Climate change and the selective signature of the Late Ordovician mass extinction

Selectivity patterns provide insights into the causes of ancient extinction events. The Late Ordovician mass extinction was related to Gondwanan glaciation; however, it is still unclear whether elevated extinction rates were attributable to record failure, habitat loss, or climatic cooling. We examined Middle Ordovician-Early Silurian North American fossil occurrences within a spatiotemporally explicit stratigraphic framework that allowed us to quantify rock record effects on a per-taxon basis and assay the interplay of macrostratigraphic and macroecological variables in determining extinction risk. Genera that had large proportions of their observed geographic ranges affected by stratigraphic truncation or environmental shifts at the end of the Katian stage were particularly hard hit. The duration of the subsequent sampling gaps had little effect on extinction risk, suggesting that this extinction pulse cannot be entirely attributed to rock record failure; rather, it was caused, in part, by habitat loss. Extinction risk at this time was also strongly influenced by the maximum paleolatitude at which a genus had previously been sampled, a macroecological trait linked to thermal tolerance. A model trained on the relationship between 16 explanatory variables and extinction patterns during the early Katian interval substantially underestimates the extinction of exclusively tropical taxa during the late Katian interval. These results indicate that glacioeustatic sea-level fall and tropical ocean cooling played important roles in the first pulse of the Late Ordovician mass extinction in Laurentia

Energetic Costs of Calcification Under Ocean Acidification

Anthropogenic ocean acidification threatens to negatively impact marine organisms that precipitate calcium carbonate skeletons. Past geological events, such as the Permian-Triassic Mass Extinction, together with modern experiments generally support these concerns. However, the physiological costs of producing a calcium carbonate skeleton under different acidification scenarios remain poorly understood. Here we present an idealized mathematical model to quantify whole-skeleton costs, concluding that they rise only modestly (up to ∼10%) under acidification expected for 2100. The modest magnitude of this effect reflects in part the low energetic cost of inorganic, calcium carbonate relative to the proteinaceous organic matrix component of skeletons. Our analysis does, however, point to an important kinetic constraint that depends on seawater carbonate chemistry, and we hypothesize that the impact of acidification is more likely to cause extinctions within groups where the timescale of larval development is tightly constrained. The cheapness of carbonate skeletons compared to organic materials also helps explain the widespread evolutionary convergence upon calcification within the metazoa

Marine extinction risk shaped by trait-environment interactions over 500 million years

Perhaps the most pressing issue in predicting biotic responses to present and future global change is understanding how environmental factors shape the relationship between ecological traits and extinction risk. The fossil record provides millions of years of insight into how extinction selectivity (i.e., differential extinction risk) is shaped by interactions between ecological traits and environmental conditions. Numerous paleontological studies have examined trait-based extinction selectivity; however, the extent to which these patterns are shaped by environmental conditions is poorly understood due to a lack of quantitative synthesis across studies. We conducted a meta-analysis of published studies on fossil marine bivalves and gastropods that span 458 million years to uncover how global environmental and geochemical changes covary with trait-based extinction selectivity. We focused on geographic range size and life habit (i.e., infaunal vs. epifaunal), two of the most important and commonly examined predictors of extinction selectivity. We used geochemical proxies related to global climate, as well as indicators of ocean acidification, to infer average global environmental conditions. Life-habit selectivity is weakly dependent on environmental conditions, with infaunal species relatively buffered from extinction during warmer climate states. In contrast, the odds of taxa with broad geographic ranges surviving an extinction ( \u3e 2500km for genera, \u3e 500km for species) are on average three times greater than narrow-ranging taxa (estimate of odds ratio: 2.8, 95% confidence interval=2.3-3.5), regardless of the prevailing global environmental conditions. The environmental independence of geographic range size extinction selectivity emphasizes the critical role of geographic range size in setting conservation priorities

A Temperature-Dependent Positive Feedback on the Magnitude of Carbon Isotope Excursions

