Diedrich, C. G. 2011. Middle Triassic horseshoe crab reproduction areas on intertidal flats of Europe with evidence of predation by archosaurs. Biological Journal of the Linnean Society103: 76–105. doi: 10.1111/j.1095-8312.2011.01635.x
Abstract - A systematically excavated track site in a 243.5 Myr old Middle Triassic (Karlstadt Formation, Pelsonian, middle Anisian) intertidal carbonate mud-flat palaeoenvironment at Bernburg (Saxony-Anhalt, central Germany) has revealed extensive horseshoe crab trackways attributable to the Kouphichnium Nopsca, 1923 ichnogenus. The exposed track bed of a Germanic Basin-wide spanned intertidal megatrack site is a mud-cracked biolaminate surface on which detailed tracks have been preserved because of rapid drying and cementation as a result of high temperatures, followed by rapid covering with a protective layer of arenitic storm or tsunami sediments. The different trackway types and their orientations have allowed a tidal sequence to be reconstructed, with the initial appearance of swimming horseshoe crabs followed by half-swimming/half-hopping limulids under the shallowest water conditions. The Bernburg trackways, which have mapped lengths of up to 40 m, were all produced by adult animals and exhibit a variety of shapes and patterns that reflect a range of subaquatic locomotion behaviour more typical of mating than of feeding activities. The closest match to the proportions and dimensions of the horseshoe crab tracks at Bernburg is provided by the largest known Middle Triassic limulid Tachypleus gadeai, which is known from the north-western Tethys in Spain. The horseshoe crab body fossils recognized in the German Mesozoic intertidal zones, instead, are from juveniles. The uniformly adult size indicated by the trackways therefore suggests that they may record the oldest intertidal reproductive zones of horseshoe crabs known from anywhere in the world, with the track-makers having possibly migrated thousands of kilometres from shallow marine areas of the north-western Tethys to reproduce in the intertidal palaeoenvironments of the Germanic Basin. Chirotherium trackways of large thecodont archosaurs also appeared on these flats where they appear to have fed on the limulids. With the tidal ebb, smaller reptiles such as Macrocnemus (Rhynchosauroides trackways) appeared on the dry intertidal flats, probably feeding on marine organisms and possibly also on horseshoe crab eggs.
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