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Field studies of Pan troglodytes reviewed and comprehensively mapped, focussing on Japan’s contribution to cultural primatology

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Abstract

Field studies done over decades of wild chimpanzees in East, Central and West Africa have yielded impressive, cumulative findings in cultural primatology. Japanese primatologists have been involved in this advance from the outset, over a wide variety of topics. Here I review the origins and development of field studies of Pan troglodytes, then assess their progress based on analogy between cultural primatology and cultural anthropology, through four stages: natural history, ethnography, ethnology, and intuition. Then, I focus on six topics that continue to yield informative debate: technology, universals, nuanced variation, archaeology, applied primatology, and ecology. Finally, I offer a map of sites of field study of wild chimpanzees. It is clear that Japanese primatologists have made a significant contribution to East–West scientific exchange, especially at the field sites of Bossou and Mahale.

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Acknowledgments

For helpful advice and many other things, sometimes over 40 years of chimpanzee field work, I thank Pamela Baldwin, Paco Bertolani, Stella Brewer, Susana Carvalho, Anthony Collins, Adriana Hernandez-Aguilar, Michael Huffman, Kevin Hunt, Agumi Inaba, Jun’ichiro Itani, Kenji Kawanaka, Kathelijne Koops, Linda Marchant, Tetsuro Matsuzawa, Norman McBeath, Michio Nakamura, Toshisada Nishida, Robert O’Malley, Alejandra Pascual-Garrido, Alex Piel, Crickette Sanz, Fiona Stewart, Yukimaru Sugiyama, Caroline Tutin, Shigeo Uehara, and Richard Wrangham; special thanks for photographs used here go to Susana Carvalho, Kim Hockings, Agumi Inaba, Kathelijne Koops, Stella Brewer Marsden, Tetsuro Matsuzawa, Michio Nakamura, Caroline Tutin; Kevin Hunt and Tetsuro Matsuzawa for helpful comments on the manuscript; Lauren Sarringhaus for pioneering contributions to an earlier, unpublished effort; Agumi Inaba for manuscript preparation; Alex Piel for enabling the mapping. This paper was originally presented as ‘Chasing after Chimpanzees: Confessions of a Chimpaholic’ at the 4th International Symposium on Primatology and Wildlife Science, Kyoto University, 21–22 July 2015.

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  1. Division of Biological Anthropology, Department of Archaeology and Anthropology, University of Cambridge, Fitzwilliam Street, Cambridge, CB2 1QH, UK

    William C. McGrew

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Correspondence to William C. McGrew.

Appendices

Appendix 1

Field sites of chimpanzee research

Listed alphabetically by researcher’s labels for sites. Latitude listed as North (positive value) or South (negative value) of the Equator, in minutes and decimal degrees. Ditto for longitude (E, positive value, or W, negative value). Most older sources list geographical coordinates in degrees and minutes. When degrees or minutes are a range, then mid-point is taken. All were converted into minutes and decimal degrees, using http://rapidtables.com/convert/degrees-minutes-seconds-to-degrees.htm.

Some researchers listed no geographical coordinates for their sites; whenever possible these were obtained by personal communication or were derived or calculated from other sources, e.g. interpolated from maps of study sites. As a last resort, I took coordinates from websites that yield them when place names are input (http://www.latlong.net, http://www.geoplaner.com, http://www.worldatlas.com). Unfortunately, for some sites (n = 11), geographical coordinates remain to be found; such sites are listed below but otherwise left blank.

Some early study sites come from Baldwin and Teleki (1973), but they gave no geographical coordinates. Ditto for Wrangham et al. (1994), who provided a longer list of sites (n = 41). Likewise, Shumaker et al. (2011) listed 59 sites where chimpanzee tool behavior has been observed, but without coordinates.

Baldwin and Teleki differentiated between short-term vs. long-term sites, but the difference seems arbitrary, so both are included here. Wrangham et al. and Shumaker et al. gave no criteria. Definition of a site is sometimes problematic, but self-identified surveys (e.g. Ogawa et al. 2006) are excluded. (For extensive coverage of surveys, see, e.g., Kormos et al. 2003).

