Ankylosaurs, like probably most other dinosaurs, were landlubbing, terrestrial animals without obvious aquatic adaptations. And yet, surprisingly, their fossils are found in marine sedimentary environments more often than most other dinosaurs (except hadrosaurs). Some, like Aletopelta, wound up in shallow or lagoonal environments - Aletopelta's carcass became a reef! - but some, like the Suncor nodosaurid, wound up far away from shore.
TAPHONOMY OF A MONODOMINANT CENTROSAURUS APERTUS (DINOSAURIA: CERATOPSIA) BONEBED FROM THE UPPER OLDMAN FORMATION OF SOUTHEASTERN ALBERTA
Authors:
Chiba et al
Abstract:
The horned dinosaur Centrosaurus apertus from the Belly River Group (Campanian) is represented by multiple articulated skulls and skeletons, and is particularly notable for its occurrence in dozens of large-scale monodominant bonebeds, which have been found in the Dinosaur Park Formation across southern Alberta and Saskatchewan. Here we present a detailed taphonomic analysis of the first large-scale Centrosaurus apertus bonebed (McPheeters bonebed) from the Oldman Formation of southeastern Alberta. The McPheeters bonebed rivals the richest bonebeds in the Dinosaur Park Formation in terms of bone density and size, and the complete disarticulation of elements. The bonebed occurs in an overbank facies and is dominated by small bone clasts, suggesting that only low energy water current contributed to the formation of the bonebed before its final burial event. Patterns of taphonomic modification suggest that bones experienced little weathering, breakage, or scavenging. In turn, these conclusions are compatible with an overall interpretation of rapid burial in humid conditions after the disarticulation of elements. These taphonomic features are virtually identical to those seen in the well-documented bonebeds of this species in the Dinosaur Park Formation, which are interpreted to represent mass death events caused by seasonal tropical storms and associated large-scale flooding. Late Cretaceous dinosaur species typically have small geographic and stratigraphic ranges defined by the extent of single geological formations. The new bonebed extends the distribution of Centrosaurus apertus to the upper Oldman Formation, which is interpreted as more inland than the coastally influenced Dinosaur Park Formation, and suggests that mass death events related to seasonal tropical storms occurred over a broader geographic area and in a greater range of paleoenvironments than previously documented.
Analysis of marrow cavity fillings as a tool to recognise diverse taphonomic histories of fossil reptile bones: Implications for the genesis of the Lower Muschelkalk marine bone-bearing bed (Middle Triassic, Żyglin, S Poland)
Author:
Kowal-Linka
Abstract:
The bone-bearing bed from Żyglin (S Poland), which likely represents the oldest Lower Muschelkalk accumulation of reptile remains in the SE part of the Germanic Basin, has been investigated in order to determine its genesis. The basic methods were supported by petrographic analysis of the marrow cavity and large inter-trabecular pore space fillings of fossil bones, which was used to check its usefulness to identify the environments where the bones were initially deposited and to decipher taphonomic histories of the remains. The bone-bearing bed is a composite deposit, which consists of three distinct layers (from bottom to top): micritic limestone (mudstone, bioturbated autochthonous mud), crinoidal limestone (grainstone to packstone, calcirudite), and shell-rich limestone (packstone to wackstone, calcirudite). The crinoidal limestone layer, the main bone-bearing bed, is recognised as the proximal tempestite deposited in the mid-ramp zone. The petrographic analysis of the fillings reveals the prevalence of minute ostracod carapaces, accompanied by other grains, embedded together with micrite to microspar. Such compositions suggest that these sediments may have been inserted into the bone pore spaces in lagoons and tidal-flat ponds. Features of some bones record their early diagenetic burial and lithification before the final redeposition. The isopachous spar, blocky spar and weathered pyrite document changes in the chemical composition of fluids that flowed into the bone interiors. The burrow found in the marrow cavity of specimen IGUAM-ZOV-7 provides evidence that some remains were inhabited by minute or juvenile invertebrates. All recognised features indicate that both invertebrate and vertebrate bioclasts, included in the tempestite, were initially deposited in various settings of the carbonate ramp and in the end redeposited as a result of a heavy storm or a hurricane. The examined bone-bearing bed represents time-averaged assemblage, which originated due to hydraulic concentration of the vertebrate bioclasts. The petrographic analysis of the fillings is a valuable tool to identify complex taphonomic pathways of vertebrate bioclasts.
