Academia.eduAcademia.edu
Vascular flora inventory and plant diversity of the Ruvubu National Park, Burundi Tatien MASHARABU Marie Josée BIGENDAKO Université du Burundi, Faculté des Sciences, Département de Biologie, B.P. 2700 Bujumbura (Burundi) tmashara@ulb.ac.be Benoît NZIGIDAHERA Institut national pour l’Environnement et la Conservation de la Nature (INECN), B.P. 2757 Bujumbura (Burundi) Balthazar MPAWENAYO Université du Burundi, Faculté des Sciences, Département de Biologie, B.P. 2700 Bujumbura (Burundi) Jean LEJOLY Université Libre de Bruxelles, Laboratoire d’écologie végétale et Biogéochimie, case postale 244, boulevard du Triomphe, B-1050 Bruxelles (Belgium) Frédéric BANGIRINAMA école normale supérieure, Département des Sciences Naturelles, B.P. 6983 Bujumbura (Burundi) Jan BOGAERT Université de Liège/Gembloux Agro-Bio Tech., Unité Biodiversité et Paysage, 2 passage des Déportés, B-5030 Gembloux (Belgium) Masharabu T., Bigendako M. J., Nzigidahera B., Mpawenayo B., Lejoly J., Bangirinama F. & Bogaert J. 2012. — Vascular flora inventory and plant diversity of the Ruvubu National Park, Burundi. Adansonia, sér. 3, 34 (1): 155-162. http://dx.doi.org/10.5252/a2012n1a17 ADANSONIA, sér. 3 • 2012 • 34 (1) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.adansonia.com 155 Masharabu T. et al. Key woRdS Akagera, Burundi, floristic inventory, plant richness, Ruvubu. ABSTRACT The Ruvubu National Park, the biggest protected area and biodiversity refuge of the country, is comparatively less studied than western Burundi near Bujum­ bura, the capital. This article reports the results of a botanical inventory of the vascular plants from the protected area, evidences species newly encountered and establishes a comparison of the floristic diversity with the Akagera National Park in Rwanda located in the same phytochorion, in the Lake Victoria regional mosaic. Records from the Ruvubu include 522 species (including infraspecific groups) from 96 families and 306 genera, representing nearly 17% of the known vascular plants of Burundi. The most common families were the Fabaceae, the Asteraceae and the Poaceae. 96 species or 18% of the flora were encountered for the first time in the park. Study of environmental determinants of Leguminosae abundance and floristic variability would provide more precisions. MoTS CléS Akagera, Burundi, inventaire floristique, richesse floristique, Ruvubu. RéSUMé Inventaire de la flore vasculaire et diversité floristique du Parc national de la Ruvubu, Burundi Comparé au Burundi occidental proche de Bujumbura, la capitale, le Parc national de la Ruvubu, la plus grande aire protégée, refuge de la biodiversité du pays, a été relativement peu étudiée. Cette étude rapporte les résultats d’un inventaire botanique des plantes vasculaires de l’aire protégée, met en relief les espèces nouvellement signalées dans le parc et établit une comparaison de la diversité floristique avec le Parc national de l’Akagera au Rwanda, localisé dans la même phytochorie, dans la mosaïque régionale du lac Victoria. La richesse spécifique de la Ruvubu fait état de 522 espèces (incluant les groupes infra­ spécifiques) réparties en 96 familles et 306 genres représentant près de 17 % des plantes vasculaires connues du Burundi. Les familles les plus nombreuses sont les Fabaceae, les Asteraceae et les Poaceae. 