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Corresponding author: Thomas L. P. Couvreur ( thomas.couvreur@ird.fr ) Academic editor: Yasen Mutafchiev
© 2022 Thomas L. P. Couvreur, Leo-Paul M. J. Dagallier, Francoise Crozier, Jean-Paul Ghogue, Paul H. Hoekstra, Narcisse G. Kamdem, David M. Johnson, Nancy A. Murray, Bonaventure Sonké.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Couvreur TLP, Dagallier L-PMJ, Crozier F, Ghogue J-P, Hoekstra PH, Kamdem NG, Johnson DM, Murray NA, Sonké B (2022) Flora of Cameroon – Annonaceae Vol 45. PhytoKeys 207: 1-532. https://doi.org/10.3897/phytokeys.207.61432
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Annonaceae is a major pantropical family with 113 genera and about 2550 species. Cameroon is one of the most biodiverse countries in Africa but its flora remains incompletely known. In this volume of the Flora of Cameroon, we describe 166 native taxa representing 163 species in 28 native genera within the family Annonaceae. A total of 22 species (about 13%) are endemic to the country. We provide keys to all native genera, species, and infraspecific taxa. For each species a detailed morphological description and a map of its distribution in Cameroon are provided. Distribution maps and diversity analyses are based on a taxonomically verified database of 2073 collections. Across Africa, Cameroon is a center of diversity for Annonaceae harboring one of the highest numbers of species and genera. For example, Cameroon harbors the highest number of African species for the only pantropical genus of Annonaceae, Xylopia. Annonaceae are found across all 10 administrative regions of Cameroon but diversity is concentrated within the tropical rain forest areas situated in the south and South-West. The areas around Bipindi and Mount Cameroon show the highest levels of diversity, but this is correlated with collection effort. Line drawings and/or photographs accompany most species. One species new to science Uvariopsis etugeana Dagallier & Couvreur sp. nov. is described. We also undertake a number of nomenclatural changes such as lectotypifications, six new synonymies and two new combinations (Uvaria anisotricha (Le Thomas) Couvreur, comb. nov.; Uvariodendron fuscum var. giganteum (Engl.) Dagallier & Couvreur, comb. nov.).
Les Annonacées sont une grande famille pantropicale avec 113 genres et 2550 espèces. Le Cameroun est l’un des pays les plus riches en biodiversité d’Afrique mais sa flore reste incomplètement connue. Dans ce volume de la Flore du Cameroun, nous décrivons 166 taxons représentant 163 espèces dans 28 genres au sein de la famille des Annonaceae. Au total, 22 espèces (environ 13%) sont endémiques du pays. Nous fournissons une clé de tous les genres et espèces et des infra-espèces au sein des genres. Pour chaque espèce une description morphologique détaillée et une carte de sa répartition au Cameroun sont fournies. Les cartes de distribution et les analyses de diversité sont basées sur une base de données taxonomiquement vérifiée de 2073 collections. À travers l’Afrique, le Cameroun est un centre de diversité pour les Annonacées abritant l’un des plus grands nombres d’espèces et de genres. Par exemple, le Cameroun abrite le plus grand nombre d’espèces africaines pour le seul genre pantropical d’Annonaceae, Xylopia. Les Annonaceae sont présentent dans les 10 régions du Cameroun, mais la plus grande diversité est concentrée dans les régions abritant la forêt tropicale humide située au sud et au sud-ouest. Les zones autour de Bipindi et du Mont Cameroun présentent les niveaux de diversité les plus élevés, mais cela est corrélé à l’effort de collecte. Des dessins et/ou des photographies accompagnent la plupart des espèces. Une espèce nouvelle pour la science Uvariopsis etugeana Dagalier & Couvreur sp. nov. est décrite. Nous entreprenons également un certain nombre de changements nomenclaturaux tels que des lectotypifications, six nouvelles synonymies et deux nouvelles combinaisons (Uvaria anisotricha (Le Thomas) Couvreur, comb. nov.; Uvariodendron fuscum var. giganteum (Engl.) Dagalier & Couvreur, comb. nov.).
Africa, botanical identification, conservation, diversity, lectotypification, new species, taxonomy, vascular plants
Annonaceae is a large pantropical family of trees, shrubs and lianas (
Morphological species descriptions were automatically generated before manual checking and editing. We used the online collaborative platform PROTEUS (
Data was gathered from two main sources. First, we used all available taxonomic revisions of African Annonaceae genera (e.g.
A first draft of descriptions for all species was generated using the package MonographaR (
A database of Annonaceae collections from Cameroon was generated in which we recorded collector number, location, coordinates, region and herbarium where the specimens are deposited. A collection represents a herbarium sample identified by having the same collector and number (when present). It may be composed of one or more specimens, and such duplicates can be deposited in different herbaria. We used several sources as primary data providers to build the database. The initial database was based on data extracted from the “Réseau Informatique des Herbiers d’Afrique” (RIHA). This database contains all specimens held in the Herbier National du Cameroun/National Herbarium of Cameroon. We extracted all Annonaceae from Cameroon. We then supplemented this database with other available databases: BRAHMS (Naturalis Biodiversity Center, Leiden, The Netherlands); TROPICOS (Missouri Botanical Garden, St Louis, USA); Kew Database (Royal Botanical Gardens, Kew, UK) and other databases (T.L.P. Couvreur (IRD); V. Droissart (IRD); D.J. Harris (E); N. Kamdem (Université de Yaoundé I)). As much as possible specimens were checked in herbaria to confirm their identification. Thus, the database contains specimens that we have seen and confirmed, and others which we did not see. For genera with recent taxonomic revisions, determinations were updated (
Collections without coordinates were georeferenced using QGIS ver. 3.2.3 (
Within this project, numerous herbaria were visited over the course of the last eight years including B, BR, BRLU, G, K, P, YA and WAG. Within the taxonomic revision of certain genera, specimen loans were made available from other herbaria (BM, MO, OWU). In addition, we used specimen scans available online from these different herbaria when possible and needed. Numerous field trips were carried out across Cameroon over a period of eight years (2012–2019), mainly in the regions Central, East, Littoral, South West and West regions. During these field trips, high quality herbarium collections were made and deposited at MPU, YA and WAG. Finally, detailed photographs of the different parts available (leaves, trunk, flowers and fruits) were made and used to illustrate species found within this flora.
The database was used to generate collection, species and genus raw diversity maps. Collection density was log transformed before plotting. After filtering for unidentified species and genera, raw diversity maps were made at 0.25° resolution sampling units (SU) were plotted using the ‘ggplot2’ package (
Collection citations and the index to numbered collections were generated using the package ‘exsic’ (
As much as possible we tried to use the original line drawings drawn by Hélène Lamourdedieu, intended by Annick Le Thomas for this flora. Thanks to Thierry Deroin, we had access to her archives at the Muséum national d’Histoire Naturelle in Paris, where we found numerous line drawings not published in the Flore du Gabon. The numbering of drawings was retained when possible. In some cases, specimens used for drawing a specific species changed (new identification). In other cases drawings came from more recent taxonomic revisions.
The IUCN conservation status of each species was downloaded from the IUCN Red List website (www.iucnredlist.org). Official IUCN published evaluations are provided here, except for the genus Monanthotaxis where the preliminary status were taken from
We document a total of 28 native genera, 167 native taxa and 163 native species of Annonaceae in Cameroon (Table
List of number of 28 native genera recorded for Cameroon plus 12 continental African genera not found in Cameroon, with known accepted number of species in Cameroon, for continental Africa (including the Gulf of Guinea Islands, but excluding Madagascar) and percentage of species for each genus found within Cameroon. Total species diversity numbers were taken from
Genus | Number of species in Cameroon | Number of species in continental Africa | Percentage in Cameroon |
---|---|---|---|
Afroguatteria1 | 1 | 3 | 33 |
Annickia | 4 | 8 | 50 |
Annona | 1 | 4 | 25 |
Anonidium | 2 | 4 | 50 |
Artabotrys | 8 | 32 | 25 |
Brieya | 1 | 2 | 50 |
Cleistopholis | 3 | 3 | 100 |
Dennettia | 1 | 1 | 100 |
Duguetia | 4 | 4 | 100 |
Greenwayodendron | 2 | 5 | 40 |
Hexalobus | 4 | 5 | 80 |
Isolona | 9 | 15 | 60 |
Letestudoxa | 2 | 3 | 67 |
Meiocarpidium | 1 | 1 | 100 |
Mischogyne | 1 | 5 | 20 |
Monanthotaxis1 | 31 | 79 | 39 |
Monodora | 6 | 14 | 43 |
Neostenanthera2 | 3 | 5 | 60 |
Piptostigma | 13 | 13 | 100 |
Polyceratocarpus | 3 | 8 | 38 |
Sirdavidia | 1 | 1 | 100 |
Sphaerocoryne1 | 1 | 2 | 50 |
Toussaintia | 1 | 4 | 25 |
Uvaria | 17 | 77 | 23 |
Uvariastrum | 3 | 5 | 60 |
Uvariodendron | 5 | 18 | 28 |
Uvariopsis | 13 | 19 | 68 |
Xylopia | 22 | 45 | 49 |
Asteranthe (3), Cleistochlamys (1), Dielsiothamnus (1), Huberantha (4), Lettowianthus (1), Lukea (2), Mkilua (1), Monocyclanthus (1), Mwasumbia (1), Ophrypetalum (1), Pseudartabotrys (1), Sanrafaelia (1). | 0 | 19 | 0 |
Total : | 163 | 399 | 41 |
Genus | Species epithet | Author(s) |
---|---|---|
Afroguatteria | discostigma | (Diels) X.Guo & R.M.K.Saunders |
Artabotrys | dielsiana | Le Thomas |
Hexalobus | bussei | Diels |
Monanthotaxis | couvreurii | P.H.Hoekstra |
Monanthotaxis | dielsiana | (Engl.) P.H.Hoekstra |
Monanthotaxis | elegans | (Engl. & Diels) Verdc. |
Monanthotaxis | hexamera | P.H.Hoekstra |
Monanthotaxis | submontana | P.H.Hoekstra |
Monanthotaxis | zenkeri | P.H.Hoekstra |
Monodora | zenkeri | Engl. |
Piptostigma | goslineanum | Ghogue, Sonké & Couvreur |
Piptostigma | longepilosum | Engl. |
Piptostigma | macrophyllum | Ghogue, Sonké & Couvreur |
Piptostigma | mayndongtsaeanum | Ghogue, Sonké & Couvreur |
Piptostigma | submontanum | Ghogue, Sonké & Couvreur |
Uvaria | mollis | Engl. & Diels |
Uvariopsis | dicaprio | Gosline & Cheek |
Uvariopsis | etugeana | Dagallier & Couvreur |
Uvariopsis | korupensis | Gereau & Kenfack |
Uvariopsis | sessiliflora | (Mildbr. & Diels) Robyns & Ghesq. |
Uvariopsis | submontana | Kenfack, Gosline & Gereau |
Uvariopsis | zenkeri | Engl. |
A total of 2073 collections were seen for this treatment (Fig.
Annonaceae have been collected from all 10 regions in Cameroon, but most sampling comes from the southern regions of the country (Fig.
In terms of species diversity, there are two main hotspots, one located in the SU around Bipindi in the northwestern South Region (with 46 species recorded), and one in the SU around Mount Cameroon in the South-West region (with 40 species recorded) (Fig.
Spatial diversity of Annonaceae in Cameroon A log transformed collection density B raw species diversity C raw generic diversity D scatter plot of number of species in function of collections per SU, with correlation parameter R and p value. Maps represent Cameroon with limits between regions. Sampling units (SU) are of 0.25.
Genus diversity is also concentrated in the Atlantic forests, with hotspots along the Cameroonian Volcanic Line, and towards the western area of the South region. Once again, the SU around Bipindi has the highest generic diversity with 19 (mean: 3.6 genera / SU) recorded genera (Fig.
Xylopia is the most collected genus across Cameroon (277 collections), followed by Monanthotaxis (196 collections) (Table
Number of collections for all 28 genera recorded in Cameroon and for the top 27 species. Values based on collections between 1861 and 2019.
Genus | # specimens | Species | # specimens |
---|---|---|---|
Xylopia | 277 | Greenwayodendron suaveolens | 119 |
Monanthotaxis | 196 | Annickia affinis | 60 |
Uvariodendron | 165 | Uvariodendron connivens | 60 |
Monodora | 156 | Monodora myristica | 54 |
Uvariopsis | 135 | Anonidium mannii | 53 |
Greenwayodendron | 134 | Xylopia aethiopica | 45 |
Piptostigma | 126 | Monanthotaxis enghiana | 43 |
Artabotrys | 119 | Meiocarpidium oliverianum | 42 |
Uvaria | 115 | Xylopia thomsonii | 41 |
Isolona | 80 | Monodora undulata | 36 |
Annickia | 73 | Monodora tenuifolia | 34 |
Neostenanthera | 62 | Xylopia quintasii | 32 |
Anonidium | 59 | Uvariodendron molundense | 31 |
Hexalobus | 53 | Neostenanthera neurosericea | 30 |
Cleistopholis | 46 | Xylopia africana | 30 |
Annona | 44 | Artabotrys aurantiacus | 28 |
Uvariastrum | 44 | Neostenanthera myristicifolia | 28 |
Meiocarpidium | 42 | Uvariodendron calophyllum | 28 |
Duguetia | 40 | Annona senegalensis subsp. oulotricha | 25 |
Polyceratocarpus | 33 | Hexalobus crispiflorus | 25 |
Sphaerocoryne | 20 | Uvariodendron fuscum | 24 |
Brieya | 15 | Uvariastrum zenkeri | 22 |
Dennettia | 7 | Uvariopsis dioica | 22 |
Letestudoxa | 6 | Polyceratocarpus parviflorus | 21 |
Afroguatteria | 4 | Sphaerocoryne gracilipes | 20 |
Mischogyne | 3 | Uvariopsis bakeriana | 20 |
Sirdavidia | 1 | Artabotrys thomsonii | 19 |
Toussaintia | 1 | Duguetia staudtii | 19 |
Sampling history of Annonaceae in Cameroon A ranked number of collections (specimens) per taxa (including subspecies and varieties) B cumulative number of Annonaceae collections (specimens) through time from 1861 to 2019 C cumulative number of Annonaceae species through time from 1861 to 2019. This graph is based on 2060 herbarium collections (with known year of collection)
A total of 95 species received a conservation status from the International Union for the Conservation of Nature, IUCN. However, we excluded the species Uvariodendron fuscum which now also includes formerly distinguished taxa (U. mirabile and U. giganteum) and thus would need to be reassessed. In addition, the name Boutiquea platypetala is considered here to be a synonym of Neostenanthera neurosericea. We nevertheless used the available assessment of B. platypetala because N. neurosericea is only known from the type specimen (single location), and this nomenclature change does not affect the assessment. Thus, just over half (94 species) of the Cameroonian species have an official IUCN conservation assessment to date. Of these, only two are liana species (Uvaria angolensis, U. chamae). Two species were evaluated using old criteria (version 2.3 and were published before 2000). Of the 94 species considered, 24 (Table
List of the 24 Annonaceae species occurring in Cameroon officially assessed (and published) as Threatened following IUCN criteria. The assessment of Neostenanthera neurosericea is based on Boutiquea platypetala which is now a synonym of the former. Uvariopsis pedunculata was assessed under the former name U. vanderystii.
Genus and species epithet | Red list category | Red list criteria | Year published | Criteria version |
---|---|---|---|---|
Duguetia dilabens | Endangered | B2ab(iii,v) | 2020 | 3.1 |
Hexalobus bussei | Endangered | B1ab(ii,iii,iv)+2ab(ii,iii,iv) | 2019 | 3.1 |
Isolona pilosa | Vulnerable | B2ab(iii) | 2019 | 3.1 |
Isolona pleurocarpa | Endangered | B2ab(iii) | 2019 | 3.1 |
Mischogyne gabonensis | Endangered | B2ab(i,ii,iii,iv,v) | 2021 | 3.1 |
Neostenanthera neurosericea (as Boutiquea platypetala) | Vulnerable | B2ab(iii) | 2014 | 3.1 |
Piptostigma calophyllum | Vulnerable | B2ab(iii) | 2019 | 3.1 |
Piptostigma giganteum | Vulnerable | B1+2c | 1998 | 2.3 |
Piptostigma goslineanum | Vulnerable | B1ab(iii)+2ab(iii) | 2019 | 3.1 |
Piptostigma longepilosum | Endangered | B2ab(iii,v) | 2019 | 3.1 |
Piptostigma macrophyllum | Vulnerable | B2ab(iii,iv) | 2019 | 3.1 |
Piptostigma oyemense | Vulnerable | B2ab(iii) | 2019 | 3.1 |
Piptostigma submontanum | Endangered | B1ab(iii)+2ab(iii) | 2019 | 3.1 |
Sirdavidia solannona | Vulnerable | D2 | 2019 | 3.1 |
Uvariodendron giganteum | Vulnerable | B2ab(iii) | 2004 | 3.1 |
Uvariopsis korupensis | Endangered | B2ab(iii) | 2014 | 3.1 |
Uvariopsis submontana | Endangered | B1ab(iii)+2ab(iii) | 2014 | 3.1 |
Uvariopsis pedunculata | Vulnerable | B2ab(iii) | 2014 | 3.1 |
Uvariopsis zenkeri | Vulnerable | B2ab(i,ii,iii,iv,v). | 2021 | 3.1 |
Xylopia africana | Vulnerable | A2c | 2014 | 3.1 |
Xylopia calva | Endangered | B2ab(iii) | 2019 | 3.1 |
Xylopia gilbertii | Vulnerable | B2ab(iii) | 2019 | 3.1 |
Xylopia mildbraedii | Vulnerable | B2ab(iii) | 2019 | 3.1 |
Xylopia paniculata | Endangered | B2ab(iii) | 2019 | 3.1 |
Based on preliminary conservation assessments (
Cameroon is a diversity hotspot for Annonaceae (Table
The increase of Cameroonian Annonaceae specimens or species through time has not been constant (Fig.
Collecting intensity across the country is highly heterogeneous, with certain regions densely collected (Bipindi region, Mount Cameroon, Yaoundé area), whereas others have few or no collections (Fig.
Just over a quarter (25.5%, 24/93) of assessed Annonaceae species occurring in Cameroon have been evaluated as threatened at global levels under IUCN criteria (Table
Cameroonian Annonaceae provide an important sample of the morphological variability encountered across the family in general. Below we provide a brief overview of this variability. All these observations concern Cameroonian Annonaceae, unless stated otherwise.
In Cameroon, Annonaceae can be large emergent trees (e.g. several species of Xylopia more than 30 m in height), understory trees (e.g. Uvariodendron sp. between 3 and 15 m in hight), shrubs (Neostenanthera neurosericea), scrambling shrubs (i.e. lianas not growing along a tree, but mainly growing on the ground producing long bended stems; some species in Uvaria) and lianas (e.g. Artabotrys, Monanthotaxis). No epiphytic species are known in the family. If we just look at the dichotomy between liana and tree habits, there are 62 liana species (38%) versus 99 tree species (62%). Some species are intermediate between a liana and tree habit. This is the case for example of Neostenanthera myristicifolia, Xylopia thomsonii, or Monodora crispata, which can appear scandent, leaning on surrounding vegetation (
Annonaceae exhibit two growth architecture patterns (
Most Annonaceae species have a smooth and cylindrical trunk with no buttresses or stilt roots. The genus Xylopia is an exception with a range of basal structures from small or large buttresses to stilt roots or no structure at all. This provides a useful character for species identification (
Leaves in Annonaceae are alternate, distichous (in a single plane) and lack stipules. The single plane disposition of the leaves confers a characteristic look to Annonaceae in the forest. Moreover, most species have clearly plagiotropic (horizontally spreading) branches (e.g. Greenwayodendron). Leaves are always entire and the margins are never serrate or dentate. Petiole length varies from 1 to 20 mm, the longer petioles being found in species such as Annona senegalensis, Cleistopholis glauca and Uvariodendron calophyllum.
The leaf blade is inserted either on the top or to the side of the petiole (see fig. 2 of
Within Cameroonian species, leaf size varies more than 20-fold, from 3–4 cm long in species such as Uvaria klaineana or Xylopia pynaertii, to up to more than 60 cm long in species such as Piptostigma submontanum or Uvariodendron fuscum var. giganteum. However, most species have intermediate-sized leaves between 10 and 30 cm long.
Most Annonaceae species have concolorous leaves, being green on both sides of the leaf blade. Some genera and species, however, can be distinguished by having discolorous leaves, with a much lighter whitish green color of the lower side of the leaf blade, e.g. Afroguatteria, Brieya, Cleistopholis, Monanthotaxis, Piptostigma, Polyceratocarpus, Neostenanthera, Sphaerocoryne, Uvaria or some species of Xylopia.
In most species, the midrib is flat or sunken on the adaxial surface. The genera Isolona and Monodora are unusual in African Annonaceae as they have a raised midrib (
Secondary venation can provide a useful taxonomic character in Annonaceae. Most species have fewer than 20 pairs of secondary veins. The genus Piptostigma however, is characterized by leaves with a generally high number of parallel secondary veins (
The inflorescences of Annonaceae species are termed monotelic, meaning that the apex of the inflorescence ends with a terminal flower (
An important character concerns the position of the inflorescences which are, in the most fundamental sense, either ‘axillary’ that is originating from an axillary meristem, or ‘terminal’ that is originating from a terminal meristem (
Most species of Annonaceae have a clearly pedicellate flower, with pedicels generally shorter than 10 cm, and in most cases between 0.2 and 2 cm long. Uvariopsis congolana has (female) pedicels up to 45 cm long that grow from the base of the trunk and along the forest floor (flagelliflory, see
In most genera, the pedicel bears a lower and upper bract conforming to Fries’s type 2, the most common situation across Annonaceae (
Annonaceae flowers are generally bisexual, with stamens and carpels within the same flower. However, some genera are androdioecious (male and bisexual flowers on different individuals, although this state needs to be confirmed with more detailed field observations), dioecious (male and female flowers on different individuals) or monoecious (male and female flowers on the same individual). In Monanthotaxis we can find bisexual and monoecious species. Several Monanthotaxis species with unisexual flowers have female flowers on the trunk while the male flowers are located high in the canopy in axils of the leaves, therefore collections often only contain male or female flowers. Monoecy has not been proven for M. cauliflora or M. pynaertii, but there are collections with both female and male flowers in other species such as M. bidaultii, M. diclina, M. letouzeyi. Uvariopsis is also monoecious, while Anonidium (but see under A. brieyi), Greenwayodendron and Polyceratocarpus are androdioecious (
The receptacle or torus which bears the stamens on the basal part and/or the carpels towards the central apical part is quite variable within Annonaceae in general (
The general floral pattern in Annonaceae is actinomorphic, cyclic and trimerous with one whorl of three sepals and two whorls of three petals each (
Sepals are mostly free or are basally fused. Sometimes it can be hard to distinguish between these two states. In Letestudoxa and some species of Uvaria, the sepals are completely fused into a tube or a “cup” and tear open (generally into three parts) during anthesis. In Xylopia the fused sepals often form a cup-shaped calyx. Petals are free in most species, but in the genera Hexalobus, Isolona and Monodora they are basally fused. In this case the petals form either in single whorl (Hexalobus, Isolona) or two differentiated whorls (Monodora). In the former case, the petals are identical in shape and size (referred to as “not differentiated” in the descriptions) and the petals are clearly fused basally; we refer to the non-fused part as “lobes” and fused part as the “tube” (
Petal shape and size are very variable across Cameroonian species ranging from 1–2 mm long in male flowers of Monanthotaxis cauliflora (female flowers are larger) to 50 mm in Anonidium mannii, to 79 mm in Xylopia mildbraedii,or even up to 100 mm in Monodora myristica, and from circular (several species in Monanthotaxis) to linear (Artabotrys, Xylopia). In some species, the inner petals are rounded and concave at the base, and form a pollination chamber (e.g. Artabotrys, Neostenanthera, Xylopia). Pollination chambers are also possible by connivance of the inner petal margins (but not fused together), for example in some species of Monodora or Uvariodendron.
Stamen number varies from six in three species of Monanthotaxis (
In Annonaceae, the carpels are generally free, that is apocarpous, within the flower (
Carpel number varies from one in Sirdavidia to over 250 in Uvariopsis dioica. We provide a count for each species based on available material, but these numbers remain estimates in most cases. Most species have fewer than 20 carpels. Genera with pseudosyncarpous fruits (see below, Annona, Anonidium, Duguetia, Letestudoxa) generally have more than 50 carpels and up to 120. Other genera such as Uvariodendron or Uvariopsis, Monanthotaxis, Neostenanthera and Uvaria also have species which can have more than 50 carpels. Carpels are generally topped by a stigma which can be variable in shape from bilobed to globose or filiform (
The ovules are either numerous and lateral in one or two rows (
Most genera have aggregated fruits composed of individual units termed “monocarps” each resulting from the fertilization of a single carpel (
Monocarps are either stipitate or sessile. When present, the stipes can be up to 5 cm long in certain species of Annickia, Neostenanthera or Uvaria, conferring to the characteristic “star shape” look to Annonaceae fruits. In Annickia, Cleistopholis, and Neostenanthera, the stipe is articulated at the apex. In some species, the stipe is short (less than 5 mm) but still present. A number of genera have sessile (no apparent stipe) or short-stipitate (less than 5 mm) monocarps, for example Mischogyne, Uvariodendron or Uvariopsis. Sessile and stipitate monocarps can occur within the same genus, for example in Neostenanthera or Uvaria. Monocarps of most Cameroonian genera are indehiscent, but in Xylopia the monocarps dehisce longitudinally along an abaxial suture.
Annonaceae seeds are quite variable in size, ranging from 3 mm to more than 40 mm long. Generally in Annonaceae, the seeds are characterized by a ruminate endosperm, resulting from the invasion of the intertegument into the endosperm (
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.
1 | Midrib of leaf blade clearly raised above | 2 |
– | Midrib of leaf blade sunken,impressed, or flat above | 4 |
2 | Petals fused at base (even just shortly); fruits in a single unit (syncarpous) | 3 |
– | Petals free; fruits in several independent monocarps (apocarpous) | Polyceratocarpus pellegrinii |
3 | Corolla lobes similar and equal in length, forming a distinct tube at the base, margins generally flat | Isolona |
– | Corolla lobes clearly differentiated into inner and outer petals; the outer ones longer than inner ones, margins generally undulate or crisped | Monodora |
4 | Liana or scrambling shrub | 5 |
– | Tree or shrub | 14 |
5 | Hook-shaped structures (modified inflorescence) present on branches even in juvenile plants | Artabotrys |
– | Hook-shaped structures absent | 6 |
6 | Indumentum of stellate and/or fasciculate hairs | Uvaria (pro parte) |
– | Indumentum (if present) of simple hairs | 7 |
7 | Anthers septate (few species) | 8 |
– | Anthers not septate (most species) | 9 |
8 | Petals subequal; stipe shorter than seeded section of monocarp; indehiscent; several seeds per monocarp | Xylopia thomsonii |
– | Outer petals longer than inner; stipe at least twice as long as seeded section of monocarp; monocarps indehiscent; seed 1 per monocarp | Neostenanthera myristicifolia |
9 | Leaves bicolored; above green, below glaucous to whitish; monocarps moniliform when more than one seed, ovules uniseriate | 10 |
– | Leaves green on both sides; monocarps not moniliform, globose to conical, ovules biseriate | 12 |
10 | Tertiary venation percurrent when viewed from below, or if venation obscure, then stamens < 35 | Monanthotaxis |
– | Tertiary venation reticulate and stamens > 40 | 11 |
11 | Inflorescences terminal (leaf opposed or extra-axillary) | Afroguatteria |
– | Inflorescences axillary | Sphaerocoryne |
12 | Receptacle columnar or elongated; petals 6 to 12 in 2 or 3 whorls | Toussaintia |
– | Receptacle convex but not columnar; petals 6 in two whorls | 13 |
13 | Sepals entirely fused, enclosing flower in bud, tearing as flower enlarges; fruits pseudosyncarpous | Letestudoxa |
– | Sepals free or basally fused, not enclosing flower in bud; fruits apocarpous | Uvaria (pro parte) |
14 | Indumentum of scale-like hairs (easily visible with a hand lens) | Meiocarpidium |
– | Indumentum (if present) of stellate, fasciculate or simple hairs | 15 |
15 | Indumentum of stellate and/or fasciculate hairs and fruits pseudosyncarpous | Duguetia |
– | Indumentum of simple hairs, or glabrous; fruits mostly apocarpous (pseudosyncarpous in Annona and Anonidium) | 16 |
16 | Sepals 2; petals 4 | Uvariopsis |
– | Sepals 3; petals 3 or 6 | 17 |
17 | Inner bark/slash yellow; petals 3, opposite the 3 sepals | Annickia |
– | Inner bark/slash cream to reddish; petals 6, or if 3 then only 2 sepals present | 18 |
18 | Petals fused into a clear tube at the base, plicate (transversely folded) in bud | Hexalobus |
– | Petals free, petals not plicate (not folded in bud) | 19 |
19 | Outer petals reduced, sepal like, smaller than inner petals | 20 |
– | Outer petals not reduced, subequal to or larger than inner petals | 21 |
20 | Secondary veins 11 to 17 pairs; inflorescence compact, generally up to than 10(–15) mm long | Brieya |
– | Secondary veins (15–)22 to 66 pairs; inflorescence compact to lax, but always longer than 16 mm | Piptostigma |
21 | Receptacle cylindrical; anther connective reduced to a tuft of hairs | Mischogyne |
– | Receptacle convex to flat; connective well developed, discoid to apiculate | 22 |
22 | Tertiary venation percurrent | 23 |
– | Tertiary venation reticulate | 25 |
23 | Leaf apex obtuse, rounded or emarginate (in Cameroonian species); fruits (pseudo)syncarpous | Annona |
– | Leaf apex acute, acuminate or caudate; fruits apocarpous | 24 |
24 | Outer petals much longer than inner petals; inner petals forming a dome over the receptacle; anthers septate; seed 1 per monocarp | Neostenanthera |
– | Petals sub equal or outer slightly longer; inner petals not forming a dome over the receptacle; anthers not septate; seeds > 1 per monocarp | Polyceratocarpus (pro parte) |
25 | Sepals reduplicate-valvate, buds ridged | Uvariastrum |
– | Sepals not reduplicate-valvate, buds not ridged | 26 |
26 | Petals homogenously red to pink, all reflexed at maturity; anthers bright yellow in vivo at maturity; carpel 1; monocarp 1 | Sirdavidia |
– | Petals green, yellow, cream, not reflexed or only curved outward; anthers not bright yellow; carpels > 1; monocarps generally more than one | 27 |
28 | Anthers septate; monocarps dehiscent | Xylopia |
– | Anthers not septate; monocarps not dehiscent | 29 |
29 | Individuals androdioecious or dioecious with separate male, female or bisexual flowers | 30 |
– | Individuals with bisexual flowers | 31 |
30 | Flowering peduncles present, > 50 mm long; stamens more than 30; fruits (pseudo)syncarpous | Anonidium |
– | Flowering peduncle absent; stamens less than 30; fruits apocarpous | Greenwayodendron |
31 | Flowering pedicels > 15 mm; sepals free; outer petals up to five times longer than inner petals | Cleistopholis |
– | Flowering pedicels < 15 mm; sepals basally fused; petals subequal | 32 |
32 | Petioles 2–5 mm long, 1–2 mm wide; petals 3(4), less than 10 mm long, basally fused | Dennettia |
– | Petioles > 4 mm long, 3–9 mm wide; petals 6, 10 mm or longer, free | Uvariodendron |
Genera in parentheses means some but not all species have the indicated trait.
Liana or scrambling to scandent shrub: Afroguatteria; Artabotrys; Uvaria; (Monodora); Monanthotaxis; (Neostenanthera); Sphaerocoryne; Toussaintia; Letestudoxa; (Xylopia).
Tree: Annickia; Annona; Anonidium; Brieya; Cleistopholis; Duguetia; Greenwayodendron; Hexalobus; Isolona; Meiocarpidium; Mischogyne; Monodora; Neostenanthera; Piptostigma; Polyceratocarpus; Uvariastrum; Uvariodendron; Uvariopsis; Xylopia.
Slash of the bark yellow: Annickia.
Stilt roots or buttresses present: (Xylopia).
Indumentum of stellate hairs: Annickia, Duguetia; Uvaria.
Indumentum of lepidote hairs: Meiocarpidium.
Hook-like structures on branches: Artabotrys.
Leaves discolorous, light green to whitish below: Afroguatteria; Cleistopholis, Monanthotaxis; Piptostigma, Sphaerocoryne; (Uvaria), (Xylopia).
Trunk whitish overall: Cleistopholis; Greenwayodendron.
Leaves with many parallel secondary veins (> 25): (Piptostigma), (Uvaria), (Uvariodendron).
Midrib clearly raised above: Isolona; Monodora; (Polyceratocarpus).
Sepals 2; petals 4: Uvariopsis; (Monanthotaxis).
Sepals reduplicate-valvate (margins folded in bud): Uvariastrum, Toussaintia.
Petals fused into a single whorl with a distinct tube: Hexalobus; Isolona.
Petals 3: Annickia; Dennettia; (Monanthotaxis).
Petals 9 to 10 inserted in 2 to 3 whorls: Toussaintia.
Petals plicate in bud, transversely folded when open: Hexalobus.
Inner petals much longer than outer: Brieya; Piptostigma.
Anthers septate: Neostenanthera; Xylopia.
Staminodes present: (Monanthotaxis); Xylopia.