Rocky shore taphonomy—A comparative study of modern and Late Cretaceous analogues
Authors:
Sørensen et al
Abstract:
Rocky shores are rare in the fossil record due to erosion under both sea-level rise and fall. In contrast, modern rocky shores are well-studied, but little is known about the evolution of their ecosystems due to the rarity of ancient counterparts. Reconstruction of these ancient ecosystems is thus essential to get an insight into their evolution. A high-diversity Late Cretaceous (Campanian) rocky shore fauna is found in southern Sweden. The original composition of the shelly fauna cannot be interpreted by direct examination of the preserved fauna due to the effects of taphonomic processes. Life and death assemblages from a modern rocky shore fauna from Thailand have previously been analysed and a hypothetical fossil assemblage was reconstructed in order to attempt an interpretation of the Campanian life assemblage. This study shows a low taxonomic agreement between the original Campanian life assemblage and the fossil assemblage, due to taphonomic processes, and high environmental fidelity with only a few out-of-habitat species represented. The modern life assemblage showed in an earlier study, a high loss of species before onset of fossilisation. This suggests that the faunal composition of the Campanian life assemblage cannot be easily reconstructed, and time averaging by generations of death assemblages makes this even more difficult. The Campanian aragonitic fauna is poorly represented and the rarity of moulds after aragonitic species is interpreted as due to taphonomic processes and not to lower richness of aragonitic species in the Cretaceous. This is supported by comparison with the high richness of aragonitic species found on a Late Cretaceous rocky shore in Germany. An originally high-diversity gastropod fauna is thus interpreted to have dominated the intertidal zone in the Campanian example, and the rare moulds of each of the aragonitic species indicate a high taphonomic loss in spite of rapid burial. Calcitic species-richness is higher in the Campanian fauna than in the modern life, death, and constructed hypothetical fossil assemblages. This is interpreted as reflecting time averaging of generations of calcitic species and low loss of calcitic species by taphonomic processes in the Campanian fauna. It is thus assumed that the original Campanian fauna experienced a change in faunal composition from a gastropod-dominated life assemblage to a bivalve-dominated fossil assemblage due to dissolution of aragonite and excellent preservation of calcite. Reconstruction of ancient rocky shore shelly faunas can thus be considerably improved by comparison with analogous modern rocky shore faunas.
Dickinsonia lift off: Evidence of current derived morphologies
Authors:
Evans et al
Abstract:
Dickinsonia, an iconic member of the Ediacara biota, is abundant in the Ediacaran deposits found at the Nilpena field site, South Australia. Despite exquisite fossil preservation at this site, many specimens of Dickinsonia appear to be incomplete, with an apparently “missing piece” on the periphery. Orientation measurements from specimens on three fossil beds suggest that these so-called “missing pieces” are aligned irrespective of the axial orientation of Dickinsonia. The nonrandom orientation of incomplete specimens matches that of other aligned structures found on two of these beds. The preferred directionality of this feature suggests the molding of incomplete specimens under the influence of current activity prior to or during burial. We propose that this feature originates where part of a Dickinsonia was lifted off of the substrate during a storm event and that sand was deposited beneath this lifted portion. This model suggests that Dickinsonia was easily separated from the sea floor and was not attached to the substrate on which it lived. This is consistent with the data from Dickinsonia footprints suggesting that Dickinsonia was mobile.
A MODEL FOR ORGANIC FOSSILIZATION OF THE EARLY CRETACEOUS JEHOL LAGERSTÄTTE BASED ON THE TAPHONOMY OF “EPHEMEROPSIS TRISETALIS”
Authors:
Pan et al
Abstract:
The taphonomic pathways of “Ephemeropsis trisetalis” nymphs (mayfly larvae) were systematically investigated based on fossils of different preservational types, collected during three high–stratigraphic-resolution (mm to cm) excavations in the Lower Cretaceous Yixian Formation in the Sihetun area of western Liaoning, China. All fossils studied are fully articulated either in three or two dimensions, which indicates that decay was terminated at a stage before the exoskeleton became disarticulated. We conclude that the Jehol organic skeletons represent at least two general types of preservation produced by pyritization and collapse/compression, respectively. The two-dimensional compressions show no evidence for authigenic minerals, but the three-dimensionally preserved fossils are wholly or partially pyritized. Our study also indicates that aluminosilicate clay and pyrite mineralization are closely associated with fossil “Ephemeropsis trisetalis” nymphs, suggesting that both clay and pyrite played important roles in lacustrine fossil preservations, as in some marine fossil Lagerstätten. We propose a general model for organic tissue fossilization in the Jehol Lagerstätte based on study of taphonomy of “Ephemeropsis trisetalis” nymphs.
