, Volume 28, Issue 6, pp 1777–1791 | Cite as

Does phenology distinguish bitter and sweet African bush mango trees (Irvingia spp., Irvingiaceae)?

  • Romaric Vihotogbé
  • Ronald G. van den Berg
  • Frans Bongers
  • Brice Sinsin
  • Marc S. M. Sosef
Original Paper


Key message

This phenological analysis of bitter and sweet bush mango trees is part of their biosystematics. It supports the species distinction hypothesis postulated by Harris (Bull J Bot Nat Belg 65(1–2):143–196, 1996 ) and Lowe et al. (Mol Ecol 9:831–841, 2000 ).


African Bush Mango trees are priority food trees in Sub-Saharan Africa. The unclear distinction between bitter and sweet fruited trees is still subject to taxonomic debate. This hinders their effective use and conservation programmes. This study investigates differences in phenological behaviour between bitter and sweet fruited populations and their taxonomic implications. Monthly phenological description data on seven populations of bitter or sweet bush mangos across Benin and Togo were used to assess within and between mango type phenological diversity, to discriminate bitter and sweet trees and to evaluate their responses to environmental factors. The phenological states differentiating bitter and sweet trees were identified and individual trees were classified based on the discriminating phenological characters. Finally, phenological variation was analyzed with time of the year, soil type, type of bush mango tree, and climatic zone. Phenological diversity varies significantly among populations. Bitter and sweet trees have consistently different phenological states. Bitter trees have a lower phenological diversity for all phenological phases throughout the year compared to sweet trees, possibly due to their limited distribution range in the study area. The tree types also differ in their reproductive responses to environmental factors, but did not respond differently to soils. These results support the hypothesis that bitter and sweet trees represent different taxa and we suggest for efficient conservation purpose to consider them as different species.


Adaptation Ecology Domestication Irvingia Phenological states Species distinction 


Author contribution statement

Romaric Vihotogbé proposed the research idea, led the field works, analyzed the data, proposed the manuscript and improved it based on the co-authors’ comments. Ronald G. Van Den Berg influenced the field work design and was thoroughly involved in the statistical analysis. Frans Bongers, Brice Sinsin and Marc Sosef validated the research idea and methodology end provided specific and significant comments on the manuscript. In addition Brice Sinsin was involved in planning and execution of field works. After the first submission, the manuscript was thoroughly revisited by each of the co-authors based on reviewers’ comments and Romaric Vihotogbé coordinated this activity. We all agree on this version being re-submitted.


The Dutch Organization for International Cooperation in Higher Education (NUFFIC, the Netherlands) funded this research and the International Foundation for Science provided significant financial support for fieldworks (IFS/Grant No: D/4672-1, Stockholm, Sweden for Romaric Vihotogbé). We are grateful to those two institutions and to their donors. We are also grateful to all local farmers that freely permitted us to study their trees. Prof. Dr. Ir. Glèlè Kakaï Romain (FSA/UAC, Benin) advised during sampling design and Wilfried Bonou (MSc: (FSA/UAC, Benin) provided assistance in the statistical analyses and we thank them in a very particular way. Lastly, the “Unanimous Reviewer 1” of this paper provided important comments and direct contributions to the paper and we are thankful to him.

Conflict of interest

I (the corresponding author), confirm that there is no conflict of interest of any kind (commercial, scientific/co-authorship) related to this article. Also, this is an original research paper that has never been published or even submitted for publication to any other journal. All the co-authors agreed on this version being re-submitted to TREES.


