Research Article |
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Corresponding author: Carmen Puglisi ( carmenpuglisi@gmail.com ) Academic editor: Brecht Verstraete
© 2025 Edgard Mestre Serra, Carmen Puglisi, Alexander G. Linan, Nattanon Meeprom, Hasina N. Rakouth, Heidi H. Schmidt, Porter P. Lowry II.
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:
Mestre Serra E, Puglisi C, Linan AG, Meeprom N, Rakouth HN, Schmidt HH, Lowry II PP (2025) A taxonomic revision of the continental African material previously included in Diospyros ferrea (Ebenaceae). Plant Ecology and Evolution 158(1): 82-99. https://doi.org/10.5091/plecevo.140561
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Background and aims – As currently circumscribed, Diospyros ferrea represents one of the most geographically widespread species in the genus and had previously been described as an ochlospecies because of its particularly challenging morphological complexity. Recent studies have shown that it likely contains multiple species that differ from populations in SW India and Sri Lanka corresponding to the type, which was collected on the Malabar coast in India. Here we present the second in a series of papers that aim to revise the taxonomy of D. ferrea, treating material from Africa.
Material and methods – This study was based on measurements and observations of herbarium specimens from K, MO, and P as well as scanned images from BM, BR, C, COI, EA, FHO, LISC, US, and WAG. Species were delimited based on a combination of fruit and leaf characters, in conjunction with eco-geographic distribution. Specimen records were georeferenced, mapped, and used to conduct risk of extinction assessments based on the IUCN Red List criteria.
Key results – Five well-delimited and geographically coherent species are recognized among the African material previously ascribed to Diospyros ferrea, including three described as new (D. angolensis, D. moutsambotei, and D. suaheliensis) and two for which new combinations are made (D. guineensis and D. smeathmannii). We provide a diagnostic key along with detailed morphological descriptions and photos illustrating the principal characters that distinguish the five species. Preliminary risk of extinction assessments indicate that only one of them is Least Concern, whereas three are Vulnerable, and one is Endangered.
Conclusion – This revision demonstrates that taxonomically challenging ochlospecies can be resolved by careful analysis of morphological characters and geographic distribution, provided adequate material is available. Studies to clarify the nature, evolutionary history, and taxonomic status of other ochlospecies recognized by Frank White (viz. D. natalensis and D. mespiliformis) should be undertaken, but may also require additional phylogenetic and/or phylogeographic data.
Africa, Diospyros, Ebenaceae, new species, ochlospecies, species complex, taxonomy
The ebony genus, Diospyros L., comprises approximately 945 almost entirely tropical species, including 795 that are currently accepted (
Recent work by
In this article, we present the results of a taxonomic revision of material from continental Africa assigned to Diospyros ferrea, particularly by
We studied collections from key herbaria with holdings from continental Africa, in particular K, MO, and P, supplemented by images of specimens deposited at several other herbaria (BM, BR, C, COI, EA FHO, LISC, US, and WAG). Species hypotheses were formulated based on a combination of leaf and fruit characters, structure of the fruiting calyx, and geography. Measurements of the vegetative parts and fruits were taken directly from exsiccatae, whereas flowers were rehydrated prior to being dissected and measured. The information provided on locality, habitat, and phenology was obtained from specimen labels. Geocoordinates were assigned post facto when latitude and longitude were not provided (cited below in square brackets); we were unable to assign coordinates to just three collections. For collections lacking precise locality data, we have also indicated this in square brackets. Occurrences were then mapped using ArcGIS v.10.6 and risk of extinction assessments were performed according to the Guidelines for Using the IUCN Red List Categories and Criteria (
| 1. | All leaves with an obtuse, acute, or acuminate apex | 2 |
| – | At least some leaves with a rounded apex (rarely slightly retuse) | 3 |
| 2. | At least some leaves asymmetric, falcate, blade with varying numbers of scattered, minute glands, tertiary venation on lower surface branched, irregularly reticulate; fruits ellipsoid, 10–14 × 5–9 mm; widespread, Senegal to NE Democratic Republic of the Congo | 4. D. smeathmannii |
| – | Leaves symmetric, not falcate, blade with an evident pair of basal glands, one usually closer to the base than the other, tertiary venation on lower surface forming a dense, nearly horizontal reticulation; fruits broadly ellipsoid, 12–15(–20) × 9–11 mm; Cameroon, NE Gabon, S Central African Republic, N Republic of the Congo, NW Democratic Republic of the Congo | 3. D. moutsambotei |
| 3. | Fruits ellipsoid to slightly obovoid, 8–12 × 6–8 mm; coastal forests N of the equator, Ivory Coast to Benin | 2. D. guineensis |
| – | Fruits globose (occasionally very weakly obloid), coastal and riverine forests S of the equator | 4 |
| 4. | Fruits 7–8 × 7–8(–9) mm, fruiting calyx ca 6 mm wide, calyx cup (base to sinus) 0.5–1 mm high, lobes rounded; Angola | 1. D. angolensis |
| – | Fruits (8–)10–11(–12) × 8–11 mm, fruiting calyx 8–10 mm wide, calyx cup (base to sinus) 2–4 mm high, lobes broadly acute; coastal and lowland riverine forest, Kenya, Tanzania, and Mozambique | 5. D. suaheliensis |
ANGOLA – District de Loanda [= Luanda Prov.] • Coastal region [without precise locality]; 1903; fr.; Gossweiler 789; holotype: P! [P04595254]; isotypes: BM, FHO, K! [K001561020].
Diospyros angolensis can be distinguished from other African species previously included in D. ferrea by the combination of leaves with a symmetric (not falcate) lamina that has dark, pitted glands in the proximal half and an obtuse to rounded (rarely acute or retuse) apex, a globose to very slightly ellipsoid fruit 7–8 × 7–9 mm, and a fruiting calyx with 3 rounded lobes 1–1.5 × 3–4 mm.
Leaves of Diospyros depicting abaxial (left) and adaxial (right) surfaces. A. D. angolensis (Gossweiler 9182, K001561021). B. D. guineensis (de Koning 6007, NY02610014). C. D. moutsambotei (Tadjouteu 570, BR0000009522992). D. D. smeathmannii (Morton SL1201, MO-2865964). E. D. suaheliensis (Faulkner 2151, K001561011). Photos by Alexander Linan.
Fruits of Diospyros. A. D. angolensis (holotype, Gossweiler 789, P04595254). B. D. guineensis (Aubréville 903, P04595333). C. D. moutsambotei (Bidault et al. 4494, MO-6956210). D. D. smeathmannii (Jongkind 4327, MO-6379106). E. D. suaheliensis (Faulkner 2151, K001561011). Photos by Porter P. Lowry II.
