Plant Ecology and Evolution 153(3): 373-389, doi: 10.5091/plecevo.2020.1754
Patterns in the alien flora of the Democratic Republic of the Congo: a comparison of Asteraceae and Fabaceae
expand article infoFarzaneh Bordbar, Pierre Meerts§
‡ Herbarium et bibliothèque de botanique africaine, Université Libre de Bruxelles, Avenue F.D. Roosevelt 50, CP 265, B-1050 Brussels, Belgium§ Universite Libre de Bruxelles, Bruxelles, Belgium
Open Access

Background and aims – This work provides the first pattern analysis of the alien flora of the Democratic Republic of the Congo (D.R. Congo), using Asteraceae and Fabaceae as a case study.

Methods – Based on herbarium collections, existing databases, and literature data, a database of 38 alien species of Asteraceae and 79 alien species of Fabaceae has been assembled. Patterns in the introduction pathway, phylogeny, life form, morpho-functional traits, geographic origin, and occurrence in D.R. Congo are explored.

Key results – America is the main source continent in both families, but Asia is also an important donor of Fabaceae. Taxonomic spectrum discrepancies between the alien and the native flora reflect the continent of origin. Sixty-six percent of alien Asteraceae have been accidentally introduced, most of which being annual weeds of disturbed soil. In contrast, 90% of alien Fabaceae have been deliberately introduced for forestry, agriculture, or environmental purposes, most of which being phanerophytes. Traits were compared between pairs of congeneric alien and native species. For Asteraceae, a sharp discrepancy was found in the life form spectrum (aliens: mostly therophytes; natives: phanerophytes). For Fabaceae, alien species had larger leaves and larger pods compared to their native congeners. The number of specimens in collections was positively correlated with the time since the date of first collection for both families. The Guineo-Congolian region has the highest number of alien Fabaceae, while alien Asteraceae are overrepresented in the Zambezian region.

Conclusions – Contrasting patterns between alien Asteraceae and Fabaceae in the flora of D.R. Congo in terms of life forms, trait divergence compared to the native flora, and occurrence, reflect the divergent biological attributes and relations to humans of the two families. The striking discrepancies between the two families call for analyses of patterns of alien flora at family level and warn against global generalisations.

Alien, Asteraceae, checklist, congeneric pairs, Fabaceae, invasion, life form, naturalization, traits, tropical Africa


  • Anonymous (2012) Synthèse des études sur les causes de la déforestation et de la dégradation des forêts en République Démocratique du Congo. Kinshasa, Ministère de l’environnement, Conservation de la Nature et Tourisme, UN-reDD Programme.
  • Ansong M., Pergl J., Essl F., Hejda M., van Kleunen M., Randall R., Pyšek P. (2019) Naturalized and invasive alien flora of Ghana. Biological Invasions 21(3): 669–683.
  • Bigirimana J., Bogaert J., De Cannière C., Bigendako M.-J., Parmentier I. (2012) Domestic garden plant diversity in Bujumbura, Burundi: Role of the socio-economical status of the neighborhood and alien species invasion risk. Landscape and Urban Planning 107(2): 118–126.
  • Binggeli P. (2011) The human dimensions of invasive plants in tropical Africa. In: Rotherham I.D., Lambert R. (eds) Invasive and introduced plants and animals: human perceptions, attitudes and approaches to management: 201–220. Abingdon, Earthscan.
  • Binggeli P., Hall J.B., Healey J.R. (1998) An overview of invasive woody plants in the tropics. School of Agricultural and Forest Sciences Publication 13. Bangor, University of Wales.
  • Blackburn T.M. Pyšek P., Bacher S., Carlton J.T., Duncan R.P., Jaros V., Wilson J.R.U., Richardson D.M. (2011) A proposed unified framework for biological invasions. Trends in Ecology & Evolution 26(7): 333–339.
