Allometry between leg and body length of insects: lack of support for the size-grain hypothesis

The size-grain hypothesis (Kaspari & Weiser, 1999) states that (1) as organisms decrease in size, they perceive their environment as being more rugose; (2) long legs allow organisms to step over obstacles but hinder them from entering small gaps; and (3) as the size of an organism decreases, the benefits of long legs begin to be outweighed by the costs of construction. Natural selection should therefore favour proportionally longer legs in larger organisms, thereby leading to a positive allometry between leg and body length (scaling exponent b > 1). Here we compare the scaling exponent of leg-to-body length relationships among insects that walk, walk and fly, and predominantly fly. We measured the lengths of the hind tibia, hind femur, and body length of each species. The taxa varied considerably in the scaling exponent b. In seven out of ten groups (Formicidae, Isoptera, Carabidae, Pentatomidae, Apidae, Lepidoptera, Odonata adult), b was significantly greater than one. However, there was no gradual decrease in b from walking to walking/flying to flying insects. The results of the present study provide no support for the size-grain hypothesis. We propose that leg length is not only affected by the rugosity of the environment, but also by (1) functional adaptations, (2) phylogeny, (3) lifestyle, (4) the type of insect development (hemimetabolism or holometabolism), and (5) constraints of gas exchange.