Curious chiral cases of caddisfly larvae: handed behavior, asymmetric forms, evolutionary history

Studies of right-left asymmetries have yielded valuable insights into the mechanisms of both development and evolution. Larvae from several groups of caddisflies (Trichoptera) build portable asymmetrical cases within which they live. In nearly all species that build spiral-walled tubular cases, the direction of wall coiling is random (equal numbers of dextral and sinistral cases within species) whereas in all species that build helicospiral, snail-like cases the direction of coiling is exclusively dextral. Asymmetrical tubes result from handed behavior, and ∼20% of larvae removed from a spiral-walled, tubular case build a replacement case of opposite chirality. So handed behavior (and hence direction of tube-wall spiraling) is likely learned rather than determined genetically. Asymmetrical larval cases appear to have evolved at least seven times in the Trichoptera, five times as spiral-walled tubes and twice as snail-like helicospiral cases. Helicospiral cases may reduce vulnerability to predation by mimicking snail shells, whereas spiral arrangements of vegetation fragments in tube walls may be more robust mechanically than other arrangements, but experimental evidence is lacking. Within one family (Phryganeidae), one or perhaps two species exhibit an excess of sinistral-walled cases, suggesting that genes that bias handed behavior in a particular direction evolved after handed behaviors already existed (genetic assimilation).