  • Borokini T.I. (2011) Invasive alien plant species in Nigeria and their effects on biodiversity conservation. Tropical Conservation Science 4(1): 103–110. 10.1177%2F194008291100400110
  • Boy G., Witt A. (2013) Invasive alien plants and their management in Africa Malta, Gutenberg Press Limited.
  • Chenje M., Mohamed-Katerere J. (2006) Invasive alien species. In: Africa Environment Outlook 2: Our Environment, Our Wealth: 331–349. Nairobi, United Nations Environment Programme.
  • Chevalier A. (1931) Le rôle de l’Homme dans la dispersion des plantes tropicales. Échanges d’espèces entre l’Afrique Tropicale et l’Amérique du Sud. Revue de botanique appliquée et d’agriculture coloniale 120: 633–650.
  • Crawford P.H.C., Hoagland B.W. (2009) Can herbarium records be used to map alien species invasion and native species expansion over the past 100 years? Journal of Biogeography 36: 651–661.
  • Dawson W., Mndolwa A.S., Burslem D.R.F.P., Hulme P.E. (2008) Assessing the risk of plant invasions from collections from tropical botanical gardens. Biodiversity Conservation 17: 1979–1995.
  • Dawson W., Burslem D.F.R.P., Hulme P.E. (2011) The comparative importance of species traits and introduction characteristics in tropical plant invasions. Diversity and Distributions 17: 1111–1121.
  • Divíšek J., Chytrý M., Beckage B., Gotelli N.J., Lososová Z., Pyšek P., Richardson D.M., Molofsky J. (2018) Similarity of introduced plant species to native ones facilitates naturalization, but differences enhance invasion success. Nature Communications 9: 4631.
  • Essl F., Dawson W., Kreft H., Pergl J., Pyšek P., van Kleunen M., Weigelt P., Mang T., Dullinger S., Lenzner B., Moser D., Maurel N., Seebens H., Stein A., Weber E., Chatelain C., Inderjit, Genovesi P., Kartesz J., Morozova O., Nishino M., Nowak P.M., Pagad S., Shu W., Winter M. (2019) Drivers of the relative richness of naturalized and invasive plant species on Earth. AoB PLANTS 11: plz051.
  • Foxcroft L.C. Richardson D.M., Wilson J.R.U. (2008) Ornamental plants as invasive aliens: problems and solutions in Kruger National Park, South Africa. Environmental Management 41: 32–51.
  • Foxcroft L.C., Richardson D.M., Rejmánek M., Pyšek P. (2010) Alien plant invasions in tropical and sub-tropical savannas: patterns, processes and prospects. Biological Invasions 12: 3913–3933.
  • Foxcroft L.C., Witt A., Lotter W.D. (2013) Icons in peril: invasive alien plants in African protected areas. In: Foxcroft L.C., Pyšek P., Richardson D.M., Genovesi P. (eds) Plant invasions in protected areas: 117–144. Dordrecht, Springer.
  • Fu Z.-X., Jiao B.-H., Nie B., Zhang G.-J., Gao T.-G. (2016) A comprehensive generic-level phylogeny of the sunflower family: Implications for the systematics of Chinese Asteraceae. Journal of Systematics and Evolution 54(4): 416–437.
  • Gallagher R.V., Randall R.P., Leishman M.R. (2015) Trait differences between naturalized and invasive plant species independent of residence time and phylogeny. Conservation biology 29(2): 360–369.
  • Gichua M., Njoroge G., Shitanda D., Ward D. (2013) Invasive species in East Africa: current status for informed policy decisions and management. Jomo Kenyatta University of Agriculture and Technology 15(1): 45–55.
  • Godoy O., Valladares F., Castro-Díez P. (2011) Multispecies comparison reveals that invasive and native plants differ in their traits but not in their plasticity. Functional Ecology 25: 1248–1259.
  • Grotkopp E., Rejmánek M. (2007) High seedling relative growth rate and specific leaf area are traits of invasive species: phylogenetically independent contrasts of woody angiosperms. American Journal of Botany 94: 526–532.
