Taxonomy and phylogenetics of Cuviera (Rubiaceae-Vanguerieae) and reinstatement of Globulostylis...

Taxonomy and phylogenetics of Cuviera(Rubiaceae–Vanguerieae) and reinstatement ofGlobulostylis with the description of three new species

BRECHT VERSTRAETE1*, OLIVIER LACHENAUD2,3, ERIK SMETS FMLS1,4,STEVEN DESSEIN3 and BONAVENTURE SONKÉ2,5,6

1Plant Conservation and Population Biology, KU Leuven, Kasteelpark Arenberg 31, PO Box 2437,3001 Leuven, Belgium2Service d’Evolution Biologique et Ecologie, Université Libre de Bruxelles, CP 160/12, 50 Avenue F.Roosevelt, 1050 Bruxelles, Belgium3National Botanic Garden of Belgium, Domein van Bouchout, 1860 Meise, Belgium4Naturalis Biodiversity Center, Leiden University, PO Box 9517, 2300 RA Leiden, The Netherlands5Plant Systematic and Ecology Laboratory, Higher Teachers’ Training College, University of YaoundeI, PO Box 047, Yaounde, Cameroon6Africa and Madagascar Department, Missouri Botanical Garden, PO Box 299, St Louis, MO63166-0299, 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. Recentmolecular studies, while providing new insights, have not yet solved all the problems. In this study, the taxonomyand 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 inCuviera, is reinstated as a distinct genus, with eight species from Central Africa. Both genera are revised; thelatter includes three new species (G. dewildeana, G. rammelooana and G. robbrechtiana) and two new combina-tions (G. leniochlamys and G. uncinula). The close relationship of Cuviera s.s. and Globulostylis to Vangueriella isshown. Six aberrant species (most from East Africa) are excluded from Cuviera, but further work is needed beforethey can be confidently assigned to other genera. © 2013 The Linnean Society of London, Botanical Journal ofthe Linnean Society, 2013, 173, 407–441.

ADDITIONAL KEYWORDS: Cameroon – Equatorial Guinea – Gabon – Tropical Africa.

INTRODUCTION

Tribe Vanguerieae are palaeotropical in distributionand among the larger tribes in Rubiaceae, comprisingapproximately 600 species in 25 currently acceptedgenera (Govaerts et al., 2012). The tribe is delineatedby a set of morphological characteristics, in particularthe specialized hood-like structure of the stigma(Lantz & Bremer, 2004), the valvate corolla lobes, the

pendulous ovules and the drupaceous fruits with one-seeded pyrenes in each locule. Generic delimitation inthe tribe is still problematic as several genera lackgood diagnostic characters. Not surprisingly, recentmolecular studies are partially in conflict withmorphological classifications (Lantz et al., 2002;Lantz & Bremer, 2004, 2005). At present, molecularsampling and tree resolution are insufficient toanswer all questions related to the generic delimita-tions in the tribe. Detailed molecular studies inparticular groups of the tribe are rare (e.g.Razafimandimbison et al., 2009) and many taxonomicproblems are still to be solved. Furthermore, many

*Corresponding author. E-mail:[email protected]

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Botanical Journal of the Linnean Society, 2013, 173, 407–441. With 5 figures

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species remain to be described from Central and WestAfrica, but also from Madagascar and Asia.

The main aim of this study is to elucidate thephylogenetic position of the African genus CuvieraDC., especially in relation to Vangueriella Verdc.Cuviera was described by de Candolle (1807) with onespecies, C. acutiflora DC. It was regarded as closelyrelated to Vangueria Juss. because of the solitarypendulous ovules in each locule. Further species wereadded 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 GlobulostylisWernham with two species, G. minor Wernham andG. talbotii Wernham, and later added a third species,G. cuvieroides Wernham (Wernham, 1918). AlthoughCuviera and Globulostylis clearly belong to the tribeVanguerieae, they were not addressed in the mono-graph of the tribe by Robyns (1928).

Hallé (1960) published a conspectus of Cuviera,including studies of inflorescence, style and pyrenemorphology, and a key to the species then known. Henoted that certain Cuviera spp. have a hairy swellingat the base of the style and suggested an infragenericgroup. Verdcourt (1987) saw the resemblance of thesespecies with Globulostylis and treated the latter as asubgenus 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–Congolianregion of Africa, but four species, poorly known and ofquestionable generic position, are found in East Tropi-cal Africa (Verdcourt & Bridson, 1991; Bridson, 1998).So far only C. physinodes K.Schum. (misidentified asC. angolensis Welw. ex K.Schum.) has been included inmolecular 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 subgenusBrachyanthus Robyns (Robyns, 1928) to generic rank.Verdcourt considered Vangueriella to be related toCuviera. Molecular analyses (Lantz et al., 2002; Lantz& Bremer, 2004) rather suggested an affinity to Can-thium Lam., but the only species included (V. spinosa(Schumach. & Thonn.) Verdc.) is atypical. We there-fore found it necessary to add data for Vangueriellawhile investigating Cuviera.

The taxonomic problems in Cuviera do not onlyinvolve generic delimitation, but frequent misidenti-fications between the species also occur in literatureand even more so in herbaria. For example, in theChecklist of Gabonese vascular plants (Sosef et al.,

2006), three out of ten species were misidentified. Thenames C. acutiflora DC., C. leniochlamys K.Schum.,C. longiflora Hiern and C. minor C.H.Wright, in par-ticular, have been commonly misapplied. A taxonomicrevision of this group was therefore necessary and isprovided here.

MATERIAL AND METHODS

To test the monophyly and systematic position ofCuviera and Vangueriella, a molecular phylogeneticstudy has been performed based on seven DNAregions (trnTL, trnLF, rpl16, petD, ITS, rpl32-trnLand accD-psaI). Ninety-two specimens of Vanguerieaerepresenting the different groups recognized in thetribe were included in the phylogenetic analysis, aswell as three outgroup species. Detailed informationon the investigated taxa can be found in Appendix 1.DNA of silica-dried and herbarium material wasextracted using the E.Z.N.A.TM HP Plant DNA MiniKit (Omega Bio-Tek). The primers for the amplifica-tion of the seven DNA regions can be found in Appen-dix 2, with their respective annealing temperatures.Purified PCR products were sent to Macrogen forsequencing (Macrogen Inc., Amsterdam, the Nether-lands). The sequences were assembled, edited andaligned using Geneious 5.4 (Drummond et al., 2011).Gaps were coded following Simmons & Ochoterena(2000). Maximum parsimony analysis was conductedusing PAUP* v.4.0b10a (Swoffort, 2002). Heuristicsearches were conducted with tree bisection–reconnection (TBR) branch swapping on 10 000random addition replicates, with five trees held ateach step. Non-parametric bootstrap analysis wascarried out to calculate the relative support for indi-vidual clades found in the parsimony analysis. Foreach of 1000 bootstrap replicates, a heuristic searchwas conducted with identical settings as in the origi-nal heuristic analysis. Phylogenetic trees were alsoestimated using probabilistic methods under Bayes-ian in the CIPRES web portal (Miller, Pfeiffer &Schwartz, 2010). Bayesian analysis was inferredusing MrBayes 3.1, running four Markov chains sam-pling every 100 generations for 5 million generations.DNA substitution models for the individual markerswere selected by preforming jModelTest 0.1.1 (Posada,2008) (Appendix 2). The concatenated data set waspartitioned and independent models were applied foreach of the partitions.

The taxonomic revision is based on the examinationof herbarium specimens in BM, BR, BRLU, E, HBG,K, LBV, MPU, P, WAG and YA. Central African col-lections of Cuviera from all these herbaria and WestAfrican collections from BM, BR, K and WAG, wereseen for this study. The distribution is based on thespecimens of the above-mentioned herbaria, comple-

408 B. VERSTRAETE ET AL.

© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 407–441

https://www.researchgate.net/publication/250676161_A_new_combination_and_key_to_the_species_of_Cuviera_subgenus_Globulostylis_Rubiaceae_Vanguerieae_from_Central_Africa?el=1_x_8&enrichId=rgreq-0b18bc98-1c94-484c-bff5-6a05dced3813&enrichSource=Y292ZXJQYWdlOzI1ODAyODE3NTtBUzoxMDQzNTcxMjMwNjc5MTBAMTQwMTg5MjA4NTQ4OQ==

https://www.researchgate.net/publication/271812698_Notes_on_African_Rubiaceae?el=1_x_8&enrichId=rgreq-0b18bc98-1c94-484c-bff5-6a05dced3813&enrichSource=Y292ZXJQYWdlOzI1ODAyODE3NTtBUzoxMDQzNTcxMjMwNjc5MTBAMTQwMTg5MjA4NTQ4OQ==

mented with data from the literature in a few cases(e.g. Hepper & Keay, 1963). Measurements, coloursand other details given in the descriptions are basedon living material, spirit and herbarium specimensand data derived from field notes. The dimensionsgiven 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 alpha-betically first by country (with a further distinctionfor Equatorial Guinea between Bioko Island and RioMuni) and then by collector, with the initials onlygiven in the case of ambiguity (e.g. J. J. F. E. de Wildeand W. J. J. O. de Wilde). The conservation status wasassessed by calculating the extent of occurrence or thearea of occupancy using the geographic informationsystem (GIS) and applying the 2001 IUCN Red ListCategories and Criteria.

RESULTS

The analysis of the combined data set using maximumparsimony and Bayesian inference yielded similartrees, of which the consensus tree is shown in Figure 1.Bootstrap values and posterior probabilities are indi-cated below the respective branches. Vanguerieae as awhole is monophyletic and strongly supported. Theconsensus tree is well resolved and contains severalstrongly supported clades, in correspondence with thefindings of Lantz & Bremer (2004).

1. The earliest branching clade contains the generaAfrocanthium, Keetia, Psydrax, Bullockia, Pep-onidium and Pyrostria, the last three forming thedioecious group of Lantz & Bremer (2004). In ouranalysis, Keetia is sister to the remainder of theclade and Afrocanthium is sister to the dioeciousgroup, whereas Lantz & Bremer (2004) foundthese two genera to be sister to each other. Rela-tionships in the dioecious group are not wellresolved; for a detailed investigation of this groupsee Razafimandimbison et al. (2009).

2. The so-called ‘spiny’ group of Lantz et al. (2002)containing most Canthium Lam. spp., Plectron-iella armata (K.Schum.) Robyns, Pygmaeotham-nus chamaedendrum (Kuntze) Robyns andVangueriella spinosa (Schumach. & Thonn.) Verdc.The name ‘spiny’ is somewhat misleading as spinytaxa are not uncommon in other clades as well(e.g. Cuviera and Rytigynia).

3. As in all previous studies of Vanguerieae, ourstudy finds the Fadogia Schweinf./RytigyniaBlume group as a clear and distinct clade in thetribe. The representatives of this group all haveunique indels in the investigated DNA regions.The internal topology of this clade is still unclear.

4. The focus of this study is on the Cuviera/Vangueriella group. It includes three well-supported subclades corresponding to Cuvierasubgenus Cuviera (excluding C. schliebenii Verdc.and C. semseii Verdc.), Cuviera subgenus Globulo-stylis and Vangueriella (excluding V. spinosa). Thelatter two subclades are weakly supported assister groups.

5. The Vangueria group includes Multidentia Gilli,Pygmaeothamnus zeyheri (Sond.) Robyns, Robyn-sia Hutch., Vangueria (in the enlarged sense ofLantz & Bremer, 2005), Vangueriopsis and twoEast African species of Cuviera (C. schliebenii andC. semseii). Vangueria is monophyletic with goodsupport but the rest of the clade is not wellresolved.

DISCUSSION

Our phylogenetic tree indicates several supragenericgroups with good support and these are not in con-tradiction with the findings of Lantz & Bremer(2004). The molecular data show a supported cladecontaining Cuviera s.l. (including subgenus Globulo-stylis) and Vangueriella (except V. spinosa) (Fig. 1). Inthis clade three distinct groups are strongly sup-ported: C. subgenus Cuviera, C. subgenus Globulosty-lis and Vangueriella.

In our opinion there are three possible options fordefining phylogenetic groups: (1) placing all species ina large genus Cuviera containing three subgeneraCuviera, Globulostylis and Vangueriella; (2) maintain-ing the taxonomy as it is now with the genera Cuvieras.l. and Vangueriella; and (3) reinstating Globulostylisas a genus and recognizing three genera. A majorinconvenience of an enlarged Cuviera is that it wouldbecome quite heterogeneous and difficult to definemorphologically; for example, in relation to Vangue-ria. Even in the current sense, Cuviera is quiteheterogeneous and not easily separated from Van-gueriella based on morphological characters. In addi-tion, the monophyly of Cuviera + Globulostylis is notsupported by molecular data. We favour option 3,which allows the formation of more homogeneous(and molecularly well-supported) groups (see Table 1for distinguishing characters); the problematic case ofCuviera le-testui Pellegr. is discussed below. We there-fore recognize three genera, Cuviera, Globulostylisand Vangueriella. These are briefly discussed further,as well as species excluded from Cuviera.