  1. Adomou AC, Sinsin B, van Der Maesen LJG (2006) Phytosociological and chorological approaches to phytogeography: a meso-scale study in Benin. Syst Geo Plants 76:155–178Google Scholar
  2. Anegbeh PO, Usoro C, Ukafor V, Tchoundjeu Z, Leakey RRB, Schreckenberg K (2003) Domestication of Irvingia gabonensis: 3. Phenotypic variation of fruits and kernels in a Nigerian village. Agrofor Syst 58:213–218CrossRefGoogle Scholar
  3. Asaah EK, Tchoundjeu Z, Atangana AR (2003) Cultivation and conservation status of Irvingiawombolu in humid lowland forest of Cameroon. Food Agric Environ 1(3–4):251–256Google Scholar
  4. Atangana AR, Tchoundjeu Z, Fondoun J-M, Asaah E, Ndoumbe M, Leakey RRB (2001) Domestication of Irvingia gabonensis: 1. Phenotypic variation in fruit and kernels in two populations from Cameroon. Agrofor Syst 53:55–64CrossRefGoogle Scholar
  5. Atangana AR, Ukafor V, Anegbeh PO, Asaah E, Tchoundjeu Z, Usoro C, Fondoun J-M, Ndoumbe M, Leakey RRB (2002) Domestication of Irvingia gabonensis: 2. The selection of multiple traits for potential cultivars from Cameroon and Nigeria. Agrofor Syst 55:221–229CrossRefGoogle Scholar
  6. Bidogeza JC, Berentsen BPM, De Graaff J, Oude Lansink AGJM (2009) A typology of farm households for the Umutara Province in Rwanda. Food Sec 1:321–335CrossRefGoogle Scholar
  7. Bloom SA (1981) Similarity indices in community studies: potential pitfalls. Eco Prog Ser 5:125–128CrossRefGoogle Scholar
  8. Booth AH (1957) The Niger, the Volta and the Dahomey Gap as geographic barriers. Evolution 12(1):48–62CrossRefGoogle Scholar
  9. Chuine I (2010) Why does phenology drive species distribution? Philo Trans Roy Soc 365:3149–3160CrossRefGoogle Scholar
  10. Chuine I, Beaubien EG (2008) Phenology is a major determinant of tree species range. Ecol Let 4(5):500–510Google Scholar
  11. Doi H, Takahash M, Atano KI (2010) Genetic diversity increases regional variation in phenological dates in response to climate change. Glob Change Biol 16:373–379CrossRefGoogle Scholar
  12. Dolor D (2011) Effect of propagation media on the germination and seedling performance of Irvingiawombolu (Vermoesen). Am J Bot. doi: 10.5251/ajbms.2011. Google Scholar
  13. Ekpe OO, Umoh IB, Eka OU (2007) Effect of a typical rural processing method on the proximate composition and amino acid profile of bush mango seeds (Irvingia gabonensis). Afr J Agric Food Nut Dev 7(1):1–12Google Scholar
  14. Giresse P (2008) Tropical and sub-tropical west Africa. Marine and continental changes during the late quaternary. Elsevier, United KingdomGoogle Scholar
  15. Goulart MF, Filho JPL, Lovato MB (2005) Phenological variation within and among populations of Plathymeniareticulata in Brazilian Cerrado, the Atlantic forest and transitional sites. Ann Bot 96:445–455PubMedCrossRefGoogle Scholar
  16. Hammer Ø, Harper DAT, Ryan PD (2001) PAST: paleontological statistics software package for education and data analysis. Palaeo Electro 4:1–9Google Scholar
  17. Harris DJ (1996) A revision of the Irvingiaceae in Africa. Bull J Bot Nat Belg 65(1–2):143–196Google Scholar
  18. Hodghin T, Brown AHD, van Hintumth JL, Morales EAV (1995) Core collection of plant genetic resources. IPGRI, Wiley-Sayce Publication, LondonGoogle Scholar
  19. Jose S (2012) Agroforestry for conserving and enhancing biodiversity. Agrofor Syst 85:1–8CrossRefGoogle Scholar
  20. Kang BT, Akinnifesi FK, Ladipo DO (1994) Performance of selected woody agroforestry species grown on an Alfisol and an Ultisol in the humid lowland of West Africa, and their effects on soil properties. J Trop For Sci 7(2):303–312Google Scholar
  21. Kengni E, Kengue J, Ebenezer EBK, Tabuna H (2011) Irvingiagabonensis, Irvingiawombolu, bush mango. Conservation and sustainable use of genetic resources of priority food tree species in sub-Saharan Africa. Bioversity International, RomeGoogle Scholar
