The 'Cambrian Explosion' describes the rapid increase in animal diversity and abundance, as manifest in the fossil record, between ~ 540 and 520 million years ago (Ma). This event, however, is nested within a far more ancient record of macrofossils extending at least into the late Ediacaran, ~571 Ma. The evolutionary events documented during the Ediacaran-Cambrian interval coincide with geochemical evidence for the modernisation of Earth's biogeochemical cycles. Holistic integration of fossil and geochemical records leads us to challenge the notion that the Ediacaran and Cambrian worlds were markedly distinct, and places biotic and environmental change within a longer-term narrative. We propose that the evolution of metazoans may have been facilitated by a series of dynamic and global changes in redox conditions and nutrient supply, which, together with potential biotic feedbacks, enabled turnover events that sustained phases of radiation. In this synthesis, we argue that early metazoan diversification should be recast as a series of successive, transitional radiations that extended from the late Ediacaran and continued through the early Palaeozoic. We conclude that while the Cambrian Explosion represents a radiation of crown-group bilaterians, it was simply one phase amongst several older, and younger, metazoan radiations.

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Ediacaran macrofossils from the Avalon Terrane (primarily eastern Newfoundland and the central U.K.) record some of the earliest large and complex multicellular organisms on Earth.Perhaps the greatest unknown regarding these fossils is their relevance to the early evolutionary history of the Kingdom Animalia. In recent years, new data and discoveries have revealed insights into Ediacaran paleobiology, taxonomic relationships, paleoecology and taphonomy, significantly refining our understanding of Avalonian ecosystems. Here, we summarise recent observational and quantitative studies, and their bearing on the current understanding of Avalonian benthic marine ecosystems. A review of existing knowledge of the biological composition of Avalonian marine assemblages demonstrates that they recorddensely-populated ecosystems inhabited by a diverse range of organisms, likely representing multiple biological Kingdoms. Appreciation of this diversity, and of the complexities it introduces to paleoecological studies, is vital when considering the relationship between macroevolution and contemporaneous climatic, tectonic and geochemical events. We then summarise current understanding of Avalonian paleoecology. Studies into locomotion, reproduction, feeding strategies, and community structure and succession reveal that these ecosystems were considerably different to Phanerozoic settings. Furthermore, we suggest that Avalonian ecosystems witnessed the appearance of novel nutrient sources, offering new opportunities and niches for benthic organisms. The suggestion that the numerically dominant rangeomorphs were osmotrophic is reviewed and appraised in light of geochemical, morphological, and biological information. Finally, the use of modern ecological metrics in the study of Ediacaran fossil assemblages is assessed. Concerns regarding the interpretation of paleoecological data are outlined in light of current taphonomic and sedimentological understanding, and these cast doubt on previous suggestions that the Avalonian assemblages were largely composed of metazoans. Nevertheless, we emphasise that if treated with necessary caution, paleoecological data can play a significant role in assisting efforts to determine the biological affinities of late Ediacaran macroscopic organisms.