Research Article |
Corresponding author: Terry W. Henkel ( terry.henkel@humboldt.edu ) Academic editor: Alfredo Vizzini
© 2019 Bart Buyck, Terry W. Henkel, Valérie Hofstetter.
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:
Buyck B, Henkel TW, Hofstetter V (2019) Epitypification of the Central African Cantharellus densifolius and C. luteopunctatus allows for the recognition of two additional species. MycoKeys 49: 49-72. https://doi.org/10.3897/mycokeys.49.32034
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Cantharellus densifolius and C. luteopunctatus are epitypified on the basis of recently collected specimens from the Central African rain forest that correspond in every way to their respective original descriptions. Sequences obtained from these new collections demonstrate that both epitypes represent distinct species that belong in different subclades of Cantharellus subg. Rubrinus. Previously, the name C. densifolius has been consistently misapplied to more or less similar species from the African woodland area, including C. densilamellatus sp. nov. which is described here, In addition, C. tomentosoides sp. nov., a rain forest species that is easily confused with C. densifolius, is described.
Cantharellales, ectomycorrhizal, tef-1, phylogeny, rain forest, taxonomy
Tropical African Cantharellus species (“chanterelles”) have been well-documented compared to other tropical regions. Nonetheless, there is a great need for sequence data to provide the foundation for unambiguous species concepts. This is due to the highly variable and potentially deceptive macromorphologies, compounded by the limited interspecific micromorphological variation among Cantharellus species (
Despite this need for sequence data, Cantharellus has been difficult to work with from a molecular standpoint. Cantharellus ribosomal genes have high rates of molecular evolution (
While phylogenetic understanding of Cantharellus in Europe and North America has recently improved (
Many of the older species names for African chanterelles have been misapplied to morphologically similar specimens gathered in dense, closed-canopy rain forest versus the surrounding seasonal woodlands, or in open woodlands of neighboring Madagascar (
Here we epitypify Cantharellus densifolius Heinem. based on recent collections made ~400 km from the type locality but in the same forest habitat. The chosen epitype, which is in perfect agreement with the original description, clearly demonstrates that the name has been misapplied to different species for decades. The new collections constitute the first records for C. densifolius since this species was collected by Mme. Goossens-Fontana in 1929 and later described by
Basidiomata were collected in the Central African Republic (RCA) during dry conditions in early May 2016 in pure Gilbertiodendron dewevrei stands of the Dzangha-Sangha Forest Reserve. In Cameroon, basidiomata were collected during the Aug.-Nov. rainy seasons of 2014, 2016, and 2017 from the Dja Biosphere Reserve, Northwest Sector near the village of Somalomo, Upper Dja River Basin, within a two km radius of a base camp located at 3°21'29.8"N, 12°43'46.9"E, 650 m a.s.l., in forests dominated by G. dewevrei. Photographs and descriptions of macromorphological features were made from fresh material in the field. Colors were compared with color plates from
Microscopic observations and measurements were made from ammoniacal Congo red mounts after a short pretreatment in a 10% aqueous KOH solution to improve tissue dissociation and matrix dissolution. Original drawings for all elements of the hymenium and pileipellis were made at a magnification of 2400× with the aid of a camera lucida. Measurements of basidiospores cite length, width and length/width ratio (Q) in this format: (minimum–) mean minus standard deviation – mean value – mean plus standard deviation (− maximum measured); basidiospore size statistics are based on 20 basidiospores measured per specimen.
Genomic DNA isolation, amplification and sequencing for the transcription elongation factor 1-alpha (tef-1) of the new Cantharellus collections were obtained as described in
Seven new sequences were produced for this study (five collections of C. densifolius, one of C. tomentosoides, and one of C. luteopunctatus). The alignment used for phylogenetic analyses included sequences of 90 Cantharellus specimens and one of Craterellus tubaeformis used for outgroup. The full alignment length was 864 base pairs. After exclusion of three spliceosomal introns, the remaining 629 characters were used for the analyses.
