Charles E. Mitchell

A comparison of crayfish burrow morphologies: Triassic and Holocene fossil, paleo‐ and neo‐ichnological evidence, and the identification of their burrowing signatures

The architectural and surficial morphologies of crayfish burrows from the Upper Triassic Chinle Formation and Holocene sediments were compared in order to determine: 1) if Triassic burrows could truly be attributed to crayfish activity; 2) how comparable the burrowing mechanisms are; and 3) whether or not a common set of burrowing signatures could be identified for both ancient and modern freshwater crayfish. Materials used in this study include burrows from the members of the Upper Triassic Chinle Formation, casts of modern burrows constructed by Procambarus clarkii Hobbs and Procambarus acutus acutus (Girard) in the laboratory, and casts of naturally constructed modern burrows of Cambarus diogenes di‐ogenes (Girard). Triassic and Holocene crayfish burrow morphologies exhibit simple to complex architectures, varying degrees of branching, chamber, and chimney development. They also exhibit relatively textured surficial morphologies (bioglyphs) such as scrape and scratch marks, mud‐ and lag‐liners, knobby ...

Late Ordovician to earliest Silurian graptolite and brachiopod biozonation from the Yangtze region, South China, with a global correlation

Late Ordovician to earliest Silurian is an important geological period marked by large geological and biological events. However, the strata and fossils of this interval are not complete in many parts of the world. Based on studies of 43 sites in South China, in particular the continuous sections on the Yangtze platform, we recognize a complete succession including seven graptolite zones and two shelly faunas. In ascending order, the graptolite zones are the Dicellograptus complanatus , Dicellograptus complexus , Paraorthograptus pacificus (including Lower Subzone, Tangyagraptus typicus Subzone and Diceratograptus mirus Subzone), Normalograptus extraordinarius–Normalograptus ojsuensis , Normalograptus persculptus , Akidograptus ascensus and Parakidograptus acuminatus zones. The shelly faunas are the Foliomena–Nankinolithus and Hirnantia faunas, which may be correlated with D. complanatus Zone and N. extraordinarius–N. ojsuensis to part of N. persculptus zones respectively. The biozonation through this interval from the Yangtze region can be correlated with that of other parts of the world such as Dob’s Linn in Scotland, Spain and Portugal, Thuringia–Saxonia–Bavaria, Bohemia, Poland, Kazakhstan, Kolyma, Malaya Peninsula, Yukon, Canadian Arctic Islands, Nevada, Argentina, Niger and Victoria, Australia. The Hirnantian Substage, which has been proposed by us recently, includes the N. extraordinarius–N. ojsuensis Zone, Hirnantia fauna and N. persculptus Zone. The base of the Hirnantian Substage is marked by the First Appearance Data (FADs) of N. extraordinarius and N. ojsuensis , which have been determined to be synchronous on a global scale.

Environmental changes in the Late Ordovician–early Silurian: Review and new insights from black shales and nitrogen isotopes

The Late Ordovician (Katian-Hirnantian) through earliest Silurian (Rhuddanian) interval was a time of varying climate and sea level, marked by a peak glacial episode in the early-mid-Hirnantian. Synthesis of recently published data permits global correlation of at least two cycles of glacial advance and retreat with a distinct interglacial period that is recognizable in sequence-stratigraphic and chemostratigraphic records in many parts of the world. A period of warming and sea-level rise during the late Katian is marked by the widespread occurrences of oceanic anoxia in paleotropical and subtropical localities, mostly confined to regions of inferred upwelling and semirestricted marine basins. Nitrogen isotope data show that the regions of oceanic anoxia were marked by intense water-column denitrification in which cyanobacteria were the principal source of fixed N. In the overlying peak glacial interval of the Hirnantian, sedimentary successions from localities representing a wide range of water depths and paleolatitudes indicate that anoxia was restricted during the early-mid-Hirnantian. The shift to more positive N isotope values also suggests less intense water-column denitrification. In the overlying late Hirnantian and early Rhuddanian, the distribution of black shales reaches its greatest extent in the studied interval. Localities showing evidence of anoxia are globally spread over all paleolatitudes and water depths for which data are available, indicating a Rhuddanian ocean anoxic event comparable to examples from the Mesozoic. It is accompanied by a return to intensely denitrifying conditions within the water column, as indicated by the shift to negative N isotope values. The two phases of Hirnantian mass extinction coincide with rapid, climate-driven changes in oceanic anoxia. The first extinction occurred at the onset of glaciation and with the loss of anoxic conditions at the end of the Katian. The second extinction occurred at the demise of glaciation and coincided with the return of anoxic conditions during the late Hirnantian–early Rhuddanian. Integration of our N isotope data with graptolite biodiversity records suggests that the extinctions were profoundly influenced by changes occurring at the base of the marine food web, i.e., redox-driven changes in nutrient cycling and primary producer communities.

