David Morgan, Ph.D.

Conservation & Science
Lester E. Fisher Center for the Study and Conservation of Apes
Research Fellow; Co-director of the Goualougo Triangle Ape Project
  • Ph.D. – Biological Anthropology, Cambridge University
  • U.K.B.S. – Biology, Western Carolina University

Areas of Expertise

  • Chimpanzee and gorilla ecology and behavior
  • Environmental influence on African great apes
  • Lowland forest ecology
  • Gorilla and chimpanzee conservation


David began his research career in 1997 as an assistant in the Republic of Congo, where he collected data on western lowland gorillas with Mbeli Bai Project in the Nouabale-Ndoki National Park. After two years, he initiated the Goualougo Triangle Chimpanzee Project to expand our understanding of the central chimpanzee. This groundbreaking study has led to numerous discoveries in chimpanzee behavior, tool-use, and conservation needs. His investigation of the social structure and spatial distribution of three communities of chimpanzees was the focus of his graduate research at Cambridge University, U.K.

For more than a decade, David and his partner in life, Crickette Sanz, Ph.D., have co-directed what is now the Goualougo Triangle Ape Project. It is the only site in all of Africa to include focused studies on sympatric chimpanzees and gorillas.

Concerned about the fate of our closest relations, he has made evaluating the impact of mechanized logging on great apes a primary interest. David believes strategies to safeguard apes in logging concessions are attainable and effort should be made to develop and evaluate them.

He considers his greatest achievement thus far to be his involvement in securing the annexation of the Goualougo Triangle onto Nouabale-Ndoki Park in 2003.


  • Morgan, D., Strindberg, S., McElmurray, P., Zambarda, A., Singono, I., Huskisson, S., Musgrave, S., Ayina, C. E., Funkhouser, J., Hellmuth, H., Joshi, P., Cassidy, R., & Sanz, C. (2023). Extending the conservation impact of great ape research: Flagship species sites facilitate biodiversity assessments and land preservation. Primates. https://doi.org/10.1007/s10329-023-01080-x  
  • Cooksey, K. E., Sanz, C., Massamba, J. M., Ebombi, T. F., Teberd, P., Abea, G., Mbebouti, G., Kienast, I., Brogan, S., Stephens, C., & Morgan, D. (2023). Predictors of respiratory illness in western lowland gorillas. Primates, 1–13. https://doi.org/10.1007/s10329-022-01045-6  
  • Wroblewski, E. E., Guethlein, L. A., Anderson, A. G., Liu, W., Li, Y., Heisel, S. E., Connell, A. J., Ndjango, J.-B. N., Bertolani, P., Hart, J. A., Hart, T. B., … Sanz, C. M., Morgan, D. B., Peeters, M., Sharp, P. M., Hahn, B. H.,… & Parham, P. (2023). Malaria-driven adaptation of MHC class I in wild bonobo populations. Nature Communications, 14(1), 1033. https://doi.org/10.1038/s41467-023-36623-9
  • Teberd, P., Sanz, C., Zambarda, A., Kienast, I., Ebombi, T. F., Abea, G., Kunz, J., Judson, K., Stephens, C., & Morgan, D. (2023). Path to Acceptance and Refined Practices for Habituating Western Lowland Gorillas. Primates. https://doi.org/10.1007/s10329-023-01086-5.
  • Osuna‐Mascaró, A. J., Ortiz, C., Stolz, C., Musgrave, S., Sanz, C. M., Morgan, D. B., & Fragaszy, D. M. (2021). Dexterity and technique in termite fishing by chimpanzees (Pan troglodytes troglodytes) in the Goualougo Triangle, Republic of Congo. American Journal of Primatology, 83(1), e23215.
  • Barratt, C. D., Lester, J. D., Gratton, P., Onstein, R. E., Kalan, A. K., McCarthy, M. S., … & Kühl, H. (2021). Quantitative estimates of glacial refugia for chimpanzees (Pan troglodytes) since the Last Interglacial (120,000 BP). American Journal of Primatology, e23320.
  • Harris, D. J., Ndolo Ebika, S. T., Sanz, C. M., Madingou, M. P., & Morgan, D. B. (2021). Large trees in tropical rain forests require big plots. Plants, People, Planet, 3(3), 282-294.
  • Lester, J. D., Vigilant, L., Gratton, P., McCarthy, M. S., Barratt, C. D., Dieguez, P., … & Arandjelovic, M. (2021). Recent genetic connectivity and clinal variation in chimpanzees. Communications biology, 4(1), 1-11.