CUVIERA SENSU STRICTO

Cuviera, now considered in a narrower sense, isrevised here and includes ten species from West and

REVISION OF CUVIERA AND GLOBULOSTYLIS 409


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.



Central Africa. Most of these species form a ratherhomogeneous group that is well characterized, in par-ticular by their striped petals, many-flowered inflo-rescences and (usually) ant holes in the twigs. Thebracts are recaulescent (Hallé, 1960), which meansthey are moved to the upper node. Therefore, the firstnode of the inflorescence is bare and the lowest bractsare inserted at the second node (or on the lateralpedicels if the inflorescence is unbranched).

Three species are more or less atypical and needsome discussion. Cuviera trilocularis Hiern lacks antholes in the twigs (which is sometimes also the case inthe type species, C. acutiflora) and has few-floweredinflorescences, but matches Cuviera in other charac-ters (e.g. style, bracts). We did not obtain enoughDNA data of this species to include it in the generalanalysis, but when analysing the trnL-F marker,C. trilocularis clearly belongs to the Cuviera s.s.clade.

Cuviera pierrei also has few-flowered inflorescencesand the style is thickened in the upper two-thirds,which is reminiscent of Globulostylis (although in thelatter the style swelling occurs in the lower half).Presumably for these reasons, it was included insubgenus Globulostylis by Onana (2008). However,the striped corolla lobes, twigs with ant holes and thelowest node of the inflorescence lacking bracts (theseare inserted halfway on the pedicels of the lateralflowers) are clear differences with Globulostylis andplace this species in Cuviera s.s.

Cuviera le-testui is certainly the most aberrantspecies (Table 1). It is presumably related to Cuvieras.s. in having ant holes in the twigs, many-floweredinflorescences and corolla lobes longer than the tube,

but differs in its bilocular ovary, plain green corollalobes and in having a pair of bracts at the lowest nodeof inflorescence (the other bracts are recaulescent andtherefore the second nodes of the inflorescence arebare; Hallé, 1960). Its peculiar characters could welljustify the creation of a new genus, but we think thiswould be premature given the lack of genetic data. Wetherefore provisionally keep it in Cuviera s.s.

GLOBULOSTYLIS

Globulostylis is reinstated with eight species, includ-ing three newly described (G. dewildeana, Fig. 2;G. rammelooana, Fig. 3; and G. robbrechtiana, Fig. 4)and two newly transferred from Cuviera (G. leni-ochlamys (K.Schum.) Sonké, O.Lachenaud & Desseinand G. uncinula (N.Hallé) Sonké, O.Lachenaud &Dessein). The main characters of Globulostylis are thefew-flowered inflorescences with a pair of bracts atthe apex of the peduncle and the style with a swelling(sometimes rather slight) in the lower half. The genusis quite homogeneous; the only slightly atypicalspecies is G. cuvieroides Wernham, which has narrowcalyx lobes.

VANGUERIELLA

A remarkable finding is that Vangueriella is polyphy-letic; most of the species studied [including V. laxi-flora (K.Schum.) Verdc., the type of the genus] form awell-supported clade related to Cuviera but V. spinosa(Schumach. & Thonn.) Verdc. (the type species ofsection Stenosepalae) is related to Canthium, asalready found by Lantz & Bremer (2004). This is not

Table 1. Main distinguishing characters between Vangueriella, Globulostylis and Cuviera

Vangueriella Globulostylis Cuviera s.s. C. le-testui

Ovary Bilocular Pentalocular Pentalocular BilocularInflorescences Many-flowered One- to five-flowered Three- to

many-floweredMany-flowered

Bracts at lowestnode

Present/absent Present Absent Present

Corolla lobes(shape)

Narrow, at most equallingtube (occasionalappendages notincluded)

Rather broadly triangular,variously shorter orlonger than tube

Narrow, longer thantube

Narrow, longerthan tube

Corolla lobes(colour)

Plain green or yellow Plain green or yellow Striped (see note underC. trilocularis)

Plain green

Style Slender Swollen in lower half(sometimes slightly so)

Slender, rarely(C. pierrei) thickenedin upper two-thirds

Slender

Ant holesin twigs

Absent Absent Usually present Present



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)].



Figure 3. Globulostylis rammelooana. A, flowering branch. Bracts are represented by dotted lines. B, stipule. C, flowerbud. 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)].



Figure 4. Globulostylis robbrechtiana. A, flowering branch. Bracts are represented by dotted lines. B, stipule. C, flowerbud. 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)].



so unexpected as V. spinosa (and several other speciesof section Stenosepalae) differ quite markedly fromthe rest of the genus in floral characters, lacking aring of deflexed hairs inside the corolla tube andhaving 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. ofsection Vangueriella (which it indeed closely resem-bles in flower, leaf and fruit). This species has thecalyx lobes narrower than usual in section Vangueri-ella, but still clearly foliaceous, and the corolla tubehas a ring of deflexed hairs inside, as in sectionVangueriella.

This shows that the circumscription of Vangueriellahas to be redefined, which will be the object of afuture study. Most species of section Stenosepalae willprobably have to be transferred to Canthium. Lantz &Bremer (2004) already transferred V. spinosa (as Can-thium stenosepalum Lantz), but this has not beenwidely 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 genusbased on morphological and/or molecular data, but atpresent cannot be transferred to other genera. Theseare C. calycosa Wernham, C. nigrescens (Scott-Elliottex Oliv.) and the four East African species (C. midge-odii Verdc., C. schliebenii Verdc., C. semseii Verdc.and C. tomentosa Verdc.). All these species differ fromboth Cuviera s.s. and Globulostylis in having theinflorescences borne on old woody twigs below theleaves. However, they do not form a homogeneousgroup and further studies, especially in the Vangueriagroup, are crucial to understand their position.

Cuviera calycosa, C. nigrescens and C. semseii areclosely related and, in fact, may even be conspecific.The first two were already indicated as the leasttypical species of the genus by Hallé (1960). Theyhave a pentalocular ovary, as usual in Cuviera, butdiffer in (1) inflorescence position, (2) pyrenes with asharp dorsal crest, (3) small, black-drying leaves withtuft domatia in nerve axils, and (4) twigs that becomewoody soon, with the bark peeling off. The molecular

data (for C. semseii only) suggest an affinity withVangueria. Further studies are required before thesespecies can be transferred to either Vangueria or anew genus.

Cuviera schliebenii and C. tomentosa have a bilocu-lar ovary and small flowers, two characters reminis-cent of Vangueriella, but the molecular data forC. schliebenii rather suggest an early branching posi-tion in the Vangueria group. Until the generic rela-tionships in this group are better understood, wecannot suggest a generic placement.

The last species, C. migeodii, is only known fromthe poor type and is therefore difficult to place, but isclearly not a Cuviera s.s. The small, black-dryingleaves with tuft domatia resemble C. nigrescens, butthe ovary is described as bilocular.

In conclusion, we here reinstate Globulostylis as agenus and we describe three new species, bringing thetotal to eight species. The remaining ten species areplaced in Cuviera s.s. Four East African species,C. calycosa and C. nigrescens are aberrant and areexcluded from Cuviera. Vangueriella is shown to beparaphyletic with V. section Stenosepalae relatedto the ‘spiny’ group and V. section Vangueriella toGlobulostylis.

TAXONOMIC TREATMENT

Keys to the genera Cuviera and Globulostylis to theirrespective species and a detailed species list, includ-ing the description of the three new Globulostylis spp.(G. dewildeana, G. rammelooana and G. robbrech-tiana) are presented. Species excluded from Cuvieraare 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, ever-green, with regular horizontal branching (Roux’sarchitectural model); vegetative parts glabrous orrarely puberulent; twigs usually with swollen and

KEY TO CUVIERA AND 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 usuallywith a median stripe ............................................................................................................Cuviera

1’. Inflorescence one- to five-flowered with a pair of bracts at lowest node (often caducous in fruit); styleswollen in the lower half (sometimes slightly so); twigs without ant holes; corolla lobes plain green oryellow, not striped .........................................................................................................Globulostylis



hollow internodes, inhabited by ants (solid in C. tri-locularis and occasionally in C. acutiflora); youngplants and coppicing shoots frequently armed withpaired supra-axillary spines that sometimes persiston the trunk. Leaves opposite, ± elliptic, shortly peti-olate; stipules small, ± basally connate, not or shortly(< 2 mm) awned, caducous to ± persistent. Inflores-cences axillary and paired at each node on youngtwigs, cymose, three- to many-flowered; bracts linearto broadly elliptic, ± leafy, but absent at lowest nodeof inflorescence (except in C. le-testui), inserted at thebase of pedicels (or towards the middle in C. pierrei).Flowers hermaphrodite. Calyx lobes three to five, freeor shortly connate at base, ± leafy, linear to broadlyovate and usually longer than the corolla, or rarelytruncate calyx (C. truncata). Corolla tube campanu-late to nearly cylindrical, glabrous outside, with aring of reflexed hairs near the middle inside; lobesfive, longer than tube, elongate, acute to long-apiculate 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 fullyexserted (frequently reflexed between the lobes).Ovary (bi-) pentalocular, obconic or turbinate, some-times five-angled; ovules solitary in each locule, pen-dulous. Disk ± flat, glabrous or hairy in the centre.Style exserted, narrowly cylindrical or (in C. pierrei)thickened in the upper two-thirds, stiffly pubescent toglabrous; stigmatic club ± mitriform, shortly (two-)five-lobed. Fruits drupaceous, apparently remaininggreen, large 2.0–3.7 × 1.8–3.3 cm, subglobose toobovoid, smooth to variously lobed, with calyxgenerally persistent (sometimes tardily caducous);pyrenes (two-)five, one-seeded, shaped like a manda-rin orange segment, usually with a faint dorsal crest,and a slight ventral indentation towards the upperthird.

Distribution and ecology: Cuviera has ten species inthe Guineo–Congolian rainforest zone (including BiokoIsland). The centre of diversity is in Lower Guinea;only two species occur in Upper Guinea (C. acutifloraand C. macroura) and only one extends into Congolia(C. angolensis). Most species are relatively lightdemanding, favouring secondary or riverine forest,although some can be found in the understory aswell. They are frequently gregarious, especiallyC. physinodes that may form dense thickets locally.

Notes: The most important characters for speciesidentification in Cuviera are the calyx lobes, thecorolla and, to some degree, the fruits. Leaves are oflimited value, as in many species they are quitevariable; for example, the leaf base can be acute tosubcordate. However, an experienced researcher can

identify most of the sterile collections if the origin istaken into account.

Several aspects of the biology of Cuviera would beinteresting to study in the field, in particular theassociation with ants, which presumably play a rolein protecting the plant (although in our experiencethey are not aggressive to humans). Whether differ-ent Cuviera spp. have different associated ants is notknown.

Not much is known about the pollination of theflowers, which are quite attractive and often fragrant(e.g. those of C. macroura smell of honeysuckle), orabout the dispersal of the fruits (although, consider-ing the size of the pyrenes, large animals are presum-ably involved).

Cuviera spp. are rarely used by humans, althoughat least two species (C. le-testui and C. longiflora)have edible fruits, and the leaves of C. angolensissubsp. latior are eaten as a vegetable in the Demo-cratic 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., Smeath-man 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-eastof Suacoco, Traub 299 (holotype: BR000000882973!;isotypes: BM000903517!, BR000000882965!, MO n.v.).

Distribution: Lower and Upper Guinea Domain: Cam-eroon, Equatorial Guinea (Rio Muni), Ghana (seeNotes), Guinea, Ivory Coast, Liberia, Nigeria, SierraLeone and Togo (see Notes).

Notes: Cuviera acutiflora is closely related toC. physinodes and C. truncata; the three species sharecoriaceous leaves and glabrous, shortly exserted styles.

Cuviera acutiflora is a widespread and polymorphicspecies, especially in West Africa where the leaf basevaries from cuneate to asymmetrically cordate; speci-mens from Cameroon and Equatorial Guinea alwayshave cordate leaf bases (which allows them to beseparated from C. physinodes). Ant holes in the twigscan be present or absent, apparently irrespective oflocality. Some Nigerian specimens are intermediatewith 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 ofC. acutiflora from Gabon (Sosef et al., 2006) refer toC. physinodes.



