anth%3491% % %%%%%%carvalho% fall%2014% · cultural primatology, relating it with social learning,...

ANTH 3491 CARVALHO FALL 2014

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PRIMATE MODELS FOR HUMAN BEHAVIOR Fall semester 2014, Mondays-Wednesdays 2.20-3.35 pm

CRN: 86936 (3 CREDITS) Classroom: 1776 G St, 105

Dr. Susana Carvalho Email: [email protected] Telephone: 202-994-4223 Office hours: Thursdays 3:00-4:00 pm and by appointment, 2114 G St NW Room 307 Course Synopsis

The course will be divided in two sections: Firstly we will introduce the theme of Cultural Primatology, relating it with social learning, and focusing on the debate about culture in non-human primates. In light of evolutionary theory, we will discuss the meaning of culture, how to recognize and to measure culture in nature, and how can we analyze culture comparatively. A key topic in this class is whether human culture is unique. We will discuss whether it makes any sense to talk about Primate Archaeology to understand the evolution of material culture not just in humans and our ancestors, but also in other primates.

Secondly, in the absence of direct behavioural evidence from our ancestors, we use indirect means to understand further our own behavioural evolution. One of the key sources of information is the study of some primate behavioural traits, especially those of our closest living relations, the apes. Here we focus on apes as models in socio-ecology, and cover such topics as bipedality, diet, technology, language, and culture. Such models are a challenge: How to model? What to model? Whom to model? Teaching will be based on lectures and discussions (seminars). Learning Objectives (Specific) By the end of this semester students will:

1) Understand the complexity of finding culture in non-human species; know the current debate on the existence of culture in non-human primates; learn which features are unique to human culture.

2) Know how to use primate modern behaviour to model and further understand the evolution of human behaviour. Knowledge and understanding of key theoretical issues grounded in evolutionary theory are expected.

Learning Objectives (General) This course will contribute to the student’s development of:

1) Reasoning on relevant theoretical issues while being able to use specific examples to illustrate key points.


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2) Expressing novel opinions about a given topic, provided that appropriate examples from the literature are given. Originality, synthesis, and critique are encouraged.

3) Scientific writing and oral presentation skills. Required Texts Only for Section 1 (Cultural Primatology): McGrew, W.C. (2004) The Cultured Chimpanzee. Reflections on Cultural Primatology. Cambridge University Press, 248 pp. Evaluation Grades are based on the performance in a final exam (essay), oral presentation, and participation in lectures and discussion seminars. A final essay on a unique topic will count for 50%; the oral presentation for 25% and, attendance to lectures and participation in the discussion seminars will count for 25% of the final grade. Note: The Essays will seek to answer an important question on the topics of the class after discussion with the instructor. Essays should be based on a critical reading of the literature and written with strong arguments supported by relevant data. For the discussion seminars all students should read the starter reference and, for each topic each student also will be assigned an individual reading, which therefore will be her/his unique contribution to the exercise. Religious Holidays It is acceptable for you to miss lecture or seminar due to observance of religious holidays. However, it is your responsibility to look ahead on the calendar and notify the instructor of any conflicts (for the entire semester) with seminars or lecture no later than the second week of class. Blackboard Once you are registered for this course, you will automatically have access to the Blackboard site associated with it. Go to https://blackboard.gwu.edu/ and sign in using your email ID and password. We will use Blackboard to communicate announcements, store important documents and external links to web sites of interest that deal with material covered in the course, and provide a way for you to check your grades as the course progresses. Academic Integrity All graded work must be completed in accordance with The George Washington University Code of Academic Integrity, available online: http://www.gwu.edu/~ntegrity/code.html Support for Students Outside the Classroom Disability Support Services (DSS). Any student who may need an accommodation based on the potential impact of a disability should contact the Disability Support Services


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office at 202-994-8250 in the Marvin Center, Suite 242, to establish eligibility and to coordinate reasonable accommodations. For additional information, please refer to http://gwired.gwu.edu/dss/ University Counseling Center (UCC). The Counseling Center (UCC) offers 24/7 assistance and referral to address students’ personal, social, career, and study skills problems. Services for students include: crisis and emergency mental health consultations; and confidential assessment, counseling services (individual and small group), and referrals. You can reach the UCC at 202-994-5300. For additional information, please refer to http://gwired.gwu.edu/counsel/CounselingServices/AcademicSupportServices Security In the case of emergency, if at all possible, the class should shelter in place. If the building that the class is in is affected, follow the evacuation procedures for the building. After the evacuation, seek shelter at a predetermined rendezvous location.


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Tentative schedule Please note that the course syllabus is a general plan for the course; deviations announced to the class by the instructor may be necessary. DATE TOPIC Aug 25 Introduction Aug 27 Primate Evolution in Ecological Context (I) Sep 1 Holiday Sep 3 Primate Evolution in Ecological Context (II) Sep 8 SECTION 1: Cultural Primatology Sep 10 Seminar#1: What is culture & Operationalizing culture Sep 15 Recognizing Culture (Methods) Sep 17 Invited lecture (TBA) Sep 22 Non-primate culture (Fish, Birds & Mammals) Sep 24 Culture in Monkeys Sep 29 Seminar#2: Are humans unique? Out 1 Culture in Cetaceans and Apes Out 6 Chimpanzee non-material culture Out 8 Chimpanzee material culture Out 13 Seminar#3: Primate Archaeology Out 15 SECTION 2: Referential Models (I) Out 20 Referential Models (II) Out 22 Seminar#4:Bipedality Out 27 Seminar#5: Culture Out 29 Seminar#6: Hunting Nov 3 Seminar#7: Cannibalism Nov 5 Seminar#8: Warfare Nov 10 Seminar#9: Technology Nov 12 Seminar#10: Family Nov 17 Seminar#11: Language Nov 19 Seminar#12: Intelligence Nov 24 Review session Nov 26 Holiday Dec 1 Oral Presentations (I) Dec 3 Oral Presentations (II)


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READING LIST

1. Introduction Primate Evolution in Ecological Context Cartmill, M. (1972). Arboreal adaptations and the origin of the order Primates. In: Tuttle, R.H. (ed.), The Functional and Evolutionary Biology of Primates, Aldine-Atherton, Chicago, pp. 97-122. Cartmill, M. (1974). Rethinking primate origins. Science 184:436-443. Sussman R. (1991). Primate origins and the evolution of angiosperms, American Journal of Primatology, 23: 209-223.