Patterns and processes of latest Ordovician graptolite extinction and recovery based on data from south China

Abstract We have studied the pattern of graptolite species turnover during the latest Ordovician mass extinction based on four continuous Ashgillian to earliest Llandovery sections together with data from more than 30 other published sections. The studied sections represent relatively shallow-water and deeper-water belts in the Yangtze Platform region. Using temporally scaled range data, species diversities and extinction and origination probabilities have been calculated using several analytical methods, including a capture-mark-recapture method. We test the statistical significance of these results and the apparent taxonomic selectivity of extinction and origination via Monte Carlo simulations and contingency analysis. Graptolite species diversity within the Yangtze Platform rose steadily during the late Ashgill, until in the mid-late Paraorthograptus pacificus Chron, when rising extinction risk overtook origination. Diversity dropped to very low levels during the early Hirnantian when extinction probabilities attained significantly elevated rates for a period of 600–900 Ky. The period of high extinction risk was followed immediately by a short period of very high origination probability. A second, short period of high extinction risk occurred at the end of Hirnantian time. The Hirnantian extinction events marked a change from relatively low, steady origination and extinction probabilities to a prolonged period of elevated extinction risk and highly variable origination probability that extended well into the Rhuddanian. Extinction and origination was highly selective during the Hirnantian and favored both the survival and diversification of the Normalograptidae relative to the Dicranograptidae, Diplograptidae, and Orthograptidae. The main phase of extinction in the latest Rawtheyan and early Hirnantian was coincident with continental glaciation in the Southern Hemisphere. The resulting changes in ocean circulation and oxygenation appear to have almost completely eliminated the preferred habitat for most graptolite species. The Yangtze Platform region, however, may have served as a refugium for many taxa that disappeared earlier in other regions as well as a host site for the initiation of graptolite rediversification. Following the end of the glaciation, conditions favorable for graptolite proliferation were restored but graptolite communities remained unstable for much of the late Hirnantian and early Rhuddanian. Accordingly, the Hirnantian mass extinction appears to have fundamentally altered graptolite species dynamics as well as clade dominance patterns. A full understanding of the history of life requires an expanded, hierarchical theory of evolution that gives to mass extinctions (and other levels of selection) an appropriate role in determining clade and diversity histories.

Application of morphologic burrow interpretations to discern continental burrow architects: Lungfish or crayfish?

A methodology for trace fossil identification using burrowing signatures is tested by evaluating ancient and modern lungfish and crayfish burrows and comparing them to previously undescribed burrows in a stratigraphic interval thought to contain both lungfish and crayfish burrows. Permian burrows that bear skeletal remains of the lungfish Gnathorhiza, from museum collections, were evaluated to identify unique burrow morphologies that could be used to distinguish lungfish from crayfish burrows when fossil remains are absent. The lungfish burrows were evaluated for details of the burrowing mechanism preserved in the burrow morphologies together forming burrowing signatures and were compared to new burrows in the Chinle Formation of western Colorado to test the methodology of using burrow signatures to identify unknown burrows. Permian lungfish aestivation burrows show simple, nearly vertical, unbranched architectures and relatively smooth surficial morphologies with characteristic quasi‐horizontal striae on...

A combined landmark and outline-based approach to ontogenetic shape change in the Ordovician trilobite Triarthrus becki

Landmark based geometric morphometrics has developed as a powerful set of statistical and visual tools for the study of the covariance patterns of organismal shape change with a range of variables or factors. The approach is limited in the kinds of shape information accessible to it, however, by the need to employ discrete landmarks as the basis for comparison. In particular, curves and complex outlines are difficult to address using strictly landmark-based methods. Information about curves may be incorporated into the study of shape by the use of semi-landmark methods, which allow information about curved surfaces to be incorporated into the framework of landmark-based geometric morphometrics. We present a discussion of several software and statistical approaches needed to carry out combined landmark and semi-landmark analysis. In particular, we demonstrate several approaches to semi-landmark alignment (including the “edgewarp” method) and compare these to standard landmark based methods utilizing a regression analysis of the Ordovician trilobite Triarthrus becki. Abundant landmarks on the cranidium of T. becki allow landmark methods to represent the shape of that structure effectively, making it a good test case for combined landmark and semi-landmark methods. We verify that patterns of ontogenetic change implied by regression models using varying combinations of landmark and semi-landmark information are consistent with one another. Thus, semi-landmark methods and standard landmark based geometric morphometric methods yield commensurate information about this ontogenetic shape transformation. These results suggests that semi-landmark methods show substantial promise for rigorously testing hypotheses that involve the comparison of shapes when an adequate set of landmarks is not available.

