Stephen Pates

Hurdiid radiodontans from the middle Cambrian (Series 3) of Utah

Abstract. n Radiodontan body elements, some belonging to Peytoia and Hurdia and some unassigned, have been reported from the Langston Formation (Spence Shale Member), Wheeler Formation, and Marjum Formation of the middle Cambrian (Series 3) of Utah. These identifications are reassessed in light of recent work on the morphology of the radiodontan Hurdia. New specimens of Hurdia are identified from the Spence Shale, representing mouthparts (oral cones), cephalic carapace H-elements, frontal appendages, and a single isolated swimming flap. The shape of the H-elements allows H. victoria Walcott, 1912 to be identified from the Spence Shale for the first time. The flap is larger and more complete than any reported from the Burgess Shale and allows for a better understanding of the morphology of Hurdia swimming flaps. A 3D model of a Hurdia frontal appendage indicates that there is only one morph of Hurdia frontal appendage found in both species, and apparent morphological differences between disarticulated appendages reflect a preservational continuum caused by varying oblique angles relative to the seafloor. Peytoia should no longer be reported from the Spence Shale, but its presence is confirmed in the Wheeler and Marjum formations. New mouthparts (oral cones) of Hurdia from the Spence Shale and Peytoia from the Marjum Formation with surface textures of submillimeter-diameter raised nodes are described. These new features have not been observed in material from the Burgess Shale and suggest slight differences in preservation.

Early fossil record of Euarthropoda and the Cambrian Explosion

Euarthropoda is one of the best-preserved fossil animal groups and has been the most diverse animal phylum for over 500 million years. Fossil Konservat-Lagerstätten, such as Burgess Shale-type deposits (BSTs), show the evolution of the euarthropod stem lineage during the Cambrian from 518 million years ago (Ma). The stem lineage includes nonbiomineralized groups, such as Radiodonta (e.g., Anomalocaris) that provide insight into the step-by-step construction of euarthropod morphology, including the exoskeleton, biramous limbs, segmentation, and cephalic structures. Trilobites are crown group euarthropods that appear in the fossil record at 521 Ma, before the stem lineage fossils, implying a ghost lineage that needs to be constrained. These constraints come from the trace fossil record, which show the first evidence for total group Euarthropoda (e.g., Cruziana, Rusophycus) at around 537 Ma. A deep Precambrian root to the euarthropod evolutionary lineage is disproven by a comparison of Ediacaran and Cambrian lagerstätten. BSTs from the latest Ediacaran Period (e.g., Miaohe biota, 550 Ma) are abundantly fossiliferous with algae but completely lack animals, which are also missing from other Ediacaran windows, such as phosphate deposits (e.g., Doushantuo, 560 Ma). This constrains the appearance of the euarthropod stem lineage to no older than 550 Ma. While each of the major types of fossil evidence (BSTs, trace fossils, and biomineralized preservation) have their limitations and are incomplete in different ways, when taken together they allow a coherent picture to emerge of the origin and subsequent radiation of total group Euarthropoda during the Cambrian.