2. Section 1 Cultural Primatology

Books on the Evolution of Culture: Boyd, R. & Richerson, P.J. (eds.) (2005) The Origin and Evolution of Cultures. Oxford University Press, 456 pp. Laland, K.N. & Galef, B.G. (eds.) (2009) The Question of Animal Culture. Harvard University Press, 351 pp. McGrew, W.C. (1992) Chimpanzee Material Culture: Implications for Human Evolution. Cambridge University Press, 277 pp. McGrew, W.C. (2004) The Cultured Chimpanzee. Reflections on Cultural Primatology. Cambridge University Press, 248 pp. Mesoudi, A. (2011) Cultural Evolution. How Darwinian Theory Can Explain Human Culture and Synthesize the Social Sciences. Univ. of Chicago Press, 264 pp. de Waal, F.B.M. (2001) The Ape and the Sushi Master. Cultural Reflections of a Primatologist. Basic Books, 433 pp. Richerson, P.J. and Boyd, R. (2005) Not By Genes Alone. How Culture Transformed Human Evolution. University of Chicago Press, 332 pp. de Waal, F.B.M. & Tyack, P.J. (eds.) (2003) Animal Social Complexity. Intelligence, Culture and Individualized Societies. Harvard University Press, 616 pp.

Seminar # 1: What is culture; operationalizing culture Starter Reference (everyone to read): Whiten, A. et al. (1999) Cultures in chimpanzees. Nature, 399:682-685.

Individually References (to be assigned for the discussion seminars): Biro, D. et al. (2003) Cultural innovation and transmission of tool-use in wild chimpanzees: Evidence from field experiments. Journal of Animal Cognition, 6: 213-223.


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Bonnie, K.E. & de Waal, F.B. (2006) Affiliation promotes the transmission of a social custom: handclasp grooming among captive chimpanzees. Primates, 47: 27-34. Hockings, K.J. et al. (2007) Chimpanzees share forbidden fruit. PLoS One, 2:e886. Hockings, K.J. et al. (2006) Road crossing in chimpanzees: A risky business. Current Biology, 16: R668-R670. Horner, V. et al. (2010) Prestige affects cultural learning in chimpanzees. PLos One, 5(5): e10625. Kamilar, J.M. & Marshack, J.L. (2012) Does geography or ecology best explain ‘cultural’ variation among chimpanzee communities? Journal of Human Evolution, 62: 256-260. Masataka , N. et al. (2009) Free-ranging macaque mothers exaggerate tool-using behaviour when observed by offspring. PLoS One, 4(3): e4768. Nakamura, M. (2004) Proximate factors of different types of grooming hand-clasp in Mahale chimpanzees: Implications for chimpanzee social customs. Current Anthropology, 44: 563-571. Russon, A.E. et al. (2007) Orangutan leaf-carrying for nest-building: Toward unravelling cultural processes. Animal Cognition, 10: 189-202. Sapolsky, R.M. & Share, J.J. (2004) A pacific culture among wild baboons: Its emergence and transmission. PLoS Biology, 2(4): E101. Schoening, C. et al. (2008) The nature of culture: Technological variation in chimpanzee predation on army ants revisited. Journal of Human Evolution, 55: 48-59. van de Waal, E., Borgeaud, C. & Whiten, A. (2013) Potent social learning and conformity shape a wild primate’s foraging decisions. Science, 6131: 483-5. Whiten, A. et al. (2007) Transmission of multiple traditions within and between chimpanzee groups. Current Biology, 17: 1038-1043. Whiten, A. et al. (2005) Conformity to cultural norms of tool use in chimpanzees. Nature, 437: 737-740. Longer References (if you are especially interested in the topic): Alvard, M. S. (2003) The adaptive nature of culture. Evolutionary Anthropology, 12:136-149. Amant, R.S., Horton, T.E. (2008) Revisiting the definition of animal tool use. Animal Behaviour, 75: 1199-1208. Boesch, C., Marchesi, P., Marchesi, N., Fruth, B. & Joulian, F. (1994) Is nut cracking in wild chimpanzees a cultural behaviour? J. Hum. Evol. 26, 325–338. Hirata, S. et al. (2001) ‘Sweet-potato washing’ revisited. In: Primate Origins of Human Cognition and Behavior, Matsuzawa, T. (ed.), pp. 487-508. Springer Kendal, R.L. et al. (2009) Identifying social learning in animal populations: A new option’bias’ method. PLoS One, 4, e6541. Koops K, McGrew WC. , Matsuzawa T (2013) Ecology of culture: do environmental factors influence foraging tool use in wild chimpanzees, Pan troglodytes verus?Animal Behaviour, 85(1): 175-185.


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Lonsdorf, E.V. (2006) What is the role of mothers in the acquisition of termite-fishing behaviors in wild chimpanzees? Animal Cognition, 9: 36-46. Lycett, S.J. (2010) The importance of history in definitions of culture: Implications from phylogenetic approaches to the study of social learning in chimpanzees. Learning & Behavior, 38: 252-264. Mesoudi, A. et al. (2006) Towards a unified science of cultural evolution. Behavioral and Behavioral Sciences, 29:329-383. Perry, S.E. (2006) What cultural primatology can tell anthropologists about the evolution of culture. Annual Review of Anthropology, 35:171-190. Tennie, C. et al. (2009) Ratcheting up the ratchet: on the evolution of cumulative culture. Philosophical Transactions of the Royal Society, B, 364: 2405-2415. Whiten, A. (2011) The scope of culture in chimpanzees, humans and ancestral apes. Philosophical Transactions of the Royal Society, B, 366: 997-1007.

Seminar # 2: Are humans unique? General Reference (everyone to read) van Schaik, C.P. et al. (2003) Orangutan cultures and the evolution of material culture. Science, 299: 102-105.