The decrease in the average magnitude of carbon isotope excursions in marine carbonates over Phanerozoic time is a longstanding unresolved problem. In addition, carbon isotope excursions commonly co-occur with oxygen isotope excursions of the same sign, implying the existence of a longstanding link between organic carbon burial fluxes and temperature. It was proposed that this connection was provided by the thermodynamic relationship between temperature and microbial respiration rates – changes in temperature drive changes in organic carbon remineralization rate and organic carbon burial efficiency. Such a mechanism provides the logic for a positive feedback affecting the magnitude of both climate changes and carbon isotope excursions. Here, we employ feedback analysis to quantify the strength of this mechanism with modifications to a simple carbon isotope mass balance framework. We demonstrate that the potential strength of this feedback is large (perhaps several permil) for plausible ranges of historical climate change. Furthermore, our results highlight the importance of the surface temperature boundary condition on the magnitude of the expected carbon isotope excursion. Comparisons of our model predictions with data from the terminal Eocene and Late Ordovician (Hirnantian) greenhouse–icehouse climate transitions suggest that these excursions might be substantially explained by such a thermodynamic microbial respiration feedback. Consequently, we hypothesize that the observed pattern of decreasing excursion magnitude toward the present might be explained at least, in part, by a decrease in the mean temperature of environments of organic carbon burial driven by long-term climate and paleogeographic trends. SOMMAIRELa diminution de l'amplitude moyenne des excursions des isotopes du carbone dans les carbonates marins au fil du Phanérozoïque est une énigme de longue date.  On note en outre que les excursions des isotopes du carbone coexistent couramment avec des excursions isotopiques de même signe de l'oxygène, ce qui implique l'existence d'un lien de longue date entre les flux d’enfouissement du carbone organique et la température.  On a suggéré que ce lien découlait de la relation thermodynamique entre la température et les taux de respiration microbienne - les changements de température déterminent le taux de reminéralisation du carbone organique et l’efficacité de l’enfouissement du carbone organique.  Un tel mécanisme peut expliquer la rétroaction positive affectant à la fois l'ampleur des changements climatiques et les excursions des isotopes du carbone.  Dans le cas présent, nous utilisons l'analyse de la rétroaction pour quantifier la robustesse de ce mécanisme avec des modifications d’un simple bilan de masse des isotopes du carbone.  Nous démontrons que la robustesse potentielle de cette rétroaction est forte (peut-être plusieurs pour mille) dans les gammes plausibles du changement climatique historique.  De plus, nos résultats mettent en évidence l'importance de la condition aux limites de la température de surface sur l'ampleur de l'excursion isotopique du carbone attendue.  Les comparaisons des prédictions de notre modèle avec les données de la fin de l'Éocène et de la fin de l’Ordovicien (Hirnantien) des transitions climatiques à effet de serre-effet/de glaciation permettent de penser que ces excursions pourraient être correctement expliquées par une telle rétroaction de la thermodynamique de la respiration microbienne.  Par conséquent, nous émettons l'hypothèse que la tendance observée de diminution de l'ampleur de l’amplitude des excursions du passé vers le présent peut s'expliquer, au moins en partie, par une diminution de la température moyenne du milieu d'enfouissement du carbone organique engendrée par des tendances climatiques et paléogéographiques à long terme

Carbonates in Skeleton-poor Seas: New Insights From Cambrian and Ordovician Strata of Laurentia

Calcareous skeletons evolved as part of the greater Ediacaran–Cambrian diversification of marine animals. Skeletons did not become permanent, globally important sources of carbonate sediment, however, until the Ordovician radiation. Representative carbonate facies in a Series 3 (510–501 Ma) Cambrian to Tremadocian succession from western Newfoundland, Canada, and Ordovician successions from the Ibex area, Utah, USA, show that, on average, Cambrian and Tremadocian carbonates contain much less skeletal material than do post-Tremadocian sediments. Petrographic point counts of skeletal abundance within facies and proportional facies abundance in measured sections suggest that later Cambrian successions contain on average <5% skeletal material by volume, whereas the skeletal content of post-Tremadocian Ordovician sections is closer to ~15%. A compilation of carbonate stratigraphic sections from across Laurentia confirms that post-Tremadocian increase in skeