Holotypus: Cameroon. Reg. South: 13 km E Sangmelima, 24.V.2016, fl., Sonké & Zemagho 6341 (BR!; iso-: BRLU!, K!, MO!, WAG!, YA!).

Habitu sarmentoso, ovario biloculare, foliis valde discoloribus subtus indumento denso lanato et persistente munitis, corolla fauce pilis brevibus inflatis munita S. nobile similis, a quae differt ramis et foliis subtus ad nervis indumento mixto pilis lanatis et rectis (nec pilis omnibus lanatis) munitis, inflorescentiis ad anthesim breviter ramosis (nec capitatis), fructibus rubro-purpureis deinde nigrescentibus (nec albis), seminibus grosse reticulatis (nec striolatis).

Sarmentose liana, c. 3 m high; stems cylindrical, 3.5–6 mm thick, covered with a dense buff indumentum of straight ± appressed hairs c. 1.5 mm long, intermingled with shorter crispate hairs. Stipules interpetiolar, pale green, 10–15 × 4–12 mm, erect, ovate and longitudinally plicate (except on buds), truncate at base, gradually acuminate at apex, uniformly pubescent outside with same indumentum as the stems, more sparsely so inside. Leaves opposite, slightly unequal; petiole (1–)1.8–5 cm long, with same indumentum as the stems; lamina (9–)10.7–19.3 × (3.8–)5.8–10.3 cm, elliptic to ovate, obtuse to rounded at base, gradually acuminate at apex, slightly coriaceous, discolorous; upper side dark green, densely pubescent on the veins with same indumentum as the stems, very sparsely so between them; margin densely ciliate with appressed hairs; lower side pale buff, with dense woolly hairs covering most of the surface (sparser on old leaves), and long straight hairs on the veins; secondary veins 13–16 pairs, weakly to moderately ascending, eucamptodromous; tertiary veins raised on the lower leaf surface and distinctly more prominent than quaternary veins, the latter densely reticulate but ± hidden by indumentum. Inflorescences axillary, paired at nodes or sometimes solitary, densely cymose, many-flowered, 1.7–5 cm long, densely pubescent with same indumentum as the stems; peduncle 0.5–2.5 cm long; primary ramifications 0.3–1 cm long. Bracts green, free, pubescent on both sides, all recaulescent except the basal pair; basal bracts broadly ovate, 7.5–12 × 5–6 mm, entire or dentate, acute at apex; median bracts broadly ovate, 6–14 × 3–7 mm, entire, acute to obtuse at apex; upper bracts narrowly elliptic, 3–6 × 0.8–2 mm, entire, acute at apex. Flowers 5-merous; pedicels 0–1 mm long, densely villose. Calyx pale green; tube c. 1 mm long, densely villose on both sides; lobes elliptic to obovate, 1.5–2.5 × 0.8–1.5 mm, obtuse and curved outwards at apex, villose on both sides, alternating with minute colleters. Corolla white; tube cylindrical or nearly so, 4–5 × 1–1.5 mm, outside mostly glabrous but sparsely appressed-pubescent at apex, inside pubescent from the upper ¼^th to the throat, the latter with short inflated unicellular hairs; lobes triangular, c. 1.3 × 1.3 mm, patent, pubescent with stiff appressed hairs outside, glabrous inside; flower buds obtuse, slightly enlarged at apex. Stamens included, attached around the upper ¼th of the corolla tube; anthers sessile, elliptic, 0.75–1 × 0.25–0.4 mm, glabrous. Ovary 1.5–2 mm long, densely villose, 2-locular. Disk cylindrical, c. 0.7 mm long, glabrous, shorter than the calyx tube. Style 4–4.5 mm long, just reaching the corolla throat, glabrous, bifid, with shortly stipitate elliptic flattened stigmas c. 1 mm long. Fruits purplish-red when young, blackish when mature, ellipsoid, with persistent calyx, 8–11 × 7–9 mm in life, 6–9 × 4.5–8 mm when dry, villose, with pedicel c. 1 mm long; mesocarp very soft, purple, c. 1.5 mm thick, with numerous darker granules. Seeds numerous, dark brown, polyedric, c. 0.6 × 0.4 mm, the surface rather coarsely reticulate.

Etymology. – The species is named after Dr Pierre Couteron (Institut de Recherche pour le Développement, Montpellier, France) in recognition for his constant support to the second and third authors.

Distribution, ecology and phenology. – Sabicea couteronii has an apparently very scattered distribution (Fig. 4), being known from one locality in south Cameroon (near Sangmelima), one in central Gabon (Waka National Park) and two in southwestern Republic of Congo (Mayombe and Chaillu massifs). It occurs in secondary forest, edges and regrowth, c. 685 m in elevation (probably also lower).

Flowers have been collected in May-June, fruits in November and January (immature) and June (mature).

Conservation status. – The extent of occurrence (EOO) of S. couteronii is calculated to be 73,728 km^2, exceeding the limit for “Vulnerable” status under subcriterion B1, while its area of occupancy (AOO) is estimated to be 16 km^2, within the limit for “Endangered” status under subcriterion B2. The species occurs in Atlantic Central Africa (Cameroon, Gabon and Republic of Congo) and is known from six specimens, representing four subpopulations. Two of them occur in protected areas, Waka National Park in Gabon and Dimonika Biosphere Reserve in Congo. The other two subpopulations occur along roads, and the Cameroonian one at least is threatened by habitat destruction due to agricultural activities. A decline in habitat extent and quality and number of individuals is therefore expected. The four subpopulations represent four locations in the sense of ICUN (2012), and the species qualifies for “Endangered” [EN B2ab(iii,v)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – Sabicea couteronii resembles both S. nobilis and S. proselyta, which both occur in its range, and at first sight might be suspected to be a hybrid between them; this is however unlikely, as it differs from both by the colour of its fruits, which are purplish-red when young and blackish when mature (vs. white or greyish-white). It is further separated from S. proselyta by its strongly discolorous leaves with dense woolly indumentum below (Table 1), and from S. nobilis by its (shortly) branched inflorescences, reticulate seed surface, and the presence of long silky hairs (in addition to woolly ones) on the stems, petioles and underside of the leaf veins.

The specimens from Gabon and Congo are either vegetative (Cusset 875) or with immature fruits (Boussengui-Nongo et al. 309, Kimpouni 600) but match very well the Cameroonian material, differing only in the longer peduncles (probably not a very significant character), and, in the case of the Gabonese material, sparser indumentum of the lower leaf surface (probably because the leaves are older). Therefore, we regard them as conspecific. Kimpouni 600 was previously cited as Pseudosabicea proselyta (Lachenaud, 2009) of which it was the only record from the Republic of Congo; however, authentic Congolese specimens of S. proselyta have since been found (see that species).

It is not known if S. couteronii is heterostylous; only the type specimen bears flowers, which are of the long-styled form.

Paratypi. – Cameroon. Reg. South: 13 km E Sangmelima, 28.VI.2015, fr., Sonké 6248 (BR, BRLU, K, MO, WAG, YA); ibid. loco, 16.V.2016, fr., Sonké & Zemagho 6334 (BR, BRLU, K, MO, WAG, YA).

Gabon. Ngounié: Parc national de Waka, 1°10′34″S 11°08′02″E, 28.XI.2007, imm.fr., Boussengui-Nongo et al. 309 (LBV).

Republic of Congo. Kouilou: env. de Dimonika, source Paris Sangha, 5.III.1980, ster., Cusset 875 (P). Niari: Rte Komono–Mossendjo, 2 km du camp SIDETRA, 8.I.1989, imm.fr., Kimpouni 600 (BRLU).

Holotypus: Gabon. Ngounié: km 43 Mouila–Yéno, 21.II.2008, Dessein et al. 2028 (BR [BR0000013945923]!; iso-: LBV!, WAG!).

Habitu sarmentoso, ovario biloculare, seminibus grosse reticulatis, foliis utrinque viridibus subtus ad nervis indumento albido valde appresso munitis, corollaque fauce pilis brevibus inflatis munita S. proselyta similis, a quae differt calyce bicolore apice viride et basi rubro-vinaceo (nec omnino viride), inflorescentiis ad anthesim glomerulatis deinde cymosis (nec semper laxe cymosis), pedunculis nullis vel brevissimis usque 0.3 cm longis (nec 0.7–3.8 cm longis), stipulis basi auriculatis et extus sparsissime pubescentibus (nec basi rotundatis et extus ± dense pubescentibus), foliis subtus venulis tertiariis conspicuis sed haud prominentibus, fructuque albissimo (nec albo-griseo) mesocarpio granulis atris nec munito.

Sarmentose liana, 2–4 m high; stems cylindrical, 2–3 mm thick, densely covered with straight whitish appressed hairs c. 1 mm long, intermingled with shorter uncinate hairs. Stipules interpetiolar, pale green, 9–14 × 6–9 mm, erect, ovate and longitudinally plicate (except on buds), slightly auriculate at base, acute or acuminate at apex, minutely and very sparsely appressed-pubescent outside (except the base with indumentum similar to the stems), sparsely villose inside. Leaves opposite, often slightly unequal; petiole 1–5 cm long, with same indumentum as the stems; lamina 8–16 × 3.5–9 cm, elliptic to ovate, acute to subcordate at base, gradually acuminate at apex, papyraceous, discolorous; upper side medium green, minutely and sparsely appressed-pubescent to glabrous (except for the midrib which often has longer hairs); lower side pale green with ± orange-tinged secondary veins, minutely and sparsely appressed-pubescent, more densely so on the veins; margin densely ciliate with appressed hairs; secondary veins 13–19 pairs, weakly ascending, forming inconspicuous loops near the margin; tertiary veins conspicuous but not or hardly raised on lower leaf surface, slightly more prominent than quaternary veins, the latter densely reticulate forming areolae c. 0.5 mm in diameter. Inflorescences axillary and paired at nodes, many-flowered, glomeruliform and 1.2–2.5 cm in diameter at anthesis, becoming laxer and conspicuously cymose, 2.5–3 cm long in the fruiting stage, densely pubescent with stiff appressed whitish hairs 0.5–0.7 mm long; peduncle 0–0.3 cm long; primary ramifications not or hardly distinct at anthesis, c. 0.5 cm long in the fruiting stage. Bracts green, free, shortly pubescent on both sides, all recaulescent except the basal pair; basal bracts ovate, lobed at base, 7–8.5 × 2.5–4 mm; median bracts elliptic to obovate, 5–6 × 1–2 mm, entire; upper bracts narrowly elliptic to linear, 1.7–4 × 0.25–0.5 mm, entire. Flowers 5-merous; pedicels 0–1 mm long at anthesis, pubescent like the inflorescence. Calyx dark pinkish-red at base and green at apex; tube 0.7–1 mm long, appressed-pubescent outside and more sparsely so inside; lobes obovate to elliptic, 2.5–4 × 1.5–2.3 mm, somewhat unguiculate at base, obtuse to acute and curved outwards at apex, covered on both sides with short patent hairs, and alternating with minute colleters. Corolla tube wine red, cylindrical, 5 × 1.5 mm, outside glabrous at base and sparsely puberulous in the upper half, inside with a ring of multicellular hairs around the upper third and short, inflated unicellular hairs at the throat, the intervening zone glabrous; lobes cream on both sides, triangular, 1.5 × 1.5 mm, reflexed, sparsely puberulous outside, inside with short inflated unicellular hairs at base and glabrous at apex; flower buds obtuse, slightly enlarged at apex. Stamens included, inserted around upper ¼th of the corolla tube with their tips just reaching throat; filaments c. 0.4 mm long; anthers cream, elliptic, c. 1.2 × 0.7 mm, glabrous. Ovary c. 1.5 mm long, densely covered with stiff appressed whitish hairs, 2-locular. Disk cylindrical, c. 0.5 mm long, glabrous, shorter than the calyx tube. Style c. 3.5 mm long, included, bifid, with elliptic flattened stigmas c. 1 mm long, these shortly stipitate. Fruits white, ellipsoid, with persistent calyx, 5–7 × 4–6 mm when dry, sparsely pubescent with appressed hairs, on accrescent pedicel 4–6.5 mm long; mesocarp very soft, entirely white (without darker granules). Seeds numerous, dark brown, polyedric, c. 0.5 × 0.3 mm, the surface prominently and rather coarsely reticulate.