Individual References (to be assigned for the discussion seminars): Bender, C.E. et al. (2009) Evidence of teaching in Atlantic spotted dolphins by mother dolphins foraging in the presence of their calves. Animal Cognition, 12: 43-53. Boyd, R., Richerson, P. J. & Henrich, J. (2011) The cultural niche: why social learning is essential for human adaptation. Proc. Natl. Acad. Sci. USA 103, 1–8. Franks, N.R. & Richardson, T. (2006) Teaching in tandem-running ants. Nature, 439: 153. Thornton, A. & McAuliffe, K. (2006) Teaching in wild meerkats. Science, 313: 227-229. Fuhrmann, D., Ravignani, A., Marshall-Pescini, S. & Whiten, A. (2014) Synchrony and motor mimicking in chimpanzee observational learning. Scientific Reports, 4, 5283. DOI: 10.1038/srep05383. Guinet, C. & Bouvier, J. (1995) Development of intentional stranding techniques in killer whale calves at Crozet Archipelago. Canadian Journal of Zoology, 73: 27-33. Hohmann, G. & Fruth, B. (2003) Culture in bonobos? Between-species and within-species variation in behavior. Current Anthropology, 44: 563-571. Hunt, G.R. & Gray, R.D. (2003) Diversification and cumulative evolution in tool manufacture by New Caledonian crows. Proceedings of the Royal Society B, 270: 867-874. Kruetzen, M. et al. (2011) Culture and geographic variation in orang-utan behavior. Current Biology, 21: 1-5. Madden, J.R. (2008) Do bowerbirds exhibit cultures? Animal Cognition, 11: 1-11. Mueller, C.A. & Cant, M.A. (2010) Imitation and traditions in wild banded mongooses. Current Biology, 20: 1171-1175.


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Nielsen M., Tomaselli, K., Mushin, I. & Whiten, A. (2014) Exploring tool innovation: a comparison of Western and Bushman children. J. Experimental Child Psychology, 126, 384-394. Santorelli, C.J. et al. (2011) Traditions in spider monkeys are biased towards the social domain. PLoS One, 6 (2): e16863. Tanaka, I. (1995) Matrilineal distribution of louse egg-handling techniques during grooming in free-ranging Japanese macaques. American Journal of Physical Anthropology, 98: 197-201. Whitehead, H. (1998) Cultural selection and genetic diversity in matrilineal whales. Science, 282: 1708-1711. (See also critique and rejoinder in Science, 284:205-) Zhang, P. et al. (2007) Habitual hot-spring bathing by a group of Japanese macaques in their natural habitat. American Journal of Primatology, 69:1425-1430. Longer References (if you are especially interested in the topic): Barrett, L. & Wursig, B. (2014) Why dolphins are not aquatic apes. Animal Behavior and Cognition, 1: 1-18. Leca, J.B. et al. (2007) Japanese macaque culture: Inter- and intra-troop variability of stone handling patterns across 10 troops. Behaviour, 144: 251-281. McGrew, W.C. (2003) Ten dispatches from the chimpanzee culture wars. In: Animal Social Complexity, de Waal, F.B.M. & Tyack, P.L. (eds.), Harvard University Press, pp. 419-439. (see also update in Laland & Galef, 2009) Perry, S.E. et al. (2003) Social conventions in wild white-faced capuchin monkeys: Evidence for behavioral traditions in a neotropical primate. Current Anthropology, 44: 241-268. Rendell, L. & Whitehead, H. (2001) Culture in whales and dolphins. Behavioral and Brain Sciences, 24: 309-324. Tennie, C. et al. (2009) Ratcheting up the ratchet: On the evolution of cumulative culture. Philosophical Transactions of the Royal Society, B, 364: 2405-2415. Whiten, A. et al. (2001) Charting cultural variation in chimpanzees. Behaviour, 138: 1481-1516.

Seminar #3: Primate Archaeology, what can tell us about the evolutionary origins of human culture? General Reference (everyone to read): Haslam, M. et al. (2009) Primate archaeology. Nature, 460: 339-344. Individual References (to be assigned for the discussion seminars): Byrne, R.W. (2007) Culture in great apes: Using intricate complexity in feeding skills to trace the evolutionary origin of human technical prowess. Philosophical Transactions of the Royal Society B, 362: 577-585.


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Carvalho, S. et al. (2009). Tool-composite reuse in wild chimpanzees: Archaeologically invisible steps in the technological evolution of early hominins? Animal Cognition, 12: 103-114. Carvalho, S. et al. (2012) Chimpanzee carrying behaviour and the origins of human bipedality. Current Biology, Goren-Inbar, N. et al. (2002) Nuts, nut-cracking, and pitted stones at Gesher Benot Ya’agov, Israel. Proceedings of the National Academy of Sciences (USA), 99: 2455-2460. Haslam M., Gumert M., Biro D., Carvalho S., Malaivijitnond S. (2013). Use-Wear patterns on wild macaque stone tools reveal their behavioural history. PLoS ONE 8(8): e72872. doi:10.1371/journal.pone.0072872 Leca, J.B. et al. (2010) Indirect social influence in the maintenance of stone-handling tradition in Japanese macaques, Macaca fuscata. Animal Behaviour, 79: 117-126. Marchant, L.F. & McGrew, W.C. (2005) Percussive technology: Chimpanzee baobab smashing and the evolutionary modelling of hominin knapping. In: Stone Knapping, Roux, V. and Bril, B. (eds.), Cambridge: McDonald Institute Monographs, pp. 341-350. McGrew, W.C. et al. (2007) Etho-archaeology of manual laterality: Well-digging by wild chimpanzees. Folia Primatologica, 78: 240-244. McGrew W.C. et al. 2014 Experimental primate archaeology: detecting stone handling by Japanese macaques (Macaca fuscata). Lithic Technology, 39: 113-121. Mercader, J. et al. (2007) 4,300-year-old chimpanzee sites and the origins of percussive stone technology. Proceedings of the National Academy of Sciences (USA), 104: 3043-3048. Roffman, I. et al. (2012) Stone tool production and utilization by bonobo-chimpanzees (Pan paniscus). Proc. Natl. Acad. Sci. (USA), 109: 14500-14503. Schick, K.A. et al. (1999) Continuing investigations into the stone-tool-making and tool-using capabilities of bonobo. Journal of Archaeological Science, 26:821-832. Stewart, F.A. et al. (2011) Living archaeology: Artefacts of specific nest site fidelity in wild chimpanzees. Journal of Human Evolution, 61: 388-395. Visalberghi, E. et al. (2009) Selection of effective stone tools by wild bearded capuchin monkeys. Current Biology, 19: 213-217. Visalberghi E, Haslam M, Spagnoletti N & Fragaszy D (2013). Use of stone hammer tools and anvils by bearded capuchin monkeys over time and space: construction of an archeological record of tool use. Journal of Archaeological Science, 40:3222-e3232.