96 espèces (18 %) de la flore sont signalées pour la première fois dans le parc. L’étude des déterminants envi­ ronnementaux de l’abondance des légumineuses et de la variabilité floristique pourrait fournir de plus amples précisions. INTRODUCTION Many studies (Lewalle 1972; Ndabaneze 1983; Bigendako 1989; Bizuru 2005) have reported that Burundi shelters several plant species on a reduced size. It is quite remarkable that western Burundi, not far from the capital Bujumbura, was more studied in relation to other remaining parts of the country. According to Bizuru (2005), areas corresponding currently to national parks and different other pro­ tected areas of Burundi have been until now very little studied. On the whole protected sites of East Africa, the Ruvubu National Park (RNP), the biggest protected area of Burundi with 50 800 ha, occupies a particular position. Although it does not shelter probably any species that does not exist elsewhere, 156 it has an interest that goes over burundian borders since it is about a rare evidence of the transition between dense forest and humid savannas. It rep­ resents the unique protected sample of the humid savanna ecosystem with Parinari-Pericopsis (Vande Weghe & Kabayanda 1992). This study carries on a botanical inventory conducted in the RNP, sanctuary and shelter of large mammals disappeared elsewhere in the country. Nzigidahera (2000) underlined indeed that the vegetation of the RNP remains little known in its whole. Masharabu et al. (2008) observe that little study have been conducted and data related to its floristic composition are lacking either insufficient. In a context of good manage­ ment, planning and biodiversity conservation, it is important to keep a better knowledge of its flora ADANSONIA, sér. 3 • 2012 • 34 (1) and that requires proceeding to a plant resource assessment. Therefore, in order to reinforce a deci­ sion making of animal communities conservation and sustainable management of the protected area system, it is important to get a better knowledge of the botanical diversity (Lubini 1982; Laurance et al. 2006). The entire species list for the RNP has been published by Masharabu et al. (2010). The present paper provides an inventory of the RNP vascular flora (Pteridophytes and Phanerophytes) and highlights species newly encountered in the park. This paper is the first to provide an inventory of plant species as complete as possible in a given protected site of Burundi. The study hypothesis was that plant diversity in RNP and Akagera National Park (ANP) in Rwanda would be similar due to their biogeographical location at the crossroads of several biogeographical influences in the Lake Victoria re­ gional mosaic. This region is the meeting place of five regional floras: Guineo­Congolian, Sudanian, Zambezian, Somalia­Masai and Afromontane. Its vegetation is a mosaic of floristically impoverished variants of the first four plus some communities which are transitional between Afromontane and lowland vegetation (White 1993). MATERIAL AND METHODS This study was carried out in the RNP using a botanical inventory in 114 vegetation plots vary­ ing between 100 and 900 m2. The inventory was conducted from December 2005 until June 2009, intermittently. The park is located in the north­ eastern Burundi (Fig. 1) between latitudes 2°54’­ 3°22’S and longitudes 30°6’­30°33’E, and covers an area of 50 800 ha. The altitude ranges from 1350 to 1836 m (Vande Weghe & Kabayanda 1992). According to Köppen classification, it has a tropical climate classified as (AW3)s. The main vegetation features are made of savannas, riparian forests and swamps. Duplicates of plant material were deposited at the Herbarium of the Université Libre de Bruxelles (BRLU), as well as at the Her­ barium of the University of Burundi (BJA). The systematic determination and verification were conducted at BJA and BRLU Herbaria as well as at the Herbarium of the National Botanic Garden of Belgium (BR) at Meise by comparison of herbarium ADANSONIA, sér. 3 • 2012 • 34 (1) DEMOCRATIC REPUBLIC OF CONGO Vascular flora inventory and plant diversity of the Ruvubu National Park, Burundi 2 RWANDA TANZANIA 1 BURUNDI N 100 km Fig. 1. — Location of the Ruvubu National Park (1) in Burundi and Akagera National Park (2) in Rwanda. specimens and by the help of literature and special­ ists. The main bibliographic references consulted during taxa identification are: Blundell (1987) and Troupin (1971, 1978, 1982, 1983, 1985, 1988). Nomenclature follows Roux (2009) for ferns and Lebrun & Stork (1991­1997) for flowering plants. The names of flowering plants were checked in African Flowering Plants Database (AFPD) avail­ able online at http://www.ville­ge.ch/musinfo/ bd/cjb/africa/recherche.php. In order to provide a floristic inventory more complete as possible, our inventory was also partially completed by specific literature citations referring to the protected area (Ndabaneze 1989; Vande Weghe & Kabayanda 1992; Nzigidahera 2000). These documents pro­ duced at University of Burundi and at the National (Burundi) Institute for Environment and Nature Conservation (INECN) in different research ac­ tivities were consulted in order to evidence species newly encountered in the protected area. The main richest families from ANP were sorted through Troupin (1966) inventory. Chi­Square test (Scher­ rer 2007) was performed to test whether within the main richest families with at least 15 species each one, the numbers of species and genera from the two parks differ significantly. 157 Masharabu T. et al. Table 1. — Taxonomic composition of the Ruvubu National Park, Burundi. Family DICOTS Acanthaceae Amaranthaceae Anacardiaceae Annonaceae Apiaceae Apocynaceae Araliaceae Asteraceae Balsaminaceae Bignoniaceae Bombacaceae Boraginaceae Caesalpiniaceae Celastraceae Chrysobalanaceae Clusiaceae Combretaceae Convolvulaceae Cucurbitaceae Euphorbiaceae Fabaceae Flacourtiaceae Gentianaceae Lamiaceae Leeaceae Loranthaceae Malvaceae Melastomataceae Meliaceae Melianthaceae Menispermaceae Mimosaceae Monimiaceae Moraceae Myristicaceae MONOCOTS Agavaceae Amaryllidaceae Anthericaceae Arecaceae Asparagaceae Asphodelaceae Balanophoraceae Colchicaceae Commelinaceae Cyperaceae PTERIDOPHYES Cyatheaceae Dennstaedtiaceae Dryopteridaceae 158 Genera Species Subspecies/ Varieties Family Genera Species Subspecies/ Varieties 11 4 4 3 5 10 1 32 1 3 1 2 3 1 2 3 2 3 2 14 22 1 1 12 1 2 5 3 2 1 3 5 1 1 1 16 4 7 3 5 11 1 57 1 3 1 2 6 4 2 3 4 7 3 26 59 1 2 16 1 2 11 7 2 1 3 8 1 10 1 – – – – – – – – – – – – – – – – – – – – 5 – – – – – – – – – – – – – – Myrsinaceae Myrtaceae Nymphaeaceae Ochnaceae Olacaceae Oleaceae Onagraceae Opiliaceae Oxalidaceae Passifloraceae Pedaliaceae Phytolaccaceae Pittosporaceae Polygalaceae Polygonaceae Primulaceae Proteaceae Ranunculaceae Rhamnaceae Rhizophoraceae Rosaceae Rubiaceae Rutaceae Sapindaceae Sapotaceae Scrophulariaceae Strychnaceae Solanaceae Sterculiaceae Thymeleaceae Tiliaceae Ulmaceae Urticaceae Verbenaceae Vitaceae 1 2 1 1 2 2 1 1 2 1 1 1 1 2 2 1 2 1 2 1 1 19 1 4 1 2 1 1 2 1 1 2 2 3 4 2 4 2 2 2 2 2 1 3 1 2 1 1 2 4 1 4 3 2 1 1 28 1 4 1 2 4 7 2 3 4 2 2 7 9 2 1 – – 1 – – – – – – – – – – – – – – – – 3 – – – – 1 – – – – – 1 – 1 2 1 1 1 3 1 1 3 6 1 2 2 1 2 3 1 1 5 15 – – – – – – 2 – – – Dioscoreaceae Dracaenaceae Hyacinthaceae Hypoxidaceae Iridaceae Orchidaceae Poaceae Smilacaceae Xyridaceae Zingiberaceae 1 1 1 1 2 2 26 1 1 2 6 1 1 1 3 2 52 1 1 3 – – – – – – – – – – 1 2 1 3 2 1 – – – Nephrolepidaceae Osmundaceae Pteridaceae 1 1 1 1 1 1 – – – ADANSONIA, sér. 3 • 2012 • 34 (1) Vascular flora inventory and plant diversity of the Ruvubu National Park, Burundi RESULTS The floristic richness of the RNP comprises 522 spe­ cies (including infraspecific groups) distributed into 96 families and 306 genera (Table 1). In total, Dicots are better represented with 78% of the whole spe­ cies. Monocots represent 20% while Pteridophytes represent only 2% of the flora. The eight following families record over a half (51%) of the whole spe­ cies from RNP: Fabaceae (59 species), Asteraceae (57 species), Poaceae (52 species), Rubiaceae (28 species), Euphorbiaceae (26 species), Acanthaceae (16 species), Lamiaceae (16 species) and Cyperaceae (15 species). Table 2 establishes a comparison of floristic richness of families recording at least 15 species each one in RNP and ANP, relatively closer and belonging to the same phytochorion in the Lake Victoria regional mosaic according to White (1993). 