  22. Ladipo DO, Fondoun JM, Ganga N (1996) Domestication of the bush mango (Irvingiaspp): some exploitable intraspecific variations in West and Central Africa. In: Leakey RRB, Temu AB, Melnyk M, Vantomme P (eds) Non-wood forest products no. 9: domestication and commercialization of non-timber forest products for agroforestry. FAO, Rome, pp 193–205Google Scholar
  23. Ladizinsky G (1998) Plant evolution under domestication. Kluwer Academic Publishers, Dordrecht (ISBN 0 412 802105, 247)CrossRefGoogle Scholar
  24. Leakey RRB (2012) Living with the trees of life. Towards the transformation of tropical Agriculture. CABI Publications, UK. ISBN 978-1-78064-099-0CrossRefGoogle Scholar
  25. Leakey RRB, Simons AJ (1998) The domestication and commercialization of indigenous trees in agroforestry for the alleviation of poverty. Agrofor Syst 38:165–176CrossRefGoogle Scholar
  26. Leakey RRB, Tchoundjeu Z (2001) Diversification of tree crops: domestication of companion crops for poverty reduction and environmental services. Exp Agric 37:279–296CrossRefGoogle Scholar
  27. Leakey RRB, Fondoun J-M, Atangana A, Tchoundjeu Z (2000) Quantitative descriptors of variation in the fruits and seeds of Irvingiagabonensis. Agrofor Syst 50:47–58CrossRefGoogle Scholar
  28. Leakey RRB, Tchoundjeu Z, Schreckenberg K, Schackleton SE, CM Schackleton (2005a) Agroforestry trees products (AFTPs): targeting poverty reduction and enhanced livelihoods. Inter J Agric Sust 3:1–23CrossRefGoogle Scholar
  29. Leakey RRB, Greenwell P, Hall MN, Atangana AR, Usoro C, Anegbeh PO, Fondoun J-M, Tchoundjeu Z (2005b) Domestication of Irvingia gabonensis: 4. Tree-to-tree variation in food-thickening properties and in fat and protein contents of Dika Nut. Food Chem 90:365–378CrossRefGoogle Scholar
  30. Lesley A, Brown N (2004) Bush Mango (Irvingia gabonensis and I. wombolu). In: Clark LE, Sunderland TCH (eds) Key non-timber forest products of central Africa: state of the knowledge. SD Publication Series, USAID, USA, pp 15–35Google Scholar
  31. Lowe AJA, Gillies CM, Wilson J, Dawson IK (2000) Conservation genetics of bush mango from central/west Africa: implications from random amplified polymorphic DNA analysis. Mol Ecol 9:831–841PubMedCrossRefGoogle Scholar
  32. Maley J (1996) The African rain forest—main characteristics of changes in vegetation and climate from the Upper Cretaceous to the Quaternary. Proc Roy Soc Edinb 104b:31–73Google Scholar
  33. Mohammadi SA, Prasanna BM (2003) Review and interpretation. Analysis of genetic diversity in crop plants—salient statistical tools and considerations. Crop Sci 43:1235–1248CrossRefGoogle Scholar
  34. Munguia-Roas M, Ollerton J, Parra-Table V, De-Nova JA (2011) Meta-analysis of phenotypic selection on flowering phenology suggests that early flowering plants are favoured. Ecol Let. doi: 10.1111/j.1461-0248.2011.01601.x Google Scholar
  35. National Research Council (2006) Lost crops of Africa; development, security, and cooperation policy and global affairs II, vegetables. The National Academies Press, WashingtonGoogle Scholar
  36. Newstrom LE, Frankie GW, Baker HG (1994) A new classification for plant phenology based on flowering patterns in lowland tropical rain forest trees at La Selva, Costa Rica. Biotropica 26(2):141–159CrossRefGoogle Scholar
  37. Nkwatoh AF, Labode P, Iyassa SM, Nkwatoh FW (2010) Evaluation of Irvingia (I. gabonensis and I. wombuluBail.) trade; a major non-timber forest product in the Ejagham forest reserve of south west Cameroon. Ethiop J Env Stud Manag 3(1):69–77CrossRefGoogle Scholar
  38. Nord EA, Lynch JP (2001) Plant phenology: a critical controller of soil resource acquisition. J Exp Bot 18:1–11Google Scholar
  39. Nya PJ, Omokaro DN, Nkang AE (2006) The effect of storage temperature and humidity on germination of Irvingia gabonensis var. excelsa. Trop Sci 46(2):64–69CrossRefGoogle Scholar
  40. Nzekwe U, Onyekwelu SSC, Umeh VC (2002) Improving the germination of Irvingia gabonensis var. excelsa seeds. Niger J Hort Sci 7:48–52Google Scholar
  41. Ogunsina BS, Koya OA, Adeosun OO (2008) “A table mounted device for cracking Dika nut (Irvingiagabonensis)” Agric Eng Int CIGR E J. Accessed 12 March 2012