  • Boesch, C., Kalan, A. K., Mundry, R., Arandjelovic, M., Pika, S., Dieguez, P., Ayimisin, E. A., Barciela, A., Coupland, C., Egbe, V. E., Eno-Nku, M., Michael Fay, J., Fine, D., Adriana Hernandez-Aguilar, R., Hermans, V., Kadam, P., Kambi, M., Llana, M., Maretti, G.,  Morgan, D., Murai, M., Neil, E., Nicholl, S., Ormsby, L.J., Orume, R., Pacheco, L., Piel, A., Sanz, C., Sciaky, L., Stewart, F.A., Tagg, N., Wessling, E.G., Willie, J. & Kühl, H. S. 2020. Chimpanzee ethnography reveals unexpected cultural diversity. Nature Human Behaviour, 1–7. https://doi.org/10.1038/s41562-020-0890-1.
  • Cooksey, K., Sanz, C., Ebombi, T. F., Massamba, J. M., Teberd, P., Magema, E., Abea, G., Peralejo, J. S. O., Kienast, I., Stephens, C., & Morgan, D. 2020. Socioecological Factors Influencing Intergroup Encounters in Western Lowland Gorillas (Gorilla gorilla gorilla). International Journal of Primatology. https://doi.org/10.1007/s10764-020-00147-6.
  • Musgrave, S., Lonsdorf, E., Morgan, D., Prestipino, M., Bernstein-Kurtycz, L., Mundry, R., & Sanz, C. 2020. Teaching varies with task complexity in wild chimpanzees. Proceedings of the National Academy of Sciences, 117(2), 969. https://doi.org/10.1073/pnas.1907476116.
  • Campbell, T. P., Sun, X., Patel, V. H., Sanz, C., Morgan, D., & Dantas, G. 2020. The microbiome and resistome of chimpanzees, gorillas, and humans across host lifestyle and geography. The ISME Journal, 1–16. https://doi.org/10.1038/s41396-020-0634-2.
  • Plenderleith, L. J., Liu, W., Learn, G. H., Loy, D. E., Speede, S., Sanz, C.M., Morgan, D.B., Bertolani, P., Hart, J.A., Hart, T.B., Hahn, B.H., Sharp, P. M. 2019. Ancient Introgression between Two Ape Malaria Parasite Species. Genome Biology and Evolution, 11(11), 3269–3274. https://doi.org/10.1093/gbe/evz244.
  • Bibollet-Ruche, F., Russell, R. M., Liu, W., Stewart-Jones, G. B. E., Sherrill-Mix, S., Li, Y., … Hahn, B. H.   2019. CD4 receptor diversity in chimpanzees protects against SIV infection. Proceedings of the National Academy of Sciences, 201821197. https://doi.org/10.1073/pnas.1821197116.
  • Morgan, D. B.   2019. Impacts of Selective Logging and Associated Anthropogenic Disturbance on Intact Forest Landscapes and Apes of northern Congo. Frontiers in Forests and Global Change, 2. https://doi.org/10.3389/ffgc.2019.00028.
  • Sherrill-Mix, S., McCormick, K., Lauder, A., Bailey, A., Zimmerman, L., Li, Y., … Bushman, F. D. 2018. Allometry and Ecology of the Bilaterian Gut Microbiome. MBio, 9(2), e00319-18. https://doi.org/10.1128/mBio.00319-18.
  • Adamescu, G. S., Plumptre, A. J., Abernethy, K. A., Polansky, L., Bush, E. R., Chapman, C. A., … Beale, C. M. 2018. Annual cycles are the most common reproductive strategy in African tropical tree communities. Biotropica. Retrieved from https://eprints.whiterose.ac.uk/129767/.
  • Marrocoli, S., Gatiso, T. T., Morgan, D., Nielsen, M. R., & Kühl, H. 2018. Environmental Uncertainty and Self-monitoring in the Commons: A Common-pool Resource Experiment Framed Around Bushmeat Hunting in the Republic of Congo. Ecological Economics, 149, 274–284. https://doi.org/10.1016/j.ecolecon.2018.03.020.
  • Strindberg, S., Maisels, F., Williamson, E. A., Blake, S., Stokes, E. J., Aba’a, R., … Wilkie, D. S. 2018. Guns, germs, and trees determine density and distribution of gorillas and chimpanzees in Western Equatorial Africa. Science Advances, 4(4), eaar2964. https://doi.org/10.1126/sciadv.aar2964.
