Why Animals Construct Helical Burrows: Construction vs.
Post-Construction Benefits
Abstract
The extended phenotype of helical burrowing behavior in animals has
evolved independently many times since first appearing after the
Cambrian explosion (~540 million years ago). 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
from the literature 10 possible hypotheses for why animals construct
helical burrows, including our own ideas. Of these, six were
post-construction hypotheses—-benefits to the creator or offspring,
realized after burrow construction—-and four were construction
hypotheses reflecting direct benefits to the creator during
construction. We examined the fit of these hypotheses to a total of 21
extant taxa and ichnotaxa representing 59–184 spp. Only two
hypotheses—-antipredator, biomechanical advantage—-could not be
rejected for any species (possible in 100% of spp.), but six of the
hypotheses could not be rejected for most species (possible in 86–100
% of spp.): microclimate buffer, reduced falling sediment (soil),
anticrowding, vertical patch, and the above two hypotheses. Four of
these six were construction hypotheses, raising the possibility that
helical burrowing might have evolved without providing post-construction
benefits. Our analysis showed that increased drainage, deposit feeding,
microbial farming, and offspring escape could not 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 would seem, at least in some cases, to provide
different advantages for different taxa. Although direct tests of many
of the hypotheses would be difficult, we nevertheless offer ways to test
some of the hypotheses for selected taxa.