It has long been assumed that the extant bilaterian phyla generally have their origin in the Cambrian explosion, when they appear in an essentially modern form. Both these assumptions are questionable. A strict application of stem- and crown-group concepts to phyla shows that although the branching points of many clades may have occurred in the Early Cambrian or before, the appearance of the modern body plans was in most cases later: very few bilaterian phyla sensu stricto have demonstrable representatives in the earliest Cambrian. Given that the early branching points of major clades is an inevitable result of the geometry of clade diversification, the alleged phenomenon of phyla appearing early and remaining morphologically static is seen not to require particular explanation. Confusion in the definition of a phylum has thus led to attempts to explain (especially from a developmental perspective) a feature that is partly inevitable, partly illusory. We critically discuss models for Proterozoic diversification based on small body size, limited developmental capacity and poor preservation and cryptic habits, and show that the prospect of lineage diversification occurring early in the Proterozoic can be seen to be unlikely on grounds of both parsimony and functional morphology. Indeed, the combination of the body and trace fossil record demonstrates a progressive diversification through the end of the Proterozoic well into the Cambrian and beyond, a picture consistent with body plans being assembled during this time. Body-plan characters are likely to have been acquired monophyletically in the history of the bilaterians, and a model explaining the diversity in just one of them, the coelom, is presented. This analysis points to the requirement for a careful application of systematic methodology before explanations are sought for alleged patterns of constraint and flexibility.

The increase in trace fossil diversity across the Neoproterozoic-Cambrian boundary often is presented in terms of tabulations of ichnogenera. However, a clearer picture of the increase in diversity and complexity can be reached by combining trace fossils into broad groups defined both on morphology and interpretation. This also focuses attention on looking for similarites between Neoproterozoic and Cambrian trace fossils. Siliciclastic sediments of the Neoproterozoic preserve elongate tubular organisms and structures of probable algal origin, many of which are very similar to trace fossils. Such enigmatic structures include Palaeopascichnus and Yelovichnus, previously thought to be trace fossils in the form of tight meanders.A preliminary two or tripartite terminal Neoproterozoic trace fossil zonation can be be recognized. Possibly the earliest trace fossils are short unbranched forms, probably younger than about 560 Ma. Typical Neoproterozoic trace fossils are unbranched and essentially horizontal forms found associated with diverse assemblages of Ediacaran organisms. In sections younger than about 550 Ma a modest increase in trace fossil diversity occurs, including the appearance of rare three-dimensional burrow systems (treptichnids), and traces with a three-lobed lower surfaces.

The earliest evolution of the animals remains a taxing biological problem, as all extant clades are highly derived and the fossil record is not usually considered to be helpful. The rise of the bilaterian animals recorded in the fossil record, commonly known as the 'Cambrian explosion', is one of the most significant moments in evolutionary history, and was an event that transformed first marine and then terrestrial environments. We review the phylogeny of early animals and other opisthokonts, and the affinities of the earliest large complex fossils, the so-called 'Ediacaran' taxa. We conclude, based on a variety of lines of evidence, that their affinities most likely lie in various stem groups to large metazoan groupings; a new grouping, the Apoikozoa, is erected to encompass Metazoa and Choanoflagellata. The earliest reasonable fossil evidence for total-group bilaterians comes from undisputed complex trace fossils that are younger than about 560 Ma, and these diversify greatly as the Ediacaran-Cambrian boundary is crossed a few million years later. It is generally considered that as the bilaterians diversified after this time, their burrowing behaviour destroyed the cyanobacterial mat-dominated substrates that the enigmatic Ediacaran taxa were associated with, the so-called 'Cambrian substrate revolution', leading to the loss of almost all Ediacara-aspect diversity in the Cambrian. Why, though, did the energetically expensive and functionally complex burrowing mode of life so typical of later bilaterians arise? Here we propose a much more positive relationship between late-Ediacaran ecologies and the rise of the bilaterians, with the largely static Ediacaran taxa acting as points of concentration of organic matter both above and below the sediment surface. The breaking of the uniformity of organic carbon availability would have signalled a decisive shift away from the essentially static and monotonous earlier Ediacaran world into the dynamic and burrowing world of the Cambrian. The Ediacaran biota thus played an enabling role in bilaterian evolution similar to that proposed for the Savannah environment for human evolution and bipedality. Rather than being obliterated by the rise of the bilaterians, the subtle remnants of Ediacara-style taxa within the Cambrian suggest that they remained significant components of Phanerozoic communities, even though at some point their enabling role for bilaterian evolution was presumably taken over by bilaterians or other metazoans. Bilaterian evolution was thus an essentially benthic event that only later impacted the planktonic environment and the style of organic export to the sea floor.

The trace fossil record is important in determining the timing of the appearance of bilaterian animals. A conservative estimate puts this time at Ϸ555 million years ago. The preservational potential of traces made close to the sediment-water interface is crucial to detecting early benthic activity. Our studies on earliest Cambrian sediments suggest that shallow tiers were preserved to a greater extent than typical for most of the Phanerozoic, which can be attributed both directly and indirectly to the low levels of sediment mixing. The low levels of sediment mixing meant that thin event beds were preserved. The shallow depth of sediment mixing also meant that muddy sediments were firm close to the sediment-water interface, increasing the likelihood of recording shallow-tier trace fossils in muddy sediments. Overall, trace fossils can provide a sound record of the onset of bilaterian benthic activity.T he appearance and subsequent diversification of bilaterian animals is a topic of current controversy (refs. 1-7; Fig. 1). Three principal sources of evidence exist: body fossils, trace fossils (trails, tracks, and burrows of animal activity recorded in the sedimentary record), and divergence times calculated by means of a molecular ''clock.'' The body fossil record indicates a geologically rapid diversification of bilaterian animals not much earlier than the Precambrian-Cambrian boundary, the so-called Cambrian explosion. The largely terminal Proterozoic Ediacaran biota remain problematic to questions of bilaterian origins (7-10). Molecular estimates (1) suggest that diversification of bilaterian groups may have commenced more than 1,000 million years ago. However, the molecular clock studies provide a considerable spread in the results, with some coming close (11) or even very close § to the pattern seen from body fossils (Fig. 1). An increase in diversity and complexity of trace fossils across the Neoproterozoic-Cambrian boundary has long been recognized (12)(13)(14). The oldest widely accepted trace fossils are no older than 555 million years old (5, 15). Broadly speaking, terminal Proterozoic trace fossils are simple, unbranched, less than a few millimeters in diameter, and were made close to the sedimentwater interface. In the Cambrian, morphological diversity increased, size range expanded, and depth of sediment penetration increased modestly (16).Although not accepted universally, bilaterians may be primitively benthic, and many of their morphological features could only have evolved in a moderately large animal with a benthic lifestyle (5,6,15). Such animals would have had the ability to burrow, and it requires special pleading to argue that they would not have produced trace fossils (5, 15). The appearance of macroscopic bilaterians, thus, is arguably recorded by terminal Proterozoic trace fossils. Conditions for the preservation of relatively surficial infaunal activity should have been particularly favorable because of the shallow depth of bioturbation at this time (5,17). However, precise mechanisms of ...