  42. Okafor JC (1975) Varietal delimitation in Irvingiagabonensis (Irvingiaceae). Bul J Bot Nat Belg 45:211–221CrossRefGoogle Scholar
  43. Okafor JC, Fernandes ECM (1987) Compound farms of South-eastern Nigeria: a predominant agroforestry homegarden system with crops and small livestock. Agrofor Syst 5:153–168CrossRefGoogle Scholar
  44. Orwa C, Mutua A, Kindt R, Jamnadass R, Simons A (2009) Agroforestree database: a tree reference and selection guide version 4.0 (
  45. Ruml M, Vulie T (2005) Importance of phenological observation and prediction in agriculture. J Agric 50(2):217–225Google Scholar
  46. Salzmann U, Hoelzmann P (2005) The Dahomey Gap: an abrupt climatically induced rain forest fragmentation in West Africa during the late Holocene. Holocene 15(2):190–199CrossRefGoogle Scholar
  47. SAS Institute Inc (1999) SAS online doc, version 8. Cary, North Carolina, USAGoogle Scholar
  48. Shiembo PN, Newton AC, Leakey RRB (1996) Vegetation propagation of Irvingia gabonensis, a West African fruit tree. For Ecol Manag 87:185–192CrossRefGoogle Scholar
  49. Statsoft (2001) STATISTICA for Windows, version 6. 2300. StatSoftInc, Tulsa. Sub Saharienne et l’Europe. FAO, AccraGoogle Scholar
  50. Tabuna H (2000) Evaluation des échanges des produits forestiers non ligneux entre l’Afrique subsaharienne et l’Europe. FAO, Bureau Régional pour l’AfriqueGoogle Scholar
  51. Tchoundjeu Z, Kengue J, Leakey RRB (2002) Domestication of Dacryodesedulis: state of the art. For Tre Liv 12:3–13CrossRefGoogle Scholar
  52. Turling N (1991) Application of the ideotypes concept in breeding for higher yield in the oil seed brassicas. Crop Field Res 26:201–219CrossRefGoogle Scholar
  53. Ude GN, Dimkpa CO, Anegbeh PO, Shaibu AA, Tenkouano A, Pillay M, Tchoundjeu Z (2006) Analysis of genetic diversity in accessions of Irvingiagabonensis (Aubry-Lecomte ex O’Rorke) Baill. Afric J Biotech 5(3):219–223Google Scholar
  54. Van Dijk JFW (1999) An assessment of non-wood forest product resources for the development of sustainable commercial extraction. In: Sunderland TCH, Clark LE, Vantomme P (eds) Non-wood forest products of Central Africa: current research issues and prospects for conservation and development. FAO Rome, Italy, pp 37–50Google Scholar
  55. Vihotogbé R, Houessou GL, Ponette Q, Codjia JTC, Le Boulengé E (2008) Ethnobotanic knowledge and endogenous conservation of Irvingiagabonensis (Aubry- Lecomte) Baill. in. Benin. Indilinga 6(2):196–209Google Scholar
  56. Vihotogbé R, van Den Berg RG, Sosef MSM (2013) Morphological Characterization of African Bush Mango Trees (Irvingia species) in the Dahomey Gap (West Africa). Gen Res Crop Evol 60:1597–1614CrossRefGoogle Scholar
  57. Vihotogbé R, GlèlèKakaï R, Bongers F, van Andel T, van Den Berg RG, Sinsin B, Sosef MSM (2014a) Impacts of the diversity of traditional uses and potential economic value on food tree species conservation status: case study of African bush mango trees (Irvingiaceae) in the Dahomey Gap (West Africa). Pl Eco Evol 147(1):109–125CrossRefGoogle Scholar
  58. Vihotogbé R, van den Berg RG, Missihoun AA, Sinsin B, Sosef MSM (2014b) Genetic diversity and difference within and between bitter and sweet African bush mango trees (Irvingia spp., Irvingiaceae) in West and Central Africa. Afric J BiotechGoogle Scholar
  59. Wolkovich EM, Cleland EE (2011) The phenology of plant invasions: a community ecology perspective. Front Ecol Env 9(5):287–294CrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2014

Authors and Affiliations

  • Romaric Vihotogbé
    • 1
    • 2
  • Ronald G. van den Berg
    • 1
  • Frans Bongers
    • 3
  • Brice Sinsin
    • 2
  • Marc S. M. Sosef
    • 1
    • 4
  1. 1.Biosystematics GroupWageningen UniversityWageningenThe Netherlands
  2. 2.Laboratory of Applied Ecology, Faculty of Agronomic SciencesUniversity of Abomey-CalaviCotonouBenin
  3. 3.Forest Ecology and Forest Management GroupWageningen UniversityWageningenThe Netherlands
  4. 4.Botanic Garden MeiseMeiseBelgium

Personalised recommendations