The typification of C. acutiflora needs further inves-tigation. Two Smeathmann sheets in BM are probablypart of the original gathering, but it is not clearwhether De Candolle saw them. There might be othersheets in P, where De Candolle worked, so we havenot 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 Oct1969, Bos 5556 (BR, K, P, WAG, YA); Ibid., 9 Nov1970, Bos 6118 (BR, K, P, WAG, YA); km 20, Campo-Kribi, 10 Nov 1970, Bos 8706 (YA); ibid., 4 Dec 1975,

KEY TO THE CUVIERA 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, ifpresent, never reflexed .................................................................................................................2

2. Calyx truncate or with short (< 2 mm) irregular lobes (Nigeria and south-west Cameroon) .C. truncata2’. Calyx lobes conspicuous, 4 mm long at least (sometimes caducous on fruit) ....................................33. Calyx lobes three(–four), the fourth lobe when present often slightly smaller; style hairy, long exserted

.................................................................................................................................................. .43’. Calyx lobes five ...........................................................................................................................54. 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. angolensis4’. Calyx lobes ovate, conspicuously widened above the base, 2.5–23.0 mm wide (Gabon to north-east

Angola) ...................................................................................................C. angolensis subsp. latior5. Leaves thick and coriaceous, tertiary nerves invisible or lax; corolla lobes not (or shortly) apiculate ..

.................................................................................................................................................. .65’. 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) ............................................... .96. Inflorescence three- to five-flowered, very lax; twigs lacking ant holes (Nigeria and south-west Cam-

eroon) .......................................................................................................................C. trilocularis6’. Inflorescence many-flowered (but often with few fruits developing); twigs usually with ant holes,

sometimes lacking in C. acutiflora ................................................................................................77. 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 AfricanRepublic) .....................................................................................................................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; fruitbroadly 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 towardsbase; anthers erect; bracts and calyx lobes ovate to lanceolate, broader, (2-)3–8 mm; fruit narrowlyellipsoid, 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 presentas 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; domatiaabsent; 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), approxi-mately 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, alsoBioko) ...................................................................................................................... .C. subuliflora

10’. Inflorescence two- or three- (five-) flowered; bracts small (< 1 cm), much shorter than the sepals; corollalobes not appendaged; calyx lobes erect on fruit (Equatorial Guinea and Gabon) .................C. pierrei



J.J.F.E. de Wilde 8706 (BR, K, P, WAG); Bimbia–Bonadikombo community forest, 27 Apr 2009, Desseinet al. 2809 (BR, MO, YA); 8 km north of Kribi, 13 Feb1997, Elad 530 (WAG); Limbe, 1904, Kalbreyer 29 (K);Victoria [= Limbe], Nov 1929, Maitland 784 (BR, K);Ambas Bay, Feb 1861, Mann 770 (K); CameroonMountains, 1862, Mann 1211 (P); Bipindi, 25 Nov2004, 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 BairroIkundi, 23 Jun 1991, Carvalho 4749 (WAG); RioNdote, 7 Jun 1999, Eneme 304 (BRLU, WAG).

IVORY COAST. Km 25 Sassandra–Gagnoa, 30 Oct1968, Breteler 5860 (BR); near Abidjan, Banco ForestPark, 9 Jan 1976, de Koning 6377 (BR); c. 140 kmnorth of Tabou, 10 Oct 1963, W.J.J.O. de Wilde 1054(K); Abidjan, 24 Aug 1955, de Wit 801 (WAG); Adio-podoumé, Oct 1960, F. Hallé 188 (K); ibid., Oct 1960,F. Hallé 198 (BR); Abouabou, 9 Jan 1959, Leeuwen-berg 2348 (K); N Sékré, Oldeman 603 (K); Yapo, 28Oct 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, Jansen1129 (BR); Gibi Mountain, 5–10 miles south-east ofSalala, 15 Jan 1970, Jansen 1708 (BR); Bendu,10 miles north of Robertsport, 30 Jan 1970, Jansen1761, (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, Linder1539 (K); Gola–Yoma National Forest, 12 Nov 1965,van Meer 274 (BR); 27 km south of Zwedru, 22 Feb1966, van Meer 481 (BR); Paynesville, 8 miles east ofMonrovia, 23 Oct 1960, Voorhoeve 95 (BR).

NIGERIA. Unnokoo, 5 Nov 1967, Ariwaodo 1080(K); Epe, 28 Nov 1994, Daramola 589 (K); Milkenhill, 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 (siteof 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, 14Dec 1943, Onochie 7495 (P); Gambari ForestReserve, south-west Ibadan, 16 Jan 1958, Onochie &de Wit 678 (WAG); W Lagos, Rowland s.n. (P); UdiForest 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, 2Apr 1967, Gledhill 531 (BR); Freetown, 24 Jan 1883,Johnston 50 (K); Leicester Peak, 4 Dec 1891, Scott-Elliott 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, Welwitsch2564 (lectotype designated here: BM000903514!;syntypes: BM000903515!, BM000903516!,K000412037!, K000412038!, LISU208621!,LISU208622!, LISU208623!, M0106328!).

Notes: Cuviera angolensis is closely related to C. longi-flora, in particular in the long-exserted hairy style.However, it is hard to understand why Hepper & Keay(1963) synonymized the two species, which have dif-ferent calyces (C. angolensis: three or four narrowlylanceolate and acute lobes; C. longiflora: five broadlyelliptic and rounded lobes). The corolla lobes are alsogenerally longer and narrower in C. longiflora.

Cuviera angolensis is, however, closely related toC. latior and, after reviewing all available material ofboth taxa, we concluded that the latter is a subspeciesof the former. The only distinction between themconcerns the calyx lobes (see Key). The two subspeciesalso have separate ranges.

The specimen McPherson 16297 referred to asC. angolensis in previous phylogenetic studies (Lantzet al., 2002; Lantz & Bremer, 2004, 2005) is actuallyC. physinodes. Several Welwitsch collections made ondifferent dates were given the same number 2564, asis usual with this collector (Albuquerque, Brummit &Figueiredo, 2009). The lectotype chosen here was col-lected in April 1855 and a complete description of thenew species is attached to the specimen, hence thelectotypification.

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. CuanzaSul, Libollo Hochland, 1954–1955, Boss 5755 (BM);Calandula Mountain, Libulo, 18 May 1915, Goss-weiler 6314 (BM).

DEMOCRATIC REPUBLIC OF THE CONGO.M’vuazi, Malanga Forest, 23 Sep 1951, Devred 727



(BR); M’vuazi, Mansiesie Forest, 25 Sep 1951, Devred755 (BR, K); Massif of Matete (Mvuazi); Thysville, 15Jun 1954, J. Dubois 23 (BR); Nzengi Nzengi forest, 1Aug 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: DEMO-CRATIC REPUBLIC OF THE CONGO. NBoyeka, 31 Aug 1914, Nannan 139 (lectotypedesignated here: BR0000008829580!; isolecto-types: 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 (holo-type: 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-eastof 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, Demo-cratic Republic of the Congo (widespread except insouth), north-east Gabon (Ivindo basin), Republic ofCongo.

Notes: Unlike the rather uniform subsp. angolensis,subsp. latior is variable, especially in the width of thecalyx lobes (some specimens approaching subsp.angolensis) and in the size of the corolla lobe append-ages that vary from 6 mm long to nearly lacking.

The leaves are sometimes shortly pubescent on thenerves beneath, which is unique in Cuviera. Thischaracter may vary on a single specimen, so Hallé’s(1960) var. hispidula has not been maintained, norvar. evorombila (separated only on account of its lesspubescent style). The tertiary veins are typicallyobscure, but in a few specimens (e.g. Gilbert 14345,M. Laurent 669) they are apparent and rather dense.

Additional specimens examined: ANGOLA. Dundo, 2Jun 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, 24Nov 1922, Le Testu 4386 (BM, BR, P); Kpalata river,

Tisserant 437 (P); Boukoko, 14 Jan 1948, Tisserant614 (BM, P); Boukoko, 8 Mar 1951, Tisserant 2030(BM, P); Gbondo river, Pudjiya, 15 Jan 1928, Tisser-ant 2483 (P); Mbaïki, 5 Jan 1961, Tisserant 3692 (P).

DEMOCRATIC REPUBLIC OF THE CONGO.Epulu, 27 Feb 1998, Amsini 097 (BR); Island ofElelwa, 19 Apr 1960, Bamps 932 (BR, K); Metmas, 13Jan 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 Kisan-gani, 9 Feb 1971, Bokdam 3085 (WAG); ibid., 1 May1971, Bokdam 3180 (WAG); Yangambi–Yaselia road,25 Jun 1963, Bolema 1144 (BR); Yangambi, 17 Jan1961, 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, 10Aug 1956, Christiaensen 1818 (BR, K); Eala,Corbisier-Baland 1399 (BR); ibid., Corbisier-Baland1524 (BR); ibid., Corbisier-Baland 1717 (BR);Gangala na Bodio, 24 Mar 1943, Cornet D’ Elzius DuChenoy, Lauwers & Offermann 808 (BR); Yangambi,1948, Gilbert 8424 (BR); Eala, 10 Sep 1937, Coûteaux325 (BR); Kiyaka, 11 Jul 1955, Devred 2140 (BR);Bas-Uele, 5 Oct 1934, Dewulf 203 (BR); ibid., 27 Dec1934, Dewulf 523 (BR); Yangambi, 3 Oct 1950, Donis2847 (BR); ibid., 8 Jan 1952, Donis 3336 (BR); ibid.,30 Jan1952, Donis 3501 (BR); Boyagidigba, 23 Apr1955, Evrard 755 (BR); Mangala (Terr. Isangi), 19Mar 1957, Evrard 2252 (BR); Befale, 2 Jan 1958,Evrard 3179 (BR, K); Eandja, 13 Feb 1958, Evrard3423 (BR, K); Djolu (Bolomba), 20 Feb 1959, Evrard5791 (BR, K); [Yionda], 15 km south of Mbandaka(Zaire), 18 Feb 1987, Evrard 10688 (BR); Epulu, 16Nov 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), 6Dec 1956, Gerard 2570 (BR); Bambesa, 31 Jan 1958,Gerard 3232 (BR); Bambesa, 12 Nov 1957, Gerard3300 (BR); Tukpwo, 12 Feb 1958, Gerard 3540 (BR);Bambesa, 8 Jan 1960, Gerard 4015 (BR); ibid., 14 Nov1963, 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, Mar1924, Goossens 4641 (BR); Busanga, Feb 1953, Gor-batoff 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);



Bambesa, 11 Jan 1940, Hendrickx 869bot (BR);Bolima, 31 Oct 1941, Hulstaert 471 (BR); Bokote,1943, Hulstaert 953 (BR); Boende, 28 Jul 1944, Hul-staert 1354 (BR, K); s.loc., Jans s.n. (BR); bank ofCongo river, near Bolombe, 2 Jan 1904, M. Laurents.n. (BR); Eala (Ikakéma), 29 Jan 1904, M. Laurents.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 Oct1906, 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); betweenLibenge and Gemena, Lebrun 1871 (BR, K); ibid.,Lebrun 2186 (BR); between Niangara and Wamba,Jul 1931, Lebrun 3228 (BR); between Walikale andKalehe, Mar 1932, Lebrun 5240 (BR); Urega, Jun1932, Lebrun 5634 (BR); between Kindu and Kata-kokombe, Lebrun 6073 (BR, K); Eala, 1936, Leemans295 (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, Lejoly1928 (BR); Kisangani, Island of Mayele near Wagenia,13 May 1978, Lejoly 3572 (BR); Kisangani, 13 Nov1981, 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. Léonard1020 (BR); Kikwit, 28 Jul 1904, Lescrauwaet 165(BR); road Hemptinne-Kamwandu (Terr. Dibaya), 21Jun 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, LisowskiB9565 (BR); Yangambi, 13 Dec 1935, J. Louis 823(BR, K, P); île ‘Esali’, en face de Yangambi, 13 Feb1936, J. Louis 1255 (BR, K); north-west Eala, 20 May1936, 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. Louis12583 (BR, K); Opala, J. Louis 14137 (BR); ÎleLotumba, 5 Apr 1939, J. Louis 14483 (BR); Lokutu, 29Oct 2004, Luke 10412Z (BR); Likimi, 13 Feb 1910,Malchair 96 (BR); Kikwit/Erco, 21 Nov 1990, Masens569 (BR); Bikoro, 8 Aug 1983, Nsola 229 (BR); Mabali,7 Apr 1985, Nsola 845 (BR); Kapanga, 1934, Overlaet1216 (BR); km 82 road Kavumu–Walikale, 27 Sep1955, 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 Mar1988, 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., VanDer Gucht 131 (BR); Saint Trudon, Van Kerkhoven824 (BR); ibid., 18 Aug 1913, Van Kerkhoven s.n. (BR);Ipamu, Jul 1922, Vanderyst 9804 (BR); ibid., Sep1921, Vanderyst 10607 (BR); ibid., Sep 1921, Vander-yst 10907 (BR); ibid., Oct 1922, Vanderyst 12293 (BR);between Lubue 1 and Luange, Oct 1922, Vanderyst12486 (BR); Hemptinne Saint Benoît, Vanderyst23677 (BR); Ibid., Vanderyst 23644/4 (BR); Mobwasa,28 Apr 1914, Vermoesen 324 (BR); Bambesa, 11 Jan1940, Vrydagh 68 (BR).

GABON. Ivindo, Makokou, 20 Nov 1985, Caraglio56 (MPU); Ipassa, 10 km south of Makokou, 21 Jul1978, Florence 1556 (P); Bélinga, Basse Nounah, 1Dec 1964, N. Hallé 3381 (P).

REPUBLIC OF THE CONGO. Dongou, 28 Jan1966, Bouquet 2099 (P); Odzala National Park, campMboko, 9 Dec 1993, Diafouka 558 B (BRLU); ibid., 17Nov 1994, Dowsett-Lemaire 1826 (BR); ibid., 21 Nov1996, 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, 13Apr 1925, Le Testu 2053 (lectotype designatedhere: P00553438!; isolectotypes: BM!,BR0000008846822!, BR0000008846839!,K000412042!, LISC002648 n.v., P00553436!,P00553437!).

Distribution: Lower Guinea Domain: DeomacraticRepublic of the Congo (Bas–Congo), EquatorialGuinea (Rio Muni), Gabon (Libreville area, DoudouMountains, Chaillu Mountains), Republic of Congo(Mayombe).

Notes: Cuviera le-testui is an aberrant species stand-ing well apart from its congeners in having a pair oflarge bracts at the apex of the peduncle and a bilocu-lar ovary. Another remarkable character is the redexudate in the bark mentioned by two different col-lectors (A.M. Louis 1684 and Senterre & Obiang4315). This is exceptional in Rubiaceae, although italso occurs in a new species of Psydrax Gaertn. (cur-rently under description).

Cuviera le-testui is newly reported from the Repub-lic of Congo and Equatorial Guinea. Material from the



latter country is sterile, but the identification makeslittle doubt.

Additional specimens examined: DEMOCRATICREPUBLIC OF THE CONGO. Kimbuya–Kingediroad, 24 Aug 1959, Compère 154 (BR, K).

EQUATORIAL GUINEA. Rio Muni: Monte Bata,1 km north of road Bata–Niefang at Santa Marta, 8Aug 2003, Senterre & Obiang 4297 (BRLU); ibid., 9Aug 2003, Senterre & Obiang 4315 (BRLU).

GABON. Mourindi, 15 Sep 2000, Bourobou Bouro-bou 247 (BR, K, P, WAG); Etéké, 15 May 1963, N.Hallé & Cours 5907 (P); Tchibanga, 2 May 1914, LeTestu 1720 (BR, BM, P); Mouloumbi, 1 Aug 1930, LeTestu 8211 (BR); Malibé, 1 km toward west, 7 May1985, A.M. Louis 1684 (WAG); near Igotchi–Mouenda,11 May 1997, McPherson 16942 (K, MO); km 23 roadsouth-east of Igotchi–Mouenda, 19 May 1997,McPherson 17047 (BR, K, MO); c. 10 km north ofLibreville, 23 Apr 1985, J.M. Reitsma & B. Reitsma811 (WAG); Mourindi, 16 Apr 2000, Sosef 1312(BRLU, WAG).

REPUBLIC OF THE CONGO. Sounda, bank ofKouilou, 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 Cam-eroon, Jan 1862, Mann 1211 (lectotype designatedhere: K000412049!; isolectotype: K000412050!).

Distribution: Lower Guinea Domain: Cameroon(widespread in the west, centre and east), CentralAfrican Republic (extreme west), Equatorial Guinea(Bioko Island), north-east Gabon (Bélinga), eastNigeria.

Notes: Cuviera longiflora is closely related to C. ango-lensis, but we cannot follow Hepper & Keay (1963) inuniting them; they especially differ in the number ofcalyx lobes (see C. angolensis) and also in the range.

The type of C. longiflora, Mann 1211 (K) has twosheets, one of which is here selected as lectotype. Notethat Mann 1211 (P) represents another species,C. acutiflora.

The species can be considered as new to Gabon, asthe two specimens cited for this country by Sosef et al.(2006) are both misidentified: McPherson 16297 isC. 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 kmsouth-west of Yaoundé, trail to Eloumden Mountain,29 Dec 1962, Breteler et al. 2318 (BR, K, P, WAG);

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, Etuge3410 (K); Laikom, 3 Dec 1998, Etuge 4538 (K); Ndi-kiniméki, 24 Dec 1927, Hédin 3 (P); Nkambe, 10 mileswest, 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 ForestReserve, 7 May 1960, Keay 37925 (BR, K, YA); Kupevillage, 17 Jan 1995, Lane et al. 298 (K); Mount Kupe,Ndum, 31 Jan 1995, Lane et al. 460 (K); km 17 BetaréOya–Meiganga, 5 km south-east of Ndokayo, 3 Nov1966, Leeuwenberg 7697 (BR, K, P, WAG, YA); bank ofSanaga, 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); nearNgoila, 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, Letouzey13450 (P, YA); Tinachong, 30 km west-north-west ofBamenda, 10 Aug 1975, Letouzey 14234 (BR, K, YA);bank of Boumba river, 14 km west-south-west of Kin-sassa, 7 Mar 1971, Letouzey & Villiers 10525 (BR, P,YA); Mount Oku, Lumutu Forest, 1 May 1998,Maisels 116 (YA); Lakom, Bamenda, May 1938, Mait-land 1361 (K); Mount Cameroon, Jan 1862, Mann1212 (K); Mapanja, 16 Mar 1992, Mbani 55 (K);between Yaoundé and Deng Deng, c. 170 km north-west 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, 14Jan 1985, Nkongmeneck 892 (YA); Bafut–NgembaForest Reserve, 19 Mar 1955, Richards 530 (K, P);Small Massaka (Rumpi hills), 19 Dec 2009, Sonkéet al. 5442 (BR, BRLU, K, MO, P, WAG, YA); NationalPark Mbam Djerem, 23 Jan 2011, Sonké et al. 5510(BR, BRLU, K, MO, YA); Bertoua–Batouri, 15 May1905, Tchinaye 111 (P, YA); south slope of MountCameroon, above Batoke, 29 Dec 1983, D.W. Thomas2844 (BR, K, P, YA); above small Koto village, 6 Mar1985, 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, Ujor30416 (K); east slope Mount Cameroun, 6 km east ofBomana, 34 km north-west of Limbe, 11 Dec 1984,Villiers 2447 (BR, YA); Mapanja, 19 Feb 1992, Wheat-ley 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).



EQUATORIAL GUINEA. Bioko: Gran Caldeira deLuba, along the river towards Mount Pisarro, Car-valho 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 May1971, 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 doubtfulrecord from Equatorial Guinea (Bioko).

Notes: Cuviera macroura is unique in the genus inhaving small glabrous pits in the nerve axils (some-times hard to see), a calyx with subulate lobes dis-tinctly fused at the base, and stipules with a broadrounded lobe (see photographs in Hawthorne &Jongkind, 2006). The species is rather variable, speci-mens from Cameroon and Nigeria generally havesmaller 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!). Bothsheets are labelled ‘Fernando Po’ [= Bioko], but theirorigin is somewhat doubtful, for Bentham (in Hooker,1849) states in the protologue of C. subuliflora: ‘Fer-nando Po, on the seashore, and, apparently the samespecies, at Abòh, growing in the water, Vogel’. Thelatter locality is in Nigeria. There are no other recordsof C. macroura on Bioko and its occurrence there is tobe regarded as doubtful.

Cuviera djalonensis A.Chev. (in Explor. Bot. AfriqueOcc. Franç.: 331. 1920.) is a nomen nudum referringto C. macroura (Hallé, 1960).

Additional specimens examined: BENIN. Lokoli, 18Dec 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, nearEdéa, 26 km north of Kribi, 18 Apr 1965, Leeuwen-berg 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 kmnorth of Douala, 9 4 1976, Letouzey 14751 (BR, P, YA);Douala–Edéa Reserve, bank of Kombe river, 3 Aug1978, 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, MortonA468 (K); Kumasi, Jun 1929, Vigne 1758 (K); s.loc.,s.d., Vigne 3008 (BM).

GUINEA. Gouam–Geasso, 23 Apr 1956, Adam12092 (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); betweenMount Nienokoue and Hana river, Taï–Tabou road, 11Aug 1962, Guillaumet 1519 (BR); Adiopodoumé, 16Aug 1961, F. Hallé 219 (BR, K); near the debarcationof the ferry from Grand Bassam to Bingerville, 24 Jul1969, 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 Oct1926, Linder 1035 (K).

NIGERIA. s.loc., Hambler 409 (K); Ibarapa, 22 Aug1966, Latilo 58826 (K, P); 19 miles from Imo river, 12Aug 1966, Okafor 60323 (K); Ikorodu, 29 Dec 1952,Onochie 32038 (K); Degema, 1916, Talbot 3632 (BM,K).

SIERRA LEONE. Mussaea, 31 Aug 1963, Haswell150 (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 desig-nated here: P03818077!; isolectotype:K000412041!); Sibange Farm, Munda, 25 Oct1881, Soyaux 307 (syntype: K000412040!).

Synonyms: Cuviera ledermannii K.Krause, Bot.Jahrb. Syst. 48: 418. 1912. syn. nov. Type: CAM-EROON. Nkolebunde [= Elephant Mountain], Oct1908, Ledermann 725, 751 (holotype: B†); ElephantMountain, 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 (RioMuni), Gabon (widespread except in the east).

Notes: This species has been overlooked since itsdescription, possibly because the Index Kewensisgives a wrong reference for the protologue. Cuvieraphysinodes is closely related to C. acutiflora and fre-quently 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-



eroon, Equatorial Guinea) can be set apart fromC. acutiflora by their cuneate leaf base (alwayscordate in local populations of C. acutiflora).

The description of C. physinodes is based on twosyntypes, Soyaux 29 (with flower buds) and Soyaux307 (with young fruits). The P sheet of the formerbears the number 23, which is evidently a copyingerror: Soyaux 23 (K) was collected on 21 Aug andis an isosyntype of Pavetta brachycoryne K.Schum.(= Rutidea glabra Hiern). In spite of this, we selectthis sheet as lectotype, as it is by far the better.

Krause (1912) already considered C. ledermannii asclosely related to C. physinodes and his good descrip-tion 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. Loberiver, 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 Dec1968, Bos 3426 (BR, K, P, WAG, YA); near Mbikilikivillage, 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 ForestReserve (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, 19Apr 2001, Ngomo 916 (BRLU); south-west part ofMonte Alén National Park, 2 km north-east of RioUolo, 24 June 2002, Senterre & Nguema 3018(BRLU); Monte Alén National Park, 12.5 km east ofMosumo, 8 Sept 2003, Senterre & Obiang 4071(BRLU).

GABON. Toucan, 11 Oct 2002, Bourobou et al. 990(WAG); km 34 Mouila–Yeno, 22 Sep 1986, Breteleret 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, Degreef181 (BR); Benga, Klaine 82 (BR, K, P); Owendo, 10Nov 1903, Klaine 3340 (K, P); road to Cap Estérias onthe right, 11 Sep 1985, A.M. Louis 1805 (WAG); Forêtdes Abeilles, east Lopé Reserve, 3 Dec 1993, McPher-son 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 Jul1986, J.M. Reitsma & B. Reitsma 2463 (WAG);Doudou Mountains National Park, c. 30 km south ofMandji, 18 Nov 2005, Sosef 2373 (WAG); Pechoudroad, southwards, 27 Oct 1990, Van Nek 123 (WAG);Cap Esterias, 1896, Klaine 538 (P); Mondah Forest,30 Aug 1969, Villiers s.n. (P).

7. CUVIERA PIERREI N.HALLÉ, Bull. Soc. Bot. France106: 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: EquatorialGuinea (Rio Muni), Gabon.

Notes: This species is unusual in Cuviera in having(two-)three- (five-) flowered inflorescences and thestyle 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 resem-blance to Globulostylis cuvieroides, because of theshape of the calyx lobes, however, the striped corollalobes, the position of the bracts (inserted halfway onthe pedicels of the lateral flowers rather than at theapex of the peduncle), and the presence of ant holes inthe twigs are clear distinctions.

The species is newly reported from EquatorialGuinea. A record from the Democratic Republic of theCongo (Compère, 1962) is an error for C. subuliflora(see that species).

Additional specimens examined: EQUATORIALGUINEA. Rio Muni: Abama (Nsork), 22 Aug 2001,Esono, Merino & Gomez 289 (BRLU); Ngoma, 15 kmsouth-east de Etembue, 12 Aug 1998, Lejoly & Elad98/158 (BRLU).

GABON. 9 km south of Kinguélé, 21 Sept 1994,Breteler et al. 12980 (WAG); Mount Mela, 26 Aug1978, 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); Libre-ville, 25 Mar 1904, Klaine 48 (BR, K, P, WAG); ibid.,12 Feb 1896, Klaine 378 (P); ibid., 15 Oct 1896, Klaine558 (P); ibid., 25 Jul 1897, Klaine 558bis (P); ibid., 24Oct 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, Wilks1299 (WAG).