Etymology. – The species epithet refers to the bicoloured calyx lobes, which are the plant's most striking character in the field.

Distribution, ecology and phenology. – Sabicea dichrosepala is apparently endemic to Gabon, where it occurs in the Cristal Mountains in the northwest and the Chaillu massif in the south (Fig. 4), two areas well-known for their endemism and floristic affinity. It might be expected in adjacent parts of Equatorial Guinea and the Republic of Congo. Although probably under-collected, it is clearly an uncommon species. It occurs in forest edges, especially along tracks or forest streams, 610–730 m in elevation.

Flowers have been collected in January-February (buds) and March (open); fruits in September (immature), February and March (mature).

Conservation status. – The extent of occurrence (EOO) of Sabicea dichrosepala is estimated to be 17,016 km^2, and its area of occupancy (AOO) to be 28 km^2, respectively within the limits for “Vulnerable” and “Endangered” status under subcriteria B1 and B2. The species is endemic to Gabon and is known from seven specimens, representing six subpopulations, one of which occurs in a protected area, the Crystal Mountains National Park. The species occurs in sparsely populated areas, and favours forest edge habitats, so is likely to benefit from small-scale forest clearing, e.g. that induced by selective logging. However, two of its known sites are very close to the Tchimbélé dam, the construction of which has presumably affected the extent and quality of habitat and number of individuals. The six subpopulations represent six locations in the sense of IUCN (2012), and the species qualifies as “Vulnerable” [VU B2ab(iii,v)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – The earliest collection of this species, J.J.F.E. de Wilde et al. 284, was cited as Pseudosabicea proselyta by Sosef et al. (2006). Sabicea dichrosepala is indeed closely related to S. proselyta (the two share, in particular, a similar seed ornamentation) but differs by the characters mentioned in Table 1, the most conspicuous of which are the bicoloured calyx, and the inflorescences that are very dense and glomeruliform in the flowering stage.

It is not known whether the species is heterostylous; the only open flowers seen (Fig. 6D) are short-styled.

Paratypi. – Gabon. Estuaire: Mts de Cristal, vers le secteur Mont Séni, Concession BSG, 0°50′52″N 10°24′12″E, 8.III.2019, fl., Bidault et al. 4401 (BRLU, LBV n.v., MO); forest exploitation Leroy, 20 km NW of Asok, 27.I.1983, buds, J.J.F.E. de Wilde et al. 284 (BR, LBV, P, WAG); Mts de Cristal, Tchimbélé-Nkol Madouaka, 17.II.2007, fr., Leal et al. 1396 (BR). Ngounié: Mouyanama falls, at base of Mt Songou, 23.II.2008, buds, Dessein et al. 2111 (LBV); Massif du Chaillu au SE localité Moudouma, 21.IX.2001, imm.fr., Mayombo-Nzengue 928 (LBV). Woleu-Ntem: 2 km N of Tchimbélé on former road to Asok, 6.II.2008, old fl., Dessein et al. 1741 (BR, LBV).

Lectotypus (designated by Hallé, 1966: 216): Cameroon. Reg. South: Lolodorf, 1896, Staudt 1 (P [P00077586]!; isolecto-: COI!, G [G00014620]!, S [S05-10061] image seen). Syntypus: Cameroon. Reg. South: Lolodorf, 1896, imm.fr., Staudt 159 (G [G00014619]!, K [K000414629]!, P [P03793513]!).

Distribution, ecology and phenology. – A common and widespread central African species, occurring from southern Cameroon to eastern D.R. Congo with an isolated population in eastern Nigeria. It grows in secondary forest, edges and regrowth, on both drained and flooded soils, 0–1250 m in altitude. For more precise information see under infraspecific taxa. Flowers and fruits throughout the year.

Conservation status. – The extent of occurrence (EOO) of Sabicea floribunda is calculated to be 2,017,090 km^2, well above the limit for “Vulnerable” status under subcriterion B1. Its area of occupancy (AOO) is estimated to be 344 km^2, which falls within the limit for “Endangered” status under subcriterion B2, but is an obvious underestimation given the low collecting density in most of its range. The species occurs from Nigeria to the Democratic Republic of Congo, and is known from 108 specimens. It is a common species of secondary forest habitats, and not in any way threatened; consequently, it is here assessed as “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – The original description of S. floribunda (Schumann, 1897) is based on two syntypes, Staudt 1 and 159. Hallé (1966: 216) cited the P sheet of Staudt 1 as “type”, which can be accepted as an implicit lectotypification.

This species is the most widespread of the group, and the only one occurring east of the Congo River. It has usually been regarded as monotypic (e.g. Hallé, 1966) but a careful examination of the material shows that it can be quite easily separated into four distinct taxa, which differ mostly in characters of the bracts and the vegetative parts (Table 2) – the flowers and fruits being apparently identical. These taxa have mostly separate distributions but locally come in contact (Fig. 8) and are here recognised as varieties. When they are better studied in the field, especially in their areas of sympatry, some of them (in particular var. anisostegia) may eventually be raised to specific rank. The illustration of S. floribunda published in Hallé (1966: 217) represents var. anisostegia.

Sabicea floribunda is very closely related to S. segregata. The two species share, in particular, lax inflorescences and corolla lobes brown at anthesis, and are not safely separable by vegetative characters. Since they frequently grow side by side, mixed collections are occasionally made (e.g. Champluvier 6019 was initially a mixture of both species). Fertile specimens can be identified with the characters mentioned in the key. The length of calyx lobes was used by Hallé (1966: 200) as the main diagnostic character, but in fact shows some (limited) overlap between the two species.

Distribution, ecology and phenology. – This taxon occurs in Atlantic Central Africa, mostly from southern Cameroon to east-central Gabon with a disjunct occurrence in southeastern Nigeria and adjacent western Cameroon (Fig. 8). Its distribution marginally overlaps with those of var. paucinervis in the west of its range and of var. anisostegia in the east. It occurs in forest edges and regrowth, often along roads, 0–510 m in elevation, and is locally abundant.

Flowers probably all year (recorded from all months except March, June and September); fruits from February to June.

Conservation status. – The extent of occurrence (EOO) of S. floribunda var. floribunda is calculated to be 187,299 km^2, well above the limit for “Vulnerable” status under subcriterion B1. Its area of occupancy (AOO) is estimated to be 84 km^2, which falls within the limit for “Endangered” status under subcriterion B2, but is an obvious underestimation, given the low collecting density in most of its range. The taxon occurs in Central Africa from Nigeria to Gabon, and is known from 26 specimens. It is a locally common plant of secondary forest habitats, and not in any way threatened; consequently it is here assessed as “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – This taxon is morphologically (and also geographically) intermediate between S. floribunda var. paludosa and var. paucinervis, differing from the former in the shape and indumentum of its bracts, and from the latter by its stems and leaves being more hairy, the latter with more numerous secondary veins.

Additional material studied. – Cameroon. Reg. South: au S de Zingui, 20.I.1962, fl., Letouzey 4043 (BR, K, P, WAG, YA); Ebemwok (55 km W Ebolowa), 13.III.1963, imm.fr., J. & A. Raynal 10435 (P, YA); Bebai, Campogebiet, 14.XII.1908, buds, Tessmann 716 (K); Bipindi, 1898, buds, Zenker 1820 (BM, G, K, P, WAG); Bipindi, 1904, fl., Zenker 3232 (BM, BR, K). Reg. Southwest: Abonando, 5.V.1902, buds, Rudatis 52 (K).

Equatorial Guinea. Centro-Sur: Monte Mitra, 28.XI.1997, fl., Obama 315 (BRLU, WAG); SE of Bata c. 50 km, Oborobikó, 25.II.1969, fl. (fallen), Sanford 6034 (K); Ayaguening (Akonangui), 11.VIII.2001, buds, Sonké 2518 (BR, BRLU); 8 km N de Mitong, 13.XII.2003, buds, Sonké 3157 (BR, BRLU, K). Litoral: rte d'exploitation chantier SEMGE vers le 1er pont sur Mitemelé, 13.VI.1988, fr., Dibata 451 (BR, K, WAG); de Sendje à Ongamnsok, 19.II.2001, imm.fr., Lejoly 01/08 (BRLU); rte de Nco vers Misobong au niveau de la grande cascade, 9.I.1998, fl., N. Nguema & Lejoly 206 (BRLU).

Gabon. Estuaire: Forêt de Mondah on road Libreville-Cap Esterias, 26.II.1999, imm.fr., Andersson & Nilsson 2277 (BR); c. 20 km NNE of Libreville, 11.III.1987, fr., J.M. Reitsma & B. Reitsma 3142 (LBV, WAG); Crystal Mountains N.P., 8 km on road from Kinguélé to Tchimbélé, 26.III.2013, imm.fr., Wieringa et al. 7801 (WAG). Moyen-Ogooué: Mboumi. 0°18–0°30S/10°45–10°48E, 16.I.2000, fl., Champluvier 6019 (BR, K, WAG). Ogooué-Lolo: Chantier Bambidie, c. 52 km on the road to Okondja-Lelama, along Sebe river, 31.X.2005, fl., Sosef et al. 2122 (BR, LBV, WAG). Ogooué-Maritime: Région du Lac Alombié, ± 7 km au N de Mpaga, 18.X.2014, buds, Lachenaud et al. 2007 (BRLU, LBV, MO). Woleu-Ntem: SEEG road near Tchimbélé, pylone 7, 6.II.2008, buds, Dessein et al. 1724 (BR, LBV, WAG); Angoum, 28.VII.1933, fl., Le Testu 9198 (BM, P); Monts de Crystal, Tchimbélé, 14.IV.2006, imm.fr., Leal et al. 1109 (BR, LBV).

