Abstract
The extended phenotype of helical burrowing behavior in animals has evolved independently many times since the Cambrian explosion (~540 million years ago [MYA]). A number of hypotheses have been proposed to explain the evolution of helical burrowing in certain taxa, but no study has searched for a general explanation encompassing all taxa. We reviewed helical burrowing in both extant and extinct animals and from the trace fossil record and compiled 10 hypotheses for why animals construct helical burrows, including our own ideas. Of these, six are post-construction hypotheses-benefits to the creator or offspring, realized after burrow construction-and four are construction hypotheses reflecting direct benefits to the creator during construction. We examine the fit of these hypotheses to a total of 21 extant taxa and ichnotaxa representing 59-184 possible species. Only two hypotheses, antipredator and biomechanical advantage, cannot be rejected for any species (possible in 100% of taxa), but six of the hypotheses cannot be rejected for most species (possible in 86%-100% of taxa): microclimate buffer, reduced falling sediment (soil), anticrowding, and vertical patch. Four of these six are construction hypotheses, raising the possibility that helical burrowing may have evolved without providing post-construction benefits. Our analysis shows that increased drainage, deposit feeding, microbial farming, and offspring escape cannot explain helical burrowing behavior in the majority of taxa (5%-48%). Overall, the evidence does not support a general explanation for the evolution of helical burrowing in animals. The function and evolution of the helix as an extended phenotype seems to provide different advantages for different taxa in different environments under different physicochemical controls (some traces/tracemakers are discussed in more detail due to their association with body fossils and well-constrained physicochemical parameters). Although direct tests of many of the hypotheses would be difficult, we nevertheless offer ways to test some of the hypotheses for selected taxa.