An updated classification of animal behaviour preserved in substrates

During the last few decades, many new ethological categories for trace fossils have been proposed in addition to the original five given by Seilacher. In this article, we review these new groups and present a version of the scheme of fossil animal behaviour originally published by Bromley updated with regard to modern ethological concepts, especially those of Tinbergen. Because some behaviours are more common in certain environments than others, they are useful in palaeoecological reconstructions, forming the original basis of the ichnofacies concept. To simplify, we summarise some ethological categories as previously done by others. However, the tracemaker’s behaviour in some cases is so distinctive that subcategories should be employed, especially in ecological interpretations of certain environments where a special behaviour may be dominant.