8. CUVIERA SUBULIFLORA BENTH., in W. J. Hooker,Niger Fl.: 407. 1849. Canthium subuliflorum(Benth.) Roberty, Bull. Inst. Franç. Afrique Noire16: 60. 1954. Type: EQUATORIAL GUINEA.Bioko, Vogel 83 (lectotype designated here:K000412045!); Excluded isolectotype: Bioko, Vogel83 (K000412044!) is Cuviera macroura.

Synonyms: Cuviera plagiophylla K.Schum., Bot.Jahrb. Syst. 33: 353. 1903. Type: CAMEROON.



Bipindi, 1897, Zenker 1350 (lectotype designatedhere: K000412043!, isolectotypes: BM!,BR000000884676!, HBG, L0058090 n.v., MA311021n.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 ofCongo.

Notes: The type of C. subuliflora, Vogel 83, has twosheets, one corresponding to the species hithertonamed C. subuliflora, the other being C. macroura. Topreserve current usage, we select the former sheet(K000412045!) as lectotype. The origin of these col-lections is somewhat doubtful (see note underC. macroura).

The fruits of C. subuliflora, with their broadly trun-cate apex and the calyx lobes patent and star-like, areunique in the genus. The style can be glabrous orpubescent, which explains discrepancies between thekeys in Hallé (1960) and Hepper & Keay (1963). Thisspecies is newly reported from the Democratic Repub-lic of the Congo (see C. pierrei above). A recordfrom Ghana (Hepper & Keay, 1963) is probably anerror for C. macroura (Hawthorne & Jongkind, 2006:628), but we could not verify the specimen. Twodifferent Talbot specimens bear the same number222: one collected in 1911 (C. subuliflora), the otherfrom 1909 (C. truncata).

Additional specimens examined: CAMEROON. Southof 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 & Thomas257 (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 duNtem, 40 km east-south-east of Campo, 10 Dec 1979,Letouzey 15342 (P, YA); Bimbia, Feb 1929, Maitland417 (K); Limbe, Oct 1929, Maitland 740 (K); Fenda,58 km east of Kribi, 1911, Mildbraed 6009 (HBG);Bibundi, Nov 1928, Mildbraed 10662 (K); Bonjare, 1May 1994, Ndam 1111 (K); Nndian–Dibonda–Ekumbako road, 24 Nov 1986, Nemba & D.W. Thomas398 (BR, K, MO, YA); Kumba–Mamfe road, 23 Jul1987, Nemba & D.W. Thomas 616 (K, MO); Kumbaarea, 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 (DjaFauna 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 & Nemba4049 (K, MO, P, YA); 10 km Ikata–Munyenge (north-east 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, Car-valho 2283-22 (K); s.loc., 1986, Carvalho 2521-3 (K);s.loc., 1986, Fernandez Casas 11544-3 (K); s.loc., Dec1860, Mann 87 (K); Rio Muni: Sendje, 13 Aug 2001,Esono & Ndong 304 (BRLU); Monte Alen, Ngomo &Ndong 232 (BRLU); Nkolentangan, 15 Feb 1908, Tess-mann 204 (K).

GABON. Km 4 Mbigou–Mimongo, 1983, J.J.F.E. deWilde 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 Aug1987, Dibata 274 (WAG); Moubigou 2, 21 May 1963,N. Hallé & Cours 6138 (P); around Libreville, 13 Jul1895, 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, MontsDoudou Reserve, 25 Mar 2000, Sosef 1004 (WAG);Saint Martin, Jul 1939, Walker 2 (P).

NIGERIA. Oban Group Forest Reserve, 11 Sep1960, Adebusuyi 44000 (K); ibid., 22 Nov 1961,Binuyo 45423 (K); Aponmu Forest Reserve, 7 Aug1963, 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 Meer892 (BR, K, WAG).

REPUBLIC OF THE CONGO. region of Kouilou(Bas–Kouilou), 26 Aug 1990, Moutsamboté & Dowsett-Lemaire 4499 (BR); Les Saras region, Congo–Cabindaroad, 8 Jun 1966, Sita 1376 (P); s.loc., Moutsamboté1401 (BR).

9. CUVIERA TRILOCULARIS HIERN, in D. Oliver, Fl.Trop. Afr. 3: 157. 1877. Type: NIGERIA. OldCalabar, 11 Apr 1863, Thomson 122 (lectotypedesignated here: K000412061!; isolectotypes:E00193711!, E00259212!).

Distribution: Lower Guinea Domain: restricted toextreme south-east Nigeria (Oban) and adjacentsouth-west Cameroon (Korup National Park).

Notes: Hiern (1877) based his specific name on thesupposedly trilocular ovary, which seems to have beena wrong observation. We dissected two ovaries (oneeach from D.W. Thomas 6797 and van Meer 1357) andfound them to be pentalocular, as is usual in thegenus.



Cuviera trilocularis is the only species of Cuviera inwhich ant holes seem to be consistently absent; it isalso unusual in its few-flowered inflorescences. Theflower colour pattern of C. trilocularis needs checkingin the field: the label of van Meer 1357 describes thecorolla as ‘greenish white inside’, not mentioningstripes but on the dried specimen the central zone ofthe 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. Thomas1181 (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 (south-west 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, vanMeer 1261 (WAG); Oban Group Forest Reserve (westblock, cut line from Pillar 51), 19 Apr 1971, van Meer1357 (BR, WAG).

10. Cuviera truncata Hutch. & Dalziel, Fl. W. Trop.Afr. 2: 118. 1931.

Type: NIGERIA. Oban, 1911, Talbot 286 (lectotypedesignated here: K000412060!; isolectotype:BM000903509!); Degema, 1916, Talbot 3686 (syn-types: BM000903507!, BM000903508!).

Distribution: Lower Guinea Domain: south-westCameroon, Nigeria.

Notes: Cuviera runcata is unique in the genus inhaving the calyx truncate or with short (< 2 mm) andirregular lobes. It further differs from C. acutiflora,its closest relative, in having larger and broaderbracts. However, some Nigerian collections (Dalziel 45and Elliott 198, both from Lokoja; Kennedy 2758 fromSapoba; Thornewill 47 from Wamba river) are inter-mediate between the two species, either by combiningcharacters of both (large bracts and long calyx lobes,or the reverse) or by having variable calyx lobes. Theycould represent hybrids; fieldwork in Nigeria wouldbe needed to solve this question. Another Talbot 222specimen belongs to C. subuliflora (see Note underthat species).

Additional specimens examined: CAMEROON. Kupe–Muanenguba, Kodmin, 20 Jan 1998, Etuge et al. 4051(YA); Eyumojock Forest Reserve (c. 42 km east ofMamfe), 9 Feb 1985, Onana 107 (BR, YA); Mamfetown, 22 Apr 1987, D.W. Thomas 7304 (BR, K, MO,WAG).

NIGERIA. s.loc., Akete 64 (K); Ganglani, 23 Nov1957, Hepper 1448 (BR, K, P); Sanga river, DogonKurmi, 30 Nov 1957, Keay 37246 (K, P); DogonKurmi, 17 Nov 1946, Keay & Onochie 21535 (K);Gangumi (between Dondere and Yandere), 8 Dec1954, Latilo & Daramola 28861 (K, P); Nimbia ForestReserve, 9 Apr 1963, Latilo 47417 (K, P, WAG); Boki(c. 18 miles from Bende/Ayuk), 28 Feb 1973, Latilo67754 (K, WAG); Dogon Kurmi, 10 Dec 1964, Olorun-femi 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 (lecto-type designated by Verdcourt, 1987: 189).

Description: Small trees or shrubs, 1.5–16.0 m high,evergreen, with regular horizontal branches (Roux’sarchitectural model); vegetative parts glabrous orrarely shortly pubescent (in some specimens ofG. rammelooana and G. robbrechtiana); twigs withoutant holes; spines absent. Leaves opposite, ± elliptic;stipules small, basally ± connate, with an awn> 2 mm long (frequently damaged). Inflorescencesaxillary and generally paired at nodes (or occasionallyterminal), on young twigs, cymose, umbelliform, one-to five-flowered, with a single pair of elliptic leafybracts at apex of peduncle (in G. cuvieroides andG. minor rarely with some additional bracts on thepedicels). Flowers hermaphrodite. Calyx lobes five,almost free, leafy, broadly elliptic or more rarely lan-ceolate (G. cuvieroides). Corolla green or yellow; tubebroadly funnel-shaped; lobes five, triangular, acute toshortly apiculate, spreading or reflexed; outside gla-brous or with five rows of hairs on the lobes andupper part of tube; inside with a ring of hairs in thelower half of the tube. Stamens five, inserted in theupper part of the tube, anthers partly to fullyexserted. Ovary pentalocular, obconic or turbinate,smooth; ovules solitary in each locule, pendulous.Disk ± flat, glabrous. Style shortly exserted, with apubescent swelling in the lower part (fusiform inG. dewildeana and G. robbrechtiana, discoid toglobose in other species), otherwise glabrous; stig-matic club ± mitriform, shortly five-lobed. Fruitdrupaceous, maturing yellow to orange (but generallyseen green), large, 1.5–3.5 × 1.2–3.5 cm, subglobose toellipsoid, smooth to conspicuously five-lobed, withcalyx persistent (rarely caducous); pyrenes five, one-seeded, shaped like a mandarin orange segment, witha slight ventral indentation towards the upper third.

Distribution and ecology: Globulostylis has eightspecies in Central Africa, all endemic to the Lower



Guinea Domain, except G. uncinula, which alsooccurs in the Congolian Domain. The centre of diver-sity is in Cameroon (seven species, including threeendemics). All Globulostylis spp. are shrubs or smalltrees of forest understory, never gregarious.

Notes: Globulostylis spp. are often difficult to separatein the absence of flowers. The leaves in particularshow little variation between the species.

The inflorescences are most often axillary, butterminal inflorescences, previously unrecorded inVanguerieae, are occasionally found in G. robbrech-tiana (e.g. Nguema & Parmentier 446, Ngomo &Ndong 439) and G. rammelooana (e.g. Sonké & Simo4671).

As for Cuviera, the pollinators or dispersors ofGlobulostylis are unknown. There are well-markeddifferences in flower characters between species witha long corolla tube, a generally weak style swelling,and an internal ring of hairs close to the base(G. dewildeana, G. leniochlamys and G. robbrech-tiana) and those with a short corolla tube, a thickstyle swelling and an internal ring of hairs near themiddle of the tube (the remaining five species), so it islikely these two groups have different pollinators. Thefunction of the style swelling is unclear, although a

role in preventing nectar robbery has been suggested(Verdcourt, 1987). No human uses of Globulostylishave 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, Mildbraed5610 (lectotype designated here: HBG!).

Distribution: Lower Guinea Domain: south-east Cam-eroon, east Gabon.

Notes: Globulostylis cuvieroides stands well apartfrom its congeners in having narrowly lanceolatecalyx lobes, a character more reminiscent of Cuviera.For that reason it shows a superficial resemblance toCuviera pierrei (see that species).

An examination of the type of C. heisteriifoliashows this to be conspecific with G. cuvieroides. Theoriginal specimen has presumably been destroyed inBerlin, so the HBG duplicate is selected as lectotype.

KEY TO THE GLOBULOSTYLIS SPECIES

1. 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

1’. Calyx lobes broadly elliptic, < 3 times longer than wide; smaller trees or shrubs, 1.5–6 m high ........22. Calyx ciliate on margin; corolla lobes pubescent outside on the midrib ...........................................32’. Calyx entirely glabrous; corolla lobes glabrous outside ...................................................................43. Flowers small; calyx 5–7 mm long; corolla tube 4–5 mm, and shorter than the lobes (Nigeria and

south-west Cameroon) .....................................................................................................G. talbotii3’. 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. dewildeana4. 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

4’. Corolla green (possibly yellow in C. leniochlamys), with tube 2–15 mm long, not or shortly exceedingcalyx; style with a strong, globose to discoid swelling; fruits smooth to weakly five-lobed and generallylarger, 19–32 × 17–27 mm when dry .............................................................................................5

5. Calyx large, with lobes 11–22 mm long .........................................................................................65’. Calyx smaller, with lobes 4–10 mm long ..................................................................................... ..76. Corolla tube c. 15 mm long, ± equalling calyx; inflorescence three- to five-flowered (but only one fruit

may develop) (south Cameroon) ...............................................................................G. leniochlamys6’. Corolla tube much shorter than calyx; inflorescence always one-flowered (Gabon to Democratic Republic

of the Congo) ............................................................................................................. ..G. uncinula7. 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. minor7’. 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



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).



