Michael J. Melchin

High-resolution, early Paleozoic (Ordovician-Silurian) time scales

For much of the geologic time scale, resolving power appears to be limited by the duration of biostratigraphic zones and subzones. Yet these zones exploit the appearance and disappearance events of only a small fraction of the species they span. Computer algorithms can sequence an order of magnitude more events and make explicit the uncertainties that arise when calibrating the resulting time line. An illustrative case history builds an Ordovician and Silurian time scale from the geologic record of the entire graptolite clade—over 1900 species from more than 400 localities and sections worldwide. The same approach can be applied to any stratigraphic interval with detailed biostratigraphic observations. It provides the foundation for time scales and paleobiologic time lines. Using logic similar to graphic correlation, optimizing algorithms search for a composite sequence of species first- and last-appearance events that minimizes the implied shortcomings of all the field observations. The algorithms minimize the number of species coexistences that are implied, but never observed, and the net adjustments needed to bring local range charts into agreement with a single composite sequence of events. After total section thicknesses have been rescaled to help normalize for variation in depositional rate, mean stratigraphic thicknesses are used to scale the intervals between adjacent events in the composite sequence. The resulting scaled composite sequence is converted to a relative geologic time scale by identifying stage and zone boundaries within the sequence of graptolite events. This relative scale is, in turn, calibrated by dated volcanic ash beds that were incorporated in the search for the optimal sequence. These dated events are also used to test for linearity of the scaled composite. The final time scale has a potential resolving power of 0.02–0.1 m.y., more than ten times better than can be achieved by traditional zones. Graptolite zones vary widely in duration from as short as 0.1 m.y. to nearly 5.0 m.y. The mean duration of zones or zonal groupings calibrated here is 1.44 m.y. in the Ordovician and 0.91 m.y. in the Silurian. The average uncertainty in locating zone boundaries in a single composite sequence is about one-fourth of the mean zone duration. The variance resulting from differences in time scales developed from differing numbers of field observations and in response to changes in the optimization criteria gives a very conservative measure of the overall robustness of the method. These differences indicate an average uncertainty in the age of graptolite zone boundaries that is more nearly equal to the mean zone duration. For zone duration, the mean uncertainty amounts to about one-half of the length of an average zone.

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.

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.

Carbon isotope chemostratigraphy of the Llandovery in Arctic Canada: Implications for global correlation and sea-level change

Abstract Stratigraphic and δ13C data from the Rhuddanian to lower Telychian succession on Cornwallis Island, Arctic Canada show evidence of a significant positive δ13C excursion in the upper Aeronian and weak positive shifts in the mid-Rhuddanian and uppermost Rhuddanian—lower Aeronian. The lower and upper Aeronian levels coincide with times of continental glaciation in Gondwana and these can be correlated with events recorded in the δ13C records from Dobs Linn (Scotland), Anticosti Island (Quebec), and Estonia. In most instances in the C-isotope records appear to coincide closely with local sea-level changes. The available data suggest that the global environmental impact of the late Aeronian glaciation was greater than that of the early Aeronian event. The data support the hypothesis that changes in degree of carbonate platform exposure and weathering resulting from of a combination of local and global (glacioeustatic) sea-level changes (particularly sea-level fall) were an important controlling factor in the generation of positive δ13C excursions in this time interval. These changes shifted the isotope value of the C-weathering flux entering shelf seas, which in turn resulted in positive δ13C excursions of varying magnitudes in shelf and deep basinal settings. These varied regionally in magnitude in close correspondence with differences in local sea-level histories.

Wenlock (Silurian) graptolites, Cape Phillips Formation, Canadian Arctic Islands

Twenty six of the more important species of monograptids (s.l.), retiolitids and Cyrtograptus (from a total fauna of 52 species) are described from Wenlock strata of the Cape Phillips Formation, Canadian Arctic Archipelago. Of this fauna, eight new species or subspecies, Monograptus firmus festinolatus, M. instrenuus, M. opimus, M. testis incomptus, Cyrtograptus falcatus, C. hamatus brevis, C. kolobus and C. pseudomancki , are described and illustrated. Wenlock biostratigraphic zones comprise the Cyrtograptus centrifugus-C. insectus Zone (earliest Wenlock), M. instrenuus-C. kolobus Zone, tentatively divisible into lower and upper subzones, C. perneri-M. opimus Zone possibly divisible into lower and upper subzones, C. lundgreni-M. testis Zone divisible into a lower M. testis incomptus Subzone and an upper M. testis testis Subzone, and the Pristiograptus ludensis Zone (latest Wenlock).

Silurian Tectonics of Western Avalonia: Strain-Corrected Subsidence History of the Arisaig Group, Nova Scotia

The Arisaig Group is a Silurian succession of predominantly shallow marine clastic sediments overlying volcanics, exposed in northern mainland Nova Scotia. Sediment accumulation provides a record of the subsidence of western Avalonia during the interval when terranes were being accreted within the Canadian Appalachians. To calculate the amount of subsidence, one must correct the measured thicknesses for the effects of tectonic strain. Deformed fossils on bedding surfaces indicate strain ratios mainly between 1.2 and 1.6. An empirical porosity-depth relationship is used to correct for compaction. The subsidence curves are then adjusted to allow for variations in water depth and eustatic sea level. The resulting curves show significant variations in subsidence rate regardless of which version of the Silurian time-scale is used. An initial episode of rapid subsidence followed eruption of Llandoverian volcanics. Slower subsidence took place in Wenlockian and Ludlovian time, with deeper-water sedimentation during an early Ludlovian eustatic high. This part of the history is consistent with thermal subsidence following an initial extensional event. A rapid increase in subsidence rate occurred during deposition of the Pridolian Stonehouse Formation. This episode was accompanied by a rapid increase in sediment supply, and a change in paleocurrent flow from southwest to northwest. The rapid Pridolian subsidence probably resulted from oblique collision between Avalonia and the Meguma Terrane to the south.

