Trees, scrambling shrubs or lianas, up to 50 m tall, monoecious, dioecious or putatively androdioecious. Indumentum, when present, of simple, fasciculate, stellate, or scale-like hairs. Leaves alternate, simple, distichous, margins entire, stipules absent. Inflorescences terminal or axillary, ramiflorous in leaf axils, on young or old leafless branches or cauliflorous, single to many-flowered, pedunculate or subsessile, bracts often present. Flowers bisexual or unisexual, actinomorphic, generally trimerous. Sepals in a single whorl, (2)3(4), valvate or imbricate in bud, free or basally to fully connate. Petals 3, 4 or 6, in 1 or 2(3) whorls, generally differentiated into an inner and outer whorl alternating with the sepals, valvate or imbricate in bud, free, basally or fully connate. Stamens 3 to numerous, inserted onto a flat or convex receptacle; anthers generally exceeded by the connective apex, which forms a a protective cover at the top of the stamen; connective apex flat, extended (tongue shaped) or absent; filaments short or absent, free or rarely fused; staminodes absent or present. Carpels 1 to numerous, free or more rarely fused (syncarpous) in flower; stigma capitate, oblong or variously folded; ovules 1 to numerous, uni- or biseriate, basal or lateral. Fruit generally apocarpous, each carpel producing a single monocarp, or more rarely pseudosyncarpous (carpels fusing during fructification) or syncarpous (unilocular fruits resulting from syncarpous flowers), indehiscent or sometimes dehiscent; monocarps 1 to numerous, sessile to long-stipitate, cylindrical, globose, ovoid, ellipsoid, club-shaped or moniliform, 3 to over 40 mm in length, usually large; seeds 1 to numerous per monocarp, uni- or biseriate, or unordered in syncarpous species, sometimes with arilor sarcotesta; endosperm ruminate, hard.

Pantropical, from the Pacific and northern Australia to South East Asia (including southern China), India, Madagascar, tropical Africa, temperate eastern North America south to Central America and South America. 113 genera, and around 2550 species.

In Cameroon, 28 genera and 163 species reported to date.

Several Annonaceae species have been introduced and are commonly cultivated across the country. Monoon longifolium (Sonn.) B.Xue & R.M.K.Saunders (=Polyalthia longifolia (Sonn.) Hook.f. & Thomson) is sold and grown as an ornamental, and planted mainly along roads in major towns and gardens. Cananga odorata (Lam.) Hook.f. & Thomson (ylang ylang) is sold and planted as an ornamental, with its large flowers emitting a strong sweet scent especially at night. Several non-native species of Annona are planted in gardens for their large sweet fruits, especially A. muricata L. Annona glabra L. is naturalized in coastal mangrove regions of West Africa including Cameroon. These non-native species are not treated here.

Afroguatteria bequaertii (De Wild.) Boutique.

Same as species.

A genus of lianas with three species from Central Africa, in the Democratic Republic of the Congo, Cameroon and Angola (Cabinda); one species in Cameroon, endemic.

This genus was phylogenetically validated (Guo et al. 2017b).

no revision has yet been published, but see Boutique (1951b) and Paiva (1966).

Cameroon. South Region; Bipindi, Zenker G.A. 2980, Apr 1904: holotype: B[B10 0153055]; isotypes: BM[BM001125042]; BR [BR000008800398]; [BR000008800398]; G[G00308361]; GOET[GOET005676]; HBG[HBG-502538]; K[K000198949]; L[L.1754813]; M[M-0107910]; P[P00363341]; S[S03-2239]; WAG[WAG0053550].

Liana, height unknown, d.b.h. unknown. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches pubescent with short sericeous hairs. Leaves: petiole 3–4 mm long, 1–2 mm in diameter, sparsely pubescent to glabrous, grooved, blade inserted on the side of the petiole, 7.5–10 cm long, 4–5 cm wide, elliptic, apex acuminate, acumen ca.1 cm long, base obtuse, subcoriaceous, below sparsely pubescent to glabrous when young, glabrous when old, above glabrous when young and old, discolorous, whitish below (both when fresh and dry); midrib flat or sunken, above glabrous when young and old, below sparsely pubescent when young, glabrous when old; secondary veins 8 to 13 pairs, glabrous above; tertiary venation reticulate. Individuals bisexual; inflorescences cauliflorous or ramiflorous on young foliate branches, leaf-opposed or extra-axillary. Flowers with 9 perianth parts in 3 whorls, 1 per inflorescence; pedicel 18–22 mm long, ca. 1 mm in diameter, pubescent; in fruit 20–40 mm long, 2–3 mm in diameter, pubescent; bracts not seen; sepals 3, valvate, free, 1–2 mm long, 1–2 mm wide, triangular, apex acute, base truncate, pubescent outside, glabrous inside, margins flat; petals free, subequal; outer petals 3, valvate, 5–6 mm long, 3–4 mm wide, ovate, apex acute, base narrowed, margins flat, pubescent outside, pubescent inside; inner petals 3, valvate, 3–3.5 mm long, 2–3 mm wide, elliptic, apex obtuse, base truncate, margins flat, pubescent outside, pubescent inside; number of stamens not counted, number of rows not seen, 2–3 mm long, oblong; anthers not septate; connective discoid, pubescence not seen; staminodes absent; carpels not seen but free. Monocarps stipitate, stipes 5–10 mm long, 1–2 mm in diameter; monocarps 3 to 7, 10–15 mm long, 5–8 mm in diameter, ellipsoid, apex apiculate, pubescent, smooth, not ribbed, color unknown; seed 1, 13–15 mm long, 5–7 mm in diameter, ellipsoid; aril absent.

endemic to Cameroon; known from the South Region.

A rare species, in primary lowland rain forests. Altitude 100–200(?) m a.s.l.

None recorded.

NE (Not Evaluated), but probably CR.

None recorded.

Afroguatteria discostigma is only known from four collections by Zenker, all collected close to the type locality in Bipindi (South Region). It remains incompletely known and measurements here are incomplete. The species can be distinguished by its almost glabrous vegetative parts (young foliate branches and underside of young leaves can be pubescent with short hairs), its small leaves that are glaucous below (both when fresh and dry) and branches drying black. The flowers are borne on terminal pedicels that appear leaf-opposed or extra-axillary, but can also be cauliflorous (Zenker 3023), the carpels have a single ovule and thus monocarps are single-seeded like in Afroguatteria bequaertii (De Wild.) Boutique) (Boutique 1951a). The altitude range given here is the one around Bipindi, but could be higher given that the mountain range Ngovayang (up to 1000 m) is very close.

A recent molecular phylogenetic study showed that this species (under the name Friesodielsia discostigma) clustered in the genus Afroguatteria being sister to the Congolese species A. bequaertii (Guo et al. 2017b). It is thus quite different genetically from the the African species of Friesodielsia (now Monanthotaxis) in which it was placed before based on morphology (van Steenis 1964).

Figure 4. 

Afroguatteria discostigma A branch B upper and lower side of leaf, notice glaucous lower side and network like secondary veins C detail of upper lower side of leaf and petiole D inner (left) and outer petals (right), inner view E stamens front and side views F carpels side view and view of ovules G infructescence with moniliform monocarps A–C from Zenker 2980 [S03-2239] reproduced from Swedish Museum of Natural History Department of Botany (S) https://plants.jstor.org/ D–G from Zenker 3023. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris.

Map 1. 

A Afroguatteria discostigma B Annickia affinis C Annickia chlorantha D Annickia letestui E Annickia polycarpa F Annona senegalensis G Annona senegalensis subsp. oulotricha H Anonidium brieyi I Anonidium mannii. White borders represent region limits in Cameroon; green patches represent protected areas (see methods and Suppl. material 1: Fig. S1).

South Region: Bipindi, 3.05°N, 10.25°E, 01 January 1905, Zenker G.A. 2102a (BM,BR,E,G,K,P); Bipindi, 3.05°N, 10.25°E, 01 January 1904, Zenker G.A. 3023 (BM,G,K,L,P,WAG); Bipindi, 3.08°N, 10.41°E, 01 March 1914, Zenker G.A. 576 (MA).

Enantia chlorantha Oliv. (≡ Annickia chlorantha (Oliv.) Setten & Maas).

Trees, up to 30 m tall, d.b.h. up to 50 cm; stilt roots or buttresses absent, slash yellow. Indumentum of simple, stellate and/or fasciculate hairs. Leaves: petiole 2–9 mm long, 1–2 mm in diameter; blade 3.5–29.5 cm long, 1.5–10.5 cm wide, elliptic to obovate, apex acuminate to acute, base narrowly cuneate to shortly attenuate to rounded; midrib sunken or flat; secondary veins 8 to 13 pairs; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, leaf opposed or extra axillary. Flowers with 6 perianth parts in 2 whorls, 1 per inflorescence; pedicel 4–19 mm long; in fruit 10–27 mm long; bracts 1 to 2, basal and one upper towards the middle or lower half of pedicel, 2–8 mm long; sepals 3, valvate, free, 5–22 mm long, triangular, apex acute, base truncate; petals free; outer petals absent; inner petals 3, valvate, 12–34 mm long, 5–19 mm wide, ovate, margins inversely Y-shaped ridged, apex acute, base broad and concave; stamens 60 to 200, in 5 to 6 rows, 2–4 mm long, linear; connective tongue shaped or flattened, glabrous; staminodes absent; carpels free, 20 to 70, ovary 2–4 mm long, stigma lobed or cylindrical, pubescent. Monocarps stipitate, stipes 6–59 mm long, 5 to 55 monocarps, 18–35 mm long, 4–14 mm in diameter, ellipsoid to obovoid, apex sometimes mucronate, smooth, glossy; seed 1, 20–30 mm long, ca. 10 mm in diameter, ellipsoid; aril absent.

A genus of eight species mostly distributed across west and central Africa, with one endemic species in Tanzania; four species occur in Cameroon, none endemic.

The genus is easily identifiable when sterile by its yellow slash, and when fertile, by its leaf opposed or extra-axillary (terminal) flowers with 3 sepals and 3 petals, and stipitate monocarps with a single seed.

Versteegh and Sosef (2007).

Angola. Cabinda, Munze, ring at Buco Zau, Gossweiler J. 6675, 11 Sep 1916: holotype K[not seen]: isotypes: BM[BM000547034]; COI[COI00004913]; LISC[LISC000073, LISC000072, LISC000075, LISC000074].

Tree, 3–30 m tall, d.b.h. 3–50 cm; stilt roots or buttresses absent, slash yellow. Indumentum of simple, bifid and fasciculate hairs; old leafless branches glabrous, young foliate branches sparsely pubescent. Leaves: petiole 2–8 mm long, 1–2 mm in diameter, sparsely pubescent, grooved, blade inserted on the side of the petiole; blade 3.5–26 cm long, 1.5–9.5 cm wide, elliptic to obovate, apex acuminate to acute, acumen 1 cm long, base narrowly cuneate to shortly attenuate, coriaceous to subcoriaceous, below pubescent when young and old with simple or bifid hairs pointing towards the leaf apex, above sparsely pubescent when young and old, concolorous; midrib sunken or flat, above sparsely pubescent to glabrous when young and old, below pubescent when young and old; secondary veins 8 to 13 pairs, sparsely pubescent below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, leaf opposed or extra axillary. Flowers with 6 perianth parts in 2 whorls, 1 per inflorescence; pedicel 7–14 mm long, 1–2 mm in diameter, pubescent; in fruit 27 mm long, 2–3 mm in diameter, pubescent; bracts 1–2, basal and one upper towards the middle of pedicel, ca. 4 mm long, ca. 2 mm wide; sepals 3, valvate, free, 7 mm long, ca. 4 mm wide, triangular, apex acute, base truncate, green, pubescent outside, glabrous inside, margins flat; petals free; outer petals absent; inner petals 3, valvate, 15–33 mm long, 5–15 mm wide, ovate to inversely Y–shaped ridged, apex acute, base broad and concave, greenish yellow, margins flat, pubescent outside, glabrous inside; stamens 110 to 175, in 5 to 6 rows, 2–4 mm long, linear; connective tongue shaped, glabrous, yellow; staminodes absent; carpels free, 35 to 70, ovary 3–4 mm long, stigma lobed, pubescent. Monocarps stipitate, stipes 10–40 mm long, 1–2 mm in diameter; monocarps 3 to 34, 20–35 mm long, 9–14 mm in diameter, ellipsoid to obovoid, apex sometimes mucronate, sparsely pubescent, smooth, glossy, black when ripe; seed 1, ca. 30 mm long, ca. 10 mm in diameter, ellipsoid; aril absent.