Androdioecious, dioecious or monoecious: Anonidium; Greenwayodendron; (Monanthotaxis); Polyceratocarpus; Uvariopsis.
Pseudosyncarpous fruits (individual monocarps visible): Annona, Anonidium, Duguetia, Letestudoxa.
Fruits syncarpous (individuals monocarps not visible) with numerous unordered seeds: Isolona; Monodora.
Monocarps moniliform: Monanthotaxis; (Xylopia).
Monocarps dehiscent: Xylopia.
Seeds arillate: (Duguetia); (Xylopia).
Seeds with a sarcotesta: (Xylopia).
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 (
no revision has yet been published, but see
≡ Cleistopholis discostigma Diels, Bot. Jahrb. Syst. 39: 474, 1907; Oxymitra discostigma (Diels) Ghesq. ex Pellegr., Bull. Soc. Bot. France, 66, 1949; Richella discostigma (Diels) R.E.Fr., in Engler & Prantl Nat. Pflanzenfam., ed. 2, 17a (2): 139, 1959; Friesodielsia discostigma (Diels) Steenis, Blumea 12: 359, 1964.
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) (
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 (
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.
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
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 Oliv. nom. illeg.; J. Linn. Soc., Bot. 9: 174–175, 1867.
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.
1 | Upper side of midrib pilose | 2 |
– | Upper side of midrib glabrous, or pubescent just at the basal part, never pilose | 3 |
2 | Lower leaf surface with simple, bifid and trifid hairs; petals pubescent at base inside; monocarps with stipes < 20 mm | A. chlorantha |
– | Lower leaf surface at least with some stellate or fasciculate hairs; petals glabrous at base inside; monocarps with stipes > 20 mm | A. polycarpa |
3 | Pubescence on lower leaf surface simple, bifid, trifid or stellate, pointing in all directions | 4 |
– | Pubescence on lower leaf surface simple or bifid, all hairs pointing towards leaf apex | A. affinis |
4 | Petiole and young shoots tomentose | A. letestui |
– | Petiole and young shoots sparsely pubescent to pubescent | A. chlorantha |
≡ Enantia affinis Exell, J. Bot. 64, Suppl.: 9, 1926.
= Enantia chlorantha var. soyauxii Engl. & Diels, Monogr. Afrik. Pflanzen.-Fam. 6: 70 1901. Type. Gabon. Estuaire, Munda, Sibange Farm, Soyaux H. 125, 21 Sep 1880: lectotype, designated by
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) (
medicine
: bark as a malaria prophylaxis; construction: house building, furniture; dyes and tannins: as a yellow dye (
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 (
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).
≡ Enantia chlorantha Oliv., J. Linn. Soc., Bot. 9: 175, 1867.
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) (
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 (
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).
≡ Enantia letestui Le Thomas, Adansonia sér. 2, 2: 306, 1962.
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.
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
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) (
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.
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).
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
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.
≡ Unona polycarpa DC., Syst. Nat. 1: 499, 1817; Coelocline polycarpa (DC.) A.DC., Mém. Anon.: 33, 1832; Melodorum polycarpum (DC.) Benth., Trans. Linn. Soc. London 23: 477–478, 1862; Xylopia? polycarpa (DC.) Oliv., Fl. Trop. Afr. 1: 32, 1868.
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) (
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).
= Guanabanus Mill. Gard. Dict. Abr., ed. 4: 2, 1754.
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 (
no recent revision, but see
1 | Leaves glabrous, elliptic in shape with an acuminate apex, petiole inserted on the side of the petiole | A. glabra |
– | Young leaves always pubescent, generally obovate (more rarely elliptic), rounded to emarginated at the apex; petiole inserted on the top of the petiole | 2 |
2 | Lower side of leaf blade tomentose with short curly hairs covering the whole blade | A. senegalensis subsp. oulotricha |
– | Lower side of leaf blade glabrescent to densely pubescent with non-curly hairs | A. senegalensis subsp. senegalensis |
= Annona arenaria var. obtusa Robyns & Ghesq., Bull. Soc. Roy. Bot. Belge 67: 22 (1934). Type. Republic of the Congo. Pool, Brazzaville, Chevalier A.J.B. 27304, Jul 1912; holotype: P[P00363246].
= Annona arenaria auct., non Thonn., Robyns & Ghesq., Bull. Soc. Roy. Bot. Belge 67: 22 (1934); Annona senegalensis var. arenaria
(Thonn.) Sillans, Bull. Mus. Natl. Hist. Nat., sér. 2, 24: 581 (1952). Type. Democratic Republic of the Congo. Kongo-Central, Temvo, Vermoesen F.M.C. 1592, 20 Fev 1919: neotype, designated by
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 (
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 (
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
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).
= Annona arenaria Thonn., Beskr. Guin. Pl. 257, 1827. non Robyns & Ghesq., Bull. Soc. Roy. Bot. Belge 67: 22, 1934. Type. Ghana. Thonning s.n.: holotype: P [Herb. Jussieu, number: 10799].
= Annona chrysophylla Bojer, Ann. Sci. Nat., Bot. sér. 2, 20: 53, 1943; Annona senegalensis var. chrysophylla
(Boj.) Sillans, Bull. Mus. Natl. Hist. Nat., sér. 2, 24: 581 (1952). Type. Comores. Anjouan [Ndzuwani, Nzwani], Bojer W. s.n., s.d.: holotype: We were not able to locate the type specimen.
= Annona senegalensis var. latifolia
Oliv., Fl. Trop. Africa: 17, 1868. Type. Uganda. Northern region, Madi, Speke & Grant s.n., s.d.: holotype: We were not able to locate the type specimen, which should be in Kew (
Annona porpetac Boiv. ex Baill.; Bull. Mens. Soc. Linn. Paris 1. 341, 1882; Annona senegalensis var. porpetac (Boiv. Ex Baill.) Diels, Notizbl. Konigl. Bot. Gart. Berlin 9: 356, 1934. Type. Madagascar. Antsiranana Province, Nossi Be, Bovin M. 2115, 1846: holotype: P[P030360].
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 (
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
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 (
No recent revision has yet been published, but see
1 | Leaves obovate to oblong-elliptic, 20–45 cm, upper bract inserted at middle of flowering pedicel; individuals male or hermaphrodite (androdioecious); sepals basally fused; outer petals 25–50 mm long, 20–40 mm wide, elliptic to obovate | A. mannii |
– | Leaves distinctly oblong, 37–50 cm long; upper bract inserted directly under the calyx, orbicular; individuals female or male (dioecious), sepals free, outer petals 35–60 mm long, 10–18 mm wide; narrowly elliptic to narrowly oblong | A. brieyi |
≡ Anonidium mannii var. brieyi (De Wild.) R.E.Fr., Acta Hort. Berg. 10: 80, 1930.
= Anonidium friesianum Exell, J. Bot. 70, Suppl. Polypet.: 211, 1932. Type. Angola. Mayombe, Buco Zau, Gossweiler J. 6690, 16 Sep 1916: lectotype, designated here: LISC[LISC000056]; isolectotypes: COI[COI00004880]; BM[BM000546826, BM000546827]; LISC[LISC000054, LISC000055, LISC000057, LISC000058, LISC000059, LISC000060, LISC000061, LISC000062, LISC000063].
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.
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
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.
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 (
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 (
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).
≡ Annona mannii Oliv., Hooker’s Icon. Pl. vol. 11: 7–8, 1867.
= Uvaria crassipetala Engl., Notizbl. Königl. Bot. Gart. Berlin 2: 292, 1899. Type. Cameroon. South-West Region, Station Johann-Albrechtshohe, Staudt A. 813, no date: holotype B destroyed, lectotype, here designated: PH[PH28648].
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) (
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).
= Ropalopetalum Griff. Not. Pl. Asiat. 4: 716, 1854.
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 (
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
1 | Upper side of midrib glabrous, or pubescent just at the basal part, never densely pubescent | 2 |
– | Upper side of midrib densely pubescent | A. thomsonii |
2 | Young foliate branches and petioles glabrous or sparsely pubescent | 3 |
– | Young foliate branches and petioles densely pubescent to tomentose | 7 |
3 | Petioles 10–15 mm long | A. congolensis |
– | Petioles less than 8 mm long | 4 |
4 | Leaves 10–20 cm m; sepals 10–15 mm long and 5–8 mm wide, apex of monocarps clearly apiculate, apicule curved | A. insignis var. insignis |
– | Leaves smaller than 13 cm; sepals < 5 mm long and < 3 mm wide, apex of monocarps rounded | 5 |
5 | Flowering pedicels 10–25 mm long; sepals minute, ca. 1 mm long and ca. 1 mm wide, petals linear, 1–2 mm wide above the broad base, pubescent; monocarps 20–40 mm 10–20 mm in diameter, warty to verrucose, faintly ribbed | A. jacquesfelicis |
– | Flowering pedicels 7–10 mm, sepals 3–5 mm long 2–3 mm wide, petals elliptic to ovate, 4–9 mm wide above the broad base, tomentose; monocarps 15–20 mm 7–13 mm in diameter, smooth, not ribbed | 6 |
6 | Inflorescence pauciflorous, 1 to 4 flowers | A. aurantiacus var. aurantiacus |
– | Inflorescence multiflorous, 6 to 15 flowers | A. aurantiacus var. multiflorus |
7 | Young foliate branches and petioles hirsute with long erect hairs | A. rufus |
– | Young foliate branches densely pubescent with appressed or shortly erect hairs | 8 |
8 | Lower side of leaf blade densely pubescent brown, base of leaves subcordate with the leaf base inserted on top of petiole, secondary veins 13 to16 pairs, inflorescences multiflorous, > 15 flowers, generally on leafless branches | A. dielsiana |
– | Lower side of leaf blade sparsely pubescent to glabrous, base of leaves decurrent to acute with the leaf base inserted on the side of petiole, secondary veins 8 to 12 pairs, inflorescences pauciflorous, < 10 flowers, generally on leafy branches | 9 |
9 | Leaves 10–20 cm, leaf base acute, sepals 10–15 mm long and 5–8 m wide, petals 30–35 mm long, 7–12 mm wide, elliptic, not tubular | A. insignis var. batesii |
– | Leaves 8–12 cm, leaf base usually decurrent (but can also be acute), sepals 2–3 mm long and 2–3 m wide, petals 5–15 mm long, 1–2 mm wide, linear, tubular in shape | A. velutinus |
= Artabotrys pynaertii De Wild., Ann. Mus. Congo Belge, Bot. sér. 5, 3(1): 78, 1909. Type. Democratic Republic of the Congo. Equateur, Eala, Pynaert L.A. 606, 15 Oct 1906: lectotype, sheet here designated: BR[BR0000008809971]; isotypes: BR[BR0000008809964, BR0000008809988]; S[S07-13416].
= Artabotrys claessensii De Wild., Bull. Jard. Bot. État Brux. 3: 262, 1911. Type. Democratic Republic of the Congo, Orientale, Yangambi, Claessen, J. 725, Jul 1910: lectotype, sheet here designated: BR[BR0000008809995]; isotype: BR[BR0000008809988].
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).
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
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.
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
The variety status is doubtful, but without further investigation, we shall follow
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).
= Artabotrys rhopalocarpus Le Thomas, Adansonia sér. 2, 6: 591, 1966. Type. Central African Republic: Lobaye, Boukoko, Tisserant C. 2242, 25 Sep 1951: lectotype, sheet here designated: P[P00364752]; isotypes: BM[BM000546867]; BR[BR0000008820822]; LISC[LISC000373]; P[P00363392, P00363390]; WAG[WAG0392422].
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.
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
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 (
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].
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
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 (
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).
= Artabotrys malchairi De Wild., Etudes Fl. Bangala & Ubangi: 312, 1911. Type. Democratic Republic of the Congo. Equateur, Environ de Likimi, Malchair L. 282, 20 Apr 1910: lectotype, sheet here designated: BR[BR0000014480478]; isotype: BR[BR0000014480461].
= Artabotrys insignis var. latifolius Pellegr., Bull. Soc. Bot. France 94: 256, 1947. Type. Gabon. Ogooué-Lolo, région de Lastoursville, Moughimba, Le Testu G.M.P.C. 8474; 27 Oct 1930: lectotype, here designated: P[P01954179]; isolectotype: BM[BM000546856].
= Artabotrys lucidus A. Chev.; Expl. Bot. Afr. Occ. Franc., 1: 9, 1920, nom. nud.
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).
Cameroon. East Region; Bitya, near Dja river, Bates G.L. 1792, Sep 1922: holotype: P[00363370].
Differs from the type variety by its densely brown tomentose and shortly hirsute branches and petioles, pubescent lower side of leaf blades and tomentose petals.
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.
Differences between var. insignis and var. batesii are quite small, and are mainly related to the pubescence of the branches, lower side of the leaves and petals (
Artabotrys dielsiana A branch B base of leaf blade; upper side C base of leaf blade; lower side D detail of hooked shaped inflorescence (sterile). Artabotrys insignis var. batesii E branch F base of leaf blade; lower side, note tomentose branches G base of leaf blade; upper side, note tomentose branches H flower, side view I flower, top view J flower, bottom view A–D Couvreur 453, Lélé, Cameroon E–J Couvreur 1044, Mt Cameroon, Cameroon. Photos Thomas L.P. Couvreur.
East Region: Village Djang 40 km west of Bertoua, 4.58°N, 13.35°E, 15 May 1962, Breteler F.J. 2956 (P,WAG). South Region: Bitye near R Ja, 3.02°N, 12.37°E, 01 September 1922, Bates G.L. 1792 (P); Rive du Ntem à Ebianemeyong 60 km east de Campo, 2.42°N, 10.33°E, 12 April 1970, Letouzey R. 10370 (P,WAG,YA).
= Artabotrys robustus Louis ex Boutique Bull. Jard. Bot. Etat Brux. 21: 107, 1950. Type. Democratic Republic of the Congo. Orientale, Yangambi, Louis J.L.P. 6077, 16 Sep 1937: lectotype, sheet here designated: BR[BR0000008820686]; isotypes: BR[BR000000882072]; K[K000795930]; NY[NY00025831]; P[P00363357].
Cameroon. Central Region; Nkidi forest, Jacques-Félix H. 2490, Nov 1938: lectotype, sheet here designated: P[P00363361]; isotypes: K; P[P00363359, P00363362].
Liana, height unknown, d.b.h. unknown. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 4–7 mm long, ca. 1 mm in diameter, glabrous, grooved, blade inserted on the side of the petiole; blade 8–13 cm long, 3–14 cm wide, oblong to elliptic, apex acute, acumen ca. 0.5 cm long, base decurrent to acute, coriaceous, below glabrous when young and old, above glabrous when young and old, shiny when dried, concolorous; midrib impressed, above glabrous when young and old, below glabrous when young and old; secondary veins 9 to 12 pairs, glabrous above; tertiary venation reticulate. Flowers bisexual with 9 perianth parts in 3 whorls. Individuals bisexual; inflorescences ramiflorous on old leafless branches, leaf opposed. Flowers with 9 perianth parts in 3 whorls, 3 to 6 per inflorescence, hook-shaped peduncle 15–20 mm long; pedicel 10–25 mm long, ca. 1 mm in diameter, glabrous; in fruit 20–30 mm long, 1–2 mm in diameter, glabrous; bracts 1 to 2, all basal, basal bracts ca. 1 mm long, ca. 1 mm wide; sepals 3, valvate, free, 1–2 mm long, ca. 1 mm wide, triangular, apex acute, base truncate, glabrous outside, glabrous inside, margins flat; petals free, sub equal; outer petals 3, 20–35 mm long, 1–2 mm wide, linear to narrowly ovate, apex rounded, base broad and concave, margins flat, pubescent outside, glabrous inside; inner petals 3, valvate, 15–25 mm long, 1–3 mm wide, linear, apex rounded, base broad and concave, margins flat, densely pubescent outside, glabrous inside; stamens 50 to 70, in 3 to 4 rows, 2–3 mm long, oblong; connective discoid, pubescent; staminodes absent; carpels free, 3 to 4, ovary 3–4 mm long, stigma cylindrical, glabrous. Monocarps stipitate, stipes 5–10 mm, ca. 6 mm in diameter, 1 to 4 monocarps, 20–40 mm long, 10–20 mm in diameter, ellipsoid, apex rounded, glabrous, warty to verrucose, faintly ribbed, color when ripe unknown; seeds 1 to 2 per monocarp, 20–25 mm long, 10–15 mm in diameter, flattened ellipsoid; aril absent.
A central African species, only known from Cameroon and the Democratic Republic of the Congo; in Cameroon known from the Central and South regions.
Artabotrys jacquesfelicis A flowering branch B outer petal, inner view C inter petal, inner view D floral receptacle, petals removed E stamens, front and back view F carpel with detail on ovules G fruiting branch H seed, section latitudinal section I seed, longitudinal section. Artabotrys velutinus J flowering branch K flower receptacle, outer petals removed L outer petal, inner view M inner petal, inner view N flowering receptical, petals removed O stamen P carpel, side view, detail of ovules A–F from Jacques Felix 2490 G–I from Tisserant 2405 J–P from Zenker 1222. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris.
A rare species across its range, in primary lowland rain forests. Altitude 500–600 m a.s.l.
None recorded.
Not evaluated.
None recorded.
Artabotrys jacquesfelicis is distinguished by its entirely glabrous branches, petioles and leaves that are shiny above in herbarium material, narrowly ellipsoid flower buds, petals with a large concave base abruptly narrowing into an upper linear section, and ellipsoid monocarps with a warty to verrucose surface. In Cameroon, this species is only known from three old collections.
South Region: River Ja Bitya, 3.02°N, 12.37°E, 01 January 1922, Bates G.L. 1699 (K); Bipindi, 3.08°N, 10.42°E, 01 January 1909, Zenker G.A. 3834 (L,P).
= Artabotrys boonei De Wild., Repert. Spec. Nov. Regni Veg. 13: 383, 1914. Syn. nov. Type. Democratic Republic of the Congo. Orientale, Nala, Boone A. 80, 1911: lectotype, sheet here designated: BR[BR0000008820365]; isotype: BR[BR0000008820372].
= Artabotrys dahomensis Engl. & Diels, Notizbl. Königl. Bot. Gart. Berlin 2: 299, 1899. Syn. nov. Type. Benin: Dahome, Newton s.n., 1886: holotype: B[B 10 0153018].
= Artabotrys setulosus Mildbr. & Diels, Bot. Jahrb. Syst. 53. 447, 1915. Type. Cameroon. East Region, Mulundou, Mildbraed G.W.J. 4999, 26 Jan 1911: lectotype, here designated: HBG[HBG502545].
Democratic Republic of the Congo. Equateur; Likimi, Malchair L. 274, 20 Avr 1910: holotype: BR[BR0000008820297]
Liana, to 20 m tall, d.b.h. 3–5 cm. Indumentum of simple hairs; old leafless branches sparsely pubescent, young foliate branches hirsute, with erect hairs. Leaves: petiole 2–4 mm long, 1–2 mm in diameter, hirsute, slightly grooved, blade inserted on top of the petiole; blade 8–14 cm long, 3.5–5.5 cm wide, oblong to obovate, apex acuminate, acumen 0.5–1 cm long, base rounded to subcordate or obtuse, papyraceous, below pubescent when young and old with long appressed hairs, above glabrous when young and old, concolorous; midrib impressed, above glabrous when young and old, below pubescent when young and old; secondary veins 9 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 3 per inflorescence, hook-shaped peduncle 4–7 mm long; pedicel 3–5 mm long, ca. 1 mm in diameter, pubescent to sparsely hirsute; in fruit 3–15 mm long, ca. 2 mm in diameter, pubescent; bracts caduceus, not seen; sepals 3, valvate, free, 5–7 mm long, 2–4 mm wide, triangular, apex acute, base truncate, green, hirsute outside, glabrous inside, margins flat; petals free, sub equal, green turning yellow; outer petals 3, 10–20 mm long, 3–4 mm wide, very narrowly elliptic to linear, apex acute, base rounded to broad and concave, green turning bright yellow, margins flat, recurved inwards in vivo, densely appressed-pubescent outside, appressed-pubescent to glabrous inside; inner petals 3, valvate, 12–20 mm long, 2–3 mm wide, very narrowly elliptic to linear, apex acuminate to acute, base broad and concave, green turning bright yellow, margins flat, recurved inwards in vivo, appressed-pubescent outside, appressed-pubescent inside; stamens 60 to 70, in ca. 5 rows, ca. 1 mm long, broad; connective discoid, glabrous, green; staminodes absent; carpels free, 8 to 10, ovary ca. 1 mm long, stigma capitate, glabrous. Monocarps sessile, 5 to 11, 12–15 mm long, 5–6 mm in diameter, ellipsoid to fusiform, apex apiculate, glabrous, smooth, not ribbed, red when ripe; seeds 1 to 2 per monocarp, 10–12 mm long, 5 mm in diameter, ellipsoid to oblong; aril absent.
A mainly central African species, from Benin to Nigeria and Cameroon to the Republic of the Congo and in the Democratic Republic of the Congo; in Cameroon known from the Central and East regions.
A fairly common species, in secondary lowland or premontane rain forests. Altitude 400–900 m a.s.l.
nginda (pygmée–bibaya) (
Not evaluated.
None recorded.
Artabotrys rufus is distinguished by the hirsute pubescence of the young foliate branches, petioles, peduncles and flowering pedicels, its leaves that are elliptic, apiculate and less than 14 cm long, with long (1–2 mm) appressed hairs on the lower side of the leaf blade and a rounded to subcordate or obtuse base, a short peduncle (generally less than 7 mm long), petals with long dense brown hairs and smooth apiculate monocarps. The tertiary venation is also clearly visible forming clear loops towards the margins of the leaves.
Artabotrys rufus resembles A. velutinus being pubescent, but the pubescence of A. velutinus is not hirsute, with shorter hairs and more tomentose on the young foliate branches and petioles. The petals are also very similar being curved inwards, giving them the appearance of a tube. It is possible that both names are synonymous. Artabotrys rufus is also very close morphologically to the west African species A. hispidus Sprague & Hutch. by its hirsute pubescence and the shape of the leaves and flowers. It is also very possible that these names are synonymous. Several specimens from Cameroon where identified as A. hispidus, but upon closer look we have identified them as belonging to A. rufus. Several authors (
We here synonymize the names A. boonei and A. dahomensis with A. rufus. The former name was considered a synonym of A. velutinus (
Artabotrys thomsonii A flowering branch B outer petal, inner view C inner petal, inner view D flowering receptacle, petals removed E stamens, front and side views F carpel, side and detail of ovules G fruiting peduncle. Artabotrys rufus H flowering branch I flowering receptacle with one outer petal removed J outer petals, outer view and inner views K inner petals, inner and outer views L flowering receptacle, petals removed M stamen N carpel, side view and detail of ovule O fruiting peduncle A–G from Le Testu 9249 H–N from Hall 3193; 15 from Hall 3528. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
More detailed studies across the range of these species (A. hispidus, A. rufus, A. rupestris and A. velutinus) are needed to determine if there is one single widespread species from west to east Africa (possibly with different infraspecific taxa), or if there are several different species possibly grouped into a species complex (
Central Region: Fébé Mount, 3.91°N, 11.48°E, 30 March 1962, Breteler F.J. 2717 (A,BR,K,P,WAG,YA); Mont Mbam Minkon on trail 5 km from Nkol Nyada village On top of small hill, 3.97°N, 11.40°E, 21 March 2013, Couvreur T.L.P. 418 (MPU,WAG,YA); Nachtigal, 4.35°N, 11.63°E, 01 July 1964, de Wilde W.J.J.O 2779 (P,WAG,YA); Nkolbison, 3.88°N, 11.45°E, 02 November 1964, de Wilde W.J.J.O 3715 (BR,K,P,WAG); Left bank Sanaga river near Ferry Nachtigal ca 20 km N of Obala, 4.34°N, 11.64°E, 29 April 1965, Leeuwenberg A.J.M. 6011 (B,BR,C,K, MO,P,WAG,YA). East Region: Dimako, 4.38°N, 13.57°E, 02 August 1961, Breteler F.J. 1752 (P,YA); Bertoua 15 km along road to Deng Deng, 4.66°N, 13.63°E, 31 August 1961, Breteler F.J. 1825 (WAG); Bamékok (Batouri), 4.2°N, 14.15°E, 16 April 1962, Breteler F.J. 2825 (P,WAG); 60 km south of Yokadouma 5 km south of Maséa village, 3.10°N, 14.84°E, 06 March 2019, Couvreur T.L.P. 1211 (MPU,WAG,YA); A 25 km au NE de Bangé km 75 route Yokadouma-Moloundou, 3.02°N, 15.12°E, 25 May 1963, Letouzey R. 5147 (P,YA); Mbatika-Malen 20 km de Moloundou route Yokadouma 2.03°N, 15.22°E, 22 April 1971, Mezili P. 193 (P,YA); Moloundou near Lokomo Bumba and Bange, 2.08°N, 15.25°E, 26 January 1911, Mildbraed G.W.J. 4350 (B,HBG); Forêt inhabitée entre Yokaduma et Assobam, 3.52°N, 15.05°E, 24 April 1911, Mildbraed G.W.J. 4999 (B,HBG).
Nigeria. Cross River State; Old Calabar, Thomson W.C 25, Feb 1863: holotype: K[K000198871].
Liana, 30 m tall, d.b.h. 10–20 cm. Indumentum of simple hairs; old leafless branches sparsely pubescent to glabrous, young foliate branches sparsely pubescent. Leaves: petiole 1–10 mm long, ca. 2 mm in diameter, pubescent to glabrous, slightly grooved, blade inserted on top of the petiole; blade 7–20 cm long, 5–10 cm wide, elliptic to oblong, apex acute to abruptly acuminate, acumen 0.5–1 cm long, base rounded to obtuse, coriaceous, below sparsely pubescent to glabrous when young, sparsely pubescent to glabrous when old, above glabrous when young and old, concolorous; midrib impressed, above densely pubescent to pubescent when young and old, below sparsely pubescent to glabrous when young and old; secondary veins 7 to 14 pairs, glabrous above; tertiary venation intermediate. Individuals bisexual; inflorescences ramiflorous on old leafless branches, leaf opposed. Flowers with 9 perianth parts in 3 whorls, 30 to 90 per inflorescence, hook-shaped peduncle 20–35 mm long; pedicel 10–20 mm long, ca. 1 mm in diameter, pubescent; in fruit 10–25 mm long, ca. 2 mm in diameter, pubescent; bracts several, basal with one towards the lower half of pedicel, soon falling, basal bracts ca. 1 mm long, ca. 1 mm wide; upper bract ca. 2 mm long, ca. 2 mm wide; sepals 3, valvate, free, 2–3 mm long, 2–3 mm wide, triangular, apex acute, base truncate, pubescent outside, glabrous inside, margins flat; petals free, sub equal; outer petals 3, 10–20 mm long, 2–3 mm wide, elliptic to narrowly elliptic, apex rounded to obtuse, base broad and concave, margins flat, not folded inwards, pubescent outside, pubescent with a glabrous base inside; inner petals 3, valvate, 12–16 mm long, 2–3 mm wide, narrowly elliptic, apex acute, base broad and concave, margins flat, not folded inwards, pubescent outside, pubescent inside; stamens 30 to 35, in 2 to 3 rows, ca. 1 mm long, cuneiform; connective discoid, glabrous; staminodes absent; carpels free, 4 to 10, ovary ca. 1 mm long, stigma coiled, sparsely pubescent. Monocarps stipitate, stipes 10–25 mm long, 2–3 mm in diameter; monocarps 1 to 7, 15–25 mm long, 12–15 mm in diameter, ellipsoid to obovoid, apex rounded, glabrous, smooth, not ribbed, green when ripe; seeds 1 (to 2) per monocarp, 10–15 mm long, 8–9 mm in diameter, ellipsoid; aril absent.
A central African species; from Nigeria to Angola (Cabinda) and in the Democratic Republic of the Congo; in Cameroon known from the Central, East, Littoral, South, South-West and West regions.
A common species, in lowland and premontane secondary or primary rain forests, along forests margins and in logging areas. Altitude 100–1000 m a.s.l.
None recorded.
Not evaluated.
food : water drinken from stem; medicine: water/sap used for liver, genital stimulant/depressants, pregnancy, antiaborifacients.
Artabotrys thomsonii is easily distinguished by the densely pubescent upper midrib which is not found in any other Cameroonian species of Artabotrys.
There seems to be confusion around the type specimen of the name Artabotrys thomsonii. In the protologue,
The collection Thomson 25 is without doubt a Thomson collection with a printed label indicating “Collected at Old Calabar, by the Rev. W.C. Thomson” followed by “Presented by Professor Balfour, Dec; 1963”. Indeed, some specimens of Thomson were forwarded by Balfour to Kew (
For what it is worth, the fruit and leaves on Mann 2310 (K [K000198872] and P [P00363393]) appear to belong to Neostenanthera myristicifolia (Oliv.) Exell also present in Nigeria.
Central Region: Yaoundé, 3.87°N, 11.52°E, 1896, Zenker G.A. 697 (P). East Region: Near Dimako 28 km SW of Bertoua, 4.38°N, 13.57°E, 01 August 1961, Breteler F.J. 1725 (BR,K,P,U,WAG,YA); Bertoua 6 km along road to Batouri and Betaré-Oya, 4.58°N, 13.68°E, 30 August 1961, Breteler F.J. 1797 (U,WAG,YA); 67 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, 3.08°N, 14.67°E, 08 March 2019, Couvreur T.L.P. 1230 (MPU,WAG,YA); Palisco forest consession 15 km along main road into consession, 3.52°N, 13.54°E, 27 March 2015, Couvreur T.L.P. 751 (WAG,YA); 20 km environ à l’ENE de Mikel village situé à 85 km au N de Moloundou sur la route de Yokadouma 2.81°N, 15.24°E, 23 February 1971, Letouzey R. 10413 (K,P,YA); Ndongo (Dja-Molundou), 2.58°N, 15.29°E, 18 March 1973, Letouzey R. 12240 (K,P,YA); Djouo (Somalomo), 3.32°N, 12.93°E, 26 February 1962, Letouzey R. 4435 (K,P,YA); A 8 km au SSW de Koso (village situé à 60 km au SSW de Batouri), 3.93°N, 14.17°E, 29 July 1963, Letouzey R. 5533 (P,YA); Entre Badekok et Mpan (50 km ENE de Lomié), 3.22°N, 15.02°E, 05 August 1963, Letouzey R. 5548 (P,YA). Littoral Region: 8 km W of Massok, 4.13°N, 10.47°E, 27 March 1965, Leeuwenberg A.J.M. 5201 (B,BR,C,GC,K,MO,P,UC,WAG,YA). South Region: Bitya near R Ja, 3.02°N, 12.37°E, 01 November 1922, Bates G.L. 1763 (P); Djoum North East Nkout Base of ridge, 2.55°N, 12.80°E, 05 December 2014, Cheek M. 17781 (K,WAG); Elephant Mont, 2.8°N, 10.01°E, 24 May 2001, van Andel T.R. 3459 (KRIBI,WAG,YA); Campo-Ma’an area 2.73°N, 9.873°E, 16 August 2001, van Andel T.R. 3872 (KRIBI,U,WAG); Nkuambe, 3.26°N, 10.46°E, 01 December 1914, Zenker G.A. 489 (P,WAG). South-West Region: Bayang Mbo Wildlife Sanctuary after Mbu river, 5.35°N, 9.497°E, 27 March 2016, Couvreur T.L.P. 1020 (WAG,YA); Ekombe, 4.48°N, 10.87°E, 16 January 1987, Etuge M. 485 (MO,P,WAG). West Region: Près Bandounga à 40 km au NW de Ndikinimeki, 4.98°N, 10.55°E, 12 February 1972, Letouzey R. 11200 (P,YA).
= Artabotrys nigericus Hutch., Bull. Misc. Inform. Kew 10: 356, 1921. Type. Nigeria. Jos North, Naraguta, Lely H.V. 541, 17 Aug 1921: holotype: K[K000198866].
= Artabotrys stenopetalus Engl. Notizbl. Königl. Bot. Gart. Berlin 2: 300, 1899. Syn. nov. Type. Cameroon. South Region, Bipindi, Zenker G.A. 1222, 1896: holotype: B[B10 0154052]; isotypes: GOET[GOET005675]; HBG[HBG502541]; K[K000198862]; M[M0107912]; MO[MO-216860]; NY[NY00025832]; P[P00363384, P00363385]; S[S07-13456]; WAG[WAG0053235]; WU[WU0025887, WU0025888].
? Artabotrys stenopetalus var. parviflorus Pellegr., Mém. Soc. Linn. Normandie 26: 7, 1924. Type. Gabon. Nyanga, Tchibanga, Le Testu G.M.P.C. 1964, 6 Jan 1915: lectotype, sheet here designated: P[P00363379]; isotypes: EA[EA000002453, EA000002452]; K[K000198861]; LISC[LISC000375]; P[P00363378, P00363380].
= Artabotrys nitidus auct. Exell Jour. of Bot. 73 Supp. Polypet. Add.: 5, 1935 (non Diels) (specimens Gossweiler 5978 [COI00070298] and 7361 [COI00070297]).
Sierre Leone: Northern Region; Falaba, Scott Elliot G.F. 5137, 5 Mar 1892: holotype: K[K000198865]; isotype: B[B 10 0154055].