Nigeria. Cross River State: Oban, 1911, buds, Talbot 228 (BM, K); New Ndebiji, 30.IV.1952, fl., Ejiofor FHI 21883 (FHI, K).

Holotypus: Cameroon. Reg. East: près Yenga Port Gentil, village situé à 35 km au NNE de Moloundou, 21.IV.1971, fl., Letouzey 10716 (BR!; iso-: K!, P!, YA!).

Ab alteribus varietatibus differt bracteis valde inaequales majoribus in inflorescentia dispersis (nec bracteis superioribus gradualiter minoribus quam inferioribus).

Etymology. – The varietal epithet anisostegia refers to the bracts of very unequal sizes, which are the main diagnostic character of this taxon.

Distribution, ecology and phenology. – This variety occurs from southeastern Cameroon to southwestern D.R. Congo, and disjunctly in eastern D.R. Congo (Fig. 8), but is mostly absent from the central depression of the Congo basin (“cuvette congolaise”) where replaced by var. paludosa. Its range closely approaches that of the latter in both Gabon and D.R. Congo, and locally overlaps that of var. floribunda in Gabon. It is found in forest edges and regrowth, 180–1250 m in elevation.

Flowers have been recorded from October to January and April to July, probably all year; fruits from January to July and in September-October, probably all year as well.

Conservation status. – The extent of occurrence (EOO) of S. floribunda var. anisostegia is calculated to be 1,349,092 km^2, well above the limit for “Vulnerable” status under subcriterion B1. Its area of occupancy (AOO) is estimated to be 124 km^2, which falls within the limit for “Endangered” status under subcriterion B2, but is an obvious underestimation, given the low collecting density in most of its range. The taxon occurs in Central Africa from Cameroon to the Democratic Republic of Congo, where it grows in secondary forest, and is known from 40 specimens. Its situation is rather comparable to that of the species as a whole, and it is assessed as “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012) for the same reasons.

Notes. – This is the most distinctive variety of S. floribunda, and further work may show it to be specifically distinct. It is best recognised by the arrangement of the bracts in the inflorescence: var. anisostegia has a few large bracts occurring at scattered nodes and the remainder much smaller, while in other varieties the bracts are more or less gradually decreasing in size from lower to upper nodes (except the lowest pair, which is often smaller than those above it). It is also usually more hairy than other varieties, especially on the stems.

Paratypi. – Angola. Cabinda: near the village Caio in the Hombe country, 9.I.1917, fl. & imm.fr., Gossweiler 7665 (BM, K).

Cameroon. Reg. Central: Réserve forestière Mbalmayo, 15.IV.1967, fl., Mezili 80 (BR, P); Atotzok, 1953, fl. & imm.fr., Surville 312 (YA). Reg. East: 22 km E of Lomié, 12.II.1977, fr., Lowe 3149 (K, YA); Near Echiambor, 18.II.1977, fr., Lowe 3500 (YA); Nkamouna, 11.VI.2004, fl., Sonké 3481 (BR, BRLU); ibid. loco, 11.VI.2004, fr., Sonké 3482 (BR, BRLU); Mbalam II: Mbarga, 24.VI.2008, fl., Sonké 4943 (BR). Reg. South: Bitye, 1917, fl., Bates 885 (BM); Bitye, 1917, buds, Bates 1120 (BM); à l'W d'Alati (100 km SE de Djoum), 12.I.1973, imm.fr., Letouzey 11832 (BR, P, YA).

Democratic Republic of Congo. Ituri: Ituri, Lodjo, near AGK mining camp, 25.I.2011, imm.fr., Bytebier et al. 3343 (BR); Ituri, mont Hoyo, 2.VIII.1975, imm.fr., Lisowski 40879 (BR). Kongo Central: Kimuenza, X–XI.1900, fl., Gillet 1749 (BR); Entre Luzumu et Sanda, vallée de la Ma Ngo, 25.II.1965, fr., Pauwels 4905 (BR). Nord-Kivu: entre Walikale et Kalehe (Kivu), III.1932, fl. & imm.fr., Lebrun 5267 (BR, WAG); Rte Kavumu-Walikale, vers km 110, env. d'Irangi, 11.III.1960, fr., Troupin 12067 (BR). Tshopo: km 58, rte de Kisangani vers Ubundu (rive gauche), 6.IV.1977, fr., Lejoly 1278 (BR, BRLU); rte Kisangani-Ubundu, km 58, 7.IV.1977, fr., Lisowski 44605 (BR, K).

Gabon. Haut-Ogooué: 5 km rte Moanda-Bakoumba, 16.IX.1970, fr., F.J. Breteler 6475 (BR, K, LBV, P, WAG); de Zanaga à Franceville, 21.X.1954, fl., Trochain 9099 (P). Ogooué-Ivindo: 7 km N of km 49 Makokou-Mekambo road which is bifurcation with Bélinga road, near Djadie R., 15.XI.1977, fl., Leeuwenberg 11545 (BR, K, P, WAG); Babièl-Nord ridge, 4 km from camp Bélinga, X.1987, fl., A.M. Louis et al. 2312 (BR, LBV, WAG); Rte de Babiel Nord, few km W of Belinga, 18.VII.1985, fl. & imm.fr., Bos et al. 10658 (BR, K, LBV, P, WAG); Bélinga, 9.VI.1978, fr., Florence 1217 (P); Belinga, 25.VII.1981, fr., Gentry 33569 (P); piste du Bouéni, 20 km SE Makokou, 11.II.1961, fr., N. Hallé 1130 (K, P); 7 km NE Makokou, 11.III.1961, fr., N. Hallé 1454 (P); Bélinga, 26.X.1964, fl., N. Hallé 2804 (K, P); ibid. loco, 28.X.1964, buds, N. Hallé 2866 (BR, P); ibid. loco, 31.X.1964, fl. & fr., N. Hallé 2951 (BR, K, P); ibid. loco, 11.XI.1964, buds, N. Hallé 3126 (P); ibid. loco, 1.I.1980, fl., Pierre-Baltus 98 (P); 25 km on road Mékambo to Madjingo, 27.XII.2000, fl., Wieringa et al. 3648 (BR, LBV, WAG). Ogooué-Lolo: Chantier Bambidie, c. 52 km on the road to Okondja-Lelama, along Sebe river, 31.X.2005, fr., Sosef et al. 2123 (LBV, WAG). Woleu-Ntem: behind the WWF-station, at entrance to Bordamur consession area, 8.X.2002, fr., Strijk 54 (WAG); ibid. loco, 1 km from Assock Begue, 8.X.2002, fl., Strijk 59 (LBV, WAG).

Republic of Congo. Bouénza: forêt de la Bouénza, piste de l'Espérance du village de M'Boumou vers la Bouénza, 16.XI.1964, buds, Bouquet 757 (P). Sangha: West slope of Mt. Nabemba, 15.XI.1991, buds, Thomas et al. 8892 (WAG).

Holotypus: Democratic Republic of Congo. Tshopo: Lilanda (Yangambi), 15.VIII.1940, fl., Germain 449 (BR!; iso-: K!).

Ab alteribus varietatibus differt bracteis ± obovatis utrinque dense pubescentibus (nec orbicularibus, ovatis vel late ellipticis, glabris vel pilis sparsis munitis).

Etymology. – The epithet paludosa refers to the fact that this taxon is most commonly found in swamp and riverine forests, while other varieties usually grow on drained soils.

Distribution, ecology and phenology. – This taxon occurs in northeastern Gabon, and in D.R. Congo where it is found mostly in the central depression of the Congo basin (“cuvette congolaise”) with isolated records further east (Kivu) and south (near the border with Angola) (Fig. 8). Its range closely approaches that of var. anisostegia in both Gabon and D.R. Congo. It grows mostly in swamp and riverine forest, but apparently also in secondary forest regrowth on drained soils, 350–1000 m in elevation.

Flowers have been recorded in January, from April to June and from August to October; fruits from April to October.

Conservation status. – The extent of occurrence (EOO) of S. floribunda var. paludosa is calculated to be 759,404 km^2, well above the limit for “Vulnerable” status under subcriterion B1. Its area of occupancy is estimated to be 64 km^2, which falls within the limit for “Endangered” status under subcriterion B2, but is an obvious underestimation, given the low collecting density in most of its range. The taxon occurs in eastern Gabon and the Democratic Republic of Congo, and is known from 21 specimens representing 15 subpopulations, one of which occurs in a protected area (Kahuzi-Biega National Park in D.R. Congo). It is found in swamp forest and secondary forest regrowth, and appears to be not rare in its range. Given its ecology and wide distribution there is no evidence of a particular threat, and it is here assessed as Least Concern [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – This taxon differs from other varieties by its more or less obovate and densely pubescent bracts (vs. orbicular, ovate or broadly elliptic, with indumentum sparse or absent) which are also usually narrower in proportion – hence a similarity with S. segregata, but that species does not occur in its range. In all other characters, especially leaves and indumentum, it comes very close to var. floribunda.

Paratypi. – Democratic Republic of Congo. Bandundu: Iküma, 13.X.1948, fl., Jans 807 (BR). Equateur: Ikelemba, VI.1934, fl., Dubois 423 (BR); rte Bikoro-Inongo, km 27, 17.X.1957, imm.fr., Evrard 2674 (BR); rte Flandria-Makako (territoire Ingende), 9.IV.1959, fl., Evrard 6054 (BR); Bondombe, 9.IX.1942, buds, Hulstaert 843 (BR); Lac Tumba-I.R.S.A.C. Mabali, 20.VI.1958, imm.fr., Thonet 221 (BR, K). Kwango: entre Popokabaka et Kimabaka, près rivière Pumbu, 18.IV.1959, fl., Pauwels 2495 (BR, P). Kwilu: vallée de la Djuma, VII.1902, imm.fr., Gentil s.n. (BR); région de Bampumu, X.1922, fl., Vanderyst 12481 (BR). Sud-Kivu: Kiaselela, territoire Shabunda, 19.IV.1959, imm.fr., A. Léonard 3900 (BR, P). Tshopo: Opala, 15.VI.1949, imm.fr., Germain 5116 (BR, WAG); Yabahondo (rive gauche du Lomami), 15.X.1952, fl. & imm.fr., Germain 8116 (BR); Yangambi, 3.VI.1958, imm.fr., A. Léonard 759 (BR, K); env. de Kisangani, à 3 km à l'E de Kabondo, 24.X.1977, buds, Lisowski 47453 (BR, K); Kisangani, Kabondo, 6.V.1979, fl. (fallen), Lisowski 52401 (BR, K); 20 km à l'W de Yangambi, 27.X.1938, fl., J. Louis 12110 (BR). Tshuapa: Befale, IX.1934, imm.fr., Dubois 767 (BR); Befale-Tolongote, 5.VI.1958, fl., Evrard 4185 (BR, K, WAG). Sine loco: sine loco, s.d., fl., Pauwels 660 (BR).