From Nigeria (one collection) to the Republic of Congo and the extreme west of the Democratic Republic of Congo; in Cameroon known from the East, South, Littoral, Center and South-West regions.

A very common species; in lowland rain forests in primary and secondary habitats. Altitude 50–650 m a.s.l.

Bololo, Bonuke, Bunuku bolobo (dial. Duala); Bululu, Mfo, Pobalo, Ufol, Moabé (dials. Ewondo, Bulu); M’Fo, Mofo, Mpuley (dial. Mab Kwasio, Foury 113, Service Forestier du Cameroun 84, Bates 1959); N’jie (Dials. Duala, Punu); Ogowa (Punguegaloa, De Wilde 8492); Moabi jaune (French); évué (dial. Bibaya, Baka).

Least Concern (LC) (Cosiaux et al. 2019a).

medicine : bark as a malaria prophylaxis; construction: house building, furniture; dyes and tannins: as a yellow dye (Versteegh and Sosef 2007).

Annickia affinis is distinguished by having overall glabrous branches and petioles and the lower side of the leaf blades which is sparsely pubescent with simple or bifid hairs pointing in the same direction. Annickia affinis is morphologically close to A. chlorantha from which it is distinguished by having a glabrous upper midrib surface (versus pilose in A. chlorantha). In addition, A. chlorantha has few simple hairs pointing in different directions combined with smaller bifid or trifid hairs.

Annickia affinis is the most common species of Annickia and is generally found as a young plant in secondary forest, or as an adult in older secondary or primary forests. For a long time (and still now) Annickia affinis was confused with A. chlorantha (or even Enantia chlorantha), but the latter name is attributed to a different and rarer species (Versteegh and Sosef 2007). Thus, most literature refers to the old name A. (Enantia) chlorantha when referring to A. affinis (the common and widespread species). Previous reports of A. chlorantha outside Nigeria and Cameroon (e.g. Gabon) refer to A. affinis.

Central Region: near Ebolbom village 3 km est of Ngoumou 2 km north west of Otélé, 3.59°N, 11.28°E, 02 May 2013, Couvreur T.L.P. 426 (WAG,YA); Ottotomo Forest Reserve 3 km after reserve base near small loggers road, 3.66°N, 11.28°E, 02 May 2013, Couvreur T.L.P. 437 (WAG,YA); Mefou Proposed National Park, 3.62°N, 11.57°E, 15 March 2004, Etuge M. 5139 (K,YA); Mbam Minkom, 3.96°N, 11.36°E, 19 September 2013, Kamdem N. 143 (YA); Nguila 1, 4.77°N, 11.75°E, 30 April 2017, Kamdem N. 521 (YA); Colline entre Tcherikoy et Sokelle II (30 km NW Eséka), 3.78°N, 10.96°E, 14 December 1973, Letouzey R. 12361 (P,YA). East Region: 77 km south of Yokadouma 30 km after Ngato 15 km after river ALPICAM ‘base de vie’ then 40 km on forestry road starting 4 km before Maséa village, 3.15°N, 14.72°E, 05 March 2019, Couvreur T.L.P. 1203 (MPU,WAG,YA); Deng Deng, 5.21°N, 13.44°E, 19 April 2016, Kamdem N. 422 (YA); 16 km E de Dimako, 4.38°N, 13.57°E, 15 December 1965, Leeuwenberg A.J.M. 7355 (BR,K,MO,P,PHA,WAG,YA); 15 km E of Dimako, 4.38°N, 13.57°E, 08 February 1966, Leeuwenberg A.J.M. 7787 (BR,C,K,MO,P,WAG,YA); Route Mintom I (70 km E de Djoum)-Alati (100 km SE de Djoum)-PK 63, 2.83°N, 13.35°E, 01 January 1973, Letouzey R. 11751 (P,YA). Littoral Region: Ebo Wildlife Reserve Djuma permanent camp On Djuma-Djuma trail, 4.33°N, 10.24°E, 14 February 2014, Couvreur T.L.P. 621 (WAG,YA); Mambe Massif above Boga village 100 km along road from Yaoundé to Ed 3.90°N, 10.77°E, 20 June 2014, Couvreur T.L.P. 657 (WAG,YA). South Region: Ebolowa, 2.96°N, 11.28°E, 01 January 1925, Bates G.L. 1959 (BM,BR,MO); on road Lolodorf-Bipindi ca half way near Mbiguiligui village (Mbikiliki), 3.16°N, 10.53°E, 26 February 2018, Couvreur T.L.P. 1153 (P,WAG,YA); 22 km east from Lélé village, 3.26°N, 10.10°E, 07 September 2013, Couvreur T.L.P. 469 (WAG,YA); ca 15 km east from Lélé village, 2.26°N, 13.29°E, 09 September 2013, Couvreur T.L.P. 492 (WAG,YA); Campo Ma’an National Park 11 km on trail from Ebinanemeyong village on road 7 km from Nyabessan to Campo town, 2.47°N, 10.33°E, 11 February 2015, Couvreur T.L.P. 671 (WAG,YA); A 6 km à l’ouest de Masea (village situé à 50 km au SSW de Yokadouma), 3.14°N, 14.86°E, 05 July 1963, Letouzey R. 5412 (P,YA); Campo-Ma’an area road Nko-elon-Mvini Akok Beryat rock, 2.36°N, 10.25°E, 30 June 2001, van Andel T.R. 3784 (KRIBI,WAG,YA); Bipindi, 3.08°N, 10.42°E, 01 January 1909, Zenker G.A. 3839 (BM,BR,K,MO,P). South-West Region: Ekundu Kundu, 5.15°N, 8.883°E, 30 April 1996, Cheek M. 8297 (K,WAG,YA); Mungo river forest reserve North of Kumba-Tombel road entered ca 05 km West of Mungo bridge, 4.73°N, 9.55°E, 24 October 1998, Cheek M. 9354 (YA); Foot of Nyale Rock, 4.98°N, 9.616°E, 17 November 1998, Cheek M. 9654 (K,YA); on trail through palm oil plantation 3 km before lava flow and Seme Beach hotel when coming from Limbe, 4.05°N, 9.076°E, 18 October 2013, Couvreur T.L.P. 519 (WAG,YA); Kupe village to Loum State Forest, 4.73°N, 9.716°E, 30 May 1996, Etuge M. 2049 (K,WAG,YA); Nyale forest and rock, 5°N, 9.633°E, 15 February 1998, Etuge M. 4235 (K,YA); Edensueh forest, 5.25°N, 9.576°E, 30 November 2000, Etuge M. 4850 (K); Kumba-Mbonge road 500 m W of Meme River bridge between Bole and Mabonji, 4.55°N, 9.25°E, 07 July 1986, Thomas D.W. 6327 (MO); Baro village, 5.27°N, 9.21°E, 03 March 1988, Thomas D.W. 7494 (K,MO,P,WAG).

Nigeria. Cross River State; Old Calabar, Thomson W.C 130, Dec 1863: holotype: K[K000380204].

Tree, 9–25 m tall, d.b.h. to 5 cm; stilt roots or buttresses absent, slash yellow. Indumentum of simple, bifid trifid, and fasciculate hairs; old leafless branches glabrous, young foliate branches sparsely pubescent to pubescent. Leaves: petiole 2–9 mm long, ca. 2 mm in diameter, pubescent, slightly grooved, blade inserted on top of the petiole; blade 7–28 cm long, 2–9.5 cm wide, elliptic to obovate, apex acuminate, acumen ca. 1 cm long, base narrowly cuneate to shortly attenuate, coriaceous to papyraceous, below densely pubescent when young, sparsely pubescent when old, hairs simple, bifid or trifid hairs pointing in all directions, above glabrous when young and old, concolorous; midrib sunken or flat, above densely pubescent (pilose) to pubescent at least towards base when young, densely pubescent to pubescent at least towards base when old (rarely glabrous), below pubescent when young and old; secondary veins 8 to 12 pairs, glabrous below; tertiary venation intermediate. Individuals bisexual; inflorescences ramiflorous on young foliate branches, leaf opposed or extra axillary. Flowers bisexual with 6 perianth parts in 2 whorls, 1 per inflorescence; pedicel 5–11 mm long, ca. 1 mm in diameter, densely pubescent; in fruit 10–15 mm long, ca. 2 mm in diameter, densely pubescent; bracts 1–2, one basal and one upper towards the middle of pedicel, basal bract 4–8 mm long; sepals 3, valvate, free, 8–12 mm long, 4–6 mm wide, triangular, apex acute, base truncate, green, pubescent outside, glabrous inside, margins flat; petals free; outer petals absent; inner petals 3, valvate, 15–29 mm long, 6–14 mm wide, elliptic to inversely Y–shaped ridged, apex acute, base broad and concave, greenish yellow, margins flat, pubescent outside, pubescent in a small triangle at the base inside; stamens 145 to 160, in 5 to 6 rows, 2 mm long, linear; connective flattened, glabrous, yellow; staminodes absent; carpels free, 20 to 35, ovary ca. 2 mm long, stigma cylindrical, pubescent. Monocarps stipitate, stipes 6–20 mm long, ca. 1 mm in diameter; monocarps 3 to 27, 10–16 mm long, 4–9 mm in diameter, ellipsoid to obovoid, apex mucronate, sparsely pubescent, smooth, glossy, green turning red to black when ripe; seed 1, ca. 20 mm long, ca. 10 mm in diameter, ellipsoid; aril absent.

Known from Nigeria (one collection); in Cameroon known from the East, South, Center and South-West regions.

Locally common when present but rare overall, in lowland and premontane rain forests, mainly in primary habitats. Altitude 150–850 m a.s.l.

Otou han (dial. bulu, Bos 6894), Otoungué (dial. Ewondo, Chevalier 33132).

Least Concern (LC) (Cosiaux et al. 2019b)

medicine : bark as an antisepctic, against fever, malaria prophylaxis; construction: house building, furniture; dyes and tannins: as a yellow dye.

Annickia chlorantha is distinguished by having sparsely pubescent branches and petioles and the lower side of the leaf blades pubescent with simple, bifid, trifid or fasciculate hairs pointing in all directions. The midrib is generally densely pubescent (pilose) at least towards the base, but some specimens are reported to be glabrous (Versteegh and Sosef 2007). See notes under A. affinis for confusions surrounding this name. Annickia chlorantha also closely resembles A. letestui, but differs by its sparsely pubescent young branches versus tomentose in A. letestui.

