Spectacular trilobite Lagerstätten occur in distinctive offshore calcareous mudstone facies through the Late Ordovician to Devonian, and reflect a combination of mass mortality or molting and burial, coupled with early diagenetic enhancement. Evidence indicates two distinct modes of burial, Type I and II assemblages, which show evidence for burial without or with seafloor disturbance, respectively. Type I assemblages suggest rapid (hours to days), but not instantaneous burial, without bottom disruption, enabling preservation of in situ behavior, including mass aggregations and molt ensembles. Most occurrences contain bedding planes in which trilobites exhibit incipient disarticulation. These assemblages were buried by cascades of flocculated sediment from hypopycnal, detached flows. Type II assemblages show well-articulated, enrolled, semi-enrolled, and outstretched trilobites in varied orientations relative to bedding. In such cases, bottom flows and seafloor disruption by storm or seismic disturbances in shallow waters suspended large amounts of flocculated muds as viscous slurries, which developed into hyperpycnal flows that entrained carcasses of trilobites and other organisms. In many cases, both Type I and II obrution was followed by additional sedimentation, geochemical zones moved upward through the sediment column, and there was little tendency to form diagenetic overprints. Alternatively, if burial was followed by an interval of sediment starvation, the sediments were bioturbated and very early diagenetic mineralization was superimposed, first, in rare cases, as mineralized soft parts in entombed carcasses, and later as pyritization of burrow linings. Development of the concretionary layers required more prolonged periods of stability of the sulfate reduction zone. Cementation of sediment shielded organism bodies from most or all effects of compaction. Thus, ironically, the best preservation of delicate remains required rapid burial, associated with mass mortality, and very low rates of background sedimentation following the event.
Determining whether morphological trends in fossil species represent evolution within a lineage or lateral shifts in morphologically variable populations through time requires a thorough examination of the details of both morphology and paleoenvironment in time and space. The purpose of this study is to explore at high resolution the relationship between morphology of the trilobite Flexicalymene granulosa and paleoenvironmental conditions in Upper Ordovician deposits of southwestern Ohio and northernKentucky. This is achieved by using geometric morphometrics to measure high-resolution morphological changes and by using gradient analysis to capture environmental gradients underlying faunal distribution patterns. Quantitatively comparing the outcomes of these two techniques provides an assessment of whether shape changes relates to environment. Results indicate that a significant amount of shape change, seen as an anteromedial movement of the eye region over time, corresponds to ordination scores. This suggests a relationship between certain aspects of morphology and environment. The combination of these quantitative techniques has provided the foundation for determining whether morphological trends within F. granulosa represent evolution or temporal shifts in geographic variation. Future work will involve examining this relationship in greater detail both geographically and stratigraphically.
Flexicalymene retrorsa minuens from the uppermost 3 m of the Waynesville Formation of the Cincinnatian Series (Upper Ordovician) of North America lived approximately 445 Ma and exhibited marked reduction in maximum size relative to its stratigraphically subjacent sister subspecies, Flexicalymene retrorsa retrorsa. Phylogenetic analysis is consistent with the notion that F. retrorsa retrorsa was the ancestor of F. retrorsa minuens. F. retrorsa minuens has been claimed to differ from F. retrorsa retrorsa"in size alone," and thus presents a plausible example of global paedomorphic evolution in trilobites. Despite strong similarity in the overall form of the two subspecies, F. retrorsa minuens is neither a dwarf nor a simple progenetic descendant of F. retrorsa retrorsa. More complex patterns of global heterochronic paedomorphosis, such as a neotonic decrease in the rate of progress along a common ontogenetic trajectory with respect to size, coupled with growth cessation at a small size, "sequential" progenesis, or non-uniform changes in the rate of progress along a shared ontogenetic trajectory with respect to size, can also be rejected. Rather, differences between these subspecies are more consistent with localized changes in rates of character development than with a global heterochronic modification of the ancestral ontogeny. The evolution of F. retrorsa minuens from F. retrorsa retrorsa was largely dominated by modifications of the development of characters already evident in the ancestral ontogeny, not by the origin of novel structures. Factors promoting size reduction in F. retrorsa minuens appear to have been specific to this subspecies, because other co-occurring taxa, including other trilobite species, do not show marked differences in mean size.
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