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Boletín de la Sociedad Argentina de Botánica

versión On-line ISSN 1851-2372

Bol. Soc. Argent. Bot. vol.49 no.1 Córdoba mar. 2014



Gilbertiodendron grandistipulatum (Leguminosae-Caesalpinioideae), a singular species from West Central Africa and new record for Congo (Brazzaville)


Manuel De La Estrella1, 2* and Juan Antonio Devesa2

1Institut de recherche en biologie végétale and Département de Sciences biologiques, Université de Montréal, 4101 Sherbrooke est, Montréal, Québec, H1X 2B2, Canada.
2Departamento de Botánica, Ecología y Fisiología Vegetal, Universidad de Córdoba, Facultad de Ciencias, Campus de Rabanales, Edificio José Celestino Mutis, 14071 Córdoba, España.


Gilbertiodendron (Leguminosae-Caesalpinioideae) is an endemic tropical African genus of ca. 30 species, most of which are trees of primary forest. The highest concentration of species and morphological variation is found in the Guineo-Congolian region, particularly in Gabon. One of those species is G. grandistipulatum, which is easily recognized by its stipules (up to 18 cm long) and flowers (adaxial petal up to 12.5 × 14 cm). The taxonomy, habitat and distribution of G. grandistipultaum are discussed, extending its known distribution to Congo. A full description, distribution map and two illustrations are provided for the first time for this species.

Key words: Congo; Detarieae; Gabon; Taxonomy; Tropical forest.

Resumen: Gilbertiodendron grandistipulatum (Leguminosae-Caesalpinioideae), una especie singular del centro-oeste de África y nuevo registro para Congo (Brazzaville). Gilbertiodendron (Leguminosae- Caesalpinioideae) es un género endémico de África con ca. 30 especies, las cuales en su mayor parte son árboles que crecen en bosques primarios. El mayor número de especies y la mayor variabilidad morfológica se encuentra en la región Guineo-Congoleña, y el particular en Gabón. Una de esas especies es G. grandistipulatum, que se reconoce fácilmente por sus estípulas (de hasta 18 cm de longitud) y flores (con pétalos adaxiales de hasta 12,5 × 14 cm). En el presente trabajo se discute la taxonomía, hábitat y distribución de G. grandistipulatum, extendiendo su área de distribución a Congo. Se proporciona una descripción completa, un mapa de distribución y dos iconografías por primera vez para esta especie.

Palabras clave: Bosque tropical; Congo; Detarieae; Gabón; Taxonomía.



Gilbertiodendron is an endemic tropical African genus of ca. 30 species, most of which are trees of primary forest (Léonard, 1957; Mackinder, 2005; Estrella et al., 2012a). Species belonging to this genus have leaflets with marginal or submarginal glands, a pair of bracteoles that enclose the flower bud, five reduced sepals, five petals, lateral and abaxial petals that are much reduced, alternate to sepals, and similar to them; and a single welldeveloped adaxial petal, the base unguiculate and apex bilobed. The androecium is usually formed by six staminodes and three stamens that are fused with the staminodes to a short fleshy tube, although there are few exceptions, such as G. splendidum, that have nine well-developed stamens. The ovary, usually located on a small stipe, develops into a pod with longitudinal and transverse nerves, and the valves twist when mature to disperse the seeds (Polhill & Raven, 1981; Tucker, 2002; Estrella et al., 2012a).
The highest concentration of species and morphological variation is found in the Guineo- Congolian region, particularly in Gabon which is a high diversity area for Legumes (Estrella et al., 2012b) and where at least 18 species have been previously reported with 10 species considered endemic or subendemic (Sosef et al., 2006). Several new species are to be described during the ongoing taxonomic revision of the genus. In 1775 Vouapa, a new genus of Caesalpinioideae from the French Guyana was published (Aublet, 1775) afterwards it was rejected against the nomina conservanda Macrolobium (Schreber, 1789; Hutchinson, 1964). New Macrolobium species growing both in tropical America and Africa were described until Palisot de Beauvois (1806) published Anthonotha, a closely related new genus of African trees. Baillon (1865) considered that Anthonotha should be included within Macrolobium and the species were treated together again. In 1949 Louis in his Essences forestières et bois du Congo(Louis & Fouarge, 1949) wondered whether the Macrolobium species growing in tropical Africa and tropical America where the same or belonged to different entities.
All species from tropical America stay in the genus Macrolobium, but in a succession of works Léonard reinstalled Anthonotha (Léonard, 1955) and published three new genera to accommodate the African species: Gilbertiodendron (Léonard, 1952a), Paramacrolobium (Léonard, 1954) and Pellegriniodendron (Léonard, 1955).
Anthonotha and two related genera Englerodendron and Isomacrolobium, have been recently revised by Breteler (2006, 2008, 2010 and 2011). Paramacrolobium is a genus with only one species (Léonard, 1957). By contrast Pellegriniodendron (one species) has been transferred to Gilbertiodendron according with recently published phylogenies (Bruneau et al., 2008) and with a morphological re-evaluation of its status (Estrella et al., 2012a).
is a genus that is important both economically (Burkill, 1995), with its species used for timber or traditional medicine, and ecologically, with many species reported as growing gregariously (Burgt et al., 2012; Estrella et al., 2012c; Estrella & Devesa, 2014).
The lack of adequate material has always been a problem faced by those wishing to study this genus (Léonard, 1957: 236). Gilbertiodendron grandistipulatum is probably the species within the genus that presents more spectacular flowers, with an adaxial petal up to 12.5 × 14 cm. The aim of this work is to provide a full comprehensive description using the modern collections available, including a distribution map and a detailed black and white line drawing and colour art illustration of the species.