The most likely tree (Fig.
Most likely tree obtained by analysis of the 91 tef-1 sequence dataset. Species names are preceded by their extraction number (see
“Pileus carnosulus, infundibuliformis, lobatus, laete ochraceus, squamulosus. Stipes solidus, pileo concolor. Lamellae confertissimae, angustissimae, furcatae, non intervenatae. Caro ochracea, sapore amaro. Sporae breviter ellipsoidae, 5,6–7 × 3,7–4,5 μm.”
DEMOCRATIC REPUBLIC OF THE CONGO. Binga, dispersed on the soil of the dry forest, Aug. 1929, Mme. Goossens-Fontana 879 (BR).
(freely translated from French) “Pileus ca. 8 cm diam., thin, deeply concave to infundibuliform, with the margin convex to stretched, irregular and wavy; surface ochraceous orange, very finely punctuated with tiny squamules that are easily detached. Stipe ca. 30 × 7 mm, cylindrical, massive, concolorous with the cap. Hymenophore composed of crowded gill-folds, less than 1 mm high, 1–4 times forking, deeply decurrent, with blunt gill edge, not interveined. Context fibrous, bright ochraceous. Taste bitter. Smell of C. cibarius. Spore print white. Exsiccatum with reddish ochre brown color.
Spores hyaline, 5.6–7 × 3.7–4.5 μm, shortly ellipsoid, thin-walled, not amyloid; apiculus small. Basidia slender, 37–48 × 6–8 μm, probably 6-spored. Hymenophoral trama pseudoparenchymatic, slightly bilateral. Pseudoparenchyma very compact. Pileipellis squamulae composed of easily detaching cells that are irregularly cylindrical, often undulating, thick-walled with a very thick yellow wall in ammonium solution; the terminal cells obtusely rounded. Clamp connections rare.”
Cantharellus densifolius. a Field habit of the epitype (BB 16.021), showing the areolate-squamose ochraceous orange pileus surface resulting from the concentrical disruption of a dark tomentum that covered initially the young pileus b Detail of the epitype hymenophore showing the remarkably low and thick, crowded, repeatedly forking gill folds without interstitial veination c Original watercolor of the holotype by Mme. Goossens-Fontana from
Cantharellus densifolius (epitype, BB 16.021). Microscopic features: a basidiospores b basidia and basidiola c distinctly thick-walled and typically sinuous-undulate hyphal extremities of the pileipellis d detail of an encrusted hypha from the pileus context. Scale bar: 10 µm but only 5 µm for basidiospores. Drawings B. Buyck.
Basidiomata solitary or in small groups. Pileus medium-sized to rather large and up to 100 mm diam., 1–2 mm thick at mid-radius, yet firm and leathery; margin undulating, irregularly waving to strongly lobed, smooth; surface layer remaining more or less continuous in the center, then disrupting toward the margin with expansion of the pileus and forming dark, more or less concentrically arranged squamules or fibres; observed under a hand lens these can be appressed and flat, or forming a woolly-cottony mass of suberect fibers, pale brown (5AB3) to warm chocolate brown or dark brown (5EF7–8, 5F4–8, 5D5–8, 5C5–6) when young, but rapidly tinged with ochraceous orange as a consequence of the exposure of the underlying pileus tissue and the yellowing tendency of the context. Hymenophore composed of very crowded (>30/mm) gill folds, which are very low (<1 mm) and thick, not interveined, often transversely fissuring over their entire height, repeatedly forking, strongly decurrent, off-white when young, then darkening to the color of coffee with copious milk, moderately to strongly yellowing upon handling. Stipe 40–60 × 4–5 mm, widening toward the hymenophore and there up to 8(–17) mm wide; surface smooth, whitish, pale brown just beneath the hymenophore. Context whitish, thin and leathery, fibrous in the stipe, faintly to strongly yellowing upon injury or handling, occasionally turning rusty brown. Odor faint. Taste mild. Spore print off-white.
