Allison Daley

In Cambrian fossil Lagerstätten like the Burgess Shale, exceptionally preserved arthropods constitute a large part of the taxonomic diversity, providing opportunities to study the early evolution of this phylum in detail. The anomalocaridids, large presumed pelagic predators, are particularly relevant owing to their unique combination of morphological characters and basal position in the arthropod stem lineage. Although isolated elements and fragmented specimens were first discovered over 100 years ago, subsequent findings of more complete bodies of Anomalocaris and Peytoia, especially in the 1980s, allowed for a better understanding of these enigmatic forms. Their evolutionary significance as stem group arthropods was further clarified by the recent discovery of a third anomalocaridid taxon, Hurdia. Here, examination of hundreds of Hurdia specimens from different stratigraphical layers within the Burgess Shale and Stephen Formation, combined with statistical analyses, provides a detailed description of the taphonomy, morphology and diversity of the genus and further elucidates anomalocaridid systematics. Hurdia is distinguished from other anomalocaridids in having mouthparts with extra rows of teeth, a large frontal carapace complex and diminutive swimming flaps with prominent setal structures. The two original species, H. victoria Walcott, 1912 and H. triangulata Walcott, 1912, are confirmed based on morphometric outline analyses of the frontal carapace components combined with stratigraphical evidence; a third species, Hurdia dentata Simonetta & Delle Cave, 1975, is synonymized with H. victoria. Morphology, preservation and stratigraphical distribution suggest that H. victoria and H. triangulata share the same type of frontal appendage; a second type of appendage, previously assigned to Hurdia (Morph A), belongs to Peytoia nathorsti. These and other morphological differences between the anomalocaridids may reflect different feeding strategies. Appendages and mouthparts of Hurdia indet. sp. are also identified from the Spence Shale Member of Utah, making Hurdia and Anomalocaris the most common and globally distributed anomalocaridid taxa.

Two species of Anomalocaris co-occur in the Emu Bay Shale (Cambrian Series 2, Stage 4) at Big Gully, Kangaroo Island. Frontal appendages of Anomalocaris briggsi Nedin, 1995, are more common than those of Anomalocaris cf. canadensis Whiteaves, 1892, at a quarry inland of the wave-cut platform site from which these species were originally described. An oral cone has the three large, node-bearing plates recently documented for Anomalocaris canadensis, confirming that Anomalocaris lacks a tetraradial ‘Peytoia’ oral cone and strengthening the case for the identity of the Australian specimens as Anomalocaris. Disarticulated anomalocaridid body flaps are more numerous in the Emu Bay Shale than in other localities, and they preserve anatomical details not recognized elsewhere. Transverse lines on the anterior part of the flaps, interpreted as strengthening rays or veins in previous descriptions of anomalocaridids, are associated with internal structures consisting of a series of well-bounded, striated blocks or bars. Their structure is consistent with a structural function imparting strength to the body flaps. Setal structures consisting of a series of lanceolate blades are similar to those of other anomalocaridids and are found in isolation or associated with body flaps. A single specimen also preserves putative gut diverticula. The morphology of the appendages, oral cone, gut diverticula and compound eyes of Anomalocaris, along with its large size, suggests that it was an active predator, and specimens of coprolites containing trilobite fragments and trilobites with prominent injuries have been cited as evidence of anomalocaridid predation on trilobites. Based on frontal appendage morphology, Anomalocaris briggsi is inferred to have been a predator of soft-bodied animals exclusively and only Anomalocaris cf. canadensis may have been capable of durophagous predation on trilobites, although predation (including possible cannibalism) by Redlichia could also explain the coprolites and damage to trilobite exoskeletons found in the Emu Bay Shale.