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In review

Ramsay, M.S., Olgun, H. & Georgiev, A.V. (in review) Primate road ecology: Current knowledge and future directions. In: McKinney, T., Waters, S. & Rodrigues, M. (Eds.) Primates in Anthropogenic Landscapes: Exploring Primate Behavioural Ecology Across Human Contexts. Springer.


Russell, R.M., Bibollet-Ruche, F., Liu, W., Sherrill-Mix, S., Li, Y., Connell, J., Loy, D.E., Trimboli, S., Smith, A.G., Avitto, A.N., Gondim, M.V.P., Plenderleith, L.J., Wetzel, K.S., Collman, R.G., Ayouba, A., Esteban, A., Peeters, M., Kohler, W.J., Miller, R.A., François-Souquiere, S., Switzer, W.M., Hirsch, V.M., Marx, P.A., Piel, A.K., Stewart, F.A., Georgiev, A.V., Sommer, V., Bertolani, P., Hart, J.A., Hart, T.B., Shaw, G.M., Sharp, P.M.,Hahn, B.H. (2021) CD4 receptor diversity represents an ancient protection mechanism against primate lentiviruses. Proc Natl Acad Sci USA, 118(13), e2025914118.

Olgun, H., Mohammed, M.K., Mzee, A.J., Green, M.E.L., Davenport, T.R.B. & Georgiev, A.V. (2021) The implications of vehicle collisions for the Endangered and endemic Zanzibar red colobus Piliocolobus kirkii. Oryx, 1-9. doi:10.1017/S0030605320000605


Pre-print: bioRxiv, doi:



Elliot, T.F., Georgiev, A.V., Lokasola, A.L. & Smith, M.E. (2020) Hysterangium bonobo: A newly described truffle species that is eaten by bonobos in the Democratic Republic of Congo. Mycologia, 112: 1045-1047. DOI: 10.1080/00275514.2020.1790234

Kuzawa, C.W., Adair, L., Bechayda, S.A., Borja, J.R.B., Carba, D.B., Duazo, P.L., Eisenberg, D.T., Georgiev, A.V., Gettler, L.T., Lee, N.R., Quinn, E.A., Rosenbaum, S., Rutherford, J., Ryan, C., McDade, T.W. (2020) Evolutionary life history theory as an organising framework for cohort studies: insights from the Cebu Longitudinal Health and Nutrition Survey. Annals of Human Biology, 47: 94-105.DOI:


Georgiev, A.V., Melvin, Z.E., Warkentin, A-S., Winder, I.C., & Kassim, A. (2019) Two cases of dead-infant carrying by female Zanzibar red colobus (Piliocolobus kirkii) at Jozani-Chwaka Bay National Park, Zanzibar. African Primates, 13: 57-60.

Rosenfield, K.A, Semple, S., Georgiev, A.V., Maestripieri, D., Higham, J.P. & Dubuc, C. (2019) Experimental evidence that female rhesus macaques (Macaca mulatta) perceive variation in male facial masculinity, Royal Society Open Science, 6: 181415


Pre-print: bioRχiv, doi:



Milich, K.M., Georgiev, A.V., Petersen, R.M., Emery Thompson, M. & Maestripieri, D. (2018) Alpha male status and availability of conceptive females are associated with high glucocorticoid concentrations in high-ranking male rhesus macaques (Macaca mulatta) during the mating season. Hormones and Behavior. 97: 5-13.

Kyweluk, M.A., Georgiev, A.V., Borja, J.B., Gettler, L.T. & Kuzawa, C.K. (2018) Menarcheal timing is accelerated by favorable nutrition but unrelated to developmental cues of mortality or familial instability in Cebu, Philippines. Evolution and Human Behaviour. 39: 76-81.

Sherrill-Mix et al. (2018) Allometry and ecology of the bilaterian microbiome. mBio. 9: e00319-18


Petersdorf, M., Dubuc, C., Georgiev. A.V., Winters, D., Higham. J.P. (2017) Is male rhesus macaque facial coloration under intrasexual selection? Behavioural Ecology. 28: 1472-1481


Liu, W. Sherrill-Mix, S., Learn, G.H., Scully, E.J., Yingying, L., Avitto, A.N., Loy, D.E., Lauder, A.P., Sundararaman, S.A., Plenderleith, L.J., Ndjango, J.N., Georgiev, A.V., Ahuka-Mundeke, S., Peeters, M., Bertolani, P., Dupain, J., Garai, C., Hart, J., Hart, T.B., Shaw, G.M., Shap, P.M., Hahn, B.H. (2017) Wild bonobos host geographically restricted malaria parasites including a putative new Laverania species. Nature Communications, 8, 1635.