Central Region: Nkolbisson shrubby low forest on summit of Mt Akockdoué Yaoundé, 3.88°N, 11.45°E, 23 May 1970, Bos J.J. 6894 (BR,MO,P,WAG); Mont Mbam Minkon on trail 3 km from Nkol Nyada village, 3.96°N, 11.40°E, 21 March 2013, Couvreur T.L.P. 414 (WAG,YA); Colline Akok Ndoue près Nkolbisson 5 km WSW Yaoundé, 3.88°N, 11.45°E, 23 May 1970, Farron C. 7335 (P,YA); Yaoundé, 3.86°N, 11.51°E, Feburary 1895, Zenker G.A. 726 (P). Littoral Region: Forêt de Ye Youme, 3.48°N, 12.3°E, 01 June 1917, Chevalier A.J.B. 33132 (P). South-West Region: Bambuko FR, 4.43°N, 9.116°E, 16 September 1951, Olorunfemi J. 30760 (K); Korup National Park, 5.28°N, 9.083°E, 03 April 1988, Thomas D.W. 7555 (MO).

Gabon. Ogooué-Lolo, Ikembélé, Le Testu G.M.P.C. 8432, Oct 1930: lectotype, here designated: P[P00267987]; isolectotypes: BM[BM000547036]; BR[BR0000006418700]; P[P00362651, P02005895, P02005896].

Tree, 2–8 m tall, d.b.h. unknown; stilt roots or buttresses absent, slash yellow. Indumentum of simple, bifid, fasciculate or stellate hairs; old leafless branches glabrous, young foliate branches tomentose. Leaves: petiole 3–8 mm long, 1–2 mm in diameter, tomentose to sparsely pubescent, cylindrical, blade inserted on top of the petiole; blade 10–29.5 cm long, 3.5–10.5 cm wide, elliptic to obovate, apex acuminate to mucronate, acumen 1–2 cm long, base cuneate to rounded to acuminate, subcoriaceous, above glabrous when young and old, below pubescent when young and old, hairs simple, bifid and stellate pointing in all directions, concolorous; midrib sunken or flat, above glabrous when young and old, below pubescent when young and old; secondary veins 9 to 13 pairs, glabrous below; tertiary venation intermediate. Individuals bisexual; inflorescences ramiflorous on young foliate branches, leaf opposed or extra axillary. Flowers with 6 perianth parts in 2 whorls, 1 per inflorescence; pedicel 4–14 mm long, ca. 2 mm in diameter, sparsely pubescent; in fruit ca. 10 mm long, ca. 2 mm in diameter, pubescent; bracts 2, one basal and one upper towards the lower half of pedicel, basal bract 2–4 mm long, 2–3 mm wide; sepals 3, valvate, free, 5–9 mm long, 3–4 mm wide, triangular, apex acute, base truncate, pubescent outside, glabrous inside, margins flat; petals free; outer petals absent; inner petals 3, valvate, 12–26 mm long, 7–12 mm wide, ovate to inversely Y–shaped ridged, apex acute, base broad and concave, yellow–green, margins flat, pubescent outside, pubescent towards margins inside; stamens 60 to 125, in 5 to 6 rows, ca. 2 mm long, linear; connective flattened, glabrous; staminodes absent; carpels free, 20 to 35, ovary ca. 3 mm long, stigma lobed, pubescent. Monocarps stipitate, stipes 8–19 mm long, ca. 1 mm in diameter; monocarps 8 to 20, 19–25 mm long, 10–14 mm in diameter, ellipsoid, apex mucronate, glabrous, smooth, glossy, green turning red to black when ripe; seeds ca. 20 mm long, ca. 10 mm in diameter, ellipsoid; aril absent.

Figure 5. 

Annickia affinis A flowering branch B detail of lower side of leaf blade C flower bud D detail of receptacle with petals removed E inner petals, inner view F stamens, note different shape of connective (discoid to apiculate) G carpel, whole and showing the single basal ovule H floral diagram I fruit. Annickia polycarpa: J details of lower side of leaf blade K flower bud L fruiting branch A–C from Le Testu 1783 D–I from Letouzey 5412 J–L from Chevalier 1611. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from Le Thomas (1969b; pl. 57, p. 311, pro parte).

From Cameroon to Gabon, and one collection in northern Republic of Congo; in Cameroon known from the South region.

A rare species; in lowland rain forests, mainly in primary habitats. Altitude 300–700 m a.s.l.

M’Fo, Mofo, Mpuley (dial. Mab Kwasio, van Andel 4216); N’jie (Dials. Duala, Punu, Bos 4962).

Least Concern (LC) (Cosiaux et al. 2019c).

None recorded.

Annickia letestui is characterized by having tomentose young foliate branches and petioles, the lower side of the leaf blades are pubescent with appressed or erect hairs that are simple, bifid, fasciculate or stellate, pointing in all directions. Versteegh and Sosef (2007) note that the indumentum is quite variable within this species, even within individuals, varying from short and appressed to erect and longer hairs.

Vernacular names are likely to apply to other species of the genus.

South Region: 15 km from Kribi Lolodorf road, 3.00°N, 10.02°E, 01 July 1969, Bos J.J. 4962 (MO,WAG); Mendoum, 2.22°N, 11.23°E, 13 February 1965, Raynal J. 13392 (P); Campo-Ma’an area Bifa, 2.67°N, 10.28°E, 13 October 2001, Tchouto Mbatchou G.P. BIFAX_150 (WAG); Campo-Ma’an area 2.73°N, 9.873°E, 16 August 2001, van Andel T.R. 3882 (WAG); Campo-Ma’an area near Boussebeliga creek bridge, 2.37°N, 9.822°E, 26 October 2001, van Andel T.R. 4216 (WAG).

Figure 6. 

Annickia letestui A flowering branch B detail of lower side of leaf blade C detail of stellate hair D flower bud E inner petal, inside view F stamen G carpel, whole and showing the single basal ovule A–G from Le Testu 8432. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from Le Thomas (1969b; pl. 56, p. 307, pro parte).

Figure 7. 

Annickia affinis A detail of the yellow bark B lower side of leaf, petiole and branch C upper side of leaf, petiole and branch D flower E detail of receptacle F and G fruits. Annickia chlorantha H lower side of leaf, petiole and branch I upper side of leaf, petiole and branch J flowers, side view, one petal removed showing receptacle K flower, top view showing the three sepals in front of the petals A no voucher B–C Couvreur 1123, Gabon D, E Couvreur 469, Lélé, Cameroon F Couvreur 519, Mt Etinde, Cameroon G Couvreur 591, Gabon. Photos Thomas L.P. Couvreur.

Sierra Leone. no region; no locality, Afzelius A. s.n., no date: holotype: B[B 10 0068937]; isotype: BM[BM000547035].

Tree, 2–20 m tall, d.b.h. unknown; stilt roots or buttresses absent, slash yellow. Indumentum of simple, bifid, fasciculate or star hairs; old leafless branches glabrous, young foliate branches densely pubescent to tomentose. Leaves: petiole 3–8 mm long, 2 mm in diameter, densely pubescent to tomentose, grooved, blade inserted on the side of the petiole; blade 5–27 cm long, 2–8 cm wide, elliptic to obovate, apex acuminate, acumen 1–2 cm long, base cuneate to acuminate, coriaceous, below pubescent when young and old, hairs mainly bifid or stellate but some simple too, pointing in all directions, above pubescent when young and old, concolorous; midrib sunken or flat, above pubescent towards base when young, pubescent towards base when old, below pubescent when young, sparsely pubescent when old; secondary veins 8 to13 pairs, pubescent below; tertiary venation intermediate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, leaf opposed or extra axillary. Flowers with 6 perianth parts in 2 whorls, 1 per inflorescence; pedicel 9–19 mm long, ca. 2 mm in diameter, pubescent; in fruit 20 mm long, 4–5 mm in diameter, pubescent; bracts 2, one basal and one upper towards the middle of pedicel, basal bract 8 mm long, 4 mm wide; sepals 3, valvate, free, 9–22 mm long, 4–5 mm wide, triangular, apex acute, base truncate, pubescent outside, glabrous inside, margins flat; petals free; outer petals absent; inner petals 3, valvate, 23–34 mm long, 8–19 mm wide, elliptic to inversely Y–shaped ridged, apex acute, base broad and concave, claw mm long, yellow, margins wavy, densely pubescent outside, glabrous inside; stamens 90 to 200, in 5 to 6 rows, 3–4 mm long, linear; connective flattened, glabrous; staminodes absent; carpels free, ca. 70, ca. ovary 3 mm long, stigma lobed, sparsely pubescent. Monocarps stipitate, stipes 19–59 mm long, 1–2 mm in diameter, monocarps 5 to 55, 18–23 mm long, 8–12 mm in diameter, obovoid, apex mucronate, sparsely pubescent, smooth, glossy, green turning red to black when ripe; seed 1, ca. 20 mm long, ca. 10 mm in diameter, ellipsoid; aril absent.

A mainly West African species, from Sierra Leone to Cameroon; in Cameroon known from the South region.

A rare species in Cameroon, in lowland and pre-montane rain forests mainly in primary habitats. Altitude 110–1400 m a.s.l.

Pola (Mvaï, Fang, Annet 174); African yellow wood, yellow wood (english); Moambe jaune (french).

Least Concern (LC) (Cosiaux et al. 2019d).

medicine : bark as an antisepctic, against fever, malaria; construction: house building, furniture; dyes and tannins: as a yellow dye.

Annickia polycarpa is distinguished by the densely pubescent to tomentose upper side of the midrib and the petioles, and generally long stipes.

South Region: Bipindi, 3.26°N, 10.20°E, 09 June 1928, Annet E. 174 (P).

Annona muricata L.

Trees, 1–10 m tall, d.b.h. 2–10 cm; stilt roots or buttresses absent. Indumentum of simple hairs. Leaves: petiole 7–20 mm long, 1–3 mm in diameter, blade 6–25 cm long, 4–19 cm wide, broadly obovate or obovate to broadly elliptic to elliptic, apex rounded or obtuse or shortly emarginated, base subcordate to rounded, discolorous, whitish below or concolorous; midrib sunken or flat; secondary veins 7 to 16 pairs; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on young and old leafless branches, leaf opposed. Flowers with 9 perianth parts in 3 whorls, 1 to 2 per inflorescence; flowering peduncle sometimes present, short; pedicel 10–25 mm long; in fruit 15–50 mm long; bracts 2, all basal, 1–5 mm long; sepals 3, valvate, free, 3–4 mm long, triangular to ovate, apex acute, base truncate; petals free; outer petals longer than inner; outer petals 3, valvate, 10–15 mm long, 8–10 mm wide, ovate, apex acute, base truncate; inner petals 3, valvate, 8–10 mm long, 3–4 mm wide, narrowly oblong or narrowly elliptic, apex acute to obtuse, base truncate; stamens numerous (not counted), in 2 to 3 rows, 2–3 mm long, linear; connective discoid, shortly pubescent; staminodes absent; carpels free, numerous (not counted), ovary 1–2 mm long, stigma capitate, glabrous or pubescent. Fruit pseudosyncarpous, 20–50 mm long, 20–50 mm in diameter, obovoid to globose; monocarps sessile, completely fused between them, numerous (not counted); seed 1, 8–10 mm long, 4–5 mm in diameter, flattened ellipsoid, irregular in shape; aril absent.

A mainly South American genus, one of the largest in Annonaceae with about 170 accepted species (Rainer 2001). In Africa, there are between three or four native species, with numerous subspecies and varieties and of which the taxonomy remains complicated (Robyns and Ghesquière 1934; Sillans 1952; Le Thomas 1969c). Annona glabra L. is probably of South American origin (Le Thomas 1969b, 1969c) but is naturalized along the coast of West and Central Africa. In Cameroon it is also found in mangrove areas, but is little collected (e.g. van der Burgt 130 (WAG)) We thus include it in the key, but do not provide a description. In addition, this genus contains the non-native edible species Annona squamosa L., A. muricata L. and A. reticulata L. (from South America), all of which can be found in cultivation (not included in the descriptions) in Cameroon.

no recent revision, but see Le Thomas (1969c), Le Thomas (1969b).

Republic of the Congo. Pool; Bord de la M’Boté, Bouquet, A. 513, 12 Sep 1964: holotype: P[P00363247].

