Craig Stanford

Professor of Biological Sciences and Anthropology
Craig Stanford
Email Office AHF B10 Office Phone (213) 740-1918

Research & Practice Areas

1. The ecology and social behavior of nonhuman primates, especially the great apes

2. The conservation biology of reptiles, especially critically endangered tortoises and turtles


Craig Stanford is Professor of Biological Sciences and Anthropology at USC.  He has conducted extensive field research on wild great apes, monkeys, and reptiles. His work has often focused on the ecological relationships among the primate species sharing a tropical forest ecosystem. He has conducted field studies in East Africa, South Asia, Southeast Asia, and Central and South America. In addition to more than 15 years field research on chimpanzee behavioral ecology in East Africa, he has recently been collaborating in studies on mountain gorillas, endangered Asian primates and other animals. The author or co-author of 16 books and more than 130 scholarly articles, he also holds a research appointment in vertebrate biology at the Los Angeles County Museum of Natural History, and is involved in the biology and conservation of endangered tortoises in southeast Asia. Stanford’s research has been profiled in the New York Times, Los Angeles Times and other national newspapers, and he is a frequent guest on television and radio programs about human origins and animal behavior, and has published widely in popular as well as scholarly literature.

Stanford has also been given every major teaching and research award offered in USC Dornsife College and continues to teach a large lecture course on human origins each year.


  • Ph.D. Anthropology, University of California, Berkeley, 1990
  • B.A. Zoology, Drew University
  • M.A. Anthropology, Rutgers University
  • Tenure Track Appointments

    • Professsor of Biological Sciences and Anthropology, USC, 2001 – 2024
    • Associate Professor of Anthropology, USC, 1997 – 2000
    • Assistant Professor of Anthropology, USC, 1992 – 1996

    Research, Teaching, Practice, and Clinical Appointments

    • Visiting Assistant Professor, Biological Anthropology, Univ of Michigan, Ann Arbor, 1989-1991
    • Research Associate in Herpetology, Los Angeles County Natural History Museum, 1998-2024
  • Summary Statement of Research Interests

    I continue to 1) direct projects in Africa and Asia with my graduate students on great ape behavioral ecology, 2) am conducting conservation biology and field research on reptiles in Mexico, acquiring land and habitat for research sites, and 3) involved in assisting a large number of students and conservation biologists working in various countries on studies of reptile biology, with many of whom I co-publish.

    Research Keywords

    Primate Behavior and Ecology, Human Evolution, Great Apes, Tropical Forest Ecology, Herpetology, Turtles, Reptiles

    Research Specialties

    1. The ecology and social behavior of nonhuman primates, especially the great apes

    2. The conservation biology of reptiles, especially critically endangered tortoises and turtles

  • Conference Presentations

    • Acquisition and protection of critical habitat for the Bolson Tortoise (Gopherus flavomarginatus) in the Bolsón de Mapimí Biosphere Reserve, Durango, Mexico , Desert Tortoise CouncilTalk/Oral Presentation, Tucson, 2018-2019
    • Land acquisition and preservation for biodiversity , Turtle Survival AllianceTalk/Oral Presentation, Tucson, 2018-2019
    • Alpha male status predicts longevity in wild male chimpanzees , American Assoc Physical AnthropologistsTalk/Oral Presentation, Invited, Knoxville TN, 2012-2013
    • Comparative aspects of the behavioral ecology of Manouria emys phayrei and M. impressa in Thailand , Turtle Survival AllianceTalk/Oral Presentation, Tucson AZ, 2012-2013
    • Chimpanzee behavior and the reconstruction of Ardipithecus ramidus , American Association of Physical AnthropologistsPoster, Minneapolis, 2010-2011
    • Great apes and the evolution of the human diet , Ancestral Health SymposiumTalk/Oral Presentation, Invited, UCLA, 2010-2011
    • The role of seasonal fallback foods in the diets of chimpanzees and gorillas in Bwindi Impenetrable National Park, Uganda. , International Primatological Society Congress, KyotoTalk/Oral Presentation, Invited, Kyoto, Japan, 2010-2011
    • Nutritional composition of actual and potential insect prey for the Kasakela chimpanzees of Gombe Stream National Park, Tanzania. , American Association of Physical Anthropologists, AlbuquerquePoster, 2009-2010
  • Book