TAPHONOMY AND DEPOSITIONAL SETTING OF THE BURGESS SHALE TULIP BEDS, MOUNT STEPHEN, BRITISH COLUMBIA
Authors:
O'Brien et al
Abstract:
Burgess Shale–type deposits represent exceptional preservational windows for examining the biodiversity and ecological structure of some of the earliest metazoan communities that evolved during the Cambrian Explosion. While much attention has been paid to the original Burgess Shale locality, the Walcott Quarry on Fossil Ridge, temporal and regional variations of the depositional environment of the Burgess Shale biota as a whole are still poorly understood. Here we present the first comprehensive taphonomic and sedimentological study of the Tulip Beds on Mount Stephen (Campsite Cliff Shale Member, Burgess Shale Formation), based on a time-averaged assemblage of nearly 10,000 specimens. The taphonomic characteristics—size sorting, resistance to decay, and potential flow alignment—and mode of deposition of this assemblage are compared specifically to those of the nearby and stratigraphically younger Walcott Quarry assemblage. Like other Burgess Shale–type deposits, the Tulip Beds consist of millimeter-laminated, event-derived claystone, but lack the thicker claystone layers and prominent carbonate interbeds that occur in the Walcott Quarry. These differences suggest a depositional environment lower in energy and possibly more distal to the Cathedral Escarpment. Overall, taphonomic analyses suggest no significant decay biases, transport, or sorting of the assemblage, and most specimens, benthic taxa in particular, appear to have been buried close to their living environments. Single bedding planes with large accumulations dominated by a single taxon, e.g., isolated claws of Anomalocaris, suggest short time-averaged assemblages with limited background sedimentation. Overall the Tulip Beds locality is environmentally and taphonomically comparable to the Walcott Quarry and biotic variations between the two sites are likely to be primary in nature, thus paving the way for more detailed paleoecological investigations in the future.
Integrated taphonomy in an open-marine platform: The Lower Cretaceous of Sierra Helada (Betic Cordillera, SE Spain)
Authors:
Giannetti et al
Abstract:
A detailed taphonomic analysis was carried out on the lower Albian deposits of the Sierra Helada section (Alicante, Betic Cordillera, southeastern Spain). Ten taphonomic characters were studied and ten skeletal concentrations were defined on the basis of taphonomic features and the dominant taxa. Cluster analysis was performed on the dataset represented by the abundance of the taphonomic characters in each skeletal concentration. This enabled the definition of four different taphonomic categories: 1) skeletal concentrations characterized by the presence of fossils preserved in life position, 2) skeletal concentrations showing very little physical reworking, 3) skeletal concentrations related to high-energy background conditions, and 4) skeletal concentrations produced by medium- to high-energy events.
Four taphofacies were defined on the basis of the main sedimentological features and the most representative skeletal concentrations. Taphofacies A represents the low energy outer platform, rich in skeletal concentrations with echinoids in life position and only slightly reworked. The second taphofacies (taphofacies B) is very rich in reworked echinoid tests and calcarenitic beds and records the transition to shallower areas, while taphofacies C shows abundant thick-bedded calcarenites and skeletal concentrations produced by sediment transport and rapid deposition. Finally, cross-bedded grainstone beds, which are rich in fine-grained fragmented, locally reoriented bioclasts (taphofacies D), record the existence of shifting sandy dunes in the shallow inner part of the platform.
Taphonomy of the Ediacaran Fossil Pteridinium Simplex Preserved Three-Dimensionally in Mass Flow Deposits, Nama Group, Namibia
Authors:
Meyer et al
Abstract:
Ediacara-type fossils are found in a diverse array of preservational styles, implying that multiple taphonomic mechanisms might have been responsible for their preservational expression. For many Ediacara fossils, the “death mask” model has been invoked as the primary taphonomic pathway. The key to this preservational regime is the replication or sealing of sediments around the degrading organisms by microbially induced precipitation of authigenic pyrite, leading toward fossil preservation along bedding planes. Nama-style preservation, on the other hand, captures Ediacaran organisms as molds and three-dimensional casts within coarse-grained mass flow beds, and has been previously regarded as showing little or no evidence of a microbial preservational influence. To further understand these two seemingly distinct taphonomic pathways, we investigated the three-dimensionally preserved Ediacaran fossil Pteridinium simplex from mass flow deposits of the upper Kliphoek Member, Dabis Formation, Kuibis Subgroup, southern Namibia. Our analysis, using a combination of petrographic and micro-analytical methods, shows that Pteridinium simplex vanes are replicated with minor pyrite, but are most often represented by open voids that can be filled with secondary carbonate material; clay minerals are also found in association with the vanes, but their origin remains unresolved. The scarcity of pyrite and the development of voids are likely related to oxidative weathering and it is possible that microbial activities and authigenic pyrite may have contributed to the preservation of Pteridinium simplex; however, any microbes growing on P. simplex vanes within mass flow deposits were unlikely to have formed thick mats as envisioned in the death mask model. Differential weathering of replicating minerals and precipitation of secondary minerals greatly facilitate fossil collection and morphological characterization by allowing Pteridinium simplex vanes to be parted from the massive hosting sandstone.
