernham and G. talbotii Wernham, and later added a third species, G. cuvieroides Wernham (Wernham, 1918). Although Cuviera and Globulostylis clearly belong to the tribe Vanguerieae, they were not addressed in the monograph of the tribe by Robyns (1928). Hallé (1960) published a conspectus of Cuviera, including studies of inflorescence, style and pyrene morphology, and a key to the species then known. He noted that certain Cuviera spp. have a hairy swelling at the base of the style and suggested an infrageneric group. Verdcourt (1987) saw the resemblance of these species with Globulostylis and treated the latter as a subgenus of Cuviera. More recently, Onana (2008) presented a conspectus of subgenus Globulostylis. Currently, Cuviera s.l. (including Globulostylis) includes 24 accepted species (Govaerts et al., 2012). The genus occurs mainly in the Guineo–Congolian region of Africa, but four species, poorly known and of questionable generic position, are found in East Tropical Africa (Verdcourt & Bridson, 1991; Bridson, 1998). So far only C. physinodes K.Schum. (misidentified as C. angolensis Welw. ex K.Schum.) has been included in molecular studies (Lantz et al., 2002; Lantz & Bremer, 2004, 2005), and its position was unresolved. Vangueriella, with 18 species in two sections (section Vangueriella and section Stenosepalae (Robyns) Verdc.), was erected by Verdcourt (1987) when he raised Vangueriopsis Robyns subgenus Brachyanthus Robyns (Robyns, 1928) to generic rank. Verdcourt considered Vangueriella to be related to Cuviera. Molecular analyses (Lantz et al., 2002; Lantz & Bremer, 2004) rather suggested an affinity to Canthium Lam., but the only species included (V. spinosa (Schumach. & Thonn.) Verdc.) is atypical. We therefore found it necessary to add data for Vangueriella while investigating Cuviera. The taxonomic problems in Cuviera do not only involve generic delimitation, but frequent misidentifications between the species also occur in literature and even more so in herbaria. For example, in the Checklist of Gabonese vascular plants (Sosef et al., 2006), three out of ten species were misidentified. The names C. acutiflora DC., C. leniochlamys K.Schum., C. longiflora Hiern and C. minor C.H.Wright, in particular, have been commonly misapplied. A taxonomic revision of this group was therefore necessary and is provided here. MATERIAL AND METHODS To test the monophyly and systematic position of Cuviera and Vangueriella, a molecular phylogenetic study has been performed based on seven DNA regions (trnTL, trnLF, rpl16, petD, ITS, rpl32-trnL and accD-psaI). Ninety-two specimens of Vanguerieae representing the different groups recognized in the tribe were included in the phylogenetic analysis, as well as three outgroup species. Detailed information on the investigated taxa can be found in Appendix 1. DNA of silica-dried and herbarium material was extracted using the E.Z.N.A.TM HP Plant DNA Mini Kit (Omega Bio-Tek). The primers for the amplification of the seven DNA regions can be found in Appendix 2, with their respective annealing temperatures. Purified PCR products were sent to Macrogen for sequencing (Macrogen Inc., Amsterdam, the Netherlands). The sequences were assembled, edited and aligned using Geneious 5.4 (Drummond et al., 2011). Gaps were coded following Simmons & Ochoterena (2000). Maximum parsimony analysis was conducted using PAUP* v.4.0b10a (Swoffort, 2002). Heuristic searches were conducted with tree bisection– reconnection (TBR) branch swapping on 10 000 random addition replicates, with five trees held at each step. Non-parametric bootstrap analysis was carried out to calculate the relative support for individual clades found in the parsimony analysis. For each of 1000 bootstrap replicates, a heuristic search was conducted with identical settings as in the original heuristic analysis. Phylogenetic trees were also estimated using probabilistic methods under Bayesian in the CIPRES web portal (Miller, Pfeiffer & Schwartz, 2010). Bayesian analysis was inferred using MrBayes 3.1, running four Markov chains sampling every 100 generations for 5 million generations. DNA substitution models for the individual markers were selected by preforming jModelTest 0.1.1 (Posada, 2008) (Appendix 2). The concatenated data set was partitioned and independent models were applied for each of the partitions. The taxonomic revision is based on the examination of herbarium specimens in BM, BR, BRLU, E, HBG, K, LBV, MPU, P, WAG and YA. Central African collections of Cuviera from all these herbaria and West African collections from BM, BR, K and WAG, were seen for this study. The distribution is based on the specimens of the above-mentioned herbaria, comple- © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS mented with data from the literature in a few cases (e.g. Hepper & Keay, 1963). Measurements, colours and other details given in the descriptions are based on living material, spirit and herbarium specimens and data derived from field notes. The dimensions given for the calyx do not include the ovary. Phytogeographical considerations follow White (1979), but we simplify his ‘(sub)centres of endemism’ into Domain and Region. Specimens are cited alphabetically first by country (with a further distinction for Equatorial Guinea between Bioko Island and Rio Muni) and then by collector, with the initials only given in the case of ambiguity (e.g. J. J. F. E. de Wilde and W. J. J. O. de Wilde). The conservation status was assessed by calculating the extent of occurrence or the area of occupancy using the geographic information system (GIS) and applying the 2001 IUCN Red List Categories and Criteria. 409 4. The focus of this study is on the Cuviera/ Vangueriella group. It includes three wellsupported subclades corresponding to Cuviera subgenus Cuviera (excluding C. schliebenii Verdc. and C. semseii Verdc.), Cuviera subgenus Globulostylis and Vangueriella (excluding V. spinosa). The latter two subclades are weakly supported as sister groups. 5. The Vangueria group includes Multidentia Gilli, Pygmaeothamnus zeyheri (Sond.) Robyns, Robynsia Hutch., Vangueria (in the enlarged sense of Lantz & Bremer, 2005), Vangueriopsis and two East African species of Cuviera (C. schliebenii and C. semseii). Vangueria is monophyletic with good support but the rest of the clade is not well resolved. DISCUSSION RESULTS The analysis of the combined data set using maximum parsimony and Bayesian inference yielded similar trees, of which the consensus tree is shown in Figure 1. Bootstrap values and posterior probabilities are indicated below the respective branches. Vanguerieae as a whole is monophyletic and strongly supported. The consensus tree is well resolved and contains several strongly supported clades, in correspondence with the findings of Lantz & Bremer (2004). 1. The earliest branching clade contains the genera Afrocanthium, Keetia, Psydrax, Bullockia, Peponidium and Pyrostria, the last three forming the dioecious group of Lantz & Bremer (2004). In our analysis, Keetia is sister to the remainder of the clade and Afrocanthium is sister to the dioecious group, whereas Lantz & Bremer (2004) found these two genera to be sister to each other. Relationships in the dioecious group are not well resolved; for a detailed investigation of this group see Razafimandimbison et al. (2009). 2. The so-called ‘spiny’ group of Lantz et al. (2002) containing most Canthium Lam. spp., Plectroniella armata (K.Schum.) Robyns, Pygmaeothamnus chamaedendrum (Kuntze) Robyns and Vangueriella spinosa (Schumach. & Thonn.) Verdc. The name ‘spiny’ is somewhat misleading as spiny taxa are not uncommon in other clades as well (e.g. Cuviera and Rytigynia). 3. As in all previous studies of Vanguerieae, our study finds the Fadogia Schweinf./Rytigynia Blume group as a clear and distinct clade in the tribe. The representatives of this group all have unique indels in the investigated DNA regions. The internal topology of this clade is still unclear. Our phylogenetic tree indicates several suprageneric groups with good support and these are not in contradiction with the findings of Lantz & Bremer (2004). The molecular data show a supported clade containing Cuviera s.l. (including subgenus Globulostylis) and Vangueriella (except V. spinosa) (Fig. 1). In this clade three distinct groups are strongly supported: C. subgenus Cuviera, C. subgenus Globulostylis and Vangueriella. In our opinion there are three possible options for defining phylogenetic groups: (1) placing all species in a large genus Cuviera containing three subgenera Cuviera, Globulostylis and Vangueriella; (2) maintaining the taxonomy as it is now with the genera Cuviera s.l. and Vangueriella; and (3) reinstating Globulostylis as a genus and recognizing three genera. A major inconvenience of an enlarged Cuviera is that it would become quite heterogeneous and difficult to define morphologically; for example, in relation to Vangueria. Even in the current sense, Cuviera is quite heterogeneous and not easily separated from Vangueriella based on morphological characters. In addition, the monophyly of Cuviera + Globulostylis is not supported by molecular data. We favour option 3, which allows the formation of more homogeneous (and molecularly well-supported) groups (see Table 1 for distinguishing characters); the problematic case of Cuviera le-testui Pellegr. is discussed below. We therefore recognize three genera, Cuviera, Globulostylis and Vangueriella. These are briefly discussed further, as well as species excluded from Cuviera. CUVIERA SENSU STRICTO Cuviera, now considered in a narrower sense, is revised here and includes ten species from West and © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 410 B. VERSTRAETE ET AL. Figure 1. Strict consensus tree of the tribe Vanguerieae based on the combined analysis of seven DNA markers. Bayesian posterior probabilities/bootstrap values are indicated below the branches. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS 411 Table 1. Main distinguishing characters between Vangueriella, Globulostylis and Cuviera Vangueriella Globulostylis Cuviera s.s. C. le-testui Ovary Inflorescences Bilocular Many-flowered Pentalocular One- to five-flowered Bilocular Many-flowered Bracts at lowest node Corolla lobes (shape) Present/absent Present Pentalocular Three- to many-flowered Absent Present Narrow, at most equalling tube (occasional appendages not included) Plain green or yellow Rather broadly triangular, variously shorter or longer than tube Narrow, longer than tube Narrow, longer than tube Plain green or yellow Plain green Slender Swollen in lower half (sometimes slightly so) Absent Absent Striped (see note under C. trilocularis) Slender, rarely (C. pierrei) thickened in upper two-thirds Usually present Corolla lobes (colour) Style Ant holes in twigs Central Africa. Most of these species form a rather homogeneous group that is well characterized, in particular by their striped petals, many-flowered inflorescences and (usually) ant holes in the twigs. The bracts are recaulescent (Hallé, 1960), which means they are moved to the upper node. Therefore, the first node of the inflorescence is bare and the lowest bracts are inserted at the second node (or on the lateral pedicels if the inflorescence is unbranched). Three species are more or less atypical and need some discussion. Cuviera trilocularis Hiern lacks ant holes in the twigs (which is sometimes also the case in the type species, C. acutiflora) and has few-flowered inflorescences, but matches Cuviera in other characters (e.g. style, bracts). We did not obtain enough DNA data of this species to include it in the general analysis, but when analysing the trnL-F marker, C. trilocularis clearly belongs to the Cuviera s.s. clade. Cuviera pierrei also has few-flowered inflorescences and the style is thickened in the upper two-thirds, which is reminiscent of Globulostylis (although in the latter the style swelling occurs in the lower half). Presumably for these reasons, it was included in subgenus Globulostylis by Onana (2008). However, the striped corolla lobes, twigs with ant holes and the lowest node of the inflorescence lacking bracts (these are inserted halfway on the pedicels of the lateral flowers) are clear differences with Globulostylis and place this species in Cuviera s.s. Cuviera le-testui is certainly the most aberrant species (Table 1). It is presumably related to Cuviera s.s. in having ant holes in the twigs, many-flowered inflorescences and corolla lobes longer than the tube, Slender Present but differs in its bilocular ovary, plain green corolla lobes and in having a pair of bracts at the lowest node of inflorescence (the other bracts are recaulescent and therefore the second nodes of the inflorescence are bare; Hallé, 1960). Its peculiar characters could well justify the creation of a new genus, but we think this would be premature given the lack of genetic data. We therefore provisionally keep it in Cuviera s.s. GLOBULOSTYLIS Globulostylis is reinstated with eight species, including three newly described (G. dewildeana, Fig. 2; G. rammelooana, Fig. 3; and G. robbrechtiana, Fig. 4) and two newly transferred from Cuviera (G. leniochlamys (K.Schum.) Sonké, O.Lachenaud & Dessein and G. uncinula (N.Hallé) Sonké, O.Lachenaud & Dessein). The main characters of Globulostylis are the few-flowered inflorescences with a pair of bracts at the apex of the peduncle and the style with a swelling (sometimes rather slight) in the lower half. The genus is quite homogeneous; the only slightly atypical species is G. cuvieroides Wernham, which has narrow calyx lobes. VANGUERIELLA A remarkable finding is that Vangueriella is polyphyletic; most of the species studied [including V. laxiflora (K.Schum.) Verdc., the type of the genus] form a well-supported clade related to Cuviera but V. spinosa (Schumach. & Thonn.) Verdc. (the type species of section Stenosepalae) is related to Canthium, as already found by Lantz & Bremer (2004). This is not © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 412 B. VERSTRAETE ET AL. Figure 2. Globulostylis dewildeana. A, flowering twig. Bracts are represented by dotted lines. B, stipule. C, flower bud. D, open flower. E, style. F, cross section of ovary. G, fruit. [Based on: J.J.F.E. de Wilde 7886 (A–F); Letouzey 9843 (G)]. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS 413 Figure 3. Globulostylis rammelooana. A, flowering branch. Bracts are represented by dotted lines. B, stipule. C, flower bud. D, open flower. E, style. F, cross section of ovary. G, fruit. H, pyrene. [Based on: Sonké & Simo 4671 (A, B); Sonké & Simo 4711 (C–F); Sonké & Taedoumg 4464 (G, H)]. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 414 B. VERSTRAETE ET AL. Figure 4. Globulostylis robbrechtiana. A, flowering branch. Bracts are represented by dotted lines. B, stipule. C, flower bud. D, calyx. E, open flower. F, style. G, cross section of ovary. H, fruit. I, pyrene. [Based on: Elad et al. 1554 (A); Sonké, Taedoumg & Simo 4868 (B–G); Lisowski 1571A (H, I)]. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS KEY TO CUVIERA AND 415 GLOBULOSTYLIS 1. Inflorescence three- to many-flowered, the lowest node lacking bracts (except C. le-testui); style slender, rarely (C. pierrei) thickened in the upper two-thirds; twigs frequently with ant holes; corolla lobes usually with a median stripe ............................................................................................................Cuviera 1’. Inflorescence one- to five-flowered with a pair of bracts at lowest node (often caducous in fruit); style swollen in the lower half (sometimes slightly so); twigs without ant holes; corolla lobes plain green or yellow, not striped .........................................................................................................Globulostylis so unexpected as V. spinosa (and several other species of section Stenosepalae) differ quite markedly from the rest of the genus in floral characters, lacking a ring of deflexed hairs inside the corolla tube and having vestigial or subulate (rather than foliaceous) calyx lobes. Interestingly, Vangueriella rufa (Robyns) Verdc., placed in section Stenosepalae by Verdcourt (1987), is sister to V. discolor (Benth.) Verdc. of section Vangueriella (which it indeed closely resembles in flower, leaf and fruit). This species has the calyx lobes narrower than usual in section Vangueriella, but still clearly foliaceous, and the corolla tube has a ring of deflexed hairs inside, as in section Vangueriella. This shows that the circumscription of Vangueriella has to be redefined, which will be the object of a future study. Most species of section Stenosepalae will probably have to be transferred to Canthium. Lantz & Bremer (2004) already transferred V. spinosa (as Canthium stenosepalum Lantz), but this has not been widely followed in the literature (e.g. Hawthorne & Jongkind, 2006 still used V. spinosa). SPECIES EXCLUDED FROM CUVIERA Six Cuviera spp. should be excluded from the genus based on morphological and/or molecular data, but at present cannot be transferred to other genera. These are C. calycosa Wernham, C. nigrescens (Scott-Elliott ex Oliv.) and the four East African species (C. midgeodii Verdc., C. schliebenii Verdc., C. semseii Verdc. and C. tomentosa Verdc.). All these species differ from both Cuviera s.s. and Globulostylis in having the inflorescences borne on old woody twigs below the leaves. However, they do not form a homogeneous group and further studies, especially in the Vangueria group, are crucial to understand their position. Cuviera calycosa, C. nigrescens and C. semseii are closely related and, in fact, may even be conspecific. The first two were already indicated as the least typical species of the genus by Hallé (1960). They have a pentalocular ovary, as usual in Cuviera, but differ in (1) inflorescence position, (2) pyrenes with a sharp dorsal crest, (3) small, black-drying leaves with tuft domatia in nerve axils, and (4) twigs that become woody soon, with the bark peeling off. The molecular data (for C. semseii only) suggest an affinity with Vangueria. Further studies are required before these species can be transferred to either Vangueria or a new genus. Cuviera schliebenii and C. tomentosa have a bilocular ovary and small flowers, two characters reminiscent of Vangueriella, but the molecular data for C. schliebenii rather suggest an early branching position in the Vangueria group. Until the generic relationships in this group are better understood, we cannot suggest a generic placement. The last species, C. migeodii, is only known from the poor type and is therefore difficult to place, but is clearly not a Cuviera s.s. The small, black-drying leaves with tuft domatia resemble C. nigrescens, but the ovary is described as bilocular. In conclusion, we here reinstate Globulostylis as a genus and we describe three new species, bringing the total to eight species. The remaining ten species are placed in Cuviera s.s. Four East African species, C. calycosa and C. nigrescens are aberrant and are excluded from Cuviera. Vangueriella is shown to be paraphyletic with V. section Stenosepalae related to the ‘spiny’ group and V. section Vangueriella to Globulostylis. TAXONOMIC TREATMENT Keys to the genera Cuviera and Globulostylis to their respective species and a detailed species list, including the description of the three new Globulostylis spp. (G. dewildeana, G. rammelooana and G. robbrechtiana) are presented. Species excluded from Cuviera are listed at the end. CUVIERA DC., Ann. Mus. Hist. Nat. 9: 222. 1807. nom. cons. Type species: Cuviera acutiflora DC., Ann. Mus. Hist. Nat. 9: 222. 1807. Description: Small trees or shrubs, 1–14 m high, evergreen, with regular horizontal branching (Roux’s architectural model); vegetative parts glabrous or rarely puberulent; twigs usually with swollen and © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 416 B. VERSTRAETE ET AL. hollow internodes, inhabited by ants (solid in C. trilocularis and occasionally in C. acutiflora); young plants and coppicing shoots frequently armed with paired supra-axillary spines that sometimes persist on the trunk. Leaves opposite, ± elliptic, shortly petiolate; stipules small, ± basally connate, not or shortly (< 2 mm) awned, caducous to ± persistent. Inflorescences axillary and paired at each node on young twigs, cymose, three- to many-flowered; bracts linear to broadly elliptic, ± leafy, but absent at lowest node of inflorescence (except in C. le-testui), inserted at the base of pedicels (or towards the middle in C. pierrei). Flowers hermaphrodite. Calyx lobes three to five, free or shortly connate at base, ± leafy, linear to broadly ovate and usually longer than the corolla, or rarely truncate calyx (C. truncata). Corolla tube campanulate to nearly cylindrical, glabrous outside, with a ring of reflexed hairs near the middle inside; lobes five, longer than tube, elongate, acute to longapiculate at apex, spreading or reflexed, glabrous, usually green with a white to pink median stripe (apparently plain green in C. le-testui). Stamens five, inserted in upper part of tube, anthers partly to fully exserted (frequently reflexed between the lobes). Ovary (bi-) pentalocular, obconic or turbinate, sometimes five-angled; ovules solitary in each locule, pendulous. Disk ± flat, glabrous or hairy in the centre. Style exserted, narrowly cylindrical or (in C. pierrei) thickened in the upper two-thirds, stiffly pubescent to glabrous; stigmatic club ± mitriform, shortly (two-) five-lobed. Fruits drupaceous, apparently remaining green, large 2.0–3.7 × 1.8–3.3 cm, subglobose to obovoid, smooth to variously lobed, with calyx generally persistent (sometimes tardily caducous); pyrenes (two-)five, one-seeded, shaped like a mandarin orange segment, usually with a faint dorsal crest, and a slight ventral indentation towards the upper third. Distribution and ecology: Cuviera has ten species in the Guineo–Congolian rainforest zone (including Bioko Island). The centre of diversity is in Lower Guinea; only two species occur in Upper Guinea (C. acutiflora and C. macroura) and only one extends into Congolia (C. angolensis). Most species are relatively light demanding, favouring secondary or riverine forest, although some can be found in the understory as well. They are frequently gregarious, especially C. physinodes that may form dense thickets locally. Notes: The most important characters for species identification in Cuviera are the calyx lobes, the corolla and, to some degree, the fruits. Leaves are of limited value, as in many species they are quite variable; for example, the leaf base can be acute to subcordate. However, an experienced researcher can identify most of the sterile collections if the origin is taken into account. Several aspects of the biology of Cuviera would be interesting to study in the field, in particular the association with ants, which presumably play a role in protecting the plant (although in our experience they are not aggressive to humans). Whether different Cuviera spp. have different associated ants is not known. Not much is known about the pollination of the flowers, which are quite attractive and often fragrant (e.g. those of C. macroura smell of honeysuckle), or about the dispersal of the fruits (although, considering the size of the pyrenes, large animals are presumably involved). Cuviera spp. are rarely used by humans, although at least two species (C. le-testui and C. longiflora) have edible fruits, and the leaves of C. angolensis subsp. latior are eaten as a vegetable in the Democratic Republic of the Congo (according to Pynaert 11, BR). 1. CUVIERA ACUTIFLORA DC., Ann. Mus. Hist. Nat. 9: 222. 1807. Type: SIERRA LEONE. s.loc., Smeathman s.n. (type: BM, see Notes). Synonyms: Cuviera africana Spreng., Syst. Veg. 1: 760. 1824. Type: not traced. Vangueriopsis coriacea Robyns, Bull. Jard. Bot. État 22: 319. 1952. Type: LIBERIA. 3 miles north-east of Suacoco, Traub 299 (holotype: BR000000882973!; isotypes: BM000903517!, BR000000882965!, MO n.v.). Distribution: Lower and Upper Guinea Domain: Cameroon, Equatorial Guinea (Rio Muni), Ghana (see Notes), Guinea, Ivory Coast, Liberia, Nigeria, Sierra Leone and Togo (see Notes). Notes: Cuviera acutiflora is closely related to C. physinodes and C. truncata; the three species share coriaceous leaves and glabrous, shortly exserted styles. Cuviera acutiflora is a widespread and polymorphic species, especially in West Africa where the leaf base varies from cuneate to asymmetrically cordate; specimens from Cameroon and Equatorial Guinea always have cordate leaf bases (which allows them to be separated from C. physinodes). Ant holes in the twigs can be present or absent, apparently irrespective of locality. Some Nigerian specimens are intermediate with C. truncata (see that species). The species has been reported from Ghana (Hepper & Keay, 1963) and Togo (Brunel, Hiepko & Scholz, 1984), which seems likely on geographical grounds, but we have not seen the specimens. Records of C. acutiflora from Gabon (Sosef et al., 2006) refer to C. physinodes. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS KEY TO THE CUVIERA 417 SPECIES 1. Ovary and fruits bilocular; inflorescence many-flowered with a pair of large bracts at the lowest node; apex of flower buds with reflexed appendages (Gabon to south-west Democratic Republic of the Congo) ......................................................................................................................................C. le-testui 1’. Ovary and fruits pentalocular; no bracts at lowest node of inflorescence; flower bud appendages, if present, never reflexed .................................................................................................................2 2. Calyx truncate or with short (< 2 mm) irregular lobes (Nigeria and south-west Cameroon) .C. truncata 2’. Calyx lobes conspicuous, 4 mm long at least (sometimes caducous on fruit) .................................... 3 3. Calyx lobes three(–four), the fourth lobe when present often slightly smaller; style hairy, long exserted .................................................................................................................................................. .4 3’. Calyx lobes five ...........................................................................................................................5 4. Calyx lobes narrowly lanceolate, of about the same width throughout, 2.5–4.0 mm wide (west Angola and south-west Democratic Republic of the Congo) ...................................C. angolensis subsp. angolensis 4’. Calyx lobes ovate, conspicuously widened above the base, 2.5–23.0 mm wide (Gabon to north-east Angola) ...................................................................................................C. angolensis subsp. latior 5. Leaves thick and coriaceous, tertiary nerves invisible or lax; corolla lobes not (or shortly) apiculate .. .................................................................................................................................................. .6 5’. Leaves not markedly thick, tertiary nerves apparent, usually dense (sometimes laxer in C. subuliflora); corolla lobes often with a filiform appendage (lacking in C. pierrei) ............................................... .9 6. Inflorescence three- to five-flowered, very lax; twigs lacking ant holes (Nigeria and south-west Cameroon) .......................................................................................................................C. trilocularis 6’. Inflorescence many-flowered (but often with few fruits developing); twigs usually with ant holes, sometimes lacking in C. acutiflora ................................................................................................7 7. Style hairy, exceeding corolla throat by 8–10 mm; corolla lobes 10–25 mm long; fruit large, 3.2–3.7 × 2.3– 3.3 cm when dry, on long (1.5–2.0 cm) and much-thickened pedicel (Nigeria to the Central African Republic) .....................................................................................................................C. longiflora 7’. Style glabrous, exceeding corolla throat by 2–3 mm only; corolla lobes 5–9 mm long; fruit smaller, 2.0–3.2 × 1.8–2.2 cm when dry, on short pedicel (0.5–1.0 cm) .......................................................... 