Why the null matters: statistical tests, random walks and evolution

A number of statistical tests have been developed to determine what type of dynamics underlie observed changes in morphology in evolutionary time series, based on the pattern of change within the time series. The theory of the ‘scaled maximum’, the ‘log-rate-interval’ (LRI) method, and the Hurst exponent all operate on the same principle of comparing the maximum change, or rate of change, in the observed dataset to the maximum change expected of a random walk. Less change in a dataset than expected of a random walk has been interpreted as indicating stabilizing selection, while more change implies directional selection. The ‘runs test’ in contrast, operates on the sequencing of steps, rather than on excursion. Applications of these tests to computer generated, simulated time series of known dynamical form and various levels of additive noise indicate that there is a fundamental asymmetry in the rate of type II errors of the tests based on excursion: they are all highly sensitive to noise in models of directional selection that result in a linear trend within a time series, but are largely noise immune in the case of a simple model of stabilizing selection. Additionally, the LRI method has a lower sensitivity than originally claimed, due to the large range of LRI rates produced by random walks. Examination of the published results of these tests show that they have seldom produced a conclusion that an observed evolutionary time series was due to directional selection, a result which needs closer examination in light of the asymmetric response of these tests.

Graptoloid diversity and disparity became decoupled during the Ordovician mass extinction

The morphological study of extinct taxa allows for analysis of a diverse set of macroevolutionary hypotheses, including testing for change in the magnitude of morphological divergence, extinction selectivity on form, and the ecological context of radiations. Late Ordovician graptoloids experienced a phylogenetic bottleneck at the Hirnantian mass extinction (∼445 Ma), when a major clade of graptoloids was driven to extinction while another clade simultaneously radiated. In this study, we developed a dataset of 49 ecologically relevant characters for 183 species with which we tested two main hypotheses: (i) could the biased survival of one graptoloid clade over another have resulted from morphological selectivity alone and (ii) are the temporal patterns of morphological disparity and innovation during the recovery consistent with an interpretation as an adaptive radiation resulting from ecological release? We find that a general model of morphological selectivity has a low probability of producing the observed pattern of taxonomic selectivity. Contrary to predictions from theory on adaptive radiations and ecological speciation, changes in disparity and species richness are uncoupled. We also find that the early recovery is unexpectedly characterized by relatively low morphological disparity and innovation, despite also being an interval of elevated speciation. Because it is necessary to invoke factors other than ecology to explain the graptoloid recovery, more complex models may be needed to explain recovery dynamics after mass extinctions.

K-bentonites and graptolites biostratigraphy in the Middle Ordovician of New York State and Quebec; a new chronostratigraphic model

Published correlations of the extensive set of altered vol- canic ash beds (K-bentonites) present in the upper Middle Ordovician rocks of New York State imply that the lower Climacograptus (Diplacanthograptus) spiniferus Zone, the entire Orthograptus ruedemanni Zone, and the upper Corynoides americanus Zone are coeval. Precise new information on the ranges of graptolite and conodont species and the position of zonal boundaries within key Mohawk Valley sections together with new data on the occurrence of K-bentonites provide the foundation for an alternative, integrated chronostratigraphic model for the Middle Ordovician of the Mohawk Valley and the St. Lawrence Lowland of Quebec. This alternative model rec- onciles the K-bentonite and biostratigraphic correlations, but remains to be independently tested

Temporal and spatial distribution of biozones and facies relative to geochemically correlated K-bentonites in the Middle Ordovician Taconic foredeep

The Middle Ordovician rocks of the Taconic foredeep in the Mohawk Valley, New York, contain numerous altered volcanic ash beds (K-bentonites). These synchronous beds have the potential to link disparate facies in a way that is not feasible with biostratigraphy or sequence stratigraphy alone. Geochemical fingerprinting of glass inclusions within volcanic quartz phenocrysts permits unambiguous matching of the compositionally unique ash layers. The resulting correlations demonstrate that time lines based on the graptolite biozones and transgressive-regressive facies patterns parallel the K-bentonite isochrons. They also demonstrate that much of the upper Trenton Group is older than previously believed and facies relations are markedly different from previous interpretations. Although some facies patterns appear to be synchronous across the region, the likely existence of substantial structural control on facies development in the basin suggests that it is unwise to attribute these patterns to eustatic sea-level changes.