Tree (?). Bark pale to dark grey, smooth to slightly rugose, lenticels pale brown, round. Young stems (sub)terete, pale to dark grey, initially covered in short, translucent hispid hairs ca 0.1 mm long and sparse, appressed, yellowish brown hairs ca 0.4 mm long. Leaves alternate; petiole 1–4 mm long, 0.5–1 mm diam., flat to sulcate above, tomentose, white to yellowish-brown hairs; lamina 1.1–5.5 × 0.7–2.8 cm, elliptic to orbicular or obovate, symmetric (never falcate), chartaceous to thinly coriaceous, with sparse indument above and below (likely caducous, t-shaped), discolourous, dark brown above, chestnut brown below, with dark, pitted glands present in varying number in the basal half of the lamina, base acute to obtuse, margin slightly undulate, thickened, apex obtuse to rounded, or rarely acute or retuse; venation pinnate, midrib slightly impressed to flat above, raised below, secondary and tertiary venation usually visible on both sides, sometimes inconspicuous above, more apparent below, darker in younger leaves and concolourous with blade in older leaves, raised below and very slightly so above; secondary venation emerging at varying angles from the midrib, veins (5)6 or 7(8) per side, weakly brochidodromous, forming irregular loops 1/2 to 2/3 of the way between the midrib and margin, the most apical smaller and more steeply angled; tertiary venation similar to the secondary in size and colour, irregularly reticulate and forming further loops between the secondary veins and the margin. Male flowers (known only from immature material) in (1 to)3(4)-flowered axillary cymes, axes densely covered in semi-appressed, brown hairs < 0.5 mm long, peduncle 2–4 mm long, pedicel < 1 mm long; calyx with dense, straight, appressed, golden to golden-brown hairs 0.25–0.5 mm long outside, sparser and shorter inside, basal portion cupuliform, ca 2 × 2–2.5 mm, lobes 3, triangular, 1.5 × 1.5–2 mm, acute; corolla with dense, appressed, light golden hairs ca 1 mm long outside (only on lobes), glabrous inside, tube narrowly ovoid, 1–1.5 × 1.5–2 mm, lobes 3, rounded-triangular, ca 2 × 1.5 mm, imbricate, apex acute; stamens 10, inserted at the base of the corolla tube, filament 0.2–0.5 mm long, anther 1–1.5 mm long; pistillode discoid, ca 0.5 × 1–1.5 mm, densely covered in erect, straight, golden hairs ca 0.75 mm long. Female flowers axillary (sometimes appearing as elongated spikes when the leaves are lost), solitary, pedicel < 1 mm long; calyx densely covered outside and inside with straight, appressed, golden hairs 0.25–0.5 mm long, basal portion cupuliform, ca 2.5 mm × ca 3.5 mm, lobes 3, triangular, valvate, ca 1.5 × 2 mm, obtuse to bluntly acute; corolla glabrous outside except for a rhomboid central area of dense, straight, appressed, golden hairs 0.3–0.5 mm long, glabrous inside, tube cylindrical, ca 4 × 2 mm, lobes 3, rounded-triangular, ca 1 × 1.5 mm, weakly imbricate, apex broadly acute; staminodes lacking; ovary globose, ca 2 × 2 mm, densely covered with straight, appressed, golden hairs ca 0.5 mm long, style 1–1.5 mm long, glabrous. Fruits axillary, solitary, pedicel 1–2 mm long; calyx nearly flat, ca 6 mm wide, with minute, dense, golden hairs ca 0.3 mm long outside, shorter inside, basal portion shallowly cupuliform, 0.5–1 × 4–5 mm, lobes 3, rounded, entire, 1–1.5 × 3–4 mm, apex rounded; fruit globose to very slightly ellipsoid, 7–8 × 7–8(–9) mm, reddish brown to orange (in vivo), surface smooth, with sparse, appressed, white hairs ca 0.3 mm long or glabrescent, style persistent at the apex. Seeds 1 or 2, 6–7 × 4–5 mm, ellipsoid or hemi-ellipsoid, irregularly rough, black, dull.
Diospyros angolensis has been collected in coastal and riverine forests, likely at comparatively low elevations, although no information is provided on any of the specimen labels.
Flowering material has been collected in September and October, and fruiting material in June.
This species is named after the only country from which it has thus far been recorded.
Diospyros angolensis has an Extent of Occurrence of 33,236 km2 and a minimum Area of Occupancy of 24 km2. The species is known from ten collections made between 1904 and 1959 representing four different occurrences, none of which fall within any present-day protected area and are therefore likely subjected to various threats such as fire and land clearing for agriculture. Targeted collecting at all these sites is needed to confirm the status of the extant subpopulations. A continuing decline is inferred for the extent of occurrence, area of occupancy, area, extent, and/or quality of habitat, number of locations, and number of mature individuals. With respect to the most serious plausible threat (land clearing), Diospyros angolensis has been recorded at 4 locations and is thus assessed as Endangered: EN B2ab(i,ii,iii,iv,v).
ANGOLA – Without locality data • Welwitsch 2527; P [P04595251]. – Bengo Province • Catete, Calucala; [09°36’36”S, 14°07’48”E]; 70 m; 15 Oct. 1929; Gossweiler 9182; BM, FHO, K [K001561021], LISC [LISC035677], US [US03133208]. – Luanda Province • [Without precise locality]; 1903; Gossweiler 2450; FHO, P [P04595256]. – Namibe Province • Serra da Chela, Bibala; [14°46’12”S, 13°22’12”E]; 400 m; 7 Sep. 1941; Gossweiler 12956; FHO, LISC [LISC035678] • Serra da Chela, Bibala; [14°46’12”S, 13°22’12”E]; 800 m; 7 Sep. 1941; Gossweiler 12956A; LISC [LISC035679] • Serra da Chela, Bibala; [14°46’12”S, 13°22’12”E]; 700 m; 10 Sep. 1941; Gossweiler 12956B; LISC [LISC035680] • Moçamedes, Camucuio, Bibala; [14°09’00”S, 13°14’24”E]; 14 Oct. 1955; Mendes 409; LISC [LISC035681] • Moçamedes, Camucuio, Bibala; [14°06’00”S, 13°13’48”E]; 730 m; 19 Sep. 1959; Teixeira 4-297; LISC [LISC035685, LISC035686] • Moçamedes, Camucuio, Bibala; [14°46’12”S, 13°22’12”E]; 730 m; 29 Sep. 1951; Teixeira 509; LISC [LISC035682] • Moçamedes, Camucuio, Bibala; [14°06’00”S, 13°13’48”E]; 900 m; 18 Sep. 1959; Teixeira 4-559; LISC [LISC035687].
Diospyros angolensis is the only species formerly included in D. ferrea known to occur in Angola. It has not been documented since 1959 and newer collections are thus urgently needed to ascertain the persistence of this species and to develop a better understanding of its biology, ecology, and conservation status.
Our attempts to locate duplicates of the type collection of Diospyros angolensis (Gossweiler 789) in the two main herbaria in Portugal with important holdings from Angola (COI and LISC) were unsuccessful.
Ferreola guineensis
Schumach. & Thonn., Beskrivelse af Guineeiske Planter: 448. 1827. (
Maba guineensis
(Schumach. & Thonn.) A.DC. (
Diospyros ferrea var. guineensis
(Schumach. & Thonn.) Bakh. (
GHANA – Volta Region • Quitta (= Keta); [5°54’S, 0°54’E]; Thonning s.n.; lectotype (designated here): C [C10003856] scan seen; isolectotypes: C [C10003857, C10003858] scans seen.