8 8. Corolla tube broad (throat 4–5 mm wide); flower buds conical, conspicuously broadening towards base; anthers reflexed between corolla lobes; bracts and calyx lobes narrow, 0.5–2.0(−4.0) mm wide; fruit broadly ellipsoid, 1.9–2.5 × 1.5–2.2 cm, drying generally lobed; leaf base variable, cuneate to subcordate (Guinea to Rio Muni) ...................................................................................................C. acutiflora 8’. Corolla tube narrow (throat c. 2 mm wide); flower buds almost cylindrical, not broadening towards base; anthers erect; bracts and calyx lobes ovate to lanceolate, broader, (2-)3–8 mm; fruit narrowly ellipsoid, c. 3 × 2 cm, drying smooth; leaf base always cuneate (south Cameroon to south Gabon) ............................................................................................................................... ..C. physinodes 9. Calyx lobes almost filiform, 5–15 mm long, < 1 mm wide, fused at base into a short cup; domatia present as small glabrous holes in axils of nerves; calyx lobes caducous on fruit (Guinea to Cameroon) ...................................................................................................................................C. macroura 9’. Calyx lobes broader, (12–)16–50 mm long, (1.2–)2.0–5.0 mm broad, not forming a cup at base; domatia absent; calyx lobes persistent (but sometimes damaged) on fruit ................................................. .10 10. Inflorescence seven- to many-flowered (but only one fruit may develop); bracts long (> 2 cm), approximately equalling the sepals; corolla lobes ending in a filiform appendage; fruit broadly truncate at apex, with the calyx lobes patent and star-like (Nigeria to south-west Democratic Republic of the Congo, also Bioko) .......................................................................................................................C. subuliflora 10’. Inflorescence two- or three- (five-) flowered; bracts small (< 1 cm), much shorter than the sepals; corolla lobes not appendaged; calyx lobes erect on fruit (Equatorial Guinea and Gabon) .................C. pierrei The typification of C. acutiflora needs further investigation. Two Smeathmann sheets in BM are probably part of the original gathering, but it is not clear whether De Candolle saw them. There might be other sheets in P, where De Candolle worked, so we have not chosen a lectotype. Additional specimens examined: CAMEROON. Batanga, Apr 1895, Bates 98 (K); 9 km north of Kribi, 27 Feb 1969, Bos 4010 (P, WAG, YA); ibid., 30 Oct 1969, Bos 5556 (BR, K, P, WAG, YA); Ibid., 9 Nov 1970, Bos 6118 (BR, K, P, WAG, YA); km 20, CampoKribi, 10 Nov 1970, Bos 8706 (YA); ibid., 4 Dec 1975, © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 418 B. VERSTRAETE ET AL. J.J.F.E. de Wilde 8706 (BR, K, P, WAG); Bimbia– Bonadikombo community forest, 27 Apr 2009, Dessein et al. 2809 (BR, MO, YA); 8 km north of Kribi, 13 Feb 1997, Elad 530 (WAG); Limbe, 1904, Kalbreyer 29 (K); Victoria [= Limbe], Nov 1929, Maitland 784 (BR, K); Ambas Bay, Feb 1861, Mann 770 (K); Cameroon Mountains, 1862, Mann 1211 (P); Bipindi, 25 Nov 2004, Sonké 3578 (BR); Douala–Nkongsamba road, north Maleke, 17 Jan 1983, Van der Zon 2021 (P, WAG, YA); Liwenyi, 31 Nov 1993, Watts 893 (K); Bipindi, 1903, Zenker 2651 (BM, BR, HBG, K, P, WAG); ibid., 1908, Zenker 3594 (BM, BR, HBG, K, P); ibid., Jun 1913, Zenker 264 (BR, P). EQUATORIAL GUINEA. Rio Muni: zona do Bairro Ikundi, 23 Jun 1991, Carvalho 4749 (WAG); Rio Ndote, 7 Jun 1999, Eneme 304 (BRLU, WAG). IVORY COAST. Km 25 Sassandra–Gagnoa, 30 Oct 1968, Breteler 5860 (BR); near Abidjan, Banco Forest Park, 9 Jan 1976, de Koning 6377 (BR); c. 140 km north of Tabou, 10 Oct 1963, W.J.J.O. de Wilde 1054 (K); Abidjan, 24 Aug 1955, de Wit 801 (WAG); Adiopodoumé, Oct 1960, F. Hallé 188 (K); ibid., Oct 1960, F. Hallé 198 (BR); Abouabou, 9 Jan 1959, Leeuwenberg 2348 (K); N Sékré, Oldeman 603 (K); Yapo, 28 Oct 1954, Roberty 15338 (K). LIBERIA. Nimba, 8 Jan 1965, Adam 20500 (K); near Tobli on the Tapeta–Chien road, 14 Jan 1967, Bos 2769 (BR); Sinoe to port, 17 Jan 1969, Jansen 1129 (BR); Gibi Mountain, 5–10 miles south-east of Salala, 15 Jan 1970, Jansen 1708 (BR); Bendu, 10 miles north of Robertsport, 30 Jan 1970, Jansen 1761, (BR); 4 miles north of Monrovia, 16 Nov 1970, Jansen 2296 (BR); 6 miles from to Monrovia, 1904, Johnston s.n. (K); Monrovia, 12 Nov 1926, Linder 1539 (K); Gola–Yoma National Forest, 12 Nov 1965, van Meer 274 (BR); 27 km south of Zwedru, 22 Feb 1966, van Meer 481 (BR); Paynesville, 8 miles east of Monrovia, 23 Oct 1960, Voorhoeve 95 (BR). NIGERIA. Unnokoo, 5 Nov 1967, Ariwaodo 1080 (K); Epe, 28 Nov 1994, Daramola 589 (K); Milken hill, Enugu, 11 Jul 1964, Daramola 55162 (WAG); Ipe–Ikum road, 15 May 1978, Daramola & Ihe 420 (WAG); Shasha Forest Reserve, 10 Nov 1961, Emwiogbon 43535 (WAG); Lagos, Oct 1905, Foster 25 (K); boundary of Nigerian College site, Ibadan (site of new sports field), Nov 1960, Hambler 1100 (BR); road from Imesi–Igtayo, 23 Oct 1972, Latilo & Fagbeni 67515 (WAG); Enugu, 21 Feb 1973, Latilo & Oguntayo 67624 (WAG); Kwanya, Dec 1892, Millen 6 (K); Unekpara road, 31 Jan 1974, Odewo 67030 (WAG); Akpaka Forest Reserve, 20 Nov 1958, Okele & Latilo 38439 (WAG); Ibadan, south reserve, 14 Dec 1943, Onochie 7495 (P); Gambari Forest Reserve, south-west Ibadan, 16 Jan 1958, Onochie & de Wit 678 (WAG); W Lagos, Rowland s.n. (P); Udi Forest Reserve (Enugu), Oct 1928, Smith 9 (K); Eket, 1913, Talbot 3300 (K); Oban, Talbot 286b (K); Owena Forest Reserve, 28 Oct 1948, Ujor 23920 (P); Ibadan, 16 May 1966, van Eijnatten 1516 (WAG); s.loc., van Meer 989 (WAG); SIERRA LEONE. s.loc., Afzelius s.n. (BM); s.loc., Don s.n. (K000412059); Mount Aureol, Freetown, 2 Apr 1967, Gledhill 531 (BR); Freetown, 24 Jan 1883, Johnston 50 (K); Leicester Peak, 4 Dec 1891, ScottElliott 3898b (K); Nyungeru, 5 Jan 1914, N.W. Thomas 7232 (K); Bo, Jan 1914, N.W. Thomas 7418 (BR); Komana, 10 Jan 1914, N.W. Thomas 7648 (K). 2. CUVIERA ANGOLENSIS WELW. EX K.SCHUM., Nat. Pflanzenfam. 4(4): 94. 1891. Type: ANGOLA. Golungo Alto, Alta Queta, Apr 1855, Welwitsch 2564 (lectotype designated here: BM000903514!; syntypes: BM000903515!, BM000903516!, K000412037!, K000412038!, LISU208621!, LISU208622!, LISU208623!, M0106328!). Notes: Cuviera angolensis is closely related to C. longiflora, in particular in the long-exserted hairy style. However, it is hard to understand why Hepper & Keay (1963) synonymized the two species, which have different calyces (C. angolensis: three or four narrowly lanceolate and acute lobes; C. longiflora: five broadly elliptic and rounded lobes). The corolla lobes are also generally longer and narrower in C. longiflora. Cuviera angolensis is, however, closely related to C. latior and, after reviewing all available material of both taxa, we concluded that the latter is a subspecies of the former. The only distinction between them concerns the calyx lobes (see Key). The two subspecies also have separate ranges. The specimen McPherson 16297 referred to as C. angolensis in previous phylogenetic studies (Lantz et al., 2002; Lantz & Bremer, 2004, 2005) is actually C. physinodes. Several Welwitsch collections made on different dates were given the same number 2564, as is usual with this collector (Albuquerque, Brummit & Figueiredo, 2009). The lectotype chosen here was collected in April 1855 and a complete description of the new species is attached to the specimen, hence the lectotypification. 2A. CUVIERA ANGOLENSIS Welw. ex K.Schum. subsp. ANGOLENSIS. Distribution: Congolia Domain: north-west Angola, Democratic Republic of the Congo (Bas–Congo). Additional specimens examined: ANGOLA. Cuanza Sul, Libollo Hochland, 1954–1955, Boss 5755 (BM); Calandula Mountain, Libulo, 18 May 1915, Gossweiler 6314 (BM). DEMOCRATIC REPUBLIC OF THE CONGO. M’vuazi, Malanga Forest, 23 Sep 1951, Devred 727 © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS (BR); M’vuazi, Mansiesie Forest, 25 Sep 1951, Devred 755 (BR, K); Massif of Matete (Mvuazi); Thysville, 15 Jun 1954, J. Dubois 23 (BR); Nzengi Nzengi forest, 1 Aug 1958, J. Dubois 328 (BR, K); Kisantu, 1900, Gillet 1076 (BR); near N’Gidinga, bank of Mosi river, Sep 1923, Gillet s.n. (BR, K). 2B. CUVIERA ANGOLENSIS SUBSP. LATIOR (WERNHAM) SONKÉ, O.LACHENAUD & DESSEIN stat. nov. Basionym: Cuviera latior Wernham, J. Bot. 56: 311. 1918. syn. nov. Type: DEMOCRATIC REPUBLIC OF THE CONGO. N Boyeka, 31 Aug 1914, Nannan 139 (lectotype designated here: BR0000008829580!; isolectotypes: BM000903511!, BR0000008829665!, K000412047!). Synonyms: Cuviera latior var. evorombila N.Hallé, Bull. Soc. Bot. France 106: 344. 1960. syn. nov. Type: GABON. Evorombil, 4 Jul 1934, Le Testu 9627 (holotype: P!; isotypes: BM!, BR!, MO!, WAG!). Cuviera latior var. hispidula N.Hallé, Bull. Soc. Bot. France 106: 344. 1960. syn. nov. Type: CENTRAL AFRICA REPUBLIC. 60 km south-east of Bambari, Gbongo river, Pudjeyo, 15 Jan. 1928, Tisserant 2404 (holotype: P!; isotypes: BM!, BR0000008846846!, BR0000008829672!, K000412046!, MO-391362!, P00553439!). Distribution: Lower Guinea and Congolia Domains: north-east Angola, Central African Republic, Democratic Republic of the Congo (widespread except in south), north-east Gabon (Ivindo basin), Republic of Congo. Notes: Unlike the rather uniform subsp. angolensis, subsp. latior is variable, especially in the width of the calyx lobes (some specimens approaching subsp. angolensis) and in the size of the corolla lobe appendages that vary from 6 mm long to nearly lacking. The leaves are sometimes shortly pubescent on the nerves beneath, which is unique in Cuviera. This character may vary on a single specimen, so Hallé’s (1960) var. hispidula has not been maintained, nor var. evorombila (separated only on account of its less pubescent style). The tertiary veins are typically obscure, but in a few specimens (e.g. Gilbert 14345, M. Laurent 669) they are apparent and rather dense. Additional specimens examined: ANGOLA. Dundo, 2 Jun 1948, Gossweiler 14023 (BM, K, P). CENTRAL AFRICAN REPUBLIC. near Bambari, 6 km on Alindao road, 20 Dec 1963, Descoings 11840 (MPU); hill near Bangui, 5 Jan 1961, Guigonis 2031 (P); 100 km south of Yalinga, road to Bangassou, 24 Nov 1922, Le Testu 4386 (BM, BR, P); Kpalata river, 419 Tisserant 437 (P); Boukoko, 14 Jan 1948, Tisserant 614 (BM, P); Boukoko, 8 Mar 1951, Tisserant 2030 (BM, P); Gbondo river, Pudjiya, 15 Jan 1928, Tisserant 2483 (P); Mbaïki, 5 Jan 1961, Tisserant 3692 (P). DEMOCRATIC REPUBLIC OF THE CONGO. Epulu, 27 Feb 1998, Amsini 097 (BR); Island of Elelwa, 19 Apr 1960, Bamps 932 (BR, K); Metmas, 13 Jan 1915, Bequaert 160 (BR); Molima (Yambuya), Bequaert 1358 (BR); Penghe, Bequaert 2461 (BR); Masisi–Walikale, 31 Dec 1914, Bequaert 6429 (BR); Manyema, 1909, Berger s.n. (BR); 5 km west of Kisangani, 9 Feb 1971, Bokdam 3085 (WAG); ibid., 1 May 1971, Bokdam 3180 (WAG); Yangambi–Yaselia road, 25 Jun 1963, Bolema 1144 (BR); Yangambi, 17 Jan 1961, Bolema 343 (BR); s.loc., Boone 57 (BR); Namoya, Kabazimba to north, Bytebier 232 (BR); Hemptinne St Benediktus, 1911, Callewaert s.n. (BR); Mutakato, km 120 Kavumu–Walikale road, 10 Aug 1956, Christiaensen 1818 (BR, K); Eala, Corbisier-Baland 1399 (BR); ibid., Corbisier-Baland 1524 (BR); ibid., Corbisier-Baland 1717 (BR); Gangala na Bodio, 24 Mar 1943, Cornet D’ Elzius Du Chenoy, Lauwers & Offermann 808 (BR); Yangambi, 1948, Gilbert 8424 (BR); Eala, 10 Sep 1937, Coûteaux 325 (BR); Kiyaka, 11 Jul 1955, Devred 2140 (BR); Bas-Uele, 5 Oct 1934, Dewulf 203 (BR); ibid., 27 Dec 1934, Dewulf 523 (BR); Yangambi, 3 Oct 1950, Donis 2847 (BR); ibid., 8 Jan 1952, Donis 3336 (BR); ibid., 30 Jan1952, Donis 3501 (BR); Boyagidigba, 23 Apr 1955, Evrard 755 (BR); Mangala (Terr. Isangi), 19 Mar 1957, Evrard 2252 (BR); Befale, 2 Jan 1958, Evrard 3179 (BR, K); Eandja, 13 Feb 1958, Evrard 3423 (BR, K); Djolu (Bolomba), 20 Feb 1959, Evrard 5791 (BR, K); [Yionda], 15 km south of Mbandaka (Zaire), 18 Feb 1987, Evrard 10688 (BR); Epulu, 16 Nov 1993, Ewango 199 (BR); s.loc., Flamigni 6122 (BR); km 37 Elundu-Kindu, 22 Jul 1957, Gaillez 65 (BR); Bambesa, 12 Feb 1953, Gerard 552 (BR); Likati, 22 Mar 1956, Gerard 2212 (BR); Madabu (Zobia), 6 Dec 1956, Gerard 2570 (BR); Bambesa, 31 Jan 1958, Gerard 3232 (BR); Bambesa, 12 Nov 1957, Gerard 3300 (BR); Tukpwo, 12 Feb 1958, Gerard 3540 (BR); Bambesa, 8 Jan 1960, Gerard 4015 (BR); ibid., 14 Nov 1963, Gerard 5706 (BR); around Yaosuka, Yangambi, May 1937, Gilbert 26 (BR); ibid., May 1937, Gilbert 26 (BR); s.loc., Mar 1938, Gilbert 974 (BR); Yaosuka, Gilbert 1341 (BR, K); ibid., Gilbert 1410 (BR); Yangambi, INEAC Forest Reserve, 1948, Gilbert 9046 (BR); Yangambi, 1949, Gilbert 9961 (BR); Botaka, Lake Leopold II, 4 Jul 1953, Gilbert 14345 (BR); Beni, Gille 255 (BR); Bikoro, Dec 1920, Goossens 2321 (BR); near Lula, Apr 1921, Goossens 2511 (BR); Lisala, Mar 1924, Goossens 4641 (BR); Busanga, Feb 1953, Gorbatoff 78 (BR); Manyema, Mutongo, 30 Oct 1957, Gutzwiller 3321 (BR); Epulu, 6 Apr 1991, Hart 1114 (BR, K); ibid., 27 Feb 1983, Hart 382bis (BR); © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 420 B. VERSTRAETE ET AL. Bambesa, 11 Jan 1940, Hendrickx 869bot (BR); Bolima, 31 Oct 1941, Hulstaert 471 (BR); Bokote, 1943, Hulstaert 953 (BR); Boende, 28 Jul 1944, Hulstaert 1354 (BR, K); s.loc., Jans s.n. (BR); bank of Congo river, near Bolombe, 2 Jan 1904, M. Laurent s.n. (BR); Eala (Ikakéma), 29 Jan 1904, M. Laurent s.n. (BR); Isangi, 13 Jan 1904, E. Laurent & M. Laurent 11 (BR); near Lié, 7 Jan 1904, M. Laurent 13 (BR); Eala, 1905, M. Laurent 669 (BR); ibid., 10 Oct 1906, M. Laurent 1216 (BR); ibid., 28 Oct 1906, M. Laurent 1256 (BR); ibid., Jun 1906, M. Laurent 1571 (BR); Yaligimba, 1988, Le Jeune 60 (BR, K, WAG); Bamania, Aug 1930, Lebrun 890 (BR); between Libenge and Gemena, Lebrun 1871 (BR, K); ibid., Lebrun 2186 (BR); between Niangara and Wamba, Jul 1931, Lebrun 3228 (BR); between Walikale and Kalehe, Mar 1932, Lebrun 5240 (BR); Urega, Jun 1932, Lebrun 5634 (BR); between Kindu and Katakokombe, Lebrun 6073 (BR, K); Eala, 1936, Leemans 295 (BR); ibid., 1936, Leemans 475 (BR); Kisangani, 500 m SE Kabondo, 30 Apr 1977, Lejoly 1424 (BR); Kisangani, near bac of Simi-Simi, Jun 1977, Lejoly 1928 (BR); Kisangani, Island of Mayele near Wagenia, 13 May 1978, Lejoly 3572 (BR); Kisangani, 13 Nov 1981, Lejoly 81/289 (BR); Kabunga, 17 Feb 1958, A. Léonard 1717 (BR); Kamisuku, 18 Aug 1959, A. Léonard 5946 (BR); between Coquilhatville [= Mbandaka] and Ileko, 15 Nov 1946, J. Léonard 1020 (BR); Kikwit, 28 Jul 1904, Lescrauwaet 165 (BR); road Hemptinne-Kamwandu (Terr. Dibaya), 21 Jun 1957, Liben 3197 (BR); River Kateba (Terr. Luisa), 16 May 1957, Liben 3314 (BR); Kisangani, Tshopo, 22 Nov 1972, Lisowski 15482 (BR, K); Ibid., 28 Jan 1973, Lisowski 16441 (BR, K); Haut Zaïre, plantation Lale–Ekili, 14 Nov 1976, Lisowski 43225 (BR); Kisangani, Tshopo, 22 Nov 1972, Lisowski B9565 (BR); Yangambi, 13 Dec 1935, J. Louis 823 (BR, K, P); île ‘Esali’, en face de Yangambi, 13 Feb 1936, J. Louis 1255 (BR, K); north-west Eala, 20 May 1936, J. Louis 1976 (BR); Yangambi, 26 Jun 1937, J. Louis 4249 (BR); ibid., 29 Nov 1937, J. Louis 6810 (BM, BR); ibid., 12 Jul 1938, J. Louis 10309 (BR, K, P); Island of Esali, 15 Oct 1938, J. Louis 11794 (BR); Yangambi, île Yalututcha, 15 Nov 1938, J. Louis 12583 (BR, K); Opala, J. Louis 14137 (BR); Île Lotumba, 5 Apr 1939, J. Louis 14483 (BR); Lokutu, 29 Oct 2004, Luke 10412Z (BR); Likimi, 13 Feb 1910, Malchair 96 (BR); Kikwit/Erco, 21 Nov 1990, Masens 569 (BR); Bikoro, 8 Aug 1983, Nsola 229 (BR); Mabali, 7 Apr 1985, Nsola 845 (BR); Kapanga, 1934, Overlaet 1216 (BR); km 82 road Kavumu–Walikale, 27 Sep 1955, Pierlot 896 (BR); Tshinganda, Mount Kahusi, km 42 Kavumu–Walikale, 13 May 1959, Pierlot 2904 (BR); Irebu, May 1904, Pynaert 397 (BR); Grelco, Quarré 2595 (BR); Kikwit, 29 May 1946, Renier 35 (BR); Dundusana, Feb 1913, Reygaert 11 (BR); near Mobwasa, Dec 1913, Reygaert 1464 (BR); Gandajika, 7 Nov 1957, Risopoulos 698 (BR); Eala, 25 Dec 1905, Seret 392 (BR); near Eala, Seret 392BIS (BR); Eala, Nov 1930, Staner 1538 (BR, P); road Weko– Bengamisa, between Weko and Yalibutu, 14 Mar 1988, Szafranski 1345 (BR); Kisangani, Tshopo area, km 8 old road to Buta, 5 Jul 1988, Szafranski 1439 (BR); Mobwasa (Itimbiri), 30 Jan 1909, Thonner 126 (BR); La Kulu, Van den Brande 340 (BR); s.loc., Van Der Gucht 131 (BR); Saint Trudon, Van Kerkhoven 824 (BR); ibid., 18 Aug 1913, Van Kerkhoven s.n. (BR); Ipamu, Jul 1922, Vanderyst 9804 (BR); ibid., Sep 1921, Vanderyst 10607 (BR); ibid., Sep 1921, Vanderyst 10907 (BR); ibid., Oct 1922, Vanderyst 12293 (BR); between Lubue 1 and Luange, Oct 1922, Vanderyst 12486 (BR); Hemptinne Saint Benoît, Vanderyst 23677 (BR); Ibid., Vanderyst 23644/4 (BR); Mobwasa, 28 Apr 1914, Vermoesen 324 (BR); Bambesa, 11 Jan 1940, Vrydagh 68 (BR). GABON. Ivindo, Makokou, 20 Nov 1985, Caraglio 56 (MPU); Ipassa, 10 km south of Makokou, 21 Jul 1978, Florence 1556 (P); Bélinga, Basse Nounah, 1 Dec 1964, N. Hallé 3381 (P). REPUBLIC OF THE CONGO. Dongou, 28 Jan 1966, Bouquet 2099 (P); Odzala National Park, camp Mboko, 9 Dec 1993, Diafouka 558 B (BRLU); ibid., 17 Nov 1994, Dowsett-Lemaire 1826 (BR); ibid., 21 Nov 1996, Lejoly 96/753 (BRLU); Kassa, 17 Dec 1926, Linder 1726 (P); bank of Lekoli river, 6 Feb 1994, Lisowski c-790 (BRLU, K). 