Liana, up to 10 m tall, d.b.h. up to 20 cm. Indumentum of simple hairs; old leafless branches sparsely pubescent, young foliate branches pubescent. Leaves: petiole 3–4 mm long, ca. 1 mm in diameter, pubescent to glabrous, grooved, blade inserted on the side of the petiole; blade 8–13 cm long, 4–5 cm wide, ovate to elliptic, apex acuminate to acute, acumen 1–1.5 cm long, base decurrent to acute, papyraceous, below pubescent to sparsely pubescent when young, glabrous when old, above pubescent when young, sparsely pubescent to glabrous when old, concolorous; midrib impressed, above glabrous when young and old, below densely pubescent to pubescent when young, glabrous to pubescent when old; secondary veins 8 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, 5 to 15 per inflorescence, hook-shaped peduncle 10–16 mm long; pedicel 2–10 mm long, ca. 1 mm in diameter, pubescent with appressed hairs; in fruit 10–20 mm long, ca. 1 mm in diameter, pubescent with appressed hairs; bracts 2, all basal, ca. 1 mm long, ca. 1 mm wide; sepals 3, valvate, free, 2–3 mm long, 2–3 mm wide, triangular, apex acute, base truncate, densely pubescent outside, glabrous inside, yellow and red at the base, margins flat; petals free, sub equal; outer petals 3, 5–15 mm long, 1–2 mm wide, linear, apex rounded, base broad and concave, margins flat but strongly recurved inwards forming a tube, densely pubescent outside, densely pubescent inside; inner petals 3, valvate, 5–15 mm long, 1–2 mm wide, linear, apex acute, base broad and concave, yellow and red at the base, margins flat but strongly recurved inwards forming a tube, tomentose outside, tomentose with a glabrous at base inside; stamens 15 to 22, in 2 rows, 1–2 mm long, oblong; connective discoid, glabrous; staminodes absent; carpels free, 7 to 12, ovary 1–2 mm long, stigma cylindrical, glabrous. Monocarps sessile, 4 to 9, 10–20 mm long, 10 mm in diameter, ellipsoid to obovoid, apex rounded, glabrous, smooth, faintly ribbed, color when ripe not seen; seeds 1 to 2 per monocarp, 5–7 mm long, 5 mm in diameter, flattened ellipsoid; aril absent.
A west and central African species, from Sierra Leone to Nigeria and Cameroon to the Republic of the Congo and in the Democratic Republic of the Congo; in Cameroon known from the Adamaoua, Central, East, North, South, South-West, and West regions.
A fairly common and widespread species; in secondary lowland premontane and montane rain forests, and in gallery forests occurring in the drier regions of the country, it is one of the few Annonaceae species (e.g. Monanthotaxis vulcanica; Xylopia africana) occurring above 2000 m in Cameroon. Altitude 400–2200 m a.s.l.
None recorded.
Not evaluated.
medicine
: water/sap/leaves used for liver, genital stimulant/depressants, pregnancy, antiaborifacients (as A. stenopetalus in
A Artabotrys velutinus B Brieya fasciculata C Cleistopholis glauca D Cleistopholis myristiciflora E Cleistopholis patens F Cleistopholis staudtii G Dennettia tripetala H Duguetia barteri I Duguetia confinis. White borders represent region limits in Cameroon; green patches represent protected areas (see methods and Suppl. material
Artabotrys velutinus is characterized by having leaf bases that are decurrent or acute, pubescent branches and petioles with appressed hairs and densely pubescent petals that are tightly recurved forming a tube. This latter character is also found in A. rufus, but in this species the pubescence is hirsute rather than appressed and the base of the leaves is rounded. Nevertheless, both species are very close morphologically.
Artabotrys velutinus belongs to a large species complex of pubescent species with petals that are tightly recurved forming a tube. We synonymize here the name A. stenopetalus. Another species name that could also be synonym is the east African species A. monteiroae Oliv. (which would be the older name). More studies are needed across the range of these species to better define the limits of these taxa.
The variety A. stenopetalus var. parviflorus is interesting. The type specimen (Le Testu 1964) appears to have a raised and grooved midrib on the upper side of the leaf blades (only seen as a scan on jstor), which is quite uncommon for African Annonaceae (
Adamaoua Region: Mbibol 40 km W de Ngaoundéré, 7.32°N, 13.58°E, 12 June 1977, Fotius G. 2660 (P,YA); Près Tekel (60 km NNO de Bagodo), 6.78°N, 13.17°E, 21 July 1966, Letouzey R. 7481 (P,YA); Boko 14 km Sud-Ouest de Ngaoundéré, 7.25°N, 13.5°E, 06 August 1981, van der Zon A.P.M. 1122 (WAG,YA). Central Region: Pentes orientales du mont Yangba (1473 m) près Nyafianga (42 km NNE de Bafia), 5.13°N, 11.35°E, 09 September 1966, Letouzey R. 7826 (K,P,YA). East Region: 27 km ENE de Mopwo (village situé au km 22 route Yokadouma-Batouri), 3.67°N, 15.08°E, 06 June 1963, Letouzey R. 5248 (K,P,YA). North Region: Mango, 8.42°N, 13.25°E, 21 July 1974, Fotius G. 2144 (YA). South Region: Rives du Ntem près du confluent de la Kye 16 km ESE d’Ambam, 2.25°N, 11.34°E, 01 February 1970, Letouzey R. 10040 (P); Bipindi, 3.08°N, 10.42°E, 3 April 1897, Zenker G.A. 1222 (B,P,WAG); Bipindi, 3.08°N, 10.42°E, 01 January 1913, Zenker G.A. 231 (P,U,WAG). South-West Region: Edip village to lake edip 2–3 km, 4.96°N, 9.65°E, 11 February 1998, Cheek M. 9143 (K,WAG,YA); Kodmin, 5°N, 9.7°E, 16 November 1998, Gosline W.G. 149 (K,YA); Bank of river Chide, 4.95°N, 9.72°E, 04 February 1998, Onana J.M. 523 (K,P,WAG,YA). West Region: Massif du Nkogam (2263 m) 25 km W de Foumban, 5.73°N, 10.67°E, 12 December 1974, Letouzey R. 13501 (P,YA).
Brieya fasciculata De Wild.
Same as species.
A genus with two species, one widespread and one restricted to northern Angola. One species in Cameroun, not endemic.
A genus easily confused with Piptostigma because of the characteristic inner petals being much longer than the outer ones in both genera, a unique feature among Cameroonian Annonaceae (
≡ Piptostigma fasciculatum (De Wild.) Boutique ex Fries, In Engler A., Prantl K. (eds) Die Natürlichen Pflanzenfamilien 17aII: 115–116, 1959.
= Piptostigma aubrevillei
Ghesq. ex. Aubrév.; Fl. For. Cote d’Ivoire 1: 98, 1936. Type. Ivory Coast. Mudjika, Aubréville A. 2115, 1932: lectotype, designated by
Democratic Republic of the Congo. Bas-Congo; Kingamu, Ganda sumi, de Briey J. 66, 14–16 Oct 1911: lectotype, sheet here designated: BR[BR-S.P.880319]; isotypes: BR[BR0000008803252, BR0000008803245, BR0000008803191].
Tree, 10–25 m tall, d.b.h. 16–50 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches pubescent. Leaves: petiole 2–5 mm long, ca. 2 mm in diameter, pubescent, grooved, blade inserted on top of the petiole; blade 12–24 cm long, 6–8 cm wide, obovate to oblanceolate, apex acuminate to obtuse, acumen 0.5–0.8 cm long, base cordate to obtuse, papyraceous, below glabrous to pubescent when young, glabrous to pubescent when old, above glabrous when young and old, discolorous, whitish below; midrib impressed, above pubescent when young and old, below pubescent when young and old; secondary veins 11 to 17 pairs, glabrous below; tertiary venation percurrent. Individuals bisexual; inflorescences ramiflorous on old leafless branches, axillary occurring on short peduncle-like bases 0–2 mm long. Flowers with 9 perianth parts in 3 whorls, 1 to 4 per inflorescence; pedicel 1–2 mm long, ca. 5 mm in diameter, pubescent; in fruit 15–25 mm long, 4–5 mm in diameter, glabrous; bracts 2, one basal and one upper towards the lower half of pedicel, basal bract 2–3 mm long, ca. 2 mm wide; upper bract ca. 1 mm long, ca. 2 mm wide; sepals 3, valvate, free, ca. 2 mm long, ca. 2 mm wide, ovate, apex acute, base truncate, brown, pubescent outside, glabrous inside, margins flat; petals free, outer petals shorter than inner; outer petals 3, sepal like, 1.5–2 mm long, 1.5 mm wide, ovate, apex acuminate, base truncate, light green, margins flat, pubescent outside, glabrous inside; inner petals 3, valvate, 38–108 mm long, 3–7 mm wide, linear, apex acute, base truncate, green, margins flat, pubescent inside, pubescent outside; stamens 30 to 40, in 4 to 5 rows, ca. 1 mm long, broad; connective discoid, glabrous, green; staminodes absent; carpels free, ca. 4, ovary ca. 2 mm long, stigma minute, densely pubescent. Monocarps sessile, 1 to 3, 42–46 mm long, 25–40 mm in diameter, ellipsoid, apex rounded, glabrous, smooth, fleshy, green when ripe; seeds 18 to 20 per monocarp, ca. 10 mm long, ca. 4 mm in diameter, ellipsoid; aril absent.
From Côte d’Ivoire to Democratic Republic of the Congo and Angola; in Cameroon known from East, South, Centre, Littoral and South-West regions.
A common species when present, in lowland to submontane rain forests in primary or secondary habitats. Altitude 250–810 m a.s.l.
baouéfou à grandes feuilles (french) (
Least Concern (LC) (
None recorded.
Brieya fasciculata is distinguished by its discolorous leaves, glaucous white below with a percurrent tertiary venation, its flowers occurring on reduced inflorescences with a short peduncle, the inner petals much longer than the outer ones, with the minute sepals and outer petals identical is shape and size, and its long green linear inner petals.
Central Region: Près Ngong (25 km NE d’Edéa), 3.75°N, 10.98°E, 12 December 1973, Letouzey R. 12345 (P,YA). East Region: 68 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, 3.08°N, 14.66°E, 08 March 2019, Couvreur T.L.P. 1231 (MPU,WAG,YA); Batéka Malen 20 km NE de Moloundou, 2.15°N, 15.35°E, 23 April 1971, Letouzey R. 10718 (P,YA); Entre Asip et Mang (60 km ENE de Lomié), 3.4°N, 14.17°E, 13 August 1963, Letouzey R. 5605 (P,YA). North-West Region: Kagwene, 6.10°N, 9.744°E, 13 June 2009, Ashworth J. 310 (K,YA). South Region: Campo Ma’an National Park 11 km on trail from Ebinanemeyong village on road 7 km from Nyabessan to Campo town, 2.48°N, 10.34°E, 11 February 2015, Couvreur T.L.P. 677 (WAG,YA); Abords de la Lobé à 50 km au SSE de Kribi et à 30 km à l’ENE de Campo, 2.51°N, 9.82°E, 23 March 1968, Letouzey R. 9132 (YA). South-West Region: Mudjika (Wudjika?), 4.29°N, 9.41°E, 01 January 1933, Aubréville A. 2115 (P); on trail from Ekongo village located 5 km before the entrance to Limbe 7 km on secondary road On flank of Mt Etinde 100 m in Mont Cameroon National Park, 4.07°N, 9.132°E, 16 October 2013, Couvreur T.L.P. 510 (WAG,YA); on trail from Ekongo village located 5 km before the entrance to Limbe 7 km on secondary road On flank of Mt Etinde 100 m in Mont Cameroon National Park, 4.07°N, 9.131°E, 16 October 2013, Couvreur T.L.P. 511 (WAG,YA); Rumpi mountains forest trail ca 5 km after Dikome Balue village ca 40 km north of Kumba, 4.93°N, 9.240°E, 10 January 2016, Couvreur T.L.P. 957 (WAG,YA); Kupe village Muanezum trail = Daniel Ajang’s Earthwatch rented area Mt 4.76°N, 9.666°E, 28 March 1996, Etuge M. 1844 (K); Kupe village, 4.77°N, 9.688°E, 28 November 1999, Gosline W.G. 234 (K); Njonji, 4.11°N, 9.016°E, 21 April 1997, Nning J. 385 (K,YA); Cameroon Mountain, 4.12°N, 9.028°E, 20 June 2001, van Andel T.R. 3732 (U,WAG).
Brieya fasciculata A flowering branch B flower, bottom view C flower, top view D detail of receptacle, all petals removed E longitudinal section of receptacle F stamen G carpel, side view and detail of ovules A from Aubréville 1500 B–G from Hallé 3166; 8 from Germain 2396. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
Brieya fasciculata A habit B leaf, upper side C leaf base, upper side D detail of leaf blade and venation; lower side E flower F detail of minute sepals and outer petal, in contrast to long inner petals G detail of receptacle H fruit with a single monocarp (others have fallen) A, H Couvreur 645, Mambe, Cameroon B, C no voucher, Rumpi Mountains, Cameroon D–G Couvreur 510, Mt Etinde, Cameroon. Photos Thomas L.P. Couvreur.
Cleistopholis glauca Pierre ex Engl. & Diels.
Trees, 15–35 m tall, d.b.h. 30–80 cm; stilt roots or buttresses absent, trunk white or brown. Indumentum of simple hairs when present, but species generally glabrous. Leaves: petiole 3–20 mm long, 1–3 mm in diameter, blade 4.5–31 cm long, 2–6.5 cm wide, elliptic to obovate to oblong, apex acuminate, base acute to rounded, discolorous, whitish below or concolorous; midrib sunken or flat; secondary veins 8 to 24 pairs; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, axillary, occurring or not on a short peduncle. Flowers with 9 perianth parts in 3 whorls, 2 to 9 per inflorescence; pedicel 10–25 mm long; in fruit 15–35 mm long; bracts 2 to 3, all basal, 1–2 mm long; sepals 3, valvate, free, 2–3 mm long, triangular to ovate, apex acute, sometimes rounded, base truncate; petals free; outer petals longer than inner; outer petals 3, valvate, 7–20 mm long, 2–7 mm wide, oblong to elliptic to obovate to linear, apex acute to rounded to obtuse, base truncate; inner petals 3, imbricate, 2–4 mm long, 2–4 mm wide, ovate to suborbicular, apex acute to obtuse, base truncate; stamens 20 to 40, in 3 to 4 rows, ca. 1 mm long, broad; connective discoid, glabrous; staminodes absent; carpels free, 10 to 24, ca. 1 mm long, stigma flat to capitate, glabrous. Monocarps stipitate, stipes 1–50 mm long, 3 to 18 monocarps, 15–30 mm long, 10–25 mm in diameter, globose to ellipsoid to obovoid, apex rounded, smooth, bumpy or constricted around the seeds, glabrous; seeds 1 to 2, 12–25 mm long, 8–12 mm in diameter, ellipsoid; aril absent.
A genus with four accepted species, two widespread, one known only from Cameroon and Gabon and one from Equatorial Guinea and possibly Cameroon. Three (four?) species in Cameroun, none endemic.
None to date, but partial treated in this present work and
1 | Trunk brown, petioles ca. 3 mm long; monocarps with stipes 49–50 mm long, thing, ca. 1 mm in diameter | C. myristiciflora |
– | Turk white, petioles generally 10–20 mm long (in C. patens can be as short as 3 mm too); monocarps with stipes 1–30 mm long, thick, 2–3 mm in diameter | 2 |
2 | Inflorescences with a distinct peduncule; monocarps ellipsoid to obovoid, drying smooth; lower leaf side glaucous, at least when fresh | C. glauca |
– | Inflorescences fasciculate, without a peduncule; monocarps globose to bilobed; lower leaf side not glaucous | 3 |
3 | Outer petals linear, 15–20 mm long; monocarps drying smooth, not bumpy; petiole 10–15 mm long | C. staudtii |
– | Outer petals oblong, elliptic or obovate, 7–12 mm long; monocarps bumpy, constricted around the seeds; petiole 3–12 mm long | C. patens |
= Cleistopholis grandiflora De Wild., Ann. Mus. Congo Belge, Bot. sér. 5, 1(1): 39, 1903. Type. Democratic Republic of the Congo. Kinshasa, Région de Kimuenza, Gérard P. s.n., Oct 1900: lectotype, sheet here designated: BR[BR0000008820327]; isotype: BR[BR0000008820655].
= Cleistopholis bequaertii De Wild., Pl. Bequaert. i.; 464, 1922. Type. Democratic Republic of the Congo. Nord-Kivu, Walikale - Lubutu, Bequaert J.C.C. 6624, 15 Jan 1915: lectotype, sheet here designated: BR[BR0000008820402]; isotype: BR[BR0000008820396].
Gabon. Estuaire; Libreville, Klaine T.-J. 376, Apr 1896: holotype: B[B 10 0154073]; isotypes: K[K000198885, K000198884]; P[P00362650, P00362649, P00362652]; MPU[MPU011662].
Tree, 10–35 m tall, d.b.h. 80 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 10–20 mm long, 1–2 mm in diameter, glabrous, grooved, blade inserted on the side of the petiole; blade 5–15 cm long, 2–5 cm wide, oblong to elliptic, apex acuminate, acumen 0.5–1.5 cm long, base decurrent to cuneate, subcoriaceous, below glabrous when young and old, above glabrous when young and old, discolorous, whitish below; midrib impressed, above glabrous when young and old, below glabrous when young and old; secondary veins 8 to 15 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on young and old leafless branches, axillary, peduncule distinct 2–10 mm long. Flowers with 9 perianth parts in 3 whorls, 2 to 8 per inflorescence; pedicel 10–18 mm long, ca. 1 mm in diameter, sparsely pubescent; in fruit 15–35 mm long, 2–3 mm in diameter, glabrous; bracts 1 to 3, all basal, basal bracts 1–2 mm long, 2 mm wide; sepals 3, valvate, free, ca. 2 mm long, ca. 2 mm wide, triangular to ovate, apex acute, base truncate, green, glabrous outside, glabrous inside, margins flat; petals free; outer petals 3, 10–15 mm long, 5–7 mm wide, oblong to elliptic, apex rounded, base truncate, green, margins flat, glabrous outside, glabrous inside; inner petals 3, imbricate, 2–4 mm long, 2–4 mm wide, ovate to orbicular, apex rounded, base truncate, green, margins flat, glabrous outside, glabrous inside; stamens 20 to 30, in 3 to 4 rows, ca. 1 mm long, broad; connective discoid, glabrous, green; staminodes absent; carpels free, 12 to 24, ovary ca. 1 mm long, stigma flat, glabrous. Monocarps stipitate, stipe 18–30 mm long, 3–4 mm in diameter; monocarps 3 to 8, 18–30 mm long, 10–15 mm in diameter, obovoid, apex rounded, glabrous, finely warty, not bumpy, green when ripe; seed (1 to) 2 per monocarp, 15–25 mm long, 10–12 mm in diameter, ellipsoid; aril absent.
Central Africa; from Cameroon to Democratic Republic of the Congo; in Cameroon known from South, Central, Littoral, South-West and East regions, with one collection from Adamaoua region.
Cleistopholis glauca A flowering branch, note long petioles B detail of inflorescence C flower, top view, note imbricate inner petals D internal petal, inner view E detail of receptacle, top view F stamen, front and side views G carpel, side view and view of ovules H fruit, note smooth monocarps I longitudinal section of seed A–G from Le Testu 8786; 8 from Klaine 41. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
A very common species, mainly growing as a pioneer species in disturbed areas and along forest margins. Altitude 100–1200 m a.s.l.
None recorded.
Least Concern (LC) (
None recorded.
Closely resembles C. patens, see below for differences.
Adamaoua Region: A 80 km au SO de Banyo-Plaine Tikar, 6.75°N, 11.82°E, 27 June 1969, Biholong M. 219 (P,YA). Central Region: Feup (Yaoundé), 3.87°N, 11.52°E, 01 July 1917, Chevalier A.J.B. 33473 (P,WAG); 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. 430 (MPU,WAG,YA); Avom, 3.87°N, 11.52°E, 01 January 1935, Foury P. 57 (P). East Region: Bertoua, 4.58°N, 13.68°E, 17 February 1960, Letouzey R. 3036 (P,YA); Région de Moloundou, 2.05°N, 15.17°E, 01 August 1949, SRFK 1372 (P,YA). Littoral Region: 18 km SEof Douala along road to Ndonga (=old to Edea), 4.05°N, 9.71°E, 20 August 1965, Leeuwenberg A.J.M. 6467 (B,BR,C,GC,K,L,LUAI,MO,P,UC,WAG,YA). South Region: 20 km from Kribi Lolodorf road, 3.05°N, 10.05°E, 09 June 1969, Bos J.J. 4784 (B,BR,K,LD,LM,MO,P,POZG,WAG,YA); on road between Campo and Kribi, 2.62°N, 9.847°E, 16 February 2012, Couvreur T.L.P. 389 (WAG,YA); Rocher du Loup km 36 road Kribi-Campo, 2.61°N, 9.85°E, 06 January 1983, de Kruif A.P.M. 1046 (MO,WAG,YA); Campo-Ma’an area 2.73°N, 9.873°E, 13 August 2001, van Andel T.R. 3846 (KRIBI,WAG,YA). South-West Region: Mungo River F.R., 4.78°N, 9.566°E, 02 December 1999, Cheek M. 10229 (K,MO,P,WAG,YA); Ntali, 5.25°N, 9.576°E, 01 December 2000, Etuge M. 4873 (K); Nyandong-forest above village, 4.98°N, 9.585°E, 20 March 2003, Etuge M. 4917 (K); Forest and forest relictss near Mundemba, 4.96°N, 8.916°E, 16 January 1985, Thomas D.W. 4200 (P); Near Mundemba town, 4.96°N, 8.916°E, 12 May 1986, Thomas D.W. 6121 (MO,P,WAG,YA).
Cameroon. South Region(?) or Equatorial Guinea. Río Muni; Campo-Gebiet; Bebao[i?], Weg nach Olonga [Manga?], Tessmann, G. 767, 6 Jan. 1909: holotype: B[B 10 0154074].
Tree, to 19 m tall, d.b.h. to 16 cm; stilt roots or buttresses absent, trunk brown. Indumentum of simple hairs (?); old leafless branches glabrous, young foliate branches pubescent (?). Leaves: petiole ca. 3 mm long, ca. 1 mm in diameter, pubescent (?), grooved, blade inserted on the side of the petiole; blade 8–12 cm long, ca. 4 cm wide, elliptic to oblong, apex acuminate, acumen ca. 1.5 cm long, base cuneate, papyraceous, below pubescent when young (?), glabrous when old, above glabrous when young and old, concolorous; midrib impressed, above glabrous when young and old, below pubescent when young (?), glabrous when old; secondary veins 10 to 13 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on young and old leafless branches, axillary, peduncule generally absent. Flowers with 9 perianth parts in 3 whorls, 4 to 9 per inflorescence, pedicel 15–27 mm long, ca. 1 mm in diameter, sparsely pubescent (?); in fruit ca. 20 mm long, 2–3 mm in diameter, glabrous (?); bracts not seen (soon falling ?); sepals 3, valvate, free, 1.5–2 mm long, ca. 1.5 mm wide, circular, apex rounded, base rounded, color unknown, pubescence not observed, margins flat; petals free; outer petals 3, valvate, 5–7 mm long, 2–3 mm wide, broadly elliptic to oblong, apex rounded, base truncate, color unknown, margins flat or wavy when dry, pubescence not observed; inner petals 3, imbricate (?), dimensions and shape not observed; stamens number not counted, row number not counted, ca. 1 mm long, broad; connective discoid, pubescence not observed, color unknown; staminodes absent (?); carpels free, 10 to 15 (?, possibly more based on the number of monocarps counted), ovary ca. 1.5 mm long, stigma flat, glabrous. Monocarps stipitate, stipes 49–50 mm long, ca. 1 mm in diameter; monocarps 17 to 18, 10–15 mm long, 10–15 mm in diameter, globose, apex rounded, glabrous (?), bumpy; seeds not seen.
Equatorial Guinea and Cameroon (?); if present in Cameroon then from South region.
A rare species, in primary submontane tierra firme forest. Altitude 750–850 m a.s.l. (altitude in Equatorial Guinea).
Akom (Equatorial Guinea) (
No assessed, but probably CR.
None recorded.
Cleistopholis myristiciflora (initially known only from the type; Tessmann 767 (B, but see below)) was originally described as being from Cameroon (
The taxonomic affinities of this species were unclear for some time but suggested to be conspecific with either C. patens or C. staudtii.
Recently, we located two specimens collected from Monte Alén in Guinea Equatorial (Senterre & Obiang 2939, 3699, BRLU) which appear to belong to C. myristicifolia (identified as such by B. Senterre). The leaves match the description and the type specimen, especially the shape and the length of the petiole being shorter (ca. 3 mm) than in the other species (> 3 mm). One specimen (Senterre & Obiang 2939), is in fruit. This single fruit is morphologically quite different than those of the other species in Cleistopholis. It is partially described here for the first time. The main difference is the length and diameter of the stipes being much longer and thinner than those from the other three species (49–50 mm long and ca. 1 mm in diameter versus 1–30 mm and 2–3 m in diameter). The number of monocarps appears to be higher with 17 to 18 counted in Senterre & Obiang 2939, versus 3 to 8 in the other species. These observations strongly support the hypothesis that C. myristiciflora is indeed a distinct and valid species. In terms of its ecology, observations from Senterre & Obiang suggest it to occur in primary submontane rain forests, occurring on gentle slopes or top of small mountains.
One specimen collected from southern coastal Gabon (Bergen 217 [WAG.1379499, WAG.1379500]) at 10 m a.s.l. was identified as C. myristiciflora by M. Fero. It is true that the inflorescences and flowers could potentially match, but the size of the tree (8 m) and leaves are different having a long petiole (> 3 mm). The ecology is also different than described above occurring along the coast on laterite soil. For now we do not consider this specimen as part of C. myristiciflora and Bergen 217 could potentially represent an undescribed Gabonese coastal species, as has been done recently in other Annonaceae genera such as Greenwayodendron littorale Lissambou, Dauby & Couvreur (
≡ Oxymitra patens Benth., Trans. Linn. Soc. London 23(3): 472, 1862.
= Cleistopholis brevipetala Exell, J. Bot. 70 (Suppl. 1): 208, 1932. Type. Angola. Cabinda, Gossweiler J. 6082, 31 Dec 1915: holotype: BM[BM000546899]; isotypes: COI[COI00004874]; LISC[LISC000068, LISC000071, LISC000069, LISC000070].
= Cleistopholis klaineana Pierre ex Engl. & Diels, Monogr. Afrik. Pflanzen.-Fam. 6: 35, 1901, Cleistopholis patens var. klaineana Pellegr., Bull. Soc. Bot. France: 57, 1949.. Type. Gabon. Estuaire, Libreville, Klaine T.J. 345, 1896: holotype: B[B 10 0154075]; isotypes: P[P00362660, P00362656].
= Cleistopholis lucens De Wild., Pl. Bequaert. i. 465, 1922. Type. Democratic Republic of the Congo. Nord-Kivu, entre Walikale et Lubutu, Bequaert J. 2774, 22 Fev 1922: holotype: BR[BR0000008820426].
= Cleistopholis pynaertii De Wild., Bull. Jard. Bot. État Bruxelles 4: 387, 1914. Type. Democratic Republic of the Congo. Equateur, Eala, Pynaert L.A. 1083, 1 Fev 1907: lectotype, sheet here designated: BR[BR0000008820419]; isotypes: BR[BR0000008820761, BR0000008820754].
= Cleistopholis verschuereni De Wild., Bull. Jard. Bot. État Brux. 4: 387, 1914. Type. Democratic Republic of the Congo. Manie Malela, Verschueren R. 358, Fev 1913: lectotype, sheet here designated: BR[BR0000008820389]; isotype: BR[BR0000008820433].
Sierra Leone. Northern Region; Bagroo River, Mann G. 828, Apr 1861: lectotype, sheet here designed: K[K000880416]; isotypes: K[K000880417]; P[P00362653].
Tree, up to 30 m tall, d.b.h. up to 60 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole (3–)10–20 mm long, ca. 2 mm in diameter, glabrous, grooved, blade inserted on the side of the petiole; blade 4.5–31 cm long, 2.5–6 cm wide, oblong to narrowly oblong or oblanceolate to narrowly oblanceolate, apex acute to acuminate, acumen 1–1.5 cm long, base cuneate to rounded, coriaceous, above glabrous when young and old, shiny when dry, below glabrous when young and old, green, concolorous; midrib impressed, above glabrous when young and old, below glabrous when young and old; secondary veins 10 to 24 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on young or old leafless branches, axillary, peduncle absent. Flowers with 9 perianth parts in 3 whorls, 2 to 9 per inflorescence; pedicel 10–25 mm long, ca. 1 mm in diameter, glabrous; in fruit 15–30 mm long, 2–3 mm in diameter, glabrous; bracts 1 to 3, all basal, 1–2 mm long, ca. 2 mm wide; sepals 3, valvate, free, 2–3 mm long, ca. 2 mm wide, triangular to ovate, apex acute, base truncate, green, glabrous outside, glabrous inside, margins flat; petals free; outer petals longer than inner; outer petals 3, 7–12 mm long, 2–4 mm wide, obovate to oblong, apex obtuse, base truncate, green, margins flat, glabrous outside, glabrous inside; inner petals 3, imbricate, 3–4 mm long, 2–3 mm wide, ovate to suborbicular, apex rounded, base truncate, green, margins flat, glabrous outside, glabrous inside; stamens 25 to 30, in 3 to 4 rows, ca. 1 mm long, broad; connective discoid, pubescent, green; staminodes absent; carpels free, ca. 10, ovary ca. 1 mm long, stigma capitate, glabrous. Monocarps stipitate (sometimes shortly so), stipes 3–12 mm long, 3–4 mm in diameter; monocarps 3 to 6, 15–23 mm long, 11–25 mm in diameter, ellipsoid to globose, apex rounded, glabrous, finely warty, constricted around seeds, bumpy; seeds 1 to 2 per monocarp, ca. 12 mm long, 8–9 mm in diameter, ellipsoid; aril absent.
In West Africa, Senegal, Sierra Leone to Nigeria, and Central Africa from Cameroon to Uganda; in Cameroon known from South, Central, Littoral, South-West and East regions.
Cleistopholis staudtii A flowering branch B flower, side view, note linear outer petals C stamen, side view D carpel, side view and detail of ovules. Cleistopholis patens E flowering branch F flower, side view G stamen, side view H carpel, side view and detail of ovules I fruit; note bumpy monocarps A–D from Letouzey 4149 E–J from Chevalier 22379. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
A very common species, mainly growing as a pioneer species in disturbed areas and along rain forest margins. Altitude 0–600 m a.s.l.
avom, sobu (pygmée Bibaya) (
Least Concern (LC) (
medicine : bark used as pain-killers, against pulmonary troubles; leaves as vermifuges, fabrifuges; construction: building materials, furniture; dyes and tannins: astringents, insecticides, arachnicides; products: fiber, pulp, wood fire, exudations-gums, resins, farming, forestry, hunting and fishing apparatus, household, domestic and personal items, pastimes-carving, musical instruments; social: religion, superstitions, sayings, aphorisms.
Cleistopholis patens closely resembles C. glauca by the shape and aspect of the flowers and the overall vegetative characters. However, both species differ by their inflorescences being pedunculate in C. glauca versus sessile, the leaves green below versus glaucous in C. glauca and the monocarps being bumpy and constricted around the seeds when dry versus to smooth and not bumpy when dry in C. glauca.
Central Region: Avome, 3.87°N, 11.52°E, 13 August 1945, Aubréville A. 41 (P). East Region: 76 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. 1202 (MPU,WAG,YA); Dja Reserve, 3.17°N, 13.18°E, 07 October 1994, Fogiel M.K. 947 (P); Deng Deng, 5.2°N, 13.51°E, 01 July 1939, Jacques-Félix H. 4630 (P,WAG); Rives de la Boumba à 14 km à l’WSW de Kinsassa village situé à 65 km au NNE de Moloundou sur route de Yokadouma 2.58°N, 15.26°E, 07 March 1971, Letouzey R. 10523 (P,YA). Littoral Region: km 19 Loum-Yabassi 3 km N of Solé, 4.61°N, 9.8°E, 30 December 1971, Leeuwenberg A.J.M. 9032 (YA,WAG). South-West Region: Southern Bakundu Forest Reserve, 4.55°N, 9.433°E, 15 June 1960, Adebusuyi J.K. 44049 (WAG); S Bakundu Forest 3 km from Kindongi Camp (8 km from road), 4.49°N, 9.374°E, 02 May 1972, Leeuwenberg A.J.M. 9785 (B,BR,C,K,M,MO,P,WAG,YA); 2 km W of km 21 Kumba-Victoria road, 4.46°N, 9.483°E, 04 May 1972, Leeuwenberg A.J.M. 9828 (B,BR,C,K,LD,M,MO,P,WAG,YA); Bibundi, 4.21°N, 8.988°E, 08 November 1928, Mildbraed G.W.J. 10640 (K); Korup National Park, 4.88°N, 8.783°E, 22 July 1983, Thomas D.W. 2329 (MO,P,WAG,YA); Limbe (Victoria), 4.01°N, 9.133°E, 25 October 1997, van der Burgt X.M. 219 (KRIBI,WAG).
≡ Oxymitra staudtii Engl. & Diels, Notizbl. Königl. Bot. Gart. Berlin 2: 297, 1899.
= Polyalthia (?) crassipes Engl. Bot. Jahrb. Syst. 34: 477, 1907. Type. Cameroon. South Region, Bipindi, Zenker G.A. 2454a, 1902: holotype B destroyed, lectotype here designated: P[01988941]; isolectotypes: MO[MO-2500050]; P[P01988940, P01988942].