Tree to shrub, 1–6(8) m tall, d.b.h. unknown; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches brown tomentose. Leaves: petiole 7–20 mm long, 2–3 mm in diameter, brown tomentose, grooved, blade inserted on top of the petiole; blade 6–20 cm long, 5–12 cm wide, obovate to elliptic, apex rounded or obtuse or shortly emarginate, base rounded to subcordate, papyraceous to coriaceous, below densely pubescent, curly hairs covering the whole leaf blade when young and old, above sparsely pubescent to glabrous when young, glabrous when old, discolorous, whitish below; midrib impressed, above glabrous when young and old, below densely pubescent when young and old; secondary veins 8 to 15 pairs, glabrous above; tertiary venation percurrent but also appearing reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, leaf opposed or extra axillary. Flowers with 9 perianth parts in 3 whorls, 1 to 2 per inflorescence; pedicel 10–25 mm long, 1–2 mm in diameter, brown tomentose; in fruit 15–30 mm long, 3–4 mm in diameter, pubescent; bracts 2, all basal, 2–5 mm long, 2–3 mm wide; sepals 3, valvate, free, 3–4 mm long, 3–4 mm wide, triangular to ovate, apex acute, base truncate, green, densely pubescent outside, glabrous inside, margins flat; petals free, inner smaller than outer; outer petals 3, 10–15 mm long, 8–10 mm wide, ovate, apex acute, base truncate, yellow to green, margins flat, tomentose outside, glabrous inside; inner petals 3, valvate, 8–10 mm long, 3–4 mm wide, narrowly oblong or narrowly elliptic, apex acute, base truncate, yellow-green, margins flat, glabrous outside, glabrous inside; stamens numerous, rows not counted, 2–3 mm long, linear; connective discoid, shortly pubescent; staminodes absent; carpels free, numerous, ovary 1–2 mm long, stigma capitate, pubescent. Fruit pseudosyncarpous, 20–50 mm long, 20–50 mm in diameter, obovoid to globose, yellow orange at maturity; individual monocarps 20 to 30, sessile, completely fused between them; apex shortly pyramidal, brown tomentose, smooth, yellow to orange when ripe; seed 1, 8–10 mm long, 4–5 mm in diameter, flattened ellipsoid; aril absent.

A west and central African subspecies distributed from Guinea to Ivory Coast and from Cameroon to the Democratic Republic of the Congo and the Central African Republic; in Cameroon known from Adamaoua, Central, East, North, North-West, South-West and West regions.

A common species; in lowland savanna regions towards the north, at higher altitudes towards the south, sometimes the dominant tree species in the savanna, reported to naturally invade certain areas (Le Thomas 1969c). Altitude 200–1300 m a.s.l.

None recorded.

Least Concern (LC) (Botanic Gardens Conservation International and IUCN SSC Global Tree Specialist Group 2019a).

None recorded, but probably same as for var. senegalensis.

Subsp. oulotricha is distinguished by the pubescence of the lower surface of leaf blades, which is tomentose with short curly hairs. Besides that, it is very close morphologically to subsp. senegalensis. The species (A. senegalensis) as a whole is very variable morphologically and widespread across the drier parts of sub-Saharan Africa (west to east), also occurring in northern Madagascar (Le Thomas 1969c). Though we have followed the classification of Le Thomas (1969c), the taxonomic limits in this group would need more in-depth studies.

Cheek et al. (2000, p. 114), in the Check list of plants of Mt Oku, reported the presence of A. chrysophylla Bojer (Brunt 234), but this specimen has now been identified as A. senegalensis subsp. oulotricha. Moreover, the former name is now a synonym of A. senegalensis subsp. senegalensis (Le Thomas 1969c).

Robyns and Ghesquière (1934) chose a neotype for the species A. arenaria (now a synonym of A. senegalensis subspecies senegalensis), thinking that no original material seen by Thonning remained (see under that name for details). However, in doing so, they chose a neotype specimen belonging to A. senegalensis subsp. oulotricha (Le Thomas 1969c). When describing this latter species, Le Thomas chose a different type than the one selected by Robyns and Ghesquière (1934) as not to “bring extra confusion to the situation” (Le Thomas 1969c).

Figure 8. 

Annona senegalensis subsp. senegalensis A flowering branch B detail of pubescence on lower side of leaf blade C leaf D detail of pubescence on lower side of leaf blade E longitudinal section of the flower F stamen G carpel, side view and view of the single basal ovule H seed. Annona senegalensis subsp. oulotricha I leaf J detail of pubescence on lower side of leaf blade K fruit branch showing two pseudosyncarpous fruits. Annona glabra L leaf M section of fruit. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from Le Thomas (1969b; pl. 56, p. 321).

Adamaoua Region: Meiganga, 6.52°N, 14.3°E, 06 February 1946, Aubréville A. 729 (P); Tignère, 7.37°N, 12.65°E, 01 March 1939, Jacques-Félix H. 3423 (P); Ngaoundéré, 7.32°N, 13.58°E, 01 June 1939, Jacques-Félix H. 4012 (P); 11 km de Ngaoundéré vers Ngakha, 7.32°N, 13.58°E, 06 March 1958, Letouzey R. 606 (P). Central Region: Mont Ngolep massif Ngoro 38 km N de Bafia, 5.09°N, 11.26°E, 21 April 1975, Ngameni B.K. 51 (P); Mont Ngoro à 58 km SW de Linte, 5.09°N, 11.26°E, 17 April 1982, Nkongmeneck B.A. 256 (P). East Region: Bétaré Oya, 5.5°N, 14.1°E, 02 March 1961, Breteler F.J. 1185 (P); Piste Moyenam Rivière Konbo, 4.58°N, 13.68°E, 29 February 1960, Letouzey R. 3187 (P); Piste Moyenam-rivière Koubou, 4.58°N, 13.68°E, 29 February 1960, Letouzey R. 3191 (P). North Region: Garoua, 9.3°N, 13.4°E, 10 February 1946, Aubréville A. 787 (P); Garoua, 9.3°N, 13.4°E, 11 February 1946, Aubréville A. 804 (P); 17 km N of Banyo along road to Mba, 6.91°N, 11.8°E, 29 February 1972, Leeuwenberg A.J.M. 9440 (WAG). West Region: Dschang, 5.45°N, 9.95°E, 13 April 1966, CNAD 317 (P); Nkounden, 5.7°N, 10.67°E, 01 May 1967, CNAD 808 (P); Bangwa ca. 15 km NW of Baganté, 5.2°N, 10.48°E, 30 April 1964, de Wilde W.J.J.O 2359 (P); Between Bangwa and Bangangté ca 8 km NW of Bangangté, 5.16°N, 10.5°E, 12 May 1964, de Wilde W.J.J.O 2589 (P,WAG); Batchingou, 5.13°N, 10.4°E, 01 January 1939, Jacques-Félix H. 3026 (P); Kontchankap, 5.58°N, 10.80°E, 01 February 1939, Jacques-Félix H. 3033 (P); Foumban, 5.72°N, 10.92°E, 01 February 1939, Jacques-Félix H. 3136 (P).

Senegal: Roussillon 69, 1798: holotype: P[P00363244].

Tree to shrub, 1–10 m tall, d.b.h. 2–10 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches brown tomentose. Leaves: petiole 10–20 mm long, 1–2 mm in diameter, brown tomentose, grooved, blade inserted on top of the petiole; blade 7–25 cm long, 4–19 cm wide, broadly obovate to broadly elliptic, apex rounded, base subcordate, subcoriaceous to coriaceous, below densely pubescent with straight hairs to glabrescent when young and old, above sparsely pubescent to glabrous when young, glabrous when old, discolorous, whitish below; midrib impressed, above glabrous when young and old, below densely pubescent when young and old; secondary veins 7 to 16 pairs, glabrous above; tertiary venation percurrent but also appearing reticulate, dense. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, leaf opposed. Flowers with 9 perianth parts in 3 whorls, 1 to 2 per inflorescence; pedicel 10–20 mm long, 1–2 mm in diameter, brown tomentose; in fruit 25–50 mm long, 3–4 mm in diameter, pubescent; bracts 2, all basal, 1–4 mm long, 2–3 mm wide; sepals 3, valvate, free, 3–4 mm long, 3–4 mm wide, broadly triangular to circular, apex acute, base truncate, green, densely pubescent outside, glabrous inside, margins flat; petals free, outer longer than inner; outer petals 3, 10–15 mm long, 8–9 mm wide, ovate, apex acute, base truncate, yellow to green, margins flat, tomentose outside, glabrous inside; inner petals 3, valvate, 8–10 mm long, 3–4 mm wide, narrowly oblong or narrowly elliptic, apex acute, base truncate, yellow-green, margins flat, glabrous outside, glabrous inside; stamens numerous (not counted), 2–3 mm long, linear; connective discoid, shortly pubescent; staminodes absent; carpels free, numerous (not counted), ovary 1–2 mm long, stigma capitate, glabrous. Fruit pseudosyncarpous; 20–50 mm long, 20–30 mm in diameter, obovoid to globose, yellow orange at maturity, monocarps sessile, numerous, apex flat, glabrous, smooth, yellow to orange when ripe; seed 1, 8–10 mm long, 4–5 mm in diameter, flattened ellipsoid; aril absent.

A west, central and east African and northern Malagasy subspecies from Senegal to Mozambique; in Cameroon known from the Adamaoua, Central, East, Far North, North, North-West, South-West and West regions.

A common species; in lowland savanna regions towards the north, at higher altitudes towards the southern region, may, sometimes be the dominant tree species across the savanna. Altitude 100–1400 m a.s.l.

Falŏ (dial. Bamileke (Burkill 1985)); pomme-cannelle du Sénégal (French); African custard-apple; wild custard apple, wild soursop (English).

food : fruit is eaten, flower for sauces, condiments, spices, flavourings; medicine: root as pain-killer, against diarrh dysentery, cholera, venereal diseases, bark used as vermifuges, diuretics, genital stimulants/depressants, lactation stimulants; construction: house building, furniture; dyes and tannins: astringents, insecticides, arachnicides; products: wood fire; fuel and lighting; social: religion, superstitions, magic.

Annona senegalensis subsp. senegalensis is distinguished by the pubescence of the lower side of the leaf blade which ranges from densely pubescent (but not tomentose) with short but straight hairs (not curly as in subsp. oulotricha) to glabrescent. See notes under subsp. oulotricha and Le Thomas (1969c) for more details. The fruits are edible.

Le Thomas (1969c p. 2) suggested that there exists a specimen of Thonning sent to Jussieu by Vahl in 1804 and present in P (under catalogue number 10779 from Herb. Jussieu) and that this would be the lectotype. However, we were not able to locate this specimen using the online scanned material.