    • Stanford, c., Allen, j., Anton, s. (2024). Biological Anthropology: The Natural History of Humankind, 4th Edition. Pearson.
    • Stanford, C. (2023). Unnatural Habitat: Native and Exotic Wildlife in Los Angeles. Heyday Books.
    • Stanford, C. (2023). The Turtle Crisis: Conservation Biology of the World’s Rarest Chelonians. NY: Mercury Press.
    • stanford, c., allen, j., anton, s. (2019). Exploring Biological Anthropology, 4th Edition. Pearson.
    • Stanford, C. (2018). The New Chimpanzee. Cambridge Mass: Harvard University Press.
    • Stanford, C. (2013). Evolution: What Everyone Needs to Know.
    • Stanford, C. B. (2012). Planet Without Apes: Is it Possible to Save Our Closest Kin?. Cambridge Mass: Harvard University Press.
    • stanford, c. b. (2010). The Last Tortoise: A Tale of Extinction in Our Time. Harvard University Press.
    • stanford, c. b. (2008). Biological Anthropology, 3rd Edition. Prentice Hall.
    • bearzi, m., stanford, c. (2008). Beautiful Minds: The Parallel Lives of Apes and Dolphins. Harvard U Press.
    • stanford, c., allen, j., anton, s., fagan, b. (2007). Biological Anthropology and Archaeology: an Integrated Approach. Prentice-Hall.
    • stanford, c. (2007). Apes of the Impenetrable Forest. Prentice Hall (Primate Field Studies Series).
    • (2005). Biological Anthropology: The Natural History of Humankind. Prentice-Hall. Second Edition.
    • (2003). Upright. Boston and New York: Houghton Mifflin.
    • none (Ed.). (1998). Chimpanzee and Red Colobus: The Ecology of Predator and Prey. Harvard University Press. Paperback edition 2001. Harvard U Press.

    Book Chapters

    • Stanford, C. (2023). Chimpanzee perspectives on Human Origins. Humans: Perspectives on Our Evolution from World Experts
    • Xuecong, L., Craig, S., Y, L. (2015). Research and conservation of Sichuan snub-nosed monkeys in Shennongjia, China. Primates of China Springer.
    • Tuma, m., Stanford, c. B. (2011). History of human interactions with North American tortoises of the genus Gopherus. Biology and Conservation of North Amer Tortoises Johns Hopkins Press.
    • reynolds and notman (Ed.). (2006). Nkrurniungi, J.B. and C.B. Stanford GIS analysis of range use by sympatric mountain gorillas and chimpanzees in Bwindi Impenetrable National Park, Uganda. In Primates in West Uganda (J. Paterson, V. Reynolds and H. Notman, eds.). pp. p. 193-205. nyc: In Primates of Western Uganda (J. Paterson, V. Reynolds and H. Notman, eds.), pp. 193-205. Kluwer-Plenum Publ. Co..
    • reynpolds and notman (Ed.). (2006). Ground-nesting by chimpanzees in Bwindi Impenetrable National Park, Uganda. In Primates of West Uganda (J. Patersonm, V. Reynolds and H. Notman, eds.). No Journal Defined.
    • gangestead and simpson (Ed.). (2006). Stanford, C. Cousins: What the great apes tell us about human origins. In The Search for a Common Language: Enviormental Writing and Education. pp. p. 35-45. No Journal Defined.

    Book Review

    • Stanford, C. (2021). Review of Chimpanzee Culture Wars (by N. Langlitz). Quarterly Review of Biology. pp. 318.
    • Craig, S. (2015). Review of State of the Apes (by Arcus Foundation) and Encountering Gorillas (by James Newman). Biological Conservation. pp. 10.
    • Stanford, C. B. (2012). Review of Chimpanzees of the Lakeshore, by T. Nishida. American J Physical Anthropology. pp. 483.
    • stanford, c. (2009). Review of Muller and Wrangham, Sexual Coercion in Primates and Humans. american scientist.
    • stanford, c. (2008). Phillosophical Primates ? (review of Primates and Philosophers, by F. de Waal). Current Anthropology.
    • stanford, c. (2008). Review of Feeding Ecology of Great Apes and Other Primates (edited by G. Hohmann et al.). Ecology.
    • unknown (Ed.). (2007). Review of Before the Dawn (by N. Wade). American Scientist. Submitted, finch, C.E., H. Kaplan and C.B. Stanford. Humans at the High Table. Scientific American. American Scientist.
    • stanford, c. (2007). Review of Before the Dawn (by N. Wade). American Scientist.
    • none (Ed.). (2006). Stanford, C.B. review of The Evolution of Thought (edited by A. E. Russon and D. R. brgun). Quarterly Review of Biology. No Journal Defined. pp. p. xx.


    • Stanford, C. (2018). War for Peace among Wild Chimpanzees. Behavioral Scientist.
    • stanford, c. (2008). Comment on Sayers and Lovejoy, “The chimpanzee has no clothes: a critical examination of Pan troglodytes in models of human evolution.”. pp. 103-104. Current Anthropology.
    • stanford, c. (2007). Comment on Alter, “The once and future apeman: reflections on chimera, human evolution, and questions of displinary coherence.”. pp. 647. Current Anthropology.