Cameroon. South-West Region; Johann-Albrechtshöhe [Kumba], Staudt A. 957, 1896: holotype: B[B 10 0154076]; isotypes: BM[BM000546890]; K[K000105343]; LE[LE00012452]
Tree, 15–30 m tall, d.b.h. up to 30 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 10–20 mm long, ca. 3 mm in diameter, glabrous, grooved, blade inserted on the side of the petiole; blade 9–17 cm long, 3–6.5 cm wide, obovate, oblong to elliptic, apex acuminate, acumen ca. 1 cm long, base rounded to acute, coriaceous, below glabrous when young and old, above glabrous when young and old, discolorous, whitish below, midrib impressed, above glabrous when young and old, below glabrous when young and old; secondary veins 10 to 12 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on young and old leafless branches, axillary, peduncle absent. Flowers with 9 perianth parts in 3 whorls, 2 to 3 per inflorescence; pedicel 12–20 mm long, ca. 1 mm in diameter, glabrous; in fruit 15–35 mm long, ca. 3 mm in diameter, glabrous; bracts 1 to 3, all basal, 1–2 mm long, 2 mm wide; sepals 3, valvate, free, ca. 2 mm long, ca. 2 mm wide, triangular to ovate, apex acute to rounded, base truncate, green, glabrous outside, glabrous inside, margins flat; petals free, outer petals longer than inner; outer petals 3, 15–20 mm long, 2–3 mm wide, linear, apex acute, base truncate, green, margins flat, glabrous outside, glabrous inside; inner petals 3, imbricate, 2–2.5 mm long, 3–3.5 mm wide, ovate, apex acute, base truncate, claw mm long, green, margins flat, glabrous outside, glabrous inside; stamens 30 to 40, in 3 to 4 rows, ca. 1 mm long, broad; connective discoid, pubescent, green; staminodes absent; carpels free, 15 to 22, ca. 1 mm long, stigma capitate, glabrous. Monocarps stipitate to sessile, stipes when present to 10 mm long, 3–4 mm in diameter, 3 to 8 monocarps, 15–20 mm long, 12–15 mm in diameter, globose, apex rounded, glabrous, striate, bumpy, constricted around seeds; seeds 1 to 3 per monocarp, 10–15 mm long, 7–10 mm in diameter, ellipsoid; aril absent.
From Cameroon to Gabon; in Cameroon known from South, Central, Littoral and South-West regions.
A common species when present, in lowland rain forests in primary or secondary habitats. Altitude 50–1000 m a.s.l.
None recorded.
Least Concern (LC) (
construction
: bark for building materials (
Easily distinguished from C. glauca and C. patens by its linear and acute outer petals, in contrast to short and rounded outer petals in the latter two. It is quite hard to distinguish these species based on sterile material alone.
Cleistopholis glauca A habit, note long drooping branches B flowers, top view, inner petals imbricate, not opened yet C fruit, note smooth monocarps D flowering branch, inner petals open revealing receptacle. Cleistopholis patens E trunk F base of leaf blades, upper view. Cleistopholis staudtii G leaf, top view, note long petioles H leaf, lower view I flowering branch, note long linear outer petals A–C Sosef 2036, Gabon D Couvreur 389, Ebodjé, Cameroon E, F Couvreur 1202, Maséa, Cameroon G–I Couvreur 570, Gabon. Photos Thomas L.P. Couvreur.
Central Region: Ca 50 km NW of Eséka W of Yaoundé, 3.65°N, 10.78°E, 25 November 1963, de Wilde W.J.J.O 1342 (B,BR,C,DES,L,LD,MO,P,U,WAG,YA); Ottotomo Forest Reserve, 3.65°N, 11.31°E, Service Forestier du Cameroun 32 (P). South Region: ca 15 km from Kribi 1 km S of Ebolowa road, 2.85°N, 10.01°E, 20 February 1970, Bos J.J. 6383 (P,WAG); Station de cacaoyer de N’koemvone 14 km On the road from Ebolowa to Ambam, 2.81°N, 11.13°E, 02 February 1975, de Wilde J.J.F.E 7947 (B,BR,K,MO,P,U,WAG,YA); Près de Bella (45 km NE de Kribi), 3.25°N, 10.2°E, 25 January 1962, Letouzey R. 4149 (P,YA); Bipindi, 3.08°N, 10.41°E, 01 January 1900, Zenker G.A. 2264 (L,P,WAG); Bipindi, 3.08°N, 10.41°E, 01 January 1902, Zenker G.A. 2454 (L,P,WAG); Bipindi, 3.08°N, 10.41°E, 01 January 1902, Zenker G.A. 2495 (L,P,WAG); Bipindi, 3.08°N, 10.41°E, 01 January 1913, Zenker G.A. 4669 (L,P); Bipindi, 3.08°N, 10.41°E, 01 January 1913, Zenker G.A. 4880 (L,P). South-West Region: Ekundu Kundu, 5.16°N, 8.874°E, 11 April 1996, Cable S. 1825 (K,YA); Kupe Mount Path to Kupe Rock, 4.75°N, 9.686°E, 24 November 1995, Cheek M. 7915 (K,P,WAG); Muanezum trail from Kupe village towards Daniel Ajang’s area 4.77°N, 9.708°E, 18 July 1996, Etuge M. 2884 (K,MO,P,WAG); Just outside Kupe village going north, 4.77°N, 9.688°E, 29 November 1999, Gosline W.G. 240 (K,MO,P,WAG,YA); Korup National Park, 5.06°N, 8.855°E, 05 December 1997, Kenfack D. 984 (MO,P,WAG); Environs of Kumba farmed land and scrub with scattered trees, 4.63°N, 9.433°E, 01 March 1984, Thomas D.W. 3271 (MO,WAG,YA).
Dennettia tripetala Baker f.
Same as species.
A genus with a single widespread species from West Africa (Sierra Leone to Nigeria) and in Cameroon. One species in Cameroon, not endemic.
Dennettia was first described by
≡ Uvariopsis tripetala (Baker f.) G.E.Schatz, Novon 13(4): 447, 2003.
Nigeria. Edo State; Benin City, Dennett R.E. 44, 1 Jan 1907: lectotype, designated by
Shrub to small tree, 2–5 m tall, d.b.h. unknown; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches glabrous to sparsely pubescent. Leaves: petiole 2–5 mm long, 1–2 mm in diameter, glabrous, slightly grooved, blade inserted on top of the petiole; blade 7.2–15.5 cm long, 3–6.8 cm wide, elliptic, apex attenuate to acuminate, acumen 0.6–1.3 cm long, base acute to decurrent, subcoriaceous, below glabrous when young and old, above glabrous when young and old; midrib sunken or flat, above glabrous when young and old, below glabrous when young and old; secondary veins 5 to 10 pairs per side, glabrous above; tertiary venation reticulate. Individuals bisexual, inflorescences ramiflorous on old leafless branches, axillary. Flowers with 6 perianth parts in 2 whorls, 1 to 4 per inflorescence; pedicel 4–9 mm long, 1–2 mm in diameter, pubescent; in fruit 5–15 mm long, 2–3 mm in diameter, glabrous to pubescent; bracts 1 to 3, all basal 0.5–2 mm long, 1.5–2 mm wide; sepals 3 (rarely 2), valvate, basally fused, 1–3 mm long, 1.5–4 mm wide, triangular, apex acute, base truncate, brown, pubescent outside, glabrous inside, margins flat; petals 3 (rarely 4, see notes), free, 7–14 mm long, 6–10 mm wide, broadly ovate, apex obtuse, base truncate, margins flat, pubescent outside, glabrous inside; stamens ca. 150, in 10 to 20 rows, 0.5–1 mm long, oblong; connective reduced or absent, glabrous; staminodes absent; carpels free, 8 to 30, ovary ca. 2–4.5 mm long, stigma globose, pubescent. Monocarps stipitate, stipes 1–3 mm long, ca. 1 mm in diameter; monocarps 1 to 8, 11–32 mm long, 5–15 mm in diameter, ovoid to oblong, apex rounded, glabrous to sparsely pubescent, verrucose, wrinkled; seeds 4 to 12 per monocarp, 4–10 mm long, 11–14 mm in diameter, ellipsoid; aril absent.
Dennetia tripetala A flowering branch B flower, petals remove, 1 sepal removed, showing receptacle C receptacle with stamens and stigmas D stamen, front view E carpel, front view F carpel, longitudinal section showing ovules. Material of drawings unknown. Author of drawings unknown, taken from
Mainly a West African species from Sierra Leone to Cameroon; in Cameroon known from the South-West region.
an uncommon species; in lowland to premontane primary or secondary rain forests. Altitude 0–1000 m a.s.l.
Bushpèpè (Westphal 9932, Pidgin English(?)); Pepperfruit (english)
Least Concern (LC) (
food
: fruit for sauces, condiments, spices, flavorings (pepper); medicine: cough, fever, toothache, diarrhea, diabetes, nausea (
Dennettia tripetala is unique in Cameroonian by being a tree with bisexual flowers having three sepals and three petals. This species resembles Uvariopsis congensis and Uvariopsis zenkeri (Uvariopsis being a genus were Dennettia was once part off, see above,
It has been reported that some D. tripetala specimens had 2 sepals and 4 petals (
South-West Region: Missellele, 4.12°N, 9.448°E, Box H.E. 3556 (BM,K); Limbe (Victoria), 4.07°N, 9.189°E, 01 April 1929, Maitland T.D. 626 (K); Buea area 4.2°N, 9.183°E, 01 January 1930, Maitland T.D. s.n. (K[K000105532]); Likomba-Pflanzung 15–35 km NE von Victoria [Limbe], 4.1°N, 9.333°E, 18 October 1928, Mildbraed G.W.J. 10515 (K); Ngandjo on Kumba Mbonge road, 4.55°N, 9.4°E, 25 February 1986, Thomas D.W. 5661 (K); Market of Victoria, 4.01°N, 9.2°E, 04 April 1978, Westphal E. 9932 (WAG).
= Pachypodanthium Engl. & Diels, Notizbl. Königl. Bot. Gart. Berlin 3: 55, 1900.
Duguetia lanceolata A.St.-Hil. (a Brazilian species).
Trees, 8–50 m tall, d.b.h. up to 50 cm; stilt roots or buttresses absent. Indumentum of stellate or fasciculate hairs. Leaves: petiole 1–10 mm long, 2–6 mm in diameter; blade 7–34 cm long, 3–8 cm wide, ovate to elliptic to obovate, apex acuminate to acute, acumen 0.5–1 cm long, base cordate to acute, discolorous, whitish below or concolorous; midrib sunken or flat; secondary veins 8 to 25 pairs; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on young and old leafless branches, leaf opposed or supra-axillary. Flowers with 9 perianth parts in 3 whorls, 2 to 5 per inflorescence; pedicel 3–22 mm long; in fruit 1–50 mm long; bracts 2, one basal and one upper in the lower half of pedicel, basal bract 7–12 mm long, upper bract similar than basal one; sepals 3, valvate, free, 6–15 mm long, apex acute, base truncate; petals free, outer petals longer than inner to sub equal; outer petals 3, valvate, free, 10–30 mm long, 4–10 mm wide, elliptic to ovate, apex acute to acuminate, base truncate; inner petals 3, imbricate, free, 4–20 mm long, 4–9 mm wide, elliptic to ovate to obovate, apex acute to acuminate, base truncate; stamens numerous, 1–2 mm long, broad; connective discoid; staminodes absent; carpels free, 50 to 125, 1.5–3.5 mm long, stigma globose. Fruit pseudosyncarpous; carpels sessile, connate or free, 60 to 125 carpels, 15–55 mm long, 7–30 mm in diameter, globose to ovoid to ellipsoid, apex domed-shaped to acute to apiculate; seed 1, 7–20 mm long, 4–13 mm in diameter, obovoid to ellipsoid; aril present, rudimentary.
A genus of 94 species, with a disjunct distribution, 89 in the Neotropics and 4 in Africa, but absent from Madagascar. All four African species are known from Cameroon, one endemic.
This genus of trees is characterized by stellate hairs on its leaves and pseudosyncarpous fruits. The only other tree genus with stellate hairs in Cameroon is Annickia, but the latter has a yellow slash and apocarpous fruits with clearly stipitate monocarps.
1 | Leaf blabes narrowly elliptic to narrowly oblong, 4 to 6 times longer than wide and leaves verruculose | D. confinis |
– | Lower side of the leaves sparsely to densely covered with appressed (flattened), stellate hairs; fruit globose or ovoid | 2 |
2 | Leaf base generally cordate, mid rid furrowed above, secondary veins weakly distinct | D. staudtii |
– | Leaf base acute, mid rid not furrowed above, secondary veins clearly distinct, forming loops | 3 |
3 | Inflorescences forming on a short peduncle in leafless parts of branches; fruiting carpels totally fused, areoles domed-shaped; seeds brown | D. barteri |
– | Inflorescences not forming a short peduncle in leafy part of branches; fruiting carpels basally fused, areoles obovoid to deltoid; seeds black | D. dilabens |
≡ Annona barteri Benth., Trans. Linn. Soc. London 23(3): 477, 1862.
= Pachypodanthium staudtii var. letestui Pellegr., Bull. Soc. Bot. France 95: 137, 1948. nom. illeg.
= Pachypodanthium tessmannii R.E.Fr., nom. nud.
Nigeria. Anambra state; Onitsha, Barter C. 445, 1858: holotype: K[K000198875].
Tree, 8–40 m tall, d.b.h. 40–60 cm; stilt roots or buttresses absent. Indumentum of stellate or fasciculate hairs; old leafless branches sparsely pubescent to glabrous, young foliate branches sparsely pubescent to densely pubescent. Leaves: petiole 3–7 mm long, 2–3 mm in diameter, densely to sparsely pubescent, grooved, blade inserted on the side of the petiole; blade 10–23 cm long, 3–7 cm wide, ovate to elliptic, apex acuminate to acute, acumen ca. 1 cm long, base cordate to acute, subcoriaceous, below densely pubescent when young and old, above glabrous when young and old, concolorous; midrib sunken or flat, not grooved, above glabrous when young and old, below densely pubescent when young and old; secondary veins 9 to 18 pairs, distinct, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless branches, appearing axillary, forming on a short peduncle 4–10 mm long. Flowers with 9 perianth parts in 3 whorls, 2 to 5 per inflorescence; pedicel 11–22 mm long, 3–4 mm in diameter, sparsely to densely pubescent; in fruit 1–25 mm long, 1–4 mm in diameter, sparsely to densely pubescent; bracts 2, one basal and one upper in the lower half of pedicel, basal bracts 7–9 mm long; sepals 3, valvate, free, 10–15 mm long, 9–10 mm wide, ovate, apex acute, base truncate, yellowish green to greyish green, densely pubescent outside, glabrous inside, margins flat; petals free, outer petals longer than inner to sub equal; outer petals 3, 10–22 mm long, 5–8 mm wide, elliptic to ovate, apex acute, base truncate, cream to white, margins flat, sparsely pubescent outside, glabrous inside; inner petals 3, imbricate, 15–17 mm long, 4–5 mm wide, elliptic to ovate, apex acute, base truncate, cream, margins flat, pubescent outside, glabrous inside; stamens numerous, ca. 1 mm long, broad; connective discoid, glabrous, red; staminodes absent; carpels free, 50 to 75, ovary ca. 1.5 mm long, stigma globose, glabrous. Fruit pseudosyncarpous, 40–200 mm in diameter, globose to depressed ovoid; individual carpels sessile, 60 to 70 carpels, completely fused, ca. 15 mm long, ca. 7 mm in diameter, globose to ovoid, apex domed-shaped, densely pubescent, longitudinally ribbed with 5 to 6 main ribs, pinkish red when ripe; seed 1 per monocarp, 7–15 mm long, 4–7 mm in diameter, ellipsoid; aril present, pale yellow.
Duguetia barteri A flower, side view B flower, bottom view C fruit, note fused monocarps into a syncarpous fruit, referred to as a pseudosyncarp and with dome shaped apex D longitudinal section of fruit, note completely fused monocarps. Duguetia confinis E leaves, upper side F base of leaf blade, upper side G base of leaf blade, lower side, note whitish pubescence representing minute stellate hairs completely covering whole lower side of leaf blade H flowering branch I flower, top view J detail of receptacle, all petals removed A, B Sosef 2138, Gabon C, D Couvreur 393, Ngovayang, Cameroon E–J Couvreur 527, Gabon. Photos Thomas L.P. Couvreur.
A central African species, from Cameroon to Gabon; in Cameroon known from East, South, Central, Littoral and South-West regions.
In periodically or permanently inundated forests. Altitude 350–600 m a.s.l.
ntom (dial. Bagali) (
Least Concern (LC) (
None recorded.
Duguetia barteri is distinguished by its elliptic to ovate leaves, the midrib not grooved above, inflorescences occurring on a short peduncle in leafless parts of the branches and its fruits with completely fused monocarps.
Central Region: Bank of Nyong river 40 km SE of Yaoundé, 3.65°N, 10.78°E, 09 November 1961, Breteler F.J. 2013 (BR,K,P,WAG,YA); Abimoa, 3.57°N, 11.62°E, 04 April 1962, de Bruijn J. s.n. (WAG[WAG0175175]); Bordure du Nyong près du lac de Nkolmaka (près route Mbalmayo-Akonolinga), 3.51°N, 11.82°E, 22 April 1954, Letouzey R. 304 (P,YA). Littoral Region: Right bank Ouem river near mouth in Sanaga R 6 km SW of Massok, 4.13°N, 10.47°E, 04 April 1965, Leeuwenberg A.J.M. 5377 (BR,K,MO,P,WAG,YA); Tissongo study area 3.57°N, 9.869°E, 01 June 1976, Waterman P.G. 874 (K). South Region: mountain chain Ngovoyang 2 km in forest from Bikiliki village situated between Bipindi and Lolodorf, 3.18°N, 10.52°E, 18 February 2012, Couvreur T.L.P. 393 (WAG,YA); Campo-Ma’an National Park, 2.38°N, 10.06°E, 01 July 2001, van Andel T.R. 3810 (KRIBI,WAG,YA). South-West Region: Ekundu Kundu, 5.15°N, 8.883°E, 30 April 1996, Cheek M. 8274 (K,WAG,YA).
≡ Pachypodanthium confine Engl. & Diels, Notizbl. Konigl. Bot. Gart. Berlin 3: 55, 1900.
= Pachypodanthium sargosii R.E.Fr., Ark. Bot. 3(2): 38, 1955; Pachypodanthium confine var. sargosii Le Thomas, Fl. Gabon 16: 106, 1969. Type. Republic of Congo: Kouilou, Sargos R. 29, 4 Mar 1920: lectotype, sheet here designated: P[P00364784]; isotype: P[P00364785].
Gabon. Estuaire; Libreville, Klaine T.-J. 217, 10 Oct 1895: lectotype, sheet here designated: P[P00315819]; isotypes: P[P00315821, P00315815].
Tree, 15–40 m tall, d.b.h. 40–85 cm; stilt roots or buttresses absent. Indumentum of stellate hairs; old leafless branches sparsely pubescent to glabrous, young foliate branches densely pubescent. Leaves: petiole 1–8 mm long, 2–6 mm in diameter, densely pubescent, cylindrical, blade inserted on the side of the petiole; blade 9–31 cm long, 3–8 cm wide, elliptic, apex acute, acumen 0.5–1 cm long, base rounded to acute, coriaceous, below densely pubescent with white erect stellate hairs covering the whole blade when young and old, above glabrous when young and old, discolorous, whitish below; midrib sunken or flat, above glabrous when young and old, below densely pubescent when young and old; secondary veins 14 to 25 pairs, indistinct, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless branches, appearing axillary, forming on a short peduncle 2–11 mm long. Flowers with 9 perianth parts in 3 whorls, 2 to 5 per inflorescence; pedicel 3–20 mm long, 2–5 mm in diameter, densely pubescent; in fruit 3–20 mm long, 2–5 mm in diameter, glabrous to densely pubescent; bracts 2, one basal and one towards the upper half of pedicel, basal bracts 5–9 mm long, 5–10 mm wide; sepals 3, valvate, free, 17–22 mm long, 13–15 mm wide, ovate, apex acute, base truncate, green, densely pubescent outside, glabrous inside, margins flat; petals free, outer petals longer than inner to sub equal; outer petals 3, 11–18 mm long, 5–9 mm wide, elliptic, apex acuminate, base truncate, cream to white, margins wavy, densely pubescent outside, glabrous inside; inner petals 3, imbricate, 11–18 mm long, 5–9 mm wide, elliptic, apex acuminate, base truncate, cream, margins flat, pubescent outside, glabrous inside; stamens 290 to 310, in 7 to 8 rows, 1–2 mm long, broad; connective discoid, glabrous, red; staminodes absent; carpels free, 100 to 125, ovary 1.4–2 mm long, stigma globose, glabrous. Fruit pseudosyncarpous, 25–50 mm in diameter, ellipsoid; individual carpels sessile, 100 to 125 carpels, completely fused, 15–12 mm long, 2–5 mm in diameter, obovoid, apex apiculate to acute, densely pubescent, longitudinally ribbed with 5 to 6 main ribs, greyish brown with purplish red or pale brown pulp when ripe; seed 1 per monocarp, 10–14 mm long, 4–8 mm in diameter, ellipsoid; aril present, red.
From Cameroon to Gabon and Republic of Congo; in Cameroon known from East, South and Littoral regions.
In lowland periodically inundated or non-inundated rain forests. Altitude 0–50 m a.s.l.
None recorded.
Least Concern (LC) (
None recorded.
Duguetia confinis is distinguished by its lower leaf surface, which is densely pubescent with erect stellate hairs completely covering the blade, and its carpels completely fused in fruit.
Duguetia staudtii A flowering branch B detail of stellate pubescence on lower side of leaf blade C flower bud, side view D carpel, side view E syncarpous fruit, referred to as a pseudosyncarpous fruit F longitudinal section of fruit. Duguetia confinis G flowering branch H detail of pubescent on lower side of blade, note that it is completely covering the lower surface I flower bud, side view J detail of flower, sepals removed K outer petal, inner view L inner petals, inner view M stamen, side and front views N carpel, side and front views O longitudinal section of fruit, note completely fused monocarps P seed A–D from Letouzey 4438 E, F from Chevalier 16224 G, H from Le Testu 1774 I–N, P from Klaine 217 O from Lecompte s.n. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
Central Region: Est N of Lom near Sanaga river (Yaundé) 250 km from Deng Deng, 4.84°N, 13.19°E, 01 March 1914, Mildbraed G.W.J. 8558 (K). Littoral Region: Au sud de Ngola (8 km E de l’embouchure de la Sanaga), 3.55°N, 9.698°E, 05 January 1974, Letouzey R. 12585 (P); Douala-Edea Reserve 10B, 5.01°N, 13.33°E, 01 April 1978, Thomas D.W. 1237 (K). South Region: ca 16 km from Kribi Ebolowa road Bidou plantation Kienke forReserve, 2.85°N, 10.01°E, 03 February 1969, Bos J.J. 3844 (WAG,YA); 20 km From Kribi N of Lolodorf road (SFIA logging road), 3.01°N, 10.05°E, 15 July 1969, Bos J.J. 5048 (BR,BR,K,WAG,YA); Mt Elephant ca 18 km SE of Kribi, 2.78°N, 10.5°E, 14 January 1970, Bos J.J. 6128 (BR,P,WAG); Near mouth of, 3.17°N, 9.961°E, 28 March 1928, Hédin L. 1690 (P,WAG); Campo-Ma’an area 2.71°N, 9.866°E, 26 October 2001, van Andel T.R. 4205 (KRIBI,WAG); Bipindi, 3.08°N, 10.42°E, 01 January 1904, Zenker G.A. 3195 (B,BR,E,G,L,M,MO,P,S).
Gabon. Ngounié; new road from Mouila to Yeno, 5 km on either side of Kembele village, Thomas D.W. & Wilks C.M. 6510, 20 Jul 1986: lectotype, sheet here designated: WAG[WAG0143388]; isotypes: MO[MO-357359]; P[P00389133]; WAG[WAG0027128].
Tree, up to 30 m tall, d.b.h. unknown; stilt roots or buttresses absent. Indumentum of stellate or fasciculate hairs; old leafless branches sparsely pubescent to glabrous, young foliate branches sparsely pubescent. Leaves: petiole 4–5 mm long, 2–3 mm in diameter, sparsely pubescent, grooved, blade inserted on the side of the petiole; blade 7–16 cm long, 2.5–6 cm wide, narrowly elliptic to narrowly obovate, apex acuminate to acute, acumen ca. 1 cm long, base acute, subcoriaceous, below sparsely pubescent when young and old, above glabrous when young and old, concolorous; midrib sunken or flat, above glabrous when young and old, below sparsely pubescent when young and old; secondary veins 8 to 15 pairs, distinct, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on young foliate branches, leaf opposed, not forming a peduncle. Flowers with 9 perianth parts in 3 whorls, 2 to 4 per inflorescence; pedicel 10–12 mm long, 1–2 mm in diameter, sparsely pubescent to densely pubescent; in fruit 10–50 mm long, 1–15 mm in diameter, sparsely pubescent to densely pubescent; bracts 2, one basal and one towards the lower half of pedicel, basal bracts 5–9 mm long, 7–9 mm wide; sepals 3, valvate, free, 12–15 mm long, 6–9 mm wide, elliptic to ovate, apex acute, base truncate, greyish green, pubescent outside, pubescent inside, margins flat; petals free, outer petals longer than inner to sub equal; outer petals 3, 12–15 mm long, 4–6 mm wide, ovate, apex acute, base truncate, white, margins flat, sparsely pubescent outside, glabrous inside; inner petals 3, imbricate, 13–15 mm long, 4–5 mm wide, elliptic, apex acute, base truncate, margins flat, pubescent outside, glabrous inside; stamens numerous, 1 mm long, broad; connective discoid, glabrous, red; staminodes absent; carpels free, ca. 75, 2–3.5 mm long, stigma globose, glabrous. Fruit pseudosyncarpous, size and shape unknown; carpels sessile, free to basally fused, unknown number of carpels, 20–35 mm long, ovary 10–30 mm in diameter, obovoid to deltoid, apex apiculate, pubescent, densely pubescent, longitudinally ribbed with 6 to 7 main ribs, color unknown; seed 1 per monocarp, 12–20 mm long, 10–13 mm in diameter, ellipsoid; aril present, color unknown.
A Duguetia dilabens B Duguetia staudtii C Greenwayodendron glabrum D Greenwayodendron suaveolens E Hexalobus bussei F Hexalobus crispiflorus G Hexalobus monopetalus H Hexalobus salicifolius I Isolona campanulata. White borders represent region limits in Cameroon; green patches represent protected areas (see methods and Suppl. material
Known from Gabon and Cameroon; in Cameroon known from South and Littoral regions.
A rare species known from four specimens; in lowland periodically inundated or non-inundated rain forests. Altitude 0–500 m a.s.l.
None recorded.
Endangered (EN) (
None recorded.
Duguetia dilabens is distinguished by its leaves that are narrowly elliptic to narrowly obovate, the midrib not grooved above and the carpels only basally fused in fruit. It was recently collected in Campo Ma’an National Park (Couvreur 692), but the sample is sterile and the identification remains doubtful, although the leaves do match the type specimen. Couvreur 692 also notes that the trunk had a bark peeling in smallish flakes.
South Region: Reserve forestière de la Kienké (Kribi-Ebolowa km 16), 3.1°N, 10.25°E, 05 January 1968, Bamps P.R.J. 1679 (BR, YA); Campo Ma’an National Park 11 km on trail from Ebinanemeyong village on road 7 km from Nyabessan to Campo town, 2.49°N, 10.34°E, 12 February 2015, Couvreur T.L.P. 692 (WAG,YA).
≡ Uvaria staudtii Engl. & Diels, Notizbl. Königl. Bot. Gart. Berlin 2: 292, 1899; Pachypodanthium staudtii Engl. & Diels, Notizbl. Königl. Bot. Gart. Berlin 3: 55, 1900.
Cameroon. South Region; near Lolodorf, Staudt A. 133, 1896: holotype: B[B 10 0154084]; isotypes: BM[BM000843984]; K[K000198873, K000198874]; P[P00315814, P00315816]; S[S02-94].
Tree, 15–50 m tall, d.b.h. 20–70 cm; stilt roots or buttresses absent. Indumentum of stellate or fasciculate hairs; old leafless branches sparsely pubescent to glabrous, young foliate branches sparsely pubescent to densely pubescent. Leaves: petiole 2–10 mm long, 2–4 mm in diameter, densely pubescent to sparsely pubescent, grooved, blade inserted on the side of the petiole; blade 13–34 cm long, 3–8 cm wide, narrowly obovate, narrowly oblong to narrowly elliptic, apex acuminate to acute, acumen 0.5–1 cm long, base cordate (more rarely acute), coriaceous, below sparsely pubescent when young, pubescent when old, above glabrous when young and old, concolorous; midrib sunken or flat, above glabrous when young and old, below sparsely pubescent when young and old; secondary veins 10 to 22 pairs, weakly distinct, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless or young foliate branches, leaf opposed or extra axillary, not forming a peduncle. Flowers with 9 perianth parts in 3 whorls, 2 to 4 per inflorescence; pedicel 6–12 mm long, 2–3 mm in diameter, sparsely pubescent to densely pubescent; in fruit 2–30 mm long, 2–6 mm in diameter, sparsely pubescent to densely pubescent; bracts 2, one basal and one towards the upper half of pedicel, basal bracts 9–12 mm long, 5–7 mm wide; sepals 3, valvate, free, 9–15 mm long, 7–10 mm wide, ovate, apex acute, base truncate, green, densely pubescent outside, glabrescent inside, margins flat; petals free, outer petals longer than inner to sub equal; outer petals 3, 7–30 mm long, 4–10 mm wide, oblong-elliptic to oblong-obovate, apex acute, base truncate, cream to white, margins flat, sparsely pubescent outside, glabrous inside; inner petals 3, imbricate, 4–20 mm long, 4–6 mm wide, elliptic to obovate, apex acute to obtuse, base truncate, cream, margins flat, pubescent outside, glabrous inside; stamens 120 to 150, in 5 to 6 rows, 1–2 mm long, broad; connective discoid, glabrous, red; staminodes absent; carpels free, 50 to 100, ovary 1–1.5 mm long, stigma globose, glabrous. Fruit pseudosyncarpous, 20–55 mm in diameter, globose to depressed ovoid; carpels sessile, basally fused, 50 to 100 monocarps, 20–55 mm long, 2–10 mm in diameter, globose to ovoid, apex acute, densely pubescent, longitudinally ribbed with 5 to 6 main ribs, red when ripe; seed 1 per monocarp, 7–13 mm long, 6–8 mm in diameter, ellipsoid; aril present, pale yellow.
A widespread species with a disjunct distribution in West (Sierra Leone, Liberia, Ivory Coast and Nigeria) and in Central Africa from Cameroon to Democratic Republic of the Congo; in Cameroon known from East, South, Central, Littoral and South-West regions.
A common species; in lowland or premontane primary and secondary non-inundated rain forests. Altitude 100–900 m a.s.l.
ntom (dial. Bagali); nto ntomba (dial. Bagielli) (
Least Concern (LC) (
medicine : bark used for pain-killers, pulmonary troubles, vermifuges, dropsy, swellings, oede gout, tumours, cancers; constructions: building materials; dyes and tannins: tannins, astringents, insecticides, arachnicides, arrow-poisons, aromatic substances, alkaloids.
Duguetia staudtii is distinguished by its narrowly obovate, narrowly oblong to narrowly elliptic leaves, the midrib that is grooved above and monocarps that are only basally fused.
Central Region: Yaoundé, 3.86°N, 11.51°E, 01 January 1935, Foury P. 69 (P,WAG). East Region: 17 km along road to Deng Deng, 4.58°N, 13.68°E, 01 September 1961, Breteler F.J. 1841 (WAG); Près Kinsassa 65 km au NNE de Moloundou sur route Yokadouma 2.63°N, 15.37°E, 04 March 1971, Letouzey R. 10509 (P,YA); Colline à l’ENE de Mbalam (140 km ESE de Djoum près Souanké-Congo), 2.22°N, 13.82°E, 20 January 1973, Letouzey R. 11867 (P,YA). Littoral Region: Douala-Edéa Reserve Tissongo study area Transect B, 3.57°N, 9.869°E, 01 June 1976, Waterman P.G. 879 (U). South Region: Bitye, 3.87°N, 11.52°E, 01 January 1919, Bates G.L. 1199 (BM,MO); 17 km east from Lélé village, 2.28°N, 13.32°E, 07 September 2013, Couvreur T.L.P. 460 (WAG,YA); 25 km east from Lélé village at end of path on Ivindo river, 2.25°N, 13.28°E, 09 September 2013, Couvreur T.L.P. 489 (WAG,YA); Sud TDC, 2.65°N, 9.9°E, 06 November 1991, Hallé F. 4220 (WAG); Ncolbew 3.28°N, 11.2°E, 26 April 1928, Hédin L. 1646 (P); Colline Ebon près Nkobiyo 25 km ENE d’Ambam, 2.45°N, 11.5°E, 21 March 1970, Letouzey R. 10181 (P,YA); Mvini 35 km east of Campo, 2.37°N, 10.09°E, 20 December 1983, Mikio K. 5 (P,YA); ca 7 km NE of Ebom, 3.11°N, 10.75°E, 01 August 1996, Parren M.P.E. 157 (KRIBI,WAG); ca 7 km NE of Ebom, 3.11°N, 10.75°E, 01 August 1996, Parren M.P.E. 212 (KRIBI,WAG); Lolodorf, 3.23°N, 10.73°E, 1896, Staudt A. 133 (P); Lolodorf, 3.23°N, 10.73°E, March 1895, Staudt A. 138 (B); Campo-Ma’an area 2.4°N, 10.1°E, 02 April 2001, van Andel T.R. 3290 (KRIBI,U,WAG,YA). South-West Region: Bayang Mbo Wildlife Sanctuary after Mbu river, 5.35°N, 9.501°E, 26 March 2016, Couvreur T.L.P. 1014 (WAG,YA); Near Mamfe, 5.75°N, 9.31°E, 19 April 1978, Thomas D.W. 384 (K).