Taphonomic traits of clay-hosted early Cambrian Burgess Shale-type fossil Lagerstätten in South China
Authors:
Forchielli et al
Abstract:
The Chengjiang and Guanshan fossil Lagerstätten of Yunnan represent key fossil deposits to understand and reconstruct early metazoan evolution. Despite extensive studies on the systematics of taxa from these Lagerstätten, the major mechanisms of exceptional soft-tissue preservation at these fossil Lagerstätten have remained controversial. The Chengjiang and Guanshan fossil Lagerstätten represent typical Burgess Shale-type deposits, influenced by late alteration and weathering that led to some major transformations in clay and iron minerals, as well as dissolution of dolomite and skeletal biominerals. The detailed geochemical screening of various fossil groups, such as brachiopods, priapulids, arthropods, and sponges, indicates a more complex preservational history than hitherto supposed. Besides oxidation processes during sub-recent/recent weathering, clay mineral formation and/or alteration, carbonization, phosphatization, and pyritization played key roles in the preservation of various metazoans. The Chengjiang fossil Lagerstätte is characterized by the existence of clay-dominated background beds (BGBs) and event beds (EBs). Fossil preservation varies considerably in the two different depositional settings, with soft-tissue preservation occurring preferentially in EBs, whereas skeletal accumulations are typical of BGBs.
The samples were analyzed and classified into three groups with different degrees of alteration and fundamentally different chemistries. The specific clay mineral composition shows increasing chlorite and goethite, decreasing kaolinite, 2M1-mica, and dolomite, and a rise in the chemical alteration of mudstones. It also indicates slight variations between the primary types of beds (EB, BGB), hinting at variations in the source of clastics or the hydrochemical environment during the deposition of strata. Furthermore, the geochemical paleoredox proxy data indicate that dysoxic or anoxic conditions were not permanently present during the deposition of EBs and BGBs in the Chengjiang and Guanshan fossil Lagerstätten.
We consider that rapid deposition in finest claystones had a greater influence on the preservation of highly volatile tissues than the presence of bottom-water anoxia, even though bottom-water anoxia and microbial sealing may further increase the probability of soft-tissue preservation. Using different geochemical proxies such as various redox indicators and iron speciation, statistics on framboid sizes, classical geological observation, and clay mineralogy, we show that iron mineral precipitation onto organic carcasses of the Chengjiang-type fossils is mostly confined to later diagenetic processes and thus is not crucial for the early fixation of easily decayable tissues in Burgess Shale-type faunas.
Experimental tumbling of echinoderms – taphonomic patterns and implications
Authors:
1. Przemysław Gorzelak (a)
2, Mariusz A. Salamon (b)
Affiliations:
a. Department of Biogeology, Institute of Paleobiology, Polish Academy of Sciences, Twarda Str. 51/55, 00–818 Warsaw, Poland
b. University of Silesia, Faculty of Earth Sciences, Department of Palaeontology and Biostratigraphy, Będzińska Str. 60, 41–200 Sosnowiec, Poland
Abstract:
Despite a wide array of published actualistic studies on echinoderm taphonomy the detailed pattern of decay and disarticulation of their skeletons is still not well understood. Here we provide results of tumbling experiments using a rotating barrel filled with artificial seawater and medium-sized quartz sand to mimic physical forces experienced by echinoderms during trasportation in high-energy conditions. In particular, we determined semi-quantitatively transportation-induced rates and patterns of damage and disintegration of freshly killed ophiuroid, asteroid and crinoid skeletons that were not allowed to decay initially. Our experiments showed that echinoderm specimens disintegrated in a characteristic sequence toward an increase of the degree of disarticulation, abrasion and roundness or thinness of echinoderm ossicles. The sequence of disintegration in crinoids began with the partial disintegration of distal arms after 2 hrs (a time equivalent to ~ 1 km of transport). The initial split of ophiuroid and asteroid arms and crinoid cirri occurred after 24 hrs (~ 12 km) and complete destruction of the asteroid mouth and ophiuroid disc area occurred after 72 hrs (~ 36 km). The duration of transport necessary to promote initial fragmentation in asteroid and ophiuroid arms and crinoid cirri into isolated ossicles was 120 hrs (~ 60 km). The complete disarticulation of crinoid, ophiuroid and asteroid arms and crinoid cirri occurred after 312 hrs (~ 156 km) and 408 hrs (~ 204 km), respectively. Although it has been argued that the quality of preservation can be a poor index of post-mortem transport, echinoderms allowed limited initial decay in the presence of rapid and relatively constant physical disturbance, an approximation of the distance of transport can be made.