    Journal Article

    • Lindken, T., Stanford (20th author), C. (2024). What factors influence the rediscovery of lost tetrapod species?. Global Change Biology. Vol. 30, pp. 1-8.
    • Stanford, C. (2021). Turtle conservation and trade in Mexico; how DNA-based tools can help to handle confiscated animals. Radiata. Vol. 30 (3), pp. 4-35.
    • Stanford, C., Co-authors, F. (2020). Turtles in Trouble: Imminent extinction risk for the world’s turtles and tortoises. Current Biology. Vol. 30 (12), pp. R721-R735.
    • Castaneda, G., Becerra, J., Valenzuela, S., Stanford, C. (2020). Conservation status of the Coahuila Box Turtle (Terrapene coahuila) in the Cuatro Ciénegas valley of Coahuila, Mexico. Chelonian Conservation and Biology. Vol. 19 (1), pp. 14-21.
    • Stanford, C. (2019). Saving the Coahuila Box Turtle through field conservation and captive breeding. Radiata. Vol. 28 (3), pp. 1-17.
    • Rhodin, A., Stanford, C., authors, F. (2018). Global conservation status of turtles and tortoises (Order Testudines). Chelonian Conservation and Biology. Vol. 17 (2), pp. 135-161.
    • McCarthy, M., Lester, J., Langergraber, K., Stanford, C., Vigilant, L. (2018). Genetic analysis suggests typical dispersal persists by chimpanzees in a fragmented forest landscape in Uganda. American Journal of Primatology. Vol. 80 (9), pp. e22902.
    • Fan, P., Li, Y., Stanford, C., Li, Z., Liu, K., Li, X. (2018). Home range variation of two different-sized groups of golden snub-nosed monkey (Rhinopithecus roxellana) in Shennongjia, China: implications for feeding competition. Zoological Research. Vol. 39, pp. 1-8.
    • Craig, S., authors, a. (2018). Top 25: The World’s Most Endangered Tortoises and Turtles.
    • Currylow, A., Mandimbihasina, A., Gibbons, P., Bekarany, E., Stanford, C., Louis, E., Crocker, D. (2017). Comparative ecophysiology of a critically endangered ectotherm: implications for conservation management. PLoS One. Vol. 2017
    • McCarthy, M., Lester, J., Stanford, C. (2017). Chimpanzees (Pan troglodytes) flexibly use cultivated species for nesting and bark feeding in a human-dominated habitat. International Journal of Primatology. Vol. 38, pp. 321-337.
    • Maureen, M., J, L., Craig, S. (2016). Chimpanzees (Pan troglodytes) flexibly use cultivated species for nesting and bark feeding in a human-dominated habitat. International Journal of Primatology. Vol. September (online), pp. not yet assigned.
    • Maureen, M., J, L., E, H., M, A., C, S., L, V. (2015). Genetic censusing identifies an unexpectedly sizeable population of chimpanzees in a fragmented forests landscape. BMC Ecology. Vol. 15, pp. 21-.
    • Liu, X., Stanford, C. B., Li, Y. (2013). Effects of group size on time budgets of Sichuan snub-nosed monkeys (Rhimpithecus roxellana) in Shennongjia National Nature Reserve, China. International J Primatology. Vol. 34, pp. 349-360.
    • Hernandez, R. A., Moore, J., Stanford, C. B. (2013). Chimpanzee nesting patterns in a dry habitat: ecological aspects and preferences. American J Primatology. Vol. 75, pp. 979-994.
    • Liu, X., Stanford, C. B., Li, J., Yao, H. (2013). Food selection by Rhinopithecus roxellana in relation to nutritional chemistry. American J Primatology. Vol. 75, pp. 860-871.
    • Stanford, C. B. (2012). Chimpanzees and the biology of Ardipithecus ramidus. Annual Reviews of Anthropology. Vol. 41, pp. 139-149.
    • Laurance, W., Stanford (52nd author), C. B. (2012). Long-term ecological changes and threats in tropical protected areas. Nature. Vol. 489, pp. 290-294.
    • Yao, H., Liu, C., Stanford, C., Yang, J. (2011). Pattern of male dispersal in Rhinopithecus roxellana. American J Primatology.
    • Wanchai, P., Stanford, C., Thirakupt, K. (2011). Home range size of the impressed tortoise, Manouria impressa, in Phu Luang Wildlife Sanctuary. Chelonian Biology and Conservation.
    • Yao, H., Liu, C., Stanford, C., Li, Y. (2011). Dynamics of the all-male unit of a provisioned Sichuan snub-nosed monkey group (Rhinopithecus roxellana) in Shennongjia Nature Reserve, China. Amer J Primtology.
    • Bearzi, M., Stanford, C. (2010). A bigger, better brain. American Scientist.
    • li, y., liu, x., stanford, c. (2009). Characteristics of a group of Hubei golden snub-nosed monkeys (Rhinopithecus roxellana hubeiensis) before and after major snow storms. American J Primatology. Vol. 71, pp. 523-526.
    • stanford, c. b., o’malley, r. c. (2008). Sleeping tree choice in Bwindi chimpanzees. American Journal of Primatology. Vol. 70, pp. 642-649.
    • bearzi, m., stanford, c. b. (2007). Dolphins and African apes: comparisons of sympatric socio-ecology. Contributions to Zoology 76(4): 235-254. Contributions to Zoology. Vol. 76 (4), pp. 235-54.
    • stanford, c. (2007). Arboreal bipedalism in wild chimpanzees: implications for models of the evolution of hominid posture and locomotion. American Journal of Physical Anthropology. Vol. 129, pp. 225-31.
    • (2006). Stanford, C.B. Review of The Cultrured Chimpanzee (by W. McGrew). American Anthropologist. No Journal Defined. pp. p. 107.
    • Arbib, M. (Ed.). (2006). Stanford, C.B. Cognition, imitation and culture in the great apes. In From Action to Language, edited by M. Arbib. Cambridge University Press. From action to language via the mirror neuron system/Cambridge University Press. pp. p. 91-109.
    • (2005). Stanford, C.B. The sympatric ecology of African great apes, with implications for the hominoid divergence. Primates. No Journal Defined. Vol. 91-101
    • (2005). Stanford, C.B. Chimpanzee behavior and ecology. Encyclopedia entry in World Book: Science Year. No Journal Defined. pp. p. n/a.
    • (2005). Stanford, C.B. Chimpanzees as Predators. In The World Atlas of Great Apes and their Conservation. UNEP. No Journal Defined. pp. p. n/a.