Thermal Maturity of Lower Paleozoic Sedimentary Successions in Arctic Canada

Mean maximum graptolite reflectance values from numerous sections in Arctic Canada range from 0.6% in Cornwallis Island and northwestern Devon Island to 4.7% in Ellesmere Island. We attribute this great lateral reflectance variation to differing burial depths of the graptolite-bearing strata beneath thick synorogenic siliciclastic covers. We attribute the low maturity of rocks in northern Cornwallis and eastern Bathurst islands to a maximum burial of about 2 km. Elsewhere, we used the paleogeographic location and proximity to the Ellesmerian overthrust wedge to interpret the measured reflectance values. In northern Ellesmere Island, where the highest graptolite reflectance values (4.7%) occur, as much as 9.6 km of synorogenic siliciclastics accumulated on a tectonically loaded carbonate shelf. Initial synorogenic siliciclastics encountered substantial submarine-to-basin relief, and thus about 2 km of sediment were deposited prior to the initiation of deposition on the adjacent drowned shelf. Also, the deep-water sequence probably was underlain by attenuated continental crust adjacent to the southeastward-advancing Ellesmerian overthrust wedge. Together, these factors caused deposition of as much as 7.6 km of sediment in western Melville Island and, as much as 9.6 km in northeastern Ellesmere Island. On the drowned shelf, synorogenic stratal thicknesses were less, a feature we attribu e to a thicker, more rigid crustal sequence and greater distance from the Ellesmerian tectonic loading. We expect liquid hydrocarbons to be generated from the organic matter in the shales in areas with a graptolite R0 maximum (GR0 max) of less than 1.7%. The presence of two types of solid bitumens having different reflectances, morphologies, and optical textures suggests that hydrocarbons were generated and migrated through the graptolite-bearing strata. We expect only gaseous hydrocarbons in areas where GR0 max exceeds 2.0% R0.

Ashgillian graptolite fauna of the Yangtze region and the biogeographical distribution of diversity in the latest Ordovician

Ashgillian graptolites have been described and recorded globally from 15 different paleoplates. The most diverse graptolite faunas are from the Yangtze region, South China, including 28 genera and 96 species. Among them, 25 genera and 73 species belong to the DDO fauna (Dicranograptidae – Diplograptidae – Orthograptidae fauna), and only three genera but 23 species belong to the N fauna (Normalograptidae fauna). Among the Yangtze graptolite fauna there are six endemic genera and 51 endemic species, which represent 21% and 52%, respectively, of the Ashgillian fauna in the region. This is an unusually high level of endemism. Endemic species are present in both the endemic and the cosmopolitan genera. A late Ashgillian stepwise extinction event has recently been recognized, based on graptolite diversity changes and graphic correlation. This begins with a major extinction from the Diceratograptus mirus Subzone to the middle Normalograptus extraordinarius-N. ojsuensis Zone, and ends with a minor extinction prior to the end of the Normalograptus persculptus Zone. Graptolite faunal replacement of the DDO fauna by the N fauna occurred throughout this interval. A comparison of Ashgillian graptolite diversity between the Yangtze region and other regions indicates that two different biogeographical realms existed in mid-Ashgillian time. A moderate-diversity graptolite fauna is present in the low-middle latitude realm, which includes South Scotland, Kazakhstan, Kolyma, Nevada, Yukon, Canadian Arctic, SE Australia. The Yangtze region was located in this realm, but was characterized by a very high-diversity fauna. Some other localities, including eastern Avalonia (Wales and England), the Argentina Precordillera, and Bohemia, which mainly represent the mid- to high-latitude realm, contain the lower-diversity mid-Ashgillian assemblages. This biogeographical distribution suggests a latitudinal diversity gradient, which may be controlled mainly by water temperature. This climate gradient becomes much less evident by late Hirnantian time in which most parts of the world have a relatively low diversity fauna totally dominated by normalograptid species, many of which appear to have been eurytopic. Throughout the Ashgillian, however, the Yangtze platform shows a high diversity and long persistence of DDO taxa the mass extinction interval. This may be a consequence of the semi-restricted nature of the basin in which conditions relatively favorable to graptolite survival and speciation existed throughout all or most of the Hirnantian.

Geobiodiversity Database: a comprehensive section-based integration of stratigraphic and paleontological data

The Geobiodiversity Database (GBDB – www.geobiodiversity.com), an integrated system for the management and analysis of stratigraphic and paleontological information, was started in 2006 and became available online in 2007. Its goal is to facilitate regional and global scientific collaborations focused on regional and global correlation, quantitative stratigraphy, systematics, biodiversity dynamics, paleogeography and paleoecology. It is unique among global, public access databases in that it is a section-based online database system, incorporating data from a wide range of disciplines of stratigraphy and paleontology, with inherent interrelationship between different kinds of data sets. It provides the capability of completely digitizing raw data, as well as integrating of different interpretations to the same paleontological and stratigraphic content. Several Windows-based visualization and analysis applications, either fully integrated with the database or supported by subset-export functions, have been developed to make the database more useful as a scientific and educational tool. The GBDB became the formal database of the International Commission on Stratigraphy (ICS) in August 2012 at the 34th International Geological Congress in Brisbane, and will produce comprehensive and authoritative web-based stratigraphic information service for global geoscientists, educators and the public.