Additional specimens examined: CAMEROON. Lomiéregion, 1911, Mildbraed 5404 (HBG); c. 15 km south-east 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 east-north-east Djoum), 9 Nov 1966, Letouzey 8325 (BR, P,YA); Bouamir, 5 Sep 2001, Nguembou 146 (BR); Akomsili-Bali (Dja Fauna Reserve), 28 Dec 2002, Nguem-bou 544 (BR, BRLU); track from Bouamir stationMbassakok (Dja Fauna Reserve), 20 Apr 2001, Sen-terre & Kouob 1007 (BRLU); ibid., Senterre & Kouob1020 (BRLU); ibid., 17 May 2001, Senterre & Kouob1233 (BRLU); ibid., 20 May 2001, Senterre & Kouob1408 (BRLU); Djolimpoum (Dja Fauna Reserve), 10Dec 1994, Sonké 1296 (BR); Mékas (Dja FaunaReserve), 12 Jan 1995, Sonké 1405 (BR, WAG);Meka’a (Dja Fauna Reserve), 15 Feb 1995, Sonké1438 (BR); Djolimpoum (Dja Fauna Reserve), 13 Apr1995, Sonké 1464 (BR); 27 km south of Djolimpoum,26 Nov 1996, Sonké 1841 (BR, K); Haut-Nyong, Soma-lomo, 12 km east of Somalomo, 8 Nov 2002, Sonkéet al. 2882 (BR, K, YA).

GABON. c. 60 km south-south-west of Moanda, 16Oct 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, 9Jun 1978, Florence 1220 (P); Levata, 29 Apr 1930, LeTestu 8059 (BM, P); Oveng, km 19 forest exploitationcamp, 9 Nov 1983, A.M. Louis et al. 525 (BR, K, P,WAG); eastern border of Lopé–Okanda Reserve, 15Jan 1993, McPherson 16062 (BR, K, MO, WAG);Oveng, c. 15 km north-west of forestry camp, 8 Feb1987, 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 kmnorth-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 May1986, Wilks 1301 (WAG).

2. GLOBULOSTYLIS DEWILDEANA SONKÉ,O.LACHENAUD & DESSEIN sp. nov. Type: CAM-EROON. between N’Kolandom and N’Koemvone,680 m alt, 2°48′N, 11°09′E, 9 Jan 1975, J.J.F.E deWilde 7886 (holotype: WAG0114671!; isotypes: BR!,K!, P!, WAG0114672!, WAG0114673!).

Diagnosis: Globulostylis dewildeana resembles G. tal-botii in its ciliate calyx and dorsally pubescent corollalobes, but differs in the larger calyx lobes (10–16 mmlong vs. 5–7 mm long), the much longer corolla tube(14.5–19.0 mm long vs. 4–5 mm long), and the stylewith a weak and fusiform basal swelling (vs. thickand subglobose in G. talbotii). Globulostylis dewil-

deana also resembles G. leniochlamys and G. robbre-chtiana in corolla shape (tube longer than broad andmuch exceeding lobes), but in these species the calyxand outside of corolla are glabrous.

Description: Shrub, 2.5–4.5 m tall; twigs glabrous,older ones with pale grey bark. Leaves with petiole7–13 mm long, glabrous; lamina elliptic to oblong–elliptic, (9.0–)17.0–23.5 × (4.3–)7.5–12.5 cm; apexacuminate with acumen 6–13 mm long; base shortlyattenuate, acute or obtuse; leaf surface glabrous onboth sides; secondary veins brochidodromous, six toeight on each side of the midrib, ascending, straightto slightly curved at the base, strongly curved somedistance from the margin to join with the adjacentvein; domatia absent. Stipules with a short triangularbase prolonged into an awn 4–9 mm. Inflorescencesaxillary paired at node, two- to four-flowered; pedun-cle short, < 6 mm; bracts triangular, c. 5 mm long,glabrous. Flowers 5-merous, subsessile; flower buds16–24 mm long, apiculate. Calyx pale green; calyxtube c. 1 mm; calyx lobes broadly elliptic to ovate,10–16 × 6–9 mm, ciliate. Corolla pale yellow to palebrownish–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 themedian keel of the lobes, prolonged on the tube for theupper third; inside with a ring of hairs near the baseof the tube. Stamens five, attached near the mouth ofthe tube; filaments short, c. 0.5 mm; anthersc. 2.5 mm, half-exserted. Ovary pentalocular, gla-brous. Disc cylindrical, c. 0.3 mm high, glabrous.Style, c. 23 mm long, with a pubescent fusiform swell-ing at the base, otherwise glabrous; stigmatic clublobed, 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 persis-tent calyx lobes; pyrenes unknown.

Distribution: Lower Guinea Domain: endemic to anarrow range south-east of Ebolowa, Cameroon.

Habitat and ecology: Evergreen forest; 650–750 m inaltitude.

Phenology: Flowers in December and January; youngfruits in December, January and March; maturefruits in April.

Etymology: The specific epithet honours Dr J. J. F. E.de Wilde of Herbarium Vadense (Wageningen, theNetherlands), a specialist on Begonia and Meliaceae,who collected most of the specimens including thetype.



Conservation status: IUCN Red List Category Criti-cally Endangered [CR B1ab (i, ii, iii)]. The species isfound far away from protected areas. The region isinterspersed with several roads along which humanactivities are intense, including housing, smallholderfarming and forest logging. The species is found inonly one location. The extent of occurrence is esti-mated to be < 100 km2. Based on the threats identi-fied, a continuing decline can be inferred in EOO,AOO and area, extent and quality of habitat.

Notes: Globulostylis dewildeana has so far been con-fused with G. talbotii in herbaria, but has much alarger corolla and calyx and a different range; it is notsympatric with any of its congeners, except the dis-tinct G. cuvieroides.

Additional specimens examined: CAMEROON. hillfacing the village of N’Kolandom, 9 Apr 1975,J.J.F.E. de Wilde 8159 (BR, K, P, WAG, YA); hill abovethe village of N’Kolandom, 12 Dec 1974, J.J.F.E. deWilde 7831A (BR, K, WAG); hill Nkolomeyan on thetrack Bihongboulou–Koungoulou, 25 km east-south-east 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 des-ignated here: K000412051!; isolectotype: BM!).

Distribution: Lower Guinea Domain: endemic to theNgowayang massif in southern Cameroon.

Notes: This species has been much mistaken in her-baria, mostly with G. rammelooana and G. robbrech-tiana that both occur in its range. It especiallyresembles the latter in corolla shape, but differs inhaving a larger calyx not or hardly exceeded by thecorolla tube, a subglobose (not fusiform) basal styleswelling, and the fruits larger, 24–28 × 17–26 mmwhen dry, smooth or nearly so. From G. ram-melooana, it differs in the larger corolla and calyx (seeKey).

The flower colour of C. leniochlamys needs checkingin the field; Schumann (1899) describes the corolla asochre yellow (‘ockergelb’), which could refer to afading flower. The original material has presumablybeen destroyed in Berlin, so the K duplicate isselected as lectotype.

Additional specimens examined: CAMEROON. fromMbikiliki to basecamp, c. 1.5 km north-west of village,9 Mar 2007, Dessein & Sonké 1448 (BR); near Mbiki-liki village, from basecamp to top of hill, 10 Mar 2007,

Dessein & Sonké 1464 (BR); Mvilé (3 km west-north-west 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 Jan2005, 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, Zenker4425 (BM, BR, HBG, K).

4. GLOBULOSTYLIS MINOR WERNHAM, Cat. Pl. Oban:50. 1913. Cuviera minor (Wernham) Verdc., KewBull. 42: 189. 1987, nom. illeg. (non C. minorC.H.Wright). Cuviera wernhamii Cheek, Pl. MountCameroon: 106. 1998. Type: NIGERIA. Oban,Talbot 247 (holotype: BM000903505!).

Distribution: Lower Guinea Domain: restricted toextreme south-east Nigeria (Oban) and neighbouringCameroon (Korup National Park, Mount Etinde).

Notes: When treated in Globulostylis, this species iscorrectly named G. minor. However, if placed inCuviera, the replacement name C. wernhamii shouldbe used (Cable & Cheek, 1998), because C. minor(Wernham) Verdc. is an illegitimate later homonym ofC. minor C.H.Wright (which is now a synonym ofC. nigrescens).

Globulostylis minor is closely related to G. ram-melooana, which is locally sympatric (see that speciesfor differences). Records from Gabon (Sosef et al.,2006) are an error for G. cuvieroides; the two species,although quite dissimilar, have been much confusedin 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 Apr2009, Dessein et al. 2876 (BR, YA); west Limbe, 15Nov 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, 28Oct 2008, Parmentier & Mambo 5115 (BRLU); Mun-demba (Korup National Park), 29 Dec 1992, Sonké381 (BR); Liwenyi, 29 Oct 1993, Tchouto 1007 (K);Idenau, 8 Nov 1993, Watts 969 (K); Limbe, 0.5 kmnorth-east of Etome, 23 Jan 1994, Wieringa 1990(WAG).

5. GLOBULOSTYLIS RAMMELOOANA SONKÉ,O.LACHENAUD & DESSEIN sp. nov. Type: CAM-EROON. 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!).



Diagnosis: Globulostylis rammelooana is closelyrelated to G. minor, from which it differs in having alarger 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 forcomparison), and also a larger corolla, with tube6–8 mm long and lobes 6–10 mm long (vs. tube2–5 mm long and lobes 3–5 mm long in G. minor). Inaddition, the leaf underside lacks the typical greyishsheen of G. minor, but instead dries olive–green toolive–brown. Globulostylis rammelooana is alsorelated to G. leniochlamys and G. robbrechtiana,which have much larger flowers. Fruiting specimensdiffer from G. leniochlamys in their smaller calyx, andfrom G. robbrechtiana in the larger and less markedlylobed fruits. The calyx lobes are more rounded andoverlapping than in G. robbrechtiana (compareFigs 3G and 4H) and the corolla is green, not yellow.

Description: Shrub, 2–4 m tall; twigs glabrous. Leaveswith petiole 5–10 mm long, glabrous; lamina ellipticto oblong–elliptic, (6.5–)8.7–20.2 × (2.5–)3.4–7.4 cm;apex acuminate with acumen 7–17 mm long; baseshortly attenuate or acute; leaf surface glabrous onboth sides; secondary veins brochidodromous, five toseven on each side of the midrib, ascending, straightto slightly curved at the base, strongly curved somedistance from the margin to join with the adjacentvein; domatia absent. Stipules deciduous, shortly tri-angular at base, prolonged into an awn of 4–7 mm.Inflorescences axillary or, more rarely, terminal, two-or three-flowered (but usually with only one fruitdeveloping), 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 mmlong, 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 hairsnear 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, gla-brous. Disc cylindrical, c. 0.7 mm high, glabrous.Style 8–10 mm long, with a conspicuous hairy globu-lar swelling c. one fifth from the base, otherwiseglabrous; stigmatic club five-lobed, exserted, lobesc. 3 mm long. Fruits subglobose to slightly ellipsoid,weakly five-lobed when dry, 19–25 mm in diameterwhen 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 Cam-eroon, occurring mostly in the South Region (Bipindito Nyabessan) with a smaller population in theSouth-West Region (Mount Etinde).


Phenology: Flowers in November–January andMarch–April; fruits in February–April and June.

Etymology: Named in honour of Professor Jan Ram-meloo, retired director of the National BotanicalGarden of Belgium, to whom all authors are indebtedfor the assistance and facilities offered by hisinstitute.

Conservation status: IUCN Red List Category Vulner-able [VU B2ab(i, ii, iii, iv, v)]. G. rammelooana isrestricted to Cameroon where it is mostly foundoutside protected areas. Logging, deforestation andurbanization are the major threats for this species.This species is known from nine locations. The area ofoccupancy is less than 500 km2 (72 km2). Based on thethreats identified, a continuing decline of EOO, AOO,area, extent and quality of habitat, number of loca-tions and number of mature individuals can beinferred.

Notes: Globulostylis rammelooana has usually beenconfused with G. leniochlamys in herbaria. Its rangeoverlaps those of G. leniochlamys and G. robbrech-tiana in the South Region of Cameroon, and ofG. minor in the South-West Region. All these speciesmostly differ in flowering characters, but the identi-fication of fruiting specimens is difficult, althoughpossible with experience with the group (see Diagno-sis and Key).

The collections from Mount Etinde lack flowers, butas they fully match G. rammelooana in other charac-ters, they are referred to this species.

Additional specimens examined: CAMEROON. Eastof Mount Elephant, 5 km south of km 18 Kribi–Ebolowa, 6 Mar 1970, Bos 6496 (WAG); near Bid-jouka, track to basecamp, 13 Mar 2007, Dessein &Sonké 1546 (BR); Mount Elephant, south-east ofKribi, 28 Apr 1970, Bos 6871 (K, WAG); near Bid-jouka, 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 May1994, Ndam 1185 (K); ibid., Ndam 1244 (K); cañon ofNtem, 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



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 Mar2008, Sonké & Simo 4711 (BR, K, MO, P, WAG, YA);Nkolembonda, 16 Jul 2007, Taedoumg 102 (BR);South Province, Campo Ma’an area, Massif desMamelles, 20 Apr 2001, Tchouto MMX110 (WAG);Mokoko Forest Reserve, Ekombe–Mofako, 23 May1994, Watts 1164 (K, SCA); Bipindi, 1909, Zenker3927 (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, Tae-doumg & Simo 4868 (holotype: BR!; isotypes: BR!,K!, MO!, P!, WAG!, YA!)

