Evolutionary Biology & Ecologyebe-Welcome.html
ebe homeebe-Welcome.html
phone: +32 (0)2 650 65 85

e-mail: Olivier.Hardy@ulb.be

Research interests

The team develops research on plant biodiversity, mostly in African rainforests:

  1. Phylogeographic studies of plant species to reconstruct past population changes and test whether past climate fluctuations have lead consistent signatures across species.

  2. Studies on seed and pollen dispersal patterns in timber tree species to assess natural regeneration processes and provide guidelines for sustainable tropical forest management.

  3. Development of DNA (meta)barcoding approaches to identify tree species, identify the diet of herbivores and develop tools for timber tracking.

  4. Molecular systematics and taxonomy of some complex plant genera.

  5. Studies of forest tree community diversity patterns from local to continental scales to assess the relative impact of habitat filtering, drift-dispersal processes and historical events on the distribution of species and the (phylogenetic) structure of species assemblages.

  6. Transcriptomic response under drought stress in African tree species

  7. Theoretical work in population genetics and community ecology developing new approaches to infer processes from patterns.

  8. Development of new software for population genetics or community data analyses (SPAGeDi, SPACoDi, AutocorQ, BiodivR, TOROCOR).




phone: +32 (0)2 650 45 11     e-mail: ekaymak@ulb.ac.be

Post-doctoral researchers

Achille BIWOLE

phone: +32 (0)2 650 45 14     e-mail: achille.biwole@gmail.com


phone: +32 (0)2 650 45 14     e-mail: simon.jansen@bfw.gv.at


phone: +32 (0)2 650 45 14     e-mail: jeremy.migliore@ulb.ac.be

PhD students


phone: +32 (0)2 650 22 67     e-mail: d.oscar.ahossou@gmail.com


phone: +32 (0)2 650 45 11     e-mail: Claire.Baudoux@ulb.ac.be

Arthur BOOM

phone: +32 (0)2 650 45 11     e-mail: arthboom@ulb.ac.be


phone: +32 (0)2 650 45 11     e-mail: Katarina.Matvijev@ulb.ac.be


phone: +32 (0)2 650 45 14     e-mail: fmonthekameni@yahoo.fr

Nicolas TEXIER

e-mail: Nicolas.Texier@ulb.ac.be


e-mail: samuelvdabeele@hotmail.com

Scientific collaborator


e-mail: mheuertz@ulb.ac.be


e-mail: bsenterre@gmail.com

Bonaventure SONKE

e-mail: bsonke_1999@yahoo.com


e-mail: Jerome.Duminil@ulb.ac.be

Previous post-doctoral researchers

Gilles DAUBY

Gabriel DEBOUT


Guillaume KOFFI

Alexandra LEY



Mohamed NEJI




Previous PhD students

Gilles DAUBY



Eben-Ezer EWEDJE

Guillaume KOFFI


Félicien TOSSO


Previous MSc students


Arthur BOOM










Romain JANS








Abubakari SAID






Publications (write an email to ohardy@ulb.ac.be for reprints)


Demenou B, Doucet J-L, Hardy OJ (2018) History of the fragmentation of the African rain forest in the Dahomey Gap: insight from the demographic history of Terminalia superba. Heredity 120:547–561. doi: 10.1038/s41437-017-0035-0

Droissart V, Dauby G, Hardy OJ, Deblauwe V, Harris DJ, Janssens S, Mackinder BA, Overgaard A-B, Sonké B, Sosef MSM, Stévart T, Svenning J-C, Wieringa JJ, Couvreur TLP (2018) Beyond trees: biogeographical regionalization of tropical Africa. Journal of Biogeography 45:1153–1167. doi: 10.1111/jbi.13190

Kasongo Yakusu E, Monthe FS, Bourland N, Hardy OJ, Louppe D, Bola Mbele Lokanda F, Hubau W, Kahindo Muhongya J-M, Van Den Bulcke J, Van Acker J, Beeckman H (2018) Le genre Entandrophragma (Meliaceae) : taxonomie et écologie d’arbres africains d’intérêt économique (synthèse bibliographique). Biotechnol. Agron. Soc. Environ. 22: 113-127.

Migliore J, Kaymak E, Mariac C, Couvreur TLP, Lissambou B-J, Piñeiro R, Hardy OJ (2018) Pre-Pleistocene origin of phylogeographical breaks in African rain forest trees: new insights from Greenwayodendron (Annonaceae) phylogenomics. Journal of Biogeography (in press)

Monthe FK, Duminil J, Kasongo Yakusu E, Beeckman H, Bourland N, Doucet J-L, Sosef MSM, Hardy OJ (2018) The African timber tree Entandrophragma congoense (Pierre ex De Wild.) A.Chev. is morphologically and genetically distinct from Entandrophragma angolense (Welw.) C.DC. Tree Genetics & Genomes 14: 66. doi: 10.1007/s11295-018-1277-6

Neji M, Serbaji MM, Hardy OJ, Chaieb M (2018) Floristic diversity and vegetation patterns along disturbance gradient in arid coasts in southern Mediterranean: Case of the Gulf of Gabès, southern Tunisia. Arid Land Research and Management 32:291-315.

Simo-Droissart M, Plunkett GM, Droissart V, Edwards MB, Farminhão JNM, Ječmenica V, D'haijère T, Lowry PPII, Sonké B, Micheneau C, Carlsward BS, Azandi L, Verlynde S, Hardy OJ, Martos F, Bytebier B, Fischer E, Stévart T (2018) New phylogenetic insights toward developing a natural generic classification of African angraecoid orchids (Vandeae, Orchidaceae). Molecular Phylogenetics and Evolution 126:241-249. doi: 10.1016/j.ympev.2018.04.021

Tosso F, Cherchye G, Hardy OJ, Daïnou K, Lognay G, Tagg N, Haurez B, Souza A, Heuskin S, Doucet J-L (2017) Characterization of animal communities involved in seed dispersal and predation of Guibourtia tessmannii (Harms) J.Léonard, a species newly listed on Appendix II of CITES. African Journal of Ecology. in press. doi: 10.1111/aje.12480