Abnormal xiphosurids, with possible application to Cambrian trilobites

Xiphosurida comprise an archetypal arthropod group of considerable interest to both biological and palaeontological researchers. This appeal is generated by a combination of unique anatomical features, utility as modern analogues for extinct arthropod groups, and an impressive fossil record. Although xiphosurids have been extensively studied, there are few published examples of abnormal specimens. Abnormalities in xiphosurids have mostly been attributed to injuries (either self-inflicted, from mating, or predation) or teratologies (developmental and genetic malfunctions). Here we summarise all previously recorded extant xiphosurid abnormalities and describe new examples of injuries and teratologies to Limulus polyphemus and Tachypleus tridentatus. Furthermore, we present the first evidence of injured fossil xiphosurids: Euproops danae and Mesolimulus walchi. We identify two main groups of telson teratologies and document new ‘U’ shaped cephalothoracic injuries to the anterior cephalothoracic margins of L. polyphemus and T. tridentatus. We show ‘V’ and ‘W’ shaped injuries to E. danae and M. walchi cephalothoracic sections. A further specimen of E. danae is described, which likely represents plastic deformation of a recently moulted exoskeleton, rather than an abnormality sensu stricto. We compare injuries on extant xiphosurids to extinct Cambrian trilobite injuries to suggest that rare cephalic injuries to trilobites were incurred during soft-shelled exoskeletal stages. Reviewing xiphosurid injuries through time is a pivotal step towards understanding how Recent and extinct arthropods responded to injuries. Russell D.C. Bicknell. Palaeoscience Research Centre, School of Environmental and Rural Science, University of New England, Armidale, New South Wales, 2351, Australia. rdcbicknell@gmail.com Stephen Pates. Department of Zoology, University of Oxford, Oxford, OX1 3PS, United Kingdom. stephen.pates@zoo.ox.ac.uk and Institute of Earth Sciences, University of Lausanne, Lausanne, CH1015, Switzerland. Mark L. Botton. Department of Natural Sciences, Fordham University, New York, NY 10023, USA. botton@fordham.edu Bicknell, Russell D.C., Pates, Stephen, and Botton, Mark L. 2018. Abnormal xiphosurids, with possible application to Cambrian trilobites. Palaeontologia Electronica 21.2.19A 1-17. https://doi.org/10.26879/866 Copyright: June 2018 Paleontological Society. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. creativecommons.org/licenses/by/4.0/ BICKNELL, PATES, & BOTTON: ABNORMAL XIPHOSURIDS

New suspension-feeding radiodont suggests evolution of microplanktivory in Cambrian macronekton

The rapid diversification of metazoans and their organisation in modern-style marine ecosystems during the Cambrian profoundly transformed the biosphere. What initially sparked this Cambrian explosion remains passionately debated, but the establishment of a coupling between pelagic and benthic realms, a key characteristic of modern-day oceans, might represent a primary ecological cause. By allowing the transfer of biomass and energy from the euphotic zone—the locus of primary production—to the sea floor, this biological pump would have boosted diversification within the emerging metazoan-dominated benthic communities. However, little is known about Cambrian pelagic organisms and their trophic interactions. Here we describe a filter-feeding Cambrian radiodont exhibiting morphological characters that likely enabled the capture of microplankton-sized particles, including large phytoplankton. This description of a large free-swimming suspension-feeder potentially engaged in primary consumption suggests a more direct involvement of nekton in the establishment of an oceanic pelagic-benthic coupling in the Cambrian.Planktonic life was well established by the Cambrian, but few nektonic filter feeders have been identified. Here, the authors provide fossil evidence that Pahvantia hastata was a suspension-feeder that likely captured much smaller plankton than any other known free-swimming animals of that time.

QUANTITATIVE ANALYSIS OF REPAIRED AND UNREPAIRED DAMAGE TO TRILOBITES FROM THE CAMBRIAN (STAGE 4, DRUMIAN) IBERIAN CHAINS, NE SPAIN

Abstract: Repaired fossil skeletons provide the opportunity to study predation rates, repair mechanisms, and ecological interactions in deep time. Trilobites allow the study of repaired damage over long time periods and large geographic areas due to their longevity as a group, global distribution, and well-preserved mineralized exoskeletons. Repair frequencies on trilobites from three sites representing offshore marine environments in the Iberian Chains (Spain) show no injuries on 45 complete redlichiid thoraces from Minas Tierga (Huérmeda Formation, Cambrian Series 2, Stage 4), or 23 complete Eccaparadoxides pradoanus thoraces from Mesones de Isuela (Murero Formation, Cambrian Series 3, Drumian). Ten injuries on 69 E. pradoanus thoraces from Purujosa (Murero Formation, Cambrian Series 3, Drumian) were noted. There is no evidence for laterally asymmetric predation or size selection on the trilobites in this study. Weak evidence for selection for the rear of the thorax is documented. A series of injured trilobites illustrates four stages of the healing process. Analysis of injury locations and frequency suggests that injuries to these trilobites are predatory in origin. Semilandmark analysis of previously described exoskeletons with unrepaired damage assigned to the ichnotaxon Bicrescomanducator serratus alongside newly collected damaged exoskeletons from Purujosa (Mansilla and Murero Formations, Stage 5, Drumian), Mesones de Isuela (Murero Formation, Drumian), and Minas Tierga (Huérmeda Formation, Stage 4) found that shapes of biotic and abiotic breaks could not be distinguished.