≡ Polyalthia sect. Afropolyalthia Engler & Prantl., Leipzig, W. Engelmann.160, 1897.
Greenwayodendron suaveolens (Engl. & Diels) Verdc.
Trees, 7–45 m tall, d.b.h. 3–125 cm; stilt roots or buttresses absent. Indumentum of simple hairs. Leaves: petiole 2–8 mm long, 1–3 mm in diameter, blade 6.5–16.2 cm long, 2–6.7 cm wide, elliptic to oblong, apex acuminate to caudate, base cuneate to rounded, concolorous; midrib sunken or flat; secondary veins 5 to 18 pairs; tertiary venation reticulate. Individuals androdioecious; male and bisexual inflorescences similar in appearance, ramiflorous on young foliate branches, leaf opposed or extra axillary. Flowers with 9 perianth parts in 3 whorls, 1 to 4 per inflorescence; pedicel 4–6 mm long; in fruit 6–13 mm long; bracts 2, one basal and one upper, 1–2 mm long; sepals 3, valvate, free, 2–4 mm long, ovate, apex acuminate, base truncate; petals free; outer petals longer than inner; outer petals 3, valvate, 8–18 mm long, 2.3–2.6 mm wide, oblong to elliptic, apex acuminate, base rounded; inner petals 3, valvate, 8–18 mm long, 1.3–2.6 mm wide, ovate to elliptic, apex acuminate, base rounded; stamens 15 to 25, in 4 to 5 rows, ovary 1–2 mm long, elongated; connective tongue-shaped, glabrous; staminodes absent; carpels free, 10 to 20, 1–2 mm long, stigma ovoid, pubescent. Fruits apocarpous, monocarps stipitate, stipes 5–10 mm long, monocarps 2 to 8, 8–21 mm long, 7–21 mm in diameter, ellipsoid to globose, apex rounded, smooth, green turning wine red when ripe; seed 1 to 4, 3–13 mm long, 3–13 mm in diameter, ellipsoid to flattened ellipsoid; aril absent.
A genus of six currently described species distributed across Africa. Two species are known from Cameroon, none endemic.
1 | Petiole and midrib glabrous above | G. glabrum |
– | Petiole and midrib pubescent or sparsely pubescent above | G. suaveolens |
Cameroon. South Region; 40 km from Kribi, 5 km. E. of Edea road, tract of Fifinda-Bella road (SFIA), Bos J.J. 6267, 6 Feb 1970: holotype WAG[WAG.1433854]; isotypes BR[BR0000014826399]; YA n.v.; WAG[WAG1433855].
Tree, 7–30 m tall, d.b.h. 3–20 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches pubescent. Leaves: petiole 3–6 mm long, 1–2 mm in diameter, glabrous, grooved, blade inserted on the side of the petiole; blade 6.5–16.2 cm long, 2.1–5.8 cm wide, elliptic to oblong, apex acuminate to caudate, acumen 0.4–2 cm long, base cuneate to rounded, papyraceous, below sparsely pubescent to glabrous when young, glabrous when old, above glabrous when young and old, concolorous; midrib impressed, above completely glabrous when young and old, below glabrous when young and old; secondary veins 5 to 7 pairs, glabrous below; tertiary venation indistinct. Individuals androdioecious; male and bisexual inflorescences similar, ramiflorous on old leafless and young foliate branches, leaf opposed or extra axillary. Flowers with 9 perianth parts in 3 whorls, 1 to 4 per inflorescence; pedicel ca. 4 mm long, ca. 1 mm in diameter, pubescent to glabrous; in fruit 6–13 mm long, 1–2 mm in diameter, pubescent to glabrous; bracts 2, one basal and one upper towards the upper half of pedicel, basal bracts 1–2 mm long, 2 mm wide; upper bracts 1–3 mm long, 1–3 mm wide; sepals 3, valvate, basally fused to free, 3 mm long, 3–4 mm wide, ovate, apex acuminate, base truncate, green, pubescent outside, glabrous inside, margins flat; petals free, sub equal; outer petals 3, 12–13 mm long, 2–2.5 mm wide, elliptic to ovate, apex acuminate, base rounded, green to light yellow, margins flat, pubescent outside, glabrous inside; inner petals 3, valvate, 12–13 mm long, 2–2.5 mm wide, ovate, apex acuminate, base rounded, green to light yellow, margins flat, pubescent outside, glabrous inside; stamens 10 to 15, in 4 to 5 rows, 1–2 mm long, elongated; connective tongue shaped, glabrous, green; staminodes absent; carpels free, 10 to 15, ovary ca. 1 mm long, stigma ovoid, pubescent. Monocarps stipitate, stipes 5–10 mm long, 1–3 mm in diameter; monocarps 2 to 8, 11–21 mm long, 11–21 mm in diameter, ellipsoid to globose, apex rounded, glabrous, smooth, smooth; seeds 1 to 4 per monocarp, 7–13 mm long, 7–13 mm in diameter, ellipsoid to flattened ellipsoid; aril absent.
Known from Cameroon and Gabon; in Cameroon known from the Littoral and South regions.
A common species when present and growing in sympatry with G. suaveolens in southern Cameroon; in lowland non-inundated primary or secondary forests. Altitude 20–750 m a.s.l.
Greenwayodendron glabrum A flowering branch B detail of lower leaf surface C detail of upper leaf surface D–G different types leaf apex H flower bud I infructescence J longitudinal section of fruit revealing seed K seed, latitudinal view L longitudinal section of seed showing ruminations A–C, G, H Letouzey 12869 D–F, I–L Bos 6267. Drawings by Hans de Vries (
None recorded, but possibly same as G. suaveolens (see below).
Least Concern (LC) (
None recorded.
This species is very close morphologically to G. suaveolens. Both species grow in sympatry in southern Cameroon. However, G. glabrum is distinguished by its glabrous petiole and upper midrib and leaf blades (versus pubescent in G. suaveolens). Studies have shown that these two species are genetically distinct at both the phylogenetic (
Central Region: Left bank Nyong R 30 km S of Edéa near bridge in road to Kribi, 3.8°N, 10.13°E, 26 April 1965, Leeuwenberg A.J.M. 5582 (B,BR,C,GC,K,LUAI,MO,P,UC,WAG,YA). Littoral Region: Ndogtima Nyong (Edéa), 3.8°N, 10.13°E, 03 February 1974, Letouzey R. 12869 (BR,P,WAG,YA). South Region: 43 kmN of Kribi 5 km E of Edea road forest track Fifinda-Bella old secondary forest, 3.21°N, 10.06°E, 06 February 1970, Bos J.J. 6267 (BR,P,WAG,YA); ca 16 km On the road from Ebolowa to Minkok, 2.98°N, 11.17°E, 12 September 1975, de Wilde J.J.F.E 8465 (B,BR,K,MO,P,WAG,YA); Mvini 35 km East of Campo, 2.39°N, 10.04°E, 19 December 1983, Kaji M. 4 (YA); Campo-Ma’an region, 2.28°N, 9.950°E, 17 January 2016, Lissambou B.J. 1745 (BRLU); Campo-Ma’an region, 2.28°N, 9.949°E, 17 January 2016, Lissambou B.J. 1748 (BRLU); Campo-Ma’an region, 2.28°N, 9.948°E, 17 January 2016, Lissambou B.J. 1775 (BRLU); Campo-Ma’an region, 2.28°N, 9.949°E, 17 January 2016, Lissambou B.J. 1788 (BRLU); Campo-Ma’an region, 2.29°N, 9.945°E, 18 January 2016, Lissambou B.J. 1807 (BRLU); Campo-Ma’an region, 2.40°N, 9.895°E, 18 January 2016, Lissambou B.J. 1828 (BRLU); Campo-Ma’an region, 2.40°N, 9.894°E, 18 January 2016, Lissambou B.J. 1830 (BRLU); Campo-Ma’an region, 3.19°N, 10.10°E, 19 January 2016, Lissambou B.J. 1855 (BRLU); Campo-Ma’an region, 3.19°N, 10.10°E, 19 January 2016, Lissambou B.J. 1856 (BRLU); Cagnon du Ntem 16 km SW de Nyabessan, 2.32°N, 10.28°E, 30 November 1982, Nkongmeneck B.A. 400 (YA).
≡ Polyalthia suaveolens Engl. & Diels, Monogr. Afr. Pfl. 6: 42., 1901.
= Polyalthia mortehanii
De Wild., Bull. Jard. Bot. État Bruxelles, 4: 384., 1914. Type. Democratic Republic of the Congo. Kasaï-Oriental: Lekimi, De Giorgi S. 1576, Dec 1913: lectotype designated by
= Polyalthia aubrevillei
Ghesquière ex Aubréville, Fl. For. Côte d’Ivoire, i. 114, 1936. Type. Cameroon. South Region: Bipindé, Urwaldgebiet, Zenker G. 1306, 1913: lectotype designated by
= Maba gossweileri Greves., J. Bot. 67 (Suppl. 2): 76., 1929. Type. Angola. Cabinda: Buco Zau - Maiombe, Gossweiler J. 6923, 8 Jan 1917: holotype BM[BM000547162]; COI[COI00004858].
= Xylopia otunga Exell., J. Bot. 69: 99, 1931. Type. Cameroon. Central: Bitye Yaoundé, Bates G.L. 1226, 1919: holotype: BM[BM000513697]; isotype LISC[LISC000385].
Gabon. Estuaire; Munda, Sibange Farm, Soyaux H. 218, 20 Feb 1881: holotype material presumably destroyed at B; lectotype, designated by
Greenwayodendron suaveolens A flowering branch B flower bud C flower at anthesis D detail of male receptacle, petals removed E detail of hermaphrodite receptacle, petals removed F inside view of outer petal G stamen H stamen I carpel J longitudinal section of carpel K fruiting branch L lateral view of seed M seed N longitudinal section of a single monocarp showing two seeds and their ruminations A–D, F–H from Le Testu 9408; E, I, J. Gilbert 936 K–N Letouzey 5322. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
Tree, 8–45 m tall, d.b.h. 10–125 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches pubescent to sparsely pubescent. Leaves: petiole 2–8 mm long, 1–3 mm in diameter, pubescent to sparsely pubescent, grooved, blade inserted on the side of the petiole; blade 5.1–15.6 cm long, 2–6.7 cm wide, elliptic to oblong, apex acuminate to caudate, acumen 0.6–1.4 cm long, base cuneate to rounded, papyraceous, below pubescent when young, glabrous to pubescent when old, above densely to sparsely pubescent when young, sparsely pubescent when old, concolorous; midrib impressed, above pubescent at least basely when young and old, below densely pubescent when young, sparsely pubescent to densely pubescent when old; secondary veins 5 to 18 pairs, glabrous below; tertiary venation indistinct. Individuals androdioecious; male and bisexual inflorescences similar, ramiflorous on old leafless and young foliate branches, leaf opposed or extra axillary. Flowers with 9 perianth parts in 3 whorls, 1 to 4 per inflorescence, pedicel 3–6 mm long, 1–2 mm in diameter, pubescent; in fruit 6–12 mm long, 2–3 mm in diameter, glabrous; bracts 2, one basal and one upper towards the upper half of pedicel, basal bracts 1–2 mm long, 2 mm wide; upper bracts 1–3 mm long, 1–3 mm wide; sepals 3, valvate, basally fused to free, 2–4 mm long, 2–4 mm wide, ovate, apex acuminate, base truncate, green, pubescent outside, glabrous inside, margins flat; petals free, sub equal; outer petals 3, 8–18 mm long, 1.3–2.6 mm wide, oblong-elliptic to ovate, apex acuminate, base rounded, green to light yellow, margins flat, pubescent outside, glabrous inside; inner petals 3, valvate, 8–18 mm long, 1.3–2.6 mm wide, elliptic to ovate, apex acuminate, base rounded, green to light yellow, margins flat, pubescent outside, glabrous inside; stamens 16 to 25, in 4 to 5 rows, 1–2 mm long, elongated; connective tongue-shaped, glabrous, green; staminodes absent; carpels free, 12 to 20, ovary 1–2 mm long, stigma ovoid, pubescent. Monocarps stipitate, stipes 5–10 mm long, 1–3 mm in diameter; monocarps 2 to 8, 8–18 mm long, 7–16 mm in diameter, ellipsoid to globose, apex rounded, glabrous, smooth, green turning wine red when ripe; seeds 1 to 4 per monocarp, 3–11 mm long, 3–11 mm in diameter, ellipsoid to flattened ellipsoid; aril absent.
Known from Nigeria to the Republic of Congo and the Democratic Republic of Congo; in Cameroon known from the East, South, Central, Littoral and South-West regions.
A very common species across the forest zone of Cameroon (growing in sympatry with G. glabrum in the south) with a wide ecologically amplitude; in lowland premontane, and sometimes in montane non-inundated primary or secondary forests. Altitude 20–1600 m a.s.l.
Moabé noir (dial. Nzime), Otunga (dial. Fang), Otungui (dial. Ewondo), Ntoulen (dial. Bassa), Botounga, Botunga (dial. Baka).
Least Concern (LC) (
Greenwayodendron suaveolens A trunk, note light grey color B leaf, upper view C flower D flower, side view E detail of fruit, with longitudinal section of one monocarp showing 2 seeds. Greenwayodendron glabrum F branch with fruit G fruit with longitudinal section of monocarp showing seed A, E Couvreur 476, Lélé, Cameroon B Couvreur 1196, Maséa, Cameroon C, D Couvreur 560, Gabon F, G Bidault 847, Gabon. Photos A–E Thomas L.P. Couvreur F, G Ehoarn Bidault, Tropicos.org, Missouri Botanical Garden.
medicine : leaves as pain-killers, against arthritis, rheumatism, fabrifuges, for menstrual cycle; bark for pregnancy, antiaborifacients, root for vermifuges, as genital stimulants/depressants, dropsy, swellings, oede gout; constructions: building materials; dyes and tannins: glycosides, saponims, steroids; products: fibre wood, farming, forestry, hunting and fishing apparatus.
See under G. glabrum. The species is here treated in the narrow sense; the former varieties gabonicum Le Thomas and usambaricum Verdc. (not recorded from Cameroon) are now raised to specific rank, as G. gabonicum (Le Thomas) Lissambou & Couvreur and G. usambaricum (Verdc.) Lissambou, Hardy & Couvreur (
Central Region: Ndanan 1, 3.62°N, 11.58°E, 21 October 2002, Cheek M. 11224 (K,YA); Mefou National Park, 3.61°N, 11.58°E, 13 March 2004, Cheek M. 27 (YA); Mefou National Park, 3.61°N, 11.58°E, 13 March 2004, Cheek M. 66 (YA); Ca 50 km S of Badjob ca 60 km SW Of Eséka Along the Njong-River, 3.68°N, 10.68°E, 19 March 1964, de Wilde W.J.J.O 2133 (B,BR,K,MO,P,WAG,YA); Yaoundé, 3.87°N, 11.52°E, 01 January 1935, Foury P. 129 (P); Ngoro, 5.06°N, 11.19°E, 29 April 2017, Kamdem N. 510 (YA); AYOS, 3.98°N, 12.36°E, 17 June 2017, Kamdem N. 560 (YA); Ossoéssam (Mbalmayo), 3.52°N, 11.5°E, 01 June 1965, Leeuwenberg A.J.M. 5755 (BR,K,P,WAG). East Region: Palisco forest consession 15 km along main road into consession, 3.48°N, 13.59°E, 27 March 2015, Couvreur T.L.P. 756 (WAG,YA); Deng Deng, 5.20°N, 13.13°E, 27 July 2014, Kamdem N. 166 (YA); Mindourou Alpicam, 4.12°N, 14.54°E, 11 December 2016, Kamdem N. 459 (YA); Colline à l’ENE de Mbalam (140 km ESE de Djoum près de Souanke-Congo, 2.22°N, 13.82°E, 20 January 1973, Letouzey R. 11866 (YA); A 6 km au Nord de Mwapak (km 43 piste Yokadouma-Lomié, 3.54°N, 14.71°E, 22 June 1963, Letouzey R. 5322 (YA). Littoral Region: Mapubi 30 km before Edea on Yaoundé-Edea road On forestry road 5 km direction to Sanaga river, 3.84°N, 10.38°E, 28 February 2018, Couvreur T.L.P. 1180 (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. 658 (WAG,YA); Olombé, 3.60°N, 9.959°E, 05 November 2014, Kamdem N. 175 (YA); Chantier Bakaka km 4 Eboné-EkoMtolo road (Eboné situated on km 11 of Nkongsamba-Loum road), 4.83°N, 9.9°E, 20 August 1971, Leeuwenberg A.J.M. 8164 (BR,K,L,MO,P,U,WAG,YA). South Region: 20 km from Kribi Lolodorf road, 3.03°N, 10.05°E, 09 June 1969, Bos J.J. 4769 (B,BR,K,LD,LM,MO,P,POZG,WAG,YA); Mt Elephant ca 18 km SE of Kribi, 2.81°N, 10.01°E, 08 January 1970, Bos J.J. 6100 (BR,C,K,LD,P,WAG,YA); hill above Nlonacko near village Ebianemeyong, 2.43°N, 10.35°E, 12 December 1998, de Wilde J.J.F.E 12163 (BR,KRIBI,MO,S,WAG); 16 km on the recently reconstructed road from Ebolowa to Minkok, 2.75°N, 11.25°E, 30 January 1975, de Wilde J.J.F.E 7940 (BR,K,MO,P,U,WAG,YA); Massif de Ngovayang village de Atog Boga, 3.25°N, 10.49°E, 30 August 2015, Droissart V. 2050 (BRLU); Massif de Ngovayang village de Atog Boga, 3.25°N, 10.49°E, 05 September 2015, Droissart V. 2159 (BRLU); A 13 km au N-NW de Djoum (UFA 09-007) vers “la tache verte” Forêt dense inondée et forêt secondaire, 2.77°N, 12.74°E, 25 April 2011, Droissart V. 834 (BRLU); Ebom, 3.1°N, 10.73°E, 20 February 1996, Elad M. 443 (KRIBI,WAG); Campo, 2.28°N, 9.950°E, 05 July 2015, Kamdem N. 329 (YA); Campo, 2.39°N, 10.02°E, 07 July 2015, Kamdem N. 349 (YA); Ma’an, 2.50°N, 10.76°E, 11 July 2015, Kamdem N. 397 (YA); Essam (Nanga Eboko), 4.68°N, 12.37°E, 13 February 1959, Letouzey R. 1106 (P); Essam (Nanga Eboko), 4.68°N, 12.37°E, 13 February 1959, Letouzey R. 1313 (P); 10 km environ à l’ESE de Campo à Kribi, 2.37°N, 9.82°E, 26 March 1968, Letouzey R. 9198 (YA); Campo-Ma’an National Park, 2.38°N, 10.06°E, 01 July 2001, van Andel T.R. 3794 (KRIBI,WAG,YA); Bipindi, 3.08°N, 10.41°E, 1897, Zenker G.A. 1278 (L,P,WAG); Bipindi, 3.08°N, 10.41°E, 1899, Zenker G.A. 2062 (L,P,WAG); Bipindi, 3.08°N, 10.42°E, 01 January 1900, Zenker G.A. 2166 (L,P,WAG). South-West Region: Bayang Mbo Wildlife Sanctuary after Mbu river, 5.35°N, 9.501°E, 25 March 2016, Couvreur T.L.P. 1002 (WAG,YA); Mokoko Forest Reserve Boa/Likinge (Bousa forest), 4.42°N, 8.972°E, 05 June 1994, Ekema S.N. 1208 (K,YA); Nguti, 5.34°N, 9.496°E, 03 June 2017, Kamdem N. 537 (YA).
Hexalobus monopetalus (A. Rich.) Engl. & Diels.
Trees, 10–40 m tall, d.b.h. 35–100 cm; stilt roots or buttresses absent, but trunk strongly fluted. Indumentum of simple hairs. Leaves: petiole 1–8 mm long, 1–4 mm in diameter; blade 3.6–36 cm long, 1.2–10 cm wide, elliptic or obovate or ovate, apex acuminate or rounded to obtuse, base cuneate or cordate, concolorous; midrib sunken or flat; secondary veins 5 to 17 pairs; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless or young foliate branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 3 per inflorescence; pedicel (0)1–15 mm long, 1–5 mm in diameter; in fruit 2–30 mm long, 1–5 mm in diameter; bracts 5 to 6, several basal and two (sometimes fused) on upper half of pedicel; sepals 3, valvate, free, 4–21 mm long, 3–14 mm wide, ovate, apex acute, base truncate; petals 6, in a single whorl and basally fused, tube 2–10 mm long, inner and outer whorl not differentiated, equal or subequal; lobes 9–80 mm long, 3–21 mm wide, margins plicate (folded in bud) or wavy; stamens numerous, in 10 to 13 rows, 1–8 mm long, elongated; connective discoid or elongated; staminodes absent; carpels free, 2 to 16, ovary 2–5 mm long, stigma bilobed or divided into two lobes with margins coiled inwards. Fruit apocarpous, monocarps stipitate or sessile, stipes 0–3 mm long; monocarps 1 to 8, 22–95 mm long, 13–65 mm in diameter, ellipsoid to cylindrical, apex rounded, smooth or rugose or warty, pubescent, orange-brown to medium brown when ripe; seeds 2 to 36, 10–40 mm long, 7–20 mm in diameter, flattened ellipsoid; aril absent.
A genus of five species, distributed across Africa. Four species are known from Cameroon, one endemic.
This genus of trees is characterized by thin plicate (folded) petals, a unique character for Annonaceae (
1 | Leaf apex rounded to obtuse; pedicel 0–2 mm long, in drier regions of northern Cameroon | H. monopetalus |
– | Leaf apex acuminate; pedicel 8–25 mm long, in wetter regions of southern Cameroon | 2 |
2 | Petiole > 2.5 mm in diameter; stamens 6–8 mm long; monocarps irregularly ribbed, rugose | H. bussei |
– | Petiole < 2.5 mm in diameter; stamens > 5 mm long; monocarps not ribbed, smooth or verrucose | 3 |
3 | Leaf blade 5–10 cm long, 1.5–4 cm wide, base cuneate; corolla lobes < 30 mm long; stamens ca. 2 mm long; carpels 3–4; monocarps verrucose | H. salicifolius |
– | Leaf blade 7–25 cm long, 2.5–8.5 cm wide, base rounded to cordate or occasionally cuneate; corolla lobes > 35 mm long; stamens 3–5 mm long; carpels 7–16; monocarps smooth | H. crispiflorus |
= Hexalobus megalophyllus Engl. & Diels, Bot. Jahrb. Syst. 39: 479, 1907. Type. Cameroon. South Region, Bipindi, Zenker G.A. 2889, 1904: holotype: B[B100154197]; isotypes: BR[BR0000015306210]; COI[COI00033178]; F; G[G00011590]; HBG[HBG518920]; K[K000198935]; L[L-0049297]; M[M0089222]; MO[MO-2246481]; P[P00315838]; S[S12-22791]; US[00098767]; WAG[WAG0053629]; WRSL; WU[WU 0025867]; Z[Z-000000827].
Cameroon. South Region; Kribi, Busse W.C.O. 3216, 1904: holotype: B[B 10 0154198].
Tree, 20–30 m tall, d.b.h. unknown; stilt roots or buttresses absent, trunk slender, not fluted. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches pubescent. Leaves: petiole 1–7 mm long, 3–4 mm in diameter, pubescent, grooved, blade inserted on the side of the petiole; blade 15.5–36 cm long, 5.5–10 cm wide, obovate, apex acuminate, acumen 0.5–1 cm long, base cordate, coriaceous, below glabrous when young and old, above sparsely pubescent to glabrous when young, sparsely pubescent to glabrous when old, concolorous; midrib sunken or flat, above pubescent when young and old, below pubescent when young and old; secondary veins 12 to 17 pairs; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless or young foliate branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 2 per inflorescence; pedicel 10–12 mm long, 4–5 mm in diameter, densely pubescent; in fruit 10–13 mm long, 2–3 mm in diameter, glabrous; bracts 5 to 6, several basal and two (sometimes fused) towards the upper half of pedicel, basal bracts 3–9 mm long, 2–5 mm wide; upper bracts 6–13 mm long, 2–3 mm wide; sepals 3, valvate, free, 16–20 mm long, 11–14 mm wide, ovate, apex acute, base truncate, brown, densely pubescent outside, glabrous inside, margins flat; petals basally fused, tube 2–5 mm long, purple, inner and outer whorl not differentiated, equal, lobes 23–40 mm long, 8–10 mm wide, elliptic, apex rounded, pale yellow, margins wavy, densely pubescent outside, densely pubescent inside, plicate; stamens numerous, 7–8 mm long, elongated; connective elongated, glabrous; staminodes absent; carpels free, 3 to 7, ovary 4–5 mm long, stigma elongate, pubescent. Monocarps stipitate, ca. 1 mm long, ca. 3 mm in diameter; monocarps 2 to 4, 53–78 mm long, ca. 40 mm in diameter, ellipsoid, apex rounded, sparsely pubescent, rugose, irregularly ribbed in reticulate pattern, orange when ripe; seeds 17 to 19 per monocarp, 23–28 mm long, 17–19 mm in diameter, flattened ellipsoid; aril absent.
Hexalobus bussei A flowering branch B stamen, front view. Hexalobus monopetalus C flowering branch D opened flower, one sepal and one petal removed E stamen, front view F portion of pedicel with three monocarps A, B from Zenker 3550 C–E from Letouzey 7301 F from Chevalier 305. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
endemic to Cameroon; known from the South region.
A rare species, in primary lowland rain forests, sometimes riverine. Altitude 0–200 m a.s.l.
None recorded.
Endangered (EN) (
None recorded.
Hexalobus bussei is distinguished by its large leaf blades (15–36 cm long) and the irregularly ridged and strongly rugose surface of its monocarps.
South Region: S bank of Kienke river just E of Kribi, 3.1°N, 10.25°E, 12 May 1969, Bos J.J. 4495 (P,WAG); 21 km from Kribi high forest exploitation N of Lolodorf road, 3.03°N, 10.05°E, 07 August 1969, Bos J.J. 5157 (BR,P,WAG,YA); ca 16 km from Kribi Lolodorf road, 3°N, 10.01°E, 19 September 1969, Bos J.J. 5370 (BR,C,K,LD,LM,MO,P,WAG,YA); Kribi, 2.95°N, 9.916°E, 01 September 1904, Busse W.C.O. 3216 (B); 31 km ESE Kribi N of Kienke River Nyabessan, 2.9°N, 10.16°E, 19 April 1968, Letouzey R. 9387 (P,YA); Bipindi, 3.08°N, 10.42°E, 01 January 1904, Zenker G.A. 2889 (B,BR,COI,F,G,L,M,MO,P,S,WAG); Bipindi, 3.08°N, 10.42°E, 26 October 1908, Zenker G.A. 3550 (BR,COI,G,M,MO,P,S); Bipindi, 3.08°N, 10.41°E, 01 January 1908, Zenker G.A. 3592 (G,K); Bipindi, 3.08°N, 10.41°E, 01 January 1909, Zenker G.A. 3889 (BR,COI,G,K,L,M,MO,P,S); Bipindi, 3.08°N, 10.42°E, 01 January 1913, Zenker G.A. 4831 (BR,G,P,S).
= Hexalobus grandiflorus Benth., Trans. Linn. Soc. London 23(3): 468, 1862. Type. Cameroon. South-West Region, “Ambas Bay”, Mann G. 709, 1861: lectotype, here designated, sheet here designated: K[K000582047]; isolectotypes: GH n.v.; K[K000105530, K000105529]; P[P00315844, P00315845].
= Hexalobus grandiflorus var. inaequilaterifolius Engl., Monogr. Afrik. Pflanzen.-Fam. 6: 57, 1901. Type. Republic of Congo: Cuvette, “Bonga, Sanga”, Schlechter F.R.R. 12685, Aug 1899: holotype: B n.v.; isotypes: BR[BR0000006915513]; WRSL n.v.
= Hexalobus lujae De Wild., Bull. Jard. Bot. État Brux. 4: 389, 1914. Type. Democratic Republic of the Congo. Kasai-Oriental, Sankuru, Luja E.P. s.n., Jun 1910: lectotype, sheet here designated: BR[BR0000008800336]; isotypes: BR[BR0000008800008, BR0000008799906].
= Hexalobus crispiflorus subsp. strigulosus R.E.Fr., Acta Horti Berg. 10: 71, 1930. Type. Cameroon. no location, Deistel H. 99, no date: holotype: B[B 10 0184706]; isotypes: B[B 10 0184707, B 10 0184708, B 10 0184706, B 10 0184709, B 10 0184710, B 10 0184711]; GH; M[M0089315, M0089316]; P[P00486245].
= Hexalobus mbula
Exell, J. Bot. 70, suppl. Polypet.: 206, 1932. Type. Angola. Cabinda, Buco Zau, Fazenda Alsyra, Gossweiler J. 6939, 20 Jan 1917: lectotype, designated by
Guinea. Labé; Fouta D’hiallon [Djallon], Heudelot, J. 865, Apr 1838: lectotype, sheet here designated: P[P00315839]; isotypes: P[P00315842, P00486270, P00315841]; G[G00011589].
Tree, 25–40 m tall, d.b.h. up to 100 cm; stilt roots or buttresses absent, trunk deeply fluted. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches densely pubescent. Leaves: petiole 2–8 mm long, 1–3 mm in diameter, densely pubescent, grooved, blade inserted on the side of the petiole; blade 7.2–25 cm long, 2.5–8.5 cm wide, ovate to obovate, apex acuminate, acumen 0.5–1.5 cm long, base cuneate to cordate, coriaceous, below glabrous when young and old, above sparsely pubescent when young, sparsely pubescent when old, concolorous; midrib sunken or flat, above pubescent when young and old, below pubescent when young and old; secondary veins 9 to 19 pairs; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless or young foliate branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 3 per inflorescence; pedicel 12–25 mm long, 1–2 mm in diameter, glabrous; in fruit 10–30 mm long, 4–5 mm in diameter, glabrous; bracts 5 to 6, several basal and two (sometimes fused) towards the upper half of the pedicel, basal bracts 3–9 mm long, 2–5 mm wide; upper bracts 8–12 mm long, 4–9 mm wide; sepals 3, valvate, free, 12–21 mm long, 9–12 mm wide, ovate, apex acute, base truncate, brown, densely pubescent outside, densely pubescent inside, margins flat; petals basally fused, tube 4–10 mm long, purple, inner and outer whorl not differentiated, sub equal; lobes 37–80 mm long, 6–21 mm wide, elliptic, apex rounded, green to bright yellow, margins wavy, pubescent outside, pubescent with glabrous base inside, plicate; stamens 190 to 210, in 10 to 13 rows, 3–5 mm long, elongated; connective hemispheric, glabrous, cream; staminodes absent; carpels free, 7 to 16, ovary 2–5 mm long, stigma bilobed, slightly capitate, pubescent. Monocarps stipitate, ca. 2 mm long, 2–3 mm in diameter; monocarps 1 to 8, (42)50–95 mm long, 35–65 mm in diameter, ellipsoid to oblong, apex rounded, pubescent to glabrous, smooth, not ribbed, rusty-brown; seeds 12 to 36 per monocarp, 28–40 mm long, 17–20 mm in diameter, flattened ellipsoid; aril absent.
Hexalobus crispiflorus A flowering branch B flower bud C receptacle, petals removed D stamen, front view E three carpels different views, one with longitudinal section showing ovules F pedicel with one monocarp, note smooth surface G longitudinal section of monocarp showing seeds. Hexalobus salicifolius H branch I a single detached monocarp, note verrucose surface A, C–F from Le Testu 8838 B from Le Testu 693 G, H from Chevalier 7471 I from Le Testu 6387 J from Zenker 2268. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
Hexalobus crispiflorus A base of trunk, note deep furrows B base of leaf blade, upper view C base of leaf blade, lower view D flower, side view E flower, top view F flowering branch G fruit and leaves, note smooth monocarp surface A Couvreur 506, Ottotomo, Cameroon B, C Couvreur 1197, Maséa, Cameroon D–F Couvreur 666, Ottotomo, Cameroon G Couvreur 446, Ottotomo, Cameroon. Photos Thomas L.P. Couvreur.
A widespread species, known from Guinea-Bissau to the Democratic Republic of Congo; in Cameroon known from the East, South, Central, Littoral and South-West regions.
A common species across the forested region of the country; in lowland or premontane periodically or non-inundated, primary or secondary rain forests, including gallery forests in savanna. Altitude 0–1000 m a.s.l.
évota, pota (dial. Bibaya, baka), owé (dial. Ewondo, Letouzey 4433); Ow (dial. Bulu, Ndoum 129); Owoe (South Province, Mildbraed 5652); Pota (dial. Bambindjere?, Harris, Fay 516, 558, 883, 1518).
Least Concern (LC) (
dyes and tannins : lycosides, saponims, steroids.
None recorded.