Shrub ca 1.5 m tall. Bark grey, rugose, densely lenticellate, lenticels brown, elliptic to round. Young stems terete, pale grey, peeling to a darker grey bark, initially covered in sparse, semi-appressed, minute, yellowish hairs ca 0.3–0.7 mm long. Leaves alternate; petiole 1–4 mm long, 0.5–1 mm diam., somewhat glossy and tightly wrinkly (in sicco), terete to flattened above, with sparse, minute, semi-erect, white to yellowish-brown hairs or glabrescent; lamina 1.3–8.6 × 1–5.5 cm, elliptic to slightly ovate or obovate, symmetric (rarely slightly asymmetric), thinly coriaceous, glabrescent above, glabrescent or with scattered, semi-appressed, yellowish hairs ca 0.5 mm long below, concolourous to very slightly discolourous, shiny, with medium green-brown to reddish brown, dark, pitted glands sometimes present in pairs near the leaf base, rarely few elsewhere on the lamina, base attenuate to (broadly) acute, margin flat to weakly undulate and revolute, slightly thickened, sometimes paler than the leaf blade, apex acute, obtuse or nearly rounded, occasionally weakly and broadly acuminate; venation pinnate, midrib slightly impressed to flat above, raised below, secondary and tertiary venation usually visible on both sides, less conspicuous above, concolourous with the blade or slightly darker, raised on both sides; secondary venation acute or decurrent to the midrib, veins 5 or 6(7) per side, weakly brochidodromous, forming irregular loops 1/2 to 2/3 of the way between the midrib and margin, the most apical smaller and more steeply angled; tertiary venation similar to the secondary in size and colour, irregularly reticulate and forming further loops between the secondary veins and the margin. Male flowers in 3- to 6-flowered axillary cymes or pseudo-umbels, axes with dense, semi-appressed, golden hairs < 0.5 mm long, peduncle to 4 mm long, pedicel to 1 mm long; calyx with straight, appressed, golden to golden brown hairs 0.25–0.75 mm long inside and outside, denser closer to the inside margin, basal portion cupuliform, 1.5–2 × 2.5–3 mm, lobes 3, triangular, valvate, 1.5 × 1.5–2 mm, acute to broadly acute; corolla glabrous inside and outside except for a rhomboid central area of dense straight, appressed, golden hairs 0.75–1.25 mm long outside, tube narrowly urceolate, 2–2.5 × 2.5–3 mm, lobes 3, ovate, 2–2.5 × 2–2.5 mm, imbricate, apex acute; stamens 9 or 10, inserted at the base of the corolla tube, filament 0.5–1 mm, anther 1–1.5 mm; pistillode discoid, ca 0.5 × 1–1.5 mm, densely covered in erect, straight golden hairs ca 0.25–0.5 mm long. Female flowers axillary (branches sometimes appearing spike-like when leaves abscise prior to anthesis), solitary, rarely in pairs, pedicel 0.5–1.5 mm long, with sparse, semi-appressed, golden hairs < 0.5 mm long; calyx moderately covered outside and inside in straight, appressed, whitish-golden hairs ca 0.25 mm long, basal portion cupuliform, 2–2.5 × ca 3 mm, lobes 3, broadly triangular, valvate, ca 1 × 2 mm, obtuse; corolla glabrous except for a rhomboid central area of dense straight, appressed, whitish-golden hairs 0.3–0.5 mm long, glabrous inside, tube cylindrical to slightly urceolate, 2.5–3 × ca 2 mm, lobes 3, broadly ovate, spreading, ca 3 × 2.5 mm, weakly imbricate, apex rounded; staminodes lacking; ovary ovoid, rounded-triangular in cross section, ca 1.5 × 1.5–2 mm, densely covered in straight, appressed, golden hairs ca 0.5 mm long, style 1–1.5 mm long, glabrous. Fruits axillary, solitary, pedicel to 0.5–1.5 mm; calyx cupuliform, 5–7 mm wide, with short, dense, appressed, pale hairs < 0.5 mm long, denser inside, basal portion cupuliform, not closely appressed to the fruit wall (in sicco), 2–3.5 × 5–7 mm, lobes 3, broadly triangular, entire, 0.5–2.5 × 4–6 mm, apex broadly acute to obtuse; fruit ellipsoid to slightly ovoid, 8–12 × 6–8 mm, orangish brown (in vivo), surface smooth, with sparse, appressed, golden hairs ca 0.5 mm long, denser toward the apex, style persistent at the apex. Seeds 1 to 3, 9–10 × 3–4.5 mm, narrowly ovoid to ovoid-wedge shaped, minutely verrucose, dark grey, dull.
Ghana, Ivory Coast, and Benin (Fig.
Diospyros guineensis is known from coastal areas, on sandy or rocky soil behind dunes, at elevations close to sea level.
Flowering material has been collected in May and August to October, and fruiting material in February and April.
Diospyros guineensis has an Extent of Occurrence of 49,057 km2 and a minimum Area of Occupancy of 48 km2. The species is known from 12 collections made between 1907 and 1975 and a more recent one made in 1994, representing eight different occurrences, none of which fall within any formally protected areas and are therefore likely subjected to various threats including fire and land clearing for agriculture. Targeted collecting at all these sites is needed to confirm the status of the extant subpopulations. A continuing decline is inferred for the extent of occurrence, area of occupancy, area, extent, and/or quality of habitat, number of locations, and number of mature individuals. With respect to the most serious plausible threat (land clearing), Diospyros guineensis has been recorded at eight locations and is thus assessed as Vulnerable: VU B2ab(i,ii,iii,iv,v).
BENIN – Littoral Department • Cotonou; [06°21’23”N, 02°18’50”E]; 20 Mar. 1910; Chevalier 23364; P [P04595323, P04595320] • Cotonou; [06°21’32”N, 02°19’35”E]; 29 Mar. 1910; Debeaux 345; K, P [P04595321, P04595322].
GHANA – Without locality data • 20 Feb. 1932; Aubréville 903; P [P04595333].
IVORY COAST – Abidjan District • Abouabo; [05°17’56”N, 04°01’10”W]; 16 Oct. 1949; Adam 6498; P [P04595328] • Port-Bouët; [05°14’30”N, 03°52’10”W]; 1932; Aubréville 1514; BR [BR0000016101500], P [P04595334; P04595335]. – Bas-Sassandra District • Sassandra Port; [04°56’24”N, 06°05’54”W]; 3 May 1907; Chevalier 17946; BR [BR0000016101487], FHO [FHO-131141], P [P04595324, P04595325, P04595326], WAG [WAG.1777958]. – Lagunes District • Entre Jacqueville et Toukouzou; [05°09’00”N, 04°47’24”W]; 0–50 m; 18 Apr. 1994; Aké Assi 18943; MO [MO-5657500]. – Montagnes District • Fabou; [04°22’03”N, 07°30’59”W]; 12 Aug. 1907; Chevalier 19932; BR [BR0000016101494], FHO [FHO-131142], P [P04595332], WAG [WAG.1777956]. – Mouri Region • Mouri, near Cape Coast; [05°07’37”N, 01°13’13”W]; 26 Apr. 1961; Hall 1907; K [K001561026]. – Woroba District • East of Tabou; [04°25’00”N, 07°20’00”W]; 10 m; 3 Sep. 1975; Beentje 853; BR [BR0000016101357], K, MO [MO-6573490], WAG [WAG.1777954, WAG.1777955] • Tabou; [04°26’17”N, 07°19’02”W]; 17 Aug. 1907; Chevalier 19961bis; P [P04595329] • Tabou; [04°25’00”N, 07°21’00”W]; 11 Sep. 1975; de Koning 6007; BR [BR0000016101340], MO [MO-6347092], WAG [WAG.1777951], NY [NY02610014].
GABON – Ogooué-Ivindo • PN d’Ivindo, station de recherche Ipassa; 00°30’10”N, 12°47’45”E; 477 m; 14 Mar. 2019; fr.; Bidault et al. 4494; holotype: MO! [MO-6956210]; isotypes: BR, BRLU [BRLU0026875], LBV, P! [P01085637], WAG.
Diospyros moutsambotei E.Mestre, Meeprom, H.N.Rakouth & Lowry can be distinguished from other African species previously included in D. ferrea by the combination of leaves with a symmetric (not falcate) lamina that has an evident pair of basal glands and an acuminate (rarely acute) apex, a broadly ellipsoid fruit 12–15 × 9–11 mm, and a fruiting calyx with 3 very broadly rounded-triangular lobes ca 2–3 × 9–11 mm.