Gabon. Ogooué-Ivindo: 10 km on road Malassa to Madouma, 8.I.2001, fl., Wieringa et al. 4028 (LBV, WAG).

Lectotypus (designated here): Cameroon. Reg. South: Gross-Batanga, 3.XII.1890, imm.fr., Dinklage 743 (P [P03793520]!; isolecto-: WAG [WAG0034480]!). Syntypus: Cameroon. Reg. South: Kribi, 24.X.1891, buds, Dinklage 1391 (P [P03793519]!, WAG [WAG0034479]!).

Distribution, ecology and phenology. – This variety occurs in southern Cameroon from Kribi to Campo, in Equatorial Guinea (Rio Muni) and in Gabon, where it is found mostly in the northwest (Crystal Mts) with one record further south in the Chaillu massif (Fig. 8). Its range locally overlaps with that of var. floribunda in Gabon and Equatorial Guinea. It grows in secondary forest regrowth, often along roads, 0–540 m in elevation.

Flowers have been recorded from March to May and October to December; fruits in August-September and from November to March.

Conservation status. – The extent of occurrence (EOO) of S. floribunda var. paucinervis is calculated to be 21,218 km^2, a little above the limit for “Vulnerable” status under subcriterion B1. Its area of occupancy (AOO) is estimated to be 80 km^2, which falls within the limit for “Endangered” status under subcriterion B2, but is certainly an underestimation given the low collecting density in most of its range. The taxon occurs in Atlantic Central Africa (Cameroon, Gabon and Equatorial Guinea) and is known from 21 specimens. It is found in secondary forest habitats where it appears to be locally common, and given its ecology there is no evidence of a particular threat; consequently, it is here assessed as “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – This taxon differs from other varieties by its leaves with fewer secondary veins (8–13 pairs only) and the generally sparser indumentum of its vegetative parts – the stems being glabrous or nearly so, and the leaves glabrous except on veins. Its original description is based on two syntypes, Dinklage 743 and Dinklage 1391 (B), both of which were presumably destroyed in the Second World War but have extant duplicates in P and WAG. The P sheet of Dinklage 743, which is the most complete, is chosen here as lectotype.

Additional material studied. – Cameroon. Reg. South: Kribi-Campo, 35 km S de Kribi, 4.I.1983, imm. fr., van der Zon 1983 (WAG); 3 km S of Kribi, Grand Batanga road, 25.X.1968, fl., Bos 3134 (BR, K, P, WAG, YA); 5 km S of Kribi, Grand Batanga road, 26.II.1969, fr., Bos 4005 (BR, P, WAG, YA); near Eboundja I, km 20 road Kribi-Campo, 4.I.1983, imm.fr., de Kruif 1020 (WAG, YA); Ebundja 2, village along road from Kribi to Campo, 17.III.2007, buds, Dessein 1640 (BR, P); 45 km on the road from Kribi to Campo, 3.XII.1974, fl., J.J.F.E. de Wilde 7784A (BR, WAG); 10 km ENE Ebodje, soit 40 km S Kribi, 7.XII.1979, fl., Letouzey 15322 (BR, P, YA); Campo Ma'an National Park, 12.II.2012, fr., Sonké 5915 (BR, MO, YA); Campo-Ma'an area, Massif des Mamelles, Along the path to Mamelles highlands, 23.IV.2001, buds, Tchouto Mbatchou et al. 3223 (WAG); Dipikar, road to centre of Island, 7.V.2001, fl., van Andel et al. 3421 (WAG); Campo Ma'an National Park, road Kribi-Ebodje Along small logger's road, c. 2.7 km from Lolabé village, 4.VIII.2001, fr., van Andel et al. 3827 (WAG).

Equatorial Guinea. Litoral: Bata-Monson-Dibolo, Estrada kms 54–55, 15.X.1991, fl., Carvalho 4887 (BR, WAG); Bata-Mari, Estrada kms 62–63, 15.XI.1991, imm.fr., Carvalho 4949 (WAG). Centro-Sur: 4 km N de Mitong, 12.XII.2003, buds, Sonké 3136 (BR, BRLU, K).

Gabon. Estuaire: 89 km along the road Libreville-Assok, Mts de Cristal, 14.VIII.1978, imm.fr., F.J. Breteler & J.J.F.E. de Wilde 7 (BR, LBV, WAG); Crystal Mountains, 3 km NE of Kinguélé, 11.IX.1994, fr., F.J. Breteler et al. 12808 (LBV, WAG); km 10 Tchimbélé-Kinguélé, 11.II.1986, fr., Breyne 5102 (BR, LBV); Crystal Mountains, 8 km on the road Tchimbélé to Kinguélé (= 74 km on SEEG road), 15.XI.2004, fl. & fr., Wieringa et al. 5440 (BR, K, LBV, WAG). Ngounié: Chantier Leroy, rte à 13 km de Masika vers Yeno sur rte Mouila-Yeno, 12.II.1988, imm.fr., A.M. Louis 2733 (LBV, WAG). Woleu-Ntem: Tchimbélé, 0.5 km S of dam, 19.III.1990, fr., Wieringa 720 (BR, LBV, WAG).

Lectotypus (designated here): Angola. Cabinda: Belize, 16.XI.1918, fl., Gossweiler 7550 (BM [BM000902997]!; isolecto-: K [K000414624]!, LISC [LISC000807, LISC000808] images seen). Syntypus: Angola. Cabinda: Belize, 12.III.1917, fr., Gossweiler 7043 (BM!, LISC [LISC000809, LISC000810, LISC000811] images seen).

= Pseudosabicea mitisphaera N. Hallé in Adansonia ser. 2, 3: 170. 1963. Holotypus: Gabon. Ogooué-Ivindo: piste du Bouéni, env. 20 km SE Makokou, 11.II.1961, N. Hallé 1129 (P [P00077593]!; iso-: BR [BR0000008852137]!, K [K000414625]!, LISC [LISC002621] image seen, P [P00077594]!).

Distribution, ecology and phenology. – Sabicea nobilis occurs in southeastern Cameroon, Gabon (mostly in the east, absent from the coastal area), southwestern Congo Republic, and Angola (Cabinda) (Fig. 9) and is locally common in its range. It is found in secondary forest, edges and regrowth, always on drained soils, 160–800 m in elevation.

Flowers have been collected from May to August and October to January; fruits from January to May and September to November.

Conservation status. – The extent of occurrence (EOO) of S. nobilis is calculated to be 238,406 km^2, well above the limit for “Vulnerable” status under subcriterion B1. Its area of occupancy (AOO) is estimated to be 132 km^2, within the limit for “Endangered” status under criterion B2, but this is an obvious underestimation since most of its range is not well explored. It occurs in Atlantic Central Africa from southeastern Cameroon to Cabinda (Angola) and is known from 43 specimens representing 30 subpopulations. It occurs in two protected areas, the Doudou Mountains National Park and Batéké Plateaux National Park in Gabon. It is a locally abundant plant of forest edge habitats, and is thus favoured by small-scale forest clearing. Furthermore, most of the area where it occurs is sparsely populated. As a result, there is no particular threat to the species, which is here assessed as “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – The original description of S. nobilis (Good, 1923) is based on two syntypes, Gossweiler 7043 and 7550, both at BM; the latter collection is here selected as lectotype, because it has a larger number of duplicates.

Additional material studied. – Cameroon. Reg. East: South of Echiambor (Lomié to Ngoila), 18.II.1978, fr., Lowe 3515 (K); Nkamouna, 11.VI.2004, buds, Sonké 3480 (BR, BRLU, K); ibid. loco., 15.VI.2004, fl., Sonké 3525 (BR, BRLU, K). Reg. South: E Sangmelima, 16.V.2016, fr., Sonké & Zemagho 6338 (BR, BRLU, K, MO, WAG, YA).

Gabon. Haut-Ogooué: c. 5–10 km of Franceville along road to Okondja, 4.III.1999, fr., Andersson & Nilsson 2316 (BR); Lekoko forest exploitation, 70 km SSW of Moanda, near the border Congo-Gabon, 13.X.1970, fr., F.J. Breteler 6871 (BR, WAG). Moyen-Ogooué: CEFA Ayem, 10 km SW Ndjolé, 10.V.1963, fl., N. Hallé 1974 (BR, K, P,WAG); ibid. loco, 10.V.1963, buds, N. Hallé 1974bis (P); ibid. loco, 6.V.1963, fr., N. Hallé 2001 (K, P). Ngounié: Moukabo, ca 25 km E. of Mouila on the road to Yeno, 27.XI.1984, fl., Arends et al. 468 (BR, LBV, WAG); au N du PN de Waka, 1°09′51″S 11°08′04″E, 24.IX.2007, fr., Boussengui-Nongo et al. 192 (LBV); km 6 Mimongo–Yéno, 21.II.2008, fr., Dessein et al. 2039 (BR, LBV, MO, WAG); Distr. de Mimongo, rte de Lébamba à Etéké km 50, 14.V.1963, fr., N. Hallé & Cours 5873 (P); rte de Lébamba à Etéké km 50, 14.V.1963, fr., N. Hallé & Cours 5874 (BR, P); Rte chantier Leroy-Massika ente Mouila et Yeno, 27.IV.1989, fr., A.M. Louis & Nzabi 3031 (LBV, WAG); Oghoubi camp, on the NE border of Waka National Park, 24.III.2007, fr., Sosef et al. 2529 (BR, LBV, WAG); 25 km on the road Ikobey to Egoubi camp, 1.IV.2004, fr., Wieringa et al. 5203 (BR, LBV, WAG). Nyanga: chantier SFN, Igotchi, 23.XI.2003, fl., van Valkenburg et al. 2604 (LBV, WAG). Ogooué-Ivindo: Masenguelani (60 km SE de Lopé), 30.I.1998, imm.fr., Binot & Lejoly 197 (BRLU); NE du parc de la Lopé, ancienne rte Lopé-Booué, 14 km du carrefour Lopé-Booué, 6.III.2010, fr., Bissiengou et al. 1053 (LBV, WAG); Mintoume, 5.II.2005, fr., Boupoya & Mbazza 186 (BRLU); La Nké, 8.X.1983, fl., Floret et al. 1735 (P, WAG); Booué, 31.VII.1966, fl., N. Hallé & Le Thomas 268 (P); Mintome, 30.III.2003, fr., Mboma et al. 61 (LBV, WAG); bac de l'Ogooué, 9.VIII.1983, fl., Sita 4983 (LBV). Ogooué-Lolo: c. 30 km E of Lastoursville, 22.XI.1991, fl., F.J. Breteler & Jongkind 10653 (BR, LBV, WAG); Bambidie, c. 30 km E of Lastoursville, 1.V.1992, fl. & fr., F.J. Breteler et al. 11205 (BR, G, LBV, WAG); Makande surroundings, c. 65 km SSW of Booué, 30.I.1999, imm.fr., F.J. Breteler et al. 14853 (BR, LBV, WAG); Makandé, Débardage Héron, 14.XI.1993, fl., Dhetchuvi et al. 11 (BRLU); Makandé, près de la station, 18.XI.1993, fl., Dhetchuvi 1308 (BR, BRLU, WAG); c. 50 km SE of Achouka, old forest in Région des Abeilles, 16.XI.1983, fr., A.M. Louis et al. 723 (BR, K, P, WAG); Camp Makande, 27.XI.1993, fl., Moungazi 928 (LBV); Forêt des Abeilles, Makandé, just NE of campsite, 12.I.2001, fl., Wieringa et al. 4075 (LBV, WAG); c. 30 km ENE of Lastoursville, east of Bambidie, 5.0 km on CEB exploration road NZ P/2, 22.III.2013, fr., Wieringa et al. 7673 (WAG). Ogooué-Maritime: Monts Doudou, W. of Doussala and Rés. de Faune de Moukalaba, 4.XII.1984, fl., Arends et al. 607 (LBV, WAG). Woleu-Ntem: Montagne Mengong sur rte Sam vers Mitzic, 25.II.1988, imm.fr., A.M. Louis 2688 (LBV, WAG).