Our data demonstrate that articulated ossicles can remain for several days, sufficient time for long (even a few hundred km) transporation. This finding illustrates that articulated echinoderm remains do not necessarily imply low energy and highlights the importance of a reliable discrimination of autochthonous and allochthonous components of fossil echinoderm assemblages. Application of isolated fossil echinoderm ossicles in e.g. paleoenvironmental and paleoecological reconstructions may lead to serious misinterpretations and should be supplemented by observations of abrasion traces.
Explaining the exceptional preservation of Ediacaran rangeomorphs from Spaniard's Bay, Newfoundland: A hydraulic model
Authors:
1. Martin D. Brasier (a, b)
2. Alexander G. Liu (c)
3. Latha Menon (a)
4. Jack J. Matthews (a)
5. Duncan McIlroy (b)
6. David Wacey (d, e)
Affiliations:
a. Department of Earth Sciences, University of Oxford, South Parks Road, Oxford OX1 3AN, UK
b. Department of Earth Sciences, Memorial University of Newfoundland, Prince Philip Drive, St John's, NL, A1B 3X5 Canada
c. Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EQ, UK
d. Department of Earth Sciences & Centre for Geobiology, Allegaten 41, University of Bergen, N-5007 Bergen, Norway
e. Australian Research Council Centre of Excellence for Core to Crust Fluid Systems, Centre for Microscopy Characterisation and Analysis & Centre for Exploration Targeting, The University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia
Abstract:
Exceptional 3-D preservation of Ediacaran rangeomorph fossils is found on a single bedding plane at Spaniard's Bay, Newfoundland. This high-quality preservation has previously been explained by entrainment of organisms within the Td-e mudstone division of a distal turbidite, followed by encasement within concretions. Our sedimentological and taphonomic analysis reveals a clear association between these fossils and evidence for erosive unidirectional flows, including scours marks, tool marks, ridge-and-groove marks, parting lineations and current crescents. We suggest an alternative sequence of events that runs broadly as follows: (i) rangeomorph discs were anchored to the seafloor during deposition of planar laminated silts (our unit 2, less than 10 mm thick; Td), now bearing pyrite framboids and pyritized organic matter; (ii) rangeomorph fronds were then felled and entrained by high velocity unidirectional currents, to lie within their own erosional scours at the top of unit 2, or to form tool marks; (iii) this topography was then draped and cast by soft-weathering sand (unit 3, Tc) associated with the growth of early diagenetic pyrite around sand grains. Pyrite grains also appear to have replaced clumps of organic matter. Fossil impressions have since been exposed by differential weathering of the ferruginous sands with respect to the silts. This new context now provides a parsimonious explanation for a range of hitherto paradoxical structures. Features previously regarded as microbial mats (‘bubble trains’) that formed in the lee of sinuous ripples on the top of unit 2 may be explained as load-casts, or by localised gas escape within areas of lowered hydraulic pressure. Rangeomorph fronds remarkably preserved in positive (rather than the more usual negative) epirelief are explained by means of sediment-casting of branches that became ruptured in the high velocity current. Paradoxical structures previously thought to be enclosing biological ‘sheaths’ around rangeomorph fronds are reinterpreted as scour marks, whereas imbricate overlaps of first order branches in Beothukis, Trepassia and Avalofractus are explained by hydraulic shear, driven by overlying currents across ruptured and deflated fronds. We find that rangeomorph bodies could be deflated, imbricated, folded over, inverted, and infilled with fine sediment. Our hydraulic model provides a null hypothesis against which future observations of rangeomorph fronds can now be tested. It removes some significant anomalies in our understanding of rangeomorph architecture, and provides a better understanding of the physical properties of their body tissues, permitting the possibility of a reasoned consideration of their puzzling biological affinities.