    Research Report

    • none (Ed.). (2005). Stanford, C. Chelonian conservation in Myanmar. Marmorata. No Journal Defined.

    Magazine/Trade Publication

    • Stanford, C. B., Tuma, M.Pancake Breakfast: Conservation of the genus Malacochersus in Africa. The Tortoise. Vol. 2, pp. 56-61.


    • Stanford, C., al, e. (2018). Turtles in Trouble: The Top 25+ Most Endangered Tortoises and Freshwater Turtles. IUCN/TCF publication.


    • 2001 Stanford, C.B. and H.T. Bunn (editors). Meat-eating and Human Evolution. Oxford University Press.
    • 2001 Stanford, C. Significant Others: the Ape-Human Continuum and the Quest for Human Nature. Basic Books.


    • 1999 Stanford, C.B. The Hunting Apes: Meat-eating and the Origins of Human Behavior. Princeton University Press. Paperback edition 2001; also printed in five foreign languages.
    • USC Raubenheimer Outstanding Senior Faculty Award, 2016-2017
    • Allan Wilson Distinguished Lecturer, New Zealand, Fall 2013
    • Traphagen Lecturer, Drew University – as outstanding alum, 2010-2011
    • USC Associates Award, 2007-2008
    • USC Center for Excellence in Teaching, Faculty Fellow, 2000-2003
    • USC or School/Dept Award for Teaching, USC General Education Teaching Award, 2000-2001
    • USC Phi Kappa Phi Faculty Recognition Award, For the books Chimpanzee and Red Colobus and The Hunting Apes., 2000
    • USC Raubenheimer Outstanding Junior Faculty Award, 1996
  • Committees

    • Chair, As Chair of DCAPT (Dornsife Committee on Appointments, Promotions and Tenure) I supervised assessments of approx 100 faculty dossiers during 2017-19 academic years., 2018-2019

    Conferences Organized

    • Co-leader, IUCN Red List Workshop, Mexico City, 2023-2024
    • Co-leader, IUCN Red List Workshop, Charleston SC, 2023-2024
    • Co-leader, IUCN Red List Workshop, Cochabamba, Bolivia, 2023-2024
    • Co-organizer, IUCN Red List Workshop, Singapore, 2018-2019

    Editorships and Editorial Boards

    • Editorial Board, Annual Editions in Physical Anthropology,, 2000 – 2013
    • Editorial Board, Paleoanthropology, 2001

    Professional Offices

    • Chair, IUCN Species Survival Commission, Specialist Group, 2017 – 2024
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