Basidiospores ellipsoid, (5.8–)6.0–6.46–6.9(–7.1) × (3.5–)3.8–4.19–4.6(–5.0) µm, Q = (1.3–)1.4–1.55–1.7(–1.8), smooth, hyaline. Basidia mostly 35–50 × 7–8 µm, (5–)6(–7)-spored; sterigmata stout but rather short. Subhymenium forming a very dense layer, not pseudoparenchymatous, but composed of mostly short cells that are not wider than the basidium base. Cystidia none. Pileipellis of loosely interwoven and much septate hyphal extremities composed of ramifying chains of distinctly thick-walled cells; terminal (but also sometimes subapical) cells subcylindrical, but often more irregularly inflated or sinuous-tortuous in outline, 5–8(–10) μm wide, mostly 25–45 µm long, often narrowing or abruptly constricted near the apex. Clamp connections absent.
CENTRAL AFRICAN REPUBLIC. Dzanga-Sangha Forest Reserve, near Bayanga, close to Bai-Hakou base camp, 02.859934N, 16.467492E, in monospecific Gilbertiodendron dewevrei forest, on bare sandy soil, 15 May 2016, 1641/Buyck 16.021 (
CENTRAL AFRICAN REPUBLIC. Dzanga-Sangha Forest Reserve, near Bayanga, in and around Bai-Hakou base camp, 02.859934N, 16.467492E, in monospecific Gilbertiodendron dewevrei forest, on bare sandy soil, 19 May 2016, Buyck 16.081/1656 (PC0142487), Buyck 16.065/1649 (PC0142488); ibid., 24 May 2016, Buyck 16.113/1672 (PC0142490); ibid., 26 May 2016, Buyck 16.137/1681 (PC0142489).
Cantharellus densifolius was originally described by
The epitype specimen selected here perfectly agrees with the original description of C. densifolius. Indeed, Heinemann (l.c.) described it as a medium-sized species with an infundibuliform, ochre-orange and finely squamulose pileus measuring ca. 80 mm diam. and ending in an irregular but stretched margin, with strongly decurrent, crowded, frequently forking and very low gill folds (< 1 mm high) with blunt edges, lacking any intervenation. Heinemann cited shortly ellipsoid basidiospores of near identical size, more precisely given by
The typical features appear to be quite constant across all specimens of C. densifolius examined here, including both the size and shape of basidiospores (Table
The form of the hyphal extremities composing the pileal tomentum is very similar to that of various other squamulose species in subg. Rubrinus sect. Isabellinus Eyssart. & Buyck, in particular those of the African C. tanzanicus Buyck & V. Hofst. (
Cantharellus densifolius has repeatedly been reported from the surrounding woodland area in Africa (e.g.
C. densifolius | |||
Holotype ( |
5.6–7 | 3.7–4.5 µm | |
Holotype ( |
5.5–6.37–7 | 3.5–4.06–4.5 | 1.3–1.57–1.8 |
Epitype: | (5.8–)6.0–6.46–6.9(–7.1) | (3.5–)3.8–4.19–4.6(–5.0) | (1.3–)1.4–1.55–1.7(–1.8) |
Buyck 16.137 | (5.4–)5.5–5.78–6.1(–6.5) | (3.5–)3.9–4.14–1.5(–1.6) | (1.2–)1.3–1.40–1.5(–1.6) |
Buyck 16.081 | (4.8–)5.4–5.78–6.1(–6.2) | (3.5–)3.8–4.02–1.5(–1.7) | (1.2–)1.3–1.44–1.5(–1.6) |
C. tomentosoides | |||
Holotype | (5.8–)6.0–6.36–6.7(–7.1) | (3.9–)4.0–4.27–4.5(–5.0) | (1.3–)1.4–1.49–1.6(–1.7) |
C. densilamellatus | |||
Holotype: | 6.7–7.05–7.4(7.9) | (3.3)3.4–3.65–3.9(4.0) | (1.7)1.8–1.94–2.1(2.3) |
C. luteopunctatus | |||
Holotype ( |
4.9–6.0 (7.5) | 3.8–4.6 (5) μm | |
Holotype ( |
5–5.97–7 | 3.5–4.19–5 | 1.2–1.42–1.6 |
Epitype / Henkel 10285 | (5.4–)5.7–6.04–6.4(–7.1) | (3.9–)4.0–4.29–4.6(–5.0) | (1.2–)1.3–1.41–1.5(–1.8) |
Henkel 10442: | (5.4–)5.4–5.94–6.5(–7.3) | (4.1–)4.2–4.33–4.5(–4.8) | (1.2–)1.3–1.37–1.5 |
Lentinus luteopunctatus Beeli, Bull. Soc Roy Bot Belge 60: 160. 1928.
