Spatial distribution patterns of Afzelia africana (Fabaceae – Detarioideae) in a tropical savanna of Benin: implications for management
cover image of Plant Ecology and Evolution 154(3)

Supplementary Files

Supplementary file 1
Supplementary file 2
Supplementary file 3
Supplementary file 4


pair correlation function
plant coexistence
plant-plant interactions
point pattern process
tree-to-tree distance

How to Cite

Atanasso, J., Salako, V., Mensah, S., Tohoun, R., Djossa, B., Glèlè Kakaï, R. and Assogbadjo, A. (2021) “Spatial distribution patterns of Afzelia africana (Fabaceae – Detarioideae) in a tropical savanna of Benin: implications for management”, Plant Ecology and Evolution, 154(3), pp. 362-375. doi: 10.5091/plecevo.2021.1713.


Background and aims – Understanding the spatial patterns and associations of tree species with their conspecific and heterospecific neighbours is critical for sustainable management of their stands. This study assessed the intra- and interspecific spatial structure of six life stages in Afzelia africana, a keystone multipurpose and endangered tree species in a tropical savanna of Benin.
Material and methods – Three plots of 4 ha each were demarcated on three sites along a conservation gradient (hunting zone – core conservation zone). Individuals of A. africana (irrespective of their diameter at breast height) and heterospecific trees (dbh ≥ 5 cm) were mapped. Tree spatial patterns and associations were determined using univariate and bivariate pair correlation functions. The distance to the nearest neighbour was further used to assess tree-to-tree distance.
Key results – We found variable spatial patterns across sites. In the core zone where wildlife density is high, most life stages had a random distribution. In contrast, in the hunting zone where wildlife density is low, the species spatial distribution changed from a predominantly aggregative pattern during early stages to a less aggregative or random spatial pattern for very large adults. Most pairs of life stages showed neutral associations, except for small and large adults, which had positive association between themselves on two sites. We also found that A. africana tree spatial distribution was unrelated to heterospecific trees.
Conclusion – We suggest that bush fire, seed dispersion, predation, and local environment would have contributed to the observed patterns.


Adomou C.A., Agbani O.P. & Sinsin B. 2011. Plants. In: Neuenschwander P., Sinsin B. & Goergen G. (eds) Protection de la nature en Afrique de l’Ouest: une liste rouge pour le Bénin. Nature Conservation in West Africa: Red List for Benin: 21–46. International Institute of Tropical Agriculture, Ibadan.

Amahowe O.I., Azihou A.F., Armand N.K., et al. 2019. Constraints of seedling survival and recruitment under adult tree of a multi-use species Afzelia africana Sm. ex Pers. in Sudanian region of Benin (West Africa). Agroforestry Systems 93(3): 1147–1156.

Amahowe O.I., Biaou S.S., Natta A.K. & Balagueman R.O. 2018. Multiple disturbance patterns and population structure of a tropical tree species, Afzelia africana (Leguminosae–Caesalpinioideae), in two contrasting bioclimatic zones of the Republic of Benin. Southern Forests: a Journal of Forest Science 80: 95–103.

Arbonnier M. 2004. Trees, shrubs and lianas of West African dry zones. CIRAD, MNHN, Montpellier.

Assédé E.P., Adomou A.C. & Sinsin B. 2012. Magnoliophyta, biosphere reserve of Pendjari, Atacora province, Benin. Check List 8(4): 642–661.

Assogbadjo A.E., Mensah S. & Glèlè Kakaï R. 2017. The relative importance of climatic gradient versus human disturbance in determining population structure of Afzelia africana in the Republic of Benin. Southern Forests: a Journal of Forest Science 79: 125–132.

Atanasso J.A., Mensah S., Azihou A.F., Djossa B.A., Glèlè Kakaï R. & Assogbadjo A.E. 2019. Heterospecific tree density and environmental factors affect Afzelia africana Sm. population structure in the Pendjari Biosphere Reserve, West Africa: implications for management and restoration. Tropical Conservation Science 12: 1–12.

Aubréville A. 1959. La flore forestière de la Côte d’Ivoire. Second edition. CTFT 3, Nogent-sur-Marne. Available from [accessed 19 Aug. 2021].

Baddeley A., Rubak E. & Turner R. 2015. Spatial point patterns: methodology and applications with R. CRC Press, Boca Raton.

Baddeley A. & Turner R. 2005. Spatstat: an R package for analyzing spatial point patterns. Journal of Statistical Software 12: 1–42.

Bai X., Queenborough S.A., Wang X., et al. 2012. Effects of local biotic neighbors and habitat heterogeneity on tree and shrub seedling survival in an old-growth temperate forest. Oecologia 170: 755–765.