Republic of Congo. Kouilou: Makaba, ancien village Manengue, 17.II.1987, fr., de Foresta 1291 (P); km 36–38, rte de Sounda à Pointe-Noire, 8.II.1966, fr., Farron 5030 (P); bord de la Loukomba, 19.I.1894, fl., Lecomte D6 (P). Lékoumou: village de Mitsiba, après Moetchè, 24.I.1965, fl., Bouquet 1087 (P).

Holotypus: Gabon. Estuaire: Nkoulounga, 11.VII.1959, N. Hallé 748 (P [P00077595]!).

Distribution, ecology and phenology. – Sabicea proselyta occurs in southern Cameroon, Equatorial Guinea (Rio Muni), western and central Gabon (but not in the lower Ogooué basin) and the Mayombe range in southwestern Republic of Congo (Fig. 10). It grows in secondary forest, edges and regrowth, on drained soils or sometimes along rivers, 50–1080 m in elevation.

Flowers have been collected in April, June-July and November; fruits in January (immature), February to May (mature), July (immature) and November (mature).

Conservation status. – The extent of occurrence (EOO) of S. proselyta is calculated to be 162,301 km^2, well above the limit for Vulnerable status under subcriterion B1. Its area of occupancy (AOO) is estimated to be 92 km^2, within the limit for “Endangered” status under criterion B2, but is evidently underestimated because most of its range is not well prospected. The species occurs in Central Africa from south Cameroon to the Congo Republic, and is known from 28 specimens representing 22 subpopulations. It occurs in four protected areas, the Monte Alén National Park in Equatorial Guinea, Waka and Doudou Mountains National Parks in Gabon, and Dimonika Biosphere Reserve in the Republic of Congo. Most of its range is sparsely populated, and because it is a plant of forest edge habitats, it is likely to benefit from small-scale forest clearing, e.g. that induced by selective logging. As a result there is no evidence of a particular threat, and the species is here assessed as “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – Although the protologue (Hallé, 1963) does not specify in which herbarium the type of S. proselyta was deposited, this collection has only one sheet in P, which is therefore considered as the holotype.

Probably because Sabicea proselyta has no very striking characters, it has often been confused with its relatives S. couteronii, S. dichrosepala, S. rubiginosa and S. sanguinosa. The differences between them are summarised in Table 1. Among the 18 Gabonese specimens cited as Pseudosabicea proselyta by Sosef et al. (2006), only nine are correctly identified; six belong to S. rubiginosa (Breteler 9576; Haegens 222; van Nek 128 & 190; Wieringa 2389; J.J.F.E. de Wilde 9688), one to S. dichrosepala (J.J.F.E. de Wilde 284) and two to S. sanguinosa subsp. sanguinosa (Sosef 1613, 1745). Another collection from the Republic of Congo, Kimpouni 600, cited by Lachenaud (2009) as Pseudosabicea proselyta, actually represents S. couteronii. The two collections from the Republic of Congo cited here are therefore the first authentic records of S. proselyta from this country.

The indumentum of the lower leaf surface is usually appressed in this species, but in two collections (J.J.F.E. de Wilde 7762 from Cameroon and van Valkenburg et al. 3140 from Gabon) it is mostly erect with appressed hairs restricted to the veins.These collections match S. proselyta in other characters and are therefore regarded as conspecific.

Additional material studied. – Cameroon. Reg. South: station du Cacaoyer de N'koemvone, S. of Ebolowa, 14 km on the road to Ambam, 28.XI.1974, fl. & fr., J.J.F.E. de Wilde 7762 (BR, K, P, WAG, YA); E Sangmelima, 16.V.2016, fr., Sonké & Zemagho 6329 (BR, BRLU, K, MO, WAG, YA); Campo Ma'an area, Onoyong, in the National Park, 20.III.2001, imm.fr., Tchouto Mbatchou et al. ONOX 93 (WAG).

Equatorial Guinea. Centro-Sur: Monte Alen, 25.III.1997, fr., Ngomo 78 (BRLU); Monte Alen, Transecto 3,900 m del transecto, 21.VII.1998, imm.fr., Ngomo & Ndong 368 (BRLU); PN de Monte Alen, dalle rocheuse d'Engong, 5 km à l'W du village d'Engong, 12.V.2002, fr., Parmentier & Esono 2780 (BRLU); PN de Monte Alen-transect de Monte Chocolate, 8.I.1998, imm. fr., Van Reeth 141 (BRLU). Litoral: Rte de Nco vers Misobong au niveau de la grande cascade, 9.I.1998, imm.fr., N. Nguema & Lejoly 202 (BRLU). Wele-Nzas: entre Nsuamayong et inselberg Akuom, 23.I.1998, imm.fr., Obama & Lejoly 656 (BRLU).

Gabon. Estuaire: km 18 Kougouleu-Kouamé road, 15.XI.1982, fl., Leeuwenberg 12511 (BR, K, LBV, WAG); km 13 Tchimbélé-Kinguélé, 7.II.2008, fr., Dessein et al. 1747 (BR, LBV, WAG); Monts de Cristal, 5 km de Kinguélé, 18.I.1968, imm.fr., N. Hallé & Villiers 4540 (P); Crystal Mountains, 250 m on transect D4, 25.XI.2000, fl., N. Nguema 1459 (BR, LBV, WAG). Moyen-Ogooué: Mboumi, 16.II.2000, imm.fr., Champluvier 6163 (BR, K); c.10 km NNW of Ndjolé, on BSO forestry road, 26.II.2008, fr., Dessein et al. 2150 (BR, LBV, WAG); C.E.F.A. Ayem, 10 km SW Ndjolé, 29.IV.1963, fr., N. Hallé 1918 (K, P); ibid. loco, 29.IV.1963, buds, N. Hallé 1918bis (BR, K, P); ibid. loco, 30.IV.1963, fl., N. Hallé 1955 (K, P); ibid. loco, 8.V.1963, fr., N. Hallé 2022 (P); Abanga, chantier CEFA, 1.VI.1963, fl., N. Hallé 2126 (P). Ngounié: Parc national de Waka, 1°10′34″S 11°08′02″E, 28.XI.2007, imm. fr., Boussengui-Nongo et al. 312 (LBV); Ovala, 16.V.1963, fr., N. Hallé & Cours 5937 (P). Nyanga: chantier SFN, 28.XI.2003, fl. & imm.fr., van Valkenburg et al. 2646 (BR, K, LBV, P, WAG). Ogooué-Maritime: logging road CBG concession eastward of Peni, 20.IV.2005, fr., van Valkenburg et al. 3140 (BR, K, WAG). Woleu-Ntem: 15 km SE of Mitzic, FOREENEX forest exploitation, road from FOREENEX forestry camp to Madouaka village, 7.XI.2009, fl., Bissiengou et al. 752 (LBV, WAG).

Republic of Congo. Kouilou: rte de Dimonika à Pounga, 24.XI.1978, fl., Cusset 514 (P). Niari: Col du Bamba, 6.I.1969, imm.fr., Attims 58 (P).

Holotypus: Gabon. Moyen-Ogooué: Mabounié, 00°46′15″S 10°32′52″E, 31.X.2014, Lachenaud et al. 2105 (BRLU!; iso-: BR [BR0000024387101]!, G!, LBV!, MO!, P!, WAG!).

Habitu sarmentoso, ovario biloculare, inflorescentiis cymosis, foliis utrinque viridibus supra inter nervis glabrescentibus, corolla fauce pilis brevibus inflatis munita, fructibusque albogriseis cum granulis atris in mesocarpio S. proselyta similis, a quae differt calycis bracteisque atro-rubris (nec viridibus), seminibus striolatis (nec grosse reticulatis), calyce tubo intus glabro (nec villoso) et lobis extus villosis (nec puberulis), foliisque subtus cum indumento suberecto pilis brevibus albidis et pilis longioribus rufescentibus intermixtis (nec indumento omnino albido et appresso, raro inter nervis suberecto).

