Intermittent and temporally variable bioturbation by some terrestrial invertebrates: implications for ichnology.

Bedding planes and vertical sections of many sedimentary rock formations reveal bioturbation structures, including burrows, produced by diverse animal taxa at different rates and durations. These variables are not directly measurable in the fossil record, but neoichnological observations and experiments provide informative analogues. Comparable to marine invertebrates from many phyla, a captive beetle larva burrowing over 2 weeks showed high rates of sediment disturbance within the first 100 h but slower rates afterwards. Tunnelling by earthworms and adult dung beetles is also inconstant-displacement of lithic material alternates with organic matter displacement, often driven by food availability with more locomotion when hungry. High rates of bioturbation, as with locomotion generally, result from internal and external drives, slowing down or stopping when needs are filled. Like other processes affecting sediment deposition and erosion, rates can drastically differ based on measured timescale, with short bursts of activity followed by hiatuses, concentrated in various seasons and ontogenetic stages for particular species. Assumptions of constant velocities within movement paths, left as traces afterward, may not apply in many cases. Arguments about energetic efficiency or optimal foraging based on ichnofossils have often overlooked these and related issues. Single bioturbation rates from short-term experiments in captivity may not be comparable to rates measured at an ecosystem level over a year or generalized across multiple time scales where conditions differ even for the same species. Neoichnological work, with an understanding of lifetime variabilities in bioturbation and their drivers, helps connect ichnology with behavioural biology and movement ecology.