  • Hammer Ø., Harper D.A.T., Ryan P.D. (2001) PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontologia Electronica 4(1): 9p. Version 3.25. Available at [accessed 1 Apr. 2019].
  • Harrison I.J., Brummett R., Stiassny M.L.J. (2016) The Congo River Basin. In: Finlayson C.M., Milton G.R., Prentice R.C., Davidson N.C. (eds) The Wetland Book: 1–18. Dordrecht, Springer Science.
  • Holou R.A.Y., Achigan-Dako E.G., Sinsin B. (2013) Ecology and management of invasive plants in Africa. In: Ose S., Singh H., Kohli R., Batish D. (eds) Invasive plant ecology: 161–174. Boca Raton, CRC Press.
  • Howard G.W., Chege F.W. (2007) Invasions by plants in the inland waters and wetlands of Africa. In: Gherardi F. (ed.) Biological invaders in inland waters: Profiles, distribution, and threats: 193–208. Dordrecht, Springer.
  • Howard G.W., Matindi S.W. (2003) Alien invasive species in Africa’s wetlands: some threats and solutions. Nairobi, IUCN Eastern Africa Regional Office
  • IUCN/PACO (2013) Invasive plants affecting protected areas of West Africa. Management for reduction of risk for biodiversity. Ouagadougou, IUCN/PACO.
  • Katz E. (1998) Plantes américaines au sud Congo. In: Chastanet M. (ed.) Plantes et paysages d’Afrique: une histoire à explorer: 283–322. Paris, Karthala, Centre de Recherches Africaines.
  • Klopper R.R., Gautier L., Chatelain C., Smith G.F., Spichiger R. (2007) Floristics of the angiosperm flora of sub-Saharan Africa: an analysis of the African plant checklist and database. Taxon 56 (1): 201–208.
  • Kull C.A., Tassin J., Moreau S., Rakoto Ramiarantsoa H., Blanc-Pamard C., Carrière S.M. (2011) The introduced flora of Madagascar. Biological Invasions 14(4): 875–888.
  • Küper W., Sommer J.H., Lovett J.C., Mutke J., Linder H.P., Beentje H., van Rompaey R.A.S.R., Chatelain C., Sosef M., Barthlott W. (2005) Africa’s hotspots of biodiversity redefined. Annals of the Missouri Botanical Garden 91: 525–536.
  • Lambertini M., Leape J., Marton-Lefevre J., Mittermeier R.A., Rose M., Robinson J.G., Stuart S.N., Waldman B., Genovesi P. (2011) Invasive: a major conservation threat. Science 333(6041): 404–405.
  • Latombe G., Pyšek P., Jeschke J.M., Blackburn T.M., Bacher S., Capinha C., Costello M.J., Fernández M., Gregory R.D., Hobern D., Hui C., Jetz W., Kumschick S., McGrannachan C., Pergl J., Roy H.E., Scalera R., Squires Z.E., Wilson J.R.U., Winter M., Genovesi P., McGeoch M.A. (2017) A vision for global monitoring of biological invasions. Biological Conservation 213: 295–308.
  • Lebrun J., Gilbert G. (1954) Une classification écologique des forêts du Congo. Bruxelles, Institut national pour l’étude agronomique du Congo belge.
  • LPWG (2017) A new subfamily classification of the Leguminosae based on a taxonomically comprehensive phylogeny: The Legume Phylogeny Working Group (LPWG). Systematics and Phylogeny 66(1): 44–77.
  • Mack R.N., Simberloff D., Lonsdale W.M., Evans H., Clout M., Bazzaz F.A. (2000) Biotic invasions: causes, epidemiology, global consequences, and control. Ecological Applications 10(3): 689–710.
  • Malele Mbala S. (2003) Situation des ressources génétiques forestières de la République démocratique du Congo. Note thématique sur les ressources génétiques forestières. Document de travail FGR/56F. Département des forêts, Division des ressources forestières. Rome, FAO.
