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      Fine-scale appendage structure of the Cambrian trilobitomorph Naraoia spinosa and its ontogenetic and ecological implications

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          Abstract

          Trilobitomorphs are a species-rich Palaeozoic arthropod assemblage that unites trilobites with several other lineages that share similar appendage structure. Post-embryonic development of the exoskeleton is well documented for some trilobitomorphs, especially trilobites, but little is known of the ontogeny of their soft parts, limiting understanding of their autecology. Here, we document appendage structure of the Cambrian naraoiid trilobitomorph Naraoia spinosa by computed microtomography, resulting in three-dimensional reconstructions of appendages at both juvenile and adult stages. The adult has dense, strong spines on the protopods of post-antennal appendages, implying a predatory/scavenging behaviour. The absence of such gnathobasic structures, but instead tiny protopodal bristles and a number of endopodal setae, suggests a detritus-feeding strategy for the juvenile. Our data add strong morphological evidence for ecological niche shifting by Cambrian arthropods during their life cycles. A conserved number of appendages across the sampled developmental stages demonstrates that Naraoia ceased budding off new appendages by the mid-juvenile stage.

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          Paleoecology of the Greater Phyllopod Bed community, Burgess Shale

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            The ontogeny of trilobite segmentation: a comparative approach

            Ontogenetic stages of trilobites have traditionally been recognized on the basis of the development of exoskeletal segmentation. The established protaspid, meraspid, and holaspid phases relate specifically to the development of articulated joints between exoskeletal elements. Transitions between these phases were marked by the first and last appearances of new trunk segment articulations. Here we propose an additional and complementary ontogenetic scheme based on the generation of new trunk segments. It includes an anamorphic phase during which new trunk segments appeared, and an epimorphic phase during which the number of segments in the trunk remained constant. In some trilobites an ontogenetic boundary can also be recognized at the first appearance of morphologically distinct posterior trunk segments. Comparison of the phase boundaries of these different aspects of segment ontogeny highlights rich variation in the segmentation process among Trilobita. Cases in which the onset of the holaspid phase preceded onset of the epimorphic phase are here termed protarthrous , synchronous onset of both phases is termed synarthromeric , and onset of the epimorphic phase before onset of the holaspid phase is termed protomeric . Although these conditions varied among close relatives and perhaps even intraspecifically in some cases, particular conditions may have been prevalent within some clades. Trilobites displayed hemianamorphic development that was accomplished over an extended series of juvenile and mature free-living instars. Although developmental schedules varied markedly among species, morphological transitions during trilobite development were generally regular, limited in scope, and extended over a large number of instars when compared with those of many living arthropods. Hemianamorphic, direct development with modest change between instars is also seen among basal members of the Crustacea, basal myriapods, pycnogonids, and in some fossil chelicerates. This mode may represent the ancestral condition of euarthropod development.
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              Relationships of Cambrian Arachnata and the systematic position of Trilobita

              Cladistic relationships of Trilobita, Naraoiidae (five ingroup taxa), Helmetiida (five ingroup taxa), Xandarellida, and the Cambrian arachnatesRetifacies, Sinoburius, Emeraldella, andSidneyiaare investigated based on 29 characters. Documentation of appendage morphology and other ventral structures inSaperionfrom the Chengjiang fauna permits an appraisal of helmetiid relationships. A monophyletic Trilobita [=“Olenellida” (Emuellida + Eutrilobita)] is defined by numerous synapomorphies, including exoskeletal calcification and dorsal eyes with calcified lenses and circumocular sutures. Helmetiida is a robust clade, resolved as (Helmetiidae (Tegopeltidae (Saperiidae + Skioldiidae))). Naraoiid monophyly is well-supported, but neither a naraoiid-trilobite nor a naraoiid-Retifaciesclade are parsimonious, the latter grouping (“Nectopleura”) being explicitly paraphyletic. A sister group relationship between Xandarellida andSinoburiusis endorsed, although character support is novel compared to previous groupings of these taxa. The fourth postantennal limb pair in trilobites, naraoiids, and apparently helmetiids is based beneath the cephalothoracic articulation. Reweighted characters favor Trilobita and Helmetiida as closest relatives, with Petalopleura and then Naraoiidae as sister groups.
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                Author and article information

                Journal
                Proc Biol Sci
                Proc. Biol. Sci
                RSPB
                royprsb
                Proceedings of the Royal Society B: Biological Sciences
                The Royal Society
                0962-8452
                1471-2954
                4 December 2019
                4 December 2019
                4 December 2019
                : 286
                : 1916
                : 20192371
                Affiliations
                [1 ]Yunnan Key Laboratory for Palaeobiology, Yunnan University , 2 North Cuihu Road, Kunming 650091, People's Republic of China
                [2 ]MEC International Joint Laboratory for Palaeobiology and Palaeoenvironment, Yunnan University , 2 North Cuihu Road, Kunming 650091, People's Republic of China
                [3 ]Department of Earth Sciences, The Natural History Museum , Cromwell Road, London SW75BD, UK
                [4 ]Department of Earth Sciences, Royal Holloway University of London , Egham, Surrey TW200EX, UK
                Author notes

                Electronic supplementary material is available online at https://doi.org/10.6084/m9.figshare.c.4723397.

                Author information
                http://orcid.org/0000-0001-6312-851X
                http://orcid.org/0000-0002-9591-8011
                Article
                rspb20192371
                10.1098/rspb.2019.2371
                6939273
                31795867
                c453bcb7-4d43-450e-b88b-748a9e69d37a
                © 2019 The Authors.

                Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited.

                History
                : 9 October 2019
                : 28 October 2019
                Funding
                Funded by: Yunnan Provincial Science and Technology Department, http://dx.doi.org/10.13039/501100008871;
                Award ID: 2015HA021
                Award ID: 2015HC029
                Award ID: 2018FA025
                Award ID: 2018IA073
                Funded by: National Natural Science Foundation of China, http://dx.doi.org/10.13039/501100001809;
                Award ID: 41472153
                Award ID: 41861134032
                Categories
                1001
                144
                Palaeobiology
                Research Article
                Custom metadata
                December 4, 2019

                Life sciences
                cambrian,chengjiang biota,micro-ct,naraoia spinosa,arthropod,gnathobase
                Life sciences
                cambrian, chengjiang biota, micro-ct, naraoia spinosa, arthropod, gnathobase

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