Treelet 2–8 m tall. Bark dark grey-brown, surface irregular, lenticels brown, round and prominent. Young stems terete, dark brown to black, with dense, persistent, shortly hispid, pale hairs ca 0.5 mm long, and sparse, caducous, longer, golden hairs < 0.1 mm long. Leaves alternate; petiole 1.5–5 mm long, 0.5–1 mm diam., terete, flat or shallowly sulcate above, with the same two types of indument as the stem; lamina 1.5–10.5 × 1–4.5 cm, ovate to elliptic, rarely obovate or suborbicular, consistently symmetric (never falcate), coriaceous, glabrous on both sides, concolourous to weakly discolourous, glossy, dark green above, pale green below, with an evident pair of glands toward the base, borne equally or unequally distant to the petiole, base obtuse to acute, margin slightly thickened, sometimes minutely revolute, apex acuminate, rarely acute, the acumen to 15 mm long and rounded; venation pinnate, midvein sulcate above, raised below, secondary and tertiary venation visible on both sides, more so below, darker in younger leaves and lighter in older leaves, raised below and slightly so above; secondary venation forming a nearly right angle with the midrib and slightly decurrent to it, veins(6)8 or 9(10) per side, brochidodromous, forming loops 2/3 or more of the way between the midrib and margin, the most apical ones smaller and more steeply angled; tertiary venation slightly finer than the secondary, forming a nearly horizontal reticulation and further loops between the secondary veins and the margin. Male flowers in dense, axillary clusters of up 9 flowers, pedicel < 1 mm; calyx moderately covered in straight, appressed, golden to golden brown hairs 0.5–0.75 mm long outside, nearly glabrous inside, basal portion cupuliform, ca 3.5 × ca 4 mm, lobes 3, broadly triangular, 1.5 × 3–3.5 mm, acute; corolla creamy white (in vivo), with dense, appressed, golden to light brown hairs 0.75–1 mm long outside, glabrous inside, tube narrowly urceolate, 4–5 × 2.5–3 mm, lobes 3, ovate, ca 3.5 × 2 mm, imbricate, apex acute; stamens 5, inserted at the base of the corolla tube, very pale cream-brown (in vivo), filament 0.5–1 mm, anther 3.5–4 mm; pistillode broadly ovoid, 1–1.5 × 1–2 mm, densely covered in erect, straight, golden hairs 0.75 mm long. Female flowers axillary, solitary or in short, few-flowered cymes, axes with sparse straight, appressed, whitish-golden hairs ca 0.25 mm long, pedicel 1–2 mm long; calyx moderately covered outside and more sparsely inside in straight, appressed, whitish-golden hairs ca 0.25 mm long, basal portion cupuliform, 4.5–5 × 3.5–4 mm, lobes 3, shallowly and broadly triangular, spreading, ca 0.5 × 3 mm, short-acuminate; corolla densely covered in straight, appressed, whitish-golden hairs to 1 mm long, becoming glabrous towards the lobe margins, glabrous inside, tube slightly urceolate, 5.5–6 × 3–3.5 mm, lobes 3(4), broadly ovate, spreading, weakly imbricate, 2–2.5 × 2–2.5 mm, apex rounded-acute; staminodes lacking; ovary ovoid, rounded-triangular in cross section, ca 3 × 2 mm, densely covered in straight, appressed, golden hairs 0.75 mm long, style 1.0–1.5 mm long, base with minute, appressed hairs, apex glabrous. Fruits axillary, solitary or in pairs, pedicel 1–1.5 mm, indumentum persistent; calyx cupuliform, 9–11 mm wide, dark brown (in sicco), with moderately dense, appressed, greyish-white hairs to 1 mm long outside, sparser, shorter and darker inside, basal portion cupuliform, 5–6 × ca 9–11 mm, lobes 3, shallow, entire, very broadly rounded-triangular, ca 2–3 × 9–11 mm, apex short-acuminate; fruit globose when immature, broadly ellipsoid when ripe, 12–15(–20) × 9–11 mm, surface smooth, with sparse, appressed, golden hairs, denser toward the apex, brown (in sicco), style persistent at the apex. Seed 1, 12–20 × 4–6 mm, narrowly obloid, slightly asymmetrical, minutely rugose, dark reddish brown and somewhat glossy (in sicco).
Photos of Diospyros moutsambotei in the field. A. Lower surface of branch with fruits. B. Upper surface of same branch. C. Fruits on lower surface of branch. D. Detail of fruits. E. Closeup of leaf base (abaxial surface) showing pair of glands. F. Inflorescence of male flowers (side view). G. Inflorescence of male flowers (top view). A–E, Bidault et al. 4494, photos by Ehoarn Bidault; F–G, Harris & Moutsamboté 9672, photos by David J. Harris.
Cameroon, Central African Republic, Gabon, Democratic Republic of the Congo, and Republic of the Congo (Fig.
Diospyros moutsambotei is known from riverine and periodically inundated forests, often associated with vegetation dominated by Gilbertiodendron dewevrei (De Wild.) J.Léonard (Fabaceae). It has been recorded at 350–700 m elevation.
Flowering material has been collected in December and from February to April, and fruiting material in February, March, May, and June.
This species is named in honour of Jean-Marie Moutsamboté (1953–2022), who was one of the most highly regarded botanists working in Central Africa. Prof. Moutsamboté contributed greatly to our knowledge of the forests of the Republic of the Congo, participating in dozens of botanical expeditions throughout the country. He served as Head of the Botany Laboratory at the Centre d’Études sur les Ressources végétales and the National Herbarium in Brazzaville from 1986 to 2001 and then taught plant biology and forestry at the Université Marien N’Gouabi before returning to the National Herbarium in 2015, where he headed the Ecology-Phytosociology unit. Jean-Marie has left an important legacy and will long be remembered by the many students and colleagues who had the pleasure of working with him.
Diospyros moutsambotei has an Extent of Occurrence of 449,586 km2 and a minimum Area of Occupancy of 92 km2. The species is known from 25 collections representing ten different occurrences, some of which fall within formally protected areas whereas others that do not are likely subjected to several threats, including land clearing for agriculture and forest exploitation. Targeted collecting at all these sites is needed to confirm the status of the extant subpopulations. A continuing decline is inferred for the extent of occurrence, area of occupancy, area, extent, and/or quality of habitat, number of locations, and number of mature individuals. With respect to the most serious plausible threat (land clearing), Diospyros moutsambotei has been recorded at ten locations and is thus assessed as Vulnerable: VU B2ab(i,ii,iii,iv,v).
GABON – Ogooué-Lolo Province • Parc National de l’Ivindo; 00°15’10”S, 12°44’16”E; 393 m; 12 Feb. 2010; Dauby 2292; BRLU, LBV, MO • Lastoursville; 00°15’19”S, 12°44’06”E; 387 m; 18 Apr. 2010; Nguema 1248; BRLU, K, LBV, MO, P • 10 km S of Majojou; [00°30’39”N, 12°48’13”E]; 500 m; 24 Feb. 1979; Florence 1676; P [P04539709]. – Woleu-Ntem Province • Minkébé area; [01°30’00”N, 12°49’00”E]; 16 May 1990; Minkébé Series W341; MO [MO-6055800], WAG [WAG.1778699, WAG.1778700] • Minkébé district, Nsye valley; [01°30’00”N, 12°48’00”E]; 21 Feb. 1990; Wieringa 599; BR [BR0000021712791], K, MO [MO-6379125], WAG [WAG.1778707].
CAMEROON – Central Region • Ndanan 2; [03°36’29”N, 11°34’41”E]; 710 m; 22 Mar. 2004; Cheek 11937; K, MO [MO-4875195], WAG [WAG.1778730] • Mefou National Park; [03°37’12”N, 11°34’48”E]; 700 m; 23 Mar. 2004; Tadjouteu 570; BR [BR0000009522992], K [K000339048], MO [MO-4811439], WAG [WAG.1778728]. – East Region • 2 km W of the Sangha River; [02°21’00”N, 16°07’00”E]; 350 m; 17 Nov. 1998; Harris 6459; E [E01021159]. – South Region • between N’Kolandom and N’Koemvone; [02°47’00”N, 11°09’00”E]; 650 m; 24 Feb. 1975; de Wilde 7995; BR [BR0000016101654], K [K001394634], MO [MO-5654747], P [P04589936], WAG [WAG.1778733, WAG.1778734].
CENTRAL AFRICAN REPUBLIC – Haute-Sangha Prefecture • Dzanga-Sangha Reserve; [02°21’00”N, 16°09’00”E]; 350 m; 9 Sep. 1988; Harris 1091; BR [BR0000100027273], MO [MO-5965327] • 45 km S of Lidjombo; [02°22’00”N, 16°09’00”E]; 350 m; 11 Feb. 1989; Harris 1823; BR [BR0000100027242], K [K001561010], MO [MO-3823625] • 40 km S of Lidjombo; [02°25’00”N, 16°09’00”E]; 350 m; 15 Feb. 1989; Harris 1876; BR [BR0000100027266], MO [MO-5965320] • 45 km S of Lidjambo; [02°21’00”N, 16°09’00”E]; 350 m; 23 Feb. 1989; Harris 1918; BR [BR000016101630], FHO [FHO-164548], K, MO [MO-5672092], WAG [WAG.1778732] • 45 km S of Lidjombo; [02°22’00”N, 16°11’00”E]; 350 m; 12 Mar. 1989; Harris 2032; BR [BR000016101623], FHO [FHO-164549], K [K001561012], MO [MO- 5672091], WAG [WAG.1778731] • 12 km S of Lidjombo; [02°34’00”N, 16°05’00”E]; 28 Mar. 1994; Harris 4854; BR [BR0000100027297], E [E01021135].
