Comparative actinopterygian fish taphonomy of laminated fossil Lagerstätten: case studies from the late Mesozoic of Eurasia

Language
en
Document Type
Doctoral Thesis
Issue Date
2016-11-28
Issue Year
2016
Authors
Chellouche, Patrick
Editor
Abstract

Laminated fossil Lagerstätten such as Plattenkalks offer a unique insight into ancient palaeo-biodiversity due to the exceptional quality of fossil preservation. Through the integrated use of qualitative, as well as quantitative taphonomical analyses of their fossil content, in combination with microfacial and geochemical investigations, the palaeoenvironmental conditions of such fossil Lagerstätten can be reconstructed. By comparing taphonomic features observed in various laminated fossil Lagerstätten, common palaeoenvironmental features which are a prerequisite for exceptional fossil preservation can be deduced. As a template for comparative taphonomy, the Upper Jurassic Wattendorf Plattenkalk of Southern Germany was chosen. Biostratinomic features of actinopterygian fish specimens, mainly of the genus Tharsis were recorded. Frequency of different grades of preservational features such as bending of the spinal column, skeletal articulation and overall completeness of the fish fossils was noted. By means of Euclidean cluster analysis, percentages of the occurrence of these features per bedding plane were grouped into clusters with similar patterns. Four different taphofacies W-A to W-D, representing various combinations of preservational features could be identified. These taphofacies indicate four energy regimes represented in the depositional environment of the Wattendorf Plattenkalk. The transition from taphofacies W-A to taphofacies W-D marks the development from only slight environmental disturbance to conditions of higher disturbance affecting the fish carcasses. At the beginning of Plattenkalk deposition, cyclic changes of the palaeoenvironment prevailed with periodic high disturbance, probably caused by storm-induced flows. In the upper part of the Plattenkalk unit, taphofacies indicative of higher disturbance dominate, suggesting a change from stable to less stable environmental conditions in the Plattenkalk basin resulting in cession of the typical Plattenkalk sedimentation. The Wattendorf Plattenkalk yielded three distinct modes of soft tissue preservation. In some fish fossils, connective tissues preserved as francolite can be observed. Soft-tissues were also preserved in acicular pyrite in a specimen of ?Palaeohirudo sp.. This modification requires slight amounts of oxygen. The third type of soft-tissue preservation is impregnation of internal voids in bones left by osteocytes and blood vessels by amorphic pyrite. Analysis of the micro- and ultrafacies of the Plattenkalk unit show rare layers of abundant coccoliths that co-occur with well-preserved fish fossils. This hints at occasional coccolithophorid blooms that might have caused fish kills. Abundant remnants of microbial mats had a positive and stabilising effect on fossil preservation. Changes in the water energy deduced from the taphonomic analysis are corroborated by microfacies analysis. Geochemical investigations of the Plattenkalk unit yielded additional palaeoenvironmental parameters for the genesis of Plattenkalks. An analysis of crystal size distribution of pyrite framboids showed anoxic conditions in the water column and in the sediment of the Wattendorf basin. Stable isotope ratios of calcitic skeletal elements of belemnites, brachiopods and bivalves found in the Plattenkalk unit show a clear thermal gradient in the water column of the Plattenkalk basin ranging from 9°C to 24°C. Combining the results of the various taphonomic, sedimentological and geochemical analyses, reconstruction of the palaeoenvironmental conditions influencing the sedimentation and early genesis of the Wattendorf Plattenkalk is possible. Based on the palaeoenvironmental information of the Wattendorf Plattenkalk gained by taphonomic analysis, these results were compared with the preservational features of actinopterygian fishes of two other laminated fossil Lagerstätten. Using quantitative taphonomy on fish specimens from the Upper Jurassic Nusplingen Plattenkalk lagerstätte of Southwestern Germany, again four different taphofacies could be established. Taphofacies N-A to taphofacies N-D show a gradual increase in extrinsic forces acting upon fish carcasses. Distribution of the taphofacies throughout the Plattenkalk succession shows only obscure cyclic changes. Low-energy taphofacies N-A to N-C prevail. High-energy taphofacies N-D, though, corresponds well with rarely occurring coarse sediment beds, colonisation horizons and scavenger activity and indicates an oxygenation of the whole water body. To test the method of quantitative taphonomic analysis on a continental depositional systems, actinopterygian fish specimens of the genus Lycoptera from the lacustrine Dawangzhangzi Bed of the Lower Cretaceous Yixian Formation of northwestern China were investigated. Euclidean cluster analysis established two taphofacies Y-A and Y-B. Taphofacies Y-A represents the lowest-energy conditions also observed in the other two fossil Lagerstätten. Taphofacies Y-B corresponds to the taphofacies W-B and N-B. Exceptionally good articulation was again likely caused by an abundance of microbial mats in a stagnant water body under inhospitable an anoxic conditions. Identification of mass mortality events of Lycoptera suggests that these conditions became on occasion widespread and even reached the uppermost part of the water column. Comparison of the preservational features of all three fossil Lagerstätten yielded common features required for exceptional fossil preservation. The presence of microbial mats is an important stabilising and encasing mechanism of carcasses. Inhospitable bottom waters caused by anoxic conditions exclude bioturbation and scavenging. The genesis of Plattenkalks is not restricted to tropical palaeoclimates. Slight wave activity and oxygen in the sediment and lower water column are not necessarily detrimental to exceptional fossil preservation and may even facilitate certain kinds of soft-tissue preservation.

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