Hexalobus crispiflorus is morphologically close to H. salicifolius, both being tall trees with deeply fluted trunks, similar flowers and growing in similar habitats. Hexalobus crispiflorus is however distinguished by its larger leaves (7.2–25 cm versus 5–10 cm long), more numerous carpels (7 to 16 versus 2 to 4 in H. salicifolius) and smooth monocarps (versus verrucose in H. salicifolius).
Central Region: Ndanan 2 to Mefou river, 3.62°N, 11.56°E, 13 October 2002, Cheek M. 11064 (K,YA); Ottotomo Forest Reserve 1 km after reserve base near small loggers road, 3.66°N, 11.28°E, 25 June 2013, Couvreur T.L.P. 446 (WAG,YA); Ottotomo Forest Reserve 3 km after reserve base near small loggers road, 3.66°N, 11.28°E, 08 September 2013, Couvreur T.L.P. 506 (WAG,YA); Ottotomo Forest Reserve 45 km South of Yaoundé 5 km on path into reserve, 3.66°N, 11.28°E, 15 January 2015, Couvreur T.L.P. 666 (WAG,YA). East Region: 73 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, 3.16°N, 14.71°E, 04 March 2019, Couvreur T.L.P. 1197 (MPU,WAG,YA); Sangha R (Ndakan), 2.78°N, 16°E, 08 March 1988, Fay J.M. 8299 (F,MO,P); West side of Sangha River, 2.35°N, 16.13°E, 01 November 1988, Harris D.J. 1518 (K,MO); West side of Sangha River, 2.35°N, 16.13°E, 02 August 1988, Harris D.J. 883 (MO,P); Rives de la Sangha près Lidjombo 120 km au N de Ouesso, 2.61°N, 16°E, 08 April 1971, Letouzey R. 10614 (P,YA); A 15 km au Sud de Djouo (20 km E de Somalomo dur le Dja), 3.32°N, 12.93°E, 25 February 1962, Letouzey R. 4433 (P,YA); South Cameroon Forest Area Molundu District Bange forest between Lokomo Bumba and Bange, 2.83°N, 15.25°E, 22 February 1911, Mildbraed G.W.J. 4539 (HBG); South Cameroon Forest Area Molundu District between Yokadouma (Post Plehn) and Assobam, 3.4°N, 14.38°E, 21 April 1911, Mildbraed G.W.J. 4996 (HBG); Réserve de Biosphère du Dja vers 500 m de la station de Bouamir, 3.19°N, 12.81°E, 27 May 2001, Senterre B. 1641 (BR); Layon nord-sud à partir de Djolimpoun (entre Somalomo-Malen), 3.33°N, 12.87°E, 13 September 1993, Sonké B. 591 (YA). Littoral Region: km 11 Loum-Solé road, 4.7°N, 9.816°E, 24 May 1972, Leeuwenberg A.J.M. 9904 (K,MO,WAG,YA). South Region: Hill roughly between Nkolandom and Nkoemvone, 2.8°N, 11.16°E, 17 January 1975, de Wilde J.J.F.E 7909 (B,BR,C,K,MO,P,U,WAG,YA); Ebolowa, 2.91°N, 11.31°E, 01 June 1911, Mildbraed G.W.J. 5652 (HBG); Ebom, 3.1°N, 10.71°E, 24 September 1996, Ndoum D. 129 (KRIBI,WAG); Kribi, 2.92°N, 9.900°E, 01 October 1997, van der Burgt X.M. 232 (KRIBI,WAG); Bisyang, 2.98°N, 9.968°E, 04 June 2006, van Velzen R. 90 (BR,G,MO,WAG). South-West Region: Melon to Nyandon ca 2 km, 4.93°N, 9.533°E, 28 November 1998, Cheek M. 9716 (K,WAG,YA); Nyasoso village on max’s trail to Mt 4.82°N, 9.701°E, 05 April 2016, Couvreur T.L.P. 1062 (WAG,YA); Ambas Bay, 4.01°N, 9.2°E, 1861, Mann G. 709 (GH,K,P).
≡ Uvaria monopetala A.Rich., Guill. & Perr., Fl. Seneg. tent.: 8, 1831. Hexalobus senegalensis A.DC., Mém. Soc. Phys. Genève 5: 213, 1832, superfluous name.
= Hexalobus monopetalus var. parvifolius Baker.f.; Macleod, Chiefs and cities of Central Africa: 301, 1912. Type. Central African Republic. Environs de Kaga M’bra, Chevalier A.J.B. 6486, 30 Nov 1902: holotype: K[K000582056]; isotypes: G[G00011614]; L[L0049298]; P; WAG[WAG0162940].
= Hexalobus tomentosus
A.Chev., Expl. bot. Afr. occ. Énum. pl. 1: 10, 1920. Type. Mali. Ségou, Sansanding, Chevalier A.J.B. 2542, 29 Sep 1899: lectotype, designated by
= Hexalobus glabrescens
Hutch. & Dalziel, Fl. W. trop. Afr. 1: 52, 1927. Type. Central African Republic: Ouham, Lere to Ham, Talbot P.A. 531, 1911: lectotype, designated by
= Hexalobus monopetalus var. obovatus Brenan, Mem. New York Bot. Gard. 8, 3: 214, 1953. Type. Zambia. North-Western, E. of Matonchi Farm, Milne-Redhead E.W.B.H. 4536, 12 Feb 1938: holotype: K[K000198933]; isotypes: BM[BM000546381]; BR[BR0000008800664]; PRE[PRE0397001-0].
= Hexalobus huillensis
(Engl. & Diels) Engl. & Diels, Monogr. Afrik. Pflanzen.-Fam. 6: 56, 1901; Uvaria huillensis Engl. & Diels, Notizbl. Königl. Bot. Gart. Berlin. 2: 296, 1899. Type. Angola. Benguela, Benguella, Huilla, Antunes J.M. 266, no date: lectotype, designated by
Senegal. Tambacounda; Galam prope Bakel, Leprieur F.M.R. s.n., 1828: lectotype, designated by
Tree to shrub, 10–15 m tall, d.b.h. up to 35 cm; stilt roots or buttresses absent, often several stemmed, not fluted. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches pubescent. Leaves: petiole 1–4 mm long, 1–2 mm in diameter, densely pubescent, grooved, blade inserted on the side of the petiole; blade 3.6–17.5 cm long, 1.2–6.5 cm wide, ovate to obovate, apex rounded to obtuse, rarely acuminate, acumen 1 cm long, base cuneate to cordate, coriaceous, below pubescent when young, glabrous when old, above pubescent when young, glabrous when old, concolorous; midrib impressed, above pubescent when young, glabrous when old, below pubescent when young, glabrous when old; secondary veins 6 to 14 pairs, below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on young foliate branches, more rarely cauliflorous, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 3 per inflorescence; pedicel sessile or short up to 2 mm long, ca. 2 mm in diameter when present, densely pubescent; in fruit 2–4 mm long, 1–2 mm in diameter, sparsely pubescent; bracts 5 to 6, several basal and two (sometimes fused) towards the upper half of pedicel, basal bracts ca. 5 mm long, ca. 4 mm wide; upper bracts ca. 5 mm long, ca. 4 mm wide; sepals 3, valvate, free, 4–7 mm long, 3–6 mm wide, ovate, apex acute, base truncate, brown, densely pubescent outside, glabrous inside, margins flat; petals basally fused, tube 3–4 mm long, inner and outer whorl not differentiated, sub equal; lobes 9–27 mm long, 3–7 mm wide, elliptic, apex rounded, cream, margins wavy, pubescent outside, pubescent inside, lobes curving inwards at the base and margins reflexed forming a hollow chamber; stamens numerous, 1–2 mm long, elongated; connective discoid, glabrous, cream-yellow; staminodes absent; carpels free, 2 to 7, ovary 2–3 mm long, stigma divided into two lobes with margins coiled inwards, pubescent. Monocarps stipitate to sessile, stipes < 3 mm long, 3–4 mm in diameter; monocarps1 to 5, 22–46 mm long, 13–22 mm in diameter, ellipsoid to cylindrical, apex rounded, sparsely pubescent, warty, constricted around the seeds, orange when ripe; seeds 2 to 8 per monocarp, 10–15 mm long, 7–10 mm in diameter, flattened ellipsoid; aril absent.
A widespread species, known from Senegal to northern South Africa, with a disjunct population in southern Angola; in Cameroon known from the North and Far-North regions.
A common species in drier regions of Africa; in woodland, savannas or gallery forests, on sandy soils or in rocky places. Altitude 200–1000 m a.s.l.
Bohili (Fulfuldé) (
Least Concern (LC) (
None recorded.
Hexalobus monopetalus is distinguished from the other Cameroonian species by being a small deciduous shrub or tree (no taller than 15 m) growing in drier areas, with sessile or subsessile flowers and small smooth monocarps (up to 46 mm long versus more than 45 mm generally).
Far-North Region: Route Lara-Guidiguis (15 km ENE de Kaele), 10.1°N, 14.45°E, 29 August 1964, Letouzey R. 6540 (P,YA); Près Bourka (65 km SS0 de Mokolo), 10.3°N, 13.56°E, 13 October 1964, Letouzey R. 7301 (P,YA); 9 km SE Guili 10 km NE Bourrah, 10.6°N, 13.74°E, 27 November 1989, Villiers J.-F. 4713 (P,YA). North Region: Ecole de faune de Garoua, 9.3°N, 13.4°E, 09 August 2000, Dong E. 393 (YA); Sanguéré (10 km Garoua), 9.27°N, 13.47°E, 01 October 1949, Malzy P. 309 (P,YA); Environs village Boulko au pied Hossere Gode 15 km NW de Poli, 8.53°N, 13.13°E, 24 October 1983, Satabié B. 702 (P,YA); Collines de Tinguelin 10 km N de Garoua, 9.3°N, 13.4°E, 26 November 1984, Satabié B. 781 (P,YA); map # NC 33 VIII Garoua, 9.93°N, 13.86°E, 13 August 1983, Thomas D.W. 2432 (MO,P,WAG,YA).
Cameroon. South Region; Bipindi, Zenker G.A. 2268, 1900: holotype: B[B 10 0154199]; isotypes: BM[BM000546383]; COI[COI00033180]; G[G00011622, G00011623]; GEOT[GOET005680]; HBG[HBG518921]; K[K000582046]; L[L 0049299]; M[M0089219]; MO n.v.; P[P00363369, P00363367]; S[S12-22767]; WAG[WAG0053744]; WU[WU0025889]; Z[Z-000034502].
Tree, up to 35 m tall, d.b.h. up to 100 cm; stilt roots or buttresses absent, trunk fluted. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches densely pubescent. Leaves: petiole 1–3 mm long, 1–2 mm in diameter, sparsely pubescent, slightly grooved, blade inserted on top of the petiole; blade 5–10 cm long, 1.5–3.5 cm wide, elliptic, apex acuminate, acumen 0.5–1 cm long, base cuneate, coriaceous, above glabrous when young and old, above sparsely pubescent when young, sparsely pubescent when old, concolorous; midrib sunken or flat, below pubescent when young and old, below pubescent when young and old; secondary veins 5 to 12 pairs; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on young foliate branches, axillary. Flowers with 9 perianth parts in 3 whorls, 1 to 3 per inflorescence; pedicel 8–15 mm long, 1–2 mm in diameter, densely pubescent; in fruit 10–13 mm long, 2–3 mm in diameter, glabrous; bracts 5 to 6, several basal and two (sometimes fused) towards the upper half of pedicel, basal bracts 5 mm long, 3 mm wide; upper bracts 5 mm long, 3 mm wide; sepals 3, valvate, free, 7–11 mm long, 4–7 mm wide, ovate, apex acute, base truncate, brown, pubescent outside, densely pubescent inside, margins flat; petals basally fused, tube 2–4 mm long, purple, inner and outer whorl not differentiated, equal; lobes 17–30 mm long, 4–8 mm wide, elliptic, apex rounded, cream to yellow, margins wavy, pubescent outside, pubescent inside, plicate; stamens numerous, in 11 to 13 rows, ca. 2 mm long, elongated; connective discoid, glabrous, cream; staminodes absent; carpels free, 3 to 4, ovary 2–3 mm long, stigma divided into two lobes with margins coiled inwards, pubescent. Monocarps stipitate, 1–2 mm long, ca. 3 mm in diameter; monocarps 2 to 4, 60–93 mm long, 40–55 mm in diameter, ovoid to oblong, apex rounded, pubescent to glabrous, verrucose, rusty-brown when ripe; seeds 15 to 15 per monocarp, 22–31 mm long, 11–18 mm in diameter, flattened; aril absent.
Known from Cameroon, Gabon and the Republic of Congo (one specimen); in Cameroon known from the East, South and Central regions.
A fairly scarce species, in primary and secondary lowland and pre-montane rain forest, occasionally in semi-deciduous forests, in periodically inundated forests and on river banks. Altitude 50–900 m a.s.l.
Ooué (dial. Jaundi, Hédin 1663); Owoé (dial. Yetbou, Médou 1703).
Least Concern (LC) (
None recorded.
Hexalobus salicifolius is morphologically close to H. crispiflorus from which it is distinguished by its smaller leaves, fewer carpels (2 to 4 versus 7 to 16 in H. crispiflorus) and verrucose monocarps (versus smooth in H. crispiflorus).
South Region: South Province 15 km from Kribi, 2.96°N, 9.933°E, 28 August 1970, Bos J.J. s.n. (WAG[WAG0162941]); Campakok (Camp Akok), 2.65°N, 9.9°E, 30 October 1991, Hallé F. 4181 (WAG); Colline près Mezese à 17 km ENE de Sangméli 2.95°N, 12.14°E, 19 October 1966, Letouzey R. 8122 (BR,P,YA); Bidou, 2.85°N, 9.991°E, 28 May 1954, Médou J. SRFK 1703 (P,YA); 22 km NE of Kribi Kribi-Bipindi road Bidou I cultivated fields NE of village, 3.00°N, 10.09°E, 04 June 2006, van Velzen R. 98 (WAG); 26 km N of Kribi SE of the toll post for Kribi-Edéa road forest at Bebamboue II, 3.05°N, 9.987°E, 26 May 2006, van Velzen R. 99 (WAG); Bipindi, 3.08°N, 10.42°E, 01 January 1900, Zenker G.A. 2268 (COI,G,G,K,L,M,MO,P,S,WAG); Bipindi, 3.08°N, 10.42°E, 01 January 1907, Zenker G.A. 3330 (BR,G,UPS).
Isolona madagascariensis (A.DC.) Engl. & Diels (a species from Madagascar).
Trees, 3–30 m tall, d.b.h. 5–60 cm; stilt roots or buttresses absent. Indumentum of simple hairs or glabrous. Leaves: petiole 1–15 mm long, 1–4 mm in diameter, blade 8.5–29 cm long, 3–15 cm wide, elliptic or obovate or oblong, apex acuminate, base decurrent to rounded or acute, concolorous; midrib raised on upper surface; secondary veins 7 to 20 pairs; tertiary venation reticulate. Inflorescences ramiflorous on old leafless or young foliate branches, axillary. Flowers bisexual with 9 perianth parts in 2 whorls, 1 to 3 per inflorescence; pedicel 1–25 mm long, 1–2 mm in diameter; in fruit 2–29 mm long, 2–5 mm in diameter; bracts 2 to 7, several basal and one upper, lower half of pedicel; sepals 3, valvate, free, 1–9 mm long, 2–5 mm wide, ovate or elliptic, apex acute or acuminate or rounded, base truncate; petals 6, basally fused, tube 3–11 mm long, inner and outer whorl not differentiated, equal; lobes 6–31 mm long, 2–12 mm wide; stamens numerous, in 3 to 4 rows, 1–2 mm long, broad; connective discoid; staminodes absent; carpels fused - syncarpous, forming a single visible gynoecium, 1–3 mm long, stigma bilobed, slightly capitate or capitate. Fruit syncarpous, forming a single visible fruit, 30–90 mm long, 15–50 mm in diameter, ovoid or ellipsoid, apex apiculate or rounded or cuspidate; seeds numerous not seriate, 8–25 mm long, 5–15 mm in diameter, ellipsoid or flattened ellipsoid; aril absent.
A genus of trees with 20 known species, 15 in Africa and 5 in Madagascar. In Cameroon nine species are known, none endemic.
Isolona, together with its sister genus Monodora, are unique in Annonaceae in having truly syncarpous flowers (fused carpels) and fruits. This translates into single fruits with unordered seeds, in contrast to other genera which have either uni- or biseriate placentation. Petals in Isolona are basally fused forming a clearly visible tube, with six equal lobes of equal length in a single whorl. In the vegetative state, Isolona and Monodora (together with Polyceratocarpus pellegrinii) are characterized by a raised midrib, in contrast to a sunken or flat midrib in all other genera found in Cameroon.
1 | Leaves and/or young foliate branches pubescent | 2 |
– | Leaves and young foliate branches completely glabrous | 3 |
2 | Leaf blade inserted on top of petiole; lobes glabrous outside, pubescent inside | I. congolana |
– | Leaf blade inserted on side of petiole; lobes pubescent on both sides | I. pilosa |
3 | Leaf blade inserted on top of petiole | 4 |
– | Leaf blade inserted on sides of petiole | 5 |
4 | Flowering pedicels 14–25 mm long, corolla smooth in dried material; corolla lobes 8–15 mm long with flat margins; fruits not ribbed | I. cooperi |
– | Flowering pedicels 3–7 mm long, corolla clearly verrucose in dried material; corolla lobes 15–25 mm long with margins curving inwards; fruits ribbed longitudinally | I. zenkeri |
5 | Sepals 4–9 mm long, papyraceous; upper bract, when present, halfway up the pedicel or subbasal, sometimes leaf-like; flowers campanulate | I. campanulata |
– | Sepals 1–3 mm long, coriaceous; upper bract absent or minute; flowers not campanulate | 6 |
6 | Base of leaf blade acute to obtuse | I. hexaloba |
– | Base of leaf blade decurrent to narrowly cuneate | 7 |
7 | Corolla lobes 4–10 times as long as wide; sepal margins glabrous | I. thonneri |
– | Corolla lobes 1.6–3.5 times as long as wide; sepal margins covered with short appressed hairs | 8 |
8 | Flowering pedicels 2–7 mm long; corolla lobes with rounded tips, narrowly elliptic to elliptic, the margins sparsely covered with short hairs | I. dewevrei |
– | Flowering pedicels 10–20 (-23) mm long; corolla lobes with acute tips, narrowly ovate to ovate, the margins glabrous | I. pleurocarpa |
= Isolona leonensis
Sprague & Hutch., Bull. Misc. Inform. Kew: 151, 1916. Type. Sierra Leone. Northern Province, Yonibana, Thomas N.W. 4230, 30 Oct 1914: lectotype, designated by
= Isolona soubreana
A.Chev., Explor. Bot. Afrique Occ. Franc. 1: 12, 1920. Type. Ivory Coast. Sassandra, Chevalier A.J.B. 19088, 23 Jun 1907: lectotype, designated by
Cameroon. Northern Region; Bangwe, Conrau G. 93, 17 Oct 1899: holotype: B[B 10 0154200]; isotype: K[K000198841].
Tree to shrub, 10–15 m tall, d.b.h. 10–15 cm; stilt roots or buttresses absent. Indumentum absent; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 3–6 mm long, 1–2 mm in diameter, glabrous, weakly grooved adaxially, blade inserted on the side of the petiole; blade 10–18 cm long, 3–7 cm wide, elliptic to obovate, apex acuminate, acumen 1 cm long, base decurrent to cuneate, subcoriaceous, below glabrous when young and old, above glabrous when young and old, concolorous; midrib raised, above glabrous when young and old, below glabrous when young and old; secondary veins 7 to 12 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 2 per inflorescence; pedicel 5–30 mm long, ca. 1 mm in diameter, glabrous; in fruit 2–5 mm long, 2–3 mm in diameter, glabrous; bracts 2 to 5, several basal and one upper towards the lower half of pedicel, basal bracts 2–3 mm long, 1–2 mm wide; upper bract sometimes leaf-like, 2–19 mm long, 1–4 mm wide; sepals 3, valvate, free, 4–9 mm long, 3–5 mm wide, elliptic to ovate, apex acute, base truncate, green speckled with red and purple, glabrous inside, glabrous outside, margins flat; petals basally fused, tube 4–7 mm long, inner and outer whorl not differentiated, equal; lobes 6–20 mm long, 2–7 mm wide, triangular, apex acute, green, turning bright yellow, margins flat, glabrous outside, glabrous inside, spreading horizontally; stamens numerous, in 3 to 4 rows, ca. 1 mm long, broad; connective discoid, glabrous, cream; staminodes absent; carpels fused into a single structure, ca. 2 mm long, stigma capitate, glabrous. Fruit syncarpous, sessile, 40–75 mm long, 20–35 mm in diameter, ovoid, apex rounded, glabrous, smooth, bumpy, green turning deep yellow when ripe; seeds not counted, 8–15 mm long, 5–10 mm in diameter, ellipsoid; aril absent.
A widespread species with a disjunct distribution in West (Sierra Leone, Liberia, Ivory Coast and Ghana) and Central Africa (Cameroon, Gabon); in Cameroon known from South, Central, Littoral and South-West regions.
An infrequent species in Cameroon; in lowland primary and secondary forests, also along rivers. Altitude 0–500 m a.s.l.
Isolona campanulata A flower, side view B flower, top view. Isolona congolana C flowering branch D base of leaf blade, top view E flower, top view F flower, bottom view G syncarpous fruit A, B no voucher, Bayang Mbo, Cameroon C–G Couvreur 1054, Mt Cameroon. Photos A, B Sonneck C–G Thomas L.P. Couvreur .
None recorded.
Least Concern (LC) (
medicine : root against rheumatism, allay fever.
This species is characterized by its large, foliaceous, and glabrous sepals, not appressed against the corolla tube, as well as the sometimes leaf-like upper bract. It is also a completely glabrous species, a character only shared with I. cooperi and I. hexaloba. The flower of I. campanulata is very distinct and uniquely campanulate.
Littoral Region: Lombe amp Tissongo Study Area Doula-Edea Reserve, 3.78°N, 10.04°E, 01 June 1976, Waterman P.G. 830 (K). South Region: ca 15 km SE of Kribi Kienke Forest Res at Bidou II, 2.86°N, 10.03°E, 30 June 1969, Bos J.J. 4947 (BR,P,WAG,YA); 10 km From Kribi Lolodorf road, 2.96°N, 9.966°E, 09 July 1970, Bos J.J. 7069 (WAG). South-West Region: 5.46°N, 9.883°E, 7 October 1899, Conrau G. 93 (B,K); 16 minutes from research station towards rivers, 5.34°N, 9.496°E, 27 November 2000, Gosline W.G. 289 (K); Ente Ekondo Nene et Loe 15 km NW Ekondo Titi, 4.69°N, 8.97°E, 03 June 1976, Letouzey R. 15078 (P,YA); Korup National Park, 5.31°N, 8.966°E, 02 July 1951, Olorunfemi J. 30662 (K); Korup National Park between the Ndian River at PAMOL field and 25 km on transect “ P ”, 5.01°N, 8.833°E, 12 April 1985, Thomas D.W. 4763 (BR,K,MO,P,WAG,YA); Takamanda Forest Reserve, 6.21°N, 9.433°E, 30 April 1987, Thomas D.W. 7354 (MO,P,WAG).
≡ Monodora congolana De Wild. & T. Durand, Bull. Soc. Roy. Bot. Belgique 38: 13, 1899.
= Isolona maitlandii Keay, Kew Bull. 7(2): 155, 1952. Type. Cameroon. North-West Province, Ngong, Maitland T.D. 1555, Jun 1931: holotype: K[K000105576]; isotypes: BM, FHO.
Democratic Republic of the Congo. Equateur; Lukandu, Dewèvre A.P. 1103, 19 Nov 1896: lectotype, sheet here designated: BR[BR0000006248932]; isotypes: BR[BR0000006248857, BR0000006249588, BR0000006249250].
Tree, 10–30 m tall, d.b.h. 5–45 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches densely pubescent. Leaves: petiole 4–7 mm long, 2–3 mm in diameter, densely pubescent, grooved, blade inserted on top of the petiole; blade 13–19 cm long, 4–5 cm wide, narrowly ovate to narrowly elliptic or oblong, apex acuminate, acumen 0.5–1 cm long, base cuneate to rounded, subcoriaceous, below sparsely pubescent when young, glabrous when old, above sparsely pubescent when young, glabrous when old, concolorous; midrib raised, above sparsely pubescent when young, glabrous when old, below sparsely pubescent when young, glabrous when old; secondary veins 11 to 14 pairs, sparsely pubescent to glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless or young foliate branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 3 per inflorescence; pedicel 10–23 mm long, 1 mm in diameter, sparsely pubescent to glabrous; in fruit 2–5 mm long, 2–3 mm in diameter, glabrous; bracts 3 to 4, several basal and one upper towards the lower half of pedicel, basal bracts 1 mm long, 1 mm wide; upper bracts 1 mm long, 1 mm wide; sepals 3, valvate, free, 3 mm long, 2 mm wide, ovate, apex acute, base truncate, green, pubescent outside, glabrous inside, margins flat; petals basally fused, tube 4–10 mm long, inner and outer whorl not differentiated, equal; lobes 9–20 mm long, 3–7 mm wide, elliptic to ovate, apex rounded, green turning red, margins wavy, glabrous outside, densely pubescent inside, spreading horizontally; stamens numerous, in 3 to 4 rows, 2 mm long, broad; connective discoid, densely pubescent, cream; staminodes absent; carpels fused into a single structure, 2 mm long, stigma bilobed, slightly capitate, densely pubescent. Fruit syncarpous, sessile, 60–80 mm long, 40–50 mm in diameter, ellipsoid, apex rounded, glabrous, smooth to verrucose, irregularly ribbed, green when ripe; seeds not counted, 15–25 mm long, 10–15 mm in diameter, ellipsoid; aril absent.
A Isolona congolana B Isolona cooperi C Isolona dewevrei D Isolona hexaloba E Isolona pilosa F Isolona pleurocarpa G Isolona thonneri H Isolona zenkeri I Letestudoxa bella. White borders represent region limits in Cameroon; green patches represent protected areas (see methods and Suppl. material
A central African species, with a disjunct distribution between the Cameroon Volcanic Line in Cameroon, and Eastern Democratic Republic of Congo, one collection from Central African Republic; in Cameroon known from the Littoral, North-West and South-West regions.
A common species when present, mainly in montane or premontane rain forests, along rivers and bush land. Altitude 800–1700 m a.s.l.
Ndin (Westphal 10012).
Least Concern (LC) (
food : fruit for condiments, spices, flavourings (Westphal 10012).
This the only species of Isolona growing above 1000 m in Cameroon. Isolona congolana is characterized by the densely pubescent inner part of the corolla tube and a glabrous outer part of the tube, a unique feature in this genus (I. pilosa is pubescent on both sides of the corolla). The leaves are also characteristic being narrowly ovate to narrowly elliptic or oblong, giving a unique aspect to the foliage.
Littoral Region: Manengouba Mt 4 km WNW Of Nkongsamba, 4.98°N, 9.900°E, 04 April 1972, Leeuwenberg A.J.M. 9550 (B,BR,C,GC,H,K,LD,M,MO,P,WAG,YA). North-West Region: Kagwene, 6.12°N, 9.734°E, 20 May 2009, Ashworth J. 196 (K,YA); Bamenda Distr Ngong, 6.58°N, 10.43°E, 01 June 1931, Maitland T.D. 1555 (BM,K); Bamenda Wae, 6°N, 10.41°E, 01 April 1931, Maitland T.D. 1596 (K). South-West Region: Nyasoso, 4.86°N, 9.7°E, 04 June 1996, Cable S. 2843 (K,MO,WAG,YA); 2 km north of Nyasoso towards Mpako, 4.84°N, 9.679°E, 04 April 2016, Couvreur T.L.P. 1054 (WAG,YA); Nyasoso, 4.83°N, 9.683°E, 19 March 1996, Etuge M. 1794 (K,WAG); White trail (above Kupe village) towards Madam 4.78°N, 9.716°E, 28 May 1996, Etuge M. 2000 (BR,K,MO,P,WAG,YA); Lake Edip, 4.95°N, 9.65°E, 21 November 1998, Etuge M. 4488 (K,WAG,YA); Etube Tape from Nyasoso, 4.83°N, 9.683°E, 06 February 1995, Lane P. 532 (K,YA); Bu 4.15°N, 9.233°E, 6 June 1898, Lehmbach H. 224 (B) . West Region: Bali Ngemba Forest Reserve, 5.82°N, 10.08°E, 13 April 2004, Etuge M. 5431 (K,WAG,YA); Along the road 6 km W of Dschang on road to Fongo Ndeng, 5.45°N, 9.95°E, 15 May 1978, Westphal E. 10012 (P,WAG).
Liberia. Montserrado; near Firestone plantations, along Dukwai road, Cooper G.P. 417, 7 May 1929: lectotype, designated by
Tree, 6–18 m tall, d.b.h. 20 cm; stilt roots or buttresses absent. Indumentum absent; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 1–5 mm long, 2–3 mm in diameter, glabrous, grooved, blade inserted on top of the petiole; blade 15–29 cm long, 6–15 cm wide, oblong to obovate, apex acuminate, acumen 1–2 cm long, base cuneate to rounded, subcoriaceous, below glabrous when young and old, above glabrous when young and old, concolorous; midrib raised, above glabrous when young and old, below glabrous when young and old; secondary veins 9 to 18 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences cauliflorous or ramiflorous on young foliate branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 2 per inflorescence; pedicel 14–25 mm long, 1 mm in diameter, glabrous; in fruit 16–27 mm long, 2 mm in diameter, glabrous; bracts 2 to 4, all basal, 1 mm long, 1mm wide; sepals 3, valvate, free, 2 mm long, 2 mm wide, ovate, apex rounded, base truncate, green, glabrous outside, glabrous inside, margins flat; petals basally fused, tube 6–11 mm long, inner and outer whorl not differentiated, equal; lobes 8–15 mm long, 4–6 mm wide, oblong, apex acute to rounded, green, margins flat, glabrous outside, glabrous inside, spreading horizontally; stamens numerous, in 3 to 4 rows, 2 mm long, broad; connective discoid, glabrous, green; staminodes absent; carpels fused into a single structure, 3 mm long, stigma capitate, glabrous. Fruit syncarpous, sessile, 30–90 mm long, 15–30 mm in diameter, ellipsoid, apex apiculate, glabrous, smooth, constricted over seeds in dried material, smooth when fresh, orange with white spots when ripe; seeds not counted, 10–15 mm long, 5–10 mm in diameter, ellipsoid; aril absent.
A mainly West African species, from Liberia to Ghana, with a few specimens from Cameroon and one from Gabon; in Cameroon known from South and South-West regions.
A rare species in Cameroon; in lowland primary and secondary forests, also along rivers, on sandy soils. Altitude 0–300 m a.s.l.
None recorded.
Least Concern (LC) (
None reported.
Isolona cooperi is distinguished by its completely glabrous leaves, young foliate branches and flowers and with the leaf blade inserted on top of the petiole. In addition, the flowers emit a very strong sweet scent, noticeable even in dried material. It has a smooth corolla in dried material and corolla lobes with straight margins. Isolona cooperi is similar to I. hexaloba by the shape of its flowers, but the latter differs by its blade inserted sideways to the petiole and the absence of the strong sweet scent. Finally, I. cooperi resembles I. campanulata by the shape of the fruits.
Isolona cooperi A flowering branch B leaf, lower view C detail of petiole and axillary inflorescence D flower bud E flower, side view F flower, semi side view G flower, bottom view H transversal section showing androecium and stigma I stamen, front view J fruit (dried) K fruit, part of pericarp removed showing ruminate section of seed (fresh) L seed with seed coat partially removed showing ruminations A–C from J.J.F.E. de Wilde 3644 D–I from fresh material collected at the Utrecht University Botanical Garden J, L from de Koning 149 K from Breteler 7458. Drawings Hans by Vries (
South Region: Bipindi, 3.08°N, 10.41°E, 01 February 1910, Zenker G.A. s.n. (F). South-West Region: Korup National Park, 5.06°N, 8.855°E, 10 March 1998, Kenfack D. 1063 (MO,WAG).
≡ Monodora dewevrei De Wild. & T. Durand, Bull. Soc. Roy. Bot. Belg., Compt. Rend. 38: 11, 1899.
Democratic Republic of the Congo. Bas Congo; Lemba-Luki, Dewèvre A.P. 365, no date: lectotype, sheet here designated: BR[BR0000008801050]; isotypes: BR[BR0000008799968, BR0000008800725, BR0000008799630].
Tree to shrub, 8–15 m tall, d.b.h. 20 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 4–15 mm long, 1–2 mm in diameter, glabrous, grooved, blade inserted on the side of the petiole; blade 10–17 cm long, 4–7 cm wide, elliptic to obovate, apex acuminate, acumen ca. 1 cm long, base decurrent to cuneate, papyraceous, below glabrous when young and old, above glabrous when young and old, concolorous; midrib raised, above glabrous when young and old, below glabrous when young and old; secondary veins 9 to 14 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 per inflorescence; pedicel 2–7 mm long, 1 mm in diameter, sparsely pubescent; in fruit 9–10 mm long, 2–3 mm in diameter, glabrous; bracts 3 to 5, several basal and one upper towards the lower half of pedicel, basal bracts 1 mm long, 0.5–1 mm wide; upper bract 1 mm long, 1 mm wide; sepals 3, valvate, free, 2–3 mm long, 3–4 mm wide, ovate, apex acuminate, base truncate, glabrous outside, glabrous inside, margins flat; petals basally fused, tube 3–4 mm long, inner and outer whorl not differentiated, equal; lobes 7–17 mm long, 5–7 mm wide, elliptic, apex rounded, green, margins flat, overall glabrous but pubescent towards margins outside and inside; margins curved outwards; stamens ca. 50, in 3 to 4 rows, 2 mm long, broad; connective discoid, glabrous; staminodes absent; carpels fused into a single structure, ca. 1 mm long, stigma capitate, glabrous. Fruit syncarpous, sessile, 60–70 mm long, 40–50 mm in diameter, ovoid, apex rounded, glabrous, smooth, not ribbed, green when ripe; seeds not counted, 10–20 mm long, 10–15 mm in diameter, ellipsoid; aril absent.