DEMOCRATIC REPUBLIC OF THE CONGO – Équateur Province • Bolomba; [00°03’44”N, 19°19’38”E]; 26 Feb. 1958; Evrard 3541; BR [BR0000016101715], WAG [WAG.1778735]. – Ituri Province • Epulu, zone de Mambasa (Ituri Forest); [01°25’00”N, 28°35’00”E]; 750 m; 5 Dec. 1987; Hart 783; MO [MO-4292715]. – Tshuapa Province • Befale; [00°22’20”N, 20°50’19”E]; 3 Jun. 1958; Evrard 4166; BR [BR0000016101692, BR0000016101708], FHO [FHO-105514].
REPUBLIC OF THE CONGO – Sangha Department • Nouabalé-Ndoki National Park; [02°10’00”N, 16°31’00”E]; 10 Jun. 2002; Harris 7961; E [E01021158] • 2.5 km NNE of Bomassa village; [02°13’30”N, 16°11’43”E]; 22 Mar. 2010; Harris & Moutsamboté 9672; E [E00397397], IEC • [Without precise locality]; [02°11’54”N, 16°17’28”E]; 22 May 2011; Ndolo Ebika 607; E [E00607099], IEC, WAG [WAG.1778729] • 33.5 km NE of Kabo; [02°01’53”N, 16°21’12”E]; 10 Nov. 2006; Nzolani Silaho 1133; E [E01021156], IEC • 55 km NE of Kabo, Nouabalé-Ndoki National Park; [02°22’17”N, 16°28’25”E]; 25 Feb. 2007; Nzolani Silaho 2724; E [E01021155], IEC • Makambi Bai; [02°09’37”N, 16°15’17”E]; 350 m; 22 Mar. 2010; Nzolani Silaho 4573; E [E01021130], IEC.
Of the taxa treated here, Diospyros moutsambotei is the only one whose distribution overlaps with that of another species, namely the widespread D. smeathmannii. It can, however, be easily distinguished by its symmetrical leaves that are not falcate (vs some or all asymmetric and falcate in D. smeathmannii), the presence of a pair of basal glands, most prominently visible on the abaxial leaf surface (vs varying numbers of scattered, minute glands), and its broadly ellipsoid fruits that measure 12–15 × 9–11 mm (vs ellipsoid and 10–14 × 5–9 mm).
Maba smeathmannii
A.DC., Prodromus Systematis Naturalis Regni Vegetabilis, vol. 8: 241. 1844 (as ‘smeathmanni’). (
Maba vacciniifolia
Benth. (in
Maba secundiflora
Hutch. (
IVORY COAST • [Without precise locality]; s.d.; Smeathman s.n.; holotype: G-DC [G00142244, mounted on 3 sheets] scan seen; isotype: BM [BM000925475] scan seen.
Shrub, tree 2–10 m tall, 4–10 cm DBH. Bark pale grey, rugose, lenticels orange-brown, longitudinally elliptic. Young stems terete, dark brown to black, initially covered in dense, hispid, short, translucent pale yellow hairs < 0.5 mm long and caducous, yellow-brown hairs 0.75–1.5 mm long. Leaves alternate; petiole 1.5–7 mm long, 0.5–1 mm diam., terete to nearly flat, occasionally narrowly sulcate, with short, erect, pale yellow hairs < 0.5 mm long; lamina 1.5–11.5 × 0.6–4 cm, elliptic to occasionally obovate, often asymmetric and somewhat falcate, chartaceous to subcoriaceous, young leaves with a very sparse indumentum above, mature leaves glabrous above except for sparse to dense, tomentose hairs ca 0.5–1.5 mm long at the base and along the midvein below, sometimes discolourous (in sicco), dull green-brown to dark brown above, glossy, similarly coloured, occasionally glossy below (in vivo), young leaves red above, pale green below, mature leaves deep green and glossy above, paler and glaucous or glossy below, with scattered minute glands in varying numbers, primarily in the lower half of the lamina, base acute, margin flat or weakly undulated, slightly thickened and revolute, apex acuminate, acute or rarely rounded, acumen to 15 mm, rounded at the tip; venation pinnate, midrib narrowly sulcate above, raised below, secondary and tertiary venation faintly visible on both sides, concolourous with the blade to slightly lighter in colour below, raised on both sides; secondary venation emerging slightly decurrently at acute angles from the midrib, veins 7 to 9 per side, brochidodromous, forming loops sometimes close to the margin, the most apical ones smaller and more steeply angled; tertiary venation less conspicuous than the secondary, irregularly reticulate, branching, and forming further loops between the secondary veins and the margin. Male flowers in 2- to 4-flowered axillary cymes, axes densely covered in semi-appressed, tomentose, golden hairs ca 0.5 mm long, peduncle 0.5–2.5 mm long, pedicel 0.5–1 mm; calyx with moderate to sparse straight, appressed, golden to golden brown hairs 0.5-0.75 mm long outside, glabrous inside, basal portion cupuliform, 1.5–2.5 × 2.5–3 mm, lobes 3, broadly rounded-triangular to triangular, slightly reflexed, 1–2 × 1.5–2.5 mm, apex broadly acute; corolla glabrous outside except for a rhomboid to narrowly-rhomboid central area with dense, straight, appressed, golden to light brown hairs 1–1.25 mm long, glabrous inside, tube cupuliform, 2.5–4 × 2.5–3 mm, lobes 3, rounded-triangular, 2–3 × 2–3 mm, slightly imbricate, apex acute; stamens (3)6 to 10(11), inserted at the base of the corolla tube, filament 0.75–1 mm long, anther 1.5–2.5 mm long; pistillode ovoid, 1.5–2 × 1.5 mm, densely covered in erect, straight, golden hairs 0.75 mm long. Female flowers axillary, solitary, pedicel 1–2.5 mm long, densely covered in semi-appressed, sericeous, brown hairs < 0.5 mm long; calyx densely covered in straight, appressed, golden hairs 0.25–0.5 mm long outside, glabrous inside, basal portion cupuliform, 2–3 × 2.5–3.5 mm, lobes 3, entire, broadly triangular, 1–1.5 × 2–3 mm, apex broadly acute; corolla glabrous except for a rhomboid central area of dense straight, appressed, golden hairs 0.75–1 mm long, glabrous inside, tube cylindrical, 2–3 × ca 2 mm, lobes ovate, slightly spreading, 2–2.5 × 1.5–2 mm, weakly imbricate, apex shortly acute; staminodes lacking; ovary ovoid, rounded-triangular in cross section, 2–2.5 × 2 mm, densely covered in straight, appressed, golden hairs ca 0.25 mm long, style ca 1 mm long, with the same indumentum as the ovary, increasingly sparse towards the stigmas. Fruits axillary, solitary, pedicel 1–1.5 mm; calyx cupuliform, 6–9 mm wide, with minute, moderately dense indumentum, sometimes glabrescent, brown (in vivo), basal portion cupuliform, 9–12 × 3.5–4 mm, lobes 3, broadly triangular, entire, 2–2.5 × 8–11 mm, apex broadly acute; fruit ellipsoid, 10–14 × 5–9 mm, orange at maturity (in vivo), surface smooth, with scattered, short, appressed, yellowish hairs, denser toward the apex, style persistent at the apex. Seeds 2, 10–11 × 4–4.5 mm, hemi-ellipsoid, surface uneven, dark grey to black, somewhat shiny.
Senegal to NE Democratic Republic of the Congo (Fig.
Diospyros smeathmannii has been collected most often on river/stream banks and in swamps, at sites from sea level to ca 500 m, rarely up to 1250 m.
Flowering material has been collected nearly year-round except in June and August, and fruiting material from September to April.
Diospyros smeathmannii has an Extent of Occurrence of 5,504,397 km2 and a minimum Area of Occupancy of 280 km2. The species is known from 87 collections, more than two-thirds of which date from prior to 1984, representing more than 50 different occurrences, some of which occur within protected areas. Targeted collecting at historic sites is needed to confirm the status of extant subpopulations. Diospyros smeathmannii is assessed as Least Concern: LC.
