Survival on a soft seafloor: life strategies of brachiopods from the Cambrian Burgess Shale

Understanding the structure of benthic communities in the Cambrian remains a major challenge. Direct evidence for species interrelationships is rare and therefore past ecological interactions typically cannot be reconstructed with great accuracy. Here we reveal the community patterns and modes of life of brachiopods - one of the most important filter-feeding groups of Cambrian ecosystems - from the Cambrian Burgess Shale Lagerstatte. Burgess Shale brachiopods attached to a range of hard substrates, including skeletal debris, conspecific brachiopods and enigmatic tubes, with an overwhelming preference for attachment on the demosponge Pirania muricata. The dominance of P. muricata as a substrate choice - even in bedding assemblages where P. muricata individuals are rare - and similarities to the gregarious attachment strategies of extant brachiopod species suggests that brachiopods larvae in the Burgess Shale community selected their attachment substrates. The distribution of brachiopod taxa is also intricately linked with the presence of suitable hard substrates, with species declining in bedding assemblages where their preferred hard substrates are absent. In addition, brachiopods in the Burgess Shale community are predominantly low epifaunal tierers and do not exploit niches high in the water column, despite the presence of suitable attachment sites. Our analysis of tiering height versus host height indicates that there is no selection by brachiopod larvae in regard to the height of attachment and individuals attached at the first point of contact with the selected substrate. Through comparisons with the 'early' Cambrian Chengjiang Biota, we confirm that by the 'middle' Cambrian (Series 3, Stage 5) brachiopods had already developed a range of attachment strategies similar to some modern brachiopod populations. Our results provide significant insight into the ecological constraints and adaptability of brachiopods in the earliest animal communities of the Cambrian. (C) 2015 Elsevier B.V. All rights reserved.