One way for animals to decrease energy expenditures is to minimize the cost
of movement. For animals dwelling on slopes, gravity can impart a large en
ergetic cost to movement. For this reason, animals traveling aboveground al
ter their movement patterns in response to the steepness of terrain (specif
ically hillslope angle) so as to minimize their energetic costs. Subterrane
an animals should also benefit from choosing optimum movement paths in rela
tion to hillslopes but concurrently must factor the cost of excavation into
their movement decisions. In cases where the excavation costs are much hig
her than the costs of working against gravity, excavation costs may overrid
e the consideration of gravitational costs and movement of subterranean ani
mals may be independent of hillslope angle. To determine the response of a
subterranean animal to hillslope angle, we excavated tunnels in the burrow
systems of 19 pocket gophers in southern California that occupied hillslope
s ranging from 2 to 30 degrees. At each excavation we measured several char
acteristics of burrow geometry and used these data in a model of pocket gop
her energetics to calculate the cost of tunnel construction at the various
hillslope angles. We found that the cost of tunnel construction was indepen
dent of hillslope angle, and that the costs of shearing soil and pushing so
il horizontally through the tunnels were 3 orders of magnitude greater than
the costs of lifting the soil against the force of gravity. Accordingly, p
ocket gopher foraging tunnels were oriented independently of the hillslope.
The decoupling of the movement patterns of subterranean animals from the e
ffects of gravity is a distinctive feature of the subterranean habit compar
ed to the movement of above-ground animals. Because of the important effect
s of tunnel construction on soil processes, this unique biological feature
of subterranean animals has implications for basic physical processes, such
as soil erosion. We found that the rate of soil flux generated by pocket g
opher activity was invariant to hillslope. This relationship is in contrast
to the most common model of soil movement generated by purely physical pro
cesses.