CAMEROON – Without locality data • May–Jun. 1946; Aubréville 118C; P [P04595314, P04595315]. – North Region • 60 km SW of Tcholliré; [08°23’15”N, 14°09’03”E]; 2 Feb. 1977; Fotius 2580; P [P04595258, P04595259].
CENTRAL AFRICAN REPUBLIC – Bamingui-Bangoran Prefecture • Manovo-Gounda-St. Floris National Park; [08°41’00”N, 21°18’00”E]; 580 m; 19 Sep. 1983; Fay 5737; MO [MO-3853649]. – Ouaka Prefecture • Riv. Fo, 30 km ouest Moroubai; 22 Apr. 1923; R.P. Tisserant 1084; FHO [FHO-128781], P [P04595255].
DEMOCRATIC REPUBLIC OF THE CONGO – Ituri Province • Epulu; [01°24’36”N, 28°34’48”E]; 21 Feb. 1914; Bequaert 2595; BR [BR0000016101722, BR0000016101739, BR0000016101746] • Bela; [00°42’00”N, 29°15’00”E]; 400 m; s.d.; Michelson 464; BR [BR0000016101685] • Zone de Mambasa (Ituri Forest); [01°10’00”N, 28°23’00”E]; 5 Mar. 1998; Ntumba 21; BR [BR0000016101678], MO [MO-5659564] • Zone de Mambasa (Ituri Forest); [01°10’00”N, 28°23’00”E]; 5 Mar. 1998; Ntumba 39; BR [BR0000016101449]. – Lomami Province • Beondo; [04°19’48”S, 20°36’00”E]; 20 Apr. 1959; Bamps 570; BR [BR0000016101777], FHO [FHO-105509] • [Without precise locality]; 7 Feb. 1914; Bequaert 2362; BR [BR0000016101753, BR0000016101760], FHO [FHO-105464].
GABON – Estuaire Province • Ovang, E of Libreville; [00°29’00”N, 09°31’00”E]; 23 Jul. 1985; Bos 10766; BR [BR0000016101647], K, MO [MO-4310171], P [P04595303], WAG [WAG.1778709, WAG.1778710] • Same locality; 1 m; 15 Oct. 2000; Breteler 15653; BR [BR0000016101463], MO [MO-6857195], WAG [WAG.1778711, WAG.1778712] • Same locality; 15 Feb. 1985; Louis 1715; BR [BR0000016101456], MO [MO-6480264], WAG [WAG.1778698] • 1 km N of the landing stage of Ovang; [00°29’00”N, 09°31’00”E]; 15 Feb. 1985; Reitsma 830; MO [MO-4285103], WAG [WAG.1778708].
GHANA – Kada Region • Kada Agric. Research Station; [06°08’24”N, 00°53’24”W]; 17 Mar. 1972; Enti Sp645; MO [MO-2094474].
GUINEA – Without locality data • Apr. 1905; Chevalier 13278; P [P04595245] • Apr. 1905; Chevalier 13320; P [P04595265] • Apr. 1905; Chevalier 13336; P [P04595246]; Chevalier 13339; FHO [FHO-131146], P [P04595266, P04595267] • Apr. 1905; Chevalier 13607; P [P04595268] • 1903; Pobéguin 851; P [P04595271]. – Boké Region • Bel-Air, vers Drameya; [10°13’35”N, 14°27’30”W]; 2 m; 21 Oct. 2016; Bidault 2498; BRLU, HNG, MO [MO-3061991], P [P01192530]. – Kankan Region • Dabola, chutes du Tinkisso; [10°43’42”N, 11°10’17”W]; 19 Apr. 1949; Adam 4535; MO [MO-2109761], P [P04595270] • Dabola, towards Mamou; [10°43’12”N, 11°24’00”W]; 20 Apr. 1949; Adam 4551; MO [MO-2109758], P [P04595269]. – Kindia Region • 4 km E of the village Taban; 09°53’32”N, 10°01’50”W; 450 m; 21 Apr. 2017; van der Burgt 2122; K [K001286269]. – Labe Region • 10 km après Misside; [12°05’15”N, 12°19’23”W]; 12 Feb. 1979; Lisowski 51718; BR [BR0000016101388, BR0000016101395] • Mont Loura; [12°04’17”N, 12°05’04”W]; 17 Feb. 1979; Lisowski 51749; BR [BR0000016101371].
GUINEA-BISSAU – Bissau Region • Bissalanca; [11°53’21”N, 14°55’32”W]; 10 Jan. 1945; Espirito Santo 1633; FHO [FHO-95112], K. – Cacheu Region • Canchungo; [12°03’36”N, 16°01’48”W]; 19 Apr. 1945; Espirito Santo 1937; FHO [FHO-95110], K, MO [MO-2427334], WAG [WAG.1777963]. – Oio Region • Farim, Begene; [12°26’24”N, 15°32’24”W]; 23 May 1947; Espirito Santo 2386; K [K001561017], MO [MO-2427339], WAG [WAG.1777962].
IVORY COAST – Bas Sassandra District • Entre Tabou et Bériby [= Béréby]; 17 Aug. 1907; Chevalier 19961 bis; P [P04595329]. – Lagunes District • Réserve de Lamto; [06°13’12”N, 05°01’12”W]; 80 m; 1 Dec. 1986; D. & L. Gautier-Béguin 446; G, MO [MO-5742662] • Réserve de Lamto; 6 Sep. 1987; D. & L. Gautier-Béguin 577; G, MO [MO-5725340]. – Sassandra-Marahoué District • Parc National de la Marahoue; [07°04’00”N, 05°55’00”W]; 300 m; 9 Feb. 1998; Jongkind 4327; BR [BR0000016101333], FHO, MO [MO-6379106], WAG [WAG.1777952]. – Vallee du Bandama District • Farakoro; [09°26’24”N, 07°06’36”W]; 21 Apr. 1972; Aké Assi 11752; G, MO [MO-04912340].
MALI – Bamoko Region • Bamako; [12°33’36”N, 08°10’48”W]; 438 m; 4 Dec. 2003; Birnbaum 699; IFAN, P [P05218584] • West of Djikoroni; [12°36’00”N, 08°02’00”W]; 31 Jan. 1973; N’Golo 1275; MO [MO-3325018], WAG [WAG.1777969] • Forêt de la Faya; [12°37’36”N, 07°33’17”W]; 7 Dec. 1978; Floret 1385; P [P04539852] • Djikoroni; [12°39’41”N, 07°54’26”W]; 31 Jan. 1973; Cissé 493C; WAG [WAG.1777967, WAG.1777968]. – Kayes Region • Cercle de Kita, Séta-Oulé (Bohké), Galerie du Dabara; 12 Apr. 1939; Dubois 154 bis; P [P00531015] • Kenieba Cercle, Falea Rockgate Mine; [12°17’42”N, 11°18’50”W]; 410 m; 27 Oct. 2012; Luke 15983; K [K001561018]. – Kita Region • Faille de Dabarako; [12°21’00”N, 09°54’36”W]; 362 m; 12 Feb. 2007; Birnbaum 1164; IFAN, P [P05217987] • Faille de Dabarako; [12°21’00”N, 09°54’36”W]; 355 m; 12 Feb. 2007; Birnbaum 1175; IFAN, P [P05218587] • Galerie du Bale; [12°04’12”N, 09°54’00”W]; 351 m; 13 Feb. 2007; Birnbaum 1183; IFAN, P [P05218427]. – Koulikoro Region • Famansa; [12°14’07”N, 08°45’17”W]; 18 May 1975; Aberlin 603; P [P04589934] • Vallée du Soulou; [12°38’39”N, 08°29’53”W]; 4 Apr. 1934; Dubois 154; P [P00531014, P00531015]. – Sikasso Region • Ninedoni; [10°55’12”N, 05°42’00”W]; 0 m; 27 Jan. 2005; Birnbaum 864; IFAN, K, P [P05218583] • Ninedoni; [10°55’12”N, 05°42’00”W]; 0 m; 27 Jan. 2005; Birnbaum 865; IFAN, K, P [P05218586]. – Unknown Region • Lido, Marigat; 23 Mar. 1976; Aberlin 603 bis; P [P04589935].
