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.
Based on the new material of seven Ordovician-Silurian boundary sections investigated recently, together with previously published data, we analyze the temporal and spatial distributions of the ...Lungmachi black shales, a key petroleum source bed widely distributed in South China. The Lungmachi black shales range in age from the Normalograptus persculptus Biozone of the uppermost Ordovician to the Spirograptus guerichi Biozone of the lower Telychian, and ten graptolite biozones can be recognized within this unit. The basal and upper contacts of the Lungmachi black shales are diachronous. The basal contact ranges from the N. persculptus to the C. cyphus biozones, a span of five graptolite biozones over two stages. The upper contact ranges from the D. pectinatus-M, argenteus Biozone to the Spirograptus guerichi Biozone, which spans four graptolite biozones over two stages. The Yichang Uplift resulted in the formation of the Hunan-Hubei Submarine High in the border area of Hubei, Hunan, and Chongqing. This is supported by a break in sedimentation in this area spanning all or part of the Hirnantian, and in many areas extending into the underlying Katian and overlying Rhuddanian. Comparison of the distribution of the Katian to Rhuddanian strata in this area indicates a growth and subsequent reduction in area of the Hunan-Hubei Submarine High particularly in the Hirnantian to early Rhuddanian. This may partly represent the influence of the process of formation and melting of ice sheet in Ordovician South Pole and consequent sea level change.
Mass extinctions disrupt ecological communities. Although climate changes produce stress in ecological communities, few paleobiological studies have systematically addressed the impact of global ...climate changes on the fine details of community structure with a view to understanding how changes in community structure presage, or even cause, biodiversity decline during mass extinctions. Based on a novel Bayesian approach to biotope assessment, we present a study of changes in species abundance distribution patterns of macroplanktonic graptolite faunas (∼447–444 Ma) leading into the Late Ordovician mass extinction. Communities at two contrasting sites exhibit significant decreases in complexity and evenness as a consequence of the preferential decline in abundance of dysaerobic zone specialist species. The observed changes in community complexity and evenness commenced well before the dramatic population depletions that mark the tipping point of the extinction event. Initially, community changes tracked changes in the oceanic water masses, but these relations broke down during the onset of mass extinction. Environmental isotope and biomarker data suggest that sea surface temperature and nutrient cycling in the paleotropical oceans changed sharply during the latest Katian time, with consequent changes in the extent of the oxygen minimum zone and phytoplankton community composition. Although many impacted species persisted in ephemeral populations, increased extinction risk selectively depleted the diversity of paleotropical graptolite species during the latest Katian and early Hirnantian. The effects of long-term climate change on habitats can thus degrade populations in ways that cascade through communities, with effects that culminate in mass extinction.
Cambrian-Devonian sedimentary rocks of the northern Canadian Cordillera record both the establishment and demise of the Great American Carbonate Bank, a widespread carbonate platform system that ...fringed the ancestral continental margins of North America (Laurentia). Here, we present a new examination of the deep-water Road River Group of the Richardson Mountains, Yukon, Canada, which was deposited in an intra-platformal embayment or seaway within the Great American Carbonate Bank called the Richardson trough. Eleven detailed stratigraphic sections through the Road River Group along the upper canyon of the Peel River are compiled and integrated with geological mapping, facies analysis, carbonate and organic carbon isotope chemostratigraphy, and new biostratigraphic results to formalize four new formations within the type area of the Richardson Mountains (Cronin, Mount Hare, Tetlit, and Vittrekwa). We recognize nine mixed carbonate and siliciclastic deep-water facies associations in the Road River Group and propose these strata were deposited in basin-floor to slope environments. New biostratigraphic data suggest the Road River Group spans the late Cambrian (Furongian) - Middle Devonian (Eifelian), and new chemostratigraphic data record multiple global carbon isotopic events, including the late Cambrian Steptoean positive carbon isotope excursion, the Late Ordovician Guttenberg excursion, the Silurian Aeronian, Valgu, Mulde (mid-Homerian), Ireviken (early Sheinwoodian), and Lau excursions, and the Early Devonian Klonk excursion. Together, these new data not only help clarify nomenclatural debate centered around the Road River Group, but also provide critical new sedimentological, biostratigraphic, and isotopic data for these widely distributed rocks of the northern Canadian Cordillera.
