Chimpanzee behavioural diversity is spatially structured and negatively associated with genetic variation
Authors: Barratt Christopher , Gratton Paolo , Marta Silvio , Onstein Renske , Fontsere Claudia , Lester Jack , Kalan Ammie , Kulik Lars , Bocksberger Gaelle , Dieguez Paula , McCarthy Maureen , Kuhlwilm Martin , Abwe Ekwoge , Agbor Anthony , Angedakin Samuel , Assumang Alfred , Aubert Floris , Ayimisin Emmanuel , Bailey Emma , Barciela Amanda , Barubiyo Donatienne , Bessone Mattia , Brazzola Gregory , Buh Valentine Ebua , Chancellor Rebecca , Cohen Heather , Corogenes Katherine , Coupland Charlotte , Danquah Emmanuel , Deschner Tobias , Dunn Andrew , Dupain Jef , Egbe Villard , Granjon Anne-Céline , Hedwig Daniela , Hermans Veerle , Hernandez-Aguilar R. Adriana , Imong Inaoyom , Jeffery Kathryn , Jones Sorrel , Junker Jesscia , Kambere Mbangi , Kambi Mohamed , Kienast Ivonne , Kujirakwinja Deo , Langergraber Kevin , Lapuente Juan , Larson Bradley , Lee Kevin , Llana Manuel , Martin Rumen , Mbi Tanyi , Meier Amelia , Morgan Bethan , Morgan David , Neil Emily , Nicholl Sonia , Nixon Stuart , Niyigaba Protais , Normand Emmanuelle , Orbell Christopher , Ormsby Lucy , Orume Robinson , Pacheco Liliana , Piel Alex , Preece Jodie , Robbins Martha , Rundus Aaron , Sanz Crickette , Schijndel Joost van , Sommer Volker , Stewart Fiona , Tagg Nikki , Tédonzong Luc Roscelin , Tickle Alexander , Vendras Elleni , Vergnes Virginie , Vyalengerera Magloire , Welsh Adam , Wessling Erin , Willie Jacob , Wittig Roman , Yuh Yisa Ginath , Yurkiw Kyle , Zuberbühler Klaus , Andres Aida , Marques-Bonet Tomas , Boesch Christophe , Arandjelovic Mimi , Kühl Hjalmar 
Published: September 2023

Abstract
The question of how behavioural diversity in humans and other animals is shaped by the combined influence of demography, genetics, culture, and the environment receives much research attention. We take a macro-ecological approach to evaluate how chimpanzee (Pan troglodytes) behavioural diversity is spatially structured and associated with genetic diversity (i.e. heterozygosity as a proxy for effective population size) and contemporary and historic environmental context. We integrate the largest available chimpanzee behavioural and genomic datasets and apply spatially explicit Bayesian Generalised Linear Mixed Models to derive marginal effects for putative drivers and range wide spatial predictions of probability to observe behavioural traits. Contrary to expectations from neutral models of behavioural evolution, we observed a negative association of genetic diversity with behavioural diversity. This result suggests that behavioural traits may impact fitness. In contrast, we observed weaker associations of chimpanzee behavioural diversity with contemporary and historic environmental context. The very strong spatial structuring of behavioural traits is consistent with cultural transmission playing a major role in shaping chimpanzee behavioural diversity. Our analytical approach can be flexibly extended by additional candidate drivers of chimpanzee behavioural diversity, and offers a novel framework for testing competing ecological-evolutionary hypotheses across a wide variety of animal cultures.