NIGERIA – Without locality data • 30 Jan. 1970; Schweinfurth 2902; K [K001561028] • 14 Nov. 1968–16 Aug. 1969; van Meer 1076; BR [BR0000016101470], K, MO [MO-6857182]. – Anambra State • Mamu Forest Reserve; [06°10’12”N, 07°10’12”E]; Jan. 1945; Burtt 21; K [K001561009]. – Bauchi State • Katagum District; [12°15’36”N, 10°21’22”E]; s.d.; Dalziel 413; FHO, K [K000350652] • Bauchi Plateau; [09°59’39”N, 09°29’12”E]; May 1928; Lely 266; FHO [FHO-32271], K [K001561008]. – Cross River State • Iyamojong F.R.; [05°58’10”N, 08°21’30”E]; 14 Apr. 1959; Binuyo 41231; BR [BR0000016101531], FHO [FHO-108729], K [K001561004], WAG [WAG.1777981]. – Kaduna State • Kaduna; [10°32’05”N, 07°33’09”E]; 9 Dec. 1949; Meikle 786; BR [BR0000016101524], K, P [P04595261, P04595263]. – Ondo State • Akure Forest Reserve, Aponmu; [07°18’09”N, 05°01’46”E]; 19 Oct. 1946; Keay 21556; FHO [FHO-74111, FHO-74112], K [K001561023] • Akure Forest Reserve, Owena; [07°18’33”N, 05°01’48”E]; 11 Dec. 1957; Keay 37273; K [K001561025] • Aponmu Pilot Sawmill Area; [07°14’26”N, 05°03’37”E]; 18 Oct. 1946; Onochie 20234; FHO [FHO-73743, FHO-73849, FHO-81184], K [K001561024]. – Oyo State • Ibadan south Reserve; [07°13’59”N, 03°58’01”E]; 22 Sep. 1948; Chizea 19480; BR [BR0000016101548]. – Plateau State • Vom; [09°44’11”N, 08°47’47”E]; 20 Apr. 1972; Gbile 1075; BR [BR0000016101517], K • Kogigiri, near reservoir above Jos; [09°54’09”N, 08°50’34”E]; 1250 m; 17 Oct. 1957; Hepper 1061; BR [BR0000016101555], K, P [P04595262] • Heipang; [09°38’44”N, 08°52’55”E]; 11 Apr. 1958; Keay 37603; FHO, K [K001561007] • Valley on Plateau; [09°53’22”N, 08°50’56”E]; 1219 m; 14 Nov. 1980; Sharland 1527; K [K001561006] • Jos, Heipan Kurmi, near Heipan Tapu; [09°03’32”N, 08°50’13”E]; 9 Nov. 1963; White 8287; FHO [FHO-114635, FHO-114671, FHO-115487, FHO-115505] • Jos plateau, Kogon Giri Forest Reserve; [09°03’32”N, 08°50’13”E]; 9 Feb. 1963; White 8304; FHO [FHO-115478]. – Taraba State • Gangumi near the bank of River Sulli; [07°21’53”N, 11°29’24”E]; 16 Dec. 1954; Latilo 28911; FHO, K [K001560122].
SENEGAL – Without locality data • 20 Mar. 1964; Berhaut 7253; BR [BR0000016101821], P [P04595250]. – Dakar Region • Sangalkam; [14°46’38”N, 17°13’36”W]; Dec. 1950; Berhaut 1030; BR [BR0000016101845], P [P04595248] • Mbao; [14°43’46”N, 17°19’26”W]; 24 Apr. 1957; Berhaut 5570; P [P04595313]. – Kédougou Region • Nepin; [12°23’24”N, 12°33’00”W]; 8 Mar. 1962; Adam 18868; MO [MO-2109623] • Cirque de Díndefelo; [12°22’02”N, 12°19’18”W]; 15 Nov. 1964; Adam 19961; K [K001561005] • Nikolo-Koba; [13°03’15”N, 12°53’59”W]; Apr. 1951; Berhaut 2927; BR [BR0000016101838], P [P04595249] • Cascade de Dindefello; [12°21’59”N, 12°19’24”W]; 21 m; 21 May 1993; Goudiaby 72; BR [BR0000016101814] • Parc National du Niokolo Koba, Mount Assirik; [12°53’00”N, 12°46’00”W]; 150 m; 5 Apr. 1993; Madsen 1490; MO [MO-5651778] • Same locality; 150 m; 5 Apr. 1993; Madsen 1498; MO [MO-5651777] • Mount Assirik; [12°53’01”N, 12°45’18”W]; 200–300 m; 11 May 1976; Tutin 38; K [K001561019]. – Kolda Region • [Without precise locality]; May 1946–Jun. 1946; Aubréville 109C; P [P04595316]. – Sangalkam Region • Sangalkam; [14°46’12”N, 17°13’12”W]; 22 Jul. 1948; Adam 1615; MO [MO-2109624], P [P04595312]. – Ziguinchor Region • Oussouye; [12°26’57”N, 16°33’42”W]; 6 Sep. 1968; Berhaut 7384; P [P04595311] • Cap Skirring; [12°21’12”N, 16°43’34”W]; 4 m; 12 Dec. 1977; Vanden Berghen 2243; BR [BR0000016101807, BR0000016101234] • Cap Skirring; [12°21’23”N, 16°43’32”W]; 10 m; 17 Sep. 1981; Vanden Berghen 4558; BR [BR0000016101197] • Cap Skirring; [12°21’12”N, 16°43’34”W]; 10 m; 19 Sep. 1981; Vanden Berghen 4599; BR [BR0000016101180] • Cap Skirring; [12°21’12”N, 16°43’34”W]; 10 m; 21 May 1982; Vanden Berghen 5040; BR [BR0000016101258] • Cap Skirring; [12°21’12”N, 16°43’34”W]; 10 m; 25 May 1982; Vanden Berghen 5103; BR [BR0000016101425] • Cap Skirring; [12°23’00”N, 16°45’00”W]; 10 m; 26 May 1982; Vanden Berghen 5110; BR [BR0000016101319], WAG [WAG.1777960] • Cap Skirring; [12°21’15”N, 16°43’41”W]; 3 m; 27 Sep. 1984; Vanden Berghen 6672; BR [BR0000016101326].
SIERRA LEONE – Western Province • John Obey Peninsula; [08°14’00”N, 13°10’00”W]; 4 Mar. 1964; Morton 901; FHO, K, WAG [WAG.1777964] • Same locality; 12 Apr. 1964; Morton SL1201; K [K001561016], MO [MO-2865964, MO-3038711], WAG [WAG.1777965].
KENYA – Coast Province • Lamu Dist., Witu Forest; 2°20’50”S, 40°29’45”E; 16 m; 17 Nov. 1988; ♂ bud, ♂ fl.; Robertson & Luke 5528; holotype: MO! [MO-4240972]; isotypes: EA, K!, US [US03133205] scan seen, WAG [WAG.1778716] scan seen.
Diospyros suaheliensis can be distinguished from other African species previously included in D. ferrea by the combination of leaves with a symmetric (not falcate) lamina that has a few dark, pitted glands toward the base and a retuse, rounded or obtuse (sometimes broadly acute) apex, a globose fruit 8–12 × 8–11 mm, and a fruiting calyx with 3 broadly triangular lobes 3–4 × 6–8 mm.