Material and Methods

This taxonomic treatment of G. grandistipulatum is based on eighteen specimens, which represents all the available specimens after consulting the following herbaria (acronyms following Index Herbariorum): A, AAU, B, BM, BR, C, COI, E, FBC, FHO, G, GH, H, HBG, K, L, LD, LISC, M, MA, MO, NY, P, PR, PRE, S, SL, U, UPS, US, WAG, WU, and Z. Seventy quantitative characters were recorded and measured using a Mitutotyo CD-15CD digital calliper (Tokyo, Japan) for all the specimens when possible. Measurements follow the common practice in morphological studies (Estrella et al., 2009). Each character was analyzed for its mean, median, range, standard deviation, and significance using the STATISTICA package ( Indumentum terminology and species descriptions follow a standard format used by Wieringa (1999) adapted for other Caesalpinioideae genera (Estrella et al., 2010). Label data from each specimen was recorded in a database from which reports and outputs were generated to produce a list of numbered collections, the representative material studied, and a file with coordinates used to produce the distribution map. Information on the uses and vernacular names is included. We also include available references for further information on these topics.

Results and Discussion

Gilbertiodendron grandistipulatum (De Wild.) J. Léonard in Bulletin du Jardin Botanique de l'État 22: 191 (1952). Macrolobium grandistipulatum De Wild. in Repertorium Specierum Novarum Regni Vegetabilis 13: 370 (1914).

Type: Congo (Kinshasa), Ganda-Sundi, 1913, De Briey 65 (lectotype designated by Léonard 1957: 237, BR!). (Figs. 1 and 2).

Fig. 1. Gilbertiodendron grandistipulatum. a: Leaf and inflorescence. b: Gland on edge of leaflet. c: Pair of stipules. d: Flower without adaxial petal. e: Staminodial tube with staminodes and base of stamens, exposed by removal of ovary. f: Bracteole outside. g: Detail of bracteole indumentum. h: Bracteole inside. i: Large, adaxial petal. j: Stamen. k: Anther ventral view. l: Anther dorsal view with indumentum. m: Gynoecium. n and o: Stigma detail. p and q: Juvenile fruit, with close-up of indumentum. r: Seed. s: Seed transversal view. Drawn from: L.J.G. van der Maesen & al. 5734, a-b, p-q; Wieringa & al. 4687, c-i, m; P. Bissiengou & al. 604, j-l; F.J. Breteler & al. 10428, n-o, r-s. Drawing by Juan L. Castillo.

Fig. 2. Gilbertiodendron grandistipulatum. Drawing by Juan L. Castillo.