Tosso F, Hardy OJ, Doucet J-L, Daïnou K, Kaymak E, Migliore J (2018). Evolution in the Amphi-Atlantic tropical genus Guibourtia (Fabaceae, Detarioideae), combining NGS phylogeny and morphology. Molecular Phylogenetics and Evolution 120: 83–93. doi: 10.1016/j.ympev.2017.11.026

Vanden Abeele S, Hardy OJ, Janssens SB (2018) Isolation of microsatellite loci in the African tree species Staudtia kamerunensis (Myristicaceae) using high-throughput sequencing. Molecular Biology Reports 45: 1539–1544. doi: 10.1007/s11033-018-4239-z

Vleminckx J, Bauman D, Hardy OJ, Drouet T (2018) Testing and interpreting the shared space-environment fraction in variation partitioning analyses of ecological data. Oikos (in press)


Blanc-Jolivet C, Kersten B, Daïnou K, Hardy O, Guichoux E, Delcamp A, Degen B (2017) Development of nuclear SNP markers for genetic tracking of Iroko, Milicia excelsa and Milicia regia. Conservation Genet Resources 9:531-533. doi:10.1007/s12686-017-0716-2

Daïnou K, Flot JF, Degen B, Blanc-Jolivet C, Doucet J-L, Lassois L, Hardy OJ (2017). DNA taxonomy in the timber genus Milicia: evidence of unidirectional introgression in the West African contact zone. Tree Genetics & Genomes 13: 90. doi:10.1007/s11295-017-1174-4

Demenou BB,  Hardy OJ (2017) Development, Characterization, and Cross-Amplification of Microsatellite Markers in the Understudied African Genus Anthonotha (Fabaceae). Applications in Plant Sciences 5(1): 1600120. doi:10.3732/apps.1600120

Donkpegan ASL, Doucet JL, Migliore J, Duminil J, Dainou K, Piñeiro R, Wieringa JJ, Champluvier D, Hardy OJ (2017) Evolution in African tropical trees displaying ploidy-habitat association: the genus Afzelia (Leguminosae). Molecular Phylogenetics and Evolution 107:270–281. doi: 10.1016/j.ympev.2016.11.004

Ewédjè EBK, Ahanchédé A, Hardy OJ (2017) Breeding system, gene dispersal and small-scale spatial genetic structure of a threatened food tree species, Pentadesma butyracea (Clusiaceae) in Benin. Conservation Genetics 18(4):799-811. doi:10.1007/s10592-017-0928-8

Ikabanga DU, Stévart T, Koffi GK, Monthe FK, Nzigou Doubindou EC, Dauby G, Souza A, M’batchi B, Hardy OJ (2017) Combining morphology and population genetic analysis uncover species delimitation in the widespread African tree genus Santiria (Burseraceae). Phytotaxa 321: 166–180. doi:10.11646/phytotaxa.321.2.2

Ley AC, Hardy OJ (2017) Hybridization and asymmetric introgression after secondary contact in two tropical African climber species, Haumania danckelmaniana and Haumania liebrechtsiana (Marantaceae). International Journal of Plant Sciences 178:421–430. doi : 10.1086/691628

Ley AC, Heuertz M, Hardy OJ (2017) The evolutionary history of central African rain forest plants: phylogeographical insights from sister species in the climber genus Haumania (Marantaceae). Journal of Biogeography 44:308–321. doi:10.1111/jbi.12902

Libalah MB, Droissart V, Sonké B, Hardy OJ, Drouet T, Pescador DS, Kenfack D, Thomas DW, Chuyong GB, Couteron P (2017) Shift in functional traits along soil fertility gradient reflects non-random community assembly in a tropical African rainforest. Plant Ecology and Evolution 150: 265–278. doi.org/10.5091/plecevo.2017.1318

Monthe FK, Hardy OJ, Doucet JL, Loo J, Duminil J (2017) Extensive seed and pollen dispersal and assortative mating in the rain forest tree Entandrophragma cylindricum (Meliaceae) inferred from indirect and direct analyses. Molecular Ecology 26: 5279–5291. doi:10.1111/mec.14241

Monthe FS, Duminil J, Tosso F, Migliore J, Hardy OJ (2017) Characterization of microsatellite markers in two African exploited trees Entandrophragma candollei and E. utile (Meliaceae). Applications in Plant Sciences 5(2): 1600130. doi:10.3732/apps.1600130

Olsson S, Seoane-Zonjic P, Bautista R, Claros MG, González-Martínez SC, Scotti I, Scotti-Saintagne C, Hardy OJ, Heuertz M (2017) Development of genomic tools in a widespread tropical tree, Symphonia globulifera L.f.: A new low-coverage draft genome, SNP and SSR markers. Molecular Ecology Resources 17(4):614-30. doi: 10.1111/1755-0998.12605

Piñeiro R, Dauby G, Kaymak E, Hardy OJ (2017) Pleistocene population expansions of shade-tolerant trees indicate fragmentation of the African rainforest during the Ice Ages. Proc. R. Soc. B 284:20171800. doi:10.1098/rspb.2017.1800

Sosef MSM, Dauby G, Blach-Overgaard A, van der Burgt X, Catarino L, Damen T, Deblauwe V, Dessein S, Dransfield J, Droissart V, Duarte MC, Engledow H, Fadeur G, Figueira R, Gereau RE, Hardy OJ, Harris DJ, de Heij J, Janssens S, Klomberg Y, Ley AC, Mackinder BA, Meerts P, van de Poel JL, Sonké B, Stévart T, Stoffelen P, Svenning J-, Sepulchre P, Zaiss R, Wieringa JJ, Couvreur TLP (2017) Exploring the floristic diversity of tropical Africa. BMC Biology 15(1):15. doi: 10.1186/s12915-017-0356-8

Tosso F., Doucet J-L, Migliore J, Daïnou K, Kaymak E, Kameni FSM, Hardy OJ (2017) Characterization of microsatellite markers in the African tropical tree species Guibourtia ehie (Fabaceae, Detarioideae). Applications in Plant Sciences 5 (7):1700023. doi:10.3732/apps.1700023