Georgiev, A.V., Ryan, C.P., Borja, J.B. & Kuzawa, C.K. (2017) Is early postnatal growth velocity, a proxy of minipubertal androgen action, related to adult second-to-fourth digit (2D:4D) ratios in men? A test in Cebu, Philippines. American Journal of Human Biology. 29: e23047

Georgiev, A .V., Ryan, C. P., Gettler, L. T., McDade, T. W. & Kuzawa, C. K. (2017) Second-to-fourth digit ratio (2D:4D) is unrelated to measures of somatic reproductive effort among young men from Cebu, The Philippines. American Journal of Physical Anthropology. 163: 437-455.


Wroblewski, E. E., Guethlein, L. A., Norman, P.J., Li, Y., Shaw, C.M., Han, A.S., Ndjango, J.N., Ahuka-Mundeke, S., Georgiev, A.V., Peeters, M., Hahn, B.H. & Parham, P. (2017) Bonobos maintain immune-system diversity with three functional types of MHC-B. The Journal of Immunology. 198:3480-3493


Ryan, C.P., Georgiev, A.V., McDade, T.W., Gettler, L.T, Eisenberg, D.T.A., Rzhetskaya, M., Hayes, M.G. & Kuzawa, C.W. (2017) Androgen receptor polyglutamine repeat length (AR-CAGn) modulates the effect of testosterone on androgen-dependent somatic traits in Filipino young adult men. American Journal of Physical Anthropology. 163: 317-327.



Georgiev, A.V., Kuzawa, C.K. & McDade, T.W. (2016) Early developmental exposures shape trade-offs between acquired and innate immunity in humans.  Evolution, Medicine, and Public Health. 256-269 doi:10.1093/emph/eow022


Moeller, A. H., Caro-Quintero, A., Mjungu, D., Georgiev, A. V., Lonsdorf, E. V., Muller, M. N., Pusey, A. E., Peeters, M., Hahn, B. H., Ochman, H. (2016) Cospeciation of gut microbiota with hominids. Science, 353: 380-382

Media coverage: UC Berkeley News :: UT News :: Science :: The Atlantic :: The Guardian ::

 LA Times :: The Scientist 

Ahuka-Mundeke, S., Lunguya-Metila, O., Mbenzo-Abokome, V., Butel, C., Inogwabini, B., Omasombo, V., Muyembe-Tamfum, J., Georgiev, A. V., Muller, M. N., Njdango, J. N., Li, Y., Delaporte, E., Hahn, B. H., Peeters, M., Ayouba, A., (2016) Genetic diversity of STLV-2 and interspecies transmission of STLV-3 in wild-living bonobos. Virus Evolution.


Georgiev, A.V., Christie, D., Rosenfield, K.A., Ruiz-Lambides, A.V., Maldonado. E., Emery Thompson, M. & Maestripieri, D. (2016) Breaking the succession rule: The costs and benefits of an alpha-status take-over by an immigrant rhesus macaque on Cayo Santiago. Behaviour. 153: 325-351

            Accompanying blog post: Part 1 | Part 2

Kuzawa, C.W., Georgiev, A.V., McDade, T.W., Augustin, S. & Gettler, L.T. (2016) Is there a testosterone awakening response in humans? Adaptive Human Physiology & Behavior. 2: 166-183


McDade, T.W, Georgiev, A.V., Kuzawa, C.W. (2016) Trade-offs in acquired and innate immune defenses in humans. Evolution, Medicine, and Public Health.