Adamaoua Region: Dodéo, 7.48°N, 12.07°E, 01 March 1939, Jacques-Félix H. 3388 (P); Bountoun Mboun mountains ca 40 km N of Ngaoundere, 7.9°N, 13.48°E, 12 April 1977, Nordal I. 929 (P). Central Region: Bibbanga, 3.72°N, 10.3°E, 09 March 1927, Hédin L. 409 (P). East Region: Bertoua-Batouri, 4.58°N, 13.68°E, 01 January 1962, Vroumsia T. 116 (P). Far-North Region: Douzeye (c Bongor), 10.1°N, 15.28°E, 08 January 1968, Achoundong G. 1385 (P); Plaine de Maroua à 5 km au NO de Maroua, 10.6°N, 14.28°E, 18 August 1964, Biholong M. 28 (P); ca 5 km W of Maroua, 10.6°N, 14.28°E, 02 September 1964, de Wilde W.J.J.O 2966 (MO); Bogo (Maroua), 10.7°N, 14.61°E, 01 May 1939, Jacques-Félix H. 3737 (P); Reserve forestière du Mayo Louti (10 km W de Mokolo), 10.7°N, 13.8°E, 10 September 1964, Letouzey R. 6779 (P); Mora, 11.0°N, 14.14°E, 01 January 1945, Vaillant A. 15 (P). North Region: Pitoa, 9.38°N, 13.50°E, 25 March 1974, Achoundong G. 3419 (P); Garoua, 9.3°N, 13.4°E, 04 August 1955, de Wit H.C.D 7182 (WAG); Ecole de faune de Garoua, 9.3°N, 13.4°E, 09 August 2000, Dong E. 391 (P). North-West Region: Piste Munka (=Munkep) 45 km NNW Wum, 6.8°N, 9.97°E, 09 July 1975, Letouzey R. 13988 (MO). South-West Region: Ndop Plain Hillside above Courtar Ndop Amp ref No 28, 6.02°N, 10.49°E, 01 March 1962, Brunt M.A. 51 (K). West Region: Bangwa, 5.2°N, 10.48°E, 12 May 1964, de Wilde W.J.J.O 2389 (P,WAG).

Anonidium mannii Engl. & Diels.

Trees, 4–30 m tall, d.b.h. up to 80 cm; stilt roots or buttresses absent. Indumentum of simple hairs. Leaves: petiole 3–10 mm long, 2–5 mm in diameter, blade 20–50 cm long, 7–18 cm wide, oblong to obovate, apex rounded or abruptly acuminate, base subcordate forming two small lobes on top of the petiole, concolorous; midrib sunken or flat; secondary veins 10 to 20 pairs; tertiary venation reticulate. Individuals androdioecious or dioecious; inflorescences cauliflorous or ramiflorous on old leafless branches, axillary. Flowers with 9 perianth parts in 3 whorls. 5 to 20 or more per inflorescence; flowering peduncle long, up to 2–4 m, woody, hanging or semi erect; pedicel 10–70 mm long; in fruit 25–100 mm long; bracts 2–4, basal or inserted along the pedicel, 1–5 mm long; sepals 3, valvate, free, 3–4 mm long, triangular to ovate, apex acute, base truncate; petals free; outer petals longer than inner; outer petals 3, valvate, 10–15 mm long, 8–10 mm wide, ovate, apex acute, base truncate; inner petals 3, valvate, 8–10 mm long, 3–4 mm wide, narrowly oblong or narrowly elliptic, apex acute to obtuse, base truncate; stamens 65 to 700, in 2 to 3 rows, 2–3 mm long, linear; connective discoid, shortly pubescent; staminodes absent; carpels free, 180 to 260, ovary 1–2 mm long, stigma capitate, glabrous or pubescent. Fruit pseudosyncarpous, 20–50 mm long, 20–50 mm in diameter, obovoid to globose; monocarps sessile, completely fused, 250 to 500; seed 1, 8–10 mm long, 4–5 mm in diameter, flattened ellipsoid; aril absent.

A genus of trees with four species distributed in Central Africa, one widespread and common across its range (A. mannii) and three others mainly in Gabon (two endemic); in Cameroon two species, none endemic.

Anonidium usambarense R.E.Fr, endemic to Tanzania, is in fact a Polyceratocarpus species (probably Polyceratocarpus scheffleri Engl. & Diels, Couvreur, pers. obs.). Finally, one study (Focho et al. 2010) reports the presence of Anonidium floribundum Pellegr. in the Mount Cameroon area. However, no herbarium collection is available to confirm this and we do not consider it present in Cameroon for now.

No recent revision has yet been published, but see Le Thomas (1969b) were most species are treated for Gabon.

Democratic Republic of the Congo. Bas-Congo; Ganda-Sundi, de Briey J. 86, 1911: lectotype, here designated: BR[BR8822635]; isolectotype: BR[BR8822642].

Tree, 15–25 m tall, d.b.h. up to 35 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches sparsely pubescent to glabrous. Leaves: petiole 3–10 mm long, 2–5 mm in diameter, sparsely pubescent, soon becoming glabrous, slightly grooved, blade inserted on top of the petiole; blade 37–50 cm long, 10–16 cm wide, oblong, apex rounded to abruptly acuminate, acumen 2–3.5 cm long, base rounded, subcordate, subcoriaceous to coriaceous, below sparsely pubescent when young, glabrous when old, above glabrous when young and old, concolorous; midrib impressed, above glabrous when young and old, below sparsely pubescent when young, glabrous when old; secondary veins 14 to 16 pairs, glabrous above; tertiary venation reticulate. Individuals dioecious; inflorescences cauliflorous or on leafless branches, axillary; peduncle 50–115 mm long, 3–10 mm in diameter, woody, hanging or semi erect from the trunk, glabrous. Flowers with 9 perianth parts in 3 whorls, 1 to 10 per inflorescence, male and female inflorescences similar; pedicel 10–35 mm long, 2–4 mm in diameter, glabrous; bracts, 1 to 2 basal and one upper towards the upper of pedicel, basal bracts 5–10 mm long, 5–10 mm wide; upper bract directly under the calyx, amplexicaul, 12–21 mm long, 10–20 mm wide; sepals 3, valvate, free, 20–40 mm long, 14–18 mm wide, triangular, apex long acuminate, gradually tapering into an acute apex, base truncate, green, densely pubescent outside, glabrous inside, margins flat; petals free, sub equal; outer petals 3, 35–60 mm long, 10–18 mm wide, narrowly elliptic to obovate, apex acute to attenuate, base attenuate, green turning yellow, margins flat, pubescent outside, pubescent inside; inner petals 3, valvate, 30–50 mm long, 9–16 mm wide, narrowly elliptic, apex acute, base truncate, green turning yellow, margins flat, pubescent outside, pubescent inside; stamens in male flowers: 500 to 700 inserted on a conical receptacle, in 14 to 17 rows, 3.5–5 mm long, oblong, connective truncate, sparsely pubescent, green to cream-yellow; carpels in female flower (see notes) 220 to 260, ovary 2–3 mm long, stigma capitate, glabrous; staminodes absent. Whole fruits not seen, label information: ca. 20 cm long, ca. 10 cm in diameter [taken from descriptions on specimens Hallé & Villiers 4505, P02032580], white [taken from Cheek 10240 P00956188]; seed 1, 40–48 mm long, 17–23 mm wide, ellipsoid; aril absent.

A central African species, known from Cameroon, Gabon, Republic of the Congo and Angola (Cabinda); in Cameroon known from the Littoral and South-West regions.

An uncommon species, in lowland primary or old secondary rain forests. Altitude 100–350 m a.s.l.

None recorded.

Not evaluated.

None recorded.

Anonidium brieyi differs from A. mannii by its usually larger leaves and the narrower inner (10–18 mm wide versus 15–25 mm in A. mannii), and the upper bract inserted just under the calyx (versus near the middle of the pedicel in A. mannii). Three collections seen have strictly female flowers (Couvreur 1132, WAG; Hallé & Villiers 4505, [P02032580]; Sita 712 [MPU411091). The first author’s field observations failed to see any stamens in the thus apparently female flowers (Fig. 10J). A note by Le Testu (Le Testu 1641 [P02032574]) also indicates that the species is dioecious. This suggests that A. brieyi is a strictly dioecious species (male and female flowers on different individuals) rather than androdioecious as in the other species of Anonidium (Le Thomas 1969b). This is the first time this is suggested to occur in this species as only male flowers were described to date. Anonidium letestui Pellegr. (endemic to Gabon) has functionally female flowers, but has a small row of sterile stamens at the base (Le Thomas 1969b), something not observed in the two female specimens examined in A. brieyi (Fig. 10J). Variation of sexual systems within genera (e.g. androdioecious and dioecious species) has been reported in African Annonaceae such as Monanthotaxis (Hoekstra et al. 2021) or Uvariopsis (Couvreur and Luke 2010).

Figure 9. 

Anonidium mannii A leaf B male inflorescence, note upper bract inserted towards lower part of pedicel C receptacle of male flower, all petals removed D stamen E receptacle of bisexual flower, note small row of stamens at base of carpels F longitudinal section of bisexual flower G stamen of bisexual flower, front view H two free carpels, showing basal ovules I fruit, note syncarpous nature, referred to as pseudosycarpous J seed, side view. Anonidium brieyi K flower, note narrow petals A from Le Testu 9169 B–D from Le Testu 9509 E–H from Le Testu 7269 I from Nigerian tree photo J from Le Testu s.n. K from Le Testu 1641. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from Le Thomas (1969b; pl. 60, p. 331).

Figure 10. 

Anonidium manni A habit, A. manni is the species with white trunk in the center B base of leaf blade, upper side C pseudosyncarpous fruit, note hand for size D Narcisse Kamdem holding an inflorescence E detail of flower, side view F detail of receptacle of bisexual flower, note 2 to 3 rows of stamens at base. Anonidium brieyi G trunk with semi erect inflorescences H leaf, upper side I flower, side view, note narrow petals J detail of receptacle in female flower, note absence of stamens at base of carpels K flower, top view showing insertion of upper bract just belong the calyx A no voucher, Gabon B, C Couvreur 449, Otottomo, Cameroon D, F Couvreur 696, Campo, Cameroon E Couvreur 1207, Maséa, Cameroon G–K Couvreur 1132, Gabon. Photos Thomas L.P. Couvreur.

Le Thomas (1969b) recognized this species as a variety of A. mannii (A. mannii var. brieyi (De Wild.) Fries), however, the morphological differences in addition to its dioecious nature described above warrant it to be retained at the species level for now. Nevertheless, a phylogeographic study of A. mannii based on hundreds of nuclear markers inferred a nested position of A. brieyi (population collected around reference number Couvreur 1132, here identified as A. brieyi) within A. mannii (Helmstetter et al. 2020). This sheds further doubt on the delimitation between these two species and needs further study.

Narrowly elliptic petals are also found in the Gabonese species A. floribundum Pellegr. (not recorded for Cameroon to date). However, this latter species is a small tree up to 4 or 5 m tall, the inflorescences are shorter with fewer flowers, and it is androdioecious (Le Thomas 1969b).

Two herbarium specimens (Cheek 10240, Cameroon; Hallé & Villiers 4505, Gabon) indicate that the collection had fruits, but we were unable to locate them. Cheek 10240 states the fruits are white. Collection Cheek 12040 was identified as A. mannii in the check list of the plants of Mt Kupe (Cheek et al. 2004, p. 236). Four of the type sheets of the name A. friesianum have seeds (Gossweiler 6690, LISC000055, LISC000058, LISC000060, LISC000063). On sheets LISC000055 and LISC000061 there are large narrow structures (18 cm by 3.5 cm) which we cannot identify, either being a dried fruit or part of the bark (?).

The sheet Cheek 10240 from P (P00956188) is a mixed collection. The leaves belong to a species of Uvariopsis (probably connivens), and not A. brieyi. The flowers however are from A. brieyi. The other sheets we have seen appear to have the correct leaves (K, WAG [WAG.1378838])

Finally, for the type of Anonidium friesianum, among the 10 sheets of Gossweiler 6690 available at we did not designate one sheet as lectotype because every sheet has important information (leaves, seeds, leaves) and should be considered as a single collection. Among the 10 sheets of Gossweiler 6690 available at LISC for the type of Anonidium friesianum, we selected a single sheet as the lectotype (LISC000056). Indeed, according to the ICBN, different sheets are considered as a single specimen only if they are cross-referenced (e.g. “Sheet I”, “Sheet II”) which is not the case here. The selected sheet (LISC000056) was the only one that had a leaf and a flower (but broken) thus being the most complete. The other sheets only had either just leaves, or inflorescences or flowers or seeds.

