After having travelled repeatedly to specific targets, individuals benefited from establishing a set of highly efficient habitual routes that would support travelling along the shortest path to reach multiple targets from multiple starting locations ( Warren, 2019). Chrastil and Warren (2014) indicated that humans rely strongly on route-based maps but incorporate metric and angular information into their movement decisions. Even though route-based maps constrain an animal's movement options to a series of pre-established routes ( Di Fiore and Suarez, 2007), they have been widely reported across animal taxa (birds: Guilford and Biro, 2014 mammals: Newmark and Rickart, 2012 Trapanese et al., 2019 insects: Mangan and Webb, 2012 Baddeley et al., 2012). Nodes are often associated with emergent trees or ridges, where animals likely increase their visual access to the surroundings and may decide where to go next ( Presotto et al., 2018). route networks Perna and Latty, 2014 Trapanese et al., 2019). Alternatively, route-based maps are composed of a series of habitually used routes that interconnect pairs of relevant locations – called ‘nodes’ – across an animal's home ranges (i.e. Calculating novel paths or detours between known locations is supported by Euclidean maps, providing a high level of navigation flexibility ( McNaughton et al., 2006 but see Warren, 2019). Euclidean maps are built upon the knowledge of distances and directions among locations within animals' habitats in a globally consistent coordinate system ( Gallistel, 1990 O'Keefe and Nadel, 1978). In turn, sophisticated cognitive maps likely support flexible and efficient movement patterns that enhance foraging efforts of individuals or groups ( Behrens et al., 2018 Milton, 1981).Įven though the concept of cognitive maps has been a subject of debate during the last decades ( Bennett, 1996 Collett and Collett, 2006 Warren, 2019), there has been a consensus postulating that navigation flexibility can be considered along a continuum with one end representing Euclidean maps and the other route-based maps ( Byrne, 2000 Warren, 2019 Warren et al., 2017). ![]() ![]() The level of sophistication of these cognitive maps has been argued to be analogous with the cognitive capacity of a species ( Poucet, 1993 Tolman, 1948 Warren, 2019). Among these navigational strategies, some species generate internal representations of the space wherein they live, known as ‘cognitive maps’ ( Tolman, 1948). celestial cues to orientate migratory routes: Foster et al., 2018 visual landmarks associated to foraging sites: Zeil, 2012 path integration using nest location as reference: Heinze et al., 2018). Animals have developed a wide variety of navigational strategies to make use of environmental information and reach relevant biological locations (e.g. Living in tropical forests involves coping with a complex matrix of environmental information characterized by highly variable spatial and temporal distributions of food resources ( Janmaat et al., 2016 Levey, 1988 Morales et al., 2010). Our findings not only expand the use of metric information during route navigation to non-human animals, but also highlight the importance of considering efficient route-based navigation as a cognitively demanding mechanism. In addition, we found that the structure of observed route networks was more complex and efficient than simulated route networks, suggesting that black howler monkeys incorporate metric information into their cognitive map. Our results indicated that black howler monkeys engaged in constrained movement patterns characterized by a high path recursion tendency, which limited their capacity to travel in straight lines and approach feeding trees from multiple directions. We simulated correlated random walks mimicking the ranging behaviour of the study subjects and tested for differences between observed and simulated movement patterns. ![]() We collected 3104 h of ranging and behavioural data on five groups of black howler monkeys ( Alouatta pigra) at Palenque National Park, Mexico, from September 2016 through August 2017. Here, we examined the properties of the cognitive map used by a wild population of primates by testing a series of cognitive hypotheses against spatially explicit movement simulations. distances and angles between locations) in route-based cognitive maps would likely enhance an animal's navigation efficiency, there has been no evidence of this strategy reported for non-human animals to date. Even though incorporating metric information (i.e. Euclidean cognitive maps) however, constrained movements along habitual routes are the most commonly reported navigation strategy. Generally, flexible navigation is hypothesized to be supported by sophisticated spatial skills (i.e. ![]() When navigating, wild animals rely on internal representations of the external world – called ‘cognitive maps’ – to take movement decisions.
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