New echinoderm holdfast discoveries from Gondwana demonstrate that pelmatozoans have been cementers able to attach to carbonate firmgrounds since the basal middle Cambrian. Echinoderms were thus colonizing shallow, high-energy environments well before the appearance of the first true carbonate hardgrounds in the Furongian. Morphological innovations and adaptations to firmground media ( = substrates) were first developed in softground, clayey, offshore environments where echinoderms cemented to shell fragments. This preadaption allowed echinoderms to quickly and effectively exploit the newly emerging hardground habitats in the Furongian to Early Ordovician.

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