Research Article |
Corresponding author: Thomas L.P. Couvreur ( thomas.couvreur@ird.fr ) Academic editor: Yasen Mutafchiev
© 2018 Suzanne Mogue Kamga, Raoul Niangadouma, Fred W. Stauffer, Bonaventure Sonké, Thomas L.P. Couvreur.
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:
Kamga SM, Niangadouma R, Stauffer FW, Sonké B, Couvreur TLP (2018) Two new species of Raphia (Palmae/Arecaceae) from Cameroon and Gabon. PhytoKeys 111: 17-30. https://doi.org/10.3897/phytokeys.111.27175
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Raphia (Arecaceae, Calamoideae) is the most diverse genus of African palms with around 20 species. Two new species from Cameroon and Gabon, Raphia gabonica Mogue, Sonké & Couvreur, sp. nov. and Raphia zamiana Mogue, Sonké & Couvreur, sp. nov. are described and illustrated. Their affinities are discussed and the conservation status of each species is assessed. For both species, distribution maps are provided. Raphia gabonica is restricted to two small populations from central Gabon, where it occurs on hillsides on tierra firme soil, and close to small streams. Its preliminary IUCN status is Endangered, being amongst the five most threatened palm species in Africa. Raphia gabonica potentially belongs to the moniliform section. Raphia zamiana is largely distributed from south Cameroon to south Gabon and is very common. It is also a multi-used palm, from which wine, grubs and construction material are extracted and sold. It generally occurs in large stands in a wide range of ecosystems such as swamps, coastal forests on partially inundated sandy soils and inundated savannahs. Its large stature, hard to access habitat (swamps) and abundant presence might have discouraged botanists to collect it until now. Raphia zamiana belongs to the taxonomically complex raphiate section.
Cameroon, Gabon, Raphia , Arecaceae , Calamoideae , IUCN conservation status, new common species
Raphia (Raphiinae, Calamoideae) is the most species-rich genus of African palms with now 22 species currently recognised to date (
The two new species reported here stem from extensive field work in Gabon, Cameroon, Republic of Congo and the Democratic Republic of Congo between September 2015 and February 2018.
Herbarium collections were made following methods described in
Gabon. Moyen Oogoué: 5 km from Alèmbé on national road in the direction to la Lope, 0.07916S, 11.00836E, 228 m a.s.l., 18 November 2015, Mogue K.S. 22 (holotype: WAG; isotypes: LBV, YA).
Raphia gabonica closely resembles R. gentiliana by its solitary stem with curly fibres and the shape of the partial inflorescences. Raphia gabonica differs from R. gentiliana in being a mainly tierra firme species (vs. a swamp species), the shape of its fruits being globose, deltoid or ovoid (vs. ellipsoid) with 11 or 12 rows of scales (vs. 9–11) and a thick mesocarp measuring >8 mm.
Stem 3–7 m tall, 20–30 cm in diameter, solitary; dead leaf sheaths persistent; trunk covered by fibres and old leaf sheaths; fibres formed through disintegration of leaf sheath margins ca. 1–2 mm in diameter, curly, dark brown to grey. Leaves 7 or 8, 8–13 m long in total, horizontal, then arching downwards towards the apex; sheath 80–140 cm long, 15–30 cm wide basally, channelled, smooth, margin fibrous, light brown with black and grey spots; petiole 1.7–4 m long, 7–10 cm in diameter basally, channelled, smooth, brown with dark and grey spots, green at younger stages; rachis 5–7 m long, channelled basally and keeled towards the apex, smooth, no spines on keel, light brown to brown, spotted black and grey; pinnae 170–195 per side, irregularly arranged in 4 planes, arching downwards towards the apex, extreme basal pinnae 55–70 cm long, 1.2–1.3 wide, filiform, middle pinnae 1.10–1.20 m long, 4.5–5 cm wide, linear, apical pinnae 9–23 cm long, 0.6–2.4 cm wide, linear, midrib prominent adaxially, brown spines along pinnae midrib and margins, older pinnae bearing spines more basally, younger ones throughout the pinnae, pinnae adaxial surface dark green, abaxial surface waxy green. Leaves next to inflorescence reduced with split sheaths. Inflorescences 5, pendulous, 1.2–1.8 m long, 9 – 10 cm (young) to 30–35 cm (mature) in diameter at base (including rachillae); light brown to brown; prophyll tubular, bearing 2 keels merging to form a pointed beak; peduncle 20–25 cm long, 7–8 cm in diameter, dorsi-ventrally compressed, smooth dark brown