3. CUVIERA LE-TESTUI PELLEGR., Bull. Soc. Bot. France 81: 142. 1934. Type: GABON. Tchibanga, 13 Apr 1925, Le Testu 2053 (lectotype designated here: P00553438!; isolectotypes: BM!, BR0000008846822!, BR0000008846839!, K000412042!, LISC002648 n.v., P00553436!, P00553437!). Distribution: Lower Guinea Domain: Deomacratic Republic of the Congo (Bas–Congo), Equatorial Guinea (Rio Muni), Gabon (Libreville area, Doudou Mountains, Chaillu Mountains), Republic of Congo (Mayombe). Notes: Cuviera le-testui is an aberrant species standing well apart from its congeners in having a pair of large bracts at the apex of the peduncle and a bilocular ovary. Another remarkable character is the red exudate in the bark mentioned by two different collectors (A.M. Louis 1684 and Senterre & Obiang 4315). This is exceptional in Rubiaceae, although it also occurs in a new species of Psydrax Gaertn. (currently under description). Cuviera le-testui is newly reported from the Republic of Congo and Equatorial Guinea. Material from the © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS latter country is sterile, but the identification makes little doubt. Additional specimens examined: DEMOCRATIC REPUBLIC OF THE CONGO. Kimbuya–Kingedi road, 24 Aug 1959, Compère 154 (BR, K). EQUATORIAL GUINEA. Rio Muni: Monte Bata, 1 km north of road Bata–Niefang at Santa Marta, 8 Aug 2003, Senterre & Obiang 4297 (BRLU); ibid., 9 Aug 2003, Senterre & Obiang 4315 (BRLU). GABON. Mourindi, 15 Sep 2000, Bourobou Bourobou 247 (BR, K, P, WAG); Etéké, 15 May 1963, N. Hallé & Cours 5907 (P); Tchibanga, 2 May 1914, Le Testu 1720 (BR, BM, P); Mouloumbi, 1 Aug 1930, Le Testu 8211 (BR); Malibé, 1 km toward west, 7 May 1985, A.M. Louis 1684 (WAG); near Igotchi–Mouenda, 11 May 1997, McPherson 16942 (K, MO); km 23 road south-east of Igotchi–Mouenda, 19 May 1997, McPherson 17047 (BR, K, MO); c. 10 km north of Libreville, 23 Apr 1985, J.M. Reitsma & B. Reitsma 811 (WAG); Mourindi, 16 Apr 2000, Sosef 1312 (BRLU, WAG). REPUBLIC OF THE CONGO. Sounda, bank of Kouilou, 10 Jun 1987, de Foresta 1265 (P); Les Saras, 30 Jun 1989, Dechamps 13234 (BR); around M’vouti, Pounga–Girard road, 10 Jun 1971, Sita 3095 (P). 4. CUVIERA LONGIFLORA HIERN, in D. Oliver, Fl. Trop. Afr. 3: 157. 1877. Type: CAMEROON. Mount Cameroon, Jan 1862, Mann 1211 (lectotype designated here: K000412049!; isolectotype: K000412050!). Distribution: Lower Guinea Domain: Cameroon (widespread in the west, centre and east), Central African Republic (extreme west), Equatorial Guinea (Bioko Island), north-east Gabon (Bélinga), east Nigeria. Notes: Cuviera longiflora is closely related to C. angolensis, but we cannot follow Hepper & Keay (1963) in uniting them; they especially differ in the number of calyx lobes (see C. angolensis) and also in the range. The type of C. longiflora, Mann 1211 (K) has two sheets, one of which is here selected as lectotype. Note that Mann 1211 (P) represents another species, C. acutiflora. The species can be considered as new to Gabon, as the two specimens cited for this country by Sosef et al. (2006) are both misidentified: McPherson 16297 is C. physinodes and McPherson 16332 is C. calycosa. Additional specimens examined: CAMEROON. Sanaga River near Goyoum, 29 Jan 1961, Breteler 963 (WAG); ibid., 29 Jan 1961, Breteler 963 (K, P); 6 km south-west of Yaoundé, trail to Eloumden Mountain, 29 Dec 1962, Breteler et al. 2318 (BR, K, P, WAG); 421 Kupe village, 24 Jan 1995, Cable et al. 764 (K); Nyasoso, 6 Feb 1995, Cheek et al. 7305 (K); Bafut– Ngemba Reserve, 12 Mar 1959, Daramola 40537 (K, P, WAG); Mount Kupe, above Nyasoso, 24 Apr 2009, Dessein et al. 2713 (BR, MO, WAG, YA); Mount Kupe, Etube village, 2 Feb 1995, Elad 101 (K); Aboh, Etuge 3410 (K); Laikom, 3 Dec 1998, Etuge 4538 (K); Ndikiniméki, 24 Dec 1927, Hédin 3 (P); Nkambe, 10 miles west, 11 Feb 1958, Hepper 1916 (K, P); Mount Bama, Jan 1939, Jacques-Félix 2986 (P); ibid., Jan 1939, Jacques-Félix 3021 (P); Koutchankap, Feb 1939, Jacques-Félix 3042 (P); Bafut–Ngemba Forest Reserve, 7 May 1960, Keay 37925 (BR, K, YA); Kupe village, 17 Jan 1995, Lane et al. 298 (K); Mount Kupe, Ndum, 31 Jan 1995, Lane et al. 460 (K); km 17 Betaré Oya–Meiganga, 5 km south-east of Ndokayo, 3 Nov 1966, Leeuwenberg 7697 (BR, K, P, WAG, YA); bank of Sanaga, between Poute and Ebaka, 1 Feb 1960, Letouzey 2901 (BR, P, YA); river Sanaga, N Goyoum, 20 Jan 1961, Letouzey 3304 (BR, P, WAG, YA); near Ngoila, 7 Feb 1973, Letouzey 11903 (BR, HBG, P, WAG, YA); track from Acha–Abaw towards lake Oku, 40 km north-east Bamenda, 5 Dec 1974, Letouzey 13450 (P, YA); Tinachong, 30 km west-north-west of Bamenda, 10 Aug 1975, Letouzey 14234 (BR, K, YA); bank of Boumba river, 14 km west-south-west of Kinsassa, 7 Mar 1971, Letouzey & Villiers 10525 (BR, P, YA); Mount Oku, Lumutu Forest, 1 May 1998, Maisels 116 (YA); Lakom, Bamenda, May 1938, Maitland 1361 (K); Mount Cameroon, Jan 1862, Mann 1212 (K); Mapanja, 16 Mar 1992, Mbani 55 (K); between Yaoundé and Deng Deng, c. 170 km northwest of Yaoundé, 1 Mar 1914, Mildbraed 8410 (HBG); Elak, 6 Nov 1996, Munyenyembe et al. 892 (K, YA); 80 km de Bertoua, route Ndemba II, 17 Jan 1956, Nana 431 (BR, P, YA); Ekona, Mount Cameroon, 14 Jan 1985, Nkongmeneck 892 (YA); Bafut–Ngemba Forest Reserve, 19 Mar 1955, Richards 530 (K, P); Small Massaka (Rumpi hills), 19 Dec 2009, Sonké et al. 5442 (BR, BRLU, K, MO, P, WAG, YA); National Park Mbam Djerem, 23 Jan 2011, Sonké et al. 5510 (BR, BRLU, K, MO, YA); Bertoua–Batouri, 15 May 1905, Tchinaye 111 (P, YA); south slope of Mount Cameroon, above Batoke, 29 Dec 1983, D.W. Thomas 2844 (BR, K, P, YA); above small Koto village, 6 Mar 1985, D.W. Thomas 4463 (MO, P, WAG, YA); Bafut– Ngemba Forest Reserve, 24 Jan 1951, Tiku 22247 (K); Bali–Nguemba Forest Reserve, 5 Jun 1951, Ujor 30416 (K); east slope Mount Cameroun, 6 km east of Bomana, 34 km north-west of Limbe, 11 Dec 1984, Villiers 2447 (BR, YA); Mapanja, 19 Feb 1992, Wheatley 14 (K). CENTRAL AFRICAN REPUBLIC. near Baboua, 3 km along the road to Besson, Descoings 12703 (MPU); Berbérati, 28 Feb 1937, Tisserant 3478 (BM, P). © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 422 B. VERSTRAETE ET AL. EQUATORIAL GUINEA. Bioko: Gran Caldeira de Luba, along the river towards Mount Pisarro, Carvalho 4268 (K); s.loc., Fernandez Casas 12167-19 (K). GABON. Bélinga hills, main ridge, 22 Nov 2007, Leal et al. 2169 (BR, MO). NIGERIA. Boshi extension Forest Reserve, 23 May 1971, van Meer 1771 (WAG). 5. CUVIERA MACROURA K.SCHUM., Bot. Jahrb. Syst. 33: 352. 1903. Type: NIGERIA. Lagos, Millen 159 (holotype: K000412062!). Distribution: Lower and Upper Guinea Domains: Benin, Cameroon (only along coast), Ghana, Guinea, Ivory Coast, Liberia, Nigeria, Sierra Leone; a doubtful record from Equatorial Guinea (Bioko). Notes: Cuviera macroura is unique in the genus in having small glabrous pits in the nerve axils (sometimes hard to see), a calyx with subulate lobes distinctly fused at the base, and stipules with a broad rounded lobe (see photographs in Hawthorne & Jongkind, 2006). The species is rather variable, specimens from Cameroon and Nigeria generally have smaller leaves and fruits than those from West Africa. Vogel 83, the type of C. subuliflora, has two sheets, one of which is C. macroura (K000412044!). Both sheets are labelled ‘Fernando Po’ [= Bioko], but their origin is somewhat doubtful, for Bentham (in Hooker, 1849) states in the protologue of C. subuliflora: ‘Fernando Po, on the seashore, and, apparently the same species, at Abòh, growing in the water, Vogel’. The latter locality is in Nigeria. There are no other records of C. macroura on Bioko and its occurrence there is to be regarded as doubtful. Cuviera djalonensis A.Chev. (in Explor. Bot. Afrique Occ. Franç.: 331. 1920.) is a nomen nudum referring to C. macroura (Hallé, 1960). Additional specimens examined: BENIN. Lokoli, 18 Dec 2002, Dan 674 (BR); Lokoli/Sisimè, 20 Dec 2002, Dan 684 (BR); Godomé, 20 Aug 1902, Estéve 115 (BM). CAMEROON. Right bank of Lokoundjé river, near Edéa, 26 km north of Kribi, 18 Apr 1965, Leeuwenberg 5620 (BR, K, P, WAG); left bank of Sanaga river, near of the mouth, 6 Jan 1974, Letouzey 12631 (YA); bank of Abo river, between Miang and Koki, 25 km north of Douala, 9 4 1976, Letouzey 14751 (BR, P, YA); Douala–Edéa Reserve, bank of Kombe river, 3 Aug 1978, D.W. Thomas 521 (K). EQUATORIAL GUINEA. Bioko: locality doubtful (see Notes above), Vogel 83 (K000412044). GHANA. Boin River Forest Reserve, 2 Jul 1956, Agbley 6263 (K); Obuasi, Jun 1936, Andoh 4218 (BR, K); near Anyinakim, 13 Jul 1954, Darko 898 (K); ± 5 miles inland from Dixcove, 31 Mar 1954, Morton A468 (K); Kumasi, Jun 1929, Vigne 1758 (K); s.loc., s.d., Vigne 3008 (BM). GUINEA. Gouam–Geasso, 23 Apr 1956, Adam 12092 (K); Sérédou, Aug 1954, Jacques-Félix 7108 (K). IVORY COAST. Lower Cavally, Prolo, 11 Aug 1909, Chevalier 19869 (K); Reserve of Boulay Island, Ebrié, 18 Nov 1956, J.J.F.E. de Wilde 818 (WAG); between Mount Nienokoue and Hana river, Taï–Tabou road, 11 Aug 1962, Guillaumet 1519 (BR); Adiopodoumé, 16 Aug 1961, F. Hallé 219 (BR, K); near the debarcation of the ferry from Grand Bassam to Bingerville, 24 Jul 1969, Versteegh & Den Outer 575 (BR). LIBERIA. Zeahtown, 1 Aug 1947, Baldwin 6967 (K); 10 miles west of Tapita, along the road to Ganta, 16 Jul 1968, Jansen 864 (BR, K); Peahtah, 13 Oct 1926, Linder 1035 (K). NIGERIA. s.loc., Hambler 409 (K); Ibarapa, 22 Aug 1966, Latilo 58826 (K, P); 19 miles from Imo river, 12 Aug 1966, Okafor 60323 (K); Ikorodu, 29 Dec 1952, Onochie 32038 (K); Degema, 1916, Talbot 3632 (BM, K). SIERRA LEONE. Mussaea, 31 Aug 1963, Haswell 150 (K); Loma Mountains, 20 Sep 1945, Jaeger 1959 (K); Tingi, 18 Dec 1965, Morton & Gledhill SL3223 (K); Binkolo, 25 Aug 1914, N.W. Thomas 1857 (K); Bumbuna, 14 Oct 1914, N.W. Thomas 3189 (K); Komorolai, 23 Dec 1915, N.W. Thomas 6914 (K). 6. CUVIERA PHYSINODES K.SCHUM., Pringsh. Jahrb. 19: 391. 1888. Type: GABON. Sibange Farm, Munda, 27 Aug 1879, Soyaux 29 (lectotype designated here: P03818077!; isolectotype: K000412041!); Sibange Farm, Munda, 25 Oct 1881, Soyaux 307 (syntype: K000412040!). Synonyms: Cuviera ledermannii K.Krause, Bot. Jahrb. Syst. 48: 418. 1912. syn. nov. Type: CAMEROON. Nkolebunde [= Elephant Mountain], Oct 1908, Ledermann 725, 751 (holotype: B†); Elephant Mountain, c. 20 km south-east of Kribi, 9 Mar 2007, Chatrou et al. 572 (neotype designated here: BR000000980675!). Distribution: Lower Guinea Domain: south Cameroon (north to Lake Tissongo), Equatorial Guinea (Rio Muni), Gabon (widespread except in the east). Notes: This species has been overlooked since its description, possibly because the Index Kewensis gives a wrong reference for the protologue. Cuviera physinodes is closely related to C. acutiflora and frequently mistaken for that species (e.g. Sosef et al., 2006); see key for differences in flower and fruit. Sterile specimens from the area of sympatry (Cam- © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS eroon, Equatorial Guinea) can be set apart from C. acutiflora by their cuneate leaf base (always cordate in local populations of C. acutiflora). The description of C. physinodes is based on two syntypes, Soyaux 29 (with flower buds) and Soyaux 307 (with young fruits). The P sheet of the former bears the number 23, which is evidently a copying error: Soyaux 23 (K) was collected on 21 Aug and is an isosyntype of Pavetta brachycoryne K.Schum. (= Rutidea glabra Hiern). In spite of this, we select this sheet as lectotype, as it is by far the better. Krause (1912) already considered C. ledermannii as closely related to C. physinodes and his good description actually leaves no doubt that they are conspecific. The syntypes have probably been destroyed in Berlin, so we designated a neotype from the same locality. Additional specimens examined: CAMEROON. Lobe river, above the Grand Batanga ferry, 24 Oct 1968, Bos 3081 (BR, K, P, WAG, YA); 13.5 km from Kribi, north of Ebolowa road, Bidou II plantation, 2 Dec 1968, Bos 3426 (BR, K, P, WAG, YA); near Mbikiliki village, 11 Mar 2007, Dessein & Sonké 1513 (BR, YA); near Bidjouka, track towards waterfall, 15 Mar 2007, Dessein & Sonké 1593 (BR, YA); near lake Tissongo, 9 May 1977, McKey 53 (P, YA); Mangombe Forest Reserve (Edéa), 21 Jan 1956, Mpom 183 (BR, YA); near Ebodje, 50 km south of Kribi, 29 Oct 1984, Nkongmeneck 822 (YA); Nkoltsia hill, 24 Apr 1974, Villiers 870 (YA); Bipindi, 1898, Zenker 1797 (BM); ibid., 1900, Zenker 2246 (BM, K, P). EQUATORIAL GUINEA. Rio Muni: Monte Alén, 19 Apr 2001, Ngomo 916 (BRLU); south-west part of Monte Alén National Park, 2 km north-east of Rio Uolo, 24 June 2002, Senterre & Nguema 3018 (BRLU); Monte Alén National Park, 12.5 km east of Mosumo, 8 Sept 2003, Senterre & Obiang 4071 (BRLU). GABON. Toucan, 11 Oct 2002, Bourobou et al. 990 (WAG); km 34 Mouila–Yeno, 22 Sep 1986, Breteler et al. 8100 (BR, K, P, WAG); Sibange (Monda), 1884, Büttner 152 (K); Monts de Cristal, inselberg Milobo, 10 km north of Mbé Akélayong, 30 Nov 2001, Degreef 181 (BR); Benga, Klaine 82 (BR, K, P); Owendo, 10 Nov 1903, Klaine 3340 (K, P); road to Cap Estérias on the right, 11 Sep 1985, A.M. Louis 1805 (WAG); Forêt des Abeilles, east Lopé Reserve, 3 Dec 1993, McPherson 16297 (K, MO, WAG); c. 25 km north of Libreville, 13 Jul 1985, J.M. Reitsma & B. Reitsma 1261 (WAG); Mondah Forest, c. 15 km north of Libreville, 30 Jul 1986, J.M. Reitsma & B. Reitsma 2463 (WAG); Doudou Mountains National Park, c. 30 km south of Mandji, 18 Nov 2005, Sosef 2373 (WAG); Pechoud road, southwards, 27 Oct 1990, Van Nek 123 (WAG); Cap Esterias, 1896, Klaine 538 (P); Mondah Forest, 30 Aug 1969, Villiers s.n. (P). 423 7. CUVIERA PIERREI N.HALLÉ, Bull. Soc. Bot. France 106: 344. 1960. Type: GABON. La Nkoulounga (north-east of Libreville), 7 Jul 1959, N. Hallé 730 (holotype: P00553435!; isotypes: K000412048!, P00553434!). Distribution: Lower Guinea Domain: Equatorial Guinea (Rio Muni), Gabon. Notes: This species is unusual in Cuviera in having (two-)three- (five-) flowered inflorescences and the style thickened in the upper two-thirds (Hallé, 1960: fig. 2L). It is presumably related to C. subuliflora, which has similar leaves. Cuviera pierrei also shows a superficial resemblance to Globulostylis cuvieroides, because of the shape of the calyx lobes, however, the striped corolla lobes, the position of the bracts (inserted halfway on the pedicels of the lateral flowers rather than at the apex of the peduncle), and the presence of ant holes in the twigs are clear distinctions. The species is newly reported from Equatorial Guinea. A record from the Democratic Republic of the Congo (Compère, 1962) is an error for C. subuliflora (see that species). Additional specimens examined: EQUATORIAL GUINEA. Rio Muni: Abama (Nsork), 22 Aug 2001, Esono, Merino & Gomez 289 (BRLU); Ngoma, 15 km south-east de Etembue, 12 Aug 1998, Lejoly & Elad 98/158 (BRLU). GABON. 