Self-regulation of trilobite diversity in Murero (middle Cambrian, Spain) due to compensatory extinction

As species accumulate in a community, competition for available ecospace is expected to prevent the addition of new species and to facilitate species extinction, thus producing a dynamic equilibrium of diversity. This mechanism remains under debate since its empirical support comes mainly from indirect or partial evidence, with very few direct tests at the species level. Here a new method is described to detect the presence of selfregulation feedbacks between species richness and turnover rates. It consists of Monte Carlo simulations which randomize the distribution of species ranges among stratigraphic intervals, providing predictions which allow the detection of genuine self-regulation feedbacks in the real data. Since the simulations include any potential bias due to preservation, sampling, or change in depositional environment, and these biases would also affect the real dataset, they are thus ruled out as explanations for any difference found. This method is applied to one of the best known fossiliferous sequences worldwide, the Rambla de Valdemiedes in Murero (RV1 section, middle Cambrian, Spain), a classic locality that has been studied for more than 150 years and which stands out due to its excellent sampling density, continuous deposition, and homogeneous fossil preservation. The results show that trilobite species richness was self-regulated due to positive feedback with extinction rate, which implies that compensatory extinction regulated this fauna in spite of the on-going Cambrian radiation. The lack of evidence of any origination feedback suggests ecological opportunities were not limiting for new species to colonize this Cambrian community