Longer References (if you are especially interested in the topic): Borgo S, Spagnoletti N, Vieu L & Visalberghi E (2013) Artifact and artifact categorization: comparing humans and capuchin monkeys. Rev. Phil. Psych. 4:375–389. Carvalho, S. et al. (2008) Chaines operatoires and resource exploitation strategies in chimpanzee nutcracking. Journal of Human Evolution, 55: 148-155.


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Davidson, I. & McGrew, W.C. (2005) Stone tools and the uniqueness of human culture. Journal of the Royal Anthropological Institute, 11: 793-817. Gumert, M.D. et al. (2009) The physical characteristics and usage patterns of stone axe and pounding hammers used by long-tailed macaques in the Andaman Sea region of Thailand. American Journal of Primatology, 71: 594-608. van Schaik, C.P. & Pradhan, G.R. (2003) A model for tool-use traditions in primates: Implications for the coevolution of culture and cognition. Journal of Human Evolution, 44: 645-664. Toth, N. & Schick, K. (2009) The Oldowan: The tool making of early hominins and chimpanzees compared. Annual Review of Anthropology, 38: 289-305. Whiten, A. et al. (2009) The evolution and cultural transmission of percussive technology: integrating evidence from paleoanthropology and primatology. Journal of Human Evolution, 57: 420-435. Wynn, T.G. & McGrew, W.C. (1989) An ape’s view of the Oldowan. Man, 24: 383-398. Wynn, T.R. et al. (2011) “An ape’s view of the Oldowan” revisited. Evolutionary Anthropology, 20: 181-197.

3. Section 2 Referential Models

Useful Books: Dominguez-Rodrigo, M. (ed.) (2012) Stone Tools and Fossil Bones. Cambridge University Press. Esp. Chapters 6 & 7. Kinzey, W.G. (ed.) (1987) The Evolution of Human Behavior: Primate Models. State University of New York Press. McGrew, W.C. (1992) Chimpanzee Material Culture. Implications for Human Evolution. Cambridge University Press. McGrew, W.C. (2004) The Cultured Chimpanzee. Reflections on Cultural Primatology. Cambridge University Press. de Waal, F.B.M. (ed.) (2001) Tree of Origin. What Primate Behavior Can Tell Us about Human Social Evolution. Harvard University Press. Positives and negatives of modelling Foley, R. (1992) Studying human evolution by analogy. In: Cambridge Encyclopedia of Human Evolution, Cambridge Univ. Press, pp. 335-340. Kinzey, W.G. (ed.) (1987) The Evolution of Human Behavior: Primate Models. State University of New York Press. Lovejoy, C.O. (2009) Reexamining human origins in light of Ardipithecus ramidus. Science, 326: 74e1-e8. (doi: 10.1126/science.1175834) McGrew, W.C. (1981) The female chimpanzee as a human evolutionary prototype. In: Woman the Gatherer, Dahlberg, F. (ed.), Yale Univ. Press, pp. 35-73.


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McGrew, W.C. (2010) In search of the last common ancestor: new findings on wild chimpanzees. Philosophical Transactions of the Royal Society, B, 365:3266-76. Moore, J. (1996) Savanna chimpanzees, referential models and the last common ancestor. In: Great Ape Societies, McGrew et al., (eds.), Cambridge UP, pp. 275-292. Rantala, M.J. (2007) Evolution of nakedness in Homo sapiens. J. of Zoology, 273: 1-7. Sayers, K. and Lovejoy, C.O. (2008) The chimpanzee has no clothes: a critical examination of Pan troglodytes in models of human evolution. Current Anthropology 49: 87-114. Stanford, C.B. (2006) The behavioral ecology of sympatric African apes: Implications for understanding fossil hominid ecology. Primates, 47: 91-101. Stanford, C.B. (2012) Chimpanzees and the behavior of Ardepithecus ramidus. Annual Review of Anthropology, 41: 139-149. Stanford, C.B. & Allen, J.S. (1991) On strategic story-telling: Current models of human behavioural evolution. Current Anthropology, 32: 58-61. Tooby, J. & DeVore, I. (1987) The reconstruction of hominid behavioural evolution through strategic modeling. In: Kinzey, pp. 183-237. Wrangham, R.W. (1987) The significance of African apes for reconstructing human social evolution. In: Kinzey, pp. 51-71.

Ape Models Kano, T. (1992) The Last Ape. Pygmy Chimpanzee Behavior and Ecology. Stanford UP. Marlowe, F.W. (2005) Hunter-gatherers and human evolution. Evolutionary Psychology, 14: 54-67. Marlow, F.W. (2010) The Hadza. Hunter-gatherers of Tanzania. Univ. Cal. Press. McGrew, W.C. (1979) Evolutionary implications of sex differences in chimpanzee predation and tool use. In: The Great Apes, Hamburg, D.A. & McCown, E.R. (eds.), Benjamin/Cummings, pp. 441-463. Robbins, M. (2007) Gorillas: Diversity in ecology and behaviour. In: Primates in Perspective, Campbell, C.J. et al. (eds.), Oxford UP, pp. 305-321. Schwartz, J. (2005) The Red Ape. Orangutans and Human Origins. (rev. edit.) Westview Press, 286 pp. Sommer, V. & Reichard, U. (2000) Rethinking monogamy: The gibbon case. In: Primate Males, Kappeler, P. (ed.), Cambridge UP, pp. 159-168. Stanford, C.B. (1998) The social behavior of chimpanzees and bonobos: Empirical evidence and shifting assumptions. Current Anthropology, 39: 399-420. Stanford, C.B. (2001) The ape’s gift. Meat-eating, meat-sharing, and human evolution. In: Tree of Origin, pp, 95-117. Stanford, C.B. (2008) Apes of the Impenetrable Forest. The Behavioral Ecology of Sympatric Chimpanzees and Gorillas. Pearson/Prentice-Hall, 140 pp. Stanford, C.B. (2012) Chimpanzees and the behavior of Ardipithecus ramidus. Annual Review of Anthropology, 41: 139-149.