Vleminckx J, Doucet J-L, Morin-Rivat J, Biwolé A, Bauman D, Hardy OJ, Fayolle A, Gillet J-F, Daïnou K, Gorel A, Drouet T (2017) The influence of spatially structured soil properties on tree community assemblages at a landscape scale in the tropical forests of southern Cameroon. Journal of Ecology 105(2):354–366. doi: 10.1111/1365-2745.12707


Abdou R, Bakasso Y, Saadou M, Baudoin JP, Hardy OJ (2016) Genetic diversity of Niger onions (Allium cepa L.) assessed by simple sequence repeat markers (SSR). Acta Horticultura 1143:77-90. doi:10.17660/ActaHortic.2016.1143.12

Daïnou K., Blanc-Jolivet CC, Degen B, Kimani PP, Ndiade-Bourobou D, Donkpegan ASL, Tosso F, Kaymak E, Bourland N, Doucet JL, Hardy OJ (2016). Revealing hidden species diversity in closely related species using nuclear SNPs, SSRs and DNA sequences - a case study in the tree genus Milicia. BMC Evolutionary Biology 16(1): 1-15. doi:10.1186/s12862-016-0831-9

Dauby G, Zaiss R, Blach-Overgaard A, Catarino L, Damen T, Deblauwe V, Dessein S, Dransfield J, Droissart V, Duarte MC, Engledow H, Fadeur G, Figueira R, Gereau RE, Hardy OJ, Harris DJ, de Heij J, Janssens S, Klomberg Y, Ley AC, Mackinder BA, Meerts P, van de Poel JL, Sonké B, Sosef MSM, Stévart T, Stoffelen  P, Svenning J-C, Sepulchre P, van der Burgt X, Wieringa JJ, Couvreur TLP (2016) RAINBIO: a mega-database of tropical African vascular plants distributions. PhytoKeys 74: 1–18. doi: 10.3897/phytokeys.74.9723

Demenou B, Piñeiro R, Hardy OJ (2016) Origin and history of the Dahomey Gap separating West and Central African rain forests: insights from the phylogeography of the legume tree Distemonanthus benthamianus. Journal of Biogeography 43:1020–1031. doi: 10.1111/jbi.12688

Duminil J, Mendene Abessolo DT, Ndiade Bourobou D, Doucet J-L, Loo J, Hardy OJ (2016) High selfing rate, limited pollen dispersal and inbreeding depression in the emblematic African rain forest tree Baillonella toxisperma – Management implications. Forest Ecology and Management 379: 20–29.

Fourdrilis S, Mardulyn P, Hardy OJ, Jordaens K, Manuel de Frias Martins A, Backeljau T (2016) Mitochondrial DNA hyperdiversity and its potential causes in the marine periwinkle Melarhaphe neritoides (Mollusca: Gastropoda). PeerJ 4:e2549. doi: 10.7717/peerj.2549

Ley AC, Hardy OJ (2016) Polymorphic microsatellite loci for Haumania danckelmaniana and transferability into H. liebrechtsiana (Marantaceae). Applications in Plant Sciences 4:1500116. doi : 10.3732/apps.1500116

Ley AC, Hardy OJ (2016). Spatially limited clonality and gene dispersal in a characteristic liana of Central African rain forests: Haumania danckelmaniana (Marantaceae). Biotropica 48:618–627. doi: 10.1111/btp.12341

Morin-Rivat J, Biwolé 1, Gorel A-P, Vleminckx J, Gillet J-F, Bourland N, Hardy OJ, Livingstone Smith A, Daïnou K, Dedry L, Beeckman H,  Doucet J-L (2016) High spatial resolution of late-Holocene human activities in the moist forests of central Africa using soil charcoal and charred botanical remains. The Holocene 26:1954–1967. doi: 10.1177/0959683616646184

Simo-Droissart M, Sonké B, Droissart V, Micheneau C, Lowry II PP, Hardy OJ, Plunkett GM, Stévart T (2016) Morphometrics and molecular phylogenetics of Angraecum section Dolabrifolia (Orchidaceae, Angraecinae). Plant Systematics and Evolution 302(8):1027-1045. doi: 10.1007/s00606-016-1315-5

Tosso F, Doucet J-L, Kaymak E, Daïnou K, Duminil J, Hardy OJ (2016) Microsatellite development in Guibourtia tessmannii (Harms) J.Leonard and cross-amplification on congeneric species uncover a polyploid complex. Applications in Plant Sciences 4(7): 1600029.


Anthony NM, Atteke C, Bruford  MW, Dallmeier F, Freedman A, Hardy OJ, Ibrahim B, Jeffery K, Johnson M, Lahm SA, Lepengue N, Lowenstein JH, Maisels F, Mboumba J-F, Mickala P, Morgan K, Ntie S, Smith TB, Sullivan JP, Verheyen E, Gonder MK (2015) Evolution and Conservation of Central African Biodiversity: Priorities for Future Research and Education in the Congo Basin and Gulf of Guinea. Biotropica 47:6–17.  doi : 10.1111/btp.12188

Biwolé AB, Morin-Rivat J, Fayolle A, Bitondo D, Dedry L, Dainou K, Hardy OJ, Doucet J-L (2015) New data on the recent history of the littoral forests of southern Cameroon: an insight into the role of historical human disturbances on the current forest composition. Plant Ecology and Evolution 148: 19–28. doi:10.5091/plecevo.2015.1011

Biwolé ABA., Daïnou K, Fayolle A, Hardy OJ, Brostaux Y, Coste SS, Delion SS, Betti J, Doucet JL (2015). Light response of seedlings of a central African timber tree species, Lophira alata (Ochnaceae), and the definition of light requirements. Biotropica 47(6): 681-688. doi:10.1111/btp.12258

Demenou BB, Migliore J, Tosso J, Kaymak E, Hardy OJ (2015) Development and characterization of 14 microsatellite markers in the African deciduous tree Terminalia superba (Combretaceae). Applications in Plant Sciences 3(12):1500070. doi: 10.3732/apps.1500070

