7. Ancient Fossil Record and Early Evolution (ca. 3.8 to 0.5 Ga)

Once life appeared, it evolved and diversified. From primitive living entities, an evolutionary path of unknown duration, likely paralleled by the extinction of unsuccessful attempts, led to a last common ancestor that was endowed with the basic properties of all cells. From it, cellular organisms derived in a relative order, chronology and manner that are not yet completely settled. Early life evolution was accompanied by metabolic diversification, i.e. by the development of carbon and energy metabolic pathways that differed from the first, not yet clearly identified, metabolic strategies used. When did the different evolutionary transitions take place? The answer is difficult, since hot controversies have been raised in recent years concerning the reliability of the oldest life traces, regardless of their morphological, isotopic or organic nature, and there are also many competing hypotheses for the evolution of the eukaryotic cell. As a result, there is a need to delimit hypotheses from solid facts and to apply a critical analysis of contrasting data. Hopefully, methodological improvement and the increase of data, including fossil signatures and genomic information, will help reconstructing a better picture of life evolution in early times as well as to, perhaps, date some of the major evolutionary transitions. There are already some certitudes. Modern eukaryotes evolved after bacteria, since their mitochondria derived from ancient bacterial endosymbionts. Once prokaryotes and unicellular eukaryotes had colonized terrestrial ecosystems for millions of years, the first pluricellular animals appeared and radiated, thus inaugurating the Cambrian. The following sections constitute a collection of independent articles providing a general overview of these aspects.