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de Waal, F.B.M. (2001) Apes from Venus: Bonobos and human social evolution. In: Tree of Origin, pp.41-68. Van Schaik, C. et al. (2004) Great ape social systems. In: The Evolution of Thought, Russon, A.E. & Begun, D.R. (eds.), Cambridge UP, pp. 190-209. White, F.J. (1996) Pan paniscus 1973 to 1996: Twenty-three years of field research. Evolutionary Anthropology, 5:11-17. Non-referential Models Foley, R.A. and Lee, P.C. (1989) Finite social space, evolutionary pathways, and reconstructing hominid behavior. Science, 243: 901-906. Laden, G. & Wrangham, R.W. (2002) The rise of the hominids as an adaptive shift in fallback foods: plant underground storage organs (USOs) and australopith origins. Journal of Human Evolution, 49: 482-498. Langdon, J.H. (1997) Umbrella hypotheses and parsimony in human evolution: A critique of the Aquatic Apes Hypothesis. Journal of Human Evolution, 33: 479-494. Martin, R.D. et al. (1994) The evolution of sexual size dimorphism in primates. In: The Differences between the Sexes, Short, R.V. & Balaban, E. (eds.), Cambridge UP, pp. 159-200. Plavcan, J.M. et al. (2005) Sexual dimorphism in Australopithecus afarensis revisited: How strong is the case for a human-like pattern of dimorphism? Journal of Human Evolution, 48: 313-320. Reno, P.L. et al. (2005) The case is unchanged and remains robust: Australopithecus afarensis exhibits on moderate skeletal dimorphism. Journal of Human Evolution, 48: 279-288. Tooby, J. & DeVore, I. (1987) The reconstruction of hominid behavioral evolution through strategic modeling. In: The Evolution of Human Behavior: Primate Models, Kinzey, W.G. (ed.), State University of New York Press, pp. 183-237. Verhaegen, M. et al. (2002) Aquaarboreal ancestors? Trends in Ecology and Evolution, 17, 212-217. Wrangham, R.W. (2005) The delta hypothesis: Hominoid ecology and hominin origins. In: Interpreting the Past, Lieberman, D.E. et al. (eds.), Brill, pp. 231-242. Wrangham, R.W. et al. (2009) Shallow-water habitats as sources of fallback foods for hominins. American Journal of Physical Anthropology, 140: 630-642. Wrangham, R.W. (2009) Catching Fire. How Cooking Made Us Human. Profile Books. Seminar # 4: Bipedality Individual References (to be assigned for the discussion seminars): Bauer, H.R. (1977) Chimpanzee bipedal locomotion in the Gombe National Park, East Africa. Primates, 18: 913-921. Carrier, D. F. (2011) The advantage of standing up to fight and the evolution of habitual bipedalism in hominins. PLoS One, 6(5): e19630.


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Carvalho, S. et al. (2012) Chimpanzee carrying behaviour and the origins of human bipedality. Current Biology, 22: R180-181. Lieberman, D.E. & Bramble, D.M. (2008) The evolution of marathon running. Sports Medicine, 37: 288-290. Mitchell, D. et al. (2009) Homeothermy and primate bipedalism: Is water shortage or solar radiation the main threat to baboon (Papio hamadryas) homeothermy? Journal of Human Evolution, 56: 439-446. Ponzer, H. et al. (2009) The metabolic cost of walking in humans, chimpanzees, and early hominins. Journal of Human Evolution, 56: 43-54. Raichlen, D.A. et al. (2008) The Laetoli footprints and early hominin locomotor kinetics. Journal of Human Evolution, 54: 112-117. Ruxton, G.D. & Wilkinson, D.M. (2011) Avoidance of overheating and selection for both hair loss and bipedality in hominins. Proceedings of the National Academy of Sciences of the USA, 108: 20965-20969. Stanford, C.B. (2006) Arboreal bipedalism in wild chimpanzees: implications for evolution of hominid posture and locomotion. American Journal of Physical Anthropology, 129: 225-231. Sylvester, A.D. & Kramer, P.A. (2008) Stand and shuffle: When does it make energetic sense? American Journal of Physical Anthropology, 135: 484-488. Thorpe, S.K.S. et al. (2007) Origin of human bipedalism as an adaptation for locomotion on flexible branches. Science, 316: 1328-1331. Videan, E.N. & McGrew, W.C. (2002) Bipedality in chimpanzee and bonobo: testing hypotheses on the evolution of bipedalism. American Journal of Physical Anthropology, 118: 184-190. Wrangham, R.W. (1980) Bipedal locomotion as a feeding adaptation in gelada baboons and its implications for hominid evolution. Journal of Human Evolution, 9: 329-331.

Seminar #5: Culture

Individual References (to be assigned for the discussion seminars): van Schaik, C.P. et al. (2003) Orangutan cultures and the evolution of material culture. Science, 299:102-105. Laland, K. & Janik, V. (2006) The animal cultures debate. Trends in Ecology and Evolution, 21:542-547. Kruetzen, M. et al. (2007) The animal cultures debate: Response to Laland and Janik. Trends in Ecology and Evolution, 22(1): 6. Luncz, L.V. et al. (2012) Evidence for cultural differences between neighboring chimpanzee communities. Current Biology, 22: 922-926. Whiten, A. et al. (2007) Transmission of multiple traditions within and between chimpanzee groups. Current Biology 17:1038-1043. Byrne, R.W. (2007) Culture in great apes: Using intricate complexity in feeding skills to trace the evolutionary origin of human technical prowess. Proceedings of the Royal Society, B, 362:577-585.