“Pileo carnoso-coriaceo, centro depresso, margine incurvato, luteo; furfuraceo brunneo, 3,5–4 cm. lato; stipite cylindrico-solido, glabro, concolori, 3 × 0,5–0,7 cm, lamellis deccurentibus, luteis; sporis ovoideis 5–6 × 3,5–4 μm, carne lutea.”
DEMOCRATIC REPUBLIC OF CONGO. Central forest district, near Djongo-Akula, dispersed on the soil of the dry Gilbertiodendron dewevrei forest, Dec. 1925, Mme. Goossens-Fontana 502 (BR).
(freely translated from French) “Pileus rather thick, 49 mm diam., soon depressed, concave with rounded, then straight margin, bright lemon yellow, punctuated – particularly in the center – with minute brownish squamules. Stipe 30 × 6 mm, [30–50 × 5–11 mm], cylindrical, solid inside, yellow, finally rusty, faintly covered from brownish scales. Gills very crowded, deeply decurrent, very narrow, 0.5–11 mm wide (sic!), yellow, irregularly forked, interconnected by rather abundant transversal anastomoses. Context firm, bright yellow, more orangish near the stipe base. Taste strong, bitter. Spore print white. Exsiccatum orangish brown-ochre.
Spores hyaline, shortly ellipsoid, 4.9–6.0 (7.5) × 3.8–4.6 (5) μm, granular inside, thin-walled, not amyloid, with a small apiculus. Basidia narrowly clavate, 30–40 × 6.7–9.5 μm, 4-spored, perhaps sometimes 6-spored. Hymenium not or only slightly accrescent. Subhymenium narrow. Pseudoparenchyma composed of very long and slender elements, mixed with some oleiferous hyphae that do not color in Congo Red. Pileipellis undifferentiated; squamules formed of hyphae united in bundles made up of yellowish to pinkish cells; terminal cells lanceolate or clavate, 6–13 μm diam. Hyphae not amyloid.”
Cantharellus luteopunctatus. a Field habit of the epitype (TH 10285) b details of younger basidiomata from specimen TH 9921, showing the gradual color change of the pileus going from pinkish brown in youngest stages to pale yellow in older stages because of the less dense squamulae; similar squamulae are present on the stipe surface. Composition based on photos by Terry Henkel and Todd Elliott c original watercolor of the holotype by Mme. Goossens-Fontana from
Cantharellus luteopunctatus. Microscopic features: a basidiospores b basidia and basidiola c detail of part of a squamula showing the terminal, thin- to slightly thick-walled hyphal extremities overlying the pileipellis. Scale bar: 10 µm but only 5 µm for basidiospores. Drawings: B. Buyck.
