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Orthrozanclus is a shell-bearing, sclerite covered Cambrian organism of uncertain taxonomic affinity, seemingly representing an intermediate between its fellow problematica Wiwaxia and Halkieria. Attempts to group these slug-like taxa into a single ‘halwaxiid’ clade nevertheless present structural and evolutionary difficulties. Here we report a new species of Orthrozanclus from the early Cambrian Chengjiang Lagerstätte. The scleritome arrangement and constitution in this material corroborates the link between Orthrozanclus and Halkieria, but not with Wiwaxia — and calls into question its purported relationship with molluscs. Instead, the tripartite construction of the halkieriid scleritome finds a more compelling parallel in the camenellan tommotiids, relatives of the brachiopods and phoronids. Such a phylogenetic position would indicate the presence of a scleritome in the common ancestor of the three major trochozoan lineages, Mollusca, Annelida and Brachiozoa. On this view, the absence of fossil Ediacaran sclerites is evidence against any ‘Precambrian prelude’ to the explosive diversification of these phyla in the Cambrian, c. 540–530 million years ago.

Wiwaxiids are a problematic group of scale-covered lophotrochozoans known from Cambrian Stages 3–5. Their imbricating dorsal scleritome of leaf-like scales has prompted comparison with various annelids and molluscs and has been used as a template to reconstruct the articulation pattern of isolated Small Shelly Fossils. The first articulated specimens of Wiwaxia from the Cambrian Stage 3 Chengjiang Konservat-Lagerstätte show that the Wiwaxia scleritome comprised nine equivalent transverse rows associated with outgrowths of soft tissue, but did not possess a separate zone of anterior sclerites. This serial construction is fundamentally incompatible with the circumferential disposition of sclerites in early molluscs, but does closely resemble the armature of certain annelids. A deep homology with the annelid scleritome must be reconciled with Wiwaxia’s mollusc-like mouthparts and foot; together these point to a deep phylogenetic position, close to the common ancestor of annelids and molluscs.

Ecological interactions, including symbiotic associations such as mutualism, parasitism and commensalism are crucial factors in generating evolutionary novelties and strategies. Direct examples of species interactions in the fossil record generally involve organisms attached to sessile organisms in an epibiont or macroboring relationship. Here we provide support for an intimate ecological association between a calcareous brachiopod (Nisusia) and the stem group mollusc Wiwaxia from the Burgess Shale. Brachiopod specimens are fixed to Wiwaxia scleritomes, the latter showing no signs of decay and disarticulation, suggesting a live association. We interpret this association as the oldest unambiguous example of a facultative ectosymbiosis between a sessile organism and a mobile benthic animal in the fossil record. The potential evolutionary advantage of this association is discussed, brachiopods benefiting from ease of attachment, increased food supply, avoidance of turbid benthic conditions, biofoul and possible protection from predators, suggesting commensalism (benefiting the symbiont with no impact for the host). While Cambrian brachiopods are relatively common epibionts, in particular on sponges, the association of Nisusia with the motile Wiwaxia is rare for a brachiopod species, fossil or living, and suggests that symbiotic associations were already well established and diversified by the "middle'' (Series 3, Stage 5) Cambrian.