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Byrne, R.W. et al. (2004) Understanding culture across species. Trends in Cognitive Sciences, 8:341-346. Bonnie, K.E. & de Waal, F. (2006) Affiliation promotes the transmission of a social custom: Handclasp grooming among captive chimpanzees. Primates, 47: 27-34. Biro, D. et al. (2003) Cultural innovation and transmission of tool-use in wild chimpanzees: Evidence from field experiments. Animal Cognition, 6: 213-223. Lycett, S.J. et al. (2010) Are behavioral differences among wild chimpanzee communities genetic or cultural? An assessment using tool-use data and phylogenetic methods. American Journal of Physical Anthropology, 142:461-467. Huffman, M.A. et al. (2008) Cultured monkeys. Social learning cast in stones. Current Directions in Psychological Science, 17: 410-414. Sapolsky, RM & Share, LJ. (2004) A pacific culture among wild baboons:its emergence and transmission. PLoS Biology, Apr., 2(4): E101. Longer Readings: O’Malley, R.C. et al. (2012) The appearance and spread of ant fishing among the Kasekela chimpanzees of Gombe. Current Anthropology, 53: 650-663. McGrew, W.C. (2007) New wine in new bottles: Prospects and pitfalls of cultural primatology. Journal of Anthropological Research, 63:167-183. Nishida, T. et al. (2009) Emergence, propagation or disappearance of novel behavioral patterns in the habituated chimpanzees of Mahale: a review. Primates, 50: 23-36.

Seminar #6: Hunting Individual References (to be assigned for the discussion seminars): Alvard, M. et al. (1997) The sustainability of subsistence hunting in the Neotropics. Conservation Biology, 11: 977-982. Gilby, I.C. et al. (2008) Economic profitability of social predation in wild chimpanzees: Individual variation promotes cooperation. Animal Behaviour, 75: 351-360. Hawkes, K. & Bird, R. (2002) Showing off, handicap signalling, and the evolution of men’s work. Evolutionary Anthropology, 11: 58-667. Kaplan, H. & Hill, H. (1985) Hunting ability and reproductive success among male Ache foragers. Current Anthropology, 26: 131-133. Milton, K. (1999) A hypothesis to explain the role of meat-eating in human evolution. Evolutionary Anthropology, 8: 11-21. Mitani, J.C. & Watts, D.P. (2001) Why do chimpanzees hunt and share meat? Animal Behaviour, 61: 915-924. Marlowe, F.W. (2010) Chapter 5. Foraging. In: Marlowe, The Hadza. University of California Press, pp. 101-131. Pruetz, J.D. & Bertolani, P. (2007) Savanna chimpanzees, Pan troglodytes verus, hunt with tools. Current Biology, 17: 412-417. Surbeck, M. & Hohmann, G. (2008) Primate hunting by bonobos at LuiKotale, Salonga National Park. Current Biology, 18: R906-908.


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Watts, D.P. (2008) Scavenging by chimpanzees at Ngogo and the relevance of chimpanzee scavenging to early hominin behavioral ecology. Journal of Human Evolution, 54: 125-133. Hill, K. & Kintigh, K. (2009) Can anthropologists distinguish good and poor hunters? Implications for hunting hypothesis, sharing conventions, and cultural transmission. Current Anthropology, 50: 369-377. Longer Readings: Alvard, M. (1993) Testing the ‘ecologically noble savage’ hypothesis: Interspecific prey choice by Piro hunters of Amazonian Peru. Human Ecology, 21: 355-387. Pobiner, B.L. et al. (2007) Taphonomic analysis of skeletal remains from chimpanzee hunts at Ngogo, Kibale Natl. Park, Uganda. J. Human Evolution, 52: 614-636. Rose, L. (1997) Vertebrate predation and food sharing in Cebus and Pan. International Journal of Primatology, 18:727-765. Bird, R.B. & Bird, D.W. (2008) Why women hunt. Risk and contemporary foraging in a Western Desert aboriginal community. Current Anthropology, 49: 655-693. Stanford, C.B. & Bunn, HT (eds.) (2001) Meat Eating and Human Evolution. Oxford UP.

Seminar #7: Cannibalism Starter References (everyone to read): White, T. D. (2001) Once were cannibals. Scientific American, Aug., pp. 48-55. Kolata, G. (1986) Anthropologists suggest cannibalism is a myth. Science 232:1497-1500. Individual References (to be assigned for the discussion seminars): Defleur, A. et al. (1999) Neanderthal cannibalism at Moula-Gueray, Ardeche, France. Science, 232: 1497-1500. Dellatore, D. et al. (2009) Two cases of mother-infant cannibalism in orangutans. Primates, 50: 277-281. Durham, W.H. (1991) Coevolution. Stanford Univ. Press, pp. 393-409. [Fore] Fowler, A. & Hohmann, G. (2010) Cannibalism in wild bonobos (Pan paniscus) at Lui Kotale. American Journal of Primatology, 71: 1-6. Goodall, J. (1977) Infant killing and cannibalism in free-living chimpanzees. Folia Primatologica, 28: 259-282. Marlar, R.A. et al. (2000) Biochemical evidence of cannibalism at a prehistoric Puebloan site in southwestern Colorado. Nature, 407: 74-78. Macbeth, H. et al. (2007) Cannibalism: No myth, but why so rare? In: Consuming the Inedible, MacClancy, J. et al. (eds.), Berghahn, pp. 189-203. Petrinovich, L. (2000) The Cannibal Within. Pp. 22-28, 35-40. [Donner Pass]


