Beyond the Burgess Shale: Cambrian microfossils track the rise and fall of hallucigeniid lobopodians

Abstract

Burgess Shale-type deposits are renowned for their exquisite preservation of soft-bodied organisms, representing a range of animal body plans that evolved during the Cambrian 'explosion'. However, the rarity of these fossil deposits makes it difficult to reconstruct the broader-scale distributions of their constituent organisms. By contrast, microscopic skeletal elements represent an extensive chronicle of early animal evolution--but are difficult to interpret in the absence of corresponding whole-body fossils. Here, we provide new observations on the dorsal spines of the Cambrian lobopodian (panarthropod) worm Hallucigenia sparsa from the Burgess Shale (Cambrian Series 3, Stage 5). These exhibit a distinctive scaly microstructure and layered (cone-in-cone) construction that together identify a hitherto enigmatic suite of carbonaceous and phosphatic Cambrian microfossils--including material attributed to Mongolitubulus, Rushtonites and Rhombocorniculum--as spines of Hallucigenia-type lobopodians. Hallucigeniids are thus revealed as an important and widespread component of disparate Cambrian communities from late in the Terreneuvian (Cambrian Stage 2) through the 'middle' Cambrian (Series 3); their apparent decline in the latest Cambrian may be partly taphonomic. The cone-in-cone construction of hallucigeniid sclerites is shared with the sclerotized cuticular structures (jaws and claws) in modern onychophorans. More generally, our results emphasize the reciprocal importance and complementary roles of Burgess Shale-type fossils and isolated microfossils in documenting early animal evolution.

Keywords: Cambrian evolutionary radiation; Lagerstätten; biostratigraphy; small carbonaceous fossils; small shelly fossils.

Figure 1.

Hallucigenia sparsa from the Burgess Shale: (a,b) Smithsonian Institution, National Museum of Natural History (NMNH) 83935 (holotype), articulated specimen, showing seven pairs of spines, partially decayed towards the rear, presumed head to the right. (a) composite image of part and counterpart; (b) enlargement of the basal part of the spines; (c,d) Royal Ontario Museum (ROM) 61513, complete specimen showing seven pairs of spines and backscatter image of boxed area (d); (e–i) ROM 57776, backscatter images (overview and close-ups of boxed areas) of spine showing four internal cones and lineations; (g) ROM 61513, backscatter image showing lineations and a distal cone; (j–o) ROM 62269, backscatter images of several spines, showing elemental distribution of carbon (l) and phosphorous (m) and details of ornamentation near spines’ mid-length (n) and base (o) (arrows indicate local disturbances in the rhomboid pattern); (p) ROM 61513, backscatter image showing details of ornamentation showing scales in positive relief (top left) and negative relief below the carbon film. Ba, basal region of spines; C, cone; Li, lineations. Scale bars: (a–d) 1000 µm; (e,j–m) 100 µm; (f–i) 50 µm; (n–p) 10 µm. (Online version in colour.)

Figure 2.

Hallucigeniid spines preserved as small carbonaceous fossils. (a–h) Isolated spines from the Mount Cap Formation (Little Bear biota), northwestern Canada: (a–b) Geological Survey of Canada (GSC) 136957, showing longitudinal striations within spine; (c–f) GSC 136958, showing that the spine does not flare at its base, where the ornament thins; (g,h) GSC 136959, cone-in-cone structure and lineations within spine, surface ornament diminishing towards apex; (i–k) GSC 136960, potential new hallucigeniid morph from the Forteau Formation, Newfoundland, showing diffuse ornament and straight sides near base. C, cone; Li, lineations. Scale bars: (a,c,g–i) 100 µm; (b,d–f,j,k) 25 µm. (Online version in colour.)