96 species (18%) were encountered for the first time in the RNP with predominance of Fabaceae (23% of the newly encountered species). The inventory of species newly encountered in the RNP is as follows: Acanthaceae. Hypoestes verticillaris (L.f.) Sol. ex Roem. & Schult., Thunbergia alata Bojer ex Sims. Amaranthaceae. Alternanthera pungens Kunth. Convolvulaceae. Dichondra repens J.R.Forst. & G. Forst., Hewittia sublobata (L.f.) Kuntze. Cucurbitaceae. Zehneria thwaitesii (Schweinf.) C.Jeffrey. Cyperaceae. Kyllinga sphaerocephala Boeckeler, Mariscus macrocarpus Kunth. dennstaedtiaceae. Histiopteris incise (Thunb.) J. Sm. dioscoreaceae. Dioscorea alata L., Dioscorea bulbifera L., Dioscorea odoratissima Pax. euphorbiaceae. Acalypha ornata Hochst. ex A.Rich., Clutia abyssinica Jaub. & Spach, Croton macrostachyus Hochst. ex Delile, Phyllanthus niruri L., Synadenium grantii Hook.f., Tragia brevipes Pax. Fabaceae. Crotalaria aculeata De Wild., Crotalaria lachnophora A.Rich., Crotalaria ononoides Benth., Crotalaria sp., Desmodium salicifolium (Poir.) DC., Desmodium setigerum (E.Mey.) Benth. ex Harv., Desmodium triflorum (L.) DC., Desmodium velutinum (Willd.) DC., Eriosema chrysadenium Taub., Eriosema rhodesicum R.E.Fr., Eriosema stanerianum Hauman, Indigofera emarginella Steud. ex A.Rich., Indigofera homblei Baker f. & Martin, Indigofera paracapitata J.B.Gillett, Indigofera rhynchocarpa Welw. ex Baker, Indigofera spicata Forssk., Indigofera zenkeri Harms ex Baker f., Rhynchosia sublobata (Schumach. & Thonn.) Meikle, Tephrosia linearis (Willd.) Pers., Teramnus labialis (L.f.) Spreng., Vigna luteola (Jacq.) Benth., Vigna parkeri Baker. Asteraceae. Bidens steppia (Steetz) Sherff, Cirsium buchwaldii O.Hoffm., Crassocephalum multicorymbosum (Klatt) S.Moore, Crassocephalum rubens (Juss. ex Jacq.) S.Moore, Crassocephalum sarcobasis (DC.) S.Moore, Erlangea cordifolia (Benth. ex Oliv.) S.Moore, Helichrysum keilii Moeser, Pleiotaxis pulcherrima Steetz, Vernonia fontinalis S.Moore, Vernonia lasiopus O.Hoffm., Vernonia ugandensis S.Moore. lamiaceae. Leucas calostachya Oliv., Solenostemon platostomoides (Robyns & Lebrun) Troupin. Boraginaceae. Cordia africana Lam., Trichodesma zeylanicum (Burm.f.) R.Br. Malvaceae. Hibiscus aponeurus Sprague & Hutch., Sida cordifolia L. Caesalpiniaceae. Cassia kirkii Oliv., Cassia mimosoides L., Cassia siamea Lam. Melastomataceae. Antherotoma naudinii Hook.f., Dissotis brazzae Cogn., Dissotis ruandensis Engl., Dissotis senegambiensis (Guill. & Perr.) Triana, Dissotis trothae Gilg, Tristemma incompletum R.Br. Commelinaceae. Commelina benghalensis L., Commelina diffusa Burm.f. subsp. diffusa, Floscopa africana (P.Beauv.) C.B.Clarke. ADANSONIA, sér. 3 • 2012 • 34 (1) Mimosaceae. Acacia sieberiana DC. 159 Masharabu T. et al. Moraceae. Ficus ovata Vahl. osmundaceae. Osmunda regalis L. Poaceae. Hyparrhenia cymbaria (L.) Stapf, Hyparrhenia newtonii (Hack.) Stapf, Hyparrhenia rufa (Nees) Stapf, Loudetia kagerensis (K.Schum.) C.E.Hubb. ex Hutch., Oplismenus compositus (L.) P.Beauv., Pennisetum polystachion (L.) Schult., Setaria kagerensis Mez, Setaria pumila (Poir.) Roem. & Schult., Urelytrum digitatum K.Schum. Polygonaceae. Rumex usambarensis (Dammer) Dammer. Rubiaceae. Agathisanthemum globosum (Hochst. ex A.Rich.) Bremek., Fadogia obovata Schweinf., Hymenodictyon floribundum (Hochst. & Steud.) Robbr., Pavetta ternifolia (Oliv.) Hiern, Spermacoce dibrachiata Oliv., Spermacoce sphaerostigma (A.Rich.) Vatke. Scrophulariaceae. Buchnera keilii Mildbr., Lindernia sp. Solanaceae. Solanum anguivi Lam. Thymeleaceae. Gnidia ericoides C.H.Wright. Tiliaceae. Triumfetta cordifolia A.Rich. Urticaceae. Urtica massaica Mildbr. Vitaceae. Rhoicissus tridentata (L.f.) Wild & R.B.Drumm. Xyridaceae. Xyris capensis Thunb. DISCUSSION The 522 taxa recorded from the RNP represent 17% of the vascular plant species so far estimated for Burundi and 17% of the flora recorded in the Lake Victoria regional mosaic. The vascular plants are estimated at about more than 3000 species for Burundi (Ndabaneze 1989) and 3000 species in 160 the Lake Victoria regional mosaic (White 1993). Ferns are weakly represented in the PNR (nine species distributed into six families) and it is not strange as the RNP is mainly made of savannas. Moreover, it is well­known that the Pteridophytes diversity in Africa is exceptionally poor (Tryon 1986; Roux 2009). Burundi comprises 32 fami­ lies of Pteridophytes distributed into 69 genera and 178 species (Roux 2009). Therefore, the Pteridophytes of the RNP represent 5% of those of the whole country. The flora of the RNP represents besides 5% of the flora of Central Africa (Burundi, Democratic Republic of Congo and Rwanda). According to Brenan (1978), floristic richness of the Cen­ tral Africa region is estimated at more or less 10 000 species. According to Léonard (1994), spermatophytes’ statistics of the flora of Central Africa include 9377 species. Burundi (27 834 km2) being confined between Eastern and Cen­ tral Africa (Bidou et al. 1991), the RNP floristic richness represents about 4.8% of the flora of Eastern Africa which is estimated according to Linder (2001), at 11 000 species. The plant communities of the RNP present rela­ tively a poor taxonomic diversity when compared with tropical forests. In the humid tropical forests, flowering plants are estimated at 7146 species in the Guianas (Guyana, Suriname, French Guiana) according to Haripersaud et al. (2010), 4300 species in French Guiana (90 000 km2) and 8000 species in Peninsular Malaysia (130 000 km2) according to Blanc (1997). The tropical forests record the largest known specific diversity and also shelter numerous even unknown species, because inven­ tories are far to be finished there (Puig 2001). Families with the greatest number of species in RNP and ANP are not statistically different in terms of species richness as well as for genera richness (p>0.05). It was hypothesized that plant diversity in RNP and ANP in Rwanda would be similar due to their biogeographical location at the crossroads of several biogeographical influ­ ences in the Lake Victoria regional mosaic. The hypothesis is accepted for species and generic richness within families in the two protected areas. However, even if they are not statistically ADANSONIA, sér. 3 • 2012 • 34 (1) Vascular flora inventory and plant diversity of the Ruvubu National Park, Burundi Table 2. — Comparison of the floristic richness of families with at least 15 species each one in Ruvubu National Park with their correspondences from Troupin (1966) in Akagera National Park (species: ddl = 7, c2 = 9.98, p > 0.05; genera: ddl = 7, c2 = 2.63, p > 0.05). Families Fabaceae Asteraceae Poaceae Rubiaceae Euphorbiaceae Acanthaceae Lamiaceae Cyperaceae Ruvubu National Park, Burundi species genera observed expected observed expected 59 57 52 28 26 16 16 15 51 59 61 23 21 19 16 19 22 32 26 19 14 11 12 6 different, the numeric gap in observed species and generic richness would be due to the fact that the ANP includes xerophytic groves and xerophytic forests not found in Ruvubu, while wetlands spread on 50% of the total