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A reappraisal of Paleozoic horseshoe crabs from Russia and Ukraine

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Abstract

Xiphosura are extant marine chelicerates that have displayed apparent morphological conservatism and remarkable survivorship across their ~ 480 Ma fossil record. The easily recognisable features that are known to even the earliest xiphosurans—a crescentic prosoma and often trapezoidal thoracetron (opisthosoma)—have generated debate surrounding their origins and taxonomic significance. This interest resulted in the description of numerous horseshoe crab species during the early to mid-twentieth century, particularly in Russia, that have remained unrevised since their original publications and unconsidered in the light of recent phylogenetic hypotheses. Here, we re-examine the non-belinurid taxa housed within the Chernyshev Central Museum for Geological Exploration in Saint Petersburg. We present the first formal diagnosis of Bellinuroopsis rossicus, erect Shpineviolimulus jakovlevi (Glushenko and Ivanov, 1961) comb. nov., to contain the species formerly described as ‘Paleolimulusjakovlevi and refer Paleolimulus juresanensis to Paleolimulidae incertae sedis. Phylogenetic analysis places S. jakovlevi at the base of Limulina. This position, coupled with a prosomal shield that is notably larger than the thoracetron, and lack of hypertrophied genal spines, suggests that this morphology may represent the ancestral austrolimulid shape. As an extension of this revision, we assessed the general austrolimulid morphological characters and uncovered two possible groups of these bizarre xiphosurids.

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Acknowledgements

We thank Tatiana Tolmacheva for access to the collection and help with the specimens. TNT is made freely available, thanks to a subsidy from the Willi Hennig Society. Finally, we thank two anonymous reviewers and the Julien Denayer (Associate Editor) for their suggested changes that clarified and re-directed the text, ultimately producing a much more informative manuscript.

Funding

This research was supported by funding from a Research Training Program Scholarship (to RDCB), a University of New England Postdoctoral Research Fellowship (to RDCB and SAB), a James R. Welch Scholarship (to RDCB), and a Betty Mayne Scientific Research Fund (to RDCB). This work was also funded by the State Program № 0135-2019-0044 of the Geological Institute of Russian Academy of Sciences (to SVN) and RFBR project No 18-04-00322 (to SVN).

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RDCB designed the study, gathered images, and wrote most of the main text. SAB conducted analyses and helped develop the Methods. SVN produced species reconstructions and wrote the Stratigraphic details section. All authors reviewed the final draft.

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Correspondence to Russell D. C. Bicknell.

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Communicated by: Julien Denayer

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ESM 1:

NEXUS file of the analysed phylogenetic matrix modified from Bicknell (2019). Originally presented in Lamsdell (2016) (NEX 75 kb)

ESM 2

Complete phylogenetic tree from strict consensus of the 7 most parsimonious trees. Node labels represent bootstrap, jackknife, and Bremer support values, respectively (PNG 198 kb)

High resolution image (EPS 5608 kb)

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Bicknell, R.D.C., Naugolnykh, S.V. & Birch, S.A. A reappraisal of Paleozoic horseshoe crabs from Russia and Ukraine. Sci Nat 107, 46 (2020). https://doi.org/10.1007/s00114-020-01701-1

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