  • Kühl, H. S. Kuhl, H. S., Kalan, A. K., Arandjelovic, M., Aubert, F., D’Auvergne, L., Goedmakers, A., Jones, S., Kehoe, L., Regnaut, S., Tickle, A., Ton, E., van Schijndel, J., Abwe, E. E., Angedakin, S., Agbor, A., Ayimisin, E. A., Bailey, E., Bessone, M., Bonnet, M., Brazolla, G., Buh, V. E., Chancellor, R., Cipoletta, C., Cohen, H., Corogenes, K., Coupland, C., Curran, B., Deschner, T., Dierks, K., Dieguez, P., Dilambaka, E., Diotoh, O., Dowd, D., Dunn, A., Eshuis, H., Fernandez, R., Ginath, Y., Hart, J., Hedwig, D., Ter Heegde, M., Hicks, T. C., Imong, I., Jeffery, K. J., Junker, J., Kadam, P., Kambi, M., Kienast, I., Kujirakwinja, D., Langergraber, K., Lapeyre, V., Lapuente, J., Lee, K., Leinert, V., Meier, A., Maretti, G., Marrocoli, S., Mbi, T. J., Mihindou, V., Moebius, Y., Morgan, D., Morgan, B., Mulindahabi, F., Murai, M., Niyigabae, P., Normand, E., Ntare, N., Ormsby, L. J., Piel, A., Pruetz, J., Rundus, A., Sanz, C., Sommer, V., Stewart, F., Tagg, N., Vanleeuwe, H., Vergnes, V., Willie, J., Wittig, R. M., Zuberbuehler, K.,Boesch, C. 2016. Chimpanzee stone throwing and accumulation: evidence for a phylogenetic origin of ritual sites. Sci. Rep. 6, 22219; doi: 10.1038/srep22219.
  • Sanz, C. M., Morgan, D. B., & Hopkins, W. D. 2016. Lateralization and performance asymmetries in the termite fishing of wild chimpanzees in the Goualougo Triangle, Republic of Congo. American Journal of Primatology, online early view, https://doi.org/10.1002/ajp.22574.
  • Cameron, K., Reed, T., Morgan, D., Ondzié, A., Sanz C., Kühl H., Olson S., Leroy E., Karesh, W. and Mundry, R. 2016. Spatial and Temporal Dynamics of a Mortality Event Among Central African Modeling Disease Spread in Apes. PLoS ONE 11(5):e0154505. doi:10.1371/journal.pone.0154505.
  • Ndolo-Ebika, Sydney, Morgan, D., Sanz, C., Harris, D. 2015. Differences between trees and free-standing stranglers in Ficus in a Congolese forest. Plant Ecology and Evolution.
  • Ebika, STN, Morgan D, Sanz C and Harris DJ. 2015. Hemi-epiphytic Ficus (Moraceae) in a Congolese Forest. Plant Ecology and Evolution 148(3): 377-386.
  • Lesnik, J. J., C. M. Sanz and D. B. Morgan. 2015. The Interdigital Brace and Other Grips for Termite Nest Perforation by Chimpanzees of the Goualougo Triangle, Republic of Congo. American Journal of Physical Anthropology 157(2): 252-259.
  • Fuenfstueck, T., M. Arandjelovic, D. B. Morgan, C. Sanz, P. Reed, S. H. Olson, K. Cameron, A. Ondzie, M. Peeters and L. Vigilant. 2015. The Sampling Scheme Matters: Pan troglodytes troglodytes and P-t. schweinfurthii Are Characterized by Clinal Genetic Variation Rather Than a Strong Subspecies Break. American Journal of Physical Anthropology 156(2): 181-191.
  • Furuichi, T., C. Sanz, K. Koops, T. Sakamaki, H. Ryu, N. Tokuyama and D. Morgan. 2015. Why do wild bonobos not use tools like chimpanzees do?. Behaviour 152(3-4): 425-460.
  • Liu, W. M., Y. Y. Li, K. S. Shaw, G. H. Learn, L. J. Plenderleith, J. A. Malenke, S. A. Sundararaman, M. A. Ramirez, P. A. Crystal, A. G. Smith, F. Bibollet-Ruche, A. Ayouba, S. Locatelli, A. Esteban, F. Mouacha, E. Guichet, C. Butel, S. Ahuka-Mundeke, B. I. Inogwabini, J. B. N. Ndjango, S. Speede, C. M. Sanz, D. B. Morgan, M. K. Gonder, P. J. Kranzusch, P. D. Walsh, A. V. Georgiev, M. N. Muller, A. K. Piel, F. A. Stewart, M. L. Wilson, A. E. Pusey, L. W. Cui, Z. L. Wang, A. Farnert, C. J. Sutherland, D. Nolder, J. A. Hart, T. B. Hart, P. Bertolani, A. Gillis, M. LeBreton, B. Tafon, J. Kiyang, C. F. Djoko, B. S. Schneider, N. D. Wolfe, E. Mpoudi-Ngole, E. Delaporte, R. Carter, R. L. Culleton, G. M. Shaw, J. C. Rayner, M. Peeters, B. H. Hahn and P. M. Sharp. 2014. African origin of the malaria parasite Plasmodium vivax. Nature Communications 5: 10.