abaxially; penduncular bracts 8–20, tubular with triangular apices, smooth, dark brown abaxially; rachis 1.40–1.60 m long, bearing numerous bracts rarely empty, 50–70 first order rachillae, moniliform in shape, circular, alternating in 2 rows on each side of the rachis, smooth, prophyllar bract at the base of first order rachillae, tubular, bearing 2 keels at the side, smooth; basal first order rachillae 30–40 m long, 1.5–2 cm in diameter excluding rachillae, 25–30 cm including second order rachillae, bud flattened; prophyllar bract at the base bearing 2 keels on both sides, other bracts bearing flowers, rarely empty; second order rachillae ca. 38 per side, basal second order rachillae 15–19 cm long, 0.3–0.5 cm in diameter, middle second order rachillae 9–12 cm long, 0.4–0.5 cm in diameter, apical second order rachillae 2.5–5 cm long, 0.3–0.4 cm in diameter, circular, alternating in 2 rows on each side of first order rachillae, bud 1.5–1.7 cm long, flattened, smooth; middle first order rachillae 27–30 cm long, 1.5–2 cm in diameter excluding rachillae, 20–22 cm including second order rachillae, bud flattened; prophyllar bract at the base bearing 2 keels on both sides, other bracts bearing flowers, rarely empty; second order rachillae ca. 33 per side, basal second order rachillae 14–15 cm long, ca. 0.5 cm in diameter, middle second order rachillae 8.5–9 cm long, ca. 0.5 cm in diameter, apical second order rachillae 2.5–3 cm long, ca. 0.5 cm in diameter, circular, alternating in 2 rows on each side of first order rachillae, bud flattened, smooth; apical first order rachillae: 17–20 cm long, 1–1.5 cm in diameter excluding rachillae, 7–10 cm including second order rachillae, bud flattened; prophyllar bract at the base bearing 2 keels on both sides, other bracts bearing flowers rarely empty; second order rachillae 20 per side, basal second order rachillae 4–7 cm long, 0.2–0.3 cm in diameter, middle second order rachillae 4–5 cm long, 0.2–0.3 cm in diameter, apical second order rachillae 2.5–3.5 cm long, 0.2–0.3 cm in diameter, circular, alternating in 2 rows on each side of first order rachillae, bud flattened, smooth, light brown. Flowers: only very old or very young flowers observed, solitary, exerted, arranged in one row (sometimes two) on each side of second order rachillae, staminate flowers distal, pistillate flowers basal, stamens 6. Fruit: ca. 4 cm long, 3.5 cm in diameter, beak 0.5 cm; globose, deltoid or ovoid; scales arranged in 11 or 12 rows, shallowly furrowed, dark green to brown when young, turning orange-red at maturity; mesocarp ca. 0.8 cm thick, yellow; seeds 1–3, circular covered in a white coating.
Raphia gabonica, illustrations. A Habit (bar = 1 m) B Details of trunk - notice curly fibres C Detail of full inflorescence D Detail of base of inflorescence E Penduncular bracts F Partial inflorescence with old flowers G Detail of basal part of rachillae with old female flower H Detail of apical part of rachillae with old male flowers (Scale bar: 1 cm) I Partial inflorescence with fruits J Fruit K Longitudinal section of fruit. Drawings based on A from Mogue 22 B–J Mogue 23. Drawings by Hans de Vries.
Raphia gabonica occurs in lowland rain forests, growing on hill sides on tierra firme and also near streams. Seen occurring together with Sclerosperma mannii H.Wendl. and Elaeis guineensis A.Chev.
Endangered: EN B2ab(ii): Raphia gabonica is a rare species collected from two close locations in central Gabon (Fig.
No uses have been reported for this species.
Raphia gabonica appears to belong to the moniliform section of
Gabon. Ngounié: Zamata village along national road N1, 1.03044S, 10.51881E, 76 m a.s.l., 19 Nov 2015, Mogue K.S. 23 (LBV, WAG,).
Cameroon: South Region: Vallée du Ntem, Nseng avion forest, Fondation agricole Samuel Menye, 1.5 km from Ma’an in the direction of the Ntem river, 2.34805N, 10.63054E, 513 m a.s.l., 25 February 2018, Mogue K. S. 44 (holotype: WAG; isotypes: K, YA).
Raphia zamiana is morphologically most similar to R. monbuttorum in their clustering habitat and straight fibres covering the trunk. Raphia zamiana differs from R. monbuttorum by the size of its stout and stiff rachillae with apical second order rachillae measuring 1 cm in diameter versus less than 1 cm in R. monbuttorum. The inflorescences of R. zamiana are pendulous even from the early stage of development as opposed to the semi obtuse erect position in R. monbuttorum.