Diagnosis: Globulostylis robbrechtiana differs from itscongeners in the long, 20- to 30-mm corolla tube,which is at least twice as long as the calyx (vs. corollatube 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 mmwhen dry). It further differs from all species (exceptpossibly G. leniochlamys) in having bright yellowflowers.

Description: Shrub 2–4(−6) m tall; twigs brown, gla-brous or sometimes sparsely hairy when young.Leaves with petiole 5–10 mm long, glabrous; laminaelliptic to oblong–elliptic, 7.3–18.8(−20.2) × 3.0–7.8 cm; apex acuminate with acumen 7–10(−17) mmlong; base shortly acute to rounded; leaf surface gla-brous, 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, stronglycurved some distance from the margin to join with theadjacent vein; domatia absent. Stipules shortly trian-gular at base, prolonged into an awn of 3–4 mm long,glabrous outside, hairy inside. Inflorescences axillaryor occasionally terminal, two–four (–six)-flowered (butusually with only one fruit developing); peduncle4–7 mm long; bracts ovate–triangular, 5–10 × 2–4 mm, glabrous. Flowers pentamerous; pedicels2–4 mm long; flower buds 22–32 mm long, acute.Calyx pale green; calyx tube 1–2(−4) mm; calyx lobesovate 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 thebase of the tube. Stamens five, inserted near the apexof the tube; filaments 1 mm; anthers c. 2.5 mm, half-exserted to included. Ovary pentalocular, glabrous.

Disc cylindrical, c. 0.5 mm high, glabrous. Style,20–32 mm long, with a pubescent fusiform swellingc. 1.5 mm above the base; stigmatic club five-lobed,just exserted, c. 2.5 mm long. Fruits subglobose, con-spicuously five-lobed, 15–22 × 12–19 mm when dry,glabrous, crowned by persistent calyx lobes, onpedicel 4–8 mm long; pyrenes five per fruit,c. 17 × 5 mm.

Distribution: Lower Guinea Domain: restricted tohilly ranges of southern Cameroon (Ngowayangmassif, Akom II area), Equatorial Guinea (MonteAlén) and northern Gabon (Crystal Mountains).Locally abundant in its range.


Phenology: Flowers mostly from May to October, withone record in March; fruits from July to December.

Etymology: Named in honour of Professor Elmar Rob-brecht, retired head of the Vascular Plants Depart-ment at the National Botanic Garden of Belgium, whomade a major contribution to the knowledge ofAfrican Rubiaceae and to whom all authors areindebted for instruction in aspects of this family.

Conservation status: IUCN Red List Category NearThreatened (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. robbrechtianais here considered to be only Near Threatened. Mostcollections have been made inside protected areas;three collection sites are outside protected areas, butthe surrounding forest seems to be rather untouchedby human activities at present. This might change inthe future, the reason why we think the category NTis the most appropriate.

Notes: The large yellow flowers of G. robbrechtianaare distinctive, but specimens in bud or fruit can bemistaken for G. leniochlamys or G. rammelooana (seethese species for differences). The ranges of the threespecies overlap in Cameroon.

Globulostylis robbrechtiana is normally glabrous,but some duplicates of Sonké, Taedoumg & Simo 4872have short patent hairs on young twigs and theunderside of midrib, which is exceptional in thegenus.

Additional specimens examined: CAMEROON. AkomII, 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,



south Efoulan, 6 Mar 2004, Sonké & Beina 3331 (BR,BRLU, K, MO, WAG, YA); Akom II, Efoulan, 24 Mar2008, Sonké et al. 4872 (BR, K, MO, P, WAG, YA);Campo Ma’an area, Efoulan, 4 Dec 2000, Tchouto3092 (WAG); Bipindi, Oct 1918, Zenker 13 (P).

EQUATORIAL GUINEA. Rio Muni: Monte Alen, 20May 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 Aug2001, 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 (MonteAlen 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., 21Oct 1993, Lejoly 93/422 (BRLU); Monte AlenNational Park, 7 Oct 1994, Lejoly 94/168 (BRLU);ibid., 7 Oct 1994, Lejoly 94/185 (BRLU); ibid., 22 Sep1997, Lisowski 1554 (BRLU); ibid., 22 Sep 1997,Lisowski 1571A (BRLU); ibid., 28 May 1997, Ngomo0173 (BRLU); ibid., 16 Jun 1998, Ngomo & Ndong348 (BRLU); ibid., 10 Aug 1998, Ngomo & Ndong 439(BRLU); 5 km west of Engong (Monte Alen NationalPark), 3 Jul 1999, N. Nguema & Parmentier 446 (BR,BRLU); s.loc., 1 Dec 1997, Obama 347 (BRLU); S duRio 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 ofNkumékié (south-east Monte Alen National Park), 8Dec 2002, Senterre & Obiang 3679 (BRLU); 8.5 kmeast de la Cabaña de Mosumo (Monte Alen NationalPark), 4 Jul 2003, Senterre & Obiang 3991 (BRLU);Monte Alen, 10 Aug 2001, Sonké et al. 2501 (BR);Transect Monte Chocolate (Monte Alen NationalPark), 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 desig-nated here: BM000903503!; isolectotypes:BM000903504!, K000412052!, K000412053!).

Distribution: Endemic to Lower Guinea Domain:restricted to extreme south-east Nigeria and adjacentsouth-west Cameroon.

Notes: Globulostylis talbotii is closely related toG. dewildeana, which has been previously mistakenfor it in herbaria, but has much smaller flowers and

a more north-westerly range. From the sympatricG. minor and G. rammelooana, it can be told apart bythe ciliate calyx and the corolla pubescent outside onthe 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 Jan1972, Leeuwenberg 9275 (BR, P, WAG, YA); MbuBakundu, 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. Cuvierauncinula 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 Repub-lic of the Congo.

Notes: G. uncinula differs from other related species(G. leniochlamys, G. minor, G. rammelooana andG. robbrechtiana) in having consistently one-floweredinflorescences and in combining a short corolla tubewith a large calyx. Fruiting specimens are difficult toseparate from G. leniochlamys, but the two specieshave different ranges.

Globulostylis uncinula, until now regarded as aGabonese endemic, is newly reported from the Demo-cratic Republic of the Congo and the Republic ofCongo.

Additional specimens examined: DEMOCRATICREPUBLIC OF THE CONGO. Imbow, 11 Feb 1959,Evrard 5689 (BR); Mompono, 25 Feb 1959, Evrard5834 (BR, K, WAG).

GABON. near Yombi (on Fougamou–Mouila road,Koumounabouali ridge), 19 Sep 1990, Breteler 13993(WAG); Mount Songou (between Poungui andDibandi), 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. Hallé724 (P); Abanga chantier CEFA, 11 Jun 1963, N.Hallé 2447 (P); Iméno Ivinzi, 23 Mar 1927, Le Testu6446 (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 ofMitzic (forestry concession Bordamur), 6 Feb 2003,Sosef et al. 1891 (WAG); c. 50 km south of Mandji



(Doudou Moutains), 17 Nov 2005, Sosef et al. 2367(WAG); Shell oil-exploitation Rabi, 1 km south-east ofShell camp., 8 Mar 1994, Wieringa 2416 (WAG); northNdouaniang (‘Sogacel’), 6 Aug 1984, Wilks 987 (WAG).

REPUBLIC OF CONGO. Maamar forest exploita-tion (towards N’Gongo-N’Dindi junction), 5 Feb 1974,Sita 3648 (P, WAG); around junction N’Dindi-N’Gongoand 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 ofVangueria lasiantha (Sond.) Sond. (Bridson, 1998).

2. CUVIERA BOLO Aubrév. & Pellegr. – now a synonymof Robynsia glabrata Hutch. (Hepper & Keay,1963).

3. CUVIERA CALYCOSA WERNHAM, J. Bot. 52: 7. 1914.Type: NIGERIA. Eket District, Talbot 3300 (lecto-type designated here: BM000903513!; isolectotypesdesignated 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 nototherwise placed (see Discussion). It differs fromC. nigrescens only (see below) in the size of calyxlobes, so an intraspecific rank might be more appro-priate. Mengé 5 (BR) is the first record from theDemocratic Republic of Congo.

4. CUVIERA MIGEODII VERDC., Fl. Trop. E. Afr.,Rubiac. (3): 771. 1991. Type: TANZANIA. Tenda-guru, Lindi District, 18 Dec 1930, Migeod 1052(holotype: BM000903510!).

Distribution: South Tanzania (Lindi District).

Notes: This species is excluded from Cuviera, but nototherwise placed (see Discussion).

5. CUVIERA NIGRESCENS (SCOTT-ELLIOTT EX OLIV.)WERNHAM, J. Bot. 49: 321. 1911. Vangueria nigre-scens 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.

Kew. 1906: 105. 1906. Type: GHANA. Kwahu, 2 Apr1900, Johnson 646 (lectotype designated here:K000412054!).

Distribution: Upper and Lower Guinea Domains:Cameroon, Central African Republic, DemocraticRepublic of the Congo, Gabon, Ghana, Guinea,Guinea-Bissau, Ivory Coast, Liberia, Nigeria andSierra Leone.

Notes: This species is excluded from Cuviera, but nototherwise placed (see Discussion). The originaldescription is based on two syntypes, one of which isselected as lectotype.

6. CUVIERA SCHLIEBENII VERDC., Kew Bull. 33: 497.1979. Type: TANZANIA. Lutamba See, Lindi Dis-trict, 1 Nov 1934, Schlieben 5576 (holotype:B100160713!; isotypes: BR0000008846815!,EA000002930!, LISC002649!, PRE0593394-0!,US00664144!).

Distribution: Mozambique and south Tanzania.


7. CUVIERA SEMSEII VERDC., Kew Bull. 12: 449. 1957.Type: TANZANIA. Mchinjiri, Rondo Plateau, LindiDistrict, 823 m, Jan 1952, Semsei 620 (holotype:EA000002929!; isotypes: BR0000008846808!,K000412039!).

Distribution: Malawi, Mozambique and southTanzania.

Notes: This species is excluded from Cuviera, but nototherwise placed (see Discussion). It is close toC. nigrescens and we are not convinced they are spe-cifically distinct.

8. CUVIERA TOMENTOSA VERDC., Kew Bull. 36: 557.1981. Type: TANZANIA. Kilwa District, SelousGame Reserve, approximately 15 km south-west ofKingupira, 8°35′S 38°27′E, 175 m, 21 Dec 1975,Vollesen 3122 (holotype: K; isotypes: C, EA, WAG).

Distribution: North Mozambique (Niassa Province) tosouth Tanzania (Lindi Region).


ACKNOWLEDGEMENTS

This research was supported financially by theResearch Foundation – Flanders (FWO, G.0343.09N),



the King Leopold-III Fund for Nature Explorationand Conservation, the Sud Expert Plantes project (no.375), the F.F.R.S.A. (Fondation pour Favoriser lesRecherches Scientifiques en Afrique), the AlbertaMennega Stichting and SYNTHESYS. B.V. holds aPhD research grant from the Agency for Innovationby Science and Technology (IWT, no. 91158). O.L. is aresearch fellow of the F.R.S.-F.N.R.S. (National Fundfor Scientific Research). B.S.’s visits to Belgium werepartly funded by the Université Libre de Bruxellesand the F.R.S.-FNRS. We thank the curators of thefollowing herbaria for their help while working intheir institutions and/or the loan of material: BM,BNRH, BR, BRLU, HBG, K, MPU, P, LBV, WAG andYA. Dr Petra De Block is acknowledged for her helpin the description of Globulostylis robbrechtiana.Murielle Simo, Quentin Luke and Olivier Maurinkindly allowed us to reproduce their photographs.B.S.’s visits to Belgium and France in 2012, duringwhich time this paper was finalized, were funded bythe ‘Institut de Recherche pour le Développement(IRD)’, under the ISMOBIAC project. Thanks are dueto Dr Pierre Couteron (Institut de Recherche pour leDéveloppement IRD – AMAP), Dr Olivier Hardy(Laboratoire d’Eco-éthologie évolutive, UniversitéLibre de Bruxelles) and Dr Tariq Stévart (Herbariumet Bibliothèque de Botanique africaine, UniversitéLibre de Bruxelles) for their assistance. We are deeplygrateful to Dr Vincent Droissart, Yves Issembé, DrSteven Janssens, Thomas Nzabi, Murielle Simo,Hermann Taedoumg and Lise Zemagho for help withthe collection of plant material.

