Paleoecology of the marine endobenthos

Endobenthic animals, which reside within the sea bottom, include stationary suspension feeders, mobile deposit feeders and both stationary and mobile carnivores. Their activities, especially with regard to dwelling, feeding, walking/crawling and resting/nesting, are recorded as trace fossils. Abundance, diversity and density of some kinds of trace fossils allow interpretation of the population strategies of the trace-makers in terms of opportunistic (r-selected) and equilibrium (K-selected) strategies. Opportunistic ichnotaxa tend to be faciesbreaking traces, which are highly localized in low-diversity, high-density trace fossil associations in rocks representing environmental extremes (e.g., variable salinities, harsh temperatures, low oxygen levels or shifting substrates). Equilibrium ichnotaxa usually are restricted to particular sedimentary facies and are characteristic of high-diversity, low-dominance trace fossil associations in sediments reflecting stable, predictable environmental conditions. The most important environmental factors influencing the composition of trace fossil assemblages in marine settings are bathymetry, substrate, oxygen and hydrodynamic energy. The four factors are closely interrelated, because as water depth increases, there is a general decrease in sediment grain size and hydrodynamic energy of the depositional environment. As depth below the water—sediment interface increases, the firmness of the sediment (due to compaction and dewatering) increases and the oxygen content of interstitial waters drops drastically. Marine ichnofacies are largely substrate-controlled. Soupgrounds are water-saturated, incompetent substrates typified by highly compressed and usually unidentifiable burrows. Softgrounds commonly contain numerous distinctive burrows and are zoned bathymetrically by the Skolithos, Cruziana, Zoophycos and Nereites Ichnofacies. Firmgrounds are characterized by stiff, compacted sediments, in which traces of the Glossifungites Ichnofacies are excavated. Hardgrounds are cemented substrates, in which bioerosion traces of the Trypanites Ichnofacies are bored. Woodgrounds are woody materials that have been exposed to the sea and bored by bivalves, which produce characteristic traces of the Teredolites Ichnofacies. Tiering of endobenthic communities is common and is related to substrate preference of the burrowers and oxygen stratification of interstitial waters.