Shells as 'extended architecture': to escape isolation, social hermit crabs choose shells with the right external architecture

Animals' cognitive abilities can be tested by allowing them to choose between alternatives, with only one alternative offering the correct solution to a novel problem. Hermit crabs are evolutionarily specialized to navigate while carrying a shell, with alternative shells representing different forms of 'extended architecture', which effectively change the extent of physical space an individual occupies in the world. It is unknown whether individuals can choose such architecture to solve novel navigational problems. Here, we designed an experiment in which social hermit crabs (Coenobita compressus) had to choose between two alternative shells to solve a novel problem: escaping solitary confinement. Using X-ray microtomography and 3D-printing, we copied preferred shell types and then made artificial alterations to their inner or outer shell architecture, designing only some shells to have the correct architectural fit for escaping the opening of an isolated crab's enclosure. In our 'escape artist' experimental design, crabs had to choose an otherwise less preferred shell, since only this shell had the right external architecture to allow the crab to free itself from isolation. Across multiple experiments, crabs were willing to forgo preferred shells and choose less preferred shells that enabled them to escape, suggesting these animals can solve novel navigational problems with extended architecture. Yet, it remains unclear if individuals solved this problem through trial-and-error or were aware of the deeper connection between escape and exterior shell architecture. Our experiments offer a foundation for further explorations of physical, social, and spatial cognition within the context of extended architecture.