Bioturbators as ecosystem engineers:assessing current models

Bioturbating organisms can dramatically alter the physical, chemical, and hydrological properties of the sediment and promote or hinder microbial growth. They are a classic example of “ecosystem engineers” as they alter the availability of resources to other species. Multiple evolutionary hypotheses evoke bioturbation as a possible driver for historical ecological change. To test these hypotheses, researchers need reliable and reproducible methods for estimating the impact of bioturbation in ancient environments. Early efforts to record and compare this impact through geologic time focused on the degree of bioturbation (e.g., bioturbation indices), the depth of bioturbation (e.g., bioturbation depth), or the structure of the infaunal community (e.g., tiering, ecospace utilization). Models which combine several parameters (e.g., functional groups, tier, motility, sediment interaction style) have been proposed and applied across the geological timescale in recent years. Here, we review all models that characterize the impact of bioturbators on the sedimentary environment (i.e., ‘ecosystem engineering'), in both modern and fossil sediments, and propose several questions. What are the assumptions of each approach? Are the current models appropriate for the metrics they wish to measure? Are they robust and reproducible? Our review highlights the nature of the sedimentary environment as an important parameter when characterizing ecosystem engineering intensity and outlines considerations for a best-practice model to measure the impact of bioturbation in geological datasets.