Author Summary Water striders are derived semi-aquatic bugs that possess a remarkable diversity of leg lengths and shapes among species and between sexes, and the selective forces shaping this diversity are well studied. The transition to living on the water surface was accompanied by dramatic changes in the size and function of their legs. The mid-legs are disproportionately long and function as oars, whereas the hind-legs are shorter and function as rudders. We present evidence demonstrating that changes in the pattern of expression and function of the Hox gene Ultrabithorax are responsible for establishing the relative size differences between mid- and hind-legs in the water strider Gerris buenoi. These changes in Ubx expression and function may have been a key event in the evolution of the distinct appendage ground plan in water striders.