Diverse bilaterian clades emerged apparently within a few million years during the early Cambrian, and various environmental, developmental, and ecological causes have been proposed to explain this abrupt appearance. A compilation of the patterns of fossil and molecular diversification, comparative developmental data, and information on ecological feeding strategies indicate that the major animal clades diverged many tens of millions of years before their first appearance in the fossil record, demonstrating a macroevolutionary lag between the establishment of their developmental toolkits during the Cryogenian [(850 to 635 million years ago (Ma)], and the later ecological success of metazoans during the Ediacaran (635 to 541 Ma) and Cambrian (541 to 488 Ma) periods. We argue that this diversification involved new forms of developmental regulation, as well as innovations in networks of ecological interaction within the context of permissive environmental circumstances.

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Ediacaran fronds at Spaniard's Bay on the Avalon Peninsula of Newfoundland exhibit exquisite, threedimensional preservation with morphological features less than 0.05 mm in width visible on the best preserved specimens. Most of the nearly 100 specimens are juvenile rangeomorphs, an extinct Ediacaran clade that numerically dominated the early evolution of complex multicellular life. Spaniard's Bay rangeomorphs are characterized by cmscale architectural elements exhibiting self-similar branching over several fractal scales that were used as modules in construction of larger structures. Four taxa of rangeomorph fronds are present -Avalofractus abaculus n. gen. et sp., Beothukis mistakensis Brasier and Antcliffe, Trepassia wardae (Narbonne and Gehling), and Charnia cf. C. masoni Ford. All of these taxa exhibit an alternate array of primary rangeomorph branches that pass off a central stalk or furrow that marks the midline of the petalodium. Avalofractus is remarkably self similar over at least four fractal scales, with each scale represented by double-sided rangeomorph elements that were constrained only at their attachment point with the higher-order branch and thus were free to rotate and pivot relative to other branches. Beothukis is similar in organization, but its primary branches show only one side of a typical rangeomorph element, probably due to longitudinal branch folding, and the position of the individual branches was moderately constrained. Trepassia shows only single-sided branches with both primary and secondary branches emanating from a central stalk or furrow; primary branches were capable of minor pivoting as reflected in bundles of secondary branches. Charnia shows only single-sided primary branches that branch from a zigzag central furrow and that were firmly constrained relative to each. This sequence provides a developmental linkage between Rangea-type and Charnia-type rangeomorphs. Avalonian assemblages show a wide array of rangeomorph constructions, but later Ediacaran assemblages contain a lower diversity of rangeomorphs represented mainly by well-constrained forms.

The latest Neoproterozoic extinction of the Ediacara biota has been variously attributed to catastrophic removal by perturbations to global geochemical cycles, 'biotic replacement' by Cambrian-type ecosystem engineers, and a taphonomic artefact. We perform the first critical test of the 'biotic replacement' hypothesis using combined palaeoecological and geochemical data collected from the youngest Ediacaran strata in southern Namibia. We find that, even after accounting for a variety of potential sampling and taphonomic biases, the Ediacaran assemblage preserved at Farm Swartpunt has significantly lower genus richness than older assemblages. Geochemical and sedimentological analyses confirm an oxygenated and non-restricted palaeoenvironment for fossilbearing sediments, thus suggesting that oxygen stress and/or hypersalinity are unlikely to be responsible for the low diversity of communities preserved at Swartpunt. These combined analyses suggest depauperate communities characterized the latest Ediacaran and provide the first quantitative support for the biotic replacement model for the end of the Ediacara biota. Although more sites (especially those recording different palaeoenvironments) are undoubtedly needed, this study provides the first quantitative palaeoecological evidence to suggest that evolutionary innovation, ecosystem engineering and biological interactions may have ultimately caused the first mass extinction of complex life.

The Ediacara biota include macroscopic, morphologically complex soft-bodied organisms that appear globally in the late Ediacaran Period (575-542 Ma). The physiology, feeding strategies, and functional morphology of the modular Ediacara organisms (rangeomorphs and erniettomorphs) remain debated but are critical for understanding their ecology and phylogeny. Their modular construction triggered numerous hypotheses concerning their likely feeding strategies, ranging from micro-to-macrophagus feeding to photoautotrophy to osmotrophy. Macrophagus feeding in rangeomorphs and erniettomorphs is inconsistent with their lack of oral openings, and photoautotrophy in rangeomorphs is contradicted by their habitats below the photic zone. Here, we combine theoretical models and empirical data to evaluate the feasibility of osmotrophy, which requires high surface area to volume (SA/V) ratios, as a primary feeding strategy of rangeomorphs and erniettomorphs. Although exclusively osmotrophic feeding in modern ecosystems is restricted to microscopic bacteria, this study suggests that (i) fractal branching of rangeomorph modules resulted in SA/V ratios comparable to those observed in modern osmotrophic bacteria, and (ii) rangeomorphs, and particularly erniettomorphs, could have achieved osmotrophic SA/V ratios similar to bacteria, provided their bodies included metabolically inert material. Thus, specific morphological adaptations observed in rangeomorphs and erniettomorphs may have represented strategies for overcoming physiological constraints that typically make osmotrophy prohibitive for macroscopic life forms. These results support the viability of osmotrophic feeding in rangeomorphs and erniettomorphs, help explain their taphonomic peculiarities, and point to the possible importance of earliest macroorganisms for cycling dissolved organic carbon that may have been present in abundance during Ediacaran times.erniettomorphs ͉ rangeomorphs ͉ Fractofusus ͉ Pteridinium

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