Balima L.H., Nacoulma B.M.I., Ekué M.R.M., Kouamé F.N. & Thiombiano A. 2018. Use patterns, use values and management of Afzelia africana Sm. in Burkina Faso: implications for species domestication and sustainable conservation. Journal of Ethnobiology and Ethnomedicine 23: 14.

Barot S. & Gignoux J. 2003. Neighbourhood analysis in the savanna palm Borassus aethiopum: interplay of intraspecific competition and soil patchiness. Journal of Vegetation Science 14: 79–88.

Barot S., Gignoux J. & Menaut J.-C. 1999a. Demography of a savanna palm tree: predictions from comprehensive spatial pattern analyses. Ecology 80: 1987–2005.

Barot S., Gignoux J. & Menaut J.-C. 1999b. Seed shadows, survival and recruitment: how simple mechanisms lead to dynamics of population recruitment curves. Oikos 86(2): 320–330.

Bationo B.A., Ouedraogo S.J. & Boussim J.I. 2000. Afzelia africana Sm. (Caesalpiniaceae): étude de la prédation des graines dans une savane boisée du Burkina Faso. Bois & Forêts des Tropiques 264: 55–56.

Bationo B.A., Ouedraogo S.J. & Guinko S. 2001. Longévité des graines et contraintes à la survie des plantules d’Afzelia africana Sm. dans une savane boisée du Burkina Faso. Annals of Forest Science 58: 69–75.

Beckman N.G. & Rogers H.S. 2013. Consequences of seed dispersal for plant recruitment in tropical forests: interactions within the seedscape. Biotropica 45: 666–681.

Bonou W., Glèlè Kakaï R., Assogbadjo A., Fonton H. & Sinsin B. 2009. Characterisation of Afzelia africana Sm. habitat in the Lama forest reserve of Benin. Forest Ecology and Management 258: 1084–1092.

Chanthorn W., Caughlin T., Dechkla S. & Brockelman W.Y. 2013. The relative importance of fungal infection, conspecific density and environmental heterogeneity for seedling survival in a dominant tropical tree. Biotropica 45: 587–593.

Chu G., Wang M. & Zhang S. 2014. Spatial patterns and associations of dominant woody species in desert–oasis ecotone of South Junggar Basin, NW China. Journal of Plant Interactions 9: 738–744.

Condit R., Ashton P.S., Baker P., et al. 2000. Spatial patterns in the distribution of tropical tree species. Science 288: 1414–1418.

Connell J.H. 1971. On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees. In: Den Boer P.J. & Gradwell G.R. (eds) Dynamics of populations. Centre for Agricultural Publishing and Documentation, Wageningen.

Diggle P. 2003. Statistical analysis of spatial point patterns. Oxford University Press Inc, New York.

Evrard Q., Hardy O.J., Tagg N. & Doucet J.-L. 2019. Removal and predation of aril-covered seeds: the case of Afzelia bipindensis (Fabaceae–Detarioidae). Plant Ecology and Evolution 152(3): 460–469.

Gérard J. & Louppe D. 2011. Afzelia africana Sm. ex Pers. In: Lemmens R.H.M.J., Louppe D. & Oteng-Amoako A.A. (eds) PROTA (Plant Resources of Tropical Africa / Ressources végétales de l’Afrique tropicale), Wageningen. Available from [accessed 12 Nov. 2020].

Getzin S., Dean C., He F., Trofymow J.A., Wiegand K. & Wiegand T. 2006. Spatial patterns and competition of tree species in a Douglas fir chronosequence on Vancouver Island. Ecography 29: 671–682.

Grégoire J.M. & Simonetti D. 2010. Interannual changes of fire activity in the protected areas of the SUN network and other parks and reserves of the West and Central Africa region derived from MODIS observations. Remote Sensing 2(2): 446–463.

Guy P.R. 1989. The influence of elephants and fire on a Brachystegia-Julbernardia woodland in Zimbabwe. Journal of Tropical Ecology 5: 215–226.

Harrison R.D., Tan S., Plotkin J.B., et al. 2013. Consequences of defaunation for a tropical tree community. Ecology Letters 16: 687–694.

Hawthorne W.D. 1995. Ecological profiles of Ghanaian forest trees. Oxford Forestry Institute, University of Oxford, Oxford.

Helm C.V. & Witkowski E. 2012. Characterising wide spatial variation in population size structure of a keystone African savanna tree. Forest Ecology and Management 263: 175–188.

Hills R. 2020. Afzelia africana. The IUCN Red List of Threatened Species 2020: e.T33032A67742420.