Some sites have ‘changed’ names (e.g. Kasoje to Mahale) and others have more than one name, sometimes by differences in spelling (Gouera vs. Gouela). Others, usually neighboring sites, may overlap, often because of differing degrees of specificity given by researchers. Finally, when the same population of chimpanzees (e.g. Kibale) yields multiple sites, I have listed different communities as study sites.

For bibliographic details of sources of geographical coordinates, see references in Appendix 2.

  1. 1.

    Abuminzok-Aninzok, Jones and Sabater Pi 1971.

  2. 2.

    Ayamiken, N +2.116667, E +10.01667, Jones and Sabater Pi 1969.

  3. 3.

    Baboon Island, N +13.63889, W −14.95944, Brewer and McGrew 1990.

  4. 4.

    Bafing, N +12.578020, W −10.307343, Duvall 2008.

  5. 5.

    Bai Hokou, N +2.85, E +16.46667, Fay and Carroll 1994.

  6. 6.

    Bakoun, N +11.9, W −12.5, Kuehl et al. 2016.

  7. 7.

    Bandafassi, N +12.53872, W −12.31060, Gaspersic and Pruetz 2011.

  8. 8.

    Bassa, N +6.25, W −10.36667, Hannah and McGrew 1987.

  9. 9.

    Bateke, S −2.167566, E +14.041160, Kuehl et al. 2016.

  10. 10.

    Belinga, N +1.1, E +13.2, McGrew and Rogers 1983.

  11. 11.

    Beni, N +0.49901, E +29.45275, Kortlandt 1966.

  12. 12.

    Bhukalai, S −5.43556, E +30.73361, Yoshikara and Ogawa 2015.

  13. 13.

    Bia, N +6.16667, W −2.5, Jeffrey 1975.

  14. 14.

    Bilenge, S −6.033333, E +29.73333, Collins and McGrew 1985.

  15. 15.

    Bili, N +4.602616, E +25.868755, Hicks et al. 2014.

  16. 16.

    Boe, N +11.750000, W −14.216666, Kuehl et al. 2016.

  17. 17.

    Bossou, N +7.6500, W −8.5000, Yamakoshi 1998.

  18. 18.

    Budongo, N +1.750000, E +31.500000, Eggeling 1947.

  19. 19.

    Bulindi, N +1.483333, E +31.48333, McLennan 2011.

  20. 20.

    Bwindi, S −1.008333, E −29.708330, Stanford and Nkurunungi 2003.

  21. 21.

    Campo, N +2.637160, E +10.463100, Hoshino 1985.

  22. 22.

    Caiquene-Cadique, N +11.21667, W −15.08333, Bessa et al. 2015.

  23. 23.

    Comoe, N +8.841651, W −3.707890, Fischer and Gross 1999.

  24. 24.

    Conkouati, S −3.800000, E +11.416667, Kuehl et al. 2016.

  25. 25.

    Dere, N +7.60849, W −8.18913, Granier et al. 2007.

  26. 26.

    Diecke, N +7.5, W −8.95, Carvalho 2011.

  27. 27.

    Dipikar, N +2.23333, E +10.05, Jones and Sabater Pi 1969.

  28. 28.

    Dja, N +3.2, E +13.21667, Dupain et al. 2004.

  29. 29.

    Djinji, Takenoshita 1996.

  30. 30.

    Djouroutou, N +5.366667, W −7.283333, Kuehl et al. 2016.

  31. 31.

    Dzanga-Ndoki, N +3.436861, E +16.346379, Lilly et al. 2002.

  32. 32.

    Eastern Congo, Rahm 1967.

  33. 33.

    Ebo, N +4.319000, E +10.330000, Morgan and Abwe 2006.

  34. 34.

    Filabanga, S −5.383333, E +30.08333, Kano 1971.

  35. 35.

    Fongoli, N +12.666, W −12.166, Pruetz and Bertolani 2007.

  36. 36.

    Gashaka, N +7.566667, E +11.700000, Pascual-Garrido et al. 2012.

  37. 37.

    Gba, N +7.316667, W −8.933333, Ohashi 2006.

  38. 38.

    Gishwati, S −1.747220, E +29.427028, Kuehl et al. 2016.

  39. 39.