9 km south of Kinguélé, 21 Sept 1994, Breteler et al. 12980 (WAG); Mount Mela, 26 Aug 1978, Breteler & J.J.F.E. de Wilde 320 (BR, K, P, WAG); around Libreville, 3 Dec 1899, Chalot 15 (P); Kinguélé road, N. Hallé & Villiers 4588 (P); Libreville, 25 Mar 1904, Klaine 48 (BR, K, P, WAG); ibid., 12 Feb 1896, Klaine 378 (P); ibid., 15 Oct 1896, Klaine 558 (P); ibid., 25 Jul 1897, Klaine 558bis (P); ibid., 24 Oct 1900, Klaine 2005 (P); ibid., Jul 1895, Thollon 216 (BR, P); Cocobeach road, 15 Jul 1986, D.W. Thomas & Wilks 6361 (K); N’Koulounga, 23 Jun 1959, Touzet 157 (P); Ngoualé, 15 km south of Ekorado, 2 May 2001, Walters et al. 582 (BR); Nombo, 22 May 1986, Wilks 1299 (WAG). 8. CUVIERA SUBULIFLORA BENTH., in W. J. Hooker, Niger Fl.: 407. 1849. Canthium subuliflorum (Benth.) Roberty, Bull. Inst. Franç. Afrique Noire 16: 60. 1954. Type: EQUATORIAL GUINEA. Bioko, Vogel 83 (lectotype designated here: K000412045!); Excluded isolectotype: Bioko, Vogel 83 (K000412044!) is Cuviera macroura. Synonyms: Cuviera plagiophylla K.Schum., Bot. Jahrb. Syst. 33: 353. 1903. Type: CAMEROON. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 424 B. VERSTRAETE ET AL. Bipindi, 1897, Zenker 1350 (lectotype designated here: K000412043!, isolectotypes: BM!, BR000000884676!, HBG, L0058090 n.v., MA311021 n.v., NY00131204 n.v., P!, S05-10683 n.v., WAG!). Distribution: Lower Guinea Domain: Cameroon, Democratic Republic of the Congo, Equatorial Guinea (Bioko and Rio Muni), Gabon, Nigeria, Republic of Congo. Notes: The type of C. subuliflora, Vogel 83, has two sheets, one corresponding to the species hitherto named C. subuliflora, the other being C. macroura. To preserve current usage, we select the former sheet (K000412045!) as lectotype. The origin of these collections is somewhat doubtful (see note under C. macroura). The fruits of C. subuliflora, with their broadly truncate apex and the calyx lobes patent and star-like, are unique in the genus. The style can be glabrous or pubescent, which explains discrepancies between the keys in Hallé (1960) and Hepper & Keay (1963). This species is newly reported from the Democratic Republic of the Congo (see C. pierrei above). A record from Ghana (Hepper & Keay, 1963) is probably an error for C. macroura (Hawthorne & Jongkind, 2006: 628), but we could not verify the specimen. Two different Talbot specimens bear the same number 222: one collected in 1911 (C. subuliflora), the other from 1909 (C. truncata). Additional specimens examined: CAMEROON. South of Bakundu, 14 Feb 1956, Binuyo & Daramola 35530 (K, P); Onge, 28 Oct 1993, Cheek et al. 5172 (K); Konye, Kumba–Mamfe road, 12 Sep 1986, Etuge & Thomas 257 (BR, K, MO, P, WAG); Boa, 1 Jun 1994, Fraser 469 (K); Nkongkengui, 12 km north-north-east of Makak, 17 Jul 1972, Letouzey 11524 (K, P, WAG, YA); chutes du Ntem, 40 km east-south-east of Campo, 10 Dec 1979, Letouzey 15342 (P, YA); Bimbia, Feb 1929, Maitland 417 (K); Limbe, Oct 1929, Maitland 740 (K); Fenda, 58 km east of Kribi, 1911, Mildbraed 6009 (HBG); Bibundi, Nov 1928, Mildbraed 10662 (K); Bonjare, 1 May 1994, Ndam 1111 (K); Nndian–Dibonda– Ekumbako road, 24 Nov 1986, Nemba & D.W. Thomas 398 (BR, K, MO, YA); Kumba–Mamfe road, 23 Jul 1987, Nemba & D.W. Thomas 616 (K, MO); Kumba area, 13 Oct 1984, Nemba & D.W. Thomas 4049 (BR); s.loc., Preuss 557 (BM); Meka’a (Dja Fauna Reserve), 12 Feb 1995, Sonké 1430 (BR, BRLU); Djomedjoh (Dja Fauna Reserve), 23 Jul 1995, Sonké 1646 (BR); Bejange, 23 Jan 2008, Sonké 4591 (BR, K, MO, P, WAG, YA); Kumba area, 13 Oct 1984, D.W. Thomas & Nemba 4049 (K, MO, P, YA); 10 km Ikata–Munyenge (northeast of Muyuka), 26 Aug 1983, D.W. Thomas 2542 (BR, K, MO, P, YA); Liwenyi, 31 Oct 1993, Watts 880 (K); Upper Boando, 21 Feb 1992, Wheatley 19 (K); Nkuambe, 1913, Zenker 346 (BR, P, WAG); Bipindi, 1911, Zenker 3985 (BM, BR, HBG, K); ibid., 1913, Zenker 4977 (BM, K); s.loc., Unknown (SCA) 2264 (YA). DEMOCRATIC REPUBLIC OF THE CONGO. Kimetete, 23 Sep 1959, P. Compère 463 (BR). EQUATORIAL GUINEA. Bioko: s.loc., 1986, Carvalho 2283-22 (K); s.loc., 1986, Carvalho 2521-3 (K); s.loc., 1986, Fernandez Casas 11544-3 (K); s.loc., Dec 1860, Mann 87 (K); Rio Muni: Sendje, 13 Aug 2001, Esono & Ndong 304 (BRLU); Monte Alen, Ngomo & Ndong 232 (BRLU); Nkolentangan, 15 Feb 1908, Tessmann 204 (K). GABON. Km 4 Mbigou–Mimongo, 1983, J.J.F.E. de Wilde et al. 545 (BR, K, WAG); Rembo, on Guidouma– Mouila road, 19 Feb 2008, Dessein et al. 2004 (BR, LBV); 23 km south-west of Koumameyong, 3 Aug 1987, Dibata 274 (WAG); Moubigou 2, 21 May 1963, N. Hallé & Cours 6138 (P); around Libreville, 13 Jul 1895, Klaine 302 (P); Libreville, 1904, Klaine 999 (K, P); Ofoubou, c. 30 km west of Mandji, 24 Jun 1991, Reitsma 3728 (WAG); c. 17 km NW Doussala, Monts Doudou Reserve, 25 Mar 2000, Sosef 1004 (WAG); Saint Martin, Jul 1939, Walker 2 (P). NIGERIA. Oban Group Forest Reserve, 11 Sep 1960, Adebusuyi 44000 (K); ibid., 22 Nov 1961, Binuyo 45423 (K); Aponmu Forest Reserve, 7 Aug 1963, Gillett 15279 (BR, K, P, WAG); ibid., Hunder 90 (WAG); Oban, 2 Dec 1911, Talbot 222 (BM, K); ibid., 1912, Talbot 2036 (BM); Degema, Talbot 3383 (BM, K); 12.5 miles Akure-Ondo, 13 Nov 1968, van Meer 892 (BR, K, WAG). REPUBLIC OF THE CONGO. region of Kouilou (Bas–Kouilou), 26 Aug 1990, Moutsamboté & DowsettLemaire 4499 (BR); Les Saras region, Congo–Cabinda road, 8 Jun 1966, Sita 1376 (P); s.loc., Moutsamboté 1401 (BR). 9. CUVIERA TRILOCULARIS HIERN, in D. Oliver, Fl. Trop. Afr. 3: 157. 1877. Type: NIGERIA. Old Calabar, 11 Apr 1863, Thomson 122 (lectotype designated here: K000412061!; isolectotypes: E00193711!, E00259212!). Distribution: Lower Guinea Domain: restricted to extreme south-east Nigeria (Oban) and adjacent south-west Cameroon (Korup National Park). Notes: Hiern (1877) based his specific name on the supposedly trilocular ovary, which seems to have been a wrong observation. We dissected two ovaries (one each from D.W. Thomas 6797 and van Meer 1357) and found them to be pentalocular, as is usual in the genus. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS Cuviera trilocularis is the only species of Cuviera in which ant holes seem to be consistently absent; it is also unusual in its few-flowered inflorescences. The flower colour pattern of C. trilocularis needs checking in the field: the label of van Meer 1357 describes the corolla as ‘greenish white inside’, not mentioning stripes but on the dried specimen the central zone of the lobes is distinctly paler. Additional specimens examined: CAMEROON. Ekundu Kundu, 26 April 1996, Cheek et al. 8186 (YA); Korup National Park, 15 Mar 1986, D.W. Thomas & Mcleod 5783 (K, YA); ibid., Mar 1979, D.W. Thomas 1181 (K); ibid., 26 Feb 1985, D.W. Thomas 4312 (K, YA); Mundemba, 28 Mar 1987, D.W. Thomas 6797 (BR, K, MO, WAG, YA); around Erat village (southwest Korup National Park), 10 Jun 1988, D.W. Thomas 8120 (K, MO, P). NIGERIA. Oban, 10 May 1912, Talbot 221 (BM, K); ibid., 1911, Talbot 272 (BM); Oban, 14 Apr 1971, van Meer 1261 (WAG); Oban Group Forest Reserve (west block, cut line from Pillar 51), 19 Apr 1971, van Meer 1357 (BR, WAG). 10. Cuviera truncata Hutch. & Dalziel, Fl. W. Trop. Afr. 2: 118. 1931. Type: NIGERIA. Oban, 1911, Talbot 286 (lectotype designated here: K000412060!; isolectotype: BM000903509!); Degema, 1916, Talbot 3686 (syntypes: BM000903507!, BM000903508!). Distribution: Lower Guinea Domain: south-west Cameroon, Nigeria. Notes: Cuviera runcata is unique in the genus in having the calyx truncate or with short (< 2 mm) and irregular lobes. It further differs from C. acutiflora, its closest relative, in having larger and broader bracts. However, some Nigerian collections (Dalziel 45 and Elliott 198, both from Lokoja; Kennedy 2758 from Sapoba; Thornewill 47 from Wamba river) are intermediate between the two species, either by combining characters of both (large bracts and long calyx lobes, or the reverse) or by having variable calyx lobes. They could represent hybrids; fieldwork in Nigeria would be needed to solve this question. Another Talbot 222 specimen belongs to C. subuliflora (see Note under that species). Additional specimens examined: CAMEROON. Kupe– Muanenguba, Kodmin, 20 Jan 1998, Etuge et al. 4051 (YA); Eyumojock Forest Reserve (c. 42 km east of Mamfe), 9 Feb 1985, Onana 107 (BR, YA); Mamfe town, 22 Apr 1987, D.W. Thomas 7304 (BR, K, MO, WAG). 425 NIGERIA. s.loc., Akete 64 (K); Ganglani, 23 Nov 1957, Hepper 1448 (BR, K, P); Sanga river, Dogon Kurmi, 30 Nov 1957, Keay 37246 (K, P); Dogon Kurmi, 17 Nov 1946, Keay & Onochie 21535 (K); Gangumi (between Dondere and Yandere), 8 Dec 1954, Latilo & Daramola 28861 (K, P); Nimbia Forest Reserve, 9 Apr 1963, Latilo 47417 (K, P, WAG); Boki (c. 18 miles from Bende/Ayuk), 28 Feb 1973, Latilo 67754 (K, WAG); Dogon Kurmi, 10 Dec 1964, Olorunfemi 55130 (P); Oban, 1909, Talbot 222 (BM). GLOBULOSTYLIS Wernham, Cat. Pl. Oban: 49. 1913. Cuviera subg. Globulostylis (Wernham) Verdc., Kew Bull. 42: 189. 1987. Type species: Globulostylis talbotii Wernham (lectotype designated by Verdcourt, 1987: 189). Description: Small trees or shrubs, 1.5–16.0 m high, evergreen, with regular horizontal branches (Roux’s architectural model); vegetative parts glabrous or rarely shortly pubescent (in some specimens of G. rammelooana and G. robbrechtiana); twigs without ant holes; spines absent. Leaves opposite, ± elliptic; stipules small, basally ± connate, with an awn > 2 mm long (frequently damaged). Inflorescences axillary and generally paired at nodes (or occasionally terminal), on young twigs, cymose, umbelliform, oneto five-flowered, with a single pair of elliptic leafy bracts at apex of peduncle (in G. cuvieroides and G. minor rarely with some additional bracts on the pedicels). Flowers hermaphrodite. Calyx lobes five, almost free, leafy, broadly elliptic or more rarely lanceolate (G. cuvieroides). Corolla green or yellow; tube broadly funnel-shaped; lobes five, triangular, acute to shortly apiculate, spreading or reflexed; outside glabrous or with five rows of hairs on the lobes and upper part of tube; inside with a ring of hairs in the lower half of the tube. Stamens five, inserted in the upper part of the tube, anthers partly to fully exserted. Ovary pentalocular, obconic or turbinate, smooth; ovules solitary in each locule, pendulous. Disk ± flat, glabrous. Style shortly exserted, with a pubescent swelling in the lower part (fusiform in G. dewildeana and G. robbrechtiana, discoid to globose in other species), otherwise glabrous; stigmatic club ± mitriform, shortly five-lobed. Fruit drupaceous, maturing yellow to orange (but generally seen green), large, 1.5–3.5 × 1.2–3.5 cm, subglobose to ellipsoid, smooth to conspicuously five-lobed, with calyx persistent (rarely caducous); pyrenes five, oneseeded, shaped like a mandarin orange segment, with a slight ventral indentation towards the upper third. Distribution and ecology: Globulostylis has eight species in Central Africa, all endemic to the Lower © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 426 B. VERSTRAETE ET AL. KEY 1. 1’. 2. 2’. 3. 3’. 4. 4’. 5. 5’. 6. 6’. 7. 7’. TO THE GLOBULOSTYLIS SPECIES Calyx lobes narrowly lanceolate, > 5 times longer than wide, 7–17 × 1.0–3.5 mm; tree 6–12(−16) m high (south-east Cameroon and Gabon) ..............................................................................G. cuvieroides Calyx lobes broadly elliptic, < 3 times longer than wide; smaller trees or shrubs, 1.5–6 m high ........2 Calyx ciliate on margin; corolla lobes pubescent outside on the midrib ...........................................3 Calyx entirely glabrous; corolla lobes glabrous outside ...................................................................4 Flowers small; calyx 5–7 mm long; corolla tube 4–5 mm, and shorter than the lobes (Nigeria and south-west Cameroon) .....................................................................................................G. talbotii Flowers large; calyx lobes 10–16 mm long; corolla tube 14.5–19.0 mm long, longer than wide, and longer than the lobes (south Cameroon, around Ebolowa) ......................................................G. dewildeana Corolla yellow, with tube 20–30 mm long, at least twice longer than calyx; style with a weak, fusiform basal swelling; fruits strongly five-lobed and rather small, 15–22 × 12–19 mm when dry (Cameroon, Equatorial Guinea, Gabon) ................................................................................... .G. robbrechtiana Corolla green (possibly yellow in C. leniochlamys), with tube 2–15 mm long, not or shortly exceeding calyx; style with a strong, globose to discoid swelling; fruits smooth to weakly five-lobed and generally larger, 19–32 × 17–27 mm when dry ............................................................................................. 5 Calyx large, with lobes 11–22 mm long .........................................................................................6 Calyx smaller, with lobes 4–10 mm long ..................................................................................... ..7 Corolla tube c. 15 mm long, ± equalling calyx; inflorescence three- to five-flowered (but only one fruit may develop) (south Cameroon) ...............................................................................G. leniochlamys Corolla tube much shorter than calyx; inflorescence always one-flowered (Gabon to Democratic Republic of the Congo) ............................................................................................................. ..G. uncinula Corolla small, tube 2–5 mm long, lobes 3–5 mm long; calyx lobes patent (Fig. 5G), 4–7 × 2.5–5.0 mm; leaves drying greyish–green below (Nigeria and south-west Cameroon) ................................G. minor Corolla larger, tube 6–8 mm long, lobes 6–10 mm long; calyx lobes erect (Fig. 5E), 6–10 × 5.5–7.0 mm; leaves drying olive–green to olive–brown below (Cameroon) ................................... ..G. rammelooana Guinea Domain, except G. uncinula, which also occurs in the Congolian Domain. The centre of diversity is in Cameroon (seven species, including three endemics). All Globulostylis spp. are shrubs or small trees of forest understory, never gregarious. Notes: Globulostylis spp. are often difficult to separate in the absence of flowers. The leaves in particular show little variation between the species. The inflorescences are most often axillary, but terminal inflorescences, previously unrecorded in Vanguerieae, are occasionally found in G. robbrechtiana (e.g. Nguema & Parmentier 446, Ngomo & Ndong 439) and G. rammelooana (e.g. Sonké & Simo 4671). As for Cuviera, the pollinators or dispersors of Globulostylis are unknown. There are well-marked differences in flower characters between species with a long corolla tube, a generally weak style swelling, and an internal ring of hairs close to the base (G. dewildeana, G. leniochlamys and G. robbrechtiana) and those with a short corolla tube, a thick style swelling and an internal ring of hairs near the middle of the tube (the remaining five species), so it is likely these two groups have different pollinators. The function of the style swelling is unclear, although a role in preventing nectar robbery has been suggested (Verdcourt, 1987). No human uses of Globulostylis have been recorded. 