Sarmentose liana, 1–2 m high, climbing or occasionally creeping on the ground; stems cylindrical, 2–5 mm thick, densely covered with c. 1 mm long rufous appressed hairs, intermingled with shorter uncinate hairs. Stipules interpetiolar, dark reddish-brown, 10–15 × 6–11 mm, erect, ovate and ± longitudinally plicate, rounded at base, acute at apex, uniformly pubescent outside with same indumentum as the stems, glabrous inside. Leaves opposite, equal; petiole 0.8–4.8 cm long, with same indumentum as the stems; lamina 6.5–17 × 2.3–7.7 cm, elliptic, acute at base, gradually acuminate at apex, slightly coriaceous, discolorous; upper side dark green, glabrous except for the midrib (and sometimes the secondary veins) with same indumentum as the stems; lower side pale green with brown-red veins, densely pubescent with both soft rufous hairs, 1–1.5 mm long appressed to half-erect, denser on the veins, and shorter whitish erect hairs 0.2–0.5 mm long; secondary veins 9–15 pairs, rather strongly ascending, eucamptodromous; tertiary veins markedly raised on lower leaf surface, much stronger than quaternary veins, the latter densely reticulate, forming areolae < 0.5 mm in diameter. Inflorescences axillary and paired at nodes, cymose or sometimes glomeruliform at anthesis, many-flowered, 1.2–6.5 cm long, densely covered with rufous hairs c. 1 mm long intermingled with shorter uncinate hairs; peduncle 0.3–3.5 cm long; primary ramifications (0–)0.2–2.3 cm long. Bracts dark red, free, appressed-pubescent outside and towards the apex inside, all recaulescent except the basal pair; basal bracts elliptic, entire or rarely dentate towards the base, 4–11 × 1.3–3.5 mm, acute at apex; median bracts elliptic, 4.5–8 × 1–1.7 mm; upper bracts narrowly elliptic, 1–4.5 × 0.2–0.8 mm. Flowers 5-merous, heterostylous; pedicels c. 0.5 mm long, densely villose. Calyx dark red, the base sometimes green or whitish; tube 0.6–0.7 mm long, villose outside like the inflorescence, glabrous inside; lobes narrowly spathulate, 1.5–3 × 0.5–1.2 mm, obtuse to subacute and curved outwards at apex, villose outside, puberulous inside except towards the base, alternating with minute colleters. Corolla pale green in bud, whitish at anthesis; tube cylindrical, 4–4.5 × 1.5 mm, outside glabrous at base and sparsely appressed-pubescent at apex, inside with a ring of multicellular hairs around the upper third and short, inflated unicellular hairs at the throat, the intervening zone with very sparse hairs; lobes triangular, 1.5–1.8 × 1.3–1.5 mm, patent, shortly appressed-hairy outside, glabrous inside; flower buds obtuse, the apex slightly enlarged and pentagonal. Stamens included and attached around the upper third of the corolla tube (long-styled flowers) or half-exserted and attached near the apex of the tube (short-styled flowers); anthers sessile, elliptic, c. 1 × 0.6 mm, glabrous. Ovary c. 1.5 mm long, densely villose, 2-locular. Disk cylindrical, c. 0.5 mm long, glabrous, slightly shorter than the calyx tube. Style c. 4.3 mm long and just reaching throat (long-styled flowers) or c. 3.5 mm long and included (short-styled flowers), bifid, with elliptic flattened stigmas c. 0.8 mm long, these shortly stipitate in long-styled flowers and sessile in short-styled flowers. Fruits whitish, ellipsoid, with persistent calyx, 5–10 × 4–8 mm in life, 4–10 × 3–9.5 mm when dry, villose, with pedicel 0.5–3.5 mm long; mesocarp very soft, whitish, c. 1.5 mm thick, with numerous darker granules. Seeds numerous, pale brown, polyedric, c. 0.8 × 0.5 mm, the surface finely striolate.

Etymology. – The epithet rubiginosa refers to the rusty-coloured indumentum of the leaves and stems.

Distribution, ecology and phenology. – Sabicea rubiginosa is endemic to west-central Gabon, south to Rabi and north to the lower Ngounié river (Fig. 10) and is locally very abundant in its range. It occurs in forest edges, especially along tracks, at low elevations (20–165 m).

Flowers have been recorded in January, March, May, July and October-November, probably most of the year; fruits from January to March, in June-July and in October-November.

Conservation status. – The extent of occurrence (EOO) of S. rubiginosa is estimated to be 4312 km^2, and its area of occupancy (AOO) to be 88 km^2, both values being within the limit for “Endangered” status under subcriteria B1 and B2. The species is endemic to west-central Gabon and is known from 30 specimens, representing five subpopulations, none of which is protected. One of these is situated in a forestry concession, which does not seem to represent a serious threat since the species occurs in forest edges and is likely to be favoured by selective logging. Two subpopulations occur in oil concessions, and one in a mining concession where an important exploitation project is scheduled; a decline in habitat extent and quality and number of individuals is therefore predicted. The five subpopulations represent five locations in the sense of IUCN (2012), and the species qualifies for “Endangered” [EN B1ab(iii,v)+2ab(iii,v)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – Sabicea rubiginosa has previously been mistaken for S. proselyta (see that species), but differs by the indumentum of its lower leaf surface, the ornamentation of its seeds, the absence of hairs inside the calyx tube, and the dark red colour of its bracts and calyces (Table 1; Fig. 12A, C). The latter character is especially conspicuous in the field, and makes S. rubiginosa a remarkably attractive plant, which could certainly be grown for ornamental purposes. The typical subspecies of S. sanguinosa also has red bracts and calyces, but differs by the hirsute and whitish (not rufous) indumentum of its vegetative parts, its leaves that are markedly bullate in the fresh state, its calyx lobes acute at apex and its generally larger bracts (Table 1). The three species, S. proselyta, S. rubiginosa and S. sanguinosa, appear to have separate ranges (Fig. 10).

Paratypi. – Gabon. Moyen-Ogooué: Mabounié, forest area south-west of camp, 6.V.2012, fl., Bidault 470 (BRLU, P); zone de Mabounié, à 45 km au SW de Lambaréné, rive N de la rivière Ngounié, 16.XI.2013, fl., Bidault et al. 1329 (BRLU); Concession froestière GWT, 5.VI.2012, fr., Boupoya & al. 727 (BRLU); Ezanga, 1.II.1991, fr., Dibata 727 (WAG); Mabounié, 13.X.2012, fl., Sonké & Ikabanga 5999 (BR, BRLU, LBV, MO); ibid. loco, 13.X.2012, fr., Sonké & Ikabanga 6001 (BR, LBV, MO); ibid. loco, 23.X.2012, fl., Sonké & Ikabanga 6132 (BR, BRLU, LBV, MO); ibid. loco, 24.X.2012, fl., Sonké & Ikabanga 6140 (BR, BRLU, LBV, MO); ibid. loco, 24.X.2012, fl., Sonké & Ikabanga 6141 (BR, BRLU, LBV, MO); ibid. loco, 25.X.2012, fl. & fr., Sonké & Ikabanga 6153 (BR, BRLU, LBV, MO); ibid. loco, 25.X.2012, fl., Sonké & Ikabanga 6155 (BR, BRLU, LBV, MO); Mabounié, 1.V.2012, buds, Stévart et al. 4125 (BRLU, LBV); ibid. loco, Ngounié River, 4.V.2012, fl., Stévart & Issembé 4176 (BRLU, LBV); Mabounié mine, near Lambaréné, 9.II.2012, imm.fr., Stévart & Droissart 4206 (BRLU, LBV, P). Ngounié: forêt sur la rive gauche (S) de l'Ikoy, à env. 10 km de l'embouchure avec la Ngounié, 26.X.2012, fl., Dauby et al. 2938 (BRLU, LBV); concession Maurel & Prom, près du Lac Ezanga, 22.XI.2013, fl., Lachenaud et al. 1531 (BR, BRLU, LBV, MO, P, WAG); Mabounié, le long de la rivière Ngounié, 12.X.2012, buds, Sonké & Ikabanga 5972 (BR, BRLU, LBV, MO). Ogooué-Maritime: Rabi, 28.III.1990, fl. & fr., F.J. Breteler et al. 9576 (BR, G, LBV, P, WAG); Rabi-Kounga, 8.VII.1998, fl. & imm.fr., F.J. Breteler et al. 14350 (BR, LBV, WAG); Rabi-Kounga, S of Rabi, 1.I.1994, fl. & fr., Haegens & van der Burgt 222 (BR, LBV, WAG); c. 15 km NW of Shell oil exploitation Rabi, 23.XI.1989, fl. & fr., J.J.F.E. de Wilde et al. 9688 (LBV, WAG); Rabi (parcelle Smithsonian), 9.XI.2014, fl., D. Nguema et al. 2843 (BRLU); Pechoud road southwards, 27.X.1990, fr., van Nek 128 (BR, LBV, WAG); Rabi-E, near Pechaud camp, 4.XI.1990, fl. & fr., van Nek 190 (BR, LBV, WAG); Rabi-Kounga, in between Shell-office and camp, 27.X.1992, fl. & fr., von Asmuth & Vosmeer 35 (WAG); Rabi-Kounga, road to Echira, 5.XI.1992, buds, von Asmuth & Vosmeer 58 (WAG); Rabi-Kounga, road to Echira, 5.XI.1992, fl., von Asmuth & Vosmeer 59 (WAG); Rabi, 4½ km on road to Divangui, 4.III.1994, fr., Wieringa & Haegens 2389 (LBV, WAG).

Holotypus: Gabon. Ogooué-Maritime: Petit Bam Bam, 21.VIII.1966, N. Hallé & Le Thomas 573 (P [P00077596]!; iso-: K [K000414623]!, MO [acc. no. MO-05067088] image seen, P [P00077597, P00077598]!).

Distribution, ecology and phenology. – This species is endemic to coastal Gabon between the Ogooué and Komo estuaries, and grows in forest edges on coastal sandy soils; for more precise information see under infraspecific taxa.

Flowers have been collected in February, from May to July and in October-November; fruits in January-February, August and October.

Conservation status. – The extent of occurrence (EOO) of S. sanguinosa is estimated to be 4009 km^2, and its area of occupancy (AOO) to be 48 km^2; both values are within the limit for “Endangered” status under subcriteria B1 and B2. The species is endemic to coastal Gabon, and is known from 14 specimens, representing nine subpopulations, four of which occur in the Wonga-Wongué Presidential Reserve. In spite of its limited range, S. sanguinosa occurs in a sparsely populated region, and is a locally abundant plant of secondary forest habitats, which benefits from small-scale forest clearing (e.g. that induced by selective logging). As a result, there is no evidence of a particular threat, and the species is here assessed as “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – This species, although endemic to Gabon, was omitted from the national checklist of Sosef et al. (2006) where two specimens (Sosef et al. 1613 and Sosef et al. 1745, both of the typical subspecies) were cited as Pseudosabicea proselyta. It indeed shows some resemblance to Sabicea proselyta and S. rubiginosa, but is quite easily separated from both by the hirsute indumentum of its vegetative parts, and its leaves that are conspicuously bullate in life; it also has generally larger bracts and calyx lobes, the latter acute at apex (Table 1).

Two subspecies are here recognised in S. sanguinosa. They differ only by the colour of their bracts and calyx lobes (dark red in subsp. sanguinosa and pale green in subsp. viridis), a character which is obvious in the field but not so in dry material, and was found to be constant in all populations studied. The two subspecies have separate ranges as far as is known, but since the intervening zone is hardly explored botanically, it is not impossible that they may come in contact.

Distribution, ecology and phenology. – This subspecies is endemic to coastal Gabon, occurring in the Wonga-Wongué reserve and slightly further north (Nyonié) (Fig. 10) and grows in forest edges on coastal sandy soils, up to 150 m in elevation.