Littoral Region: Loum Forest Reserve, 4.73°N, 9.731°E, 16 April 2005, Onana J.M. 3101 (K). South-West Region: Banga, 4.55°N, 9.416°E, 01 March 1956, Binuyo A. 35606 (FHO); Southern Bakundu FR, 4.48°N, 9.350°E, 13 March 1948, Brenan J.P.M. 9410 (K,P); Just west of Loum, 4.73°N, 9.729°E, 03 December 1999, Cheek M. 10240 (K,MO,P,WAG,YA); S Bakundu FR, 4.49°N, 9.374°E, 09 April 1951, Olorunfemi J. 30508 (K); 5 km S of Kumba on Buea/Douala Road, 4.65°N, 9.39°E, 21 June 1983, Thomas D.W. 2188 (MO,P,WAG).

Nigeria. Cross River State; Old Calabar, Mann G. 2231, 1863: lectotype, sheet here designated: K[K000198881]; isotypes: B[B100153003]; K[K000198882]; P[P00363249].

Tree, 8–30 m tall, d.b.h. 30–80 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches sparsely pubescent to glabrous. Leaves: petiole 3–7 mm long, 2–3 mm in diameter, sparsely pubescent to glabrous, slightly grooved, blade inserted on top of the petiole; blade 20–45 cm long, 7–18 cm wide, obovate to oblong-elliptic, apex rounded to abruptly acuminate, acumen 2–3 cm long, base rounded to shortly attenuate generally covering part of the petiole, subcoriaceous to coriaceous, below sparsely pubescent when young, glabrous when old, above glabrous when young and old, concolorous; midrib impressed, above glabrous when young and old, below sparsely pubescent when young, glabrous when old; secondary veins 10 to 20 pairs, glabrous above; tertiary venation reticulate, indistinct. Individuals androdioecious; inflorescences cauliflorous or on old leafless branches, axillary; peduncle 0.3–4 m long, up to 2–3 cm in diameter towards the base, woody, hanging, sparsely pubescent to glabrous. Flowers with 9 perianth parts in 3 whorls, 10 to 20 per inflorescence, male and female inflorescences similar; pedicel 10–85 mm long, 2–4 mm in diameter, glabrous; in fruit 25–100 mm long, 10–20 mm in diameter, woody, glabrous; bracts 2–4, several basal and one upper towards the middle of pedicel, basal bracts 7–12 mm long, 7–12 mm wide; upper bract 10–12 mm long, 12–25 mm wide, clasping the pedicel; sepals 3, valvate, basally fused, 15–22 mm long, 17–20 mm wide, triangular to ovate, apex acute, base truncate, green, tomentose outside, glabrous inside, margins flat; petals free, sub equal; outer petals 3, 25–50 mm long, 20–40 mm wide, elliptic to obovate, apex acute or rounded, base truncate, yellow to green, margins flat, pubescent outside, pubescent inside; inner petals 3, valvate, 12–35 mm long, 15–25 mm wide, elliptic, apex rounded, base attenuate to truncate, yellow-green, margins flat, pubescent outside, pubescent inside; stamens in male flowers: numerous on a conical receptacle, 4–4.5 mm long, oblong, connective truncate, sparsely pubescent, green to cream–yellow; stamens in hermaphrodite flowers: 65 to 100, in 3–4 rows, 2–2.5 mm long, oblong; connective truncate, sparsely pubescent, green to cream - yellow; staminodes absent; carpels free, 180–210, ovary 3–4 mm long, stigma capitate, glabrous. Fruit pseudosyncarpous, 250–500 mm long, 100–300 mm in diameter; individual monocarps 150 to 180, sessile, completely fused between them; apex shortly pyramidal, glabrous, smooth, glossy, yellow at maturity with white pulp; seed 1, 30–50 mm long, 17–30 mm in diameter, flattened to oblong; aril absent.

A central African species, from southeastern Nigeria to the Republic of Congo, and in the Democratic Republic of the Congo; in Cameroon known from the East, South, Central, Littoral, South-West and West regions.

A widespread and very common species across its range; in evergreen or semi-deciduous primary, old or young secondary, lowland or premontane rain forests. Altitude: 0–1600 m a.s.l.

mbé, nbwé, ombé (dial. Bagali, Baka), ébom, ében, ébon ntangan (dial. Ewondo, Hochuli 4).

Least Concern (LC) (Harvey-Brown 2019a).

Food : fruit is eaten; medicine: bark used against arthritis, rheumatism, stomach troubles, diarrho dysentery, menstrual cycle, antidotes, paralysis, epilepsy, convulsions, spasms; social: religion, superstitions, magic.

Anonidium mannii is easily recognizable by its characteristic large leaf blades with a slightly cordate base covering part of the petiole, its long and woody inflorescences hanging from the trunk or old leafless branches and its syncarpous fruits up to 50 cm long and 30 cm large. It is morphologically close to A. brieyi but differs mainly by the wider inner and outer petals, and slightly smaller leaf blades and individuals being androdioecious (see notes under A. brieyi).

Central Region: Eastern sector of M’fou Nat Park Footpath running E of bridge SE of Ndanan 1, 3.61°N, 11.58°E, 22 October 2002, Cheek M. 11248 (K,YA); near Otele, 3.43°N, 11.14°E, 25 February 2007, Couvreur T.L.P. 106 (WAG,YA); Mont Mbam Minkon on trail 3 km from Nkol Nyada village, 3.96°N, 11.40°E, 21 March 2013, Couvreur T.L.P. 415 (WAG,YA); Ottotomo Forest Reserve on top of small hill in front of reserve house, 3.65°N, 11.29°E, 25 June 2013, Couvreur T.L.P. 452 (WAG,YA); on trail to Oveng Lodge hotel near parking just behind the village of Oveng 30 km on road from Mbalmayo to Sangmeli 3.41°N, 11.70°E, 09 February 2014, Couvreur T.L.P. 609 (WAG,YA); Ottotomo Forest reserve 7 km north-west from Ngoumou 30 km south west from Yaoundé, 3.65°N, 11.28°E, 24 February 2016, Couvreur T.L.P. 987 (WAG,YA); SSW of M’Balmayo, 3.52°N, 11.5°E, 27 February 1964, de Wilde W.J.J.O 1968 (B,B,BR,K,MO,P,WAG,YA); Mbam-Minkom Village de Nkolniada, 3.96°N, 11.40°E, 26 July 2012, Droissart V. 1416 (MO); Nanga Eboko, 4.68°N, 12.36°E, 17 February 1927, Hédin L. 36 (P); Mbam Minkom, 3.96°N, 11.36°E, 19 September 2013, Kamdem N. 142 (YA); Nkila, 4.68°N, 12.37°E, 12 March 1959, Letouzey R. 1547 (P,YA); Ekom, 3.85°N, 11.7°E, 16 February 1947, Letouzey R. 194 (P). East Region: Toungrélo, 4.33°N, 13.53°E, 09 January 1962, Breteler F.J. 2454 (K,P,WAG); 81 km south of Yokadouma 30 km after Ngato 15 km after river ALPICAM ‘base de vie’ then 40 km on forestry road starting 4 km before Maséa village, 3.17°N, 14.69°E, 05 March 2019, Couvreur T.L.P. 1207 (MPU,WAG,YA); Somalomo, 3.32°N, 12.71°E, 18 March 2016, Kamdem N. 415 (YA); Lomié, 3.11°N, 13.58°E, 01 December 2016, Kamdem N. 454 (YA). Littoral Region: Nkam, 4.35°N, 10.67°E, 13 June 1927, Hédin L. 1337 (P). South Region: Campo Ma’an National Park 11 km on trail from Ebinanemeyong village on road 7 km from Nyabessan to Campo town, 2.48°N, 10.33°E, 13 February 2015, Couvreur T.L.P. 696 (WAG,YA); 17 km On the newly reconstructed road from Ebolowa to Minkok, 2.75°N, 11.25°E, 29 January 1975, de Wilde J.J.F.E 7930 (BR,K,MO,P,U,WAG,YA); Massif de Ngovayang village de Atog Boga, 3.25°N, 10.49°E, 05 September 2015, Droissart V. 2158 (BRLU); Essam, 4.68°N, 12.37°E, 13 February 1959, Letouzey R. 1276 (P); Essam (Nanga Eboko), 4.68°N, 12.37°E, 14 February 1959, Letouzey R. 1385 (P); Campo-Ma’an area Nsengou, 2.18°N, 10.58°E, 05 February 2001, Tchouto Mbatchou G.P. 3129 (KRIBI,WAG). South-West Region: Colline de Bokwa 42 km SE Mamfe, 5.71°N, 9.643°E, 07 December 1986, Achoundong G. 1325 (YA); Nyasoso, 4.86°N, 9.7°E, 03 June 1996, Cable S. 2801 (K,YA); Nature trail, 4.81°N, 9.683°E, 15 January 1995, Cheek M. 6009 (K,WAG); Kupe Mount, 4.82°N, 9.683°E, 20 November 1995, Cheek M. 7896 (K,WAG,YA); Nyasoso village at base of My Kupe forest reserve along nature trail, 4.82°N, 9.686°E, 04 April 2016, Couvreur T.L.P. 1053b (WAG,YA); Cameroon Mountain, 4.08°N, 9.1°E, 29 December 1983, Thomas D.W. 2850 (MO,WAG). West Region: Près Bandounga à 40 km au NW de Ndikinimeki, 4.98°N, 10.55°E, 12 February 1972, Letouzey R. 11202 (P,YA).

Lianas, up to 30 m tall, d.b.h. up to 20 cm; stilt roots or buttresses absent. Indumentum of simple hairs or absent. Leaves: petiole 1–15 mm long, 1–2 mm in diameter; blade 7–26 cm long, 2.5–14 cm wide, elliptic to ovate to obovate to oblong, apex acuminate to acute, base decurrent to subcordate, concolorous; midrib sunken or flat; secondary veins 7 to 16 pairs; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, leaf opposed or extra axillary. Flowers with 9 perianth parts in 3 whorls, 1 to 90 per inflorescence; pedicel 2–25 mm long; in fruit 2–25 mm long; bracts 2, all basal, minute, soon falling; sepals 3, valvate, free, 1–15 mm long, triangular, apex acute, base truncate; petals free, sub equal; outer petals 3, valvate, 5–35 mm long, 1–14 mm wide, ovate to elliptic to linear to tubular, apex acute to rounded, base broad and concave; inner petals 3, valvate, 5–30 mm long, 1–9 mm wide, ovate to elliptic to linear to tubular, apex acute to rounded, base broad and concave, forming a pollination chamber over the receptacle; stamens 15 to 70, in 2 to 5 rows, 2–3 mm long, linear or cuneiform; connective discoid, glabrous or pubescent; staminodes absent; carpels free, 3 to 32, ovary 1–4 mm long, stigma bilobed or cylindrical, pubescent or glabrous. Monocarps sessile or substipitate, stipe, when present 1–25 mm long, 1 to 20 monocarps, 6–60 mm long, 5–25 mm in diameter, ellipsoid to obovoid, apex rounded to apiculate, smooth or verrucose; seed 1 to 2, 5–25 mm long, 5–15 mm in diameter, ellipsoid or flattened ellipsoid; aril absent.

Artabotrys odoratissimus R.Br., nom. illegit. (≡ Annona hexapetala L.f., ≡ Artabotrys hexapetalus (L.f.) Bhandari).

A genus of lianas with around 105 species distributed across the paleotropics in South East Asia, Australia, Madagascar and Africa (Chen et al. 2019); eight species occur in Cameroon, one endemic.

Genus easily identifiable by its lianescent habit with the presence of characteristic inflorescences in form of a hook (the peduncle) and flowers that have a broad and concave base.

To date there are no taxonomic revisions for Artabotrys in Africa, but see Le Thomas (1969b), Boutique (1951b) and Paiva (1966).