    Gombe, S −4.695511, E +29.61643, Goodall 1996.

  40. 40.

    Goualougo, N +2.100000, E +16.866667, Sanz et al. 2014.

  41. 41.

    Gouela, N +7.59716, W −8.38007, Granier et al. 2007.

  42. 42.

    Grebo, N +5.405889, W −7.732278, Kuehl et al. 2016.

  43. 43.

    Guga, Takenoshita 1996.

  44. 44.

    Haut Niger, N +10.63333, W −10.66667, Fleury-Brugiere et al. 2002.

  45. 45.

    Hoima, N +1.51667, E +31.35, McClennan 2008.

  46. 46.

    Ipassa, N +0.5666667, E +12.86667, Hladik 1973.

  47. 47.

    Ishasha, S −0.6166667, E +29.65, Sept 1992.

  48. 48.

    Issa, S −5.833333, E +30.93333, Stewart and Piel 2014.

  49. 49.

    Kabogo, S −7.129425, E +29.694487, Kuehl et al. 2016.

  50. 50.

    Kabogo Point, S −5.5, E +29.78333, Kano 1971.

  51. 51.

    Kahuzi-Biega, S −2.155465, E +28.726501, Yamagiwa 1996.

  52. 52.

    Kalinzu, N −0.283, E +30.117, Aruga et al. 2015.

  53. 53.

    Kambi, Nissen 1931.

  54. 54.

    Kanfarande, N +10.837576, W −14.53931, Leciak et al. 2005.

  55. 55.

    Kanka Sili, Dunnett et al. 1970.

  56. 56.

    Kanton, Kortlandt and Holzhaus 1987.

  57. 57.

    Kanyawara, N +0.5667, E +30.3667, Wrangham 1996.

  58. 58.

    Kasakati, S −5.4, E +30.1667, Izawa and Itani 1966.

  59. 59.

    Kasokwa, N +1.65, E +31.65, Reynolds et al. 2003.

  60. 60.

    Kayan, N +12.8833, W −12.293810, Kuehl et al. 2016

  61. 61.

    Kharakhena, N +12.91417, W −11.53639, Boyer-Ontl and Pruetz 2014.

  62. 62.

    Kibira, S −2.95, E +29.45, Hakizimana et al. 2015.

  63. 63.

    Kilimi, N +9.716667, W −12.53333, Harding 1984.

  64. 64.

    Korup, N +5.286061, E +9.057099 Kuehl et al. 2016.

  65. 65.

    Kpala, N +7.166667, W −8.683333, Ohashi 2015.

  66. 66.

    Kyambura, S −0.15, E +30.033, Krueger et al. 1998.

  67. 67.

    La Belgique-Dja, N +3.416670, E +13.15, Deblauwe and Janssens 2008.

  68. 68.

    Loango, S −2.090000, E +9.48, Oelze et al. 2014.

  69. 69.

    Lobeke, N +2.291667, E +15.8667, Usongo 1998.

  70. 70.

    Lope, S −0.166700, E +11.583300, Tutin 1996.

  71. 71.

    Lossi, N +0.2333333, E +14.5, Bermejo and Illera 1999.

  72. 72.

    Ma’an, Matthews and Matthews 2002.

  73. 73.

    Mahale, S −6.1167, E +29.7333, Nishida 1996.

  74. 74.

    Maiombe, S −4.666667, E +12.75, Ron 2011.

  75. 75.

    Mayang, N +2.083333, E +9.916667, Gonzalez-Kirchner and de la Maza 1992.

  76. 76.

    Mbe, N +6.257967, W +9.041748, Kuehl et al. 2016.

  77. 77.

    Mboete, N +1.316667, E +9.6, Takemoto et al. 2005.

  78. 78.

    Mbomo, N +0.42866, E +14.69668, Kano and Asato 1994.

  79. 79.

    Minkebe, N +1.671667, E +12.75972, Huijbregts et al. 2003.

  80. 80.

    Mitumba, S −4.633333, E +29.633333, Mjungu et al. 2006.

  81. 81.

    Mondika, N +2.366667, E +16.26667, Lilly et al. 2002.

  82. 82.

    Moukalaba, S −2.366667, E +10.5, Nishimura et al. 2003.