Donkpegan ASL, Doucet J-L, Dainou K, Hardy OJ (2015) Microsatellite Development and Flow Cytometry in the African Tree Genus Afzelia (Fabaceae, Caesalpinioideae) Reveal a Polyploid Complex. Applications in Plant Sciences 3(1): 1400097. doi: 10.3732/apps.1400097

Duminil J, Daïnou K, Kombi Kaviriri D, Gillet P, Loo J, Doucet J-L, Hardy OJ (2015) Relationships between population density, fine-scale genetic structure, mating system and pollen dispersal in a timber tree from African rainforests. Heredity 116, 295–303 doi: 10.1038/hdy.2015.101

Duminil J, Mona S, Mardulyn P, Doumenge C, Walmacq F, Doucet J-L, Hardy OJ (2015) Late Pleistocene molecular dating of past population fragmentation and demographic changes in African rain forest tree species supports the forest refuge hypothesis. Journal of Biogeography 42:1443–1454. doi:10.1111/jbi.12510

Ewedje E-EBK, Ahanchédé A, Hardy OJ, Ley AC (2015) Reproductive biology of Pentadesma butyracea (Clusiaceae), source of a valuable non timber forest product in Benin. Plant Ecology and Evolution 148: 213–228.

Hardy OJ (2015) Population genetics of autopolyploids under a mixed mating model and the estimation of selfing rate. Molecular Ecology Resources 16: 103–117. doi: 10.1111/1755-0998.12431

Piñeiro R, Micheneau C, Dauby G, Hardy OJ (2015). Isolation, characterization and cross-species amplification of nuclear microsatellites in the African tree genus Greenwayodendron (Annonaceae). Journal of Tropical Forest Science 28(2):121-131.

Piñeiro R, Staquet A, Hardy OJ (2015) Isolation of nuclear microsatellites in the African timber tree Lophira alata and cross-amplification in L. lanceolata. Applications in Plant Sciences 3 (10): 1500056.

Tosso F, Daïnou K, Hardy OJ, Sinsin B, Doucet JL (2015). Le genre Guibourtia Benn., un taxon à haute valeur commerciale et sociétale (synthèse bibliographique). Biotechnol. Agron. Soc. Environ. 19: 71-88.


Dainou K, Mahy G, Duminil J, Dick C, Doucet J-L, Donkpégan A, Pluijgers M, Sinsin B, Lejeune P, Hardy OJ (2014) Speciation slowing down in widespread and long-living tree taxa: insights from the tropical timber tree genus Milicia (Moraceae). Heredity 113: 74-85. doi:10.1038/hdy.2014.5

Dauby G, Duminil J, Heuertz M, Koffi G, Stévart T  & Hardy OJ (2014) Congruent phylogeographic patterns of eight tree species in Atlantic Central Africa provide insights on the past dynamics of forest cover. Molecular Ecology 23:2299-2312. doi: 10.1111/mec.12724

Dellicour S, Kastally C, Hardy OJ, Mardulyn P (2014) Comparing phylogeographic hypotheses by simulating DNA sequences under a spatially explicit model of coalescence. Molecular Biology and Evolution 31:3359-3372.

Dellicour S, Mardulyn P, Hardy OJ, Hardy C, Roberts SPM, Vereecken NJ (2014). Inferring the mode of colonization of a solitary bee of a rapid range expansion from multi-locus DNA sequence variation. Journal of Evolutionary Biology 27:116–132. doi:10.1111/jeb.12280

Donkpegan ASL, Hardy OJ, Lejeune P, Oumorou M, Daïnou K, Doucet J-L (2014) Un complexe d’espèces d’Afzelia des forêts africaines d’intérêt économique et écologique (synthèse bibliographique). Biotechnol. Agron. Soc. Environ. 18(2): 233-246. 

Fayolle A, Swaine MD, Bastin J-F, Bourland N, Comiskey JA, Dauby G, Doucet J-L, Gillet J-F, Gourlet-Fleury S, Hardy OJ, Kirunda B, Kouamé FN, Plumptre AJ (2014) Continental-scale patterns of forest tree species composition across tropical Africa. Journal of Biogeography 41:2320–2331. doi:10.1111/jbi.12382

Ley AC, Dauby G, Köhler J, Wypior C, Röser M, Hardy OJ (2014) Comparative phylogeography of eight herbs and lianas (Marantaceae) in central African rainforests. Frontiers in Genetics 5:403. doi: 10.3389/fgene.2014.00403

Ley AC, Hardy OJ (2014) Contrasting patterns of gene flow between sister plant species in the understorey of African moist forests – the case of sympatric and parapatric Marantaceae species. Molecular Phylogenetics & Evolution 77: 264-274. doi: 10.1016/j.ympev.2014.04.026

Manel S, Couvreur TLP, Couteron P, Munoz F, Hardy OJ, Sonké B (2014) Characterizing the phylogenetic tree community structure of a protected tropical rain forest area in Cameroon. PLoS ONE, 9(6): e98920. doi:10.1371/journal.pone.0098920

Parmentier I, Réjou-Méchain M, Chave J, Vleminckx J, Thomas DW, Kenfack D, Chuyong GB, Hardy OJ (2014) Prevalence of phylogenetic clustering at multiple scales in an African rain forest tree community. Journal of Ecology 102: 1008–1016. doi:10.1111/1365-2745.12254

Simo-Droissart M, Sonké B, Droissart V, Geerinck D, Micheneau C, Lowry II PP, Plunkett GM, Hardy OJ, Stévart T (2014) Taxonomic Revision of the Continental African Species of Angraecum section Pectinaria (Orchidaceae). Systematic Botany 39:725-739. doi: 10.1600/036364414X682184

Vleminckx J, Drouet T, Amani C, Lisingo J, Lejoly J, Hardy OJ (2014) Impact of fine-scale edaphic heterogeneity on tree species assembly in a central African rainforest. Journal of Vegetation Science 26: 134–144. doi: 10.1111/jvs.12209

Vleminckx J, Morin-Rivat J, Biwolé AB, Daïnou K, Gillet J-F, Doucet J-L, Drouet T, Hardy OJ (2014) Soil charcoal to assess the impacts of past Human disturbances on tropical forests. PLoS ONE 9(11): e108121. doi:10.1371/journal.pone.0108121


Budde KB, González-Martínez SC, Hardy OJ, Heuertz M (2013) The ancient tropical rainforest tree Symphonia globulifera L. f. (Clusiaceae) was not restricted to postulated Pleistocene refugia in Atlantic Equatorial Africa. Heredity 111, 66-76.

Duminil J, Brown RP, Ewédjè E-EBK, et al. (2013) Large-scale pattern of genetic differentiation within African rainforest trees: Insights on the roles of ecological gradients and past climate changes on the evolution of Erythrophleum spp (Fabaceae). BMC Evolutionary Biology 13, 195.

Hardy OJ, Born C, Budde K, et al. (2013) Comparative phylogeography of African rain forest trees: A review of genetic signatures of vegetation history in the Guineo-Congolian region. Comptes Rendus - Geoscience 345, 284-296.

Hutsemékers V, Hardy OJ, Vanderpoorten A (2013) Does water facilitate gene flow in spore-producing plants? Insights from the fine-scale genetic structure of the aquatic moss Rhynchostegium riparioides (Brachytheciaceae). Aquatic Botany 108, 1-6.

Ley AC, Hardy OJ (2013) Improving AFLP analysis of large-scale patterns of genetic variation - A case study with the Central African lianas Haumania spp (Marantaceae) showing interspecific gene flow. Molecular Ecology 22, 1984-1997.

Parmentier I, Duminil J, Kuzmina M, et al. (2013) How effective are DNA barcodes in the identification of African rainforest trees? PLoS ONE 8, e54921.

Piñeiro R, Micheneau C, Dauby G, Hardy OJ (2013) Isolation of nuclear microsatellite loci in the African tree Scorodophloeus zenkeri (Fabaceae). Conservation Genetics Resources 5, 219-221.

Simo-Droissart M, Micheneau C, Sonké B, et al. (2013) Morphometrics and molecular phylogenetics of the continental African species of Angraecum section Pectinaria (Orchidaceae). Plant Ecology and Evolution 146, 295-309.


Anthony NM, Mickala P, Abernethy KA, et al. (2012) Biodiversity and conservation genetics research in central Africa: new approaches and avenues for international collaboration. Conservation Genetics Resources 4, 523-525.

Assogbadjo A, Fandohan B, Glèlè Kakai R, et al. (2012) Genetic evidence of the contribution of ethnic migrations to the propagation and persistence of the rare and declining scrambling shrub. Human Ecology 40, 117-128.

Baraloto C, Hardy OJ, Paine CET, et al. (2012) Using functional traits and phylogenetic trees to examine the assembly of tropical tree communities. Journal of Ecology 100, 690-701.

Daïnou K, Laurenty E, Mahy G, et al. (2012) Phenological patterns in a natural population of a tropical timber tree species, Milicia excelsa (Moraceae): evidence of isolation by time and its interaction with feeding strategies of dispersers. American Journal of Botany 99, 1453-1463.

Dauby G, Hardy OJ (2012) Sampled-based estimation of diversity sensu stricto by transforming Hurlbert diversities into Effective Number of Species. Ecography 35, 661-672.

Droissart V, Hardy OJ, Sonké B, Dahdouh-Guebas F, Stévart T (2012) Subsampling herbarium collections to assess geographic diversity gradients: a case-study with endemic Orchidaceae and Rubiaceae in Cameroon. Biotropica 44, 44-52.

Duminil J, Kenfack D, Viscosi V, Grumiau L, Hardy OJ (2012) Testing species delimitation in sympatric species complexes: the case of an African tropical tree, Carapa spp. (Meliaceae). Molecular Phylogenetics and Evolution 62, 275-285.

Ewedje E-E, Parmentier I, Natta A, Ahanchédé A, Hardy OJ (2012) Morphological variability of the tallow tree, Pentadesma butyracea Sabine (Clusiaceae), in Benin. Genetic Resources and Crop Evolution 59, 625-633.

Hardy OJ, Couteron P, Munoz FJ, Ramesh BR, Pélissier R (2012) Phylogenetic turnover in tropical tree communities: impact of environmental filtering, biogeography and mesoclimate niche conservatism. Global Ecology and Biogeography 21, 1007-1016.

Hardy OJ, Pavoine S (2012) Assessing phylogenetic signal with measurement error: a comparison of Mantel tests, Blomberg et al.'s K, and phylogenetic distograms. Evolution 66, 2614-2621.

Koffi KG, Heuertz M, Jans R, et al. (2012) Characterization of new microsatellite loci isolated from Santiria trimera (Burseraceae). American Journal of Botany 99, e334-e336.

Mouquet N, Devictor V, Meynard CN, et al. (2012) Ecophylogenetics: advances and perspectives. Biological Reviews 87, 769-785.


Micheneau C, Dauby G, Bourland N, Doucet J-L, Hardy OJ (2011) Development and characterization of microsatellite loci in Pericopsis elata (Fabaceae) using a cost-efficient approach. American Journal of Botany 98:e268-e270. doi:10.3732/ajb.1100070

Réjou-Mechain M, Flores O, Bourland N, Doucet J-L, Fétéké R F, Pasquier A, Hardy OJ (2011) Spatial aggregation of tropical trees at multiple spatial scales. Journal of Ecology 99, 1373–1381. doi: 10.1111/j.1365-2745.2011.01873.x

Droissart V, Hardy OJ, Sonké B, Dahdouh-Guebas F, Stévart T (2011) Subsampling Herbarium Collections to Assess Geographic Diversity Gradients: A Case-Study with endemic Orchidaceae and Rubiaceae in Cameroon. Biotropica (in press) 43: doi: 10.1111/j.1744-7429.2011.00777.x

Pearcy M, Hardy OJ, Aron S (2011). Automictic parthenogenesis and rate of transition to homozygosity. Heredity 107:187-188. DOI: 10.1038/hdy.2010.172

Eastman JM, Paine CET, Hardy OJ (2011) spacodiR: structuring of phylogenetic diversity in ecological communities. Bioinformatics 17:2437-2438. doi: 10.1093/bioinformatics/btr403