area. The current size of the ANP has been fixed at 108 500 ha in 1997 after an encroachment of practically ⅔ of the total area with negative effects on biodiversity (Buda et al. 2005). The Fabaceae family has the greatest number of species in RNP. It is also the third biggest family of flowering plants worldwide after Orchids and Asteraceae (Mabberley 1997; Lewis et al. 2005). Fabaceae and Asteraceae are basically the first predominant families in Central Africa (Burundi, Democratic Republic of Congo and Rwanda) in terms of species richness listed by Léonard (1994). Particularly, according to Rundel (1989), Arianoutsou & Thanos (1996), the Leguminosae taxa (Fabaceae, Caesalpiniaceae & Mimosaceae) are ecologically very successful in terms of dominance and productivity. Leguminosae, through their symbiotic abilities to fix atmospheric nitrogen, have adaptations to establish on soils with lim­ ited nitrogen availability and play an important role in colonizing disturbed ecosystem, including those that are fire prone (Arianoutsou & Thanos 1996), of which the RNP. That would justify partially their abundance in the RNP and within the newly encountered species. Nevertheless, the study of environmental determinants of Legumi­ nosae abundance and floristic variability would provide more precisions. ADANSONIA, sér. 3 • 2012 • 34 (1) 21 33 30 17 12 12 11 5 Akagera National Park, Rwanda species genera observed expected observed expected 65 85 95 28 25 29 23 31 73 83 86 33 30 26 23 27 24 39 40 19 13 16 12 5 25 38 36 21 15 15 13 6 Acknowledgements Authors are grateful to the Government of Burundi for its support as well as to the INECN staff based at the surroundings of the RNP for field assistance. Many thanks to the staff of the National Botanic Garden of Belgium at Meise, particularly to the Engineer Luc Pauwels who helped to identify the Poaceae and to Daniel Geerinck who provided con­ viviality to the first author for accessing to Meise. REFERENCES ARIANOUTSOU M. & THANOS C. A. 1996. — Legumes in the fire­prone Mediterranean regions: an example from Greece. International Journal of Wildland Fire 6: 77­82. BIDOU J. E., NDAYIRUKIYE S., NDAYISHIMIYE J. P. & SIRVEN P. 1991. — Géographie du Burundi. Hatier, Paris, 288 p. BIGENDAKO M. J. 1989. — Recherche ethnopharmacognosique sur les plantes utilisées en médecine traditionnelle au Burundi occidental. Thèse de doctorat, Université Libre de Bruxelles, 352 p. BIZURU E. 2005. — étude de la flore et de la végétation des marais du Burundi. Thèse de doctorat, Université Libre de Bruxelles, 311 p. BLANC P. 1997. — La biodiversité tropicale mémoire des changements du passé. Lettre PIGB-PMRC 6: 30­38. BLUNDELL M. 1987. — Collins Photo Guide to the Wild flowers of East Africa. Harper Collins Publishers, London, 464 p. BRENAN J. P. M. 1978. — Notes on Mimosoideae XIII. New species of Entada & Acacia from Africa. Kew Bulletin 32: 545­550. BUDA P., SIMM H., MUGABUKOMEYE B., HAKIZIMANA E. 2005. — Parc national de l’Akagera. Plan d’aménagement et de gestion 2006-2010. Draft. GTZ/DED & ORTPN, Kigali, 96 p. 161 Masharabu T. et al. HARIPERSAUD P., TER STEEGE H., GRANVILLE J.-J. DE, CHEVILLOTTE H. & HOFF M. 2010. — Species abun­ dance, distribution and diversity in time and space after centuries of botanical collecting in the Guianas. Taxon 59 (2): 592­597. LAURANCE W. F., ALONSO A., LEE M. & CAMPBELL P. 2006. — Challenges for forest conservation in Gabon, Central Africa. Futures 38: 454­470. LEBRUN J. P. & STORK A. L. 1991­1997. — énumération des plantes à fleurs d’Afrique tropicale. Conservatoire et Jardin Botanique de la ville de Genève, 4 volumes. LÉONARD J. 1994. — Statistiques des spermatophytes de la flore d’Afrique centrale de 1940 à 1990. Bulletin du Jardin Botanique national de Belgique 63: 181­194. LEWALLE