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Petrinovich, L. (2000) The Cannibal Within. Pp. 91-101. [Aztec] Reinhard, K.J. (2006) A coprological view of ancestral pueblo cannibalism. American Scientist, 94: 254-261. Watts, D.P. & Mitani, J. C. (2000) Infanticide and cannibalism by male chimpanzees at Ngogo, Kibale National Park, Uganda. Primates, 41: 357-366. Longer Readings: Darling, J.A. (1998) Mass inhumation and the exhumation of witches in the American Southwest. American Anthropologist, 100: 732-752. DeGusta, D. (1999) Fijian cannibalism: Osteological evidence from Novatu. American Journal of Physical Anthropology, 110: 215-241. Carbonell, E. et al. (2010) Cultural cannibalism as a paleoeconomic system in the European Lower Pleistocene. Current Anthropology 51, 539-549. Harner, M. (1977) The ecological basis for Aztec sacrifice. American Ethnologist, 4: 117-135. Kantner, J. (1999) Survival cannibalism or socio-political intimidation? Explaining post mortem mutilation in the American Southwest. Human Nature, 10: 1-50 Seminar #8: Warfare Starter Reference (everyone to read): Mitani, J. et al. (2010) Lethal intergroup aggression leads to territorial expansion in wild chimpanzees. Current Biology 20: R507-R508. Individual References (to be assigned for the discussion seminars): Aureli, F. et al. (2006) Raiding parties of male spider monkeys: Insights into human warfare? American Journal of Physical Anthropology, 131: 486-497. Fashing, P. (2001) Male and female strategies during intergroup encounters in guerezas: evidence for resource mediated through males and a comparison with other primates. Behavioral Ecology and Sociobiology, 50: 219-230. Kelly, R. (2005) The evolution of lethal intergroup violence. Proceedings of the National Academy of Sciences (USA), 102: 15294-15298. Nunn, C. & Deaner, R. (2004) Patterns of participation and free riding in territorial conflicts among free-ranging lemurs (Lemur catta). Behavioral Ecology and Sociobiology, 57: 50-61. Roscoe, P. (2007) Intelligence, coalitional killing, and the antecedents of war. American Anthropologist, 109: 485-495. Silk, J. (2002) Practice random acts of aggression and senseless acts of intimidation: the logic of status contests in small groups. Evolutionary Anthropology, 11: 221-225. Williams, J. et al. (2002) Female competition and male territorial behaviour influence female chimpanzees’ ranging patterns. Animal Behaviour, 63: 347-360.


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Wilson, M. et al. (2001) Does participation in intergroup conflict depend on numerical assessment, range location, or rank for wild chimpanzees? Animal Behaviour 61: 1203-1216. Wilson, M.L. et al. (2001) Chimpanzees and the mathematics of battle. Proceedings of the Royal Society of London, Biological Sciences, B 269: 1107-1112. Wrangham, R. (1999) Is military incompetence adaptive? Evolution and Human Behavior, 20: 3-14. Wrangham, R. et al. (2006) Comparative rates of violence in chimpanzees and humans. Primates, 47:14-26. Longer Readings: Kitchen, D. et al. (2004) Factors mediating inter-group encounters among savannah baboons. Behaviour, 141: 197-218. Perry. S. (1996) Intergroup encounters in wild white-faced capuchins (Cebus capucinus). International Journal of Primatology, 17: 309-330. Wadley, R. (2003) Lethal treachery and the imbalance of power in warfare and feuding. Journal of Anthropology Research, 59: 531-554. Watts, D. & Mitani, J. (2001) Boundary patrols and intergroup encounters in wild chimpanzees. Behaviour, 138: 299-327. Watts, D. et al. (2006) Lethal intergroup aggression by chimpanzees in Kibale National Park, Uganda. American Journal of Primatology, 68: 161-180. Wilson, M. & Wrangham, R. (2003) Intergroup relations in chimpanzees. Annual Review of Anthropology, 32: 363-392. Wrangham, R. (1999) Evolution of coalitionary killing. Yearbook of Physical Anthropology 42:1-39. (See Am. J. Phys. Anthropol.) Wrangham, R. (2004) Collective violence: comparisons between youths and chimpanzees. Annals of the New Academy of Sciences, 1036: 233-256. Seminar #9: Technology Starter Reference (everyone to read): McGrew, W. (2010) Chimpanzee technology. Science 328: 579-580. Individual References (to be assigned for the discussion seminars): Biro D, Haslam M, Rutz C. (2013) Tool use as adaptation. Phil Trans R Soc B 368: 20120408. (Introduction, but all the articles in this issue are relevant). Gumert, M.D. et al. (2009) The physical characteristics and usage patterns of stone axe and pounding hammers used by long-tailed macaques in the Andaman Sea region of Thailand. American Journal of Primatology, 71: 594-608. Haslam, M. et al. (2009) Primate archaeology. Nature, 460: 339-344. Haslam M., Gumert M., Biro D., Carvalho S., Malaivijitnond S. (2013). Use-Wear patterns on wild macaque stone tools reveal their behavioural history. PLoS ONE 8(8): e72872. doi:10.1371/journal.pone.0072872


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Hernandez-Aguilar, A. et al. (2007) Savanna chimpanzees use tools to harvest the underground storage organs of plants. Proceedings of the National Academy of Sciences (USA), 104:19210-19213. McGrew, W.C. (1987) Tools to get food: The subsistants of Tasmanian aborigines and Tanzanian chimpanzees compared. Journal of Anthropological Res., 43: 247-258. Mercader, J. et al. (2007) 4,300-year-old chimpanzee sites and the origins of percussive stone technology. Proc. Natl. Acad. Sciences (USA), 104: 3043-3048. Osvath, M. (2009) Spontaneous planning for future stone throwing by a male chimpanzee. Current Biology, 19: R190-191. Ottoni, E.B. & Izar, P. (2008) Capuchin monkey tool use: Overview and implications. Evolutionary Anthropology, 17: 171-178. Pruetz, J.D. & Bertlolani, P. (2007) Savanna chimpanzees hunt with tools. Current Biology, 17: 412-417. Roffman, I. et al. (2012) Stone tool production and utilization by bonobo-chimpanzees (Pan paniscus). Proc. Natl. Acad. Sci. (USA), 109: 14500-14503. Sanz, C. et al. (2009) Design complexity in termite-fishing tools of chimpanzees (Pan troglodytes). Biology Letters, 5: 293-296. Seed, A. & Byrne, R.W. (2010) Animal tool-use. Current Biology, 20: 1032-1039. Longer Readings: Sanz, C.M. & Morgan, D.B. (2007) Chimpanzee tool technology in the Goualougo Triangle, Republic of Congo. Journal of Human Evolution, 52: 420-433. Shumaker, R.W. et al. (2011) Animal Tool Behavior. 2nd edit. Johns Hopkins UP. Mannu, M. & Ottoni, E. (2009) The enhanced tool-kit of two groups of wild bearded capuchin monkeys in the Caatinga: Tool making, associative use, and secondary tools. American Journal of Primatology, 71: 242-251. Wynn, T.G. et al. (2011) “An ape’s view of the Oldowan” revisited. Evolutionary Anthropology, 20:181-197. Seminar # 10: Family Individual References (to be assigned for the discussion seminars): Bender, C.E. et al. (2009) Evidence of teaching in Atlantic spotted dolphins by mother dolphins foraging in the presence of their calves. Animal Cognition, 12: 43-53. Burkart, J. et al. (2009) Cooperative breeding and human cognitive evolution. Evolutionary Anthropology, 18: 175-186. Gomes, C.M. & Boesch, C. (2009) Wild chimpanzees exchange meat for sex on a long-term basis. PLoS One, 4(4)e5116- Haak, W. et al. (2008) Ancient DNA, strontium isotopes, and osteological analyses shed light on social and kinship organization of the later Stone Age. Proceedings of the National Academy of Sciences USA, 105: 18226-18231.