Droissart V, Sonké B, Hardy OJ, Simo M, Taedoumg H. Nguembou CK, Stévart T (2011) Do plant families with contrasting functional traits show similar patterns of endemism? A case study with Central African Orchidaceae and Rubiaceae. Biodiversity and Conservation 20: 1507-1531. doi: 10.1007/s10531-011-0042-z

Schaefer H, Hardy OJ, Silva L, Barraclough T, Savolainen V (2011) Testing Darwin’s naturalisation hypothesis in the Azores. Ecology Letters 14:389-396. doi: 10.1111/j.1461-0248.2011.01600.x

Réjou-Méchain M, Hardy OJ (2011) Properties of similarity indices under niche-based and dispersal-based processes in communities. The American Naturalist 177: 589-604. doi: 10.1086/659627

Duminil J, Koffi GK, Debout G, Sebastiani F, Vendramin GG, Heuertz M, Gonzalez-Martinez SC, Hardy OJ (2011). Isolation of SSR markers for two African tree tropical species Erythrophleum suaveolens and E. ivorense (Caesalpinioideae). American Journal of Botany 98: E106-E108. doi: 10.3732/ajb.1000482

Bourguignon T, Šobotník J, Lepoint G, Martin J-M, Hardy OJ, Dejean A, Roisin Y (2011) Feeding ecology and phylogenetic structure of a complex neotropical termite assemblage, revealed by nitrogen stable isotope ratios. Ecological Entomology 36: 261-269. doi: 10.1111/j.1365-2311.2011.01265.x

Koffi KG; Hardy OJ, Doumenge C, Cruaud C, Heuertz M (2011) Diversity gradients and phylogeographic patterns in a widespread African tree typical of mature rainforests, Santiria trimera (Burseraceae). American Journal of Botany 98: 254-264. doi: 10.3732/ajb.1000220

Vanderpoorten A, Hardy OJ, Lambinon J, Raspé O (2011) Two reproductively isolated cytotypes and a swarm of highly inbred, disconnected populations : a glimpse into Salicornia’s evolutionary history… and challenging taxonomy. Journal of Evolutionary Biology 142: 204-208. doi: 10.1111/j.1420-9101.2010.02198.x  

Parmentier I, Harrigan RJ, Buermann W, Mitchard ETA, Saatchi S, Malhi Y, Bongers F, Hawthorne WD, Leal ME, Lewis SL, Nusbaumer L, Sheil D, Sosef MSM, Affum-Baffoe K, Bakayoko A, Chuyong GB, Chatelain C, Comiskey JA, Dauby G, Doucet J-L, Faucet S, Gautier L, Gillet J-F, Kenfack D, Kouamé FN, Kouassi EK, Kouka LA, Parren MPE, Peh KS-H, Reitsma JM, Senterre B, Sonké B, Sunderland TCH, Swaine MD, Tchouto MGP, Thomas D, Van Valkenburg JLCH, Hardy OJ (2011). Predicting alpha diversity of African rain forests: models based on climate and satellite-derived data do not perform better than a purely spatial model. Journal of Biogeography 38: 1164-1176. doi: 10.1111/j.1365-2699.2010.02467.x

Van Rossum F, Stiers I, Van Geert A, Triest L, Hardy OJ (2011) Fluorescent dye particles as pollen analogues for measuring pollen dispersal in an insect-pollinated forest herbs. Oecologia 165: 663–674. doi: 10.1007/s00442-010-1745-7.

Leducq J-B, Llaurens V, Castric V, Saumitou-Laprade P, Hardy OJ, Vekemans X (2011) The effect of balancing selection on spatial genetic structure within populations: theoretical investigations on the self-incompatibility locus and empirical studies in Arabidopsis halleri. Heredity 106: 319-329. doi:10.1038/hdy.2010.68

Debout GDG, Doucet J-L, Hardy OJ (2011) Population history and dispersal ecology inferred from spatial genetic structure of a Central African timber tree, Distemonanthus benthamianus (Caesalpinioideae). Heredity 106:88-99. doi:10.1038/hdy.2010.35

Saad L, Tiébré M-S, Hardy OJ, Mahy G, Vanderhoeven S (2011) Patterns of hybridization and hybrid survival in the invasive alien Fallopia (Polygonaceae) complex. Plant Ecology & Evolution 144: 12-18. doi:10.5091/plecevo.2011.444.


Ndiade-Bourobou D, Hardy OJ, Favreau B, Moussavou H, Nzengue E, Mignot A, Bouvet JM (2010) Long-distance seed and pollen dispersal inferred from the spatial genetic structure at chloroplast and nuclear microsatellite markers in the very low density rainforest tree, Baillonella toxisperma Pierre in Central Africa. Molecular Ecology 19: 4949-4962.

Duminil J, Heuertz M, Doucet J-L, Bourland N, Cruaud C; Gavory F; Doumenge C, Navascués M, Hardy OJ (2010) CpDNA-based species identification and phylogeography: application to a complex of African tropical tree species. Molecular Ecology 19: 5469–5483. DOI: 10.1111/j.1365-294X.2010.04917.x

Daïnou K, Bizoux J-P, Doucet J-L, Mahy G, Hardy OJ, Heuertz M (2010) Forest refugia revisited: SSRs and cpDNA sequences support historical isolation in a wide-spread African tree with high colonization capacity, Milicia excelsa (Moraceae). Molecular Ecology 19:4462-4477.

Ley A, Hardy OJ (2010) Species delimitation in the Central African herbs Haumania (Marantaceae) using georeferenced nuclear and chloroplastic DNA sequences. Molecular Phylogenetics and Evolution 57:859–867. doi: 10.1016/j.ympev.2010.08.027

Koffi KG, Heuertz M, Doumenge C, Onana J-M, Gavory F, Hardy OJ (2010) A combined analysis of morphological traits, chloroplast and nuclear DNA sequences within Santiria trimera (Burseraceae) suggests several species following the Biological Species Concept. Plant Ecology & Evolution 143:160–169.

