19Many species of terrestrial animals, including primates, live in varied association with 20 the aquatic (e.g., riverine or coastal) environment. However, the benefits that each 21 species receive from the aquatic environment are thought to vary depending on their 22 social and ecological characteristics, and thus, elucidating those benefits to each species 23 is important for understanding the principles of wild animal behaviour. In the present 24 study, to gain a more complete picture of aquatic environment use, including social and 25 ecological factors in primates, factors affecting riverine habitat utilization of two 26 macaque species (Macaca nemestrina and M. fascicularis) were identified and 27 qualitative comparisons were made with sympatric proboscis monkeys (Nasalis 28 larvatus), which have different social and ecological characteristics. Temporal variation 29 in sighting frequency of macaques at the riverbanks was positively related to the fruit 30 availability of a dominant riparian plant species and negatively related to the river water 31 level which affects the extent of predation pressure. Riverine utilization of macaques 32 was greatly influenced by distribution and abundance of food (especially fruit) 33 resources, possibly in association with predation pressure. Additionally, qualitative 34 ecological comparisons with sympatric proboscis monkeys suggest that the drivers of 35 riverine utilization depend on the feeding niches of the species, and different anti-36 predator strategies resulting from their differing social structures. 37 38 Keywords 39Anti-predator strategy, aquatic environment, foraging strategy, sleeping site, southern 40 pig-tailed macaque 43 Animals often live in forests characterized as mosaic habitats comprising both terrestrial 44 and aquatic (e.g., riverine or coastal) environments while varying their associations with 45 the aquatic environment. In addition to direct relationships such as obtaining food from 46 the aquatic environment, e.g., bears eating salmon 1 ; racoons eating mussels 2 , the aquatic 47 environment has an indirect impact on terrestrial animals by bringing about 48 environmental heterogeneity. Differences in terrestrial flora, which are due to varying 49 water abundance and light intensity at the border between aquatic and terrestrial 50 environments, affect foraging behaviour of animals that use plants as a food source. In 51 addition, such boundaries restrict the distribution and movement patterns like a river-52 barrier 3 and thus are constraints on habitat utilization. Aquatic environments are, 53 therefore, undoubtedly important for terrestrial animals across many taxonomic groups 54 and geographic areas.
55Primates are a primary example of terrestrial animals that rely on the aquatic 56 environment. Kempf 4 presented a comprehensive review of primate aquatic behaviours 57 with the conclusions that the use of aquatic resources and the aquatic environment affect 58 various aspects of primate life, including feeding, traveling, predation avoidance, ...