Large euarthropod carapaces from a high latitude Cambrian (Drumian) deposit in Spain

Deposits preserving non-biomineralized tissues and animals provide an unrivalled opportunity to study the evolution and radiation of early animal life. Numerous sites of Cambrian age are known from North America (Laurentia) and South China (East Gondwana), which provide a high resolution picture of the fauna at low latitudes. By contrast, our knowledge of Cambrian animals from higher latitudes is relatively poor. This patchiness in our knowledge of animal life during the radiation of animals in the Cambrian period limits our ability to understand and detect palaeogeographic trends and does not provide a full appreciation of animal diversity at this time. Here we report a new middle Cambrian (Drumian) site preserving lightly sclerotized euarthropod carapaces, sponges and palaeoscolecids near the village of Mesones de Isuela in the Iberian Chains (Spain). We describe three bivalved euarthropod carapace morphs, two comparable to those described from the only other high latitude Drumian deposit, the Jince Formation (Czechia), and one distinct from previous discoveries. These new findings highlight the importance of high latitude Gondwana Konservat Lagerstatten for understanding the palaeogeographical aspect of the radiation of early animals and suggest that bivalved euarthropods at high latitudes were larger than those at lower latitudes during the Cambrian

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

Vertically migrating Isoxys and the early Cambrian biological pump.

The biological pump is crucial for transporting nutrients fixed by surface-dwelling primary producers to demersal animal communities. Indeed, the establishment of an efficient biological pump was likely a key factor enabling the diversification of animals over 500 Myr ago during the Cambrian explosion. The modern biological pump operates through two main vectors: the passive sinking of aggregates of organic matter, and the active vertical migration of animals. The coevolution of eukaryotes and sinking aggregates is well understood for the Proterozoic and Cambrian; however, little attention has been paid to the establishment of the vertical migration of animals. Here we investigate the morphological variation and hydrodynamic performance of the Cambrian euarthropod Isoxys. We combine elliptical Fourier analysis of carapace shape with computational fluid dynamics simulations to demonstrate that Isoxys species likely occupied a variety of niches in Cambrian oceans, including vertical migrants, providing the first quantitative evidence that some Cambrian animals were adapted for vertical movement in the water column. Vertical migration was one of several early Cambrian metazoan innovations that led to the biological pump taking on a modern-style architecture over 500 Myr ago

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

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.Department of Zoology, University of Oxford, Reino UnidoInstitute of Earth Sciences, University of Lausanne, SuizaPaleoscience Research Centre School of Environmental and Rural Science, University of New England, AustraliaUnidad de Zaragoza, Instituto Geológico y Minero de España, EspañaUnidad Asociada en Ciencias de la Tierra, Universidad de Zaragoza, Españ

First report of paired ventral endites in a hurdiid radiodont.

BACKGROUND: Radiodonta, large Palaeozoic nektonic predators, occupy a pivotal evolutionary position as stem-euarthropods and filled important ecological niches in early animal ecosystems. Analyses of the anatomy and phylogenetic affinity of these large nektonic animals have revealed the origins of the euarthropod compound eye and biramous limb, and interpretations of their diverse feeding styles have placed various radiodont taxa as primary consumers and apex predators. Critical to our understanding of both radiodont evolution and ecology are the paired frontal appendages; however, the vast differences in frontal appendage morphology between and within different radiodont families have made it difficult to identify the relative timings of character acquisitions for this body part. RESULTS: Here we describe a new genus of hurdiid, Ursulinacaris, from the middle Cambrian (Miaolingian, Wuliuan) Mount Cap Formation (Northwest Territories, Canada) and Jangle Limestone (Nevada, USA). Ursulinacaris has the same organisation as other hurdiid frontal appendages, with elongate endites on the first five podomeres in the distal articulated region and auxiliary spines on the distal margin of endites only. Unlike all other hurdiid genera, which possess a single row of elongated and blade-like ventral endites, this taxon uniquely bears paired slender endites. CONCLUSION: The blade-like endite morphology is shown to be a hurdiid autapomorphy. Two other frontal appendage characters known only in hurdiids, namely auxiliary spines on the distal margin of endites only, and elongate endites on the first five podomeres in the distal articulated region only, predate this innovation

Intraspecific variation of early Cambrian (stage 3) arthropod Retifacies abnormalis revealed by morphometric analyses

Retifacies abnormalis is a large artiopodan euarthropod known only from the famous fossil deposits of the Chengjiang biota, China (Cambrian Series 2, Stage 3). It is well known for its pronounced reticulated ornamentation that covers the entire dorsal surface of the exoskeleton. Here 109 new specimens of R. abnormalis from multiple deposits are reported. Some larger specimens display a distinct carapace ornamentation to what was previously known. By qualitatively separating specimens into two groups (‘Morph A’, ‘Morph B’) and analyzing the shape of the body, pygidium, and shape of the polygons in the reticulation, using linear and geometric morphometrics and elliptical Fourier analysis, the two morphs are shown to overlap in morphospace and display similar length:width ratios of body parts, rather than form two distinct clusters. The differences are interpreted as intraspecific rather than as diagnosing two species. As Morph B are only found in larger size classes, R. abnormalis ornamentation differences are interpreted to have developed during ontogeny, but are not thought to represent sexual dimorphs

First report of paired ventral endites in a hurdiid radiodont

Radiodonta, large Palaeozoic nektonic predators, occupy a pivotal evolutionary position as stem-euarthropods and filled important ecological niches in early animal ecosystems. Analyses of the anatomy and phylogenetic affinity of these large nektonic animals have revealed the origins of the euarthropod compound eye and biramous limb, and interpretations of their diverse feeding styles have placed various radiodont taxa as primary consumers and apex predators. Critical to our understanding of both radiodont evolution and ecology are the paired frontal appendages; however, the vast differences in frontal appendage morphology between and within different radiodont families have made it difficult to identify the relative timings of character acquisitions for this body part