Palaeogeographic evolution of the marine Middle Triassic marine Germanic Basin changements – With emphasis on the carbonate tidal flat and shallow marine habitats of reptiles in Central Pangaea

Abstract

More than seventy-five vertebrate track-sites have been found in Central Europe in 243–246.5 m.y. old Triassic coastal intertidal to sabkha carbonates. In the western part of the very flat Triassic intracontinental Germanic Basin, the carbonate strata contain at least 22 laterally extensive track horizons (called megatracksites). In contrast, in the eastern part of the basin only six megatracksites extended to near the centre of the Basin during marine low stands. Marine ingression and the development of extensive coastal marine environments began during the Aegean (Anisian) stage. This incursion began in the region of the eastern Carpathian and Silesian gates and spread westward due to the development of a tectonically controlled intracratonic basin. The tectonic origin of this basin made it susceptible to tsunamis and submarine earthquakes, which constituted very dangerous hazards for coastal terrestrial and even marine reptiles. The shallow sea that spread across the Germanic Basin produced extensive tidal flats that at times formed extensive inter-peninsular bridges between the Rhenish and Bohemian Massifs. The presence of these inter-peninsular bridges explains the observed distribution and movement of reptiles along coastal Europe and the northern Tethys Seaway during the Middle Triassic epoch. Two small reptiles, probably Macrocnemus and Hescherleria, left millions of tracks and trackways known as Rhynchosauroides and Procolophonichnium in the Middle Triassic coastal intertidal zone. The great abundance of their tracks indicates that their trackmakers Macrocnemus and Hescherleria were permanent inhabitants of this environment. In sharp contrast, tracks of other large terrestrial reptiles are quite rare along the coastal margins of the Germanic Basin, for example the recently discovered archaeosaur tracks and trackways referable to Isochirotherium, which most probably were made by the carnivore Ticinosuchus. Smaller medium-sized predatory thecodont reptiles, possibly Euparkeria, probably made the Brachychirotherium trackways that have been found across much of Central Europe. Large lepidosaurs such as Tanystrophaeus probably hunted in the tidal ponds and channels, where they locally produced Synaptichnium tracks. Recently discovered tracks made by a basal prosauropod are the world's oldest record of this group of dinosaurs, occurring in beds that have an age of about 243.5 Ma. (Pelsonian substage). This shows that very large prosauropods existed much earlier than was previously believed. These prosauropod tracks, along with tracks of small bipedal dinosaurs found in the Alps and Eastern France, show that by the middle part of the Middle Triassic the radiation and diversification of dinosaurs was already in progress. In the Germanic Basin, aquatic-adapted paraxial swimming sauropterygians are not known to have left tracks, except for occasional subaquatic swimming scratch-mark “trackways” within the coastal tidal flat zone. Marine-adapted aquatic reptiles migrated into the Germanic Basin with increasing frequency in the upper part of the Middle Triassic, when the bathymetry of the Germanic Basin was at its deepest following a strong regression that occurred due to basin uplift in the middle part of the Middle Triassic. These large marine reptiles included Pistosaurus, the ichthyosaurs Cymbospondylus or Mixosaurus, and many placodonts such as Cyamodus, Placodus and Paraplacodus, which fed on macroalgae and seem to have been the Triassic sea cows of their day. The distribution of these reptiles was mainly controlled by tectonics, but eustatic changes in sea level also were important and produced widespread environmental changes across the tidal flats up until their disappearance in the Germanic Basin in the late Middle Triassic. The initial break-up of Pangaea already had started in Middle Triassic time, and this event had begun to drastically change environments all over Central Europe. It is very interesting that dinosaurs began to diversify at exactly this time, and it seems likely that this was a direct reaction to these environmental changes. It can be inferred that the earliest dinosaurs must have started to evolve in the late Early Triassic, because in Europe it can be demonstrated that at least two main dinosaur groups already were present and clearly differentiated by the middle part of the Middle Triassic, and all three of the major groups of dinosaurs (theropods, sauropods and ornithischians) had diversified and spread globally throughout terrestrial habitats by the end of the Triassic Period. Six new palaeogeographic maps, representing time intervals from the Aegean to the Illyrian (Anisian) stages, show these important environmental changes in detail and explain the direction and timing of terrestrial reptile exchanges between the Central Massif, Rhenish Massif, and Bohemian Massif, and also the direction and timing of marine reptile exchanges between the Alps of Central Pangaea and the ancient northern Tethys Ocean and Germanic Basin Sea.