Stem 3–8 m tall, 30–40 cm in diameter, clustering; dead leaf sheaths persistent, trunk hidden in dead leaves and fibres; fibres formed through disintegration of the leaf sheath, ca. 1 cm in diameter, straight with pointed tips, brown to black. Leaves 10–12, 12–21 m long in total, horizontal and then arched downwards towards apex; sheath 90–150 cm long, channelled, smooth, margin fibrous, orange-yellow, spotted with black, white and or grey dots; petiole 4–11 m long, 5–35 cm in diameter towards the base, channelled basally and elliptic apically, smooth, green, spotted with dark and grey; rachis 7–13 m long, 10–11 cm in diameter, elliptic basally and keeled towards the apex, smooth abaxially, spiny adaxially (spines on keel), light green to green; pinnae 147–268 per side, irregularly arranged in 4 planes, arching downwards towards the apex, pinnae adaxial surface green, abaxial surface waxy green; extreme basal pinnae 0.80–1.26 m long, 7–30 mm wide, filiform, middle pinnae 1.50–1.90 m long, 5–9 cm wide, linear, apical pinnae 20–72 cm long, 1.5–3.6 cm wide, linear, midrib prominent adaxially, spines along pinnae midrib and margins, brown to black. Leaves subtending inflorescence reduced (1.20–1.40 m long).
Inflorescences 3 or 4, pendulous, 1.55–2.80 m long in total, 17–44 cm (mature) in diameter at base (including rachillae); young inflorescences light green to purple green, older ones light brown to grey-brown. prophyll 18–20 cm long, 13.7 cm diameter, tubular, bearing 2 keels merging to form a pointed beak; peduncle 26–30 cm long, 10–13.5 cm diameter, dorsi-ventrally compressed, smooth; penduncular bracts several, tubular, with triangular apices, smooth, dark brown abaxially; rachis 1.25–2.60 m long, bearing numerous bracts rarely empty, 50–70 first order rachillae, raphiate shape, dorsi-ventrally compressed, alternating in 2 rows on each side of the rachis, smooth; prophyllar bract found at the base of first order rachillae, tubular, bearing 2 keels at the sides, smooth; basal first order rachillae 0.54–1.05 m long, 4.5–7 cm in diameter excluding rachillae, ca. 14 cm in diameter including rachillae, bud flattened; prophyllar bract bearing 2 keels on both sides, subsequent bracts bearing flowers, rarely empty; second order rachillae 48–65, basal second order rachillae 23–35 cm long, 2–2.5 cm in diameter; middle second order rachillae 15–27 cm long, 2 cm in diameter; apical second order rachillae 9.5–15 cm long, 1.5–1.6 cm in diameter, dorsi-ventrally compressed, alternating in 2 rows on each side of first order rachillae, bud flattened, smooth; middle first order rachillae 39–45 cm long, 3–4 cm in diameter excluding rachillae, 12 cm in diameter including second order rachillae, bud flattened; prophyllar bract at the base bearing 2 keels on both sides, subsequent bracts bearing flowers rarely empty; second order rachillae 32–50, basal second order rachillae 16 cm long, 1.5 cm in diameter, middle second order rachillae 13 cm long, 1.2 cm in diameter; apical second order rachillae ca. 10 cm long, ca. 1 cm in diameter, dorsi-ventrally compressed, alternating in 2 rows on each side of first order rachillae, bud flattened, smooth; apical first order rachillae 25–27 cm long, ca. 2.5 cm in diameter excluding rachillae, 12 cm in diameter including second order rachillae, bud flattened; prophyllar bract at the base bearing 2 keels on both sides; second order rachillae 12–30, basal second order rachillae ca. 10 cm long, ca. 1 cm in diameter; middle second order rachillae ca. 8 cm long, 1 cm in diameter; apical second order rachillae ca. 6 cm long, ca. 1 cm in diameter, dorsi-ventrally compressed, alternating in 2 rows on each side of first order rachillae, bud flattened, smooth; second order rachillae sometimes three times the usual size (more than 25 cm long, 2.5cm wide at the apex). Inflorescence bud ca. 5 cm long, ca. 1.2 cm wide, buds of basal and medial first order rachillae sometimes elongated. Flowers solitary, exerted, inserted in two rows on each side of second order rachillae, staminate flowers distal, pistillate flowers basal. Staminate flower 13–18.5 mm long, 7.5–11.5 mm wide, stalk ca. 1 mm long; subtending bracteole 4.5–13.5 mm long, 7–11.5 mm wide, tubular, bicarinate, margins entire, smooth, with a conspicuous wide apical slit on one side, displaying conspicuous longitudinal veins on the outer side, bracteole completely covering the calyx; calyx 5–11.5 mm long, 5–7 mm wide, fused >2/3 of its length, tubular, bearing 2 or 3 shallow