1. GLOBULOSTYLIS CUVIEROIDES WERNHAM, J. Bot. 56: 313. 1918. Cuviera cuvieroides (Wernham) Onana, Kew Bull. 63: 402. 2008 publ. 2009. Type: CAMEROON. Bitye, 1911, Bates 1016 (holotype: BM000903506!). Synonyms: Cuviera heisteriifolia Mildbr., Notizbl. Bot. Gart. Berlin-Dahlem 9: 205. 1924. syn. nov. Type: CAMEROON. Ebolowa, 17 Jun 1911, Mildbraed 5610 (lectotype designated here: HBG!). Distribution: Lower Guinea Domain: south-east Cameroon, east Gabon. Notes: Globulostylis cuvieroides stands well apart from its congeners in having narrowly lanceolate calyx lobes, a character more reminiscent of Cuviera. For that reason it shows a superficial resemblance to Cuviera pierrei (see that species). An examination of the type of C. heisteriifolia shows this to be conspecific with G. cuvieroides. The original specimen has presumably been destroyed in Berlin, so the HBG duplicate is selected as lectotype. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS 427 Figure 5. Flowers of several Cuviera and Globulostylis spp. A, Cuviera angolensis subsp. latior. B, Cuviera subuliflora. C–D, Cuviera longiflora. E, Globulostylis rammelooana. F, Globulostylis robbrechtiana. G, Globulostylis minor. H, Globulostylis cuvieroides. Photographs by Quentin Luke (A), Steven Dessein (B), Murielle Simo (C–F) and Olivier Maurin (G–H). © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 428 B. VERSTRAETE ET AL. Additional specimens examined: CAMEROON. Lomié region, 1911, Mildbraed 5404 (HBG); c. 15 km southeast of Somalomo (Dja Fauna Reserve), 9 Nov 2002, Davis 3049 (BR, K); Bouamir, Dja Fauna Reserve, Mar 1996, Fogiel 586 (K); Meyos Mela (32 km eastnorth-east Djoum), 9 Nov 1966, Letouzey 8325 (BR, P, YA); Bouamir, 5 Sep 2001, Nguembou 146 (BR); Akom sili-Bali (Dja Fauna Reserve), 28 Dec 2002, Nguembou 544 (BR, BRLU); track from Bouamir station Mbassakok (Dja Fauna Reserve), 20 Apr 2001, Senterre & Kouob 1007 (BRLU); ibid., Senterre & Kouob 1020 (BRLU); ibid., 17 May 2001, Senterre & Kouob 1233 (BRLU); ibid., 20 May 2001, Senterre & Kouob 1408 (BRLU); Djolimpoum (Dja Fauna Reserve), 10 Dec 1994, Sonké 1296 (BR); Mékas (Dja Fauna Reserve), 12 Jan 1995, Sonké 1405 (BR, WAG); Meka’a (Dja Fauna Reserve), 15 Feb 1995, Sonké 1438 (BR); Djolimpoum (Dja Fauna Reserve), 13 Apr 1995, Sonké 1464 (BR); 27 km south of Djolimpoum, 26 Nov 1996, Sonké 1841 (BR, K); Haut-Nyong, Somalomo, 12 km east of Somalomo, 8 Nov 2002, Sonké et al. 2882 (BR, K, YA). GABON. c. 60 km south-south-west of Moanda, 16 Oct 1970, Breteler 6947 (BR, K, P, WAG); Oveng, chantier Rougier–Ocean, 7 May 1985, Breteler & A.M. Louis 859 (WAG); road Mouila–Yéno, 20 Feb 2008, Dessein et al. 2017 (BR, LBV); Bélinga, Mayibout I, 9 Jun 1978, Florence 1220 (P); Levata, 29 Apr 1930, Le Testu 8059 (BM, P); Oveng, km 19 forest exploitation camp, 9 Nov 1983, A.M. Louis et al. 525 (BR, K, P, WAG); eastern border of Lopé–Okanda Reserve, 15 Jan 1993, McPherson 16062 (BR, K, MO, WAG); Oveng, c. 15 km north-west of forestry camp, 8 Feb 1987, J.M. Reitsma & B. Reitsma 2923 (WAG); Oveng, c. 25 km west-south-west of Mintsic, 9 Nov 1986, J.M. Reitsma, B. Reitsma. & Mezui 2571 (WAG); c. 40 km north-east of Mitzic (forestry concession Bordamur), 10 Feb 2003, Sosef et al. 1961 (WAG); Langoue Bai (Ivindo National Park), 27 Nov 2002, Stone & Niangadouma 3529 (BR); Nombo village, 22 May 1986, Wilks 1301 (WAG). 2. GLOBULOSTYLIS DEWILDEANA SONKÉ, O.LACHENAUD & DESSEIN sp. nov. Type: CAMEROON. between N’Kolandom and N’Koemvone, 680 m alt, 2°48′N, 11°09′E, 9 Jan 1975, J.J.F.E de Wilde 7886 (holotype: WAG0114671!; isotypes: BR!, K!, P!, WAG0114672!, WAG0114673!). Diagnosis: Globulostylis dewildeana resembles G. talbotii in its ciliate calyx and dorsally pubescent corolla lobes, but differs in the larger calyx lobes (10–16 mm long vs. 5–7 mm long), the much longer corolla tube (14.5–19.0 mm long vs. 4–5 mm long), and the style with a weak and fusiform basal swelling (vs. thick and subglobose in G. talbotii). Globulostylis dewil- deana also resembles G. leniochlamys and G. robbrechtiana in corolla shape (tube longer than broad and much exceeding lobes), but in these species the calyx and outside of corolla are glabrous. Description: Shrub, 2.5–4.5 m tall; twigs glabrous, older ones with pale grey bark. Leaves with petiole 7–13 mm long, glabrous; lamina elliptic to oblong– elliptic, (9.0–)17.0–23.5 × (4.3–)7.5–12.5 cm; apex acuminate with acumen 6–13 mm long; base shortly attenuate, acute or obtuse; leaf surface glabrous on both sides; secondary veins brochidodromous, six to eight on each side of the midrib, ascending, straight to slightly curved at the base, strongly curved some distance from the margin to join with the adjacent vein; domatia absent. Stipules with a short triangular base prolonged into an awn 4–9 mm. Inflorescences axillary paired at node, two- to four-flowered; peduncle short, < 6 mm; bracts triangular, c. 5 mm long, glabrous. Flowers 5-merous, subsessile; flower buds 16–24 mm long, apiculate. Calyx pale green; calyx tube c. 1 mm; calyx lobes broadly elliptic to ovate, 10–16 × 6–9 mm, ciliate. Corolla pale yellow to pale brownish–green; tube slightly funnel-shaped, 14.5– 19.0 × 6.0–9.5 mm; lobes narrowly triangular, 7–13 mm long, keeled outside, shortly apiculate (c. 2.5 mm); outside with five rows of hairs on the median keel of the lobes, prolonged on the tube for the upper third; inside with a ring of hairs near the base of the tube. Stamens five, attached near the mouth of the tube; filaments short, c. 0.5 mm; anthers c. 2.5 mm, half-exserted. Ovary pentalocular, glabrous. Disc cylindrical, c. 0.3 mm high, glabrous. Style, c. 23 mm long, with a pubescent fusiform swelling at the base, otherwise glabrous; stigmatic club lobed, exserted, c. 2 mm long. Fruits yellow, oval– obovate to subglobose, up to 35 × 30 mm when fresh (25 × 20 mm when dry), glabrous, crowned by persistent calyx lobes; pyrenes unknown. Distribution: Lower Guinea Domain: endemic to a narrow range south-east of Ebolowa, Cameroon. Habitat and ecology: Evergreen forest; 650–750 m in altitude. Phenology: Flowers in December and January; young fruits in December, January and March; mature fruits in April. Etymology: The specific epithet honours Dr J. J. F. E. de Wilde of Herbarium Vadense (Wageningen, the Netherlands), a specialist on Begonia and Meliaceae, who collected most of the specimens including the type. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS 429 Conservation status: IUCN Red List Category Critically Endangered [CR B1ab (i, ii, iii)]. The species is found far away from protected areas. The region is interspersed with several roads along which human activities are intense, including housing, smallholder farming and forest logging. The species is found in only one location. The extent of occurrence is estimated to be < 100 km2. Based on the threats identified, a continuing decline can be inferred in EOO, AOO and area, extent and quality of habitat. Dessein & Sonké 1464 (BR); Mvilé (3 km west-northwest of Ngovayang), 29 Nov 2005, Sonké 4170 (BR, BRLU, K, MO, WAG, YA); Bibondi, 21 Jan 2005, Sonké & Nguembou 3722 (BR, K, YA); ibid., 29 Jan 2005, Sonké & Nguembou 3865 (BR, K, MO, YA); 3 km north-west of Mbikiliki, 21 Jan 2006, Sonké & Djuikouo 4334 (K, MO, YA); Mbikiliki, 20 Jan 2008, Taedoumg 210 (BR); Lokundje, Oct 1919, Zenker 89 (BM); Ibid., 1911, Zenker 4003 (BM, BR, HBG); ibid., 19 Feb 1912, Zenker 4003b (K); Ibid., 1912, Zenker 4425 (BM, BR, HBG, K). Notes: Globulostylis dewildeana has so far been confused with G. talbotii in herbaria, but has much a larger corolla and calyx and a different range; it is not sympatric with any of its congeners, except the distinct G. cuvieroides. 4. GLOBULOSTYLIS MINOR WERNHAM, Cat. Pl. Oban: 50. 1913. Cuviera minor (Wernham) Verdc., Kew Bull. 42: 189. 1987, nom. illeg. (non C. minor C.H.Wright). Cuviera wernhamii Cheek, Pl. Mount Cameroon: 106. 1998. Type: NIGERIA. Oban, Talbot 247 (holotype: BM000903505!). Additional specimens examined: CAMEROON. hill facing the village of N’Kolandom, 9 Apr 1975, J.J.F.E. de Wilde 8159 (BR, K, P, WAG, YA); hill above the village of N’Kolandom, 12 Dec 1974, J.J.F.E. de Wilde 7831A (BR, K, WAG); hill Nkolomeyan on the track Bihongboulou–Koungoulou, 25 km east-southeast of Ebolowa, Letouzey 9843 (BR, P, YA). 3. GLOBULOSTYLIS LENIOCHLAMYS (K.SCHUM.) SONKÉ, O.LACHENAUD & DESSEIN comb. nov. Cuviera leniochlamys K.Schum., Bot. Jahrb. Syst. 28: 79. 1899. syn. nov. Type: CAMEROON. Bipindi, 19 Oct 1897, Zenker 1571 (lectotype designated here: K000412051!; isolectotype: BM!). Distribution: Lower Guinea Domain: endemic to the Ngowayang massif in southern Cameroon. Notes: This species has been much mistaken in herbaria, mostly with G. rammelooana and G. robbrechtiana that both occur in its range. It especially resembles the latter in corolla shape, but differs in having a larger calyx not or hardly exceeded by the corolla tube, a subglobose (not fusiform) basal style swelling, and the fruits larger, 24–28 × 17–26 mm when dry, smooth or nearly so. From G. rammelooana, it differs in the larger corolla and calyx (see Key). The flower colour of C. leniochlamys needs checking in the field; Schumann (1899) describes the corolla as ochre yellow (‘ockergelb’), which could refer to a fading flower. The original material has presumably been destroyed in Berlin, so the K duplicate is selected as lectotype. Additional specimens examined: CAMEROON. from Mbikiliki to basecamp, c. 1.5 km north-west of village, 9 Mar 2007, Dessein & Sonké 1448 (BR); near Mbikiliki village, from basecamp to top of hill, 10 Mar 2007, Distribution: Lower Guinea Domain: restricted to extreme south-east Nigeria (Oban) and neighbouring Cameroon (Korup National Park, Mount Etinde). Notes: When treated in Globulostylis, this species is correctly named G. minor. However, if placed in Cuviera, the replacement name C. wernhamii should be used (Cable & Cheek, 1998), because C. minor (Wernham) Verdc. is an illegitimate later homonym of C. minor C.H.Wright (which is now a synonym of C. nigrescens). Globulostylis minor is closely related to G. rammelooana, which is locally sympatric (see that species for differences). Records from Gabon (Sosef et al., 2006) are an error for G. cuvieroides; the two species, although quite dissimilar, have been much confused in herbaria. Additional specimens examined: CAMEROON. c. 10 km west of Limbe, above the village of Batoke, 24 Oct 2002, Davis 3007 (BR); Mount Etinde, 29 Apr 2009, Dessein et al. 2876 (BR, YA); west Limbe, 15 Nov 1985, Gentry & D.W. Thomas 52842A (K, MO, WAG); Upper Boando, 6 Dec 1993, Lighava 31 (K); chimpanzee camp, 19 km from Mundemba village, Korup National Park, CTFS forest dynamic plot, 28 Oct 2008, Parmentier & Mambo 5115 (BRLU); Mundemba (Korup National Park), 29 Dec 1992, Sonké 381 (BR); Liwenyi, 29 Oct 1993, Tchouto 1007 (K); Idenau, 8 Nov 1993, Watts 969 (K); Limbe, 0.5 km north-east of Etome, 23 Jan 1994, Wieringa 1990 (WAG). 5. GLOBULOSTYLIS RAMMELOOANA SONKÉ, O.LACHENAUD & DESSEIN sp. nov. Type: CAMEROON. Nkolembonda, 476 m alt, 2°47′58,6”N, 10°01′15,9”E, 13 Mar 2008, Sonké & Simo 4671 (holotype: BR!; isotypes: K!, MO!, P!, WAG!, YA!). © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 430 B. VERSTRAETE ET AL. Diagnosis: Globulostylis rammelooana is closely related to G. minor, from which it differs in having a larger calyx with erect lobes 6–10 × 5.5–7.0 mm (vs. patent lobes 4–7 × 2.5–5.0 mm; see Fig. 5E and G for comparison), and also a larger corolla, with tube 6–8 mm long and lobes 6–10 mm long (vs. tube 2–5 mm long and lobes 3–5 mm long in G. minor). In addition, the leaf underside lacks the typical greyish sheen of G. minor, but instead dries olive–green to olive–brown. Globulostylis rammelooana is also related to G. leniochlamys and G. robbrechtiana, which have much larger flowers. Fruiting specimens differ from G. leniochlamys in their smaller calyx, and from G. robbrechtiana in the larger and less markedly lobed fruits. The calyx lobes are more rounded and overlapping than in G. robbrechtiana (compare Figs 3G and 4H) and the corolla is green, not yellow. Description: Shrub, 2–4 m tall; twigs glabrous. Leaves with petiole 5–10 mm long, glabrous; lamina elliptic to oblong–elliptic, (6.5–)8.7–20.2 × (2.5–)3.4–7.4 cm; apex acuminate with acumen 7–17 mm long; base shortly attenuate or acute; leaf surface glabrous on both sides; secondary veins brochidodromous, five to seven on each side of the midrib, ascending, straight to slightly curved at the base, strongly curved some distance from the margin to join with the adjacent vein; domatia absent. Stipules deciduous, shortly triangular at base, prolonged into an awn of 4–7 mm. Inflorescences axillary or, more rarely, terminal, twoor three-flowered (but usually with only one fruit developing), peduncle 0–5 mm long; bracts ovate, 5–8 × 2.5–3.5 mm, glabrous. Flowers pentamerous; pedicels 0–2 mm long; flower buds green, 10–13 mm long, acute. Calyx pale green; calyx tube 1.5–2.0 mm; calyx lobes broadly elliptic, erect, 6–10 × 5.5–7.0 mm, glabrous. Corolla green; tube campanulate, 6–8 × 5– 8 mm, outside glabrous, inside with a ring of hairs near the basal third of the tube; lobes triangular, 6–10 mm long. Stamens five, inserted in the throat; filaments 1.5 mm; anthers c. 2 mm, just exserted, reflexed between the lobes. Ovary pentalocular, glabrous. Disc cylindrical, c. 0.7 mm high, glabrous. Style 8–10 mm long, with a conspicuous hairy globular swelling c. one fifth from the base, otherwise glabrous; stigmatic club five-lobed, exserted, lobes c. 3 mm long. Fruits subglobose to slightly ellipsoid, weakly five-lobed when dry, 19–25 mm in diameter when dry, glabrous, crowned by persistent calyx lobes, on pedicel 2–4 mm long; pyrenes five per fruit, 2.4 × 0.7 mm. Distribution: Lower Guinea Domain: endemic to Cameroon, occurring mostly in the South Region (Bipindi to Nyabessan) with a smaller population in the South-West Region (Mount Etinde). Habitat and ecology: Evergreen forest; 40–476 m in altitude. Phenology: Flowers in November–January and March–April; fruits in February–April and June. Etymology: Named in honour of Professor Jan Rammeloo, retired director of the National Botanical Garden of Belgium, to whom all authors are indebted for the assistance and facilities offered by his institute. Conservation status: IUCN Red List Category Vulnerable [VU B2ab(i, ii, iii, iv, v)]. G. rammelooana