TL;DR: To place the Burgess Shale polychaetes in some geological perspective other Cambrian worms, including a polychaete from the Spence Shale of Utah, are briefly redescribed and the late Precambrian (Ediacarian) worms Dickinsonia, Spriggina and Marywadea are assessed.
Abstract: The Burgess Shale (Middle Cambrian) polychaetes Canadia spinosa Walcott, Burgessochaeta setigera (Walcott) gen. nov. and Peronochaeta dubia (Walcott) gen. nov. are redescribed on the basis of Walcott's type specimens and on much additional material. Two new polychaetes Insolicorypha psygma gen. et sp. nov. and Stephenoscolex argutus gen. et sp. nov. are described. A poorly preserved specimen of unknown generic affinity is described as type A. The polychaetes are preserved as thin films that adhere to both sides of the split in the rock so that part and counterpart may be available. In C. spinosa, B. setigera and I. psygma, parts of the bodies such as the fascicles of setae are separated by thin layers of sediment that apparently seeped in during turbulent transport in turbidites or mudflows. The bodies therefore lie on two or more planes of microbedding and the factors that control exposure across a specimen are discussed. Aspects of the palaeoecology of the Burgess Shale are reviewed, including the distance the biota was transported prior to burial, the reasons for the exquisite preservation, and the effects of sedimentary compaction. C. spinosa was characterized by broad notosetae that extended across the dorsum, and large fascicles of neurosetae. Lobate branchiae were situated in the inter-ramal spaces. The prostomium bore a pair of elongate tentacles and the straight gut had an eversible unarmed proboscis. Several lines of evidence suggest that C. spinosa was an active benthonic swimmer. B. setigera was peculiar in possessing identical notosetae and neurosetae along the entire body. Long anterior tentacles, possibly of peristomial origin, may have been used in feeding. Indirect evidence indicates that B. setigera inhabited a burrow which it might have excavated with its proboscis. P. dubia may also have burrowed but it had uniramous parapodia bearing simple and acicular setae. The prostomium bore a pair of short appendages. I. psygma had extended neuropodia bearing cirri and elongate setae. The notopodia were reduced and cirri appear to have been wanting. The peculiar prostomium carried a pair of appendages. I. psygma is regarded as a pelagic polychaete. S. argutus possessed uniramous parapodia with simple stout setae. The bilobed prostomium bore at least one pair, and perhaps three pairs, of short appendages. Type A was the largest of the Burgess Shale polychaetes and had prominent setae on at least the anterior section of the body. Type A was a sediment eater but the feeding habits of the other polychaetes are uncertain. Particular attention is given to the influence of decay on the Burgess Shale polychaetes. To place the Burgess Shale polychaetes in some geological perspective other Cambrian worms, including a polychaete from the Spence Shale of Utah, are briefly redescribed and the late Precambrian (Ediacarian) worms Dickinsonia, Spriggina and Marywadea are assessed. Contrary to the findings of other workers, no convincing evidence for placing these latter worms in the polychaetes is forthcoming.
TL;DR: It is demonstrated that constraining cladograms in such a manner that wiwaxiid sclerites, canadiid notochaetae and chrysopetalid paleae are homologous leads to results that are markedly unparsimonious.
Abstract: Wiwaxia corrugata and the indisputable polychaetes of the Middle Cambrian Burgess Shale, particularly Canadia spinosa, have figured prominently in recent hypotheses about the early evolution of polychaete annelids. Based on similarities between the sclerites of Wiwaxia and the notochaetae of Canadia with the broad notochaetae (paleae) of Recent chrysopetalid polychaetes, these two fossil taxa have been variously treated as closely related to the most highly derived stem forms of the polychaete (and annelid) crown group or as members of a specific, Recent subgroup within Polychaeta, the order Phyllodocida. Chrysopetalidae is a member of Phyllodocida, which is part of the major polychaete clade Aciculata; the latter two taxa are distinguished by four and six well defined autapomorphic characters, respectively. The best preserved or otherwise appropriate fossils of Wiwaxia corrugata, Canadia spinosa and the other polychaetes of the Burgess Shale have been studied in detail in order to determine whether they possess any characters that could support the homology of wiwaxiid sclerites, canadiid notochaetae and chrysopetalid paleae. Most of these fossil taxa have significant autapomorphies but the specific characters of the Aciculata and Phyllodocida are entirely absent. It is demonstrated that constraining cladograms in such a manner that wiwaxiid sclerites, canadiid notochaetae and chrysopetalid paleae are homologous leads to results that are markedly unparsimonious. Furthermore, Canadia and the other polychaetes of the Burgess Shale cannot be referred to any extant subgroup within the Polychaeta and cannot be used to polarize character evolution within the annelid crown group. Apart from its dubious sclerites, Wiwaxia has no characters that could indicate any close relationship with Polychaeta or Annelida.