“…The Expensive-Tissue Hypothesis (ETH) states that the high energetic expenditure of larger brains requires a matching decrease in other energetic-consuming organisms (e.g., guts) [ 15 , 36 , 37 , 38 , 39 , 41 , 42 ]. However, studies have refuted the ETH [ 43 , 44 , 45 , 46 , 47 ] and proposed the trade-off hypothesis, which suggests that instead of tissue investment, a series of trade-offs with other energetically costly traits such as body maintenance, locomotion, development, and reproductive investment (e.g., testes mass) can compensate for the cost of increased brain size [ 39 , 43 , 44 , 45 , 46 , 47 , 48 , 49 , 50 , 51 ]. Nonetheless, the gut plays an important role in brain size evolution and can be viewed as one of these energy trade-offs.…”