As editor

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)





Olgun, H., Mohammed, M.K., Mzee, A.J., Green, M.E.L., Davenport, T.R.B. & Georgiev, A.V. (2020) The impact and implications of vehicle collisions on the endangered and endemic Zanzibar red colobus (Piliocolobus kirkii). bioRxiv, doi:


Rosenfield, K.A, Semple, S., Georgiev, A.V., Maestripieri, D., Higham, J.P. & Dubuc, C. (2017) Experimental evidence that female rhesus macaques (Macaca mulatta) perceive variation in male facial masculinity. bioRχiv, doi:

Peer-reviewed articles


Kuzawa, C.W., Adair, L.Bechayda, S.A.Borja, J.R.B.Carba, D.B.Duazo. P.L.Dan T. A. Eisenberg, D.T.A., Georgiev, A.V.Gettler, L.T., Lee, N.R., Quinn, E.A., Rosenbaum, S., Rutherford, J.N., Ryan, C.P. & 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




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


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