Basidiomata scattered to occasionally caespitose. Pileus up to 65(–75) mm diam., initially broadly convex with shallow depression, extending outward and upward with age, becoming increasingly infundibuliform with downturned margin, deep golden yellow (2–3A4), beset with minute, conical erect tufts, these flesh-brown, more concentrated over the disc but extending and gradually more widely dispersed toward margin; intervening surface shiny-glabrous; margin irregularly crenulate, slightly wavy. Hymenophore composed of very thin, crowded, ridge-like gill folds, creamish to nearly concolorous with the pileus surface (2–3A3), occasionally developing tannish overtones with age (3–4A3), decurrent and fairly abruptly demarcated from the sterile stipe surface, discoloring slowly darker yellow to orangish where injured, repeatedly forking, also abundantly cross-connected between stipe and pileus margin at almost right angles, becoming increasingly tortuous and anastomosed with advanced age; edges even and concolorous. Stipe subequal or slightly tapering toward the base, (17–)24–43 × 4–8(–10) mm, concolorous with pileus, beset apically with conical, flesh-brown, erect tufts, longitudinally striate below; extreme base often developing some white mycelial tissue at the soil interface. Context solid, yellow, unchanging to increasingly yellow. Odor mildly chanterelle-like. Taste mild, nutty, pleasant, somewhat tardily acrid in young specimens. Spore print not obtained.
Basidiospores short-ellipsoid to ellipsoid, (5.4–)5.7–6.04–6.4(–7.1) × (3.9–)4.0–4.29–4.6(–5.0) µm, Q = (1.24)1.29–1.41–1.53(–1.79), smooth. Basidia quite short, mostly 30–40(–50) × 6–7 µm, (5–)6-spored. Cystidia none. Subhymenium cells mostly hardly wider than the basidium base, but locally more inflated parts make it somewhat intermediate between distinctly pseudoparenchymatous and filamentous. Pileipellis composed periclinal thin-walled hyphae of variable diameter, but most ca. 10 µm wide, that locally emit anticlinal tufts of short-septate chains of more or less inflated cells, with the largest cells in these chains distinctly zebroid incrusted and the more terminal cells distinctly thick-walled (up to 1 µm thick); terminal cells 30–60 µm long, mostly (6–)10–15 µm wide, subcylindric or clavulate to lageniform, with obtusely rounded to attenuated tips, never remarkably undulate or irregular in outline. Clamp connections absent.
CAMEROON. East Region: Dja Biosphere Reserve, Northwest Sector near the village of Somalomo, Upper Dja River Basin, within 2 km radius of Dja base camp located at 3°21'29.8"N, 12°43'46.9"E, 650 m a.s.l., 2 km SW of Dja base camp, under Gilbertiodendron dewevrei, coll. T. Henkel, 22 Nov 2016, TH 10285 (
CAMEROON. East Region: Dja Biosphere Reserve, Northwest Sector near the village of Somalomo, Upper Dja River Basin, 1.4 km SW of Dja base camp, under G. dewevrei, coll. T. Henkel, 2 Sep 2014, TH 9921 (
Cantharellus luteopunctatus has long been considered as “uncomfortably close” to C. densifolius.
Close reading reveals some differences between the original descriptions for C. luteopunctatus and C. densifolius. Apart from the difference in pileus color, the second most important difference, also noted by
Our collections also demonstrate a difference in hymenophore color between the species, something that is not evident from Heinemann’s descriptions. Heinemann described C. luteopunctatus as having a yellow hymenophore, but does not indicate the color for the hymenophore of C. densifolius, although the original watercolor clearly shows it to be more or less ochraceous (see Fig.
Other considerable differences between C. luteopunctatus and C. densifolius include the surface structures of the pileus and stipe. In C. luteopunctatus, distinct central pileal squamae are erect, flesh brown to pinkish brown, and strongly separated and paler toward the margin. The pinkish color of the squamae was also mentioned in the original description of
Micromorphologically, the basidiospores are nearly identical in both species (Table
A final remark concerns the edibility of these Central African chanterelles: In Cameroon C. luteopunctatus basidiomata are mild-flavored and consumed by the indigenous Baka, while C. densifolius slowly develops a very strong bitterness and is not consumed by the Baka (T. Henkel, pers. obs.). While the bitter taste was also noted in the original description (
Cantharellus tomentosoides is similar to C. densifolius in its low, blunt and crowded gill folds, overall yellowish brown color, identical basidiospores, and same habitat, but differs in its mostly smaller basidioma size, pileus surface texture, slightly more yellowish olive hymenophore color, and less thick-walled, less sinuous pileipellis extremities.
