Morphological integration and pleiotropy in the adaptive body shape of the snail-feeding carabid beetle Damaster blaptoides.

The snail-feeding carabid beetle Damaster blaptoides exhibits diverse head and thorax morphologies, and these morphotypes are linked with two alternative feeding behaviours. Stout-shaped beetles feed on snails by crushing the shells, whereas slender-shaped beetles consume snails by inserting their heads into the shells. A trade-off exists between these feeding strategies. Because intermediate-shaped beetles are less proficient in these two behaviours, stout-slender morphological divergence occurs between related species feeding on land snails. To examine the genetic basis of these morphotypes, we conducted morphological analyses and quantitative trait locus (QTL) mapping using backcross offspring between the stout and slender subspecies. The morphological analyses showed that the width and length of the beetle body parts were correlated with each other; in particular, the head width (HW) and thorax length (TL) were strongly negatively correlated. QTL mapping showed that QTLs for HW and TL are located in close proximity to one another on the longest linkage group and that they have positive and negative additive genetic effects. Our results suggest that the adaptive phenotypic sets of a wide head and short thorax and a narrow head and long thorax are based on the closeness of these QTLs. Morphological integration between the head and thorax may play an important role in the adaptive divergence of these beetles.