A widespread species, but with few overall specimens, from Liberia to Nigeria, and from Cameroon to Democratic Republic of Congo; in Cameroon in the South and South-West regions.
Isolona dewevrei A flowering branch B flower (side view) C flower top view D transversal section of flower E detail of stig top view F transversal section of seed G fruit (left) and longitudinal section of fruit (right) H seed, side view (top), detail of hilum (bottom). Modified from Aké Assi (1963, fig. 1, p. 14).
A rare species in Cameroon known from two specimens; in lowland primary and secondary forests. Altitude 0–860 m a.s.l.
None recorded.
Least Concern (LC) (
None reported.
Isolona dewevrei resembles I. thonneri in leaf and fruit shape, but the former can be distinguished by its elliptic and shorter corolla lobes with hairy margins and its flowering pedicel sparsely covered with short hairs. It is, however, very hard to distinguish both species apart based on fruit or vegetative characters alone.
South Region: Ngongondje hill near Akonetye 2 30'S of Ebolowa, 2.5°N, 11.13°E, 28 August 1979, Koufani A. 123 (P,YA). South-West Region: Piste Munkep-Gayama 40 km NNW Wum, 6.73°N, 9.95°E, 08 July 1975, Letouzey R. 13984 (K,MO,YA).
≡ Monodora hexaloba Pierre, Fig. Herb. L. Pierre, del. E. Delpy 5/1896, 1896.
= Isolona bruneelii De Wild., Ann. Mus. Congo Belge, Bot. ser. 5, 3(1): 82, 1909. Type. Democratic Republic of the Congo. Orientale, Dikila, Bruneel A.C.E. s.n., Dec 1906: lectotype, sheet here designated: BR[BR0000008800152] ; isotypes: BR[BR0000008799692, BR0000008799364, BR0000008799302]; S[S10-20956].
= Isolona seretii De Wild., Ann. Mus. Congo Belge, Bot. ser. 5, 3[1]: 82, 1909. Type. Democratic Republic of the Congo. Equateur, near Nala, Seret F. 792, Mar 1907: lectotype, sheet here designated: BR[BR0000008800992]; isotype: BR[BR0000008801326].
= Isolona solheidii De Wild., Ann. Mus. Congo Belge, Bot. sér. 5, 3: 83, 1909. Type. Democratic Republic of the Congo. Equateur, surroundings of Yambuya, Solheid A.F. s.n., 1906: lectotype, sheet here designated: BR[BR0000008799395]; isotype: BR[BR0000008799722].
= Isolona seretii var. grandifolia De Wild., Ét. Fl. Bang. Ub.: 313, 1911. Type. Democratic Republic of the Congo. Orientale, Mobwasa, Claessens J. 615, Apr 1910: holotype: BR.
= Isolona pleurocarpa subsp. nigerica Keay, Kew Bull. 7: 157, 1952. Type. Nigeria. Ijebu State, Shasha Forest Reserve, Richards P.W. 3343, 8 Apr 1935: holotype: BM[BM000546386]; isotypes: BR[BR0000014130090]; G[G00011543, G00011544]; MO[MO-2246487]; S[S10-20983].
Gabon. Estuaire; Environs de Libreville, Klaine T.-J. 360, 17 Feb 1896: lectotype, sheet here designated: P[P00363270]; isotypes: B; K[K00198842]; P[P00363269, P00363268]; WAG[WAG0251603].
Tree to shrub, 15–30 m tall, d.b.h. up to 50 cm; stilt roots or buttresses absent, trunk deeply fluted. Indumentum absent; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 2–4 mm long, 2–3 mm in diameter, glabrous, grooved, blade inserted on the side of the petiole; blade 10–28 cm long, 3–11 cm wide, ovate to elliptic, apex acuminate, acumen 1–2 cm long, base obtuse to acute, coriaceous, below glabrous when young and old, above glabrous when young and old, concolorous; midrib raised, above glabrous when young and old, below glabrous when young and old; secondary veins 8 to 16 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 2 per inflorescence; pedicel 7–30 mm long, ca. 1 mm in diameter, glabrous; in fruit 20–29 mm long, 3–5 mm in diameter, glabrous; bracts 3 to 5, several basal and one upper towards the lower half of pedicel, basal bracts ca. 1 mm long, 0.5 mm wide; upper bract 2–5 mm long, ca. 1 mm wide; sepals 3, valvate, free, 1–3 mm long, 2–4 mm wide, elliptic, apex acuminate, base truncate, green, glabrous outside, glabrous inside, margins flat; petals basally fused, tube 4–10 mm long, inner and outer whorl not differentiated, equal; lobes 6–25 mm long, 4–12 mm wide, elliptic to ovate, apex acute to rounded, dark red, margins flat to wavy, glabrous outside, glabrous inside, spreading horizontally; stamens numerous, in 3 to 4 rows, 2 mm long, broad; connective discoid, glabrous, green; staminodes absent; carpels fused into a single structure, ca. 1 mm long, stigma bilobed, slightly capitate, glabrous. Fruit syncarpous, sessile, 30–70 mm long, 25–40 mm in diameter, ovoid, apex rounded, glabrous, smooth, bumpy, irregularly and transversely ribbed, light green to dark purple when ripe; seeds not counted, 8–15 mm long, 4–6 mm in diameter, ellipsoid; aril absent.
A widespread species from Nigeria to the Democratic Republic of Congo and northern Angola (one specimen); in Cameroon known from Adamaoua, East, South, Central, Littoral and South-West, North-West regions.
A relatively rare species in Cameroon; in lowland primary and secondary evergreen forests, but also in semi-deciduous forests, also along rivers. Altitude 0–700 m a.s.l.
None recorded.
Least Concern (LC) (
None reported.
Isolona hexaloba is distinguished by the short and grooved petiole with the leaf blade inserted on its sides. The corolla lobes are coriaceous and can be quite variable in shape, ranging from elliptic to obovate, with a narrowed base. It is a polymorphic species and has been described under several names, now all reduced to synonymy (
Central Region: Tibati près Mbatimbang, 6.14°N, 12.48°E, 04 December 1959, Letouzey R. 2392 (P,YA). East Region: A 25 km environ à l’ENE de Mikel village situé à 85 km au N de Moloundou sur la route de Yokadouma 2.93°N, 15.33°E, 24 February 1971, Letouzey R. 10419 (P,WAG,YA); A 14 km à l’Ouest de Yenga Port Gentil village situé à 35 km au NNE de Moloundou, 2.35°N, 15.35°E, 21 April 1971, Letouzey R. 10703 (BR,COI,K,P,WAG,YA); A 20 km au Sud de Mboy I (45 km à l’Est de Yokadouma), 3.38°N, 15.13°E, 16 May 1963, Letouzey R. 5072 (K,P,WAG,YA). Littoral Region: Douala (route Razel), 4.05°N, 9.71°E, 01 January 1955, Endengle E. SRFK 2121 (P,YA); Roue forestière SNCB (km 36 vers Ndoksom) environ 25 km Sud Yabassi, 4.31°N, 9.958°E, 11 May 1976, Letouzey R. 14910 (C,K,MO,P,WAG,YA); Tissongo, 3.58°N, 9.9°E, 01 August 1976, McKey D.B. 245 (K). South Region: ca 15 km east from Lélé village, 2.28°N, 13.32°E, 10 September 2013, Couvreur T.L.P. 495 (WAG,YA); near Bipaga II km 40 road Kribi-Edéa, 3.15°N, 10.01°E, 30 December 1982, de Kruif A.P.M. 998 (MO,WAG,YA); 17 km S of the Lobe river along the road to Campo, 2.81°N, 10.13°E, 18 March 1975, de Wilde J.J.F.E 8088 (BR,K,MO,P,U,WAG,YA); A l’Ouest d’Alati (100 km SE de Djoum), 2.2°N, 13.42°E, 13 January 1973, Letouzey R. 11840 (K,P,YA); 22 km on road Kribi to Campo 12 km past Gross Batanga, 2.76°N, 9.881°E, 24 February 1994, Wieringa J.J. 2327 (MPU,U,WAG); Bipindi, 3.08°N, 10.41°E, 01 November 1901, Zenker G.A. s.n. (P). South-West Region: Korup National Park, 5.01°N, 8.783°E, 31 October 2005, van der Burgt X.M. 791 (BR,G,K,MO,P,WAG,YA). West Region: Bali- Ngemba FR, 5.81°N, 10.08°E, 13 April 2002, Onana J.M. 2030 (K,WAG).
Isolona hexaloba A, B flowering branch C transversal section of flower showing androecium and stigma D flower (bottom view) E stamens inner row (2 top); stamens of inner most row (top) and outermost row (bottom) F carpel and detail of ovules G fruits, the one on the right opened to show seeds H seed (left) and transversal section of seed showing ruminate endosperm (right) A, C–F from Le Testu 5862 B Le Testu 5836 G, H Klaine 360. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
Isolona hexaloba A trunk, note deeply furrowed B base of leaf blade, note raise midrib C flower, top view D flower, side view E fruiting branch. Isolona pleurocarpa F trunk G bade of leaf blade, notice decurrent base H flower, top view, fallen on ground I syncarpous fruit; fallen on ground A Sosef 2032, Gabon B–D Couvreur 561, Gabon E Texier 2347, Gabon F–I Couvreur 402, Ngovayang, Cameroon. Photos A–D, F–I Thomas L.P. Couvreur E Nicolas Texier, Tropicos.org, Missouri Botanical Garden.
= Isolona theobromina Exell, J. Bot. 64 (Suppl. 1): 10, 1926. Type. Angola. Cabinda, Pango Munga, Gossweiler J. 6112, 7 Jan 1916: holotype: BM[BM000889332]; isotypes: COI[COI00077211]; LISJC n.v.; LISC[LISC000094, LISC000095, LISC000096].
Democratic Republic of the Congo. Kasai Oriental; Lualaba, Ledermann C. 11, Mar 1906: holotype: B[B 10 0154216]; isotype: K n.v.
Tree, 13 m tall, d.b.h. up to 50 cm; stilt roots or buttresses absent, trunk not fluted. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches densely pubescent. Leaves: petiole 3–12 mm long, 3–4 mm in diameter, densely pubescent, grooved, blade inserted on the side of the petiole; blade 19–27 cm long, 6–10 cm wide, obovate, apex acuminate, acumen 1–2 cm long, base rounded to cordate, papyraceous, below densely pubescent when young, densely pubescent when old, above sparsely pubescent to glabrous when young, sparsely pubescent to glabrous when old, concolorous; midrib raised above, above sparsely pubescent when young, sparsely pubescent when old, below densely pubescent when young, densely pubescent when old; secondary veins 15 to 20 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 2 per inflorescence; pedicel 2–4 mm long, 1–2 mm in diameter, densely pubescent; in fruit 2–8 mm long, 2–3 mm in diameter, pubescent; bracts 2 to 4, all basal, 2–4 mm long, 1–2 mm wide; sepals 3, valvate, free, 2–5 mm long, 2–4 mm wide, ovate, apex acuminate, base truncate, densely pubescent outside, glabrous inside, margins flat; petals basally fused, tube 5–10 mm long, inner and outer whorl not differentiated, equal; lobes 8–13 mm long, 3–5 mm wide, elliptic, apex acute, yellow, margins flat, densely pubescent outside, pubescent with base glabrous inside, curving inwards over the receptacle; stamens numerous, in 3 to 4 rows, 2 mm long, broad; connective discoid, glabrous; staminodes absent; carpels fused into a single structure, 2 mm long, stigma bilobed, glabrous. Fruit syncarpous, sessile, 30–60 mm long, 20–40 mm in diameter, ellipsoid, apex cuspidate, sparsely pubescent, longitudinally ribbed, color unknown; seeds not counted, 13–15 mm long, 6–8 mm in diameter, flattened ellipsoid; aril absent.
Isolona letestui (not in Cameroon) A flowering branch B opened flower showing androecium and stigma C carpel (left) and transversal section of carpel (right) D stamens of innermost row (2 top) and stamen of outermost row (bottom). Isolona pilosa E flowering branch F opened flower showing androecium and stigma G stamen of outermost row (left) and innermost row (right) A–D from Le Testu 1252; E from Le Testu 8740 F, G from Le Testu 8602. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
A central African species with a disjunct distribution, from Cameroon to Gabon and the Republic of Congo, also present in the Democratic Republic of Congo; in Cameroon known from the extreme East region.
A rare species in Cameroon; in lowland rain forests or swampy areas. Altitude 100–450 m a.s.l.
None recorded.
Vulnerable B2ab(iii) (
None reported.
Isolona pilosa is the most pubescent species of the genus. It is distinguishable by its densely pubescent leaf midrib on the upper side (even in older leaves), as well as its short and densely hairy flowering pedicels. The corolla lobes are densely hairy on the outside and near the margins on the inside; the inner part of the tube is glabrous, which distinguishes it from I. congolona which is pubescent on the inner surface of the tube but glabrous on the outside.
Several specimens collected by Harris DJ are reported from The Lobéké National Park in East Cameroon, but we were not able to see them and verify their identification (e.g. 6408, 6538, 6627).
East Region: Region near Station Molundu Dscha (Ngoko) Nginda 21 km north Molundu, 2.2°N, 15.2°E, 07 January 1911, Mildbraed G.W.J. 4193 (HBG).
= Isolona leucantha Diels, Bot. Jahrb. Syst. 39: 484, 1907. Type. Cameroon. South Region, Bipindi, Zenker G.A. 3038, Apr 1904: holotype: B[B 10 0154212]; isotypes: BM[BM000546385]; BR[BR0000008802644]; COI[COI00077204]; E[E00259306]; G[G00011566]; GOET[GOET005681]; HBG[HBG518922]; K[K000199011]; L[L-0182762]; M[M0089224]; MO[M0089224]; P[P00363267]; S[S-G-7462]; WAG[WAG0000090]; WU[WU0025864].
Cameroon. South Region; Bipindi, Zenker G.A. 3217, Jul 1904: holotype: B[B 10 0154211]; isotypes BR[BR0000008498991]; G[G00011576 G00011761]; K[K000198837] ; S[S10-21236]; WU[WU0025885].
Tree, 15–30 m tall, d.b.h. up to 60 cm; stilt roots or buttresses absent, trunk fluted. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 4–12 mm long, 1 mm in diameter, glabrous, grooved towards the base, blade inserted on the side of the petiole; blade 8.5–15.5 cm long, 3–6 cm wide, elliptic to obovate, apex acuminate, acumen 1–1.5 cm long, base decurrent to cuneate, papyraceous to subcoriaceous, below glabrous when young and old, above glabrous when young and old, concolorous; midrib raised above, above glabrous when young and old, below glabrous when young and old; secondary veins 9 to 12 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 per inflorescence; pedicel 1–2 mm long, 1 mm in diameter, glabrous; in fruit 30 mm long, 3–4 mm in diameter, glabrous; bracts 3 to 7, several basal and one upper towards the lower half of pedicel, basal bracts ca. 1 mm long, ca. 1 mm wide; upper bracts ca. 1 mm long, ca. 1 mm wide; sepals 3, valvate, free, 2–3 mm long, 2–3 mm wide, ovate, apex acuminate, base truncate, green, glabrous outside, glabrous inside, margins flat; petals basally fused, tube 6–15 mm long, red, inner and outer whorl not differentiated, equal; lobes 10–23 mm long, 5–10 mm wide, ovate, apex acute, bright green-white to yellow, margins wavy, glabrous outside, glabrous inside, spreading horizontally; stamens numerous, in 3 to 4 rows, ca. 2 mm long, broad; connective discoid, glabrous, yellow; staminodes absent; carpels fused into a single structure, ca. 2 mm long, stigma bilobed, slightly capitate, glabrous. Fruit syncarpous, sessile, ca. 50 mm long, ca. 40 mm in diameter, globose, apex rounded, glabrous, rugulose, longitudinally 6–8 ribbed, green when unripe; seeds not counted, 8–10 mm long, 5–7 mm in diameter, ellipsoid; aril absent.
Known from Nigeria and Cameroon; in Cameroon known from the South and South-West regions.
An infrequent species; in lowland rain forests on non-inundated soils. Altitude 0–550 m a.s.l.
Avom (van Andel 4177).
Endangered (EN) (
None reported.
Isolona pleurocarpa is distinguished by the combination of these characters: young leaves glabrous, leaf blade inserted on the side of the petiole and decurrent to narrowly cuneate at base, midrib proximally depressed above, corolla lobes narrowly ovate to ovate with a narrowed base and an acute apex, undulate-wavy on the margins when dried.
South Region: NE of Mt Elephant ca 20 km SE of Kribi, 2.8°N, 10.03°E, 10 February 1970, Bos J.J. 6298 (WAG); mountain chain Ngovoyang 42 km in forest from Bikiliki village situated between Bipindi and Lolodorf, 3.18°N, 10.53°E, 19 February 2012, Couvreur T.L.P. 402 (WAG,YA); Colline Nkolo Manga (20 km SE Kribi), 2.95°N, 9.916°E, 16 April 1968, Letouzey R. 9341 (P,WAG); Elephant Mont, 2.8°N, 10.01°E, 22 October 2001, van Andel T.R. 4177 (KRIBI,WAG,YA); Bipindi, 3.08°N, 10.41°E, 1895, Zenker G.A. 1716 (B,G,M,P,WAG); Bipindi, 3.08°N, 10.41°E, 01 January 1918, Zenker G.A. 22 (P,WAG); Mbiave, 3.21°N, 10.61°E, 01 January 1913, Zenker G.A. 267 (A,B,BR,C,G,M,MO,U,WAG); Bipindi, 3.08°N, 10.41°E, 01 April 1904, Zenker G.A. 3038 (B,COI,G,K,L,M,MO,P,S,WAG); Bipindi, 3.08°N, 10.41°E, 01 July 1904, Zenker G.A. 3217 (B,BR,G,K,L,M,S,WAG); Bipindi, 3.08°N, 10.41°E, 01 January 1908, Zenker G.A. 3433 (BR,COI,G,G,L,M,M,MO,P,S); Bipindi, 3.08°N, 10.41°E, 01 January 1908, Zenker G.A. 3540 (G,K,L,M,M,MO); Bipindi, 3.08°N, 10.41°E, 01 January 1909, Zenker G.A. 3921 (B,BR,COI,COI,G,L,M,M,MO,P,S); Bipindi, 3.08°N, 10.41°E, 01 January 1913, Zenker G.A. 4704 (BM,BR,G,K,L,M,P,S); Bipindi, 3.06°N, 10.38°E, 01 November 1919, Zenker G.A. 95 (BM). South-West Region: Southern Bakundu Forest 3 km from Kindongi Camp, 4.55°N, 9.416°E, 02 May 1972, Leeuwenberg A.J.M. 9784 (B,BR,C,H,K,LD,M,MO,P,WAG,YA); Korup National Park, 5.06°N, 8.783°E, 13 April 1978, Thomas D.W. 349 (K).
≡ Monodora thonneri De Wild. & T. Durand, Bull. Soc. Roy. Bot. Belg., Compt. Rend. 38: 12, 1899.
= Diospyros oblongicarpa Gürke, Bot. Jahrb. Syst. 43: 200, 1909. Type. Cameroon. South Region, Bipindi, Zenker G.A. 3471, 1908: holotype B n.v.: isotype: K[K000199009]; US[US03899523]; WU[WU0040298].
Democratic Republic of the Congo. Equateur; Massanga (près de Monveda), Thonner F. 104, 24 Sep 1896: lectotype, designated by
Tree to shrub, 3–10 m tall, d.b.h. up to 25 cm; stilt roots or buttresses absent, trunk not fluted. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 3–8 mm long, 1–2 mm in diameter, glabrous, grooved, blade inserted on the side of the petiole; blade 11–20 cm long, 4–7.5 cm wide, elliptic to obovate, apex acuminate, acumen 1–2 cm long, base decurrent to cuneate, coriaceous to subcoriaceous, below glabrous when young and old, above glabrous when young and old, concolorous; midrib raised above, above glabrous when young and old, below glabrous when young and old; secondary veins 9 to 12 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 2 per inflorescence; pedicel 5–18 mm long, ca. 1 mm in diameter, glabrous; in fruit 8–10 mm long, 3–4 mm in diameter, glabrous; bracts 3 to 7, several basal and one upper towards the lower half of pedicel, basal bracts ca. 1 mm long, ca. 1 mm wide; upper bracts ca. 2 mm long, 1 mm wide; sepals 3, valvate, free, 2–3 mm long, 1–2 mm wide, ovate, apex acute, base truncate, dark green, glabrous outside, glabrous inside, margins flat; petals basally fused, tube 3–6 mm long, inner and outer whorl not differentiated, equal; lobes 14–31 mm long, 3–5 mm wide, linear to lorate (strap-shaped), apex acute, green, margins flat, glabrous outside, glabrous inside, pendulous; stamens numerous, in 3 to 4 rows, ca. 2 mm long, broad; connective discoid, glabrous; staminodes absent; carpels fused into a single structure, ca. 2 mm long, stigma bilobed, slightly capitate, sparsely pubescent. Fruit syncarpous, sessile, 40–60 mm long, 20–35 mm in diameter, ellipsoid, apex rounded, glabrous, smooth, not or faintly ribbed, color unknown; seeds not counted, 15–18 mm long, 8–9 mm in diameter, ellipsoid; aril absent.
Isolona zenkeri A flowering branch B flower C fruit. Isolona campanulata D flowering branch; E flower E corolla lobe opened G fruit. Isolona thonneri H flowering branch I flower J fruit A from Le Testu 5117 B from Le Testu 8001 C from Klaine 2675 D–G from Aubréville 6 H, I from Vrydagh 34 J from Lebrun 2032. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
A central African species, known from Cameroon to Gabon and the Democratic Republic of Congo; in Cameroon known from the South and East regions.
An infrequent species; in lowland rain forests, near rivers and swamps. Altitude 450–750 m. a.s.l.
None recorded.
Least Concern (LC) (
None reported.
Isolona thonneri is characterized by long, narrowly elliptic to linear and glabrous corolla lobes. In the absence of flowers, it is hard to distinguish from I. dewevrei.
East Region: Près de Banana 10 km ENE de Moloundou, 2.08°N, 15.28°E, 17 April 1971, Letouzey R. 10682 (P,WAG,YA); near Ndongo ca 45 km WNW of Moloundou, 2.16°N, 14.83°E, 15 March 1973, Letouzey R. 12085 (K,P); Près Ndongo à 45 km WNW de Moloundou, 2.15°N, 14.86°E, 16 March 1973, Letouzey R. 12111 (BR,K,P,WAG,YA); près Ndongo à 40 km WNW de Moloundou, 2.15°N, 14.86°E, 16 March 1973, Letouzey R. 12115 (BR,K,P,WAG,YA). South Region: Colline Ongongondje près Akonekye 15 km NW d’Ambam, 2.46°N, 11.16°E, 23 March 1970, Letouzey R. 10205 (BR,COI,K,P,WAG,YA); Inselberg d’Akookas pres du village d’Akookas 38 km au sud est d’Ebolowa, 2.71°N, 11.27°E, 15 March 2001, Parmentier I. 1943 (BRLU,WAG); Inselberg d’Akookas pres du village d’Akookas 38 km au sud est d’Ebolowa, 2.71°N, 11.27°E, 15 March 2001, Parmentier I. 1961 (BRLU,WAG).
Cameroon. South Region; Bipindi, Zenker G.A. 1186, 1896: holotype: B[B 10 0154218]; isotypes: BM n.v., G[G00011574]; K[K000199013]; WU[WU0025863].
Tree to shrub, 7–15 m tall, d.b.h. up to 15 cm; stilt roots or buttresses absent, trunk not fluted. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches glabrous. Leaves: petiole 2–6 mm long, ca. 2 mm in diameter, glabrous, slightly grooved, blade inserted on top of the petiole; blade 16–23 cm long, 6.5–8.5 cm wide, oblong to oblanceolate, apex abruptly acuminate, acumen 1–2 cm long, base rounded to acute, coriaceous, below glabrous when young and old, above glabrous when young and old, concolorous; midrib raised above, above glabrous when young and old, below glabrous when young and old; secondary veins 11 to 13 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on young foliate branches, axillary. Flowers with 9 perianth parts in 2 whorls, 1 to 2 per inflorescence; pedicel 3–7 mm long, ca. 1 mm in diameter, glabrous; in fruit 5–15 mm long, 2–3 mm in diameter, glabrous; bracts 2 to 4, all basal, 1 mm long, 1 mm wide; sepals 3, valvate, free, 2–5 mm long, 2–4 mm wide, ovate, apex acute, base truncate, green to brown-red, glabrous outside, glabrous inside, margins flat; petals basally fused, tube 4–7 mm long, inner and outer whorl not differentiated, equal; lobes 15–25 mm long, 3–4 mm wide, lorate (strap-shaped) to oblong, apex acute, light yellow to light green, margins curved inwards, glabrous outside, glabrous inside, erect over receptacle, verrucose when dried; stamens ca. 40, in 3 to 4 rows, ca. 2 mm long, broad; connective discoid, glabrous, cream; staminodes absent; carpels fused into a single structure, ca. 3 mm long, stigma bilobed, slightly capitate, glabrous. Fruit syncarpous, sessile, 30–65 mm long, 15–30 mm in diameter, ellipsoid to globose, apex rounded, glabrous, smooth, faintly ribbed to longitudinally ribbed, green turning yellow when ripe; seeds not counted, 15–20 mm long, 8–10 mm in diameter, ellipsoid; aril absent.
Known from Cameroon to the west of Republic of Congo; in Cameroon known from the South and Littoral regions.
A fairly infrequent species, growing in lowland rain forests. Altitude 0–800 m a.s.l.
None recorded.
Least Concern (LC) (
None reported.
Isolona zenkeri is characterized by its coriaceous corolla lobes with incurved margins, erect above the receptacle, and verrucose when dried.
South Region: Bipinde, 3.26°N, 10.20°E, 16 June 1918, Annet E. 359 (WAG); 12 km from Kribi N of Ebolowa road between Kribi aifield and Kienke R, 2.88°N, 9.983°E, 18 June 1969, Bos J.J. 4866 (BR,K,LM,MO,P,POZG,WAG,YA); 6 km N of km Kribi-Lolodorf, 3.08°N, 10.25°E, 12 March 1970, Bos J.J. 6522 (BR,K,P,WAG,YA); Campo-Ma’an area Bibabimvoto, 2.25°N, 10.36°E, 01 February 2000, Elad M. 1269 (WAG); Campo-Ma’an area Ebianemeyong, 2.46°N, 10.29°E, 24 May 2002, Elad M. 1545 (KRIBI,WAG); 3 km S of Kwambo and 6 km WSW of Bipindi, 3.05°N, 10.36°E, 19 January 1987, Manning D. 1453 (MO); Campo-Ma’an area Bifa, 2.65°N, 10.28°E, 12 October 2001, Tchouto Mbatchou G.P. BIFAX_2 (WAG); Campo-Ma’an area Bibabimvoto, 2.21°N, 10.01°E, 13 May 2000, Tchouto Mbatchou G.P. 2855 (KRIBI,WAG,YA); Campo-Ma’an area Bibabimvoto, 2.25°N, 10.04°E, 24 August 2000, Tchouto Mbatchou G.P. 3009 (KRIBI,WAG,YA); Bipindi, 3.08°N, 10.41°E, 1896, Zenker G.A. 1186 (B,BM,G,K); Bipindi, 3.08°N, 10.42°E, 01 January 1907, Zenker G.A. 3375 (P); Bipindi, 3.08°N, 10.41°E, 01 January 1908, Zenker G.A. 3471 (US); Bipindi, 3.08°N, 10.41°E, 01 January 1912, Zenker G.A. 4405 (G,K,MO).
Letestudoxa bella Pellegr.
Lianas, to 40 m tall, d.b.h. up to 4 cm; stilt roots or buttresses absent. Indumentum of simple hairs. Leaves: petiole 3–12 mm long, 1–6 mm in diameter, pubescent to glabrous, slightly grooved, blade inserted on top of the petiole; blade 5–28 cm long, 3–12 cm wide, elliptic to obovate to oblong, apex acuminate to emarginate, base rounded to cordate; secondary veins 11 to 20 pairs; tertiary venation percurrent to indistinct. Individuals bisexual; inflorescences ramiflorous on old leafless branches, extra axillary. Flowers with 9 perianth parts in 3 whorls, 1 per inflorescence; pedicel 3–11 mm long, 1–7 mm in diameter; in fruit 3–7 mm long, 1–3 mm in diameter; bracts 2, one basal and one upper towards the lower half of pedicel; sepals 3, valvate, completely fused forming a nearly closed cup but tearing open at anthesis, 10–20 mm long; petals free, outer petals longer than inner; outer petals 3, imbricate, 30–55 mm long, 15–55 mm wide, ovate, apex acute to rounded, base attenuate; inner petals 3, imbricate, 15–35 mm long, 10–25 mm wide, elliptic to ovate, apex acute, base attenuate to acute; stamens up to 800, in 16 to 20 rows, 2–10 mm long, broad; connective flattened, pubescent, red; staminodes absent; carpels free, 150–175, 2–2.5 mm long, stigma capitate, pubescent. [Fruits only known from L. bella] Fruit pseudosyncarpous ca. 45 mm long, ca. 50 mm in diameter, globose; individual carpels sessile, 125 to 150 carpels, apex rounded to apiculate; seed 1, 15–16 mm long, 4–8 mm in diameter, ellipsoid; aril present.
A genus of lianas with three known species, from Angola (Cabinda), Cameroon, Gabon and Republic of Congo; in Cameroon two species, none endemic.
Lestestudoxa is distinguished by the lianescent habit and pseudosyncarpous fruits (carpels fusing after pollination to form a single fruiting unit, similar to those of the genus Duguetia, but the latter being trees) and the sepals completely fused around the floral bud and tearing at anthesis. The only other lianescent Annonaceae liana in Africa with pseudosyncarpous fruits is Pseudartabotrys, a monospecific genus endemic to Gabon (
1 | Leaves generally elliptic (sometimes obovate), 5–18 cm long; upper bract not clasping base of flower buds | L. bella |
– | Leaves generally obovate (sometimes oblong), 15–28 cm long; upper bract clasping (amplexicaul) around the base of flower buds | L. lanuginosa |
= Letestudoxa grandifolia Pellegr., Bull. Mus. Natl. Hist. Nat. 26: 656, 1920. Type. Gabon. Nyanga, Ilou Micongo, Le Testu G.M.P.C. 1442, 4 Nov 1908: holotype: P[P00364780]; isotype: BM[BM000546387].
= Pachypodanthium simiarum Exell & Mendonça, J. Bot. 74 (Suppl.): 14, 1936. Type. Angola. Maiombe, Belize, Gossweiler J. 6971, 16 Fev 1917: holotype: BM[BM000067635]; isotypes: COI[COI00004878]; LISC[LISC000102, LISC000101, LISC000098, LISC000099, LISC000100].
Gabon. Nyanga; Midounga, near Tchibanga, Le Testu G.M.P.C. 1637, 24 Oct 1910: holotype: P[P00364779]; isotype: BM[BM000546388].
Liana, 16–40 m tall, d.b.h. up to 4 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches densely pubescent. Leaves: petiole 3–10 mm long, 1–4 mm in diameter, pubescent to glabrous, slightly grooved, blade inserted on top of the petiole; blade 5–18 cm long, 3–8 cm wide, elliptic to obovate, apex acuminate to emarginate, acumen 0.5–1 cm long, base rounded to cordate, coriaceous, below densely pubescent when young, densely pubescent when old, above glabrous when young and old, concolorous; midrib sunken or flat, above glabrous when young and old, below pubescent when young and old; secondary veins 12 to 20 pairs, pubescent below; tertiary venation percurrent to indistinct. Individuals bisexual; inflorescences ramiflorous on old leafless branches, extra axillary. Flowers with 9 perianth parts in 3 whorls, 1 per inflorescence; pedicel 3–6 mm long, 1–3 mm in diameter, pubescent; in fruit 3–7 mm long, 1–3 mm in diameter, pubescent; bracts 2, one basal and one upper towards the lower half of pedicel, basal bract 5–7 mm long, 3–5 mm wide; upper bract 6–12 mm long, 4–7 mm wide, not clasping the flower bud; sepals 3, valvate, completely fused, tearing at anthesis, 10–20 mm long, base truncate, brown, densely pubescent outside, densely pubescent inside, margins flat; petals free, outer petals longer than inner; outer petals 3, 30–55 mm long, 15–55 mm wide, ovate, apex acute, base attenuate, yellow to orange with pinkish margins, margins crisped, densely pubescent outside, pubescent inside; inner petals 3, imbricate, 15–30 mm long, 10–25 mm wide, elliptic to ovate, apex acute, base attenuate to acute, yellow to orange with red marginal zone, margins wavy, densely pubescent outside, pubescent inside; stamens numerous, 2–3 mm long, broad; connective flattened, pubescent, red; staminodes absent; carpels free, around 175, ovary ca. 2 mm long, stigma capitate, pubescent. Fruit pseudosyncarpous, ca. 45 mm long, ca. 50 mm in diameter in total, globose; individual carpels sessile, 125 to 150 carpels, obovoid to obtrulloid, apex rounded to apiculate, sparsely pubescent, 6 to 7 ribbed, green turning red when ripe; seed 1 per monocarp, 15–16 mm long, 4–8 mm in diameter, ellipsoid; aril present, light brown.