MG450685 (tef-1).
In reference to the species’ resemblance to its woodland sister-species, C. tomentosus.
CENTRAL AFRICAN REPUBLIC. Dzanga-Sangha Forest Reserve, near Bayanga, close to Bai-Hakou base camp, 02.859934N, 16.467492E, in monospecific Gilbertiodendron dewevrei forest, on bare sandy soil along trail at the entrance of the camp, 14 May 2016, Buyck 16.007 (PC0142485). MycoBank MBT 828890.
Basidiomata in small clusters, up to 40 mm high. Pileus 20–30 mm diam., thin and leathery, wavy with inrolled margin, young entirely hirsute-rugose, remaining lacerate-fibrillose to cottony in the center, elsewhere slightly rugose but lacking well-defined appressed scales, overall pale grayish brown with dark brown center, very early on becoming narrowly but strongly depressed centrally. Stipe slender, 6 mm diam., 20–30 mm high, rapidly elongating while pileus is still small, paler to concolorous with pileus margin, occasionally white at base; interior distinctly fistulose. Hymenophore composed of very crowded (up to 40/cm), low but comparatively thick and blunt gill folds, these 1 mm high, repeatedly forking, frequently fissuring over their full height, yellow with brownish tint, transitioning to warm egg-yolk yellow near extreme margin. Context leathery, whitish in the pileus, almost concolorous with the stipe surface, yellowing slowly. Odor agreeable, typical. Taste mild. Spore print not obtained.
Basidiospores short-ellipsoid to ellipsoid, (5.8–)6.0–6.36–6.7(–7.1) × (3.9–)4.0–4.27–4.5(–5.0) µm, Q = (1.3–)1.4–1.49–1.6(–1.7), smooth, hyaline. Basidia short and narrow, 30–38 × 6–8 µm, mostly five-spored. Subhymenium pseudoparenchymatous, composed of short, barely inflated cells that are slightly wider than the basidium base. Cystidia none. Pileipellis composed of almost thin-walled to slightly refringent hyphal extremities, mostly 4–8 µm wide; terminal cells rather short, mostly 20–40(–50) µm long, subcylindrical, regular in outline, broadly rounded at the apex; walls refringent, not thickened. Clamp connections absent.
Cantharellus tomentosoides is a rain forest species that is phylogenetically sister to C. tomentosus Eyssart. & Buyck (Fig.
Cantharellus tomentosoides resembles C. densifolius in its similarly crowded gill folds, overall yellowish brown coloration and identical basidiospores, but differs in its mostly smaller size, different texture of pileus surface, slightly different color of hymenophore, and thinner-walled hyphal extremities at the pileus surface. Additionally, these two species are phylogenetically distinct (Fig.
Cantharellus densilamellatus resembles C. densifolius in its overall yellowish to orange-brown color, but differs in its thinner and comparatively well-developed gill folds with less blunt edges, smaller size, nearly thin-walled, less undulate hyphal extremities at the pileus surface, more elongate basidiospores, and its seasonal woodland habitat.
JX193014 (published in
“densilamellatus”; referring to the relatively close spacing of the gill folds.
TANZANIA. Msanga village, in very degraded woodland with Brachystegia, 24 April 1998, Buyck 98.013 (PC0084126). MycoBank MBT 828893.