J. 1972. — Les étages de végétation du Burundi occidental. Bulletin du Jardin Botanique national de Belgique 42 (1/2): 1­247. LEWIS G., SHRIRE B., MACKINDER B. & LOCK M. 2005. — Legumes of the World. Royal Botanic Gardens Kew, Richmond, Surrey, 592 p. LINDER H. P. 2001. — Plant diversity and endemism in sub­Saharan tropical Africa. Journal of Biogeography 28: 169­182. LUBINI A. 1982. — Végétation messicole et postculturale des sous-régions de Kisangani et de la Tshopo (Haut-Zaïre). Thèse de doctorat, université de Kisangani, 489 p. MABBERLEY D. J. 1997. — The Plant Book. A Portable Dictionary of the Vascular Plants. 2nd edition. Cam­ bridge University Press, 858 p. MASHARABU T., LEJOLY J., BIGENDAKO M. J., BOGAERT J. & GODART M.-F. 2008. — Diversité floristique du Parc national de la Ruvubu (Burundi). Bulletin scientifique de l’INECN 6: 2­7. MASHARABU T., BIGENDAKO M. J., LEJOLY J., NKENGURUTSE J., NORET N., BIZURU E. & BOGAERT J. 2010. — étude analytique de la flore et de la végétation du Parc national de la Ruvubu, Burundi. International Journal of Biological and Chemical Sciences 4 (4): 834­856. NDABANEZE P. 1983. — La Flore graminéenne du Burundi. Taxonomie et écogéographie. Thèse de doctorat, uni­ versité de Liège, 293 p. NDABANEZE P. 1989. — Catalogue des graminées du Burundi. Nouvelle série n° 132. Les éditions de Lejeunia, 127 p. NZIGIDAHERA B. 2000. — Analyse de la diversité bio­ logique végétale nationale et identification des prio­ rités pour sa conservation. INECN, Bujumbura, Burundi, 126 p. PUIG H. 2001. — Diversité spécifique et déforestation : l’exemple des forêts tropicales humides du Mexique. Bois et forêts des tropiques 268 (2): 41­55. ROUX J. P. 2009. — Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and Neighbouring Islands. Strelitzia 23. South African National Biodi­ versity Institute, 296 p. RUNDEL P. W. 1989. — Ecological success in relation to plant form and function in the woody legumes, in STIRTON C. H. & ZARUCCHI J. L. (eds), Advances in Legume Biology. Monographs in Systematic Botany from the Missouri Botanical Garden 29: 377­398. SCHERRER B. 2007. — Biostatistique. Vol. 1, 2e édition. Les éditions de la Chenelière inc., Montréal, 816 p. TROUPIN G. 1966. — étude phytocénologique du Parc national de l’Akagera et du Rwanda oriental. Recherche d’une méthode d’analyse appropriée à la végétation d’Afrique intertropicale. Publication n° 2. INRS, Butare, 293 p. TROUPIN G. 1971. — Syllabus de la flore du Rwanda. Spermatophytes. Musée royal de l’Afrique centrale. Tervuren (Belgique), 356 p. TROUPIN G. 1978. — Flore du Rwanda. Spermatophytes. Vol. 1. Série in 8° Sciences écon. n° 9. Annales Musée royal de l’Afrique centrale, Tervuren, 413 p. TROUPIN G. 1982. — Flore des plantes ligneuses du Rwanda. Musée royal de l’Afrique centrale, Tervuren, 747 p. TROUPIN G. 1983. — Flore du Rwanda. Spermatophytes. Vol. 2. Série in 8° Sciences écon. n° 13. Annales Musée royal de l’Afrique centrale, Tervuren, 603 p. TROUPIN G. 1985. — Flore du Rwanda. Spermatophytes. Vol. 3. Série in 8° Sciences écon. n° 15. Annales Musée royal de l’Afrique centrale, Tervuren, 729 p. TROUPIN G. 1988. — Flore du Rwanda. Spermatophytes. Vol. 4. Musée royal de l’Afrique centrale, Tervuren, 651 p. TRYON R. 1986. — The biogeography of species, with spe­ cial reference to ferns. The Botanical Review 52:117­156. VANDE WEGHE J. P. & KABAYANDA A. 1992. — Le Parc national de la Ruvubu et sa région limitrophe : étude d’identification de la Ruvubu : étude n° ET/44/2/92­ SEP.MINATE­CEE. 195 p. WHITE F. 1993. — The AETFAT chorological classification of Africa: history, methods and applications. Bulletin du Jardin Botanique national de Belgique 62: 225­281. Submitted on 20 October 2010; accepted on 24 March 2011. 162 ADANSONIA, sér. 3 • 2012 • 34 (1)