  • Masocha M., Skidmore A.K., Poshiwa X., Prins H.H.T. (2011) Frequent burning promotes invasions of alien plants into a mesic African savanna. Biological Invasions 13: 1641–1648.
  • Mbale H.K., Mukendi M.T., Bongo G.N., Kikufi A.B., Lukoki F.L. (2019) Floristic inventory of invasive alien aquatic plants found in some Congolese rivers, Kinshasa, Democratic Republic of the Congo. Asian Journal of Environment & Ecology 11(4): 1–15.
  • McGeoch M.A., Genovesi P., Bellingham P.J., Costello M.J., Mcgrannachan C.M., Sheppard A. (2016) Prioritizing species, pathways, and sites to achieve conservation targets for biological invasion. Biological Invasions 18: 299–314.
  • Moravcová L., Pyšek P., Jarošík V., Pergl J. (2015) Getting the right traits: reproductive and dispersal characteristics predict the invasiveness of herbaceous plant species. PLOS ONE 10(4): e0123634.
  • Noba K., Bassene C., Ngom A., Gueye M., Camara A.A., Kane M., Ndoye F., Dieng B., Rmballo R., Bâ N., Bodian M.Y., Sane S., Diop D., Kontai S.A., Kane,A., Mbaye M.S., Bâ A. (2017) Invasive plants of West Africa: concepts, overviews and sustainable management. Advances in Recycling Waste Management 2: 121.
  • Obiri J.F. (2011) Invasive plant species and their disaster-effects in dry tropical forests and rangelands of Kenya and Tanzania. Journal of Disaster Risk Studies 3(2): 417–428.
  • Pagad S., Genovesi P., Carnevali L., Schigel D., McGeoch M.A., (2018) Introducing the global register of introduced and invasive species. Scientific Data 5: 170202.
  • Peel M.C., Finlayson B.L., McMahon T.A. (2007) Updated world map of the Köppen-Geiger climate classification. Hydrology and Earth System Sciences 11: 1633–1644.
  • Potapov P.V., Turubanova S.A., Hansen M.C., Adusei B., Broich M., Altstatt A., Mane L., Justice C.O. (2013) Quantifying forest cover loss in Democratic Republic of the Congo, 2000–2010, with Landsat eTM+ data. Remote Sensing of Environment 122: 106–116.
  • Pyšek P., Richardson D.M. (2007) Traits associated with invasiveness in alien plants: where do we stand? In: Nentwig W. (ed.) Biological Invasions: 97–126. Berlin, Springer-Verlag.
  • Pyšek P., Richardson D., Rejmánek M., Webster G., Williamson M., Kirschner J. (2004) Alien plants in checklists and floras: towards better communication between taxonomists and ecologists. Taxon 53(1): 131–143.
  • Pyšek P., Richardson D.M., Pergl J., Jarosík V., Sixtová Z., Weber E. (2008) Geographical and taxonomic biases in invasion ecology. Trends in Ecology & Evolution 23(5): 237–244.
  • Pyšek P., Jarošík V., Pergl J. (2011) Alien plants introduced by different pathways differ in invasion success: unintentional introductions as a threat to natural areas. PLOS ONE 6(9): e24890.
  • Pyšek P., Pergl J., Essl F., Lenzner B., Dawson W., Kreft H., Weigelt P., Winter M., Kartesz J., Nishino M., Antonova L.A., Barcelona J.F., Cabezas F.J., Cárdenas D., Cárdenas-Toro J., Castaño N., Chacón E., Chatelain C., Dullinger S., Ebel A.L., Figueiredo E., Fuentes N., Genovesi P., Groom Q.J., Henderson L., Inderjit, Kupriyanov A., Masciadri S., Maurel N., Meerman J., Morozova O., Moser D., Nickrent D., Nowak P.M., Pagad S., Patzelt A., Pelser P.B., Schulze M., Seebens H., Shu W., Thomas J., Velayos M., Weber E., Wieringa J.J., Baptiste M.P., van Kleunen M. (2017) Naturalized alien flora of the world: species diversity, taxonomic and phylogenetic patterns, geographic distribution and global hotspots of plant invasion. Preslia 89: 203–274.