lobes, margins entire to slightly rough, smooth; conspicuous longitudinal veins on outer side; corolla 3, 8.5–15(–20) mm long, 4–6 mm wide, basally connate for 1/3 of their length, oblong, apex slightly blunt to acuminate, margins entire, smooth, stiff, displaying a conspicuous longitudinal venation on the inner side; stamens 11–18, filaments 1–4(–6) mm long, 1–1.7 mm wide, free, basally adnate to the petals for 1–2 mm, cream white to pale pink; anthers 4–8.8 mm long, 1–1.5 mm wide, sagittate-elongate, medifixed, pale yellow; pistillode absent. Pistillate flowers 15–25 mm long, 10–13 mm wide; outer subtending bracteole 12–19 mm long, 10–13 mm wide, tubular, bicarinate, margins entire, with one wide apical slit, smooth; inner subtending bracteole 5–9 mm long, tubular, margins entire, one side longer, smooth, sometimes tearing; calyx 8.5–16 mm long, fused >2/3 of its length, tubular, 3 shallow lobes or the latter rarely absent, margins entire, smooth, with longitudinal veins conspicuous on both sides; corolla 5–8 mm long, fused, 3 lobes with margins serrated, lobes sometimes slightly acuminate, margins entire, smooth, conspicuous longitudinal veins; staminodial ring with 17–19 fused staminodes, 2–5 mm long, adnate to petals for 1–4 mm; anthers sagittate, 0.5–1 mm long; gynoecium 11–18 mm long, 5–6 mm wide, ovary 9–12 mm, 5–6 mm wide, ovate to oblong, completely covered with scales, developing at ¾ height of the gynoecium, larger scales at mid portion to base; style absent or very short; stigma ca. 1 mm long, papillae not observed but hair-like prolongations present on stigma. Fruits 4–8.7 cm long, 3.5–4.7 cm wide, beak 0.5–0.9 cm long; oblong, scales arranged in 11 or 12 rows, length of scale 16–20 mm, width of scale 15–20 mm, diamond shaped, apex texture rough, shallowly furrowed, green, beak pointed, inflated in the middle; mesocarp yellow when young, orange yellow when mature; seed 1, oblong, with ruminations.
Raphia zamiana, illustrations. A Habit in savannah B Habitat in forest, trunk not visible C Detail of the trunk, with inflorescences and straight fibres D Inflorescence E Partial inflorescence, young F Detail of rachillae G Detail of basal part of 2nd order rachillae, showing small rachillae bracts encircling young flowers H Detail of partial inflorescence (×5) I Female flower ×5 J Female inner bract ×5 K Female calyx ×5 L Female corolla ×5 M Detail of staminodial ring and staminodes N Male flower ×5 O Male calyx ×5 P Male corolla ×5 Q male stamens ×6 R Detail of stamen of male flower ×12 S Infructescence T Fruit, longitudinal section. Drawings based on: A Couvreur 1122 B–D, S, T Mogue; 17 E–R Mogue 44. Drawings by Hans de Vries.
Raphia zamiana. A Habitat along the road, with Raoul Niangadouma for scale (Oyem, Gabon) B Close-up of pendulous and “raphiate” type partial inflorescence (coastal forests near Kribi, Cameroon) C Habitat in the savannahs of Lopé National Park, Gabon D Habitat in swamp (Ma’an, Cameroon). Photos: Thomas L.P. Couvreur.
Raphia zamiana is an Atlantic central African rain forest species, endemic to lower Guinea. It appears to have a wide ecological amplitude, growing in rain forest swamps on periodically inundated soils, in coastal forests on inundated sandy soils or in open vegetation like savannahs associated with inundated soils (e.g. Lope National Park, Gabon, Figs
Lower Guinea subcentre of endemism (
Least Concern. The extent of occurrence of Raphia zamiana is estimated to be 128,243,063 km2 (far exceeding the 20,000 km2 upper limit for Vulnerable status under criterion B1) and the area of occupancy is estimated to be 36,000 km2 (far exceeding the 2,000 km2 upper limit for Vulnerable status under criterion B2). Moreover, this species is known from 9 locations, the upper limit for sub criterion a- of criterion B2 for the Vulnerable status. However, Raphia zamianais a widespread and common species in its area of occurrence. It is very common along road sides and is abundant growing in dense colonies. To date, it has not been collected within a protected area, but populations were seen in Campo National Park (Cameroon). Incredibly, its first collection dates to 2012. Botanists might have confused it with the well-known and widespread species Raphia hookeri (although both species are very different in their morphology). Given that collecting Raphias is a hard task because of their massive stature (
The name of this species is derived from its vernacular name in Beti (south Cameroon, north Gabon): Zam.