Houehanou T.D., Assogbadjo A.E., Glèlè Kakaï R., Houinato M. & Sinsin B. 2011. Valuation of local preferred uses and traditional ecological knowledge in relation to three multipurpose tree species in Benin (West Africa). Forest Policy and Economics 13: 554–562.

Jansen P.A. & Zuidema P.A. 2001. Logging, seed dispersal by vertebrates, and natural regeneration of tropical timber trees. The cutting edge: conserving wildlife in logged tropical forests. Columbia University Press, New York.

Janzen D.H. 1970. Herbivores and the number of tree species in tropical forests. The American Naturalist 104: 501–528.

Jurinitz C.F., de Oliveira A.A. & Bruna E.M. 2013. Abiotic and biotic influences on early‐stage survival in two shade‐tolerant tree species in Brazil’s Atlantic Forest. Biotropica 45: 728–736.

Kanagaraj R., Wiegand T., Comita L.S. & Huth A. 2011. Tropical tree species assemblages in topographical habitats change in time and with life stage. Journal of Ecology 99: 1441–1452.

Kuznetsova A., Brockhoff P.B. & Christensen R.H. 2017. lmerTest package: tests in linear mixed effects models. Journal of Statistical Software 82(13): 1–26.

Le N., Thi D., Van Thinh N. & Mitlöhner R. 2016. Effect of disturbance regimes on spatial patterns of tree species in three sites in a tropical evergreen forest in Vietnam. International Journal of Forestry Research 2016: 4903749.

Ledo A., Montes F. & Condés S. 2012. Different spatial organisation strategies of woody plant species in a montane cloud forest. Acta Oecologica 38: 49–57.

Lejeune P. 2001. Arpent 2.0, logiciel de saisie de données d’arpentage–Guide d’utilisation. Note Technique Forestière 5: 1–12.

Liu J., Kuang W., Zhang Z., et al. 2014. Spatiotemporal characteristics, patterns, and causes of land-use changes in China since the late 1980s. Journal of Geographical Sciences 24: 195–210.

Martínez I., Taboada F.G., Wiegand T. & Obeso J.R. 2013. Spatial patterns of seedling-adult associations in a temperate forest community. Forest Ecology and Management 296: 74–80.

Martínez I., Wiegand T., González-Taboada F. & Obeso J.R. 2010. Spatial associations among tree species in a temperate forest community in north-western Spain. Forest Ecology and Management 260: 456–465.

McIntire E.J. & Fajardo A. 2009. Beyond description: the active and effective way to infer processes from spatial patterns. Ecology 90: 46–56.

Mensah S., Houehanou T.D., Assogbadjo A.E., Anyomi K.A., Ouedraogo A. & Glèlè Kakaï R. 2016. Latitudinal variation in the woody species diversity of Afzelia africana Sm. habitats in West Africa. Tropical Ecology 57(4): 717–726.

Mensah S., Houehanou T.D., Sogbohossou E.A., Assogbadjo A.E. & Glèlè Kakai R. 2014. Effect of human disturbance and climatic variability on the population structure of Afzelia africana Sm. ex Pers. (Fabaceae–Caesalpinioideae) at country broad-scale (Bénin, West Africa). South African Journal of Botany 95: 165–173.

Mensah S., Noulekoun F., Dimobe K., et al. 2020. Revisiting biotic and abiotic drivers of seedling establishment, natural enemies and survival in a tropical tree species in a West Africa semi-arid biosphere reserve. Journal of Environmental Management 276: 111268.

Metz M.R., Sousa W.P. & Valencia R. 2010. Widespread density‐dependent seedling mortality promotes species coexistence in a highly diverse Amazonian rain forest. Ecology 91: 3675–3685.

Muvengwi J., Mbiba M., Chikumbindi J., Ndagurwa H.G. & Mureva A. 2018. Population structure and spatial point-pattern analysis of a mono stand of Acacia polyacantha along a catena in a savanna ecosystem. Forest Ecology and Management 409: 499–508.

Nacoulma B.M.I., Lykke A.M., Traoré S., Sinsin B. & Thiombiano A. 2017. Impact of bark and foliage harvesting on fruit production of the multipurpose tree Afzelia africana in Burkina Faso (West Africa). Agroforestry Systems 91: 565–576.

Onana J. & Devineau J.-L. 2002. Afzelia africana Sm. ex Pers. dans le Nord-Cameroun. Etat actuel des peuplements et utilisation pastorale. Revue d’élevage et de médecine vétérinaire des pays tropicaux 55: 39–45.

Partey S.T., Avornyo F., Ouédraogo M. & Zougmoré R. 2018. Candidate fodder species for goat production in Northern Ghana. CCAFS, Wageningen.