Tree or shrub 2–12 m tall. Bark pale grey to brownish grey, rugose, lenticels brown, round to longitudinally elliptic. Young stems terete, brown, then pale grey, initially covered in dense, hispid, very short, translucent pale yellow hairs and golden yellow, longer and caducous hairs 0.75–1 mm long. Leaves alternate; petiole 2–8 mm long, 0.5–1.5 mm diam., flat to deeply sulcate, with the same indumentum as the stem or glabrescent; lamina 0.8–8.5 × 0.8–5.6 cm, elliptic to orbicular or slightly obovate, symmetric (never falcate), chartaceous to coriaceous, glabrous above and below, concolourous to slightly discolourous, pale grey-green to dark brown above, often dull-glaucous, pale, dark or reddish brown below, with dark, pitted glands occasionally present in very small numbers in the basal part, base, acute to rounded, rarely cuneate, margin slightly to majorly undulate, thickened, often lighter than the rest of the lamina, apex retuse, rounded, obtuse or less frequently broadly acute; venation pinnate, midrib narrowly impressed to flat above, raised below, secondary and tertiary venation visible on both sides, concolourous with blade, raised on both sides; secondary venation emerging at irregularly acute and decurrent angles from the midrib, (5)6 or 7(8) veins per side, brochidodromous, forming irregular loops close to the margin, the most apical ones smaller and more steeply angled; tertiary venation similar to the secondary in size and colour, irregularly reticulate and forming further loops between the secondary veins and the margin. Male flowers in short, 3-flowered axillary cymes, axes with moderately dense, semi-appressed, golden hairs <0.5 mm long, peduncle 1–3 mm long, pedicel < 1 mm long; calyx with moderately straight, appressed, golden brown hairs 0.25–0.5 mm long outside, sparse and limited to the margin inside, basal portion cupuliform, ca 2.5 × 2.5–3 mm, lobes 3, round to round-triangular, ca 1 × 2.5 mm, apex shortly acute; corolla with dense, appressed straight, golden inside 0.5–1.5 mm long outside on the lobes, glabrous elsewhere, tube shortly campanulate 3 × 2.5 mm, lobes 3, entire, ovate, spreading, 2.5–3 × 2–2.5 mm, apex acute; stamens 9 or 10, inserted at the base of the corolla tube, filament 1.0–1.5 mm long, anther 1.75–2.3 mm long; pistillode discoid, 3- to 6-lobed, ca 0.5 × 1–1.1 mm, densely covered in erect, straight golden hairs ca 0.25 mm long. Female flowers axillary, solitary, subsessile; calyx moderately covered outside and inside in straight appressed golden hairs 0.25–0.75 mm long, basal portion cupuliform, ca 3 × ca 2.5 mm, lobes 3, triangular, ca 1 × 2 mm, acute (often broadly so); corolla and staminodes not seen; ovary obovoid, ca 3 × 3 mm (post receptivity), covered toward the apex with straight, appressed, golden hairs ca 0.25 mm long, style ca 1 mm long, glabrous. Fruits axillary, solitary, pedicel to 1 mm long; calyx cupuliform, 8–10 mm wide, with dense to very sparse, minute, golden hairs 0.25–0.5 mm long, basal portion shallowly cupuliform to flat, 2–4 × 7–11 mm, lobes 3, broadly triangular, margins ciliate, entire, 3–4 × 6–8 mm, apex broadly acute; fruit globose, (8–)10–11(–12) × 8–11 mm, bright red at maturity (in vivo), surface smooth, sparsely tomentose, style persistent at the apex. Seeds 4, 7–7.5 × 3.5–6 mm, ellipsoid-wedge shaped, rugose, dark grey, slightly shiny.
Kenya, Tanzania, and Mozambique (Fig.
Diospyros suaheliensis is known from coastal and lowland riverine forest at 0–250 m elevation.
Flowering material has been collected from September to January, and fruiting material in February, May, July, and November.
The name chosen for this new species reflects its geographic distribution, which corresponds very closely to the Swahilian regional centre of endemism, as defined by
Diospyros suaheliensis has an Extent of Occurrence of 404,496 km2 and a minimum Area of Occupancy of 76 km2. The species is known from 20 collections representing seven different occurrences, some of which fall within formally protected areas whereas others that do not are likely subjected to several threats, including land clearing for agriculture and fire. Targeted collecting at historic sites is needed to confirm the status of the extant subpopulations. A continuing decline is inferred for the extent of occurrence, area of occupancy, area, extent, and/or quality of habitat, number of locations, and number of mature individuals. Regarding the most serious plausible threat (land clearing for agriculture), Diospyros suaheliensis has been recorded at seven locations and is thus assessed as Vulnerable: VU B2ab(i,ii,iii,iv,v).
KENYA – Kwale District • South of Kinondo; [04°24’00”S, 39°33’00”E]; 5 m; 2 Mar. 1977; Faden 77/643; FHO [FHO0080385], K, US [US03133207] • Kaya Kinondo; [04°23’00”S, 39°32’00”E]; 5 m; 10 Nov. 1992; Harvey 45; MO [MO-5663315] • Kaya Mgawani; [04°24’00”S, 39°32’00”E]; 7 m; 4 Jan. 1992; Luke 3036; K, MO [MO-5659987], US [US03133204] • Kaya Kinondo; [04°23’39”S, 39°32’50”E]; 16 Jul. 1987; Robertson 4922; K [K001561013]. – Lamu District • 3 km E of Tana river in Home forest; [02°00’00”S, 40°11’00”E]; 25 m; 14 Feb. 1973; Katende K1775; MO [MO-2786187] • Lango la Simba; [02°14’00”S, 40°12’00”E]; 12 m; 30 Nov. 2004; Luke 10739; K [K001561015], MO [MO-2758137] • Boni Forest Reserve; [01°43’00”S, 41°14’30”E]; 12 m; 29 Nov. 1988; Robertson 5594; EA, K, MO [MO-4240965], US [US03133206]. – Tana District • Tana Delta; [02°22’48”S, 40°18’00”E]; 5 m; 4 May 2011; Leauthaud 145; EA, K, MO [MO-6854947)].
MOZAMBIQUE – Without locality data • 3 Nov. 1905; Le Testu 863; BM, BR [BR0000016101593], FHO, MO [MO-2198289], P [P04595252, P04595253]. – Cabo Delgado Province • Nangororo; [12°25’45”S, 40°30’28”E]; 24 Nov. 1959; Gomes 4526; BR [BR0000016101579], K [K001561001] • Quirimba; [12°04’00”S, 40°21’00”E]; 60 m; 5 Dec. 2003; Luke 9867; MO [MO-5761467].
TANZANIA – Lindi District • Kilwa Kisiwani Island; [08°57’33”S, 39°29’32”E]; 0 m; 17 Dec. 2003; Abeid 1631; MO [MO-5901562] • Lake Lutamba; [09°49’59”S, 39°47’00”E]; 800 ft; Nov. 1953; Eggeling 6730; K, TFD • Lake Lutamba; [10°01’48”S, 39°27’36”E]; 240 m; 5 Nov. 1984; Mwasumbi 12684; DSM, K, MO [MO-2758149] • Lutambasee; [09°33’36”S, 39°36’00”E]; 240 m; 9 Nov. 1934; Schlieben 5586; FHO [FHO0080387], K [K001561002], WAG [WAG.1778713], MO [MO-1703876]. – Liwale District • Selous Game Reserve; [08°27’24”S, 38°32’32”E]; 125 m; 31 Jul. 1975; Vollesen 2605; DSM, K [K001561003], WAG [WAG.1778714] • Selous Game Reserve; 125 m; 6 Oct. 1975; Vollesen 2752; DSM, K, WAG [WAG.1778715] • Selous Game Reserve; 125 m; 6 Sep. 1976; Vollesen 3979; DSM, K, WAG [WAG.1778717]. – Muheza District • Kigombe; [05°09’19”S, 39°05’46”E]; 0 m; 20 May 1958; Faulkner 2151; K [K001561011].
Diospyros suaheliensis displays some morphological resemblance to D. angolensis, sharing leaves that usually have a rounded apex and generally spherical fruits. These similarities may be linked to the coastal and riverine environments they both occupy. Further investigations into the evolutionary origins of these shared and possibly adaptive traits as well as their environmental correlates are warranted.
We wish to thank the staff and curators of the herbaria that provided loans and access to their collections (BM, FHO, K, MO, P, WAG), Fátima Sales (COI) and Maria Cristina Duarte (LISC) for advice of Gossweiler’s collections, Toral Shah for co-supervising the first author’s M.Sc. project, Roy Gereau for helpful discussions, the Royal Botanic Gardens, Kew for project support, and the Fondation Franklinia for financial support.