Macrolobium le-testui Pellegr. in Bulletin de la Société Botanique de France 69: 745 (1922). Type: Gabon, Ngounié, Pays Itsogho, Moundou, 4 Nov. 1916, Le Testu 2177 (holotype P!; isotypes, BR!, WAG!).  Tree, to ca. 25 m tall and 50 cm dbh. Twigs and branches glabrous to pubescent. Stipules in pairs, covered with hairs -mainly at base and margins- to glabrous, with parallel nerves, 75-180 × 19-30(- 43) mm, oblong-lanceolate, persistent, completely fused. Leaves alternate, 6-7(-8)-jugate, (25-)36-85 × 19-30(-43) cm, largest pair of leaflets is the upper or previous pair; petiole 6-17 mm long, 5-11.5 mm diameter, ± terete, densely covered with hairs to glabrous; rachis (12.5-)24-38(-47) cm long, rounded abaxially, canaliculate and slightly articulated at the insertion of each pair of leaflets adaxially, pubescent to glabrous; stipels absent; petiolules 2.5-7.5 mm long; leaflets opposite, coriaceous, 12-29 pairs of main lateral veins, brochidodromous, pubescent to glabrous at petiolules and midrib on leaflet blade in the abaxial surface; 0-3 glands along each margin of the leaflet, and a gland at or near the midrib top; midrib prominent on the abaxial face, rounded, slightly canaliculate in adaxial face; base of the leaflet asymmetric, proximal half of the leaflet lamina wider at base than distal half, proximal margin inserted at 2-4 mm below the distal margin insertion; basal leaflet 9.5-19(-23) × 3-5.5(-7.5) cm, oblong, apex acuminated (up to 3 mm long), terminating in a gland at the top of the midrib; largest leaflet 12-31(-38) × 4-9(-11.5) cm, oblong-obovate, apex emarginate to slightly acuminate (up to 7 mm long), terminating in a gland at the top of the midrib. Inflorescence a raceme or panicle, 24-40 cm long, axillary, sessile or slightly pedunculated; rachis densely covered with hairs ca. 0.1 mm long, with sparse hairs up to 0.5 mm long intermixed, 4-6 lateral branches, each 130-180 mm long and 11-30-flowered. Bracts 18-19 × 9-10 mm, caducous, foliaceous, ovate-lanceolate, cup-shaped in the middle, densely hairy on outer surface, inner surface glabrous with hairs located at the margin; pedicel 19-32(-47) mm long, 2.5-4.5 mm diameter, with 4 ridges when dehydrated, densely covered with short and longer hairs; bracteoles red to brown outside, inside more pinkish-white, 33-44 × 18-26 mm, caducous, oblong, cupuliform concave and slightly beaked in outline, outside densely covered with short and longer hairs, inside glabrous, edge with a ciliated margin, with a beak 2.3-4 mm long culminating in a gland at apex. Receptacle white, 3.5-6 × 5-8 mm, slightly hairy at the margins to glabrous. Sepals 5, pink red with white margin, 17-21 × 3-6 mm, triangular, broadest at base, glabrous, adaxial sepals united for 11-17.5 mm covering part of the adaxial petal claw; the other 3 sepals free. Petals 5, alternate to sepals; adaxial petal large, white to cream with pink to red nerves, deeply bilobed, 100-125 × 105-140 mm, including claw 18-28 × 3.5-6 mm, glabrous except the external face of the claw -near the petal blade- which is pubescent to glabrescent; lateral and abaxial petal pairs pinkish white, almost equal in size to sepals, 13-27 × 3-7 mm, triangular, broadest at base, glabrous. Fertile stamens 3, alternate with abaxial petals, filaments 49-62 mm long, long hairy at lower 3/4; anthers pinkish-white, 12-14 × 4-6 mm, glabrous to slightly hairy at the point of filament insertion on external face; staminodes 6, filiform, fused into a short intrastaminal tube, 3-8 mm long, glabrous. Ovary 13- 21 mm long, rectangular-rhombic in shape, densely hispid, stipe 1-2 mm long, hispid; style 35-55 mm long, white, lower third hairy; stigma 1-1.2 mm in diameter, capitate. Pod 20-30 × 8-9 cm (measure based in literature records), oblong, up to 7-seeded, broadly beaked, sparsely hairy, 2 longitudinal veins. Seeds suborbicular, wrinkled, brown-black, ca. 35 × ca. 25 × ca. 7.5 mm.

Distribution. Species found in Gabon, Congo (Brazzaville), and Congo (Kinshasa). The species has been previously reported from Gabon and Congo (Kinshasa) (Lock, 1989: 12;, V-2013), and although the Congo (Brazaville) specimen recollected by J. Koechlin 7863 (P) was dated in 1957 the species has not been previously reported from the country (Fig. 3).

Fig. 3. Known distribution of Gilbertiodendron grandistipulatum (circles).

Habitat. Primary and secondary forest, river banks and swampy areas; 40-650 m.

Phenology. Flowering specimens are recorded from October to December; the only fruiting specimen is the type but unfortunately we don't have the information on which month it was collected.

Vernacular Names. Gabon: Pokussa des marais (Pellegrin, 1949: 59). Congo (Kinshasa): M'Pokussa, Possa (Léonard, 1952b: 435).

Uses. Wood quite dense and it could be used in joinery (Pellegrin, 1949: 59; Léonard, 1952b: 435).

