Does brain size of Asiatic toads (<i>Bufo gargarizans</i>) trade-off with other energetically expensive organs along altitudinal gradients?

Brain size variation is often attributed to energetic trade-offs with other metabolically expensive tissues and organs, which is a prediction of the expensive brain hypothesis (EBH). Here we examine Asiatic toads (Bufo gargarizans) along altitudinal gradients and test size trade-offs between the brain and four visceral organs (heart, liver, alimentary tract, and kidney) with altitude. Body size and scaled mass index (a proxy for total energy intake) decline with altitude, implying stronger energetic constraints at high altitudes. Relative brain size decreases along altitudinal gradients, while visceral organs mostly increase in relative sizes. Using structural equation modeling, a significant negative relationship between brain size and a latent variable "budget," which represents the energy allocation to the four visceral organs, is detected among high-altitudinal toads. Heart appears to have the largest and most consistent response to changes in energy allocation. No such relationships are observed among toads at middle- and low-altitudes, where high energy intake may allow individuals to forego energetic trade-offs. When applying EBH to poikilotherms, a great emphasis should be placed on total energy intake in addition to energy allocation. Future research on EBH will benefit from more intra-specific comparisons and the evaluation of fitness consequences beyond energy limitation.