  83. 83.

    Mt. Alen, N +1.71667, E +10.55, Sabater Pi 1974.

  84. 84.

    Mt. Assirik, N +12.8833, W −12.7667, McGrew et al. 1981.

  85. 85.

    Mt. Bagnomba, Gaspersic 2007.

  86. 86.

    Mt. Cameroon, N +4.217340, E +9.172730, Kuehl et al. 2016.

  87. 87.

    Mt. Nimba, N +7.566667, W −8.50, Granier and Martinez 2011.

  88. 88.

    Mt. Sangbe, N +7.952841, W −7.262975, Kuehl et al. 2016.

  89. 89.

    Ndakan, N +2.333333, E +16.16667, Fay and Carroll 1994.

  90. 90.

    Ndoumbi, N +0.3666667, E +14.53333, Bermejo and Illera 1999.

  91. 91.

    Nerebili, N +10.059594, W −12.745334, Nissen 1931.

  92. 92.

    Ngel Nyaki, N +6.883333, E +11.26667, Dutton and Chapman 2014.

  93. 93.

    Ngogo, N +0.546562, E +30.432130, Mitani 2009.

  94. 94.

    Ngotto, N +4.016667, E +17.06667, Hicks et al. 2005.

  95. 95.

    Ngoubunga, Fay and Carroll 1994.

  96. 96.

    Nguye, S −5.21667, E +30.45139, Yoshikara and Ogawa 2015.

  97. 97.

    Nouabale-Ndoki, N +2.583333, E +16.6, Kuroda 1996.

  98. 98.

    Ntale, N +5.25, E +9.566667, Ingmanson 1997.

  99. 99.

    Nyungwe, S −2.490000, E +29.292778, Gross-Camp and Kaplin 2005.

  100. 100.

    Odzala, N +1.15, E +15.46667, Devos et al. 2002.

  101. 101.

    Okorobiko, N +1.466667, E +9.866667, Jones and Sabater Pi 1969.

  102. 102.

    Otzi, N +3.708333, E +31.8667, Davenport et al. 2001.

  103. 103.

    Petit Loango, S −2.283333, E +9.583333, Furuichi 2000.

  104. 104.

    Pongara, N +0.302249, E +9.311996, Petre et al. 2007.

  105. 105.

    Rubi-Tele, N +2.647449, E +24.856460, Kuehl et al. 2016.

  106. 106.

    Rubondo, S −2.3, E +31.83333, Matsumoto-Oda 2000.

  107. 107.

    Sangaredi, N +11.100000, W −13.766667, Kuehl et al. 2016.

  108. 108.

    Sapo, N +5.411114, W −8.414625, Anderson et al. 1983.

  109. 109.

    Sebitoli, N +0.633333, E +30.4, Bortolamiol et al. 2014.

  110. 110.

    Semliki, N +0.8333, E +30.3333, Hunt and McGrew 2002.

  111. 111.

    Seringbara, N +7.219052, W −8.493388, Koops et al. 2013.

  112. 112.

    Sobeya, W +10.259386, W −11.709126, Kuehl et al. 2016.

  113. 113.

    Tai, N +5.8667, W −7.3333, Boesch 1996.

  114. 114.

    Tenkere, N +9.783333, W −12.00, Hanson-Alp et al. 2003.

  115. 115.

    Tiwai, N +7.683333, W −11.3333, Hanson-Alp et al. 2003.

  116. 116.

    Tongo, S −1.3, E +29.13333, Lanjouw 2002.

  117. 117.

    Toro, Bolwig 1959.

  118. 118.

    Tubila, S −5.016667, E +30.1, Ogawa et al. 2011.

  119. 119.

    Ugalla, S −5.316700, E +30.616700, Moore 1996.

  120. 120.

    Yeale, N +7.555, W −8.465278, Shimada 2000.

Acknowledgements

I thank colleagues who supplied supplementary information in response to my enquiries: Jim Anderson, Chris Duvall, Michael Fay, Nicolas Granier, Marcel Hladik, Marie-Claude Huynen, Ellen Ingmanson, Jim Moore, Alejandra Pascual-Garrido, Tamar Ron, Janette Wallis, Michael Wilson.