Cameroon. Central Region; Yaoundé, Zenker G.A. 690, 1896: holotype: B[B 10 0153007]; isotypes: BM[BM000546848]; COI[COI00004927]; P[P00363375, P00363376]; K[K000198859, K000198860].

Liana, height unknown, d.b.h. unknown. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches sparsely pubescent with short appressed hairs. Leaves: petiole 3–5 mm long, 1–2 mm in diameter, pubescent with short appressed hairs to glabrous, grooved, blade inserted on the side of the petiole; blade 7.5–10 cm long, 2.5–6 cm wide, oblong to elliptic, apex acuminate, acumen 0.5–1 cm long, base cuneate to rounded, coriaceous to subcoriaceous, below sparsely pubescent with short appressed hairs to glabrous when young, glabrous when old, above glabrous when young and old, concolorous; midrib sunken or flat, above glabrous when young and old, below pubescent with short appressed hairs when young and old; secondary veins 10 to 12 pairs, glabrous above; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless branches, leaf opposed. Flowers with 9 perianth parts in 3 whorls, 1 to 4 per inflorescence; hook-shaped peduncle 15–20 mm long, sparsely pubescent; pedicel 7–10 mm long, 1–2 mm in diameter, sparsely pubescent; in fruit 7–13 mm long, 2–3 mm in diameter, glabrous; bracts all basal, minute; sepals 3, valvate, free, 3–5 mm long, 2–3 mm wide, triangular, apex acute, base truncate, green, pubescent outside, glabrous inside, margins flat; petals free, sub equal, green turning red-orange; outer petals 3, 15–30 mm long, 4–9 mm wide, narrowly elliptic to narrowly ovate, apex attenuate, base broad and concave, white to light green, margins flat, tomentose outside, tomentose with a glabrous base inside; inner petals 3, valvate, 15–30 mm long, 3–6 mm wide, linear to narrowly elliptic, apex acute, base broad and concave, white to light green, margins flat, tomentose outside, tomentose with a glabrous base inside; stamens 15 to 20, in 2 to 3 rows, 2–3 mm long, cuneiform; connective discoid, glabrous; staminodes absent; carpels free, 8 to 10, ovary ca. 2 mm long, stigma cylindrical, glabrous. Monocarps sessile, 4 to 6, 15–20 mm long, 7–13 mm in diameter, ellipsoid to oblong, apex rounded, glabrous, smooth, red when ripe, not ribbed; seeds 1 to 2 per monocarp, 10–13 mm long, 5–8 mm in diameter, flattened ellipsoid; aril absent.

A central African species, from Cameroon to the Republic of Congo and the Democratic Republic of the Congo; in Cameroon known from the Central, East, Littoral, South, South-West and West regions.

A common species across its range; in sub montane (sometimes lowland) secondary or primary rain forests. Altitude (100)500–1600 m a.s.l.

None recorded.

Not evaluated.

None recorded.

Artabotrys aurantiacus is distinguished by its sparsely pubescent to glabrous branches and leaf blades (upper and lower sides), with leaves that are relatively small but wide (less than 10 cm and up to 6 cm wide) and oblong to elliptic in shape with a cuneate to rounded base. The flowers have narrowly elliptic petals and the monocarps are smooth, ellipsoid with a rounded apex.

Central Region: Badjob, 3.68°N, 10.68°E, 21 December 1963, de Wilde W.J.J.O 1602 (BR,MO,P,WAG,YA); Bank Nyong River near the new bridge ca 65 km SSW of Eséka, 3.46°N, 10.5°E, 17 June 1964, de Wilde W.J.J.O 2720 (WAG); Yaoundé, 3.87°N, 11.52°E, 1896, Zenker G.A. 690 (B,K,P). East Region: Ebaka (Bertoua), 4.93°N, 13.32°E, 24 May 1961, Breteler F.J. 1429 (BR,K,M,P,WAG,YA); Ndo Riv (Bertoua), 4.58°N, 13.68°E, 12 December 1961, Breteler F.J. 2210 (BR,K,P,WAG,YA); Doumé Riv (Batouri), 4.23°N, 13.45°E, 15 April 1962, Breteler F.J. 2799 (K,P,WAG); Goyoum, 5.22°N, 13.38°E, 29 January 1961, Breteler F.J. 968 (A,BR,K,M,P,WAG); Nguélémendouka, 4.38°N, 12.92°E, 04 April 1962, de Bruijn J. s.n. (WAG[WAG0175010]); Rives du Dja près Ndongo à 40 km WNW de Moloundou, 2.15°N, 14.86°E, 18 March 1973, Letouzey R. 12141 (P,WAG,YA); Berge arbustive et broussailleuse du fleuve Sanaga au Nord de Goyoum, 5.24°N, 13.36°E, 29 January 1961, Letouzey R. 3309 (P,YA); Rives du Dja entre les rivières Meu et Edjune, 3.41°N, 13.33°E, 12 April 1961, Letouzey R. 3772 (P,YA); Rives de la Kadei entre Mindourou et Dongongo (40 km SSE de Batouri), 4.13°N, 14.60°E, 25 April 1962, Letouzey R. 4859 (P,YA); Betare Oya, 5.59°N, 14.08°E, Tisserant C. 3651 (P). Littoral Region: Manengouba mount base 4 km WNW Of Nkongsamba, 4.96°N, 9.883°E, 09 September 1971, Leeuwenberg A.J.M. 8319 (B,BR,MO,P,U,WAG,YA). South Region: Ebom, 3.1°N, 10.73°E, 13 August 1996, Elad M. 510 (WAG); Nyabesan, 2.4°N, 10.4°E, 05 March 1963, Raynal J. 10240 (P,YA). South-West Region: Likombe, 4.11°N, 9.183°E, 19 February 1995, Cable S. 1309 (K,MO,WAG,YA); Likombe, 4.11°N, 9.183°E, 19 February 1995, Cable S. 1310 (K,WAG,YA); Road to NLO Mt from Kodmin, 5°N, 9.683°E, 23 January 1998, Cheek M. 9063 (K,YA); Nyasoso, 4.81°N, 9.683°E, 08 February 1995, Elad M. 132 (K,YA); Mt Cameroon south slope Transect 8, 4.07°N, 9.015°E, 16 November 1985, Gentry A.H. 52942 (MO,P); Nzee Mbeng trail from Ngomin to Nzee Mbeng, 5.83°N, 9.716°E, 10 February 1998, Gosline W.G. 99 (K,YA); Ndum, 4.83°N, 9.7°E, 31 January 1995, Groves M. 21 (K,MO,WAG,YA); Nyasoso, 4.81°N, 9.683°E, 08 February 1995, Groves M. 77 (K,YA); South slope of mount north of Mt Etinde Forest, 4.08°N, 9.133°E, 20 March 1988, Nemba J. 953 (MO,P). West Region: Bali Ngemba grassland and forest patches northeast of Mantum, 5.82°N, 10.08°E, 12 April 2004, Etuge M. 5373 (K,MO,P,WAG,YA); Dschang, 5.45°N, 9.95°E, 01 May 1960, Jacques-Félix H. 5211 (K,P,WAG); Ngwenfon 35 km NW de Foumban, 5.72°N, 10.92°E, 11 December 1974, Letouzey R. 13495 (P,YA).

Figure 11. 

Artabotrys aurantiacus var. aurantiacus A flowering branch B outer petal, inner view C inner petal, outer and inner views D flowering receptacle with petals removed E stamen F carpel, side view and detail of basal ovules G fruiting branch. Var. multiflorus H flowering branch, note the numerous flowers. Artabotrys insignis var. insignis I flowering branch J outer petal, inner view K inner petal, inner view L flower, whole M stamen N carpel, side view, and detail of ovules O fruiting branch, note long apiculate apex of monocarps A–F from Le Testu 8499 G from Le Testu 4430 H from Le Testu 7116 I–N from Le Testu 8674; 15 from Berteler 2959. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from Le Thomas (1969b; pl. 26, p. 143).

Gabon. Ogooué-Lolo; Lastoursville, Le Testu G.M.P.C. 7116, Mar 1929: lectotype, sheet here designated: P[P02034091]; isotypes: BR[BR0000008820792, BR0000008809940, BR0000008809933]; IFAN[IFAN01625]; LISC[LISC000367]; P[P02034088]; K[K000198858].

Differs from the type variety by the presence of numerous densely packed flowers (6–15 versus 1–4).

A central African species, from Cameroon to the Republic of Congo and the Democratic Republic of the Congo, the multiflorus variety is known from Cameroon and Gabon; in Cameroon known from the Central, East, Littoral, South-West and West regions.

A rare variety; in sub montane (sometimes lowland) secondary or primary rain forests. Altitude 700–1600 m a.s.l.

None recorded.

Not evaluated.

None recorded.

Map 2. 

A Artabotrys aurantiacus B Artabotrys aurantiacus var. multiflorus C Artabotrys congolensis D Artabotrys dielsianus E Artabotrys insignis F Artabotrys insignis var. batesii G Artabotrys jacques-félicis H Artabotrys rufus I Artabotrys thomsonii. White borders represent region limits in Cameroon; green patches represent protected areas (see methods and Suppl. material 1: Fig. S1).

The variety status is doubtful, but without further investigation, we shall follow Le Thomas (1969b). The main difference between the two varieties is the number of flowers per inflorescence (see above). Le Thomas (1969b) also mentions that var. multiflorus has smaller flowers (smaller than 13 mm) and pedicels of 1 cm long. However, our measurements were not able to confirm these two latter differences.

Central Region: Près Yaoundé, 3.86°N, 11.45°E, 11 March 1981, Meijer D. 15033 (MO,WAG). West Region: Bali- Ngemba FR, 5.81°N, 10.08°E, 13 April 2002, Onana J.M. 2027 (K,WAG,YA).

Democratic Republic of the Congo. Equateur; Lukolela, Dewèvre A.P. 819, 31 Mar 1896: holotype: BR[BR0000008820808]; isotype: B[B 10 0153012].

Liana, height unknown, d.b.h. ca. 8 cm. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 10–15 mm long, ca. 2 mm in diameter, glabrous, grooved, blade inserted on the side of the petiole; blade 8–26 cm long, 4–11 cm wide, elliptic to obovate, apex acute to acuminate, acumen 0.5–1 cm long, base acute to decurrent, coriaceous, below glabrous when young and old, above glabrous when young and old, concolorous; midrib impressed, above glabrous when young and old, below glabrous when young and old; secondary veins 8 to 13 pairs, glabrous above; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless branches, leaf opposed. Flowers with 9 perianth parts in 3 whorls, 1 to 3 per inflorescence, hook-shaped peduncle 10–15 mm long; pedicel 10–25 mm long, 2–3 mm in diameter, glabrous; in fruit 12–20 mm long, 2–3 mm in diameter, glabrous; bracts 1 to 2, all basal, basal bracts 2–3 mm long, 1–2 mm wide; sepals 3, valvate, free, 5–8 mm long, 3–7 mm wide, triangular, apex acute, base truncate, pubescent outside, glabrous inside, margins flat; petals free, inner shorter than outer, green turning yellow; outer petals 3, 15–25 mm long, 8–14 mm wide, ovate, apex acute, base broad and concave, margins flat, tomentose outside, tomentose with a glabrous base inside; inner petals 3, valvate, 8–17 mm long, 5–8 mm wide, elliptic to rhombic, apex acute, base broad and concave forming a chamber over the receptacle, margins flat, tomentose outside, tomentose with a glabrous base inside; stamens numerous, number of rows unknown, 2–3 mm long, oblong; connective discoid, glabrous; staminodes absent; carpels free, 15 to 20, ovary 3–4 mm long, stigma tubular, sparsely pubescent. Monocarps stipitate, stipes 1–2 mm long, 1–3 mm in diameter, but gradually widening into seed-bearing part; monocarps 15 to 20, to 60 mm long, ca. 25 mm in diameter, obovoid, apex acute to rounded, glabrous, smooth, glossy, green when ripe; seeds 1 to 2 per monocarp, 19–22 mm long, 10–15 mm in diameter, ellipsoid, laterally flattened; aril absent.

