Palaeophysiology of terrestrialisation in the Chelicerata

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Abstract
The wide range of organs of respiration (book-gills, book-lungs, sieve- and tube-tracheae), reproduction, sensory perception, etc., among the chelicerates indicates that the major groups made the transition to land life independently. The fossil record is patchy for most chelicerate groups, certain intervals (e.g. Westphalian) being particularly rich in chelicerate bearing Lagerstatten while in others (e.g. Mesozoic) they are sparse. Due, apparently, to their unusual hyaline exocuticle, scorpions are better preserved than other arthropods, and show a fairly continuous record from fully aquatic forms in the Silurian, to both aquatic and terrestrial faunas in the Carboniferous. In particular, new and well-preserved material of the earliest demonstrably terrestrial scorpions from the Lower Carboniferous of East Kirkton, West Lothian, suggests that book-lungs, at least in the scorpions, developed directly from book-gills by suturing of the covering plate (Blattfuss of the related eurypterids) to leave stigmata for diffusion of air. This evidence supports the ideas of early authors that the scorpion mesosomal ‘sternites’ are fused plates, contra Kjellesvig-Waering (1986) who envisaged the plates being lost to reveal true sternites beneath. The fossil evidence also indicates that by the Triassic at least two scorpion lineages had evolved intra-‘sternite’ stigmata.

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