Mechanics and energetics of excavation by burrowing wolf spiders, Geolycosa spp

Burrowing wolf spiders, Geolycosa sp. (Araneae:Lycosidae), excavate vertical burrows and inhabit them throughout their lives or, in the case of males, until they mature and wander in search of mates. Three species: G. fatifera Kurata, G. missouriensis Banks, and G. rogersi Wallace were studied to understand how and at what expense the burrowing is accomplished. Normal and high-speed videography coupled with scanning electron microscopy revealed (a) that the convex surfaces of the two fangs, together, constitute the digging tool, (b) that boluses of soil are transported to the burrow entrance on the anterior surfaces of the chelicerae held there by the pedipalps, and (c) that each bolus is either incorporated into the growing turret or flung away, propelled by the forelegs. To elucidate the energetics of burrow construction, burrow volumes were calculated and then the costs associated with dislodging, elevating, and throwing the known volumes of soil were measured. A typical Geolycosa burrow, at a volume of 23.6 ± 2.0 ml and a depth of 13.2 ± 0.7 cm, required the removal of 918 boluses each weighing about 34 mg. The aggregate dislodging cost was close to 1.9 Joules in sand/sandy loam and 5.6 J in clayey subsoil, the work against gravity necessary to raise all of the boluses to the surface was about 0.13 J, and the aggregate cost of flinging the boluses was close to 0.014 J. Assuming that the ratio of external work to metabolic cost of external work is between 0.20 and 0.25 in spiders, the real cost of burrow construction would be between 8 J and 29 J, depending primarily on soil type. This is a small but not negligible cost when placed in the context of reproductive effort: a single Geolycosa egg, dozens to hundreds of which are produced in a clutch, contains about 10 J.