  • Sanz, C. M., I. Deblauwe, N. Tagg and D. B. Morgan. 2014. Insect prey characteristics affecting regional variation in chimpanzee tool use. Journal of Human Evolution 71: 28-37.
  • Wilson, M. L., Boesch, C., Fruth, B., Furuichi, T., Gilby, I. C., Hashimoto, C., … Wrangham, R. W. 2014. Lethal aggression in Pan is better explained by adaptive strategies than human impacts. Nature, 513(7518), 414–417. doi:10.1038/nature13727.
  • Tranquilli, S., M. Abedi-Lartey, K. Abernethy, F. Amsini, A. Asamoah, C. Balangtaa, S. Blake, E. Bouanga, T. Breuer, T. M. Brncic, G. Campbell, R. Chancellor, C. A. Chapman, T. R. B. Davenport, A. Dunn, J. Dupain, A. Ekobo, M. Eno-Nku, G. Etoga, T. Furuichi, S. Gatti, A. Ghiurghi, C. Hashimoto, J. A. Hart, J. Head, M. Hega, I. Herbinger, T. C. Hicks, L. H. Holbech, B. Huijbregts, H. S. Kuhl, I. Imong, S. L.-D. Yeno, J. Linder, P. Marshall, P. M. Lero, D. Morgan, L. Mubalama, P. K. N’Goran, A. Nicholas, S. Nixon, E. Normand, L. Nziguyimpa, Z. Nzooh-Dongmo, R. Ofori-Amanfo, B. G. Ogunjemite, C.-A. Petre, H. J. Rainey, S. Regnaut, O. Robinson, A. Rundus, C. M. Sanz, D. T. Okon, A. Todd, Y. Warren and V. Sommer. 2014. Protected Areas in Tropical Africa: Assessing Threats and Conservation Activities. Plos One 9(12): DOI: 10.1371/journal.pone.0114154.
  • Fuenfstueck, T., M. Arandjelovic, D. B. Morgan, C. Sanz, T. Breuer, E. J. Stokes, P. Reed, S. H. Olson, K. Cameron, A. Ondzie, M. Peeters, H. S. Kuehl, C. Cipolletta, A. Todd, S. Masi, D. M. Doran-Sheehy, B. J. Bradley and L. Vigilant. 2014. The Genetic Population Structure of Wild Western Lowland Gorillas (Gorilla gorilla gorilla) Living in Continuous Rain Forest. American Journal of Primatology 76(9): 868-878.
  • Hopper, L.M., Morgan, D.B., & Ross, S.R. 2014. The next direction for primatology? A commentary on Setchell (2013). International Journal of Primatology. 35(2), 341-348.
  • Bryne, R., Sanz, C., Morgan, D. 2013. Chimpanzees plan their tool use. In: Sanz, C., Call, J., Boesch, C. (Eds.), Tool use in animals. Cambridge University Press, Cambridge UK.
  • Morgan, D., Sanz, C., Greer, D., Rayden, T., Maisels, F. & Williamson, E.A. 2013. Great Apes and FSC: Implementing ‘Ape Friendly’ Practices in Central Africa’s Logging Concessions. Gland, Switzerland: IUCN/SSC Primate Specialist Group. 36 pp.
  • Sanz, S., Morgan, D. 2013. The social context of chimpanzee tool use. In: Sanz, C., Call, J., Boesch, C. (Eds.), Tool use in animals. Cambridge University Press, Cambridge UK.
  • Morgan, D.B., Sanz, C.M., Onononga, J.R., Eyana Ayina, C., Strindberg, S. 2012. Great Apes and Mechanized Logging in the Kabo Concession. In Clark, C. J. and Poulsen, J. R. (Eds.). Tropical Forest Conservation and Industry Partnership: An Experience from the Congo Basin Conservation Science and Practice. Conservation and Science Practice, Wiley-Blackwell.
  • Vallo, P., Petr??elková, K.J., Profousová, I., Petrášová, J., Pomajbíková, K., Leendertz, F., Hashimoto, C., Simmons, N., Babweteera, F., Machanda, Z., Piel, A., Robbins, M., Boesch, C., Sanz, C., Morgan, D., Sommer, V., Furuichi, T., Fujita, S., Matsuzawa, T.H., M.A., Modrý, D. 2012. Molecular diversity of entodiniomorphid ciliate Troglodytella abrassarti and evaluation of its coevolution with chimpanzees. American Journal of Physical Anthropology.
