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      Exploring the use of micro-computed tomography (micro-CT) in the taxonomy of sea cucumbers: a case-study on the gravel sea cucumber Neopentadactylamixta (Östergren, 1898) (Echinodermata, Holothuroidea, Phyllophoridae)

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          Abstract

          Sea cucumber taxonomy and systematics has in the past heavily relied on gross external and internal anatomy, ossicle assemblage in different tissues, and molecular characterisation, with coloration, habitat, and geographical and bathymethric distribution also considered important parameters. In the present paper, we made these observations and techniques in detail and complemented them with the novel technique of micro-computed tomography of the calcareous ring. We investigated a single European species, the so-called gravel sea cucumber, Neopentadactyla mixta (Östergren, 1898), using recently collected material from the Chausey Islands, Normandy, France. We redescribed the species, illustrated its ossicle assemblage through scanning electron microscopy, and visualised the calcareous ring through stacking photography and through micro-CT scanning. Additionally, a DNA fragment of 955 base pairs of the 18S ribosomal RNA gene was sequenced from one specimen, which showed a high similarity with the only sequence of N. mixta publicly available. We completed this integrative study by providing a detailed distribution of the occurrence of N. mixta based on published, verifiable accounts.

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          Molecular phylogeny of extant Holothuroidea (Echinodermata).

          Sea cucumbers (Holothuroidea) are a morphologically diverse, ecologically important, and economically valued clade of echinoderms; however, the understanding of the overall systematics of the group remains controversial. Here, we present a phylogeny of extant Holothuroidea assessed with maximum parsimony, maximum likelihood, and Bayesian approaches using approximately 4.3kb of mt- (COI, 16S, 12S) and nDNA (H3, 18S, 28S) sequences from 82 holothuroid terminals representing 23 of the 27 widely-accepted family-ranked taxa. Currently five holothuroid taxa of ordinal rank are accepted. We find that three of the five orders are non-monophyletic, and we revise the taxonomy of the groups accordingly. Apodida is sister to the rest of Holothuroidea, here considered Actinopoda. Within Actinopoda, Elasipodida in part is sister to the remaining Actinopoda. This latter clade, comprising holothuroids with respiratory trees, is now called Pneumonophora. The traditional Aspidochirotida is paraphyletic, with representatives from three orders (Molpadida, Dendrochirotida, and Elasipodida in part) nested within. Therefore, we discontinue the use of Aspidochirotida and instead erect Holothuriida as the sister group to the remaining Pneumonophora, here termed Neoholothuriida. We found four well-supported major clades in Neoholothuriida: Dendrochirotida, Molpadida and two new clades, Synallactida and Persiculida. The mapping of traditionally-used morphological characters in holothuroid systematics onto the phylogeny revealed marked homoplasy in most characters demonstrating that further taxonomic revision of Holothuroidea is required. Two time-tree analyses, one based on calibrations for uncontroversial crown group dates for Eleutherozoa, Echinozoa and Holothuroidea and another using these calibrations plus four more from within Holothuroidea, showed major discrepancies, suggesting that fossils of Holothuroidea may need reassessment in terms of placing these forms with existing crown clades.
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            Actinien, Echinodermen und Würmer des Adriatischen- und Mittelmeers : nach eigenen Sammlungen / beschrieben von Adolph Eduard Grube.

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              System of the class Holothuroidea.

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                Author and article information

                Contributors
                Role: ConceptualizationRole: Formal analysisRole: InvestigationRole: SupervisionRole: Writing - original draft
                Role: Data curationRole: Formal analysisRole: MethodologyRole: Software
                Journal
                Zookeys
                Zookeys
                2
                urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E
                urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2021
                04 August 2021
                : 1054
                : 173-184
                Affiliations
                [1 ] Scientific Service of Patrimony, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, 1000, Brussels, Belgium Scientific Service of Patrimony, Royal Belgian Institute of Natural Sciences Brussels Belgium
                [2 ] Joint Experimental Molecular Unit, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, 1000, Brussels, Belgium Joint Experimental Molecular Unit, Royal Belgian Institute of Natural Sciences Brussels Belgium
                Author notes
                Corresponding author: Yves Samyn ( ysamyn@ 123456naturalsciences.be )

                Academic editor: D.V. Spiegel

                Article
                67088
                10.3897/zookeys.1054.67088
                8357699
                34393568
                b5fc2a88-6c90-4e06-9a77-abdb2555d548
                Yves Samyn, Gontran Sonet, Cedric d'Udekem d'Acoz

                This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 19 April 2021
                : 07 June 2021
                Funding
                Belgian Science Policy Office
                Categories
                Research Article
                Phyllophoridae
                Taxonomy
                Theory & Methodology
                Cenozoic
                Europe

                Animal science & Zoology
                integrative taxonomy,micro-ct,normandy,scanning electron microscopy
                Animal science & Zoology
                integrative taxonomy, micro-ct, normandy, scanning electron microscopy

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