References

Baldwin LA, Teleki G (1973) Field research on chimpanzees and gorillas: an historical, geographical, and bibliographical listing. Primates 14:315–330

Kormos R, Boesch C, Bakarr MI, Butynski TM (eds) (2003) West African chimpanzees. IUCN, Gland, p 219

Ogawa H, Moore J, Kamenya S (2006) Chimpanzees in the Ntakata and Kakung areas, Tanzania. Primate Conserv 21:97–101

Shumaker RW, Walkup KR, Beck BB (2011) Animal tool behavior. The use and manufacture of tools. Johns Hopkins University Press, Baltimore

Wrangham RW, McGrew WC, de Waal FBM, Heltne PG (eds) (1994) Chimpanzee cultures. Harvard University Press, Cambridge MA, pp xxii-xxiii

Appendix 2

Sources of geographical coordinates of chimpanzee field sites

Whenever possible, primary sources authored by researchers were used, but publications of some sites do not give geographical coordinates. Then, secondary sources were used, whenever possible (see Appendix 1 for more details).

Anderson JR, Williamson EA, Carter J (1983) Chimpanzees of Sapo forest, Liberia: density, nests, tools and meat-eating. Primates 24:594–601 (Sapo)

Aruga N, Hodaka M, Furuichi T, Hashimoto C (2015) Root eating by wild chimpanzees (Pan troglodytes) in the Kalinzu forest, Uganda: possible medicinal plant use. Pan Afr News 22:7–10 (Kalinzu)

Bermejo M, Illera G (1999) Tool-set for termite-fishing and honey extraction by wild chimpanzees in Lossi forest, Congo. Primates 40:619–627 (Lossi, Ndoumbi)

Bessa J, Sousa C, Hockings KJ (2015) Feeding ecology of chimpanzees (Pan troglodytes verus) inhabiting a forest-mangrove-savanna-agricultural matrix at Caiquene-Cadique, Cantanhez National Park, Guinea-Bissau. Am J Primatol 77:651–665 (Caiquene-Cadique)

Boesch C (1996) Appendix: great ape field sites. Tai. In: McGrew WC, Marchant LF, Nishida T (eds) Great ape societies. Cambridge University Press, Cambridge, pp 317–318 (Tai)

Bolwig N (1959) A study of the nests built by mountain gorilla and chimpanzee. South Afr J Sci 55:286–291 (Toro)

Bortolamiol S, Cohen M, Potts K, Pennec F, Rwaburindore P, Kasenene J, Seguya A, Vignaud Q, Krief S (2014) Suitable habitats for endangered frugivorous mammals: small-scale comparison, regeneration forest and chimpanzee density in Kibale National Park, Uganda. PLoS One 9:e102177 (Sebitoli)

Boyer-Ontl KM, Pruetz JD (2014) Giving the forest eyes: the benefits of using camera traps to study unhabituated chimpanzees (Pan troglodytes verus) in southeastern Senegal. Int J Primatol 35:881–894 (Kharakhena)

Brewer SM, McGrew WC (1990) Chimpanzee use of a tool set to get honey. Folia Primatol 54:100–104 (Baboon Island)

Carvalho S (2011) Diecke forest, Guinea: delving into chimpanzee behavior using stone tool surveys. In: Matsuzawa T, Humle T, Sugiyama Y (eds) The chimpanzees of Bossou and Nimba. Springer, Tokyo, pp 301-312 (Diecke)

Chancellor RL, Rundus AS, Nyandwi S (2012) The influence of seasonal variation on chimpanzee (Pan troglodytes schweinfurthii) fallback food consumption, nest group size, and habitat use in Gishwati, a montane rain forest fragment in Rwanda. Int J Primatol 33:115–133 (Gishwati)

Collins DA, McGrew WC (1985) Chimpanzees’ (Pan troglodytes) choice of prey among termites (Macrotermitinae) in western Tanzania. Primates 26:375–389 (Bilenge)

Davenport TRB, Lodu E, Howard PC, Plumptre AJ (2001) Robust chimpanzee Pan troglodytes schweinfurthii in Otzi Forest Reserve, northern Uganda. Afr Primates 5:46–48 (Otzi)

