Early Cambrian Problematic Lophotrochozoans and Dilemmas of Scleritome Reconstructions

Abstract: The emergence and radiation of metazoan body plans around the Precambrian/Cambrian boundary, some 500-600 million years ago, seems to be concordant with the appearance and diversification of preservable hard parts. Several Precambrian soft-bodied, multicellular organisms most likely represent stem-group bilaterians, but their fossil record is rather sparse.  In contrast, the Cambrian fossil record is comparably rich – comprising hard part, trace fossil and delicate soft tissue preservation – and most animal phyla that we know of today had evolved by the end of the Cambrian. Consequently, this time represents an important period in the early evolution of metazoan life forms. Most skeletal remnants of invertebrate organisms from this period are preserved in incomplete, disarticulated sclerite assemblages, and the true architecture of the original skeletal structure, the scleritome, may therefore be hard to discern. Many scleritomous taxa have been suggested to be members of the lophotrochozoan clade, while their exact position within this group remains unclear. Such taxa are often referred to as Problematica. This thesis deals with some problematic scleritomous early Cambrian lophotrochozoans, and as such also addresses the dilemmas of scleritome reconstructions. In the first part, completely disarticulated calcareous sclerites from the lower Cambrian of North Greenland are described as Trachyplax arctica. Hypothetical scleritome reconstruction alternatives and comparisons to other scleritome-bearing taxa are discussed, but the lack of articulated material obscures any satisfactory conclusions regarding phylogenetic affinities and the original morphology of the organism. The other part of the thesis focuses on some minute, organophosphatic scleritomous metazoans, tommotiids, found in lower Cambrian limestone successions in South Australia – Paterimitra pyramidalis and Kulparina rostrata – their scleritome architecture and their phylogenetic relationship with paterinid brachiopods. The oldest brachiopod from South Australia, Askepasma saproconcha, and the slightly younger Askepasma toddense are also described and discussed. Based on articulated specimens, recently described partial scleritomes of the tommotiid Eccentrotheca helenia and similarities in shell ultrastructure with both Eccentrotheca and Askepasma, Paterimitra is interpreted as a stem-group brachiopod and reconstructed as a bilaterally symmetrical, sessile, filter feeder with a tubular/conical scleritome. The morphological similarities with Paterimitra point in the same direction for the slightly older Kulparina.