The role of abiotic factors in the Cambrian Substrate Revolution: A review from the benthic community replacements of West Gondwana

The Cambrian Substrate Revolution refers to a substantial and “rapid” change to the nature of marine sedimentary substrates in the early Cambrian and is widely interpreted as a biologically-driven event, a direct response to evolutionary innovations in metazoan burrowing and the development of new shelly faunas. However, abiotic factors such as tectonic and climatic evolution also had the potential to restructure Cambrian substrates, and are here shown to be more plausible drivers of change in the benthic faunas of western Gondwana. The western Mediterranean region underwent a southward drift during Cambrian times, which drove a switch from subtropical carbonates to temperate siliciclastic substrates with short-term episodes of temperate carbonate productivity. As a result, microbial and shelly carbonates disappeared diachronously in a stepwise manner across the lower–middle Cambrian boundary interval. Archaeocyathan-microbial reefs were replaced by chancelloriid-eocrinoid-(spiculate) sponge meadows, in which the stepwise immigration of new echinoderm taxa was primarily controlled by extensional tectonic events, first recorded in rifting settings and later in passive-margin platforms. Availability of new kinds of substrate was thus the primary factor that controlled where and when evolutionary innovations in benthic strategies arose. Examples of this include the early Cambrian colonization of phosphatic hardgrounds and thrombolite crusts by chancelloriids, archaeocyathan and spiculate sponges, and the exploitation by benthos to the increasingly widespread availability of shelly grounds and carbonate firmgrounds by early-diagenetic cementation. A microbial mat/epifaunal antagonistic relationship is demonstrated for echinoderm pelmatozoans based on the non-overlapping palaeogeographic distributions of microbial reefs and mats versus mud-sticker pelmatozoans. Cambrian benthic communities thus evolved in parallel with substrates in response to abiotic factors rather than being the primary drivers of substrate change.