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APPENDIX 1ACCESSIONS INCLUDED IN THE PHYLOGENETIC ANALYSIS

Taxon* Voucher trnTF rpl16 petD ITSrpl32-trnL

accD-psaI

Afrocanthiumgilfillanii (N.E.Br.)Lantz

Lemaire &Verstraete 32(BR)

JQ958219 JQ958112 JQ958036 – – –

Afrocanthiumlactescens (Hiern)Lantz

Kuchar 23003(BR)

JQ958221 JQ958113 JQ958037 – – –

Afrocanthiummundianum(Cham. &Schltdl.) Lantz


JQ958220 JQ958114 JQ958038 – – –

Bullockiapseudosetiflora(Bridson)Razafim., Lantz &B.Bremer

Bidgood et al.4959 (BR)

JQ958222 JQ958115 JQ958039 JQ957968 – –

Bullockia setiflora(Hiern) Razafim.,Lantz & B.Bremer


JQ958223 JQ958116 JQ958040 JQ957969 – –

Canthium ciliatum(D.Dietr.) Kuntze


JQ958230 JQ958122 JQ958049 – – –



APPENDIX 1 Continued


accD-psaI

Canthiumcoromandelicum(Burm.f.) Alston

Andreasen 36(UPS)

AJ620122 AJ876810 – AJ315081 – –

Canthium glaucumHiern

Kuchar 17410(BR)

JQ958231 JQ958123 JQ958046 JQ957970 – –

Canthium inerme(L.f.) Kuntze


JQ958232 JQ958124 JQ958047 JQ957971 – –

CanthiumkuntzeanumBridson


JQ958233 JQ958125 JQ958050 – – –

Canthium spinosum(Klotzsch ex Eckl.& Zeyh.) Kuntze


JQ958235 JQ958126 JQ958048 JQ957972 – –

Canthiumsuberosum Codd

Burrows &Burrows 9858(BNRH)

JQ958234 JQ958127 – – – –

Cuviera acutifloraDC.

Dessein et al.2809 (BR)

JQ958270 JQ958157 JQ958076 JQ957973 JQ958201 JQ957948

Cuviera angolensissubsp. latior(Wernham) Sonké,O.Lachenaud &Dessein

Amsini 97 (BR) JQ958271 – JQ958077 JQ957974 JQ958197 JQ957951

Cuviera longifloraHiern


JQ958272 JQ958159 JQ958079 JQ957975 JQ958199 JQ957953

Cuviera physinodesK.Schum.

Chatrou 572 (BR) JQ958273 JQ958158 JQ958078 JQ957976 JQ958198 JQ957949

Cuviera physinodesK.Schum.


JQ958274 JQ958160 JQ958080 JQ957977 JQ958200 JQ957950

Cuviera schliebeniiVerdc.


JQ958276 – JQ958082 JQ957978 – JQ957962

Cuviera semseiiVerdc.


JQ958280 – JQ958086 JQ957979 JQ958205 JQ957965

Cuviera subulifloraBenth.


JQ958275 JQ958161 JQ958081 JQ957980 JQ958202 JQ957952

Fadogia ancylanthaSchweinf.


JQ958236 JQ958132 – – – –

Fadogia cienkowskiiSchweinf.

Dessein et al. 258(BR)

JQ958244 JQ958128 JQ958052 – – –

Fadogia fuchsioidesSchweinf. ex Oliv.


JQ958239 – JQ958053 – – –

Fadogia homblei DeWild.


JQ958240 JQ958129 JQ958054 JQ957981 JQ958179 JQ957933

Fadogia salictariaS.Moore

Malaisse 13490(BR)

JQ958238 JQ958133 – – – –

Fadogia stenophyllasubsp. odorata (K.Krause) Verdc.

Lovett 2267 (BR) JQ958241 JQ958130 JQ958055 JQ957982 – –





accD-psaI

Fadogia tetraquetraK.Schum. &K.Krause


JQ958242 JQ958131 JQ958056 – – –

Fadogia triphyllaBaker


JQ958237 JQ958134 – – – –

Fadogia verdickiiDe Wild. &T.Durand

Schaijes 1502(BR)

JQ958243 JQ958135 – JQ957984 – –

Fadogiellastigmatoloba(K.Schum.)Robyns

Gillett 17403(BR)

JQ958245 JQ958136 – JQ957983 – –

Globulostylisleniochlamys(K.Schum.) Sonké,O.Lachenaud &Dessein


JQ958269 JQ958152 JQ958070 JQ957985 JQ958189 JQ957947

Globulostylis minorWernham


JQ958261 JQ958151 JQ958075 JQ957986 JQ958190 JQ957939

GlobulostylisrammelooanaSonké,O.Lachenaud &Dessein


JQ958267 JQ958153 JQ958071 JQ957987 JQ958193 JQ957942


Sonké et al. 4650(BR)

JQ958266 – – – JQ958191 JQ957943



JQ958264 – – JQ957988 – JQ957941



JQ958265 – – – JQ958192 JQ957940

GlobulostylisrobbrechtianaSonké,O.Lachenaud,Dessein & DeBlock


JQ958268 JQ958154 JQ958072 JQ957989 JQ958194 JQ957944

Globulostylisuncinula (N.Hallé)Sonké,O.Lachenaud &Dessein


JQ958262 JQ958155 JQ958073 JQ957990 JQ958195 JQ957946





accD-psaI

Globulostylisuncinula (N.Hallé)Sonké,O.Lachenaud &Dessein


JQ958263 JQ958156 JQ958074 JQ957991 JQ958196 JQ957945

Ixora finlaysonianaWall. ex G.Don

VanCaekenberghe54 (BR)

JQ958210 JQ958105 JQ958028 – – –

Keetia gueinzii(Sond.) Bridson


JQ958211 JQ958106 JQ958029 – – –

Keetia venosa (Oliv.)Bridson


JQ958212 JQ958107 – JQ957992 – –

Multidentia crassa(Hiern) Bridson &Verdc.


JQ958277 JQ958162 JQ958083 – – –

Multidentiadichrophylla(Mildbr.) Bridson


JQ958278 JQ958163 JQ958084 JQ957993 JQ958204 JQ957963

Peponidium sp. De Block et al.2487 (BR)

JQ958224 JQ958117 JQ958041 – – –

Plectroniella armata(K.Schum.)Robyns


JQ958229 JQ958121 JQ958051 JQ957994 – –

Pseudomussaendaflava Verdc.

VanCaekenberghe60 (BR)

JQ958208 JQ958104 JQ958026 – – –

Psydraxfragrantissima(K.Schum.)Bridson


JQ958213 JQ958109 JQ958030 JQ957995 – –

Psydrax kraussioides(Hiern) Bridson


JQ958214 – JQ958031 – – –

Psydrax livida(Hiern) Bridson


JQ958215 JQ958108 JQ958032 JQ957996 – –

Psydrax locuples(K.Schum.)Bridson


JQ958216 JQ958110 JQ958033 JQ957997 – –

Psydrax obovata(Klotzsch ex Eckl.& Zeyh.) Bridson


JQ958217 – JQ958034 JQ957998 – –

Psydrax parviflora(Afzel.) Bridson


JQ958218 JQ958111 JQ958035 JQ957999 – –

Pygmaeothamnuschamaedendrum(Kuntze) Robyns


JQ958228 JQ958120 JQ958045 JQ958000 – –

Pygmaeothamnuszeyheri (Sond.)Robyns


JQ958279 – JQ958085 JQ958001 JQ958203 JQ957964

Pyrostria bibracteata(Baker) Cavaco


JQ958225 JQ958118 JQ958042 JQ958002 – –





accD-psaI

Pyrostria hystrix(Bremek.) Bridson


JQ958226 – JQ958043 JQ958003 – –

Pyrostria serpentinaLantz, Klack. &Razafim.

De Block et al.2423 (BR)

JQ958227 JQ958119 JQ958044 – – –

Razafimandimbisoniahumblotii (Drake)Kainul. &B.Bremer

Tosh et al. 263(BR)

JQ958209 – JQ958027 – – –

Robynsia glabrataHutch.

Hall &Amponsah46545 (K)

AJ620170 – – AJ617774 – –

Rytigyniamembranacea(Hiern) Robyns

Lachenaud et al.739 (BR)

JQ958246 JQ958137 JQ958057 JQ958004 – JQ957934

Rytigynia monantha(K.Schum.)Robyns

Niyongabo 53(BR)

JQ958247 JQ958138 JQ958058 – – JQ957935

Rytigynia neglecta(Hiern) Robyns


JQ958248 JQ958139 JQ958060 JQ958005 – JQ957936

Rytigynia rubraRobyns


JQ958249 JQ958140 JQ958059 JQ958006 JQ958180 JQ957938

Rytigyniaumbellulata(Hiern) Robyns


JQ958250 JQ958141 JQ958061 JQ958007 – JQ957937

Vangueria agrestis(Schweinf. exHiern) Lantz

Lejoly 82/390(BR)

JQ958281 JQ958164 JQ958087 JQ958008 – –

Vangueria bowkeri(Robyns) Lantz


JQ958284 JQ958170 JQ958097 JQ958009 – –

Vangueriacinerascens (Welw.ex Hiern) Lantz


JQ958282 JQ958165 JQ958088 JQ958010 – –

Vangueria discolor(De Wild.) Lantz

Bidgood et al.6094 (BR)

JQ958283 – JQ958089 JQ958011 – –

Vangueria dryadumS.Moore


JQ958285 JQ958166 JQ958090 – – –

Vangueria infaustaBurch.


JQ958289 JQ958171 JQ958093 JQ958012 JQ958206 JQ957966

Vangueria lasiantha(Sond.) Sond.


JQ958287 JQ958168 JQ958091 – – –

Vangueria latifolia(Sond.) Sond.


JQ958296 JQ958169 JQ958101 JQ958013 – –

Vangueriamacrocalyx Sond.


JQ958288 JQ958172 JQ958098 JQ958014 JQ958207 JQ957967





accD-psaI

VangueriamadagascariensisJ.F.Gmel.


JQ958290 – JQ958094 – – –

Vangueriamicropyren(Verdc.) Lantz


JQ958292 JQ958173 – – – –

Vangueriapallidiflora(Bullock) Lantz

Ntemi Sallu et al.309 (BR)

JQ958293 JQ958174 JQ958096 JQ958015 – –

Vangueria parvifoliaSond.


JQ958294 JQ958175 JQ958099 – – –

Vangueria pygmaeaSchltr.


JQ958297 JQ958177 JQ958102 – – –

Vangueria randiisubsp. chartacea(Robyns) Verdc.


JQ958291 – JQ958095 – – –

VangueriasoutpansbergensisN.Hahn


JQ958295 JQ958176 JQ958100 – – –

Vangueria thamnus(Robyns) Lantz


JQ958298 JQ958178 JQ958103 – – –

Vangueria triflora(Robyns) Lantz


JQ958286 JQ958167 JQ958092 – – –

Vangueriellachlorantha(K.Schum.) Verdc.


JQ958251 JQ958146 JQ958062 JQ958016 JQ958181 JQ957954

Vangueriellachlorantha(K.Schum.) Verdc.


JQ958252 JQ958147 JQ958063 JQ958017 JQ958182 JQ957955

Vangueriella discolor(Benth.) Verdc.

Jongkind 7887(WAG)

JQ958256 JQ958142 JQ958067 JQ958018 JQ958185 JQ957958

Vangueriellalaxiflora(K.Schum.) Verdc.


JQ958254 JQ958149 JQ958064 JQ958019 JQ958183 –

Vangueriellanigerica (Robyns)Verdc.

Jongkind 3089(WAG)

JQ958258 JQ958144 JQ958068 JQ958020 JQ958186 JQ957959

Vangueriellanigricans (Robyns)Verdc.


JQ958253 JQ958148 JQ958066 JQ958021 – JQ957956

Vangueriellaolacifolia (Robyns)Verdc.


JQ958255 JQ958150 JQ958065 JQ958022 JQ958184 JQ957957

Vangueriella rufa(Robyns) Verdc.


JQ958257 JQ958143 – JQ958023 JQ958188 JQ957960

Vangueriella spinosa(Schumach. &Thonn.) Verdc.

Merello et al.1494 (K)

AJ620182 – – AJ315085 – –





accD-psaI

Vangueriellavanguerioides(Hiern) Verdc.

Jongkind 7855(WAG)

JQ958259 – – JQ958024 – –

Vangueriellavanguerioides(Hiern) Verdc.

Kpadeyeah 10(WAG)

JQ958260 JQ958145 JQ958069 JQ958025 JQ958187 JQ957961

VangueriopsisshimbaensisA.P.Davis &Q.Luke

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 2DNA 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 trnTFa1trnTLi

55 °C GTR + G Lantz & Bremer, 2004

trnL-trnF trnLFctrnLFf

55 °C GTR + G Lantz & Bremer, 2004

rpl16 L16exon1 (pcr)RPL16-18R (seq)1067F

55 °C GTR + I + G Lantz & Bremer, 2005

petD petB1411petD738

55 °C GTR + G Löhne & Borsch, 2005

Internal transcribedspacer (ITS)

ITS1ITS4

50 °C HKY + G White et al., 1990

rpl32-trnL rpl32F 55 °C F81 + I Shaw et al., 2007accD-psaI accD769F

psa175R55 °C GTR + I Tosh et al., 2009



Taxonomy and phylogenetics of Cuviera (Rubiaceae-Vanguerieae) and reinstatement of Globulostylis...

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