Cyrtoctenus wittebergensissp. nov. (Chelicerata: Eurypterida), a large sweep-feeder from the Carboniferous of South Africa

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Abstract
Cyrtoctenus wittebergensissp. nov. is described from a unique holotype from the Witteberg Group of the Cape Supergroup. It is a giant hibbertopteroid eurypterid having combs and specialised movable spines of crytoctenid type (Størmer & Waterston 1968) on the more distal podomeres of the second to fourth prosomal appendages. The function of the combs and their associated movable spines is discussed and it is suggested that together they formed a unique adaptation of eurypterid structures to sweep filter-feeding, the combs forming the filters and the spines the cleaners. The digestive tract is remarkably preserved and shows a spiral valve, posterior to the stomach, which is interpreted as an adaptive feature in this large arthropod to increase the absorptive area of the gut.The new evidence provided by the South African specimen has required the re-interpretation of the disarticulatedCyrtoctenusspecimens previously described from Europe.Disjecta membrarecently obtained from the Tournaisian of Foulden, Berwickshire, which may belong toCyrtoctenus, are described and show characters previously unknown in Scottish material but similar to certain features in the South African specimen.The taxonomic relationships within the Hibbertopteroidea are discussed in the light of the new combination of characters found inC. wittebergensis. Two families are recognised in the superfamily, the Hibbertopteridae, includingHibbertopterusandCampylocephalus, and the new family Cyrtoctenidae which is here erected to includeCyrtoctenus, Dunsopterusand possibly alsoHastimima.

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