  • RStudio Team (2015) RStudio: Integrated Development for R. Version 1.1.442. RStudio, Inc., Boston, MA. Available at [accessed 7 Apr. 2018].
  • Randall R.P. (2017) A global compendium of weeds. Ed.3. Perth, CABI.
  • Randall J.M., Morse L.E., Benton N., Hiebert R., Lu S., Killeffer T. (2008) The invasive species assessment protocol: a tool for creating regional and national lists of invasive nonnative plants that negatively impact biodiversity. Invasive Plant Science and Management 2008 1: 36–49.
  • Rejmánek M., Huntley B.J., Le Roux J.J., Richardson D.M. (2016) A rapid survey of the invasive plant species in western Angola. African Journal of Ecology 55: 56–69.
  • Richardson D.M., Binggeli P., Schroth G. (2004) Invasive agroforestry trees: problems and solutions. In: Schroth G., de Fonseca G.A.B., Harvey C.A., Gascon C., Vasconcelos H., Izac A.-M.N. (eds) Agroforestry and biodiversity conservation in tropical landscapes: 371–396. Washington D.C., Island Press.
  • Richardson D.M, Foxcroft L.C., Latombe G., Le Maitre D.C., Rouget M., Wilson J.R. (2020) The Biogeography of South African Terrestrial Plant Invasions. In: van Wilgen B.W., Measey J., Richardson D.M., Wilson J.R., Zengeya T.A. (eds) Biological invasions in South Africa: 67–96. Cham, Springer Nature Switzerland.
  • Robyns W. (1948) Les territoires phytogéographiques du Congo belge et du Ruanda-Urundi. Atlas Général du Congo belge. Bruxelles, Institut Royal Colonial Belge.
  • Roussel B., Juhe-Beaulaton D. (1992) Les plantes américaines en Afrique tropicale. Les Cahiers d’Outre-Mer 179–180: 373–386.
  • Seburanga J.L., Bizuru E., Mwavu E.N., Kampungu K.G., Gatesire T., Kaplin B.A., (2016) A new method for post-introduction risk assessment of biological invasions among introduced shrubs in developing countries. Environmental Management 57: 572–584.
  • Seebens H., Blackburn T.M., Dyer E.E., Genovesi P., Hulme P.E., Jeschke J.M., Pagad S.N., Pyšek P., Kleunen M.V., Winter M., Ansong M., Arianoutsou M., Bacher S., Blasius B., Brockerhoff E.G., Brundu G., Capinha C., Causton C.E., Celesti-Grapow L., Dawson W., Dullinger S., Economo E.P., Fuentes N.R., Guénard B., Jäger H., Kartesz J.T., Kenis M.S., Kühn I., Lenzner B., Liebhold A.M., Mosena A., Moser D., Nentwig W., Nishino M., Pearman D.A., Pergl J., Rabitsch W.B., Rojas-Sandoval J., Roques A.R., Rorke S.L., Rossinelli S., Roy H.E., Scalera R., Schindler S., Štajerová K., Tokarska-Guzik B., Walker K.W., Ward D.F., Yamanaka T., Essl F. (2018) Global rise in emerging alien species results from increased accessibility of new source pools. Proceedings of the National Academy of Sciences of the United States of America 115(10): E2264–E2273.
  • Sosef M.S.M., Dauby G., Blach-Overgaard A., van der Burgt X., Catarino L., Damen T., Deblauwe V., Dessein S., Dransfield J., Droissart V., Duarte M.C., Engledow H., Fadeur G., Figueira R., Gereau RE., Hardy O.J., Harris D.J., de Heij J., Janssens S., Klomberg Y., Ley AC., Mackinder BA., Meerts P., de Poel J.L.V., Sonké B., Stévart T., Stoffelen P., Svenning J.C., Sepulchre P., Zaiss R., Wieringa J.J., Couvreur T.L.P. (2017) Exploring the floristic diversity of tropical Africa. BMC Biology 15(1): 15.