is restricted to Cameroon where it is mostly found outside protected areas. Logging, deforestation and urbanization are the major threats for this species. This species is known from nine locations. The area of occupancy is less than 500 km2 (72 km2). Based on the threats identified, a continuing decline of EOO, AOO, area, extent and quality of habitat, number of locations and number of mature individuals can be inferred. Notes: Globulostylis rammelooana has usually been confused with G. leniochlamys in herbaria. Its range overlaps those of G. leniochlamys and G. robbrechtiana in the South Region of Cameroon, and of G. minor in the South-West Region. All these species mostly differ in flowering characters, but the identification of fruiting specimens is difficult, although possible with experience with the group (see Diagnosis and Key). The collections from Mount Etinde lack flowers, but as they fully match G. rammelooana in other characters, they are referred to this species. Additional specimens examined: CAMEROON. East of Mount Elephant, 5 km south of km 18 Kribi– Ebolowa, 6 Mar 1970, Bos 6496 (WAG); near Bidjouka, track to basecamp, 13 Mar 2007, Dessein & Sonké 1546 (BR); Mount Elephant, south-east of Kribi, 28 Apr 1970, Bos 6871 (K, WAG); near Bidjouka, track towards waterfall, 15 Mar 2007, Dessein & Sonké 1602 (BR); Boa, 2 May 1994, Ekema 852 (K); Boa, 4 May 1994, Ekema 922 (K); Etinde, Fraser 347 (K); Mvini (Mivini), 35 km east of Campo, 1 Dec 1983, Kaji 205 (YA); Etinde, Ndam 1132 (K); ibid., 2 May 1994, Ndam 1185 (K); ibid., Ndam 1244 (K); cañon of Ntem, 29 km south-west of Nyabessan, 1 Dec 1982, Nkongmeneck 422 (BR, YA); Bipindi, 25 Nov 2004, Sonké & Nguembou 3585 (BR, BRLU, K, MO, WAG, YA); Mingli II (6 km north of Bipindi), 15 Jan 2005, Sonké & Nguembou 3656 (BR, BRLU, K, MO, WAG, YA); west Ngoyang, 18 Sep 2005, Sonké 3988 (BR, K, MO, YA); 3 km west-north-west of Bidjouka, 16 Jun © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS 2006, Sonké & Taedoumg 4464 (BR, YA); Lambi, massif of Ngovayang, 20 Feb 2008, Sonké 4650 (BR); Nkolembonda, Mount Elephant, 13 Mar 2008, Sonké & Simo 4680 (BR, K, MO, P, WAG, YA); Ibid., 16 Mar 2008, Sonké & Simo 4711 (BR, K, MO, P, WAG, YA); Nkolembonda, 16 Jul 2007, Taedoumg 102 (BR); South Province, Campo Ma’an area, Massif des Mamelles, 20 Apr 2001, Tchouto MMX110 (WAG); Mokoko Forest Reserve, Ekombe–Mofako, 23 May 1994, Watts 1164 (K, SCA); Bipindi, 1909, Zenker 3927 (BR, K). 6. GLOBULOSTYLIS ROBBRECHTIANA SONKÉ, O.LACHENAUD, DESSEIN & DE BLOCK sp. nov. Type: CAMEROON. Efoulan, Akom II, 894 m alt, 2°44′56.6”N, 10°32′03”E, 24 May 2008, Sonké, Taedoumg & Simo 4868 (holotype: BR!; isotypes: BR!, K!, MO!, P!, WAG!, YA!) Diagnosis: Globulostylis robbrechtiana differs from its congeners in the long, 20- to 30-mm corolla tube, which is at least twice as long as the calyx (vs. corolla tube 2–19 mm long, not or shortly exceeding calyx) and in the rather small and strongly five-lobed fruits, 15–22 × 12–19 mm when dry (vs. 19–35 × 17–35 mm when dry). It further differs from all species (except possibly G. leniochlamys) in having bright yellow flowers. Description: Shrub 2–4(−6) m tall; twigs brown, glabrous or sometimes sparsely hairy when young. Leaves with petiole 5–10 mm long, glabrous; lamina elliptic to oblong–elliptic, 7.3–18.8(−20.2) × 3.0– 7.8 cm; apex acuminate with acumen 7–10(−17) mm long; base shortly acute to rounded; leaf surface glabrous, or sometimes pubescent on the midrib below; secondary veins brochidodromous, prominent, (four–) six to nine on each side of the midrib, ascending, straight to slightly curved at the base, strongly curved some distance from the margin to join with the adjacent vein; domatia absent. Stipules shortly triangular at base, prolonged into an awn of 3–4 mm long, glabrous outside, hairy inside. Inflorescences axillary or occasionally terminal, two–four (–six)-flowered (but usually with only one fruit developing); peduncle 4–7 mm long; bracts ovate–triangular, 5–10 × 2– 4 mm, glabrous. Flowers pentamerous; pedicels 2–4 mm long; flower buds 22–32 mm long, acute. Calyx pale green; calyx tube 1–2(−4) mm; calyx lobes ovate to elliptic, erect, 6–12 × 3.5–9.0 mm, glabrous. Corolla yellow; tube funnel-shaped, 20–30 × 13– 20 mm at throat; lobes triangular, 5–10 mm long; outside glabrous; inside with a ring of hairs near the base of the tube. Stamens five, inserted near the apex of the tube; filaments 1 mm; anthers c. 2.5 mm, halfexserted to included. Ovary pentalocular, glabrous. 431 Disc cylindrical, c. 0.5 mm high, glabrous. Style, 20–32 mm long, with a pubescent fusiform swelling c. 1.5 mm above the base; stigmatic club five-lobed, just exserted, c. 2.5 mm long. Fruits subglobose, conspicuously five-lobed, 15–22 × 12–19 mm when dry, glabrous, crowned by persistent calyx lobes, on pedicel 4–8 mm long; pyrenes five per fruit, c. 17 × 5 mm. Distribution: Lower Guinea Domain: restricted to hilly ranges of southern Cameroon (Ngowayang massif, Akom II area), Equatorial Guinea (Monte Alén) and northern Gabon (Crystal Mountains). Locally abundant in its range. Habitat and ecology: Evergreen forest; 460–1110 m in altitude. Phenology: Flowers mostly from May to October, with one record in March; fruits from July to December. Etymology: Named in honour of Professor Elmar Robbrecht, retired head of the Vascular Plants Department at the National Botanic Garden of Belgium, who made a major contribution to the knowledge of African Rubiaceae and to whom all authors are indebted for instruction in aspects of this family. Conservation status: IUCN Red List Category Near Threatened (NT). Despite the restricted EOO (11106, 36 km2), which qualifies for Vulnerable, and AOO (72 km2), which meets the threshold for Endangered, and a low number of locations (six), G. robbrechtiana is here considered to be only Near Threatened. Most collections have been made inside protected areas; three collection sites are outside protected areas, but the surrounding forest seems to be rather untouched by human activities at present. This might change in the future, the reason why we think the category NT is the most appropriate. Notes: The large yellow flowers of G. robbrechtiana are distinctive, but specimens in bud or fruit can be mistaken for G. leniochlamys or G. rammelooana (see these species for differences). The ranges of the three species overlap in Cameroon. Globulostylis robbrechtiana is normally glabrous, but some duplicates of Sonké, Taedoumg & Simo 4872 have short patent hairs on young twigs and the underside of midrib, which is exceptional in the genus. Additional specimens examined: CAMEROON. Akom II, 24 Apr. 2007, Droissart & Simo 438 (BRLU); Ebianemeyong, 24 May 2002, Elad et al. 1558 (WAG); ibid., 24 May 2002, Elad et al. 1574 (WAG); Engon, © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 432 B. VERSTRAETE ET AL. south Efoulan, 6 Mar 2004, Sonké & Beina 3331 (BR, BRLU, K, MO, WAG, YA); Akom II, Efoulan, 24 Mar 2008, Sonké et al. 4872 (BR, K, MO, P, WAG, YA); Campo Ma’an area, Efoulan, 4 Dec 2000, Tchouto 3092 (WAG); Bipindi, Oct 1918, Zenker 13 (P). EQUATORIAL GUINEA. Rio Muni: Monte Alen, 20 May 1992, Carvalho 5115 (WAG); Bata-Monte Alen, Subida al Monte Alen, 27 May 1993, Carvalho 5310 (WAG); Transect of Monte Chocolate, 16 Nov 2002, Desmet et al. 9 (BR); Engong, 9 May 2002, Esono & Parmentier 589 (BRLU); Monte Chocolate, 11 Aug 2001, Esono & Senterre 272 (BRLU); Monte Mitra (Monte Alen National Park), 30 Sep 2005, Leal et al. 806 (BR, MO); Ibid., 30 Sep 2005, M.E. Leal et al. 807 (BR, MO); between 0 and 1 km west of Alen (Monte Alen National Park), 15 Oct 1993, Lejoly 93/315 (BRLU); ibid., 15 Oct 1993, Lejoly 93/339 (BRLU); ibid., 21 Oct 1993, Lejoly 93/377 (BRLU); ibid., 21 Oct 1993, Lejoly 93/422 (BRLU); Monte Alen National Park, 7 Oct 1994, Lejoly 94/168 (BRLU); ibid., 7 Oct 1994, Lejoly 94/185 (BRLU); ibid., 22 Sep 1997, Lisowski 1554 (BRLU); ibid., 22 Sep 1997, Lisowski 1571A (BRLU); ibid., 28 May 1997, Ngomo 0173 (BRLU); ibid., 16 Jun 1998, Ngomo & Ndong 348 (BRLU); ibid., 10 Aug 1998, Ngomo & Ndong 439 (BRLU); 5 km west of Engong (Monte Alen National Park), 3 Jul 1999, N. Nguema & Parmentier 446 (BR, BRLU); s.loc., 1 Dec 1997, Obama 347 (BRLU); S du Rio Laña, près de la Cabaña Ecofac de Misergue (south-east Monte Alen National Park), 13 Jul 2002, Senterre & Obiang 3323 (BRLU); 2 km north-east of Nkumékié (south-east Monte Alen National Park), 8 Dec 2002, Senterre & Obiang 3679 (BRLU); 8.5 km east de la Cabaña de Mosumo (Monte Alen National Park), 4 Jul 2003, Senterre & Obiang 3991 (BRLU); Monte Alen, 10 Aug 2001, Sonké et al. 2501 (BR); Transect Monte Chocolate (Monte Alen National Park), 8 Jan 1998, Van Reeth 145 (BRLU). GABON. Crystal mountains, 15 Nov 2000, N. Nguema 1304, 1391 (WAG); Tchimbélé, 17 Dec 1998, Wieringa 244 (WAG). 7. GLOBULOSTYLIS TALBOTII WERNHAM, Cat. Pl. Oban: 50. 1913. Cuviera talbotii (Wernham) Verdc., Kew Bull. 42: 189. 1987. Type: NIGERIA, Oban, 4 Apr 1912, Talbot 2051 (lectotype designated here: BM000903503!; isolectotypes: BM000903504!, K000412052!, K000412053!). Distribution: Endemic to Lower Guinea Domain: restricted to extreme south-east Nigeria and adjacent south-west Cameroon. Notes: Globulostylis talbotii is closely related to G. dewildeana, which has been previously mistaken for it in herbaria, but has much smaller flowers and a more north-westerly range. From the sympatric G. minor and G. rammelooana, it can be told apart by the ciliate calyx and the corolla pubescent outside on the midrib of the lobes. Additional specimens examined: CAMEROON. Boa, Ekema 1131b (K, SCA); ibid., Ekema 1212 (K, SCA); western side of Mount Koupé, near Mbule, 24 Jan 1972, Leeuwenberg 9275 (BR, P, WAG, YA); Mbu Bakundu, Mamfe road, Nemba & Mambo 714 (K); Boa, Pouakouyou 76 (K, SCA); Bonjare, Tchouto 1162 (K, SCA); Korup National Park, Jan 1972; D.W. Thomas 593 (BR); south end of Korup National Park, Mar 1986, D.W. Thomas & McLeod 5725 (K). 8. GLOBULOSTYLIS UNCINULA (N.HALLÉ) SONKÉ, O.LACHENAUD & DESSEIN comb. nov. Cuviera uncinula N.Hallé, Bull. Soc. Bot. France 106: 343. 1960. syn. nov. Type: GABON. La Nkoulounga (north-east Libreville), 3 Jul 1959, N. Hallé 719 (holotype: P00553433!; isotype: P00553432!). Distribution: Lower Guinea and Congolia Domains: main range in Gabon and adjacent Republic of Congo, with a smaller area in the central Democratic Republic of the Congo. Notes: G. uncinula differs from other related species (G. leniochlamys, G. minor, G. rammelooana and G. robbrechtiana) in having consistently one-flowered inflorescences and in combining a short corolla tube with a large calyx. Fruiting specimens are difficult to separate from G. leniochlamys, but the two species have different ranges. Globulostylis uncinula, until now regarded as a Gabonese endemic, is newly reported from the Democratic Republic of the Congo and the Republic of Congo. Additional specimens examined: DEMOCRATIC REPUBLIC OF THE CONGO. Imbow, 11 Feb 1959, Evrard 5689 (BR); Mompono, 25 Feb 1959, Evrard 5834 (BR, K, WAG). GABON. near Yombi (on Fougamou–Mouila road, Koumounabouali ridge), 19 Sep 1990, Breteler 13993 (WAG); Mount Songou (between Poungui and Dibandi), 23 Feb 2008, Dessein et al. 2116 (BR, LBV); base of Mount Iboundji, 3 Mar 2008, Dessein et al. 2337 (BR, LBV); La Nkoulounga, 3 Jul 1959, N. Hallé 724 (P); Abanga chantier CEFA, 11 Jun 1963, N. Hallé 2447 (P); Iméno Ivinzi, 23 Mar 1927, Le Testu 6446 (BM, BR, P); east chantier Mitendi, 21 Oct 1999, Sosef et al. 583 (WAG); Mount Iboundji, 10 Feb 2000, Sosef et al. 720 (BRLU, WAG); c. 40 km north-east of Mitzic (forestry concession Bordamur), 6 Feb 2003, Sosef et al. 1891 (WAG); c. 50 km south of Mandji © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS (Doudou Moutains), 17 Nov 2005, Sosef et al. 2367 (WAG); Shell oil-exploitation Rabi, 1 km south-east of Shell camp., 8 Mar 1994, Wieringa 2416 (WAG); north Ndouaniang (‘Sogacel’), 6 Aug 1984, Wilks 987 (WAG). REPUBLIC OF CONGO. Maamar forest exploitation (towards N’Gongo-N’Dindi junction), 5 Feb 1974, Sita 3648 (P, WAG); around junction N’Dindi-N’Gongo and N’Tiétié, 10 Dec 1974, Sita 3816 (P); Bambama, 6 May 1983, Sita 4790 (BR, P). SPECIES EXCLUDED FROM CUVIERA 1. CUVIERA AUSTRALIS K.Schum. – now a synonym of Vangueria lasiantha (Sond.) Sond. (Bridson, 1998). 2. CUVIERA BOLO Aubrév. & Pellegr. – now a synonym of Robynsia glabrata Hutch. (Hepper & Keay, 1963). 3. CUVIERA CALYCOSA WERNHAM, J. Bot. 52: 7. 1914. Type: NIGERIA. Eket District, Talbot 3300 (lectotype designated here: BM000903513!; isolectotypes designated here: BM000903512!, K000412057!, K000412058!). Distribution: Lower Guinea and Congolia Domains: Cameroon, Democratic Republic of the Congo, Gabon, Nigeria, Republic of the Congo. Notes: This species is excluded from Cuviera, but not otherwise placed (see Discussion). It differs from C. nigrescens only (see below) in the size of calyx lobes, so an intraspecific rank might be more appropriate. Mengé 5 (BR) is the first record from the Democratic Republic of Congo. 4. CUVIERA MIGEODII VERDC., Fl. Trop. E. Afr., Rubiac. (3): 771. 1991. Type: TANZANIA. Tendaguru, Lindi District, 18 Dec 1930, Migeod 1052 (holotype: BM000903510!). Distribution: South Tanzania (Lindi District). Notes: This species is excluded from Cuviera, but not otherwise placed (see Discussion). 5. CUVIERA NIGRESCENS (SCOTT-ELLIOTT EX OLIV.) WERNHAM, J. Bot. 49: 321. 1911. Vangueria nigrescens Elliott ex Oliv., Bot. J. Linn. Soc. 30: 81. 1895. Type: SIERRA LEONE. Fabala, Mar 1892, Scott-Elliot 5736 (lectotype designated here: K000412056!); in forest near Kafogo, 6 Apr 1892, Scott-Elliot 5610 (syntype: K00412055!). Synonyms: Cuviera trichostephana K.Schum., Bot. Jahrb. Syst. 23: 461. 1897. Type: SIERRA LEONE. s.loc., Scott-Elliot s.n. (K). Cuviera minor C.H.Wright, Bull. Misc. Inform. 433 Kew. 1906: 105. 1906. Type: GHANA. Kwahu, 2 Apr 1900, Johnson 646 (lectotype designated here: K000412054!). Distribution: Upper and Lower Guinea Domains: Cameroon, Central African Republic, Democratic Republic of the Congo, Gabon, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, Nigeria and Sierra Leone. Notes: This species is excluded from Cuviera, but not otherwise placed (see Discussion). The original description is based on two syntypes, one of which is selected as lectotype. 6. CUVIERA SCHLIEBENII VERDC., Kew Bull. 33: 497. 1979. Type: TANZANIA. Lutamba See, Lindi District, 1 Nov 1934, Schlieben 5576 (holotype: B100160713!; isotypes: BR0000008846815!, EA000002930!, LISC002649!, PRE0593394-0!, US00664144!). Distribution: Mozambique and south Tanzania. Notes: This species is excluded from Cuviera, but not otherwise placed (see Discussion). 7. CUVIERA SEMSEII VERDC., Kew Bull. 12: 449. 1957. Type: TANZANIA. Mchinjiri, Rondo Plateau, Lindi District, 823 m, Jan 1952, Semsei 620 (holotype: EA000002929!; isotypes: BR0000008846808!, K000412039!). Distribution: Tanzania. Malawi, Mozambique and south Notes: This species is excluded from Cuviera, but not otherwise placed (see Discussion). It is close to C. nigrescens and we are not convinced they are specifically distinct. 