Flowers collected in January (buds), February, and from May to July; fruits in January-February and August.

Conservation status. – The extent of occurrence (EOO) of S. sanguinosa subsp. sanguinosa is estimated to be 860 km^2, and its area of occupancy (AOO) to be 32 km^2; both values fall within the limit for “Endangered” status under subcriteria B1 and B2. The taxon is endemic to coastal Gabon, and is known from nine specimens, representing five subpopulations, all but one of which occur in the Wonga-Wongué Presidential Reserve. Its situation is rather comparable to that of the species as a whole, and it is assessed as “Least Concern” [LC] for the same reasons using the IUCN Red List Categories and Criteria (IUCN, 2012).

Additional material studied. – Gabon. Estuaire: Savane Boyam, 25.I.1997, fl.b., Delègue 1.Trs (LBV); Oyane, X.1990, st., Minkoue 55 (LBV); Réserve de Wonga Wongué, 23.I.1986, fr., A.M. Louis & J.M. Reitsma 2004 (LBV, WAG); Nyonyie survey, sondage W, 28.VI.1990, fl., Wilks 2028 (WAG); Nyonyie, Transect M2, 15.VII.1990, fl., Wilks 2252 (WAG). Moyen-Ogooué: Eastern part of the Presidential Reserve Wonga-Wongué, c. 100 km S of Libreville, 28.II.1983, fl. & fr., J.J.F.E. de Wilde et al. 815 (BR, K, LBV, WAG). Ogooué-Maritime: à ± 10km au SSE de Batanga, 21.V.2001, fl., Sosef 1613 (K, LBV, P); ca 10 km à l'Est de Batanga, 26.V.2001, fl., Sosef 1745 (K, LBV, P).

Holotypus: Gabon. Ogooué-Maritime: région du Lac Alombié, ± 7 km au N de Mpaga, 0°50′20″S, 9°27′14″E, 14.X.2014, fl. & fr., Lachenaud, Ikabanga & Lafferty 1918 (BRLU!; iso-: BR!, G!, LBV!, MO!, P!, WAG!).

A typo differt calycis bracteisque pallide viridibus, nec atro-rubris.

Distribution, ecology and phenology. – This subspecies is endemic to the lower Ogooué delta in Gabon, between the coast and Lake Alombié (Fig. 10); it has thus a more southern range than subsp. sanguinosa. It occurs in the same habitat as the latter, and is locally very abundant.

Flowers recorded in October-November, fruits in October (probably in other months as well).

Conservation status. – The extent of occurrence (EOO) of S. sanguinosa subsp. viridis is estimated to be 483 km^2, and its area of occupancy (AOO) to be 16 km^2, both values being within the limit for “Endangered” status under subcriteria B1 and B2. The taxon is endemic to the Ogooué delta in Gabon, and is known from five specimens, representing four subpopulations. It is not known from any protected area, but probably occurs in the Wonga-Wongué Presidential Reserve since it has been collected very close to its limits. In spite of its restricted range, it occurs in a sparsely populated area, and is a locally abundant plant of forest edge habitats, which benefits from small-scale forest clearing (e.g. that induced by selective logging). As a result, there is no evidence of a particular threat, and the taxon is assessed as “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Paratypi. – Gabon. Ogooué-Maritime: Préfecture de M'paga, à env. 7 km au N du lac Alombié, au S de la réserve de Wonga Wongué, 11.X.2014, fl. & fr., Bidault et al. 1691 (BR, BRLU, MO, P); delta de l'Ogooué en face de Port-Gentil, au S de la Pointe Fétiche, 22.XI.2016, fl., Lachenaud et al. 2270 (BR, BRLU, G, K, LBV, MO, P, WAG); env. 5 km au SE de Mbilapé, 24.XI.2016, fl., Lachenaud et al. 2314 (BR, BRLU, G, LBV, MO, P, WAG); riv. de Kendié en aval de Mbilapé, rive E, 25.XI.2016, fl., Lachenaud et al. 2335 (BR, BRLU, LBV, MO, WAG).

Lectotypus (designated here): Gabon or Equatorial Guinea: River Muni, VIII.1862, Mann 1766 (K [K000414626]!; isolecto-: P [P00077599]!).

= Sabicea henningsiana Büttner in Bot. Vereins Prov. Brandenburg 31: 79. 1889. Lectotypus (designated here): Gabon. Estuaire: Sibangefarm, IX.1884, fl., Büttner 437 (K!). Syntypus: Gabon. Estuaire: Sibangefarm, X.1884, Büttner 443 (B†).

Distribution, ecology and phenology. – This species occurs in southern Cameroon where rare, Equatorial Guinea (Rio Muni), Gabon where very common and widespread (though absent in southeast, extreme northeast, and lower Ogooué basin), and southwestern Republic of Congo (Fig. 14). It is found in secondary forest, edges and regrowth, and in fringing forest on inselbergs, always on drained soils, on various substrates (sandy or clayish), 0–1080 m in altitude.

Flowers and fruits have been recorded in all months.

Conservation status. – The extent of occurrence (EOO) of S. segregata is estimated to be 250,602 km^2, and its area of occupancy (AOO) to be 396 km^2. The latter value falls within the limit for “Vulnerable” status under criterion B2, but is an obvious underestimation given the low collecting density in most of its range and the fact that the species is generally common there. The species occurs from south Cameroon to the Republic of Congo, and is known from 124 specimens representing 71 subpopulations. It occurs in several protected areas: Campo Ma'an National Park in Cameroon, Monte Alén National Park in Equatorial Guinea, Lopé, Waka, Doudou Mountains and Ivindo National Parks in Gabon, and Dimonika Biosphere Reserve in the Republic of Congo. In view of its wide distribution, abundance, and preference for degraded forest habitats, it is here assessed as “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes. – This species is closely related to S. floribunda, both having the corolla lobes brown at anthesis, unlike other species of the group. The differences between them are summarised in the key. The calyx lobes of S. segregata are usually longer and narrower than in other species of the group, but there is limited overlap with S. floribunda in this character.

Most collections of S. segregata, including the type, have stems with appressed hairs, but a variant with hirsute stems occurs in Gabon, particularly in the east of the country. The fruits apparently turn from whitish to red to purple-black; however, since the last colour is much less commonly reported than the other two, it is not sure whether this sequence is always followed.

The original description of S. segregata (Hiern, 1877) does not mention the type number, but the only possible candidates are Mann 1766 in Kew, and Mann s.n. in Paris, the latter being almost certainly an unnumbered duplicate of the former. The Kew sheet is here selected as lectotype. It is not known on which side of the Muni River, which forms the border between Equatorial Guinea and Gabon, this material was collected.

The description of S. henningsiana (Büttner, 1889) is based on two syntypes, Büttner 437 and Büttner 443, both from Sibang in Gabon. The original sheets of both specimens have presumably been destroyed in Berlin, but a duplicate of the former remains in Kew, and is here selected as lectotype.

Additional material studied. – Cameroon. Reg. South: Campo-Ma'an area, Ebodje, Along the beach, 20.II.2000, fr., Elad et al. 1367 (WAG); Campo-Ma'an area, Mvini, Bigan, Along Trasect7, 16.II.2000, imm.fr., Tchouto Mbatchou et al. 2524 (WAG); Bipindi, s.d., buds, Zenker s.n. (BR, P).

Equatorial Guinea. Centro-Sur: Parc National de Monte Alen, dalle rocheuse d'Engong, 5 km à l'W du village d'Engong, 12.V.2002, fr., Parmentier & Esono 2781 (BRLU); 4 km N de Mitong, 12.XII.2003, imm.fr., Sonké 3138 (BR, BRLU, K); 6 km N de Mitong, 12.XII.2003, fl., Sonké 3151 (BR, BRLU, K); Engong, 14.VIII.2001, fl., Sonké & Esono 2546 (BR, BRLU, WAG); Engong, 14.VIII.2001, fr., Sonké & Esono 2552 (BR, BRLU, WAG). Litoral: Bata-Monson-Dibolo: Estrada kms 54–55, 15.X.1991, fl., Carvalho 4889 (BR, BRLU, WAG); Bata-Nasanga: Estrada km 14, 17.X.1991, fl., Carvalho 4890 (WAG); Bata-Mari: Estrada kms 62–63, 15.XI.1991, imm.fr., Carvalho 4952 (WAG); Engong, 28.VII.1999, fr., Eneme 439 (BRLU); Etembue, 15.IX.2001, buds, Esono & Ndong 335 (BRLU); Ndote Nord, 28.VIII.1997, fr., Lisowski M-153 (BRLU). Reg. unknown: Akanabot, 15.IX.2001, fr., Esono & Ndong 351 (BRLU).