Notes. Gilbertiodendron grandistipulatum is here illustrated for the first time and the species is easily recognized by its stipules (up to 18 cm long) and flowers (adaxial petal up to 12.5 × 14 cm). Although G. grandiflorum, a species growing in West Central Africa, also presents large adaxial petals (up to 9 × 11 cm) the stipules are smaller (2-2.5 cm long) and the leaves only has 2 pairs of leaflets instead of 6-7(-8) in G. grandistipulatum. Macrolobium grandistipulatum, in which the former combination is based, was described from material collected by De Brieyi in Congo (Kinshasa) by De Wildeman (1914: 370), who referred it to a single collection: "Belgisch-Kongo: Ganda-Sundi, 1913 (Comte de Briey, no. 65)". Because this author does not indicate this specimen as nor indicate where it was deposited it is not clear the holotype. Léonard (1957: 237) selected De Briey 65 (BR) as the lectotype of M. grandistipulatum. Eight years later Pellegrin (1922: 745) described Macrolobium le-testui and he stated: "Pays Itsogho. Moundou (Icobé), le 4 novembre 1916 (Le Testu, nº 2177)". Since Pellegrin worked in P, the original sheet at P, with the more detailed label data by G. Le Testu, constituted the holotype of this species. After the study of the type material we conclude, as other authors did previously, that this name should be placed as a synonym of G. grandistipulatum.

Additional specimensstudied. CONGO (BRAZZAVILLE): Mabombo, (droite du Niari), Madingou, J. Koechlin 7863 (P). CONGO (KINSHASA): Ganda-Sundi, 4º52'S, 12º52'E, De Briey 65 (BR); Ganda-Sundi, 4º52'S, 12º52'E, Gilbert 612 (BR). GABON: Ngounié, 37 km E. of Mouila, along the road to Yeno, 1º40'S, 11º20'E, J.C. Arends, J.J.F.E. de Wilde & A.M. Louis 480 (WAG); Ngounié, village de Ndjemba, entré vers chantier EFT (Exploitation Forestière de Tsanba), route Fougamou-Lambaréné, 1º03'S, 10º30'E, P. Bissiengou, F.J. Breteler, R. Niangadouma & J.-N. Boussiengui 604 (WAG); Moyen- Ogooué,Missanga, 5-15 km NW of Ndjolé, 0º5'S, 10º45'E, F.J. Breteler & C.C.H. Jongkind, J.J. Dibata 10428 (WAG); Estuaire, 5 km au NW de Mebba, 0º28'N, 9º45'E, J.J. Floret & A.M. Louis 1455 (P); Ogooué-Ivindo km 18, piste d'le Invindo vers Lastoursville, 0º12'S, 12º20'E, J.J. Floret, A.M. Louis & A. Moungazi 1910 (P); Ngounié Moundou (Icobé), 1º15'S, 11º7'E, M.G. Le Testu 2177 (BR, P, WAG), Ngounié, region de Mouila, Mitingo (Wano), 1º58'S, 11º30'E, M.G. Le Testu 6376 (BM, BR, MA, P, WAG), Ogooué-Lolo, Lastoursville, 0º50'S, 12º40'E, M.G. Le Testu 7570 (BM, BR, P), Woleu-Ntem, Afaghé Nzoc, 0º40'N, 11º7'E, M.G. Le Testu 9368 (BM, BR, P); Estuaire, Near Mvoum, 18 km NE of Ntoum, 0º30'N, 9º50'E, A.M.Louis, F.J. Breteler, J. de Bruijn 289 (WAG); Ogooué-Lolo, km 13 on Chantier 19, SBL, branching off from km 21 Lastoursville to Koulamoutou, 0º56'S, 12º35'E, L.J.G. van der Maesen & A.M. Louis, J. de Bruijn 5734 (BR, WAG); Ogooué-Ivindo, Lope Reserve, L.J.T. White 1118 (PRE, WAG), 1227 (WAG); Ngounié, road Mouila to Yombi, 1º43'S, 10º47'E, J.J. Wieringa, T. Nzabi, J.-N. Boussiengui 4687 (WAG).


The authors wish to thank the staff of the cited herbaria for their support of our visit and/or loan of material. Frans Breteler kindly provides us photographs of G. grandistipulatum, and we thank Juan L. Castillo for the illustrations. This work was financed by the Spanish Science and Innovation Ministry (CGL2008-02982-C03-03/ CLI, CGL2011-28613-C03-02 and CGL2009- 07405/BOS). Manuel de la Estrella was funded by a Juan de la Cierva grant (JCI-2009-05243) and a NSERC Canada grant to Anne Bruneau, and visited K, P, and WAG herbaria under the FP7 Europeanfunded Integrated Infrastructure Initiative grant SYNTHESYS, GB-TAF-1006, FR-TAF 2194, and NL-TAF 2377 projects, respectively, and he thanks Anne Bruneau for her support.


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Recibido el 29 de julio de 2013,
aceptado el 25 de noviembre de 2013.

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