Leaf and flower consumption modulate the drinking behavior in a folivorous-frugivorous arboreal mammal

Abstract

Water is vital for the survival of any species because of its key role in most physiological processes. However, little is known about the non-food-related water sources exploited by arboreal mammals, the seasonality of their drinking behavior and its potential drivers (including diet composition, temperature, and rainfall). We investigated this subject in 14 wild groups of brown howler monkeys (Alouatta guariba clamitans) inhabiting small, medium, and large Atlantic Forest fragments in southern Brazil. We found a wide variation in the mean rate of drinking among groups (range=0-16 records/day). Streams (44% of 1,258 records) and treeholes (26%) were the major types of water sources, followed by bromeliads in the canopy (16%), pools (11%), and rivers (3%). The type of source influenced whether howlers used a hand to access the water or not. Drinking tended to be evenly distributed throughout the year, except for a slightly lower number of records in the spring than in the other seasons, but it was unevenly distributed during the day. It increased in the afternoon in all groups, particularly during temperature peaks around 15:00 and 17:00. We found via generalized linear mixed modelling that the daily frequency of drinking was mainly influenced by flower (negatively) and leaf (positively) consumption, whereas fruit consumption, fragment size, rainfall, and mean ambient temperature played negligible roles. The influence of leaf consumption is compatible with the metabolite detoxification hypothesis, which states that the processing of this fibrous food requires the ingestion of larger volumes of water to help in the detoxification/excretion of its metabolites. In sum, we found that irrespective of habitat size and climatic conditions, brown howlers seem to seek a positive water balance by complementing preformed and metabolic water with drinking water, even when it is associated with a high predation risk in terrestrial sources.