The Fire Bay Formation of Trettin (1998), Clements Markham belt, Ellesmere Island, Canada, includes volcanic rocks described as Silurian in age based on Llandovery graptolites in adjacent clastic ...rocks. New field observations suggest the Llandovery fossils are from packages of the Silurian Danish River and/or Lands Lokk formations that are fault-bounded rather than stratigraphically tied to Ordovician sections that contain a 470.0 ± 0.2 Ma lithic tuff, volcaniclastic units with maximum depositional ages (MDAs) of 466 ± 2 and 462 ± 2 Ma based on detrital zircon, volcanic clasts with ages of 498 ± 6, 478 ± 4, and 477 ± 8 Ma, and Ordovician conodonts and graptolites of Darriwilian and Sandbian age, respectively. Since the Fire Bay Formation of Trettin (1998) lacks a type section and is fault-bounded with ambiguous age relationships, Ordovician volcanic units and fault-bounded clastic rocks correlated with the Hazen Formation are both included in the Fire Bay assemblage following the original interpretations of Trettin and Nowlan (1990). The Fire Bay assemblage records juvenile Ordovician arc magmatism proximal to the Pearya terrane. The adjacent Lands Lokk Formation yields bimodal age peaks at 440-430 and 465 Ma, MDA of 424 ± 3 Ma, and εHf(t) values of -5 to +10. The signature matches Ordovician Pearya units and Silurian circum-Arctic arc sources but there is no evidence for Silurian arc magmatism between the Pearya terrane and Laurentian margin, compatible with Pearya accretion during oblique Ordovician arc collision and Silurian sinistral translation along the northern Laurentian margin.
Early Paleozoic bottom waters were mainly ferruginous, like the Neoproterozoic, with a shift to increased euxinia in the Devonian.
The extent to which Paleozoic oceans differed from Neoproterozoic ...oceans and the causal relationship between biological evolution and changing environmental conditions are heavily debated. Here, we report a nearly continuous record of seafloor redox change from the deep-water upper Cambrian to Middle Devonian Road River Group of Yukon, Canada. Bottom waters were largely anoxic in the Richardson trough during the entirety of Road River Group deposition, while independent evidence from iron speciation and Mo/U ratios show that the biogeochemical nature of anoxia changed through time. Both in Yukon and globally, Ordovician through Early Devonian anoxic waters were broadly ferruginous (nonsulfidic), with a transition toward more euxinic (sulfidic) conditions in the mid–Early Devonian (Pragian), coincident with the early diversification of vascular plants and disappearance of graptolites. This ~80-million-year interval of the Paleozoic characterized by widespread ferruginous bottom waters represents a persistence of Neoproterozoic-like marine redox conditions well into the Phanerozoic.
An exceptional fauna of retiolitine graptolites from Aeronian and lowermost Telychian strata in Arctic Canada provides significant new insights into the phylogeny and history of diversity of ...retiolitine graptolites. All specimens were isolated by dissolution of calcite concretions. The results indicate that retiolitines emerged within the lower Aeronian and reached a higher than expected level of diversity and disparity of forms by mid-Aeronian time. The uppermost Aeronian is almost totally devoid of preserved graptolites in Arctic Canada and, therefore, our material provides few new insights into retiolitine morphology or diversity through that interval. Specimens assigned to Pseudoretiolites? sp. occur in well-dated lower Aeronian strata, thus representing the lowest known biostratigraphic occurrences of retiolitines globally. This taxon appears to be morphologically primitive in that the sicula is completely preserved, with preservation of the proximal regions of theca 11, as well as distal thecal fusellum on mature specimens. Cladistic analysis of the Llandovery retiolitines shows that Pseudoretiolites is a stem genus for all of the other retiolitine taxa, which comprise two clades: one consisting of Pseudoplegmatograptus, Retiolites, and Stomatograptus and their derivatives; and the other includes Rotaretiolites, Aeroretiolites, Aeroretiolites?, Eorograptus, Paraplectograptus, Paraplectograptus?, and Sokolovograptus, and all of those other taxa that had previously been placed in the Plectograptinae. We follow the recent proposal that all of the taxa traditionally assigned to the Retiolitidae be assigned to the subfamily Retiolitinae. The following new taxa are described: Pseudoretiolites hyrichus n. sp., Eorograptus spirifer n. sp., and Aeroretiolites cancellatus n. gen. n. sp.
Stratigraphic and δ
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C data from the Rhuddanian to lower Telychian succession on Cornwallis Island, Arctic Canada show evidence of a significant positive δ
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C 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 δ
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C records from Dob's 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 δ
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C 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 δ
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C 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.