Gabon. Estuaire: Mondah Forest on Cape Esterias, c. 22 km on the road from Libreville, 3.IX.1978, fl. & fr., F.J. Breteler et al. 408 (BR, K, WAG); plantations de Assoukou, près du poste de Kango, 2.X.1912, buds, Chevalier 26847 (K, P); plantation Stéphane, sur la Bokoué, 9.X.1912, fl., Chevalier 26989 (P); vieux chantier ‘Sango’, Ntoum, 9.X.1987, fl., Dibata 320 (BR, WAG); Nkoulounga, 31.I.1961, fl., N. Hallé 1024 (P); sine loco, 15.X.1896, fl., Klaine 100 (P); sine loco, 12.IX.1896, fl., Klaine 123 (P); Env. de Libreville, 18.VIII.1898, fl. & fr., Klaine 377 (P); env. de Libreville, 1897, fr., Klaine 771 (P); Env. de Libreville, 15.VI.1897, fr., Klaine 962 (P); env. de Libreville, 5.X.1901, fl. & fr., Klaine 2432 (BR, K, P); km 31 Kougouleu-Atogafina, NE of Kouamé, 19.XI.1982, imm.fr., Leeuwenberg 12554 (LBV, WAG); rte Cap Santa-Clara, 6 km à droite, 31.I.1985, fr., A.M. Louis & J.M. Reitsma 1677 (BR, LBV, WAG); Forêt Classée de la Mondah, 0°32′48″N 9°19′40″E, 20.II.1994, fr., Lowry 4566 (LBV); c. 15 km N. of Libreville, 18.III.1987, fr., J.M. & B. Reitsma 3166 (LBV, WAG); forêt de la Mondah. 5 km on off road to Santa Clara from Cap Esterias road, 11.XI.1988, fl., van der Maesen et al. 5440 (WAG). Haut-Ogooué: concession CEB, W of Okandja, 19.XI.2015, imm. fr., Texier & Akouangou 174 (BRLU). Moyen-Ogooué: Mabounié, à 45 km au SW de Lambaréné, rives de la Ngounié, 15.X.2012, fl., Bidault et al. 845 (BRLU, LBV, P); ibid. loco, 16.X.2012, buds, Bidault et al. 906 (BRLU, LBV); ibid. loco, 15.XI.2013, fl., Bidault et al. 1316 (MO); au S de Lambaréné, entre 5 et 10 km depuis la ville, entre l'Ogooué et la rte de Fougamou, 13.IV.2015, fr., Bidault et al. 1922 (BR, BRLU); Concession forestière GWT (Global Wood and Trading), 5.VI.2012, imm.fr., Boupoya & al. 744 (BRLU); Mboumi, 16.I.2000, fr., Champluvier 6008 (BR); ibid. loco, 16.I.2000, fr., Champluvier 6019B (BR); c.10 km NNW of Ndjolé, on BSO forestry road, 26.II.2008, fl., Dessein et al. 2148 (BR, LBV, MO, WAG); Ayem, 10 km SW de Ndjolé, 11.IV.1963, fl., N. Hallé 1621 (P); Ayem, 10 km SW de Ndjolé, 30.IV.1963, fl. & fr., N. Hallé 1952 (P); Ayem, 10 km SW de Ndjolé, 6.V.1963, fl., N. Hallé 2007 (P); N of lake Azingo, 4.VI.2014, imm.fr., Libalah & Ikabanga 3 (BR, BRLU); Mabounié, 18.X.2012, fl., Sonké et al. 6108 (BR, BRLU); ibid. loco, 23.X.2012, fl., Sonké & Ikabanga 6127 (BR, BRLU); ibid. loco, 23.X.2012, fl., Sonké & Ikabanga 6137 (BR, BRLU); ibid. loco, 23.X.2012, fl. & fr., Sonké & Ikabanga 6139 (BR, BRLU, P); Rougier Logging Concession, Haut Abanga. East of Ndjolé, 17.II.2012, fr., Stévart et al. 4301 (BRLU, LBV, P); Ndjolé, I.1895, fr., Thollon 117 (P); roadside between Ndjole town and railway station, 22.VII.1986, fr., Thomas & Wilks 6589 (LBV, P, WAG). Ngounié: Waka forest exploitation road, 19.XI.1984, fl., Arends et al. 301 (BR, K, LBV, P, WAG); PN de Waka, 1°10′34″S 11°08′02″E, 28.XI.2007, imm.fr., Boussengui-Nongo et al. 313 (LBV); Mouyanama falls, at base of Mt Songou, 22.II.2008, fr., Dessein et al. 2041 (BR, LBV); entre Pagha & Mouteti, 26.XI.1924, fl., Le Testu 5104 (BM, BR, K, P, WAG); Parc national de Waka, près du Camp Oghoubi, 24.VI.2006, fr., J. Mayombo et al. 1559 (LBV); 25 km on the road Ikobey to Egoubi camp, 1.IV.2004, fr., Wieringa et al. 5204 (LBV, WAG); road Mouila to Yeno, c. 2 km before Issinga, 8.III.2013, fr., Wieringa et al. 7169 (WAG). Nyanga: rte Tchibanga-Ndende, 25.X.2009, fl., Bissiengou et al. 540 (BR, LBV, WAG); 48 km SSW of Doussala, road Doussala-Igotshi, 21.III.1988, fr., J.J.F.E. de Wilde et al. 9485 (BR, K, LBV, WAG); Tchibanga, IX.1915, fl., Le Testu 2120 (BM, BR, K, P); concession SFN, 30.X.2003, fl. & imm.fr., van Valkenburg et al. 2523 (BR, K, LBV, P, WAG); forestry concession of Bakker, 6 km N of Igotchi, 29.X.2003, fl., Wieringa et al. 5025 (BR, K, LBV, WAG); forestry concession of Bakker, 15 km NNE of Igotchi, 31.X.2003, fl., Wieringa et al. 5047 (LBV, WAG). Ogooué-Ivindo: concession forestière Rougier-Ivindo. En bordure du fleuve, au N de la concession, 29.X.2015, fl., Bidault et al. 2269 (BR); Mintoume, 5.II.2005, fr., Boupoya & Mbazza 185 (BRLU); station de recherche de Mpassa, 10 km S de Makokou, 9.III.1984, fl., Doumenge 191 (LBV, P); M'passa, Ile aux chauve-souris, 17.III.1977, fr., Florence 55 (LBV, P); M'passa, 1.VI.1977, fl., Florence 496 (LBV, P); La Nké, ancienne zone d'exploitation, 8.X.1983, buds, Floret et al. 1732 (WAG); piste du Bouéni, 20 km SE Makokou, 11.II.1961, fr., N. Hallé 1131 (P); Makokou, Mission Catholique, 16.II.1961, imm.fr., N. Hallé 1185 (P); ± 50 km SE Makokou, 19.X.1964, buds, N. Hallé 2695 (P); 9 km SSW of Makokou, near Ivindo R., right bank, 2.XI.1977, fl., Leeuwenberg 11418 (BR, K, WAG); Mintome, 30.III.2003, fr., Mboma et al. 64 (LBV, WAG); Ivindo National Park, rte Langoué, 11.IV.2004, fr., Moungazi 1557 (BR, LBV, WAG); Ivindo National Park, Massouna 2000, 0°08.90′S 12°27.10′E, 6.XII.2003, imm.fr., D. Nguema Ekomo 507 (LBV) & 508 (LBV); bac de l'Ogooué, 19.VIII.1983, imm.fr., Sita 4964 (LBV); Kongou waterfall area, 00°17 ′25″N 012°34′22″E, 8.II.2018, imm.fr., Texier et al. 2282 (MO); Lopé, West Woods, 26.X.1990, buds, L. White 172 (LBV); Lope Reserve, 15.X.1995, buds, L. White 1495 (LBV, WAG). Ogooué-Lolo: Bambidie, c. 30 km E of Lastoursville, 2.V.1992, fl., F.J. Breteler et al. 11248 (LBV, WAG); Makande surroundings, c. 65 km SSW of Booué, 2.II.1999, imm.fr., F.J. Breteler et al. 14892 (WAG); Makandé, débardage Héron, 20.XI.1993, fl., Dhetchuvi et al. 1372 (BRLU); Vers 10 km rte Makandé-Gongué, 3.XII.1993, fl. & imm.fr., Dhetchuvi et al. 1597 (BR, BRLU, K, WAG); Lastoursville, concession forestière CEB, 30.XI.2012, fl., Ikabanga & Haurez 343 (BRLU); c. 50 km SE of Achouka, old forest in Région des Abeilles, 17.XI.1983, fl., A.M. Louis et al. 756 (BR, K, P, WAG); Camp Makande, 16.XII.1993, imm. fr., Moungazi 967 (LBV); Camp concession SEEF, area of Mount Ngouadi, 00°16′12″S 013°05′00″E, 10.III.2017, fr., Texier et al. 707 (LBV, MO); ibid. loco, 00°17′20″S 013°10′40″E, 11.III.2017, fl., Texier et al. 795 (LBV, MO); ibid. loco, 11.III.2017, fr., Texier et al. 798 (LBV, MO); SIAEFG logging concession, 30 km N of Pana, 01°28′44″S 12°32′55″E, 9.XII.2017, fl., Texier et al. 1564 (LBV, MO); S of Road Lopé-Lastoursville, c. 60 km, 00°43′21″S 012°28′30″E, 18.XII.2017, imm.fr., Texier et al. 1761 (LBV, MO). Ogooué-Maritime: Toucan, 12.X.2002, fl., Bourobou et al. 993 (LBV, P, WAG); Rabi, c. 1°55′S 9°52′E, 13.X.2002, fr., Bourobou et al. 1022 (LBV); Gamba, 24.IX.1968, buds, Breteler & van Raalte 5675 (BR, WAG); Foot of Doudou Mountains, 25 – 35 km W of Mandji, 15.II.2008, fr., Dessein et al. 1862 (BR, LBV); between Rabi and Echira, 27.III.1990, fr., F.J. Breteler et al. 9557 (LBV, WAG); Rabi-Kounga, near Rabi 44, 1.I.1994, fr., Haegens & van der Burgt 226 (LBV, WAG); Rabi-Kounga, 15.X.1991, fr., Schoenmaker 33 (WAG); Pechoud road southwards, 27.X.1990, fl., van Nek 124 (BR, LBV, WAG); Rabi, 4½ km on road to Divangui., 4.III.1994, fr., Wieringa & Haegens 2387 (BR, LBV, WAG). Woleu-Ntem: c. 6 km SSW of Mitzic, FOREENEX forest exploitation, 6.XI.2009, fl., Bissiengou et al. 703 (LBV, WAG); Parc des Monts de cristal, 13.II.2010, fl., Bissiengou & Niangadouma 975 (LBV, WAG); between Zomoko and Saint Germain, c. 10 km SE of Mitzic, 17.IV.1988, fr., F.J. Breteler & Dibata 8771 (BR, K, LBV, WAG); 5–10 km E of Saint Germain, E of Okano River, 20.IV.1988, fl., F.J. Breteler & Dibata 8866 (BR, K, LBV, WAG); Mts de Cristal, inselberg Milobo, 10km N Mbé Akélayong, 50km W Medouneu, 3.XII.2001, fl., Degreef 254 (BR); SEEG road near Tchimbélé, pylone 7, 6.II.2008, fr., Dessein et al. 1723 (BR, LBV); Angoum, VII.1933, buds, Le Testu 9199 (BM, P); Along forestry road Oveng-Mitsic, 28.III.1985, fl., J.M. Reitsma & B. Reitsma 731 (LBV, WAG); Tchimbélé Dam, Crystal Mountains, road N. of dam., 12.XI.2000, fl., Walters et al. 499 (BR, LBV, WAG).

Republic of Congo. Kouilou: rte de Dimonika à Pounga, 24.XI.1978, fr., Cusset 520 (K, P, WAG); env. de Dimonika, M'Bulu Pambu, 7.III.1980, fr., Cusset 956 (P); env. de Dimonika, après Makaba carrefour, 13.XII.1982, fr., Cusset 1173 (P); Makaba, ancien village Manengue, 17.II.1987, fr., de Foresta 1296 (P); rte de Sounda (Pointe-Noire), 8.II.1966, imm.fr., Farron 5024 (P); near Koulila, 6.XII.1990, imm.fr., LaCroix 5005 (K); Kakamoeka, 11.X.1990, buds, Lisowski B-8005 (BR, K, WAG); en allant vers Makaba, village à 17 km de Dimonika, 4.XII.1972, fr., Makany 2045 (P); N'Tiétié, forêt à 3 km du village, 8.XII.1974, fl., Sita 3807 (BR, P). Niari: Mayombe au col de Bamba, 8.XII.1990, fl. & imm.fr., Dowsett-Lemaire 1392 (BR).