Yedmel MS, Kadio AA, Hardy OJ, Kouame NF, Barbier NS (2010) Impact du feu sur la structure et la composition floristique d’un peuplement forestier de Côte d’Ivoire et interaction avec les traitements sylvicoles. Revue canadienne de recherche forestière 40:1904-1913.

Segelbacher G, Cushman SA, Epperson BK, Fortin M-J, Francois O, Hardy OJ, Holderegger R, Taberlet P, Waits LP, Manel S. (2010) Applications of Landscape Genetics in Conservation Biology: Concepts and Challenges. Conservation Genetics 11:375–385.

Dauby G, Duminil J, Heuertz M, Hardy OJ (2010) Chloroplast DNA polymorphism and phylogeography of a Central African tree species widespread in mature rainforests: Greenwayodendron suaveolens (Annonaceae). Tropical Plant Biology 3: 4-13.

Vanderpoorten A, Raspé O, Risterrucci AM, Gohy L, Hardy O.J. (2010) Identification and characterization of eight nuclear microsatellite loci in the glasswort genus Salicornia (Chenopodiaceae). Belgian Journal of Botany 142: 204-208.

Hutsemekers V, Hardy OJ, Mardulyn P, Shaw AJ, Vanderpoorten A (2010) Macroecological patterns of genetic structure and diversity in the aquatic moss Platyhypnidium riparioides. New Phytologist 185: 852-864.


Hardy O.J. (2009) How fat is the tail? Heredity 103, 437–438.

Bizoux J-P., Daïnou K., Bourland N., Hardy O.J., Heuertz M., Mahy G., Doucet JL. (2009) Spatial genetic structure in Milicia excelsa (Moraceae) indicates extensive gene dispersal in a low density wind pollinated tropical tree. Molecular Ecology 18:4398-4408.

Duminil J., Hardy O. J., Petit R.J. (2009) Plant traits correlated with generation time directly affect inbreeding depression and mating system and indirectly genetic structure. BMC Evolutionary Biology 9:177.

Kyndt T, Assogbadjo AE, Hardy OJ, Glele Kakaï R, Sinsin B, Van Damme P, Gheysen G. (2009) Spatial genetic structuring of baobab (Adansonia digitata L., Malvaceae) in the traditional agroforestry systems of West Africa. American Journal of Botany 96: 950-957.

Navascués M., Hardy O. J., Burgarella C. (2009) Characterization of demographic expansions from pairwise comparisons of linked microsatellite haplotypes. Genetics 181: 1013-1019.

Parmentier I., Hardy O. J. (2009) The impact of ecological differentiation and dispersal limitation on species turnover and phylogenetic structure of inselberg’s plant communities. Ecography 32: 613-622.


Hardy O. J. (2008) Testing the spatial phylogenetic structure of local communities : statistical performances of different null models and test statistics on a locally neutral community. Journal of Ecology 96: 914–926.

Hardy O. J., L. Jost. (2008) Interpreting and estimating measures of community phylogenetic structuring. Journal of Ecology 96: 849–852

Hutsemékers V., Risterucci A.M., Ricca M., Boles S., Hardy O. J., Shaw A.J., Vanderpoorten, A. (2008) Identification and characterization of nuclear microsatellite loci in the aquatic moss Platyhypnidium riparioides (Brachytheciaceae). Molecular Ecology Resources 8: 1130–1132.

Dick C. W., Hardy O. J., Jones F. A., Petit R. (2008) Spatial scales of pollen and seed-mediated gene flow in tropical rain forest trees. Tropical Plant Biology 1: 20–33.

Born C., Hardy O. J., Ossari S., Attéké C., Wickings E. J., Chevallier M.-H., Hossaert-McKey M. (2008) Small-scale spatial genetic structure in the Central African rainforest tree species, Aucoumea klaineana: a hierarchical approach to infer the impact of limited gene dispersal, population history and habitat fragmentation. Molecular Ecology 17: 2041-2050.

Vanderpoorten, A., N. Devos, B. Goffinet, O. J. Hardy, A. J. Shaw (2008). The barriers to oceanic island radiation in bryophytes: insights from the phylogeography of the moss Grimmia montana. Journal of Biogeography 35: 654–663.

Hardy O. J., Pearcy M., Aron S. (2008). Small-scale spatial genetic structure in an ant species with sex-biased dispersal. Biological Journal of the Linnean Society 93: 465–473.


Basset, Y., Corbara, B., Barrios, H., Cuénoud, P., Leponce, M., Aberlenc, H.-P., Bail, J., Bito, D., Bridle, J. R., Castaño-Meneses, G., Cizek, L., Cornejo, A., Curletti, G., Delabie, J. H. C., Dejean, A., Didham, R. K., Dufrêne, M., Fagan, L. L., Floren, A., Frame, D. M., Hallé, F., Hardy, O. J., Hernandez, A., Kitching, R. L., Lewinsohn, T. M., Lewis, O. T., Manumbor, M., Medianero, E., Missa, O., Mitchell, A. W., Mogia, M., Novotny, V., Oliveira, E. G. d., Ødegaard, F., Orivel, J., Ozanne, C. M. P., Pascal, O., Pinzón, S., Rapp, M., Ribeiro, S. P., Roisin, Y., Roslin, T., Roubik, D. W., Samaniego, M., Schmidl, J., Sørensen, L. L., Tishechkin, A., Van Osselaer, C. & Winchester, N. N. (2007) IBISCA-Panama, a large-scale study of arthropod beta-diversity and vertical stratification in a lowland rainforest: rationale, description of study sites and field methodology. Bulletin de l'Institut Royal des Sciences Naturelles de Belgique-Entomologie 77: 39-69.

Tiébré M.-S., Bizoux J.-P., Hardy O.J., Bailey J. P., Mahy G (2007). Hybridization and morphogenetic variation in the invasive alien Fallopia (Polygonaceae) complex in Belgium (Western Europe). American Journal of Botany 94: 1900-1910.

Cloutier D., Hardy O. J., Caron H., Ciampi A. Y., Degen B., Kanashiro M., Schoen D. J. (2007). Low inbreeding and high pollen dispersal distances in populations of two Amazonian forest tree species Biotropica 39: 406–415.