  • Turbelin A.J., Malamud B.D., Francis R.A. (2017) Mapping the global state of invasive alien species: patterns of invasion and policy responses. Global Ecology and Biogeography 26(1): 78–92.
  • Useni Sikuzani Y., Sambiéni Kouagou R., Maréchal J., Ilunga wa Ilunga E., Malaisse F., Bogaert J., Munyemba Kankumbi F. (2018) Changes in the spatial pattern and ecological functionalities of green spaces in Lubumbashi (the Democratic Republic of Congo) in relation with the degree of urbanization. Tropical Conservation Science 11: 1–17. 10.1177%2F1940082918771325
  • van Kleunen M., Dawson W., Essl F., Pergl J., Winter M., Weber E., Kreft H., Weigelt P., Kartesz J., Nishino M., Antonova L.A., Barcelona J.F., Cabezas F.J., Cárdenas D., Cárdenas-Toro J., Castaño N., Chacón E., Chatelain C., Ebel A.L., Figueiredo E., Fuentes N., Groom Q.J., Henderson L., Inderjit, Kupriyanov A., Masciadri S., Meerman J., Morozova O., Moser D., Nickrent D.L., Patzelt A., Pelser P.B., Baptiste M.P., Poopath M., Schulze M., Seebens H., Shu W.S., Thomas J., Velayos M., Wieringa J.J.,Pyšek P. (2015) Global exchange and accumulation of non-native plants. Nature 525(7567): 100–103.
  • van Kleunen M., Pyšek P., Dawson W., Essl F., Kreft H., Pergl J., Weigelt P., Stein A., Dullinger S., König C., Lenzner B., Maurel N., Moser D., Seebens H., Kartsz J., Nishino M., Aleksanyan A., Ansong M., Antonova L.A., Barcelona J.F., Breckle S., Brundu G., Cabezas F.J., Cardenas D., Cardenas-Toro J., Castano N., Chacon E., Chatelain C., Conn B., de Sa Dechoum M., DufourDror J.M., Ebel A.L., Figueiredo E., Fragman-Sapir O., Fuentes N., Groom Q.J., Henderson L., Inderjit, Jogan N., Krestov P., Kupriyanov A., Masciadri S., Meerman J., Morozova O., Nickrent D., Nowak A., Patzelt A., Pelser P.B., Shu W.S., Thomas J., Uludag A., Velayos M., Verkhosina A., Villasenor J.L., Weber E., Wieringa J.J., Yazlik A., Zeddam A., Zykova E., Winter M. (2019) The Global Naturalized Alien Flora (GloNAF) database. Ecology 100: e02542.
  • White F. (1983) Unesco/AETFAT/UNSO vegetation map of Africa. Scale 1: 5 000 000 (in colour). Paris, Unesco.
  • Wild H. (1968) Weeds and aliens in Africa: the American immigrant. Salisbury, University College of Rhodesia.
  • Witt A.B.R., Beale T., Van Wilgen B.W. (2018) An assessment of the distribution and potential ecological impacts of invasive alien plant species in eastern Africa. Transactions of the Royal Society of South Africa 73(3): 217–236.
  • Witt A.B.R., Shackleton R.T., Beale T., Nunda W., Van Wilgen B.W. (2019) Distribution of invasive alien Tithonia (Asteraceae) species in eastern and southern Africa and the socio-ecological impacts of T. diversifolia in Zambia. Bothalia 49(1): a2356.
  • Zachariades C., van Rensburg S.J., Witt A. (2013) Recent spread and new records of Chromolaena odorata in Africa. In: Zachariades C., Strathie L.W., Day M.D., Muniappan R. (eds) Proceedings of the Eighth International Workshop on Biological Control and Management of Chromolaena odorata and other Eupatorieae: 20–27. Pretoria, ARC-PPRI.