Deblauwe I, Janssens GPJ (2008) New insights into prey choice by chimpanzees and gorillas in southeast Cameroon: the role of nutritional value. Am J Primatol 135:42–5 (La-Belgique-Dja)

Devos C, Gatti S, Levrero F (2002) New record of algae feeding and scooping by Pan t. troglodytes at Lokoue Bai in Odzala National Park, Republic of Congo. Pan Afr News 9:19–21 (Odzala)

Dunnett S, van Orshoven J, Albrecht H (1970) Peaceful co-existence between chimpanzee and man in West Africa. Bijdr Tot Dierkd 40:148–153 (Kanka Sili)

Dupain J, Guislain P, Nguenang M, de Vleeschouwer K, van Elsacker L (2004) High chimpanzee and gorilla densities in a non-protected area on the northern periphery of the Dja Faunal Reserve, Cameroon. Oryx 38:209–216 (Dja)

Dutton P, Chapman H (2014) New tools suggest local variation in tool use by a montane community of the rare Nigeria-Cameroon chimpanzee, Pan troglodytes ellioti, in Nigeria. Primates 56:89–100 (Ngel Nyaki)

Duvall CS (2008) Human settlement ecology and chimpanzee habitat selection in Mali. Landsc Ecol 23:699–716 (Bafing)

Eggeling WJ (1947) Observations on the ecology of the Budongo rainforest, Uganda. J Ecol 34:20–87 (Budongo)

Fay JM, Carroll RW (1994) Chimpanzee tool use for honey and termite extraction in central Africa. Am J Primatol 34:309–317 (Bai Hokou, Ndakan, Ngoubunga)

Fischer F, Gross M (1999) Chimpanzees of the Comoe National Park, Cote d’Ivoire. Pan Afr News 6:19–20 (Comoe)

Fleury-Brugiere M-C, Brugiere D (2002) A population density estimate of the chimpanzees in the Haut-Niger National Park, Republic of Guinea. Pan Afr News 9:1–3 (Haut-Niger)

Furuichi T (2000) Possible case of predation on a chimpanzee by a leopard in the Petit Loango Reserve, Gabon. Pan Afr News 7:21–23 (Petit Loango)

Gaspersic M (2007) Savanna chimpanzees and baobab fruits. Bol Asoc Primatol Esp 14:18–21 (Mt. Bagnomba)

Gaspersic M, Pruetz JD (2011) Chimpanzees in Bandafassi arrondissement, southeastern Senegal; field surveys as a basis for the sustainable community-based conservation. Pan Afr News 18:23–25 (Bandafassi)

Gonzalez-Kirchner JP, de la Maza MS (1992) Sticks used by wild chimpanzees: a new locality in Rio Muni. Folia Primatol 58:99–102 (Mayang)

Goodall J (1996) Appendix: great ape study sites. Gombe. In: McGrew WC, Marchant LF, Nishida T (eds) Great ape societies. Cambridge University Press, Cambridge, pp 311-312 (Gombe)

Granier N, Huynen M-C, Matsuzawa T (2007) Preliminary surveys of chimpanzees in Gouela area and Dere forest, the Nimba Mountains Biosphere Reserve, Republic of Guinea. Pan Afr News 14:20–22 (Dere, Gouela)

Granier N, Martinez L (2011) Conservation issues in the Nimba Mountains. In: Matsuzawa T, Humle T, Sugiyama Y (eds) The chimpanzees of Bossou and Nimba. Springer, Tokyo, pp 381–392 (Mt. Nimba)

Gross-Camp N, Kaplin BA (2005) Chimpanzees (Pan troglodytes) seed dispersal in an afromontane forest: microhabitat influences on the postdispersal fate of large seeds. Biotropica 37:641–649 (Nyungwe)

Hannah AC, McGrew WC (1987) Chimpanzees using stones to crack open oil palm nuts in Liberia. Primates 28:31–46 (Bassa)

Hanson-Alp R, Bakarr MI, Lebbie A, Bangura KI (2003) Sierra Leone. In: Kormos R, Boesch C, Bakarr MI, Butynski TM (eds), West African chimpanzees. IUCN, Gland, Switzerland, pp 77-87 (Tenkere, Tiwai)