Hardy O. J., B. Senterre (2007). Characterising the phylogenetic structure of communities by an additive partitioning of phylogenetic diversity. Journal of Ecology 95: 493–506.

Valbuena-Carabaña M., González-Martínez S.C., Hardy O.J., Gil L. (2007). Fine-scale spatial genetic structure in mixed oak stands with different levels of hybridization. Molecular Ecology 16: 1207-1219.


Pearcy M, O. J. Hardy, S Aron (2006). Thelytokous parthenogenesis and its consequences on inbreeding in an ant. Heredity 96: 377-382.

Hardy, O. J., L. Maggia, E. Bandou, P. Breyne, H. Caron, M-H. Chevallier, A. Doligez, C. Dutech, A. Kremer, C. Latouche-Hallé, V. Troispoux, V. Veron, B. Degen (2006). Fine-scale genetic structure and gene dispersal inferences in ten neotropical tree species. Molecular Ecology 15: 559-571.


Michaux, J.R., O.J. Hardy, F. Justy, P. Fournier, A. Kranz, M. Cabria, A.Davison, R. Rosoux & R. Libois (2005). Conservation genetics and population history of the threatened European mink Mustela lutreola, with an emphasis on the West European population. Molecular Ecology 14: 2373–2388.

Parmentier, I., T. Stévart & O. J. Hardy (2005). The inselberg flora of Atlantic Central Africa. I. Determinants of species assemblages. Journal of Biogeography 32: 685-696.


Hardy, O. J., S. C. González-Martínez, B. Colas, H. Fréville, A. Mignot & I. Olivieri (2004). Fine-scale genetic structure and gene dispersal in Centaurea corymbosa (Asteraceae) II. Correlated paternity within and among sibships. Genetics 168: 1601–1614.

Heuertz, M., S. Fineschi, M. Anzidei, R. Pastorelli, D. Salvini, L. Paule, N. Frascaria-Lacoste, O. J. Hardy, X. Vekemans, G. G. Vendramin. (2004). Chloroplast DNA variation and postglacial recolonization of common ash (Fraxinus excelsior L.) in Europe. Molecular Ecology 13: 3437-3452.

Kelly, B. A., O. J. Hardy & J.-M. Bouvet (2004). Temporal and spatial genetic structure of Vitellaria paradoxa (shea nut tree) in agroforestry system in southern Mali. Molecular Ecology 13: 1231-1240.

Heuertz, M., J.-F. Hausman, O. J. Hardy, G. G. Vendramin, N. Frascaria-Lacoste & X. Vekemans (2004). Nuclear microsatellites reveal contrasting patterns of genetic structure between western and southeastern european populations of the common ash (Fraxinus excelsior L.). Evolution 58: 976–988.

Vekemans, X. & O. J. Hardy (2004). New insights from fine-scale spatial genetic structure analyses in plant populations. Molecular Ecology 13: 921-935.

Hardy, O. J. & B. Sonké (2004). Spatial pattern analysis of tree species distribution in a tropical rainforest in Cameroon: assessing the role of limited dispersal and niche differentiation. Forest Ecology and Management, 197: 191-202.

Hardy, O. J., S. C. González-Martínez , H. Fréville, G. Boquien, A. Mignot, B. Colas & I. Olivieri (2004). Fine-scale genetic structure and gene dispersal in Centaurea corymbosa (Asteraceae) I. Pattern of pollen dispersal. Journal of Evolutionary Biology 17: 795-806.


Heuertz, M., X. Vekemans, J. F. Hausman, M. Palada & O. J. Hardy (2003). Estimating seed vs. pollen dispersal from spatial genetic structure in the common ash. Molecular Ecology 12: 2483-2495.

Fenster, C. B., X. Vekemans & O. J. Hardy (2003). Quantifying gene flow from spatial genetic structure data in a metapopulation of Chamaecrista fasciculata (Leguminosae). Evolution 57: 995-1007.

Hardy, O. J. (2003). Estimation of pairwise relatedness between individuals and characterisation of isolation by distance processes using dominant genetic markers. Molecular Ecology 12: 1577-1588.

Hardy, O. J., N. Charbonnel, H. Fréville & M. Heuertz (2003). Microsatellite allele sizes : a simple test to assess their significance on genetic differentiation. Genetics 163: 1467–1482


Vanderhoeven, S., O. J. Hardy, X. Vekemans, C. Lefèbvre, M. De Loose, J. Lambinon, & P. Meerts (2002). A morphometric study of populations of the Centaurea jacea complex (Asteraceae) in Belgium. Plant Biology 4: 403-412.

Hardy, O. J. & X. Vekemans (2002). SPAGeDi : a versatile computer program to analyse spatial genetic structure at the individual or population levels. Molecular Ecology Notes 2: 618-620.

Hardy, O. J., M. De Loose, X. Vekemans & P. Meerts (2001). Allozyme segregation and inter-cytotype reproductive barriers in the polyploid complex Centaurea jacea. Heredity 87: 136-145.

Hardy, O. J. & X. Vekemans (2001). Patterns of allozyme variation in diploid and tetraploid Centaurea jacea at different spatial scales. Evolution 55: 943-954.

Tiedemann, R., O. J. Hardy, X. Vekemans & M.C. Milinkovitch (2000). Higher impact of female than male migration on population structure in large mammals. Molecular Ecology 9: 1159-1164.

Hardy, O. J., S. Vanderhoeven, P. Meerts, & X. Vekemans (2000). Spatial autocorrelation of allozyme and quantitative markers within a natural population of Centaurea jacea (Asteraceae). Journal of Evolutionary Biology 13: 656-667.

Hardy, O. J., S. Vanderhoeven, M. De Loose, & P. Meerts (2000). Ecological, morphological and allozymic differentiation between diploid and tetraploid knapweeds (Centaurea jacea) from a contact zone in the Belgian Ardennes. New Phytologist 146: 281-290.

Hardy, O. J. & X. Vekemans (1999). Isolation by distance in a continuous population: reconciliation between spatial autocorrelation analysis and population genetics models. Heredity, 83: 145-154.

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  1. Olivier Hardy Group     Plant population genetics and community diversity in tropical rainforests

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