Figure 12. 

Artabotrys congolensis A flowering branch B sepals outer and inner views C outer petals, outer and inner views D side view of outer and inner petals showing the pollination chamber formed by the inner petals E inner petals, outer and inner views F stamen G fruit, notice the stipes gradually widening into the seed bearing part of the monocarps H longitudinal section of a monocarp showing 2 seeds A–F from Tisserant 2242 G, H from Hallé 3451. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from Le Thomas (1967c).

A central African species, from Cameroon to the Republic of Congo and in the Democratic Republic of the Congo; in Cameroon known from the Central, East, Littoral, South and South-West regions.

A common species; in lowland or premontane secondary or primary rain forests. Altitude 400–1000 m a.s.l.

None recorded.

Not evaluated.

None recorded.

Artabotrys congolensis is distinguished by its completely glabrous branches, petioles and leaf blades, its outer petals that are broad and the inner petals much narrower and its sessile fruits with a stipes gradually widening into the seed-bearing part of the monocarp.

Le Thomas described Artabotrys rhopalocarpus (Le Thomas 1967c) which was later synonymized with A. congolensis (Bamps and Le Thomas 1989).

Central Region: near Ebolbom village 4 km est of Ngoumou 3 km north west of Otélé, 3.60°N, 11.28°E, 02 May 2013, Couvreur T.L.P. 431 (MPU,WAG,YA). East Region: Sur axe Lomié-Ngoila-Souanké à 15 km au SSW de Ngola, 2.51°N, 13.86°E, 22 February 1973, Letouzey R. 12016 (P,YA). Littoral Region: Tissongo, 3.57°N, 9.869°E, 09 July 1976, McKey D.B. 111 (K). South Region: Ebolowa-Yaoundé, 3.00°N, 10.92°E, 12 January 1914, Mildbraed G.W.J. 7727 (B). South-West Region: Nyasoso, 4.86°N, 9.7°E, 04 June 1996, Cable S. 2851 (K,YA); Along path from village Mt Etinde summit, 4.05°N, 9.15°E, 02 December 1993, Cable S. 332 (K); Kupe Rock saddle, 4.78°N, 9.716°E, 11 July 1996, Cable S. 3804 (K,YA); Kupe village, 4.78°N, 9.716°E, 17 July 1996, Cable S. 3894 (K,YA); Upper Boando, 4.05°N, 9.153°E, 08 December 1993, Cable S. 475 (K,YA); Mabeta Moliwe reserve 3–5 km east of Limbe, 4.00°N, 9.256°E, 03 July 1992, Cheek M. 3470 (P); Kupe village, 4.77°N, 9.688°E, 29 November 1999, Gosline W.G. 241 (K); Pente E Mont 6 km E Bomana 35 km NW Limbé Alt 950 m, 4.27°N, 9.112°E, 11 December 1984, Villiers J.-F. 2441 (P,YA); Saddle between Mt Etinde and Cameroon, 4.08°N, 9.116°E, 28 October 1992, Wheatley J.I. 644 (K,YA).

Cameroon. South Region; Bipindi, Zenker G.A. 510, 1 Jan 1914: lectotype, sheet here designated: P[P00363381]; isotypes: B[B 10 0153019]; BR[BR0000008820693]; P[P00363377]; U[U 0000237]; WAG[WAG0000084,WAG0000085].

Figure 13. 

Artabotrys dielsiana A branch B detail of the pubescence on the lower side of leaf blade C Inflorescence D flower E sepal, outside side F outer petals inner and outside view G inner petals inner and lateral side views H receptacle with stamens and carpels (stigmas showing) A–H from Zenker 510. Drawings D. Godor de Mauroy, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from Le Thomas (1969a, pl. 1, p. 441).

Liana, height unknown, d.b.h. unknown. Indumentum of simple hairs; old leafless branches pubescent, young foliate branches tomentose to densely pubescent with long, up to 2 mm, ferruginous hairs. Leaves: petiole 2–3 mm long, 1–2 mm in diameter, densely pubescent, grooved, blade inserted on top of the petiole; blade 15–24 cm long, 6–7 cm wide, elliptic, apex acuminate to acute, acumen 0.5–1 cm long, base subcordate, coriaceous, below densely pubescent when young and old, above sparsely pubescent when young, glabrous when old, concolorous; midrib impressed, above glabrous or sparsely pubescent when young, glabrous when old, below densely pubescent when young and old; secondary veins 13 to 16 pairs, glabrous above; tertiary venation reticulate. Individuals bisexual; inflorescences, ramiflorous on old leafless branches, generally without leaves, leaf opposed. Flowers with 9 perianth parts in 3 whorls, 10 to 15 per inflorescence, hook-shaped peduncle 12–17 mm long; pedicel 3–4 mm long, ca. 1 mm in diameter, densely pubescent; bracts ca. 2, all basal, basal bracts 5 mm long, 4 mm wide; sepals 3, valvate, free, 5–12 mm long, 3–7 mm wide, triangular, apex acute, base truncate, densely pubescent outside, glabrous inside, margins flat; petals free, sub equal; outer petals 3, 20–35 mm long, 6–10 mm wide, elliptic, apex acute, base broad and concave, margins flat, densely pubescent outside, glabrous inside; inner petals 3, valvate, 15–25 mm long, 6–9 mm wide, elliptic, apex acute, base broad and concave, margins flat, densely pubescent outside, pubescent inside; stamens 50 to 70, in 3 to 4 rows, 1–2 mm long, cuneiform; connective discoid, pubescent; staminodes absent; carpels free, 25 to 32, ovary 1–2 mm long, stigma cylindrical, glabrous. Fruits unknown.

endemic to Cameroon; known from the South region.

Rare species; in primary lowland rain forest. Altitude 0–200 m a.s.l.

None recorded.

Not evaluated.

None recorded.

Artabotrys dielsiana is characterized by its densely pubescent young foliate branches with longish ferruginous hairs, leaf blades with a subcordate leaf base and pubescent below but glabrous above (including the midrib), and tightly packed flowers mainly borne on leafless branches (Le Thomas 1969a). This species has only been collected four times.

South Region: Lélé, 2.29°N, 13.34°E, 06 September 2013, Couvreur T.L.P. 453 (WAG,YA); Campo-Ma’an area 2.28°N, 9.866°E, 03 October 2001, van Andel T.R. 4128 (KRIBI,U,WAG); Bipindi, 3.08°N, 10.41°E, 01 January 1914, Zenker G.A. 2087 (WAG); Bipindi, 3.08°N, 10.42°E, 01 January 1914, Zenker G.A. 510 (B,P,U,WAG).

Cameroon. South Region; Bipindi, Zenker G.A. 2801, 1904: lectotype, sheet here designated: B[B 10 0153021]; isotypes: B[B 10 0153022]; COI[COI00004928]; GOET[GOET005674]; HBG[HBG502547]; K[K000198855]; MO[MO-216862]; P[P00363364]; S[S-G-7465]; WAG[WAG0053175]; WU[WU0025886].

Liana, up to 10 m tall, d.b.h. unknown. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches pubescent with short appressed hairs. Leaves: petiole 2–3 mm long, 1–2 mm in diameter, sparsely pubescent to glabrous, grooved, blade inserted on the side of the petiole; blade 10–20 cm long, 3.5–6 cm wide, ovate to elliptic, apex acuminate to acute, acumen 0.5–1 cm long, base acute, subcoriaceous, below sparsely pubescent when young, glabrous when old, above glabrous when young and old, concolorous; midrib impressed, above sparsely pubescent to glabrous when young, glabrous when old, below sparsely pubescent to pubescent when young, sparsely pubescent when old; secondary veins 9 to 12 pairs, glabrous above; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless branches. Flowers with 9 perianth parts in 3 whorls, leaf opposed, 1 to 3 per inflorescence, hook-shaped peduncle 16–25 mm long; pedicel 2–5 mm long, 1–2 mm in diameter, sparsely pubescent with short appressed hairs; in fruit 2–3 mm long, ca. 2 mm in diameter, glabrous; bracts 2(?), all basal, basal bracts not seen; sepals 3, valvate, free, 10–15 mm long, 5–8 mm wide, triangular, apex acute, base truncate, green turning light reddish, pubescent outside, glabrous inside, margins flat; petals free, sub equal; outer petals 3, 30–35 mm long, 7–12 mm wide, elliptic, apex acute, base attenuate (rounded), green, margins flat but recurved outwards in vivo, pubescent to densely pubescent outside, pubescent to sparsely pubescent inside; inner petals 3, valvate, 15–25 mm long, 2–6 mm wide, elliptic to oblong, apex acute, base broad and concave, margins flat, but recurved outwards in vivo, densely pubescent to pubescent outside, pubescent to sparsely pubescent inside; stamens numerous, number of rows not seen, 2 mm long, cuneiform; connective discoid, glabrous; staminodes absent; carpels free, 12 to 17, ovary 3–4 mm long, stigma cylindrical, pubescent. Monocarps sessile, 7 to 9, 20–25 mm long, 10–12 mm in diameter, ellipsoid, apex long apiculate and slightly curved, glabrous, smooth, not ribbed; red when ripe, seeds 2 per monocarp, 8–11 mm long, 4–6 mm in diameter, flattened ellipsoid; aril absent.

A west and central African species, from Sierra Leone to Benin and from Cameroon to the Democratic Republic of the Congo; in Cameroon known from Central, East, South and South-West regions.

A fairly common species in Cameroon, in secondary rain forests a long fringes of forests, in swampy regions too. Altitude 100–800 m a.s.l.

None recorded.

Not evaluated.

None recorded.

Artabotrys insignis var. insignis is distinguished by its overall relatively glabrous branches and petioles, long (> 10 cm) elliptic leaves with an acute leaf base (more rarely rounded), long (10–15 mm) triangular sepals, long and wide petals, and glabrous smooth fruits with a distinctive long curved apicule.

Central Region: Route Ndanan I-Ndangan I, 3.62°N, 11.58°E, 10 March 2004, Cheek M. 11641 (K,YA); Yangafok II 25 km ENE de Bafia, 4.93°N, 11.37°E, 26 November 1969, Letouzey R. 9607 (P,YA). East Region: 82 km south of Yokadouma 30 km after Ngato 15 km after river ALPICAM ‘base de vie’ then 40 km on forestry road starting 4 km before Maséa village, 3.15°N, 14.73°E, 06 March 2019, Couvreur T.L.P. 1214 (MPU,WAG,YA). South Region: Campo Ma’an National Park 11 km on trail from Ebinanemeyong village on road 7 km from Nyabessan to Campo town, 2.46°N, 10.35°E, 14 February 2015, Couvreur T.L.P. 710 (WAG,YA); Ebom, 3.1°N, 10.73°E, 25 February 1997, Elad M. 580 (KRIBI,WAG); Ebom, 3.1°N, 10.71°E, 26 February 1997, Parren M.P.E. 4 (KRIBI,WAG); Bipindi, 3.08°N, 10.41°E, 01 January 1904, Zenker G.A. 2801 (B,BR,K,L,P,WAG); Bipindi, 3.08°N, 10.42°E, 01 January 1907, Zenker G.A. 3320 (P). South-West Region: Mount Cameroon National Park on the Bomona trail behind Bomona village 10 km NW from Idenau, 4.29°N, 9.096°E, 03 April 2016, Couvreur T.L.P. 1044 (MPU,WAG,YA); Korup National Park, 5.28°N, 9.083°E, 03 April 1988, Thomas D.W. 7578 (MO,P).