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Hockings, K.J. et al. (2007) Chimpanzees share forbidden fruits. PLoS One, Issue 9, e886- Langergraber, K. et al. (2007) The limited impact of kinship on cooperation in wild chimpanzees. Proceedings of the National Academy of Science USA, 104: 7786-7790. Lonsdorf, E.V. et al. (2004) Sex differences in learning in chimpanzees. Nature, 428: 715-716. Mitani, J.C. & Watts, D. (1997) The evolution of non-maternal caretakers among the anthropoid primates…Behavioral Ecology and Sociobiology, 40: 213-220. Muller, M. et al. (2009) Testosterone and paternal care in East African foragers and pastoralists. Proceedings of the Royal Society, B, 276: 347-354. Murray, C.M. et al. (2008) Adult male chimpanzees inherit maternal ranging patterns. Current Biology, 18:R166-167. Surbeck, M. et al. (2010) Mothers matter! Maternal support, dominance status, and mating success in male bonobos. Proceedings of the Royal Society, B, 278: 590-598. Seminar # 11: Language Individual References (to be assigned for the discussion seminars): Slocombe, K. & Zuberbuehler, K. (2007) Chimpanzees modify recruitment screams as a function of audience composition. Proeedings of the. National Academy of Sciences USA, 104: 17228-17233. Slocombe, K. & Zuberbuehler, K. (2005) Functional referential communication in a chimpanzee. Current Biology, 15: 1779-1784. Clarke, E. et al. (2006) The syntax and meaning of wild gibbon songs. PLOS One, 20 Dec.;1;e73. Corballis, M.J. (1999) The gestural origins of language. American Scientist, 87: 138-145. Boesch, C. (1991) Symbolic communication in wild chimpanzees. Human Evolution, 6: 81-90. Hauser, M.D. et al. (2002) The faculty of language: What it is, who has it, and how did it evolve? Science, 298: 1569-1579. Pagel, M. (2009) Human language as a culturally transmitted replicator. Nature Reviews Genetics, 10: 405-415. Pollick, A.S. & de Waal, F.B.M. (2007) Ape gestures and language evolution. Proceedings of the National Academy of Sciences USA, 104: 8184-8189. Savage-Rumbaugh, E.S. et al. (1978) Symbolic communication between two chimpanzees. Science, 201: 641-644. Seyfarth, R.M. et al. (1980) Monkey responses to three different alarm calls: Evidence of predator classification and semantic communication. Science, 210: 801-803. Stout, D. & Chaminade, T. (2012) Stone tools, language and the brain in human evolution. Philosophical Transactions of the Royal Society, B, 367-75-87. Longer Readings:


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Aiello, L.C. & Dunbar, R.I.M. (1993) Neocortex size, group size, and the evolution of language. Current Anthropology, 334: 184-193. Crockford, C. et al. (2004) Wild chimpanzees produce group-specific calls: A case for vocal learning? Ethology, 110: 221-243. Lieberman, P. (2007) Evolution of human speech. Current Anthropology, 48: 39-66. MacLarnon, A.M. & Hewitt, G.P. (1999) The evolution of human speech: The role of enhanced breathing control. American Journal of Physical Anthropology, 109: 341-364. Seminar #12: Intelligence Individually References (to be assigned for discussion seminar): Byrne, R.W. & Corp, N. (2004) Neocortex size predicts deception rate in primates. Proceedings of the Royal Society of London, B, 271: 1693-1699. Di Fiore, A. & Suarez, S.A. (2007) Route-based travel and shared routes in sympatric spider and woolly monkeys: cognitive and evolutionary implications. Animal Cognition, 10: 317-329.

Emery, N.J. & Clayton, N.S. (2004) The mentality of crows: convergent evolution of intelligence in corvids and apes. Science, 306: 1903-1907.

Herrman, E. et al. (2007) Humans have evolved specialized skills of social cognition: The cultural intelligence hypothesis. Science, 317: 1360-1366.

Janmaat, K.R.L. et al. (2006) Primates take weather into account when searching for fruits. Current Biology, 16:1232-1237. Mendes et al. (2007) Raising the level: Orangutans use water as a tool. Biology Letters, 3: 453-455. Normand, E. et al. (2009) Forest chimpanzees remember the location of numerous fruit trees. Animal Cognition, 12: 797-807. Proctor, D. et al. (2013) Chimpanzees play the ultimatum game. Proceedings of the National Academy of Sciences USA, 110: 1978-

Rosati, A.G. et al. (2007) The evolutionary origins of human patience: Temporal preferences in chimpanzees, bonobos and human adults. Current Biology, 17: 1663-1668.

Russon, A.E. & Galdikas, B. (1993) Imitation in ex-captive orangutans. Journal of Comparative Psychology, 107: 147-161.

Salwiczek, L.H. et al. (2012) Adult cleaner wrasse outperform capuchin monkeys, chimpanzees, and orangutans in a complex foraging task derived from cleaner-client reef fish cooperation. PLoS One, 7(11): e49068. Classic References: Byrne, R.W. (2001) Social and technical forms of social intelligence. In: Tree of Origin, de Waal, F. (ed.), Harvard Univ. Press, pp. 145-172. Dunbar, R. (1988) The social brain hypothesis. Evolutionary Anthropology, 6: 178-190. Milton, K. (1981) Distribution of tropical food plants as an evolutionary stimulus to primate mental development. American Anthropologist, 83: 534-548.


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Humphrey, N.K. (1976) The social function of intellect. In: Growing Points in Ethology, Bateson, P.P.G. & Hinde, R.A. (eds.), Cambridge Univ. Press, pp. 303-317. Whiten, A. & Byrne, R.W. (eds.) (1997) Machiavellian Intelligence II: Extensions and Evaluations. Cambridge Univ. Press.

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