Basidiomata small to medium-sized. Pileus up to 60 mm diam., first centrally depressed and with a downturned margin, then becoming more depressed as the margin spreads out, fleshy in the center, but increasingly thin fleshed toward the margin and there often striate or radially splitting; margin regular to slightly wavy-lobed; surface dry, with a pale to dark brown to reddish brown tomentum (5DEF6–8) contiguous over disc, toward margin tomentum separating concentrically into a pale yellowish cream (3A2–3) areolate pattern. Hymenophore composed of thin, well-developed gill folds, 2–3 mm high, densely spaced (> 10/cm) but not crowded, forking, not anastomosing, often splitting transversely through their entire height, uniformly pale yellow (3A4), brighter than the pileus margin and stipe. Stipe central, up to 40 mm long, 6–11 mm wide, subcylindrical to slightly wider near the base, rapidly elongating before the pileus margin starts to spread, concolorous with the pileus margin, distinctly finely squamulose over apical portion, compact in section. Context off-white to pale cream, weakly yellowing. Odor fruity. Taste mild. Spore print off-white to very pale yellowish.
Basidiospores narrowly ellipsoid to almost elongate, often reniform or peanut shaped, 6.7–7.05–7.4(–7.9) × (3.3–)3.4–3.65–3.9(–4.0) µm, Q = (1.7–)1.8–1.94–2.1(–2.3), smooth. Basidia rather short, 35–50(–58) × 6–7 µm, (4–)5–6-spored; basidioles mostly clavate. Cystidia none. Subhymenium pseudoparenchymatous, composed of irregular, slightly inflated cells. Pileipellis a cutis of interwoven hyphal extremities forming slender chains of subcylindrical cells, these quite regular in outline, with thin to very slightly thickened and refringent walls; terminal cells (25–)30–45(–65) × (3–)4–7 µm, subcylindrical or sometimes very slightly inflated in the lower or middle portion, obtusely rounded at the tip or slightly constricted subapically. Clamp connections absent.
Cantharellus densifolius has long been the only available name for yellowish brown, clampless chanterelles in Africa with a squamulose pileus and crowded gill folds. Indeed, the holotype collection of C. densilamellatus reported here was initially identified as C. densifolius in
Phylogenetic analysis including the newly obtained sequence data demonstrated that C. densifolius and C. luteopunctatus, here epityped, belong in the same subgenus but in different subclades. Additionally, the name C. densifolius has been consistently misapplied to at least one, and possibly several, similar taxa from the African woodland area (De Kesel pers. comm.,
B. Buyck thanks the ATM of the Paris’ Museum and “l’Institut Ecologie et Environnement” (CNRS-INEE) for funding the field trip with Shelly Masi to Africa; Shelly is thanked for all the practical help and sharing her experience. Terence Fuh and the staff of the Primate Habituation Programme of the Dzanga-Ndoki National Park, Réserve Spéciale de Foret Dense de Dzanga-Sangha at Bayanga, as well as all staff, Aka guides and visitors of the Bai Hokou field station for their logistical support, field assistance and their very enjoyable company during our stay. This research was made possible through research permit 034/MENESR/DIRCAB/DGESRSTI/DRSTSPI/SSSTI/16 from the “Ministère de l’éducation nationale, de l’enseignement supérieur et de la recherche scientifique” of the Central African Republic. Funding was provided to T.W. Henkel by National Geographic Society’s Committee for Research and Exploration grant 9235-13 and National Science Foundation grant DEB-1556338. In Cameroon the Ministry of Research and Scientific Innovation issued research permits. The Conservator of the Dja Biosphere Reserve, Mr. Ndinga Hilaire, and his staff greatly assisted the fieldwork in the Dja. Congo Basin Institute staff provided much logistical support. Field assistance in Cameroon was provided by Mei Lin Chin, Todd Elliott, Camille Truong, Bryn Dentinger, Cathie Aime, Rachel Koch, Blaise Jumbam, Olivier Séné, Carolyn Delevich, Kennan Mighell, Jessie Uehling, Noah Siegel, Alamane Gabriel (a.k.a. Sikiro), and Essambe Jean-Pierre (a.k.a. Papa Chef). The first author thanks the Curator of the Herbarium (BR) of the Botanic Garden Meise, Belgium, for permission to reproduce the original watercolors by Mme. Goossens-Fontana.