Harding RSO (1984) Primates of the Kilimi area, northwest Sierra Leone. Folia Primatol 42:96–114 (Kilimi)

Hakizimana D, Hambuckers A, Brotcorne F, Huynen MC (2015) Characterization of nest sites of chimpanzees (Pan troglodytes schweinfurthii) in Kibira National Park, Burundi. Afr Primates 10:1–12 (Kibira)

Hicks TC, Fouts RS, Fouts DH (2005) Chimpanzee (Pan troglodytes troglodytes) tool use in the Ngotto forest, Central African Republic. Am J Primatol 65:221–237 (Ngotto)

Hicks TC, Tranquilli S, Kuehl H, Campbell G, Swinkels J, Darby L, Boesch C, Hart J, Menken SBJ (2014) Absence of evidence is not evidence of absence: discovery of a large, continuous population of Pan troglodytes schweinfurthii in the Central Uele region of northern DRC. Biol Conserv 171:107–113 (Bili)

Hladik CM (1973) Alimentation et activite d’un groupe die chimpanzes reintroduits en foret Gabonaise. Terre Vie 27:343–413 (Ipassa)

Hoshino J (1985) Feeding ecology of mandrills (Mandrillus sphinx) in Campo Animal Reserve, Cameroon. Primates 26:248–273 (Campo)

Huijbregts B, de Wachter P, Ndong Obiang LS, Akou ME (2003) Ebola and the decline of gorilla Gorilla gorilla and chimpanzee Pan troglodytes populations in Minkebe forest, north-eastern Uganda. Oryx 37:437–443 (Minkebe)

Hunt KD, McGrew WC (2002) Chimpanzees in the dry habitats of Assirik, Senegal and Semliki Wildlife Reserve, Uganda. In: Boesch C, Hohmann G, Marchant LF (eds) Behavioural diversity in chimpanzees and bonobos. Cambridge University Press, Cambridge pp 35–51 (Semliki)

Ingmanson E (1997) Chimpanzee ant-dipping tools from west Cameroon. Am J Primatol 42:117 (abstract) (Ntale)

Izawa K, Itani J (1966) Chimpanzees of the Kasakati Basin, Tanganyika. I. Ecological study in the rainy season 1963-64. Kyoto Univ Afr Studies 1:73–156 (Kabogo Point, Kasakati)

Jeffrey SM (1975) Notes on mammals from the high forest of western Ghana (excluding Insectivora). Bull l’Inst Fond Afr Noire 37:950–973 (Bia)

Jones C, Sabater Pi J (1969) Sticks used by chimpanzees in Rio Muni, West Africa. Nature 223:100–101 (Ayamiken, Okorobiko)

Jones C, Sabater Pi J (1971) Comparative ecology of Gorilla gorilla (Savage and Wyman) and Pan troglodytes (Blumenbach) in Rio Muni, West Africa. Bibl Primatol 13:1–96 (Abuminzok-Aninzok)

Kano T (1971) The chimpanzee of Filabanga, western Tanzania. Primates 12:229–246 (Filabanga)

Kano T, Asato R (1994) Hunting pressure on chimpanzees and gorillas in the Motaba river area, northeastern Congo. Afr Study Monogr 15:143–162 (Mbomo)

Koops K, McGrew WC, Matsuzawa T (2013) Ecology of culture: do environmental factors influence foraging tool use in wild chimpanzees (Pan troglodytes verus)? Anim Behav 85:175–185 (Seringbara)

Kortlandt A (1967) Experimentation with chimpanzees in the wild. In: Starck D, Schneider R, Kuhn H-J (eds) Neue Ergebnisse der Primatologie. Gustav Fischer Verlag, Stuttgart, pp 208–224 (Beni)

Kortlandt A, Holzhaus E (1987) New data on the use of stone tools by chimpanzees in Guinea and Liberia. Primates 28:473–496 (Kanton)

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McGrew, W.C. Field studies of Pan troglodytes reviewed and comprehensively mapped, focussing on Japan’s contribution to cultural primatology. Primates 58, 237–258 (2017). https://doi.org/10.1007/s10329-016-0554-y

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