Article: A hexapod from the early Devonian Windyfield Chert, Rhynie, Scotland

New diagnostic morphological features discovered following a re-examination and further preparation of the holotype of Leverhulmia mariae Anderson and Trewin from the Early Devonian (Pragian) Windyfield chert are described and discussed here. The discovery of anterior (thoracic?) leg appendages with attached pretarsal elements comprising a fixed median unguis, a pair of articulated lateral ungues and possible unguitractor plate has allowed us to reclassify this enigmatic fossil as Hexapoda incertae sedis. We interpret the holotype as part of an abdomen bearing at least five pairs of segmented leglets. The condition of the ungues on the anterior appendages is strongly reminiscent of Archaeognatha and Zygentoma (= Thysanura s.s.), and the segmented leglets on the abdomen show similarities with certain fossil 'apterygotes' known from the Carboniferous. Leverhulmia is the third hexapod species found in the cherts at Rhynie, the other forms being the collembolan Rhyniella praecursor Hirst and Maulik, and the pterygote insect Rhyniognatha hirsti Tillyard. Showing closer affinities with Archaeognatha and Zygentoma, Leverhulmia may well represent the earliest fossil apterous insect known to date.