Maestripieri, D. & Georgiev, A.V. (2016) What cortisol can tell us about the costs of sociality and reproduction among free-ranging rhesus macaque females on Cayo Santiago. Invited review. American Journal of Primatology. 78: 92-105


Georgiev, A.V., Emery Thompson, M., Mandalaywala, T.M. & Maestripieri, D. (2015) Oxidative stress as an indicator of the costs of reproduction among free-ranging rhesus macaques. Journal of Experimental Biology. 218: 1981-1985


Georgiev, A.V., Muehlenbein, M.P., Prall, S.P., Emery Thompson, M. & Maestripieri, D. (2015) Male quality, dominance rank and mating success in free-ranging rhesus macaques. Behavioral Ecology. 26: 763-722



Georgiev, A.V., Russell, A.F., Emery Thompson, M., Otali, E., Muller, M.N. & Wrangham, R.W. (2014) The foraging costs of mating effort in male chimpanzees (Pan troglodytes schweinfurthii). International Journal of Primatology, 35, 725-745


Emery Thompson, M. & Georgiev, A.V. (2014) The high price of success: costs of mating effort in male primates. International Journal of Primatology, 35, 609-627

Georgiev, A.V. & Emery Thompson, M. (Eds.) (2014) The High Price of Success: Costs of Male Mating Effort in Primates. Special Issue of International Journal of Primatology, 35 (3-4)


Liu, W., Li, Y., Shaw, K.S., Learn, G.H., Plenderleith, L.J., Malenke, J.A., Sundararaman, S.A., Ramirez, M.A., Crystal, P.A., Smith, A.G., Bibollet-Ruche, F., Ayouba, A., Locatelli, S., Esteban, A., Mouacha, F., Guichet, E., Butel, C., Ahuka-Mundeke, S., Inogwabini, B.I., Ndjango, J.B.N., Speede, S., Sanz, C.M., Morgan, D.B., Gonder, M.K., Kranzusch, P.J., Walsh, P.D., Georgiev, A.V., Muller, M.N., Piel, A.K., Stewart, F.A., Wilson, M.L.,. Pusey, A.E., Cui, L., Wang, Z., Färnert, A., Sutherland, C.J., Nolder, D., Hart, J.A., Hart, T.B., Bertolani, P., Gillis, A., LeBreton, M., Tafon, B., Kiyang, J., Djoko, C.F., Schneider, B.S., Wolfe, N.D., Mpoudi-Ngole, E., Delaporte, E., Carter, R., Culleton, R.L., Shaw, G.M., Rayner, J.C., Peeters, M., Hahn, B.H. & Sharp, P.M. (2014) African origin of the malaria parasite Plasmodium vivax. Nature Communications, 5, doi:10.1038/ncomms4346

           Media coverage: ScienceDaily :: International Business Times :: U Penn News 



Georgiev. A.V., Klimczuk, A.C.E, Traficonte, D.M. & Maestripieri, D. (2013) When violence pays: A cost-benefit analysis of aggressive behavior in animals and humans. Evolutionary Psychology, 11: 678-699



Georgiev, A. V., Emery Thompson, M., Lokasola, A. L. & Wrangham, R. W. (2011) Seed predation by bonobos (Pan paniscus) at Kokolopori, Democratic Republic of Congo. Primates, 52: 309-314



Liu, W, Li, Y., Learn, G. H., Rudicell, R. S., Robertson, J. D., Keele, B. F., Ndjango, J-B. N., Sanz, C. M., Morgan, D. B., Locatelli, S., Gonder, M. K., Kranzusch, P. J., Walsh, P. D., Delaporte, E., Mpoudi-Ngole, 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 gorillas. Nature, 476, 420-427


Media coverage: Nature News :: BBC News :: BBC Earth :: New York Times :: :: Not Exactly Rocket Science Blog


Georgiev, A. V., Lokasola, A. L., Nkanga, L., Lokondja, A., Nsala, J., Likenge, J., Ilanga-Bomanga, A. & Likenge, J. (2010) New observations of the terrestrial holoparasite Chlamydophytum aphyllum Mildbr. and its consumption by bonobos at Kokolopori, Democratic Republic of Congo. African Journal of Ecology, 48, 849-852



PhD dissertation


Georgiev, A.V. (2012) Energetic Costs of Reproductive Effort in Male Chimpanzees. PhD dissertation. Harvard University, Cambridge, MA.

Book reviews



Georgiev, A. V. (2012) Book review: The Chimpanzees of Bossou and Nimba: Tetsuro Matsuzawa, Tatyana Humle and Yukimaru Sugiyama (Eds.) 2011 Springer. Primate Eye, No. 107



Georgiev, A. V. (2011). Book review: Cynthia J. Moss, Harvey Croze, and Phyllis C. Lee (eds.): The Amboseli Elephants: A Long-Term Perspective on a Long-Lived Mammal.  The University of Chicago Press, Chicago and London, USA, 2011, 383 pp. (USD 65.00, hardcover). International Journal of Primatology, 32, 1242-1244

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