Energetically costly weaponry in the large morph of male stag beetles

Males of many species develop disproportionately large weapons, and frequently the weapon size increases more quickly with body size in small than in large males—a trend often considered to result from the depletion of developmental resources limiting male weapon exaggeration. Based on the cost-minimization hypothesis, a shallower weapon-body size allometric slope in large than in small males could be selected for if the energetic maintenance costs of having oversized weapons increase faster in large than in small males. Whether males of different size groups that differ in the weapon-body size allometric relationship also differ in the energetic costs of maintaining their weapons has not yet been investigated. Consequently, we studied this using the stag beetle (Cyclommatus mniszechi), in which males are larger in size and have larger mandibles than do females and are divided into two morphs: majors are larger and have longer mandibles than minors. In both male morphs, mandible size increases disproportionately with their body size, but it increases more quickly in minors than in majors. We measured the standard metabolic rate (SMR) of the three types of these beetles (the two male morphs and females) to investigate whether the energetic maintenance cost of carrying weapons differs among them. Body weight explained 80% of the variation in the SMR in the three types of beetles, and SMR increased more quickly with body weight in major males than in minor males or females. Weapon and body size also jointly explained approximately 80% of the variation in the SMR. Whereas SMR increased with body size in all three types of beetles (with a higher rate of increase in major males than in minor males or females), only the major males' SMR increased with weapon size. Overall, being heavier and larger and carrying oversized weapons are energetically costly in major males, something which could constrain the exaggeration of their weapons.