  • Morgan, D.B., Sanz, C.M., Stokes, E.J. 2012. The Importance of Set Aside Areas for the Conservation of Great Apes in the Ndoki Region: Preservation of the Goualougo and Djeke Triangle Forests. In Clark, C. J. and Poulsen, J. R. (Eds.). Tropical Forest Conservation and Industry Partnership: An Experience from the Congo Basin Conservation Science and Practice. Conservation and Science Practice, Wiley-Blackwell.
  • Sanz, C., Morgan D.B. (2011). Elemental variation in the termite fishing of wild chimpanzees (Pan troglodytes). Biological Letters, DOI: 10.1098/rsbl.2011.0088. 2011. Elemental variation in the termite fishing of wild chimpanzees (Pan troglodytes).
  • Gillespie, T.R., Morgan, D., Deutsch, J. C., Kuhlenschmidt, M. S., Salzer, J. S., Cameron, K., Reed, T., Sanz, C.   2010. A Legacy of Low-Impact Logging does not Elevate Prevalence of Potentially Pathogenic Protozoa in Free-Ranging Gorillas and Chimpanzees in the Republic of Congo: Logging and Parasitism in African Apes. EcoHealth, 6: 557-564.
  • Sanz, C., Schöning, C. Morgan, D. 2010. Chimpanzees prey on army ants with specialized tool set. American Journal of Primatology, 71:1’8.
  • Sanz, C., Morgan, D. B. 2010. Complexity of chimpanzee tool using behaviors. In: The Mind of the Chimpanzee: Ecological and Experimental Perspectives (Ed. by Lonsdorf, E. V., Ross, S. R. & Matsuzawa, T.. Chicago, IL: University of Chicago Press. pp. 127-140.
  • Neel, C., Etienne, L., Li, Y. Y., Takehisa, J., Rudicell, R. S., Bass, I. N., Moudindo, J., Mebenga, A., Esteban, A., Van Heuverswyn, F., Liegeois, F., Kranzusch, P. J., Walsh, P. D., Sanz, C. M., Morgan, D. B., Ndjango, J. B. N., Plantier, J. C., Locatelli, S., Gonder, M. K., Leendertz, F. H., Boesch, C., Todd, A., Delaporte, E., Mpoudi-Ngole, E., Hahn, B. H. Peeters, M. 2010. Molecular Epidemiology of Simian Immunodeficiency Virus Infection in Wild-Living Gorillas. Journal of Virology, 84(3): 1464-1476.
  • Liu, W., Li, Y., Learn, G. H., Rudicell, R. S., Robertson, J. D., Ndjango, J. B. N., Sanz, C. M., Morgan, D. B., Locatelli, S., Gonder, M. K., Kranzusch, P. J., Walsh, P. D., Delaporte, E., Mpoudi-Ngolle, E., Georgiev, A. V., Muller, M. N., Shaw, G. M., Peeters, M., Sharp, P. M., Rayner, J. C., Hahn, B. H. 2010. Origin of the human malaria parasite Plasmodium falciparum in western gorillas (Gorilla gorilla). Nature, 467: 420-427.
  • Sanz, C., Call, J., and Morgan, D.  2009. Design complexity in termite-fishing tools of chimpanzees (Pan troglodytes). Biology Letters, 5: 293-296.
  • Sanz, C. M. and Morgan, D. B. 2009. Flexible and Persistent Tool-using Strategies in Honey-gathering by Wild Chimpanzees. International Journal of Primatology, 30: 411-427.
  • Morgan, D.B. and Sanz, C.M. 2007. Best Practice Guidelines for Reducing the Impact of Commercial Logging on Great Apes in Western Equatorial Africa. Gland, Switzerland: IUCN SSC Primate Specialist Group (PSG). 32 pp.
  • Sanz, C.M. and Morgan, D.B. 2007. Chimpanzee tool technology in the Goualougo Triangle, Republic of Congo. Journal of Human Evolution, 52(4): 420-433.
  • Sanz, C.M, Morgan, D.B., Strindberg, S., and Onononga, J.R. 2007. Distinguishing between the nests of sympatric chimpanzees and gorillas. Journal of Applied Ecology, 44(2): 263-272.
  • Walsh, P.D., Breuer,T., Sanz, C.M., Morgan, D.B., and Doran-Sheehy, D. 2007. Potential for Ebola transmission between gorilla and chimpanzee social groups. American Naturalist, 169(5): 684-689.
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