This palm is massive thus providing large amounts of thatching material. Its petiole and rachis commonly referred to as ‘bamboo’ are used for house construction, beds, chairs, baskets and mats. These are generally sold along the roads in south Cameroon. In Gabon, its fruits are sold in markets. These are boiled and said to cure hypertension and diabetes. Finally, this species is also used for wine tapping and as a source of grubs.
Zam (Beti).
Raphia zamiana belongs to the raphiate section (
Raphia zamiana is a very conspicuous palm in southern Cameroon and eastern Gabon, being common along roads and in swamps. In addition, we report several important uses, being one of the most useful Raphias (Mogue, personal observation). However, up to now, it remained uncollected, stressing once again that new species well known to local people have yet to be scientifically discovered and described. A similar situation was recorded for the Vietnamese endemic palm Licuala centralis (
Cameroon. Central Region. near Ebolbom village, 3 km east of Ngoumou, 2 km northwest of Otélé, 3.599720N, 11.287700E, 700 m a.s.l., 2 May 2013, Couvreur T.L.P. 427 (WAG, YA). South Region: Mvila, Biyeyem, 2.514020N, 11.081930E, 573 m a.s.l., 19 Sep 2015, Mogue K.S. 15, (WAG, YA); Mvila, Biyeyem, 2.514020N, 11.081930E, 573 m a.s.l., 19 Sep 2015, Mogue K.S. 16 (WAG, YA); mountain chain Ngovoyang, 1.5 km in forest from Bikiliki village situated between Bipindi and Lolodorf, 3.181570N, 10.536960E, 460 m a.s.l., 17 Feb 2012, Couvreur, T.L.P.392 (WAG, YA); About 20 km N from Kribi, 3 km N of Longji, N7 road towards Edea, 3.146810N, 9.959510E, 0 m a.s.l., 27 Feb 2018, Mogue K.S. 45 (WAG, YA).
Gabon. Woleu-Ntem: Oyem, 2–3 km from main road in the direction of Konosoville, 01.59849N, 011.62298E, 651 m a.s.l., 12 Nov 2015, Mogue K.S. 17 (LBV, WAG, YA); Ogooué-Ivindo: 180 km on main road from Lastoursville to Lopé, 0.147300S, 11.726011E, 280 m a.s.l., 9 Jun 2016, Couvreur T.L.P. 1122 (LBV,WAG, YA). Ogooué-Maritime: Lagune de Fernan Vaz. Koundakoua, 1.4487220S, 9.2066110E, 3 m a.s.l., 20 Nov 2016, Bidault E. 2722 (BR, BRLU, LBV, MO, P, WAG). Ngounié: Mouila, 19 km from national road, 2.254280S, 11.142840E, 133 m a.s.l., 20 Nov 2015, Mogue K.S. 24 (LBV, WAG, YA).
We are very grateful to Richard Oslisly and his keen naturalist eye for providing detailed locality information about a “weird tierra firme Raphia” near Lope which led to the discovery of Raphia gabonica. We are grateful to the Centre National de la Recherche Scientifique et Technique (CENAREST) in Gabon and the Agence National des Parques Nationaux (ANPN) for delivering the necessary research permits for our field work in Gabon in 2015 and 2016 (permits AR0020/16; AR0036/15 (CENAREST) and AE16014 (ANPN)). Prof. Paul Henri Bourobou Bourobou is thanked for his assistance in obtaining these research permits. Fieldwork in Cameroon was undertaken under the “accord cadre de cooperation” between the IRD and Ministère de la Recherche Scientifique et Technique (MINRESI). This study was supported by Agropolis Fondation (RAPHIA project) under the reference ID 1403-026 through the « Investissements d’avenir » program (Labex Agro:ANR-10-LABX-0001-01), under the frame of I-SITE MUSE (ANR-16-IDEX-0006) and the Agence Nationale de la Recherche (grant number ANR-15-CE02-0002-01), both to TLPC; and an International Palm Society (IPS) grant to SMK. We are grateful to Larry Noblick for commenting on an earlier version. We thank Hans de Vries for his high quality drawings. Finally, we are grateful to Jean Grégoire (aka Petit Jean) Kayoum for his driving skills and help in the field when in Gabon and Narcisse Kandem for field assistance in Cameroon.