The Middle Cambrian metazoanWiwaxia corrugata(Matthew) from the Burgess Shale andOgygopsisShale, British Columbia, Canada

Abstract

A detailed redescription of the Middle Cambrian metazoan Wiwaxia corrugata (Matthew) is given with the aid of photographs, mostly taken under ultraviolet radiation, and explanatory camera lucida drawings. Wiwaxia is known only from the Stephen Formation with four localities within the Bathyuriscus-Elrathina Zone, the celebrated Burgess Shale (Phyllopod bed) and Ogygopsis Shale and two localities that appear to expose strata relatively high in the Formation, and a new locality in the underlying Glossopleura Zone. Specimens from the Ogygopsis Shale consist almost entirely of isolated sclerites, whereas in the Phyllopod bed the species is also known from entire specimens, semi-isolated assemblages and isolated soft-parts. T he description here is based almost entirely on Phyllopod bed m aterial. Wiwaxia was originally largely covered by sclerites. On the basis of shape and arrangement five distinct zones are recognizable: dorsal, upper lateral, lower lateral, anterior and ventro-lateral. In addition, there was a row of elongate dorso-lateral spinose sclerites running along either side of the body. Each sclerite was inserted separately via a root-like structure. Growth of the animal appears to have occurred by moulting of the sclerites; one unique juvenile specimen appears to be preserved in the act of moulting. The ventral surface was apparently an area of soft tissue devoid of sclerites. Little is known of the internal anatomy, although anteriorly there was a feeding apparatus consisting of two rows of posteriorly directed teeth. Most stages of growth are known varying from presumed juveniles, which at their smallest may have lacked spines, to adults over 50 mm long. Wiwaxia appears to have been an epifaunal deposit feeder that crawled across the sea-bed, although smaller juveniles may have been infaunal. The dorso-lateral spines may have provided protection against predators and the existence of broken spines suggests the successful deterrence of attack. The affinities of Wiwaxia do not appear to lie with the polychaetous annelids. While the possibilities of convergence cannot be eliminated, there appears to be a significant similarity between Wiwaxia and molluscs that suggests a related derivation from a turbellarian ancestor. Nevertheless, Wiwaxia has a distinctive bodyplan and as such cannot be accommodated in any known phylum. While Wiwaxia is unique to the Stephen Formation isolated sclerites from Lower Cambrian rocks appear to represent earlier wiwaxiids, although these sclerites show differences in their size and composition as com pared with Wiwaxia . Wiwaxia enhances knowledge of early metazoan evolution and ecology in the Cambrian. In particular, it gives fresh insights into the origin and relative success of certain metazoan bodyplans, the origin of trace fossils, and the importance of predation in Cambrian communities.

Mouthparts of the Burgess Shale fossils Odontogriphus and Wiwaxia : implications for the ancestral molluscan radula

“…Most chitons, like Neopilina, are deposit feeders [32,33]-reflected by the substantial radular musculature, a limited number of teeth (less than 20) per row and teeth that rotate relative to their neighbours [28]. These traits are shared by Odontogriphus and Wiwaxia (the musculature, at least in Odontogriphus, indicated by the large alary process): this substantiates previous speculation [1] that the organisms were deposit feeders.…”

New records of Burgess Shale-type taxa from the middle Cambrian of Utah

“…In contrast to W. corrugata (Conway Morris, 1985) spines are more densely arrayed (approximately 12 versus an average of approximately 8) and less elongate. On the unproven assumption that the isolated sclerites (Fig.…”

“…Remarks.-Wiwaxiids are best known from articulated material of Wiwaxia corrugata (Matthew, 1899) from the Phyllopod Bed of the Burgess Shale (Conway Morris, 1985). Also occurring in this deposit in moderate abundance are isolated sclerites (and rarely partial associations), detached from the cataphract scleritome either by deciduous action or upon death and post-mortem scattering (see also Butterfield, 1990;Mankiewicz, 1992).…”

“…In contrast, articulated wiwaxiids are very rare. Apart from the Burgess Shale examples (Conway Morris, 1985) and recent discoveries from the Xiaoshiba Lagerstätte and the Kaili Formation (Zhao et al, , 2005, Pl. II, fig.…”

Colour in Burgess Shale animals and the effect of light on evolution in the Cambrian

“…For example, predation on W. corrugata is inferred from broken spines (Conway Morris 1985). The coloration of W. corrugata, C. spinosa and M. splendens was probably aposematic (provided a visual warning of their`spines'): this is suggested by the occurrence of gratings on defensive parts or`armour', as in Aphrodita australis.…”