Perry G., Enright N., Miller B. & Lamont B. 2009. Nearest‐neighbour interactions in species‐rich shrublands: the roles of abundance, spatial patterns and resources. Oikos 118: 161–174.

R Core Team 2017. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from [accessed 1 Jul. 2021].

Riginos C., Karande M.A., Rubenstein D.I. & Palmer T.M. 2015. Disruption of a protective ant–plant mutualism by an invasive ant increases elephant damage to savanna trees. Ecology 96: 654–661.

Ripley B.D. 1981. Spatial statistics. Hayward Wiley, New York.

Salako V.K., Kénou C., Dainou K., Assogbadjo A.E. & Glèlè Kakaï R. 2019. Impacts of land use types on spatial patterns and neighbourhood distance of the agroforestry palm Borassus aethiopum Mart. in two climatic regions in Benin, West Africa. Agroforestry Systems 93(3): 1057–1071.

Sasaki T., Konno M., Hasegawa Y., et al. 2019. Role of mycorrhizal associations in tree spatial distribution patterns based on size class in an old-growth forest. Oecologia 189(4): 971–980.

Sinsin B., Matig O.E., Assogbadjo A., Gaoué O. & Sinadouwirou T. 2004. Dendrometric characteristics as indicators of pressure of Afzelia africana Sm. dynamic changes in trees found in different climatic zones of Benin. Biodiversity & Conservation 13: 1555–1570.

Sinsin B., Sogbohossou E.A., Nobime G. & Adi M. 2008. Dénombrement aérien de la faune dans la Réserve de Biosphère de la Pendjari. GTZ, Pendjari project.

Sinsin B., Tehou A.C., Daouda I. & Saidou A. 2002. Abundance and species richness of larger mammals in Pendjari National Park in Benin. Mammalia 66(3): 369–380.

Sokpon N., Affoukou M., Amahowe I., Gandji L., Gnonlonfin L. & Sossou B. 2008. Dynamique spatio-temporelle des formations végétales du Complexe Parc National de la Pendjari, zones cynégétiques de la Pendjari et de l’Atacora. CENAGREF, Benin.

Stoyan D. & Stoyan H. 1994. Fractals, random shapes, and point fields: methods of geometrical statistics. John Wiley & Sons Inc., Chichester.

Svátek M., Rejžek M., Kvasnica J., Řepka R. & Matula R. 2018. Frequent fires control tree spatial pattern, mortality and regeneration in Argentine open woodlands. Forest Ecology and Management 408: 129–136.

Tchamba M.N. & Seme P.M. 1993. Diet and feeding behaviour of the forest elephant in the Santchou Reserve, Cameroon. African Journal of Ecology 31: 165–171.

Tehou C., Kossou E., Mensah G., Houinato M. & Sinsin B. 2012. Identification et caractérisation des formations végétales exploitées par l’éléphant Loxodonta africana dans la Réserve de Biosphère de la Pendjari au Nord-Ouest de la République du Bénin. Pachyderm 52: 36–48.

Trolliet F., Bauman D., Forget P.-M., Doucet J-L., Gillet J.-F. & Hambuckers A. 2019. How complementary are large frugivores for tree seedling recruitment? A case study in the Congo Basin. Journal of Tropical Ecology 35: 223–236.

Uriarte M., Swenson N.G., Chazdon R.L., et al. 2010. Trait similarity, shared ancestry and the structure of neighbourhood interactions in a subtropical wet forest: implications for community assembly. Ecology Letters 13: 1503–1514.

White F. 1986. The taxonomy, chorology and reproductive biology of southern African Meliaceae and Ptaeroxylaceae. Bothalia 16: 143–168.

Wiegand T. & Moloney K.A. 2004. Rings, circles, and null‐models for point pattern analysis in ecology. Oikos 104: 209–229.

Yuan Z., Wei B., Chen Y., Jia H., Wei Q. & Ye Y. 2018. How do similarities in spatial distributions and interspecific associations affect the coexistence of Quercus species in the Baotianman National Nature Reserve, Henan, China. Ecology and Evolution 8: 2580–2593.

Zhang Z.-H., Hu G., Zhu J.-D., Luo D.-H. & Ni J. 2010. Spatial patterns and interspecific associations of dominant tree species in two old-growth karst forests, SW China. Ecological Research 25: 1151–1160.

Zhu Y., Queenborough S., Condit R., Hubbell S., Ma K. & Comita L. 2018. Density‐dependent survival varies with species life‐history strategy in a tropical forest. Ecology Letters 21: 506–515.

Creative Commons License

This work is licensed under a Creative Commons Attribution 4.0 International License.