8. CUVIERA TOMENTOSA VERDC., Kew Bull. 36: 557. 1981. Type: TANZANIA. Kilwa District, Selous Game Reserve, approximately 15 km south-west of Kingupira, 8°35′S 38°27′E, 175 m, 21 Dec 1975, Vollesen 3122 (holotype: K; isotypes: C, EA, WAG). Distribution: North Mozambique (Niassa Province) to south Tanzania (Lindi Region). Notes: This species is excluded from Cuviera, but not otherwise placed (see Discussion). ACKNOWLEDGEMENTS This research was supported financially by the Research Foundation – Flanders (FWO, G.0343.09N), © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 434 B. VERSTRAETE ET AL. the King Leopold-III Fund for Nature Exploration and Conservation, the Sud Expert Plantes project (no. 375), the F.F.R.S.A. (Fondation pour Favoriser les Recherches Scientifiques en Afrique), the Alberta Mennega Stichting and SYNTHESYS. B.V. holds a PhD research grant from the Agency for Innovation by Science and Technology (IWT, no. 91158). O.L. is a research fellow of the F.R.S.-F.N.R.S. (National Fund for Scientific Research). B.S.’s visits to Belgium were partly funded by the Université Libre de Bruxelles and the F.R.S.-FNRS. We thank the curators of the following herbaria for their help while working in their institutions and/or the loan of material: BM, BNRH, BR, BRLU, HBG, K, MPU, P, LBV, WAG and YA. Dr Petra De Block is acknowledged for her help in the description of Globulostylis robbrechtiana. Murielle Simo, Quentin Luke and Olivier Maurin kindly allowed us to reproduce their photographs. B.S.’s visits to Belgium and France in 2012, during which time this paper was finalized, were funded by the ‘Institut de Recherche pour le Développement (IRD)’, under the ISMOBIAC project. Thanks are due to Dr Pierre Couteron (Institut de Recherche pour le Développement IRD – AMAP), Dr Olivier Hardy (Laboratoire d’Eco-éthologie évolutive, Université Libre de Bruxelles) and Dr Tariq Stévart (Herbarium et Bibliothèque de Botanique africaine, Université Libre de Bruxelles) for their assistance. We are deeply grateful to Dr Vincent Droissart, Yves Issembé, Dr Steven Janssens, Thomas Nzabi, Murielle Simo, Hermann Taedoumg and Lise Zemagho for help with the collection of plant material. REFERENCES Albuquerque S, Brummit R, Figueiredo E. 2009. Typification of names based on the Angolan collections of Friedrich Welwitsch. Taxon 58: 641–646. Bridson DM. 1998. Rubiaceae (Tribe Vanguerieae). In: Pope GV, ed. Flora Zambesiaca, Vol. 5, part 2. Kew: Royal Botanic Gardens, 211–377. Brunel J-F, Hiepko P, Scholz H. 1984. Flore analytique du Togo. Englera 4: 1–751. Cable S, Cheek M. 1998. Plants of Mount Cameroon: a conservation checklist. London: Kew Publishing. de Candolle AP. 1807. 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Journal of Botany, British and Foreign 52: 7–8. Wernham HF. 1918. New Rubiaceae from the Belgian Congo. Journal of Botany, British and Foreign 56: 309–314. White F. 1979. The Guineo–Congolian Region and its relationship to other phytochoria. Bulletin du Jardin botanique national de Belgique 49: 11–55. White T, Bruns T, Lee S-T, Taylor J. 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis M, Gelfand D, Sninsky J, White T, eds. PCR protocols: a guide to methods and applications. San Diego, CA: Academic Press, 315–322. APPENDIX 1 ACCESSIONS 435 INCLUDED IN THE PHYLOGENETIC ANALYSIS Taxon* Voucher trnTF rpl16 petD ITS rpl32trnL accDpsaI Afrocanthium gilfillanii (N.E.Br.) Lantz Afrocanthium lactescens (Hiern) Lantz Afrocanthium mundianum (Cham. & Schltdl.) Lantz Bullockia pseudosetiflora (Bridson) Razafim., Lantz & B.Bremer Bullockia setiflora (Hiern) Razafim., Lantz & B.Bremer Canthium ciliatum (D.Dietr.) Kuntze Lemaire & Verstraete 32 (BR) Kuchar 23003 (BR) JQ958219 JQ958112 JQ958036 – – – JQ958221 JQ958113 JQ958037 – – – Lemaire & Verstraete 33 (BR) JQ958220 JQ958114 JQ958038 – – – Bidgood et al. 4959 (BR) JQ958222 JQ958115 JQ958039 JQ957968 – – Lemaire & Verstraete 197 (BR) Lemaire & Verstraete 281 (BR) JQ958223 JQ958116 JQ958040 JQ957969 – – JQ958230 JQ958122 JQ958049 – – – © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 436 B. VERSTRAETE ET AL. APPENDIX 1 Continued Taxon* Voucher trnTF rpl16 petD ITS rpl32trnL accDpsaI Canthium coromandelicum (Burm.f.) Alston Canthium glaucum Hiern Canthium inerme (L.f.) Kuntze Andreasen 36 (UPS) AJ620122 AJ876810 – AJ315081 – – Kuchar 17410 (BR) Lemaire & Verstraete 235 (BR) Lemaire & Verstraete 241 (BR) Lemaire & Verstraete 187 (BR) Burrows & Burrows 9858 (BNRH) Dessein et al. 2809 (BR) Amsini 97 (BR) JQ958231 JQ958123 JQ958046 JQ957970 – – JQ958232 JQ958124 JQ958047 JQ957971 – – JQ958233 JQ958125 JQ958050 – – – JQ958235 JQ958126 JQ958048 JQ957972 – – JQ958234 JQ958127 – – – – JQ958270 JQ958157 JQ958076 JQ957973 JQ958201 JQ957948 JQ958271 – JQ958077 JQ957974 JQ958197 JQ957951 JQ958159 JQ958079 JQ957975 JQ958199 JQ957953 JQ958158 JQ958078 JQ957976 JQ958198 JQ957949 JQ958274 JQ958160 JQ958080 JQ957977 JQ958200 JQ957950 JQ958276 – JQ958082 JQ957978 – JQ957962 JQ958280 – JQ958086 JQ957979 JQ958205 JQ957965 JQ958275 JQ958161 JQ958081 JQ957980 JQ958202 JQ957952 JQ958236 JQ958132 – – – – JQ958244 JQ958128 JQ958052 – – – JQ958239 – JQ958053 – – – JQ958240 JQ958129 JQ958054 JQ957981 JQ958179 JQ957933 JQ958238 JQ958133 – – – – JQ958241 JQ958130 JQ958055 JQ957982 – – Canthium kuntzeanum Bridson Canthium spinosum (Klotzsch ex Eckl. & Zeyh.) Kuntze Canthium suberosum Codd Cuviera acutiflora DC. Cuviera angolensis subsp. latior (Wernham) Sonké, O.Lachenaud & Dessein Cuviera longiflora Hiern Cuviera physinodes K.Schum. Cuviera physinodes K.Schum. Cuviera schliebenii Verdc. Cuviera semseii Verdc. Cuviera subuliflora Benth. Fadogia ancylantha Schweinf. Fadogia cienkowskii Schweinf. Fadogia fuchsioides Schweinf. ex Oliv. Fadogia homblei De Wild. Fadogia salictaria S.Moore Fadogia stenophylla subsp. odorata (K. Krause) Verdc. Dessein et al. JQ958272 2713 (BR) Chatrou 572 (BR) JQ958273 Dessein et al. 1416 (BR) Burrows & Burrows 9839 (BNRH) Burrows & Burrows 11303 (BNRH) Dessein et al. 2004 (BR) Dessein et al. 1101 (BR) Dessein et al. 258 (BR) Dessein et al. 1083 (BR) Lemaire & Verstraete 22 (BR) Malaisse 13490 (BR) Lovett 2267 (BR) © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS 437 APPENDIX 1 Continued Taxon* Voucher trnTF Fadogia tetraquetra K.Schum. & K.Krause Fadogia triphylla Baker Fadogia verdickii De Wild. & T.Durand Fadogiella stigmatoloba (K.Schum.) Robyns Globulostylis leniochlamys (K.Schum.) Sonké, O.Lachenaud & Dessein Globulostylis minor Wernham Globulostylis rammelooana Sonké, O.Lachenaud & Dessein Globulostylis rammelooana Sonké, O.Lachenaud & Dessein Globulostylis rammelooana Sonké, O.Lachenaud & Dessein Globulostylis rammelooana Sonké, O.Lachenaud & Dessein Globulostylis robbrechtiana Sonké, O.Lachenaud, Dessein & De Block Globulostylis uncinula (N.Hallé) Sonké, O.Lachenaud & Dessein Lemaire & JQ958242 Verstraete 223 (BR) Dessein et al. 226 JQ958237 (BR) Schaijes 1502 JQ958243 (BR) rpl16 petD ITS rpl32trnL accDpsaI JQ958131 JQ958056 – – – JQ958134 – – – – JQ958135 – JQ957984 – – Gillett 17403 (BR) JQ958245 JQ958136 – JQ957983 – – Dessein et al. 1448 (BR) JQ958269 JQ958152 JQ958070 JQ957985 JQ958189 JQ957947 Dessein et al. 2876 (BR) Dessein et al. 1546 (BR) JQ958261 JQ958151 JQ958075 JQ957986 JQ958190 JQ957939 JQ958267 JQ958153 JQ958071 JQ957987 JQ958193 JQ957942 Sonké et al. 4650 (BR) JQ958266 – – – JQ958191 JQ957943 Sonké et al. 4671 (BR) JQ958264 – – JQ957988 – JQ957941 Sonké et al. 4711 (BR) JQ958265 – – – JQ958192 JQ957940 Sonké et al. 4868 (BR) JQ958268 JQ958154 JQ958072 JQ957989 JQ958194 JQ957944 Dessein et al. 2116 (BR) JQ958262 JQ958155 JQ958073 JQ957990 JQ958195 JQ957946 © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 438 B. VERSTRAETE ET AL. APPENDIX 1 Continued Taxon* Voucher trnTF rpl16 petD ITS rpl32trnL accDpsaI Globulostylis uncinula (N.Hallé) Sonké, O.Lachenaud & Dessein Ixora finlaysoniana Wall. ex G.Don Dessein et al. 2337 (BR) JQ958263 JQ958156 JQ958074 JQ957991 JQ958196 JQ957945 Van Caekenberghe 54 (BR) Lemaire & Verstraete 97 (BR) Dessein et al. 1261 (BR) Dessein et al. 1085 (BR) JQ958210 JQ958105 JQ958028 – – – JQ958211 JQ958106 JQ958029 – – – JQ958212 JQ958107 – JQ957992 – – JQ958277 JQ958162 JQ958083 – – – Dessein et al. 2949 (BR) JQ958278 JQ958163 JQ958084 JQ957993 JQ958204 JQ957963 De Block et al. 2487 (BR) Lemaire & Verstraete 174 (BR) Van Caekenberghe 60 (BR) Lemaire & Verstraete 163 (BR) JQ958224 JQ958117 JQ958041 – – – JQ958229 JQ958121 JQ958051 JQ957994 – – JQ958208 JQ958104 JQ958026 – – – JQ958213 JQ958109 JQ958030 JQ957995 – – JQ958214 – JQ958031 – – – JQ958215 JQ958108 JQ958032 JQ957996 – – JQ958216 JQ958110 JQ958033 JQ957997 – – JQ958217 – JQ958034 JQ957998 – – JQ958218 JQ958111 JQ958035 JQ957999 – – JQ958228 JQ958120 JQ958045 JQ958000 – – JQ958279 – JQ958085 JQ958001 JQ958203 JQ957964 JQ958225 JQ958118 JQ958042 JQ958002 – – Keetia gueinzii (Sond.) Bridson Keetia venosa (Oliv.) Bridson Multidentia crassa (Hiern) Bridson & Verdc. Multidentia dichrophylla (Mildbr.) Bridson Peponidium sp. Plectroniella armata (K.Schum.) Robyns Pseudomussaenda flava Verdc. Psydrax fragrantissima (K.Schum.) Bridson Psydrax kraussioides (Hiern) Bridson Psydrax livida (Hiern) Bridson Psydrax locuples (K.Schum.) Bridson Psydrax obovata (Klotzsch ex Eckl. & Zeyh.) Bridson Psydrax parviflora (Afzel.) Bridson Pygmaeothamnus chamaedendrum (Kuntze) Robyns Pygmaeothamnus zeyheri (Sond.) Robyns Pyrostria bibracteata (Baker) Cavaco Dessein et al. 1285 (BR) Lemaire & Verstraete 14 (BR) Lemaire & Verstraete 287 (BR) Lemaire & Verstraete 188 (BR) Dessein et al. 1345 (BR) Lemaire & Verstraete 142 (BR) Lemaire & Verstraete 2 (BR) Burrows & Burrows 10784 (BNRH) © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS 439 APPENDIX 1 Continued Taxon* Voucher trnTF rpl16 petD ITS rpl32trnL accDpsaI Pyrostria hystrix (Bremek.) Bridson Lemaire & Verstraete 270 (BR) De Block et al. 2423 (BR) JQ958226 – JQ958043 JQ958003 – – JQ958227 JQ958119 JQ958044 – – – Tosh et al. 263 (BR) JQ958209 – JQ958027 – – – Hall & Amponsah 46545 (K) Lachenaud et al. 739 (BR) AJ620170 – – AJ617774 – – JQ958246 JQ958137 JQ958057 JQ958004 – JQ957934 Niyongabo 53 (BR) JQ958247 JQ958138 JQ958058 – – JQ957935 Dessein et al. 2961 (BR) Dessein et al. 2541 (BR) Lachenaud et al. 852 (BR) JQ958248 JQ958139 JQ958060 JQ958005 – JQ957936 JQ958249 JQ958140 JQ958059 JQ958006 JQ958180 JQ957938 JQ958250 JQ958141 JQ958061 JQ958007 – JQ957937 JQ958281 JQ958164 JQ958087 JQ958008 – – JQ958284 Lemaire & Verstraete 233 (BR) Dessein et al. 202 JQ958282 (BR) JQ958170 JQ958097 JQ958009 – – JQ958165 JQ958088 JQ958010 – – JQ958283 – JQ958089 JQ958011 – – JQ958285 JQ958166 JQ958090 – – – JQ958289 JQ958171 JQ958093 JQ958012 JQ958206 JQ957966 JQ958287 JQ958168 JQ958091 – – – JQ958296 JQ958169 JQ958101 JQ958013 – – JQ958288 JQ958172 JQ958098 JQ958014 JQ958207 JQ957967 Pyrostria serpentina Lantz, Klack. & Razafim. Razafimandimbisonia humblotii (Drake) Kainul. & B.Bremer Robynsia glabrata Hutch. Rytigynia membranacea (Hiern) Robyns Rytigynia monantha (K.Schum.) Robyns Rytigynia neglecta (Hiern) Robyns Rytigynia rubra Robyns Rytigynia umbellulata (Hiern) Robyns Vangueria agrestis (Schweinf. ex Hiern) Lantz Vangueria bowkeri (Robyns) Lantz Vangueria cinerascens (Welw. ex Hiern) Lantz Vangueria discolor (De Wild.) Lantz Vangueria dryadum S.Moore Vangueria infausta Burch. Vangueria lasiantha (Sond.) Sond. Vangueria latifolia (Sond.) Sond. Vangueria macrocalyx Sond. Lejoly 82/390 (BR) Bidgood et al. 6094 (BR) Lemaire & Verstraete 289 (BR) Lemaire & Verstraete 13 (BR) Lemaire & Verstraete 209 (BR) Lemaire & Verstraete 69 (BR) Lemaire & Verstraete 114 (BR) © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 440 B. VERSTRAETE ET AL. APPENDIX 1 Continued rpl16 petD ITS rpl32trnL accDpsaI – JQ958094 – – – JQ958173 – – – – Ntemi Sallu et al. JQ958293 309 (BR) JQ958174 JQ958096 JQ958015 – – Lemaire & Verstraete 265 (BR) Lemaire & Verstraete 42 (BR) Lemaire & Verstraete 98 (BR) Lemaire & Verstraete 285 (BR) Lemaire & Verstraete 25 (BR) Lemaire & Verstraete 53 (BR) Dessein et al. 1632 (BR) JQ958294 JQ958175 JQ958099 – – – JQ958297 JQ958177 JQ958102 – – – JQ958291 – JQ958095 – – – JQ958295 JQ958176 JQ958100 – – – JQ958298 JQ958178 JQ958103 – – – JQ958286 JQ958167 JQ958092 – – – JQ958251 JQ958146 JQ958062 JQ958016 JQ958181 JQ957954 Dessein et al. 1654 (BR) JQ958252 JQ958147 JQ958063 JQ958017 JQ958182 JQ957955 Jongkind 7887 (WAG) Lachenaud et al. 689 (BR) JQ958256 JQ958142 JQ958067 JQ958018 JQ958185 JQ957958 JQ958254 JQ958149 JQ958064 JQ958019 JQ958183 – Jongkind 3089 (WAG) JQ958258 JQ958144 JQ958068 JQ958020 JQ958186 JQ957959 Dessein et al. 1574 (BR) JQ958253 JQ958148 JQ958066 JQ958021 – JQ957956 Dessein et al. 2415 (BR) JQ958255 JQ958150 JQ958065 JQ958022 JQ958184 JQ957957 Lachenaud et al. 1119 (BR) Merello et al. 1494 (K) JQ958257 JQ958143 – JQ958023 JQ958188 JQ957960 AJ620182 – – AJ315085 – – Taxon* Voucher Vangueria madagascariensis J.F.Gmel. Vangueria micropyren (Verdc.) Lantz Vangueria pallidiflora (Bullock) Lantz Vangueria parvifolia Sond. Lemaire & JQ958290 Verstraete 189 (BR) Dessein et al. 253 JQ958292 (BR) Vangueria pygmaea Schltr. Vangueria randii subsp. chartacea (Robyns) Verdc. Vangueria soutpansbergensis N.Hahn Vangueria thamnus (Robyns) Lantz Vangueria triflora (Robyns) Lantz Vangueriella chlorantha (K.Schum.) Verdc. Vangueriella chlorantha (K.Schum.) Verdc. Vangueriella discolor (Benth.) Verdc. Vangueriella laxiflora (K.Schum.) Verdc. Vangueriella nigerica (Robyns) Verdc. Vangueriella nigricans (Robyns) Verdc. Vangueriella olacifolia (Robyns) Verdc. Vangueriella rufa (Robyns) Verdc. Vangueriella spinosa (Schumach. & Thonn.) Verdc. trnTF © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441 REVISION OF CUVIERA AND GLOBULOSTYLIS 441 APPENDIX 1 Continued Taxon* Voucher trnTF rpl16 petD ITS rpl32trnL accDpsaI Vangueriella vanguerioides (Hiern) Verdc. Vangueriella vanguerioides (Hiern) Verdc. Vangueriopsis shimbaensis A.P.Davis & Q.Luke Jongkind 7855 (WAG) JQ958259 – – JQ958024 – – Kpadeyeah 10 (WAG) JQ958260 JQ958145 JQ958069 JQ958025 JQ958187 JQ957961 Luke 8316 (UPS) AJ620183 AJ876875 – AJ617778 – – *Information on taxon, voucher and GenBank numbers for trnT-F, rpl16, petD, internal transcribed spacer (ITS), rpl32-trnL, accD-psaI. –, not sequenced. APPENDIX 2 DNA MARKERS WITH THEIR RESPECTIVE PRIMER PAIRS, ANNEALING TEMPERATURE, SUBSTITUTION MODEL AND REFERENCES TO THE SEQUENCES OF THE PRIMERS Marker Primer pairs Annealing Model Reference trnT-L trnTFa1 trnTLi trnLFc trnLFf L16exon1 (pcr) RPL16-18R (seq) 1067F petB1411 petD738 ITS1 ITS4 rpl32F accD769F psa175R 55 °C GTR + G Lantz & Bremer, 2004 55 °C GTR + G Lantz & Bremer, 2004 55 °C GTR + I + G Lantz & Bremer, 2005 55 °C GTR + G Löhne & Borsch, 2005 50 °C HKY + G White et al., 1990 55 °C 55 °C F81 + I GTR + I Shaw et al., 2007 Tosh et al., 2009 trnL-trnF rpl16 petD Internal transcribed spacer (ITS) rpl32-trnL accD-psaI © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441