The rise and early evolution of animals: where do we stand from a trace-fossil perspective?

The trace-fossil record provides a wealth of information to track the rise and early evolution of animals. It comprises the activity of both hard- and soft-bodied organisms, is continuous through the Ediacaran (635-539 Ma)- Cambrian (539-485 Ma) transition, yields insights into animal behaviour and their role as ecosystem engineers, and allows for a more refined characterization of palaeoenvironmental context. In order to unravel macroevolutionary signals from the trace-fossil record, a variety of approaches is available, including not only estimation of degree of bioturbation, but also analysis of ichnodiversity and ichnodisparity trajectories, and evaluation of the occupation of infaunal ecospace and styles of ecosystem engineering. Analysis of the trace-fossil record demonstrates the presence of motile benthic bilaterians in the Ediacaran, mostly feeding from biofilms. Although Ediacaran trace fossils are simple and emplaced at or immediately below the sediment surface, an increase in ichnofossil complexity, predation pressure, sediment disturbance and penetration depth is apparent during the terminal Ediacaran. Regardless of this increase, a dramatic rise in trace fossil diversity and disparity took place during the earliest Cambrian, underscoring that the novelty of the Fortunian (539-529 Ma) cannot be underestimated. The Fortunian still shows the persistence of an Ediacaran-style matground ecology, but is fundamentally characterized by the appearance of new trace-fossil architectural plans reflecting novel ways of interacting with the substrate. The appearance of Phanerozoic-style benthic ecosystems attests to an increased length and connectivity of the food web and improved efficiency in organic carbon transfer and nutrient recycling. A profound reorganization of the infaunal ecospace is recorded in both high-energy sand-dominated nearshore areas and low-energy mud-dominated offshore environments, during the early Cambrian, starting approximately during Cambrian Age 2 (529-521 Ma), but continuing during the rest of the early Cambrian. A model comprising four evolutionary phases is proposed to synthetize information from the Ediacaran-Cambrian trace-fossil record. The use of a rich ichnological toolbox; critical, systematic and comprehensive evaluation of the Ediacaran-Cambrian trace-fossil record; and high-resolution integration of the ichnological dataset and sedimentological information show that the advent of biogenic mixing was an important factor in fully marine environments at the dawn of the Phanerozoic.