Letestudoxa lanuginosa A flowering branch, with one open flower and one flower bud, note sepals completely fused B detail of lower side of leaf blade showing dense pubescence C flower, bottom view D stamens, side and front views E carpels, note long elongated stigma F longitudinal section of receptacle G flower diagram. Letestudoxa bella H detail of lower side of leaf blade showing pubescence I flower bud, note sepals completely fused A–G from Le Testu 9320; 8–9 from Le Testu 8362. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
Known from Angola, Cameroon to Gabon and in the Republic of Congo; in Cameroon known from the South region.
Growing in lowland primary and secondary rain forests. Altitude 50–750 m a.s.l.
None recorded.
Not evaluated.
None reported.
Letestudoxa bella is characterized by its mostly elliptic leaves (sometimes obovate too) which are generally smaller (5–18 cm) than those of L. lanuginosa (15–28 cm). In addition, in fresh material, L. lanuginosa has a bullate upper surface (versus a more leathery smooth upper surface in L. bella). In flower, L. lanuginosa is characterized by the upper bract clasping the flower bud, which is not the case in L. bella.
South Region: 26 km E of confluent Ntem River and Akom River near Ebolowa, 2.29°N, 11.86°E, 05 March 1970, Letouzey R. 10097 (BR,P,P,YA); Tom (Nyabessan), 2.43°N, 10.52°E, 04 March 1963, Raynal J. 10195 (P,YA).
Gabon. Woleu-Ntem; Ncout, Le Testu G.M.P.C. 9320, 13 Oct 1938: lectotype, sheet here designated: P[P00364781]; isotypes: P[P00364782, P00364783].
Liana, 20 m tall, d.b.h. 2–3 cm in diameter; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches densely pubescent to tomentose, young foliate branches densely pubescent to tomentose. Leaves: petiole 6–12 mm long, 3–6 mm in diameter, densely pubescent, slightly grooved, blade inserted on top of the petiole; blade 15–28 cm long, 7–12 cm wide, obovate to sometimes oblong, apex rounded or emarginate or obcordate or mucronate, acumen 0.1–1.9 cm long, base cordate, coriaceous, below densely pubescent when young, densely pubescent when old, above glabrous when young and old, concolorous; midrib sunken or flat, above densely pubescent when young and old, below densely pubescent when young, densely pubescent when old; secondary veins 11 to 18 pairs, densely pubescent below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old leafless branches, extra axillary. Flowers with 9 perianth parts in 3 whorls, 1 per inflorescence; pedicel 3–11 mm long, 4–7 mm in diameter, densely pubescent; in fruit unknown; bracts 2, one basal and one upper towards the upper half of pedicel, basal bract 8–13 mm long, 10–15 mm wide; upper bract 8–18 mm long, 6–10 mm wide, clasping the flower bud; sepals 3, valvate, completely fused, tearing at anthesis, 10–20 mm long, base truncate, brown, pubescent outside, pubescent inside, margins flat; petals free, outer petals longer than inner; outer petals 3, imbricate, 30–50 mm long, 30–40 mm wide, ovate, apex rounded to acute, base attenuate, orange to red, margins crisped, densely pubescent inside, pubescent outside; inner petals 3, imbricate, 25–35 mm long, 15–25 mm wide, elliptic to ovate, apex acute, base attenuate, claw ca. 5 mm long, orange to red, margins wavy, densely pubescent inside, pubescent outside; stamens 750 to 800, in 16 to 20 rows, 7–10 mm long, broad; connective flattened, pubescent; staminodes absent; carpels free, 150, ovary 2–2.5 mm long, stigma capitate, pubescent. Fruits unknown.
Letestudoxa bella A leaf, upper side B fruit, a pseudosyncarpous fruit C fruit, top view, note the large sepal remains. Letestudoxa lanuginosa D habit, liana climbing on tree trunk E leaf, upper side F leaf, lower side G detail of young flower bud, not completely enclosing sepals A–C Couvreur 600, Gabon D–G Couvreur 1148, Ma’an, Cameroon. Photos Thomas L.P. Couvreur.
A Letestudoxa lanuginosa B Meiocarpidium oliverianum C Mischogyne gabonensis D Monanthotaxis bicornis E Monanthotaxis bokoli F Monanthotaxis capea G Monanthotaxis cauliflora H Monanthotaxis congoensis I Monanthotaxis couvreurii. White borders represent region limits in Cameroon; green patches represent protected areas (see methods and Suppl. material
Known from southern Cameroon to Gabon; in Cameroon known from the South region.
A frequent species when present; growing on non-inundated soils, in primary or secondary forests, sometimes near rivers and swampy areas. Altitude 100–600 m a.s.l.
None recorded.
Not evaluated.
None reported.
See under L. bella for differences between the two species. This species can be confused with Uvaria bipindensis (also a liana) by the size of its leaves (to 27 cm long) and cordate shape of the leaf base. They differ however but the presence of stellate hairs in Uvaria bipindensis.
South Region: Ma’an 3 km on small road towards Ntem river, 2.34°N, 10.63°E, 25 February 2018, Couvreur T.L.P. 1148 (MPU,WAG,YA); 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. 697 (WAG,YA); 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. 698 (WAG,YA); Campo-Ma’an area Boucle du Ntem, 2.34°N, 10.58°E, 16 February 2001, Tchouto Mbatchou G.P. 3166 (KRIBI,WAG).
Meiocarpidium oliverianum (Baillon) D.M.Johnson & N.A.Murray
Same as species.
A monotypic genus from the Central Atlantic African region (Lower Guinea).
Meiocarpidium is characterized by the presence of peltate scale-like hairs, a character unique among Central African Annonaceae.
≡ Unona oliveriana Baillon, Adansonia 8: 307, 1868; Unona lepidota Oliv. Fl. Trop. Afr. 1: 36, 1868; Meiocarpidium lepidotum (Oliv.) Engl. & Diels, Notizbl. Königl. Bot. Gart. Berlin 3: 55, 1900.
= Uvaria zenkeri Engl., Notizbl. Königl. Bot. Gart. Berlin 2: 293, 1899. Type. Cameroon. South Region, Bipindi, Zenker G.A. 1864, 1896: lectotype, here designated: K[K000198806]; isolectotypes: K[K000198807]; HBG[HBG502509, HBG502510]; WU[WU0025880, WU0025879].
Equatorial Guinea. Rio Muni; Muni River, Mann G. 1774, Aug 1862: lectotype, sheet here designated: K[K000795931]; isotypes: K[K000795932]; P[P00362615].
Tree, 8–20 m tall, d.b.h. 20 cm; stilt roots or buttresses absent. Indumentum of peltate scale-like hairs; old leafless branches glabrous, young foliate branches pubescent. Leaves: petiole 9–11 mm long, 2 mm in diameter, pubescent, grooved, blade inserted on top of the petiole; blade 15–20 cm long, 5–7 cm wide, oblong to obovate, apex acuminate to acute, acumen 1–2 cm long, base decurrent to acute, coriaceous, below densely pubescent when young giving a silvery color, pubescent when old, above glabrous when young and old, discolorous, whitish below; midrib impressed above, above glabrous when young and old, below pubescent when young and old; secondary veins 13 to 17 pairs, glabrous below; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, leaf opposed. Flowers with 9 perianth parts in 3 whorls, 1 per inflorescence; pedicel 20–30 mm long, 1–2 mm in diameter, densely pubescent; in fruit 20–30 mm long, 2–3 mm in diameter, pubescent; bracts 2, one basal and one upper towards the upper half of pedicel, basal bract 8–9 mm long, 1 mm wide; upper bract 2–3 mm long, 1 mm wide; sepals 3, valvate, free, 2–3 mm long, 1–2 mm wide, triangular, apex acute, base truncate, green, pubescent outside, densely pubescent inside, margins flat; petals free, sub equal; outer petals 3, valvate, 15–25 mm long, 10–15 mm wide, ovate, apex acute, base truncate, cream, margins flat, densely pubescent outside, pubescent inside; inner petals 3, valvate, 15–25 mm long, 5–10 mm wide, ovate, apex acute, base narrowed into a claw 3–5 mm long, cream, margins flat, pubescent outside, glabrous inside; stamens 90 to 100, in 5 to 6 rows, 3–4 mm long, broad; connective discoid, glabrous, cream; staminodes absent; carpels free, 3 to 5, ovary 7–8 mm long, stigma capitate, pubescent. Monocarps sessile, 3 to 4, 40–60 mm long, 20–30 mm in diameter, cylindrical, apex rounded to apiculate, pubescent, smooth, silver-green when ripe; seeds 14 to 16 per monocarp, 10–20 mm long, 5–10 mm in diameter, flattened ellipsoid to oblong; aril absent.
Meiocarpidium oliverianum A flowering branch B detail of lower leaf surface showing lepidote (scale like) hairs C outer petal, outer side D inner petal, outer side E detail of receptacle, side view, 3 petals removed F detail of receptacle, semi top view, all petals removed G stamens, side and front views H carpel, side view, and detail of ovules I fruit J longitudinal section of a monocarp A–H from Zenker 3027 I–J from Letouzey 5473. Drawings by Hélène Lamourdedieu, Publications Scientifiques du Muséum national d’Histoire naturelle, Paris; modified from
Known from Cameroon, Equatorial Guinea Gabon and Republic of Congo, one collection for the Democratic Republic of Congo; in Cameroon known from East, South, Littoral and Central regions.
A frequent species when present; growing on swampy, periodically inundated or well-drained soils, in primary or secondary forests, sometimes near rivers and swampy areas. Altitude 50–500 m a.s.l.
mambéléngé, mandongé (dial. Baka bibaya).
Not evaluated (probably LC).
medicine : bark used to allay fever; construction: wood used to make spoons.
Meiocarpidium oliverianum is easily identifiable by the presence of peltate scale-like hairs on the lower side of the leaf blade, calyx and fruits, giving these parts a silvery aspect and color.
The species was previsously known under the name Meiocarpidium lepidotum (Oliv.) Engl. & Diels, but the name Meiocarpidium oliverianum of
Central Region: Bank Nyong River near the new bridge ca 65 km SSW of Eséka, 3.65°N, 10.78°E, 17 June 1964, de Wilde W.J.J.O 2735 (B,BR,K,MO,P,WAG,YA); Bafia region, 4.75°N, 11.23°E, 18 December 1957, de Wit H.C.D 7948 (WAG). East Region: Berbérati, 4.58°N, 13.68°E, 01 March 1963, BPFV 2664 (P); 47 km NW of Bertoua near road from Mbang to Abaka, 4.58°N, 13.68°E, 22 May 1961, Breteler F.J. 1398 (A,BR,K,M,P,UC,WAG,YA); Bertoua 9 km along road to Deng Deng, 4.58°N, 13.68°E, 31 August 1961, Breteler F.J. 1817 (A,BR,K,M,P,WAG,YA); 70 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, 04 March 2019, Couvreur T.L.P. 1193 (MPU,WAG,YA); Yokadouma 3.52°N, 15.05°E, 11 September 1939, Jacques-Félix H. 4904 (P); Deng Deng, 5.20°N, 13.13°E, 27 July 2014, Kamdem N. 169 (YA); sur la route de Esseleké, 4.58°N, 13.68°E, 18 August 1955, Nana P. 213 (P,YA); km 38 de la route de Deng-Deng, 4.58°N, 13.68°E, 17 September 1955, Nana P. 257 (P,YA); Environ de Landjwe 25 km SW Yokadouma 3.47°N, 14.93°E, 16 June 1984, Satabié B. 767 (P,YA). Littoral Region: Mapubi 30 km before Edea on Yaoundé-Edea road On forestry road 5 km direction to Sanaga river, 3.84°N, 10.38°E, 28 February 2018, Couvreur T.L.P. 1182 (MPU,WAG,YA). South Region: 6 km N of km Kribi-Lolodorf, 3.08°N, 10.25°E, 12 March 1970, Bos J.J. 6523 (WAG); Near village Oveng 27 km from Sangmélima along road to Yaoundé, 3.09°N, 11.90°E, 20 March 1962, Breteler F.J. 2646 (BR,K,L,P,U,WAG,YA); Campo Ma an National Park 26 km after Ntem river, 3.07°N, 14.64°E, 08 March 2019, Couvreur T.L.P. 376 (WAG,YA); on road from Lolodorf to Mekalat just after 1sty village Along the Malange river, 3.19°N, 10.71°E, 20 February 2012, Couvreur T.L.P. 407 (WAG,YA); 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. 709 (WAG,YA); Station de cacaoyer de N’koemvone 14 km On the road from Ebolowa to Ambam, 2.81°N, 11.13°E, 27 September 1974, de Wilde J.J.F.E 7576 (BR,K,MO,P,WAG,YA); Ebolowa SW of Mbalmayo, 2.96°N, 11.28°E, 27 February 1964, de Wilde W.J.J.O 1942 (BR,K,MO,P,WAG,YA); Près Akak 10 km W Sangmeli 2.96°N, 11.88°E, 11 March 1970, Letouzey R. 10153 (P,YA); Ebom, 3.05°N, 10.71°E, 29 August 1996, Ndoum D. 88 (KRIBI,WAG); Campo-Ma’an area 2.39°N, 10.07°E, 12 June 2001, van Andel T.R. 3619 (KRIBI,WAG,YA); Campo-Ma’an area Akom II, 2.8°N, 10.53°E, 18 August 2001, van Andel T.R. 3931 (U,WAG,YA); Bipindi, 3.08°N, 10.42°E, 1898, Zenker G.A. 1864 (P); Bipindi, 3.08°N, 10.42°E, 01 January 1902, Zenker G.A. 2505 (P); Bipindi, 3.08°N, 10.42°E, 01 January 1903, Zenker G.A. 2521 (L,P,WAG); Bipindi, 3.08°N, 10.42°E, 01 January 1904, Zenker G.A. 2947 (L,P,WAG); Bipindi, 3.08°N, 10.42°E, 01 January 1904, Zenker G.A. 3221 (L,P,WAG); Bipindi, 3.08°N, 10.42°E, 01 January 1911, Zenker G.A. 4209 (P,U); Bipindi, 3.08°N, 10.42°E, 01 January 1918, Zenker G.A. 60 (P).
Mischogyne michelioides Exell.
Genus description for Cameroon same as species.
A genus of trees or shrubs with five known species (
The genus Mischogyne is easily identified when fertile by the presence of a torus, an extended receptacle, and several elongated cylindrical to ovoid carpels, which are divergent from each other.
≡ Mischogyne elliotiana var. gabonensis Pellegr. ex Le Thomas, Flore du Gabon 16: 291, 1969.
Gabon. Nyanga; Mayumba, Le Testu G.M.P.C. 1768, 26 Aug 1914: lectotype, designated by
Tree, 3–25 m tall, d.b.h. up to 25 cm; stilt roots or buttresses absent. Indumentum of simple hairs; old leafless branches glabrous, young foliate branches shortly pubescent. Leaves: petiole 5–10 mm long, ca. 2 mm in diameter, sparsely pubescent to glabrous, cylindrical, blade inserted on the side of the petiole; blade 8–16 cm long, 4–7 cm wide, obovate, apex acuminate, acumen 1–3 cm long, base cuneate, papyraceous to subcoriaceous, 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 6 to 9 pairs, glabrous above; tertiary venation reticulate. Individuals bisexual; inflorescences ramiflorous on old or young foliate branches, extra-axillary. Flowers with 9 perianth parts in 3 whorls, 1 per inflorescence; pedicel 5–12 mm long, 0.5–1 mm in diameter, pubescent; in fruit 7–15 mm long, 2–3 mm in diameter, glabrous; bracts few, reduced to a tuff of hairs, all basal; sepals, valvate, completely fused in bud, tearing at anthesis into 2 or 3 parts, 7–13 mm long, 3–5 mm wide, triangular, apex acute, base truncate, green, tomentose on both sides, margins flat; petals free, equal; outer petals 3, valvate, 14–16 mm long, 3–5 mm wide, narrowly elliptic, apex acute, base truncate, bright white, margins flat, densely pubescent outside, tomentose inside; inner petals 3, valvate, 14–16 mm long, 3–5 mm wide, narrowly elliptic, apex acute, base truncate, bright white, margins flat, densely pubescent outside, tomentose inside; stamens 50 to 60, in 3 to 4 rows, 1–3 mm long, elongated; connective tongue shaped, pubescent, cream; staminodes absent; carpels free, 3–7, ovary 5–6 mm long, stigma bilobed, densely pubescent. Monocarps sessile, 1 to 3, 60–65 mm long, 28–32 mm in diameter, cylindrical, apex rounded, glabrous, smooth, 1-ribbed, yellow when ripe; seeds 6 per monocarp, ca. 20 mm long, ca. 15 mm in diameter, ellipsoid, covered with an indumentum of fine white hairs; aril absent.
Meiocarpidium oliverianum A detail of lower side of leaf showing lepidote (scale-like) hairs; white bow zoomed in) B flowering branch C fruiting branch. Mischogyne gabonensis D base of trunk E base of leaf blade, upper side F base of leaf bade, lower side G flower bud, note sepals tearing H detail of flower, top view I detail of flower, all petals and sepals fallen, note elongated carpels J longitudinal section of monocarp, note hairy seeds A–C Sosef 2317 D–I Couvreur 1033, Mt Cameroon, Cameroon J Kenfack 1545; Mt Cameroon, Cameroon. Photos A–H Thomas L.P. Couvreur I David Kenfack, Tropicos.org, Missouri Botanical Garden.
Cameroon, Gabon and Bioko island (Equatorial Guinea); in Cameroon known from the South and South-West regions.
A rare species in Cameroon (or at least uncommonly collected); in lowland primary or old secondary rain forests. Altitude 0–500 m a.s.l.
None recorded.
Endangered (EN) (
None reported.
Mischogyne gabonensis is morphologically similar to the West African endemic M. elliotiana (of which it was formerly regarded as a variety) in the shape and size of its flowers and monocarps (
South Region: Environs de Nko’olong 30 km E Campo, 2.40°N, 10.03°E, 21 November 1992, Satabié B. 978 (YA). South-West Region: Mount Cameroon National Park Bakinguili trail above Bakinguili village, 4.09°N, 9.054°E, 02 April 2016, Couvreur T.L.P. 1033 (WAG,YA); Fako Njonji Njonji along path to the lake Low canal forest, 4.15°N, 9.066°E, 26 September 2001, Kenfack D. 1545 (MO).
= Clathrospermum Planch. ex Benth., Gen. Pl. 1(1): 29, 1862.
= Enneastemon Exell J. Bot. 70 (Suppl. 1): 209, 1932.
= Exellia Boutique, Bull. Jard. Bot. État Bruxelles 21: 117, 1951.
= Atopostema Boutique, Bull. Jard. Bot. État Bruxelles 21: 121, 1951.
= Gilbertiella Boutique, Bull. Jard. Bot. État Bruxelles 21: 124, 1951.
Scrambling shrubs or lianas, up to 30 m tall, d.b.h. up to 11 cm; stilt roots or buttresses absent. Indumentum of simple hairs. Leaves: petiole 1–11 mm long, 1–3 mm in diameter; blade 3–35 cm long, 1.4–12.5 cm wide, linear to elliptic to ovate to obovate to oblong, apex acuminate to acute, base decurrent to subcordate, discolorous, whitish below; midrib sunken or flat; secondary veins 6 to 23 pairs; tertiary venation percurrent. Individuals unisexual or bisexual; inflorescences cauliflorous or ramiflorous on old or young foliate branches, axillary, leaf opposed or extra axillary. Flowers with (6-)9 perianth parts in 2 or 3 whorls, 1 to more than 50 per inflorescence with a peduncle from 1 to 70 mm long; pedicel 1–60 mm long; in fruit 2–55 mm long; bracts 2 to 3, one or two basal and one inserted on the pedicel at varying levels; sepals 3, valvate, free, 0.5–10 mm long, ovate to elliptic to triangular, apex acute or obtuse, base truncate; petals free; outer petals 3 or 6 (when petals in one whorl), valvate, 1–50 mm long, 1–25 mm wide, ovate to elliptic, apex acute to rounded, base truncate; inner petals (0-)3, valvate, 0.5–25 mm long, 0.3–10 mm wide, ovate to elliptic to rhombic, apex acute to rounded, base truncate; stamens 1 to more than 125, in 1 to 6 rows, 1–2 mm long, linear or cuneiform to clavate; anthers sometimes connate apically covering connectives, connective discoid to truncate to absent, glabrous or pubescent; staminodes 6 to 12(13) or absent; carpels free, 3 to 150, 1–4 mm long, stigma bilobed or cylindrical or elongate and flattened at top, pubescent or glabrous. Monocarps sessile or stipitate, stipes 1–25 mm long; monocarps 2 to 25, 6–60 mm long, 5–25 mm in diameter, moniliform, cylindrical to ellipsoid, apex rounded to apiculate, smooth or verrucose or weakly torulose; seeds 6–25 mm long, 4–11 mm in diameter, ellipsoid; aril absent.
Monanthotaxis congoensis Baill.
A genus of scrambling shrubs or lianas with 79 known continental African species, from West, Central and East Africa, plus around 23 endemic species from Madagascar. In Cameroon 26 species are known, six endemic.
The genus Monanthotaxis is easily identified when sterile by its lianescent or scrambling habit, discolorous leaves with very glaucous lower surface, and percurrent (parallel) tertiary venation. Some species are unisexual and have cauliflorous inflorescences; some are confirmed as monoecious, but for others, even though male and female flowers are known, it remains to be confirmed if they occur on the same or different individuals. The stamen number is highly variable within the genus ranging from 1 or 2 to over 100 (
In the checklist to the plants of Mt Cameroon (
African species (excluding Madagascar) are revised in
The species of Monanthotaxis are variable in their vegetative characters (
1 | Young foliate branches covered with erect hairs longer than 0.5 mm, normally around 1 mm long | 2 |
– | Young foliate branches almost glabrous, or covered with appressed to ascending hairs, or with erect hairs shorter than 0.5 mm. (if in doubt choose this option: M. diclina has dense yellow-brown indumentum ascending to erect hairs and M. letouzeyi has ascending to erect hairs just shorter than 0.5 mm) | 7 |
2 | Inflorescences cauliflorous or axillary | 3 |
– | Inflorescences extra–axillary | 4 |
3 | Petiole 5–7 mm long; flowers unisexual; female inflorescences cauliflorous; petals < 7 mm | M. pynaertii |
– | Petioles 2.5–6 mm long; flowers bisexual, all axillary; petals > 8 mm | M. filamentosa |
4 | Leaves smaller than 17 cm; stamens 22 to 34; carpels glabrous; monocarps glabrous or at most with few scattered hairs on the stipe | 5 |
– | At least several leaves larger than 18 cm; stamens 90 to 120; carpels pubescent; monocarps pubescent | 6 |
5 | Leaf apex normally obtuse; outer petals 15–19 mm long; carpels 27 to 38; seeds cylindrical, 14–21 mm long; stipes 7–10(–25) mm long | M. bokoli |
– | Leaf apex acute; outer petals 5.8–6.7 mm long; carpels 12 to 24; seeds ellipsoid, 7–8 mm long; stipes 3–4 (–6.5) mm long | M. ferruginea |
6 | Hairs on young foliate branches around 1.5 mm long; leaves normally oblong to slightly obovate with broad leaf base and acute apex; outer petals 21–50 mm long; thecae large, covering more than half of the stamen length; carpels 22 to 24 | M. hirsuta |
– | Hairs on young foliate branches around 1 mm long; leaves normally oblong–oblanceolate with narrow leaf base and acuminate apex; outer petals 12–22 mm long; thecae small, covering less than half of the stamen length; carpels 40 to 60 | M. enghiana |
7 | Leaves linear to narrowly elliptic, at least 5 times longer than wide, widest in middle or lower half of the leaf, secondary veins almost perpendicular to the primary vein (>75) | M. sterilis |
– | Leaves oblong, elliptic, obovate or oblanceolate, if > 5 times longer than wide then widest in upper half of the leaf and secondary veins forming acute angle with the primary vein (<60) | 8 |
8 | Inflorescences leaf-opposed or extra–axillary, but not consistently a few mm above the leaf axils | 9 |
– | Inflorescences cauliflorous, axillary or supra–axillary and then consistently 1–8 mm above the leaf axils | 15 |
9 | Upper bract leaf-like, 7–15 mm long; stamens 15 | M. vulcanica |
– | Upper bract not leaf-like, shorter than 5 mm or absent; stamens 9 or more than 5 mm or absent; stamens 9 or more than 23 | 10 |
10 | Young foliate branches with sparse hairs not covering the entire surface; old leafless branches light brown or reddish brown | 11 |
– | Young foliate branches with dense hairs completely covering the surface; old branches dark brown to blackish (or grey in M. dielsiana) | 12 |
11 | Branches light brown; stamens 80 to 125; carpels and monocarps hairy | M. gracilis |
– | Branches reddish brown; stamens 23 to 24; carpels and monocarps glabrous | M. laurentii |
12 | Inflorescence a 1 to 8 flowered glomerule–like rhipidium; pedicels 1.2–2(–5) mm long; stamens 9; staminodes 6 | M. elegans |
– | Inflorescence a solitary flower or a 2 to 4 flowered fascicle; pedicels 5–33 mm long; stamens more than 35; staminodes absent | 13 |
13 | Petiole 0.8–1.5 mm diam.; pedicels 0.4–0.5 mm diam.; stamens 36 to 48; carpels 11 to 14 | M. montana |
– | Petiole 1.6–2.8 mm diam.; pedicels 1.4–2.1 mm diam.; stamens >60; carpels 41 to 50 | 14 |
14 | Young foliate branches orange brown, densely covered with appressed, orange brown hairs; stamens c. 65 | M. dielsiana |
– | Young foliate branches brown, densely covered with appressed, pale brown hairs; stamens >100 | M. glaucifolia |
15 | Inflorescences cauliflorous, at least most of them (in some species with unisexual flowers the male inflorescences are axillary on foliate branches) | 16 |
– | Inflorescences axillary or supra–axillary, rarely some inflorescences cauliflorous; flowers always bisexual | 21 |
16 | Young foliate branches densely covered with erect or curly hairs 0.4–0.6 mm long | 17 |
– | Young foliate branches covered with appressed to ascending hairs 0.1–0.2 mm long | 18 |
17 | Young foliate branches with reddish brown hairs; male flowers with 31 to 40 stamens in four whorls; 0 staminodes; carpels c. 120; seeds globose | M. letouzeyi |
– | Young foliate branches with yellow-brown hairs; male flowers with 6 stamens in one whorl and 12 small staminodes in an external whorl; carpels 80 to 100; seeds ellipsoid | M. diclina |
18 | Young foliate branches with very short yellow-brown hairs; petiole long and less than 1 mm in diam.; carpels 65 to 85 | M. submontana |
– | Young foliate branches with very short reddish brown or dark brown hairs; petiole thick, more than 1 mm in diam.; carpels 20 to 60(–76) | 19 |
19 | Flowers unisexual; female inflorescences cauliflorous in many-flowered panicle–like rhipidia, male inflorescences axillary; stamens 6; staminodes 12; carpels 48 to 60; leaf base rounded to cuneate; secondary veins 11 to 16 | M. cauliflora |
– | Flowers bisexual; inflorescences a solitary flower, a few flowered fascicle or a short glomerule; stamens 9; staminodes 9; carpels 20 to 34; leaf base rounded to subcordate; secondary veins 7 to 14 | 20 |
20 | Leaf base rounded or subcordate; flower buds ovoid, with outer petals and base of the 3 inner petals visible; monocarps with 1 seed; stipes 1–3.5 mm long | M. klainei |
– | Leaf base narrowly subcordate; flower buds globose, with inner petals entirely covered by outer petals; monocarps with 1 to 3 seeds; stipes 4.5–12 mm long | M. whytei |
21 | Leaf underside with dense appressed hairs more than 1 mm long, giving a silky appearance; inflorescences raceme-like or paniculate rhipidia; all 6 petals in one whorl | 22 |
– | Leaf underside with different pubescence, not silky, the hairs either shorter or not appressed; inflorescences 1-flowered or in few-flowered fascicle-like rhipidia; petals in 2 whorls of 3 | 23 |
22 | Leaf underside with yellowish hairs; inflorescence a many–flowered panicle-like rhipidium; flower buds depressed-globose | M. paniculata |
– | Leaf underside with whitish hairs; inflorescence a 4–10 flowered raceme-like rhipidium; flower buds deltoid–ovoid | M. congoensis |
23 | Young foliate branches covered with white-yellowish hairs; stamens 6; filaments >1 mm long, much longer than half the total length of the stamen | M. hexamera |
– | Young foliate branches covered with yellowish-brown, reddish brown or dark brown hairs; stamens 8 to 35; filaments <1 mm long, occupying about half or less than the total length of the stamen | 24 |
24 | Young foliate branches covered with appressed hairs | 25 |
– | Young foliate branches covered with ascending to erect hairs | 27 |
25 | Young foliate branches covered with yellowish–brown hairs, 0.2–0.4 mm long; stamens 15 to 17; thecae converging on top of the stamen, leaving only a small part of connective visible from above, filament wider than connective | M. bicornis |
– | Young foliate branches covered with reddish–brown hairs 0.1–0.2 mm long; stamens 8 to 12; thecae on sides of the stamen (not on top) with connective clearly visible from above, as wide as or wider than filament | 26 |
26 | Leaf base cuneate or sometimes rounded, with slightly thickened black margin; secondary veins forming an acute angle with primary vein, straight at their base; petals 6 in 2 whorls; 1 ovule; monocarps with 1 seed | M. vogelii |
– | Leaf base rounded, sometimes with thick globose glands at the margin; secondary veins curving from base; petals 3 to 4; 3 to 4 ovules; monocarps with 1 to 4 seeds | M. tripetala |
27 | Young foliate branches with dense erect hairs, 0.3–0.4 mm long; stamens 35 in three to four whorls, sparsely hairy; carpels 16 | M. zenkeri |
– | Young foliate branches with ascending or erect hairs 0.1–0.3 mm long; stamens 8 to 24 in one or two whorls, glabrous, papillate or with a few hairs on the inner side of the connective; carpels 6 to 21 | 28 |
28 | Flower bud just before anthesis with only 3 petals visible, inner petals completely covered by outer petals, normally clearly different in shape and smaller; stamens 15 to 24, glabrous, thecae converging on top of the stamen, connective much narrower than width of filament | M. pellegrinii |
– | Flower bud just before anthesis with inner petals at least partly visible, only slightly differing from outer petals in shape and size; stamens 8 to 15, papillate or with a few hairs on the inner side of the connective, thecae not converging on top of the stamen, connective clearly visible from above and as wide or wider than filament | 29 |
29 | Petiole 3–5 mm long, 0.8–0.9 mm thick inflorescences cauliflorous, ramiflorous or axillary; flower buds ovoid; stamens 13 to 15, basally connate | M. couvreurii |
– | Petiole 4.5–8 mm long, 1.2–2.6 mm thick; inflorescences axillary or slightly supra-axillary; flower buds rounded or slightly ovoid; stamens 9, free | 30 |
30 | Tertiary venation on upper leaf side strongly raised; inflorescences normally 3 to 16 flowered, with a sympodial rachis 3–17 mm long; ovules 2 to 3; monocarps smooth | M. foliosa |
– | Tertiary venation on upper leaf side flat; inflorescences 1 to 4 flowered, sympodial rachis absent; ovules 4 to 6; monocarps strongly tuberculate-rugulose | M. capea |
≡ Popowia bicornis Boutique, Bull. Jard. Bot. État Bruxelles 21: 115.
Democratic Republic of the Congo. Orientale; Yamboa, Louis J.L.P. 8957, 21 Apr 1938: lectotype, chosen by
Liana, 30 m tall, d.b.h. unknown. Indumentum of simple hairs; old leafless branches glabrescent with lenticels, young foliate branches yellow-brown, with dense appressed yellow-brown hairs 0.2–0.4 mm long. Leaves: petiole 3–5 mm long, 0.6–1 mm in diameter, densely pubescent, cylindrical, blade inserted on the side of the petiole; blade 6–12 cm long, 1.9–3.5 cm wide, oblong to elliptic, apex acuminate, acumen 0.5–1.5 cm long, base cuneate, papyraceous, below densely pubescent when young, densely pubescent to sparsely pubescent when old, above glabrous when young and old, discolorous, whitish below; midrib impressed, above sparsely pubescent to glabrous when young and old, below densely pubescent when young and old; secondary veins 11 to 14 pairs, glabrous above; tertiary venation percurrent. Individuals bisexual; inflorescences ramiflorous on young foliate branches, axillary. Flowers with 9 perianth parts in 3 whorls, 1 to 3 per inflorescence; pedicel 15–60 mm long, 0.3–0.6 mm in diameter, sparsely pubescent to densely pubescent; basal bract ca. 1 mm long, ca. 0.5 mm wide; upper bract 0.5–1 mm long, 0.5–1 mm wide; sepals 3, valvate, free, ca. 1 mm long, 2–3 mm wide, ovate, apex acute, base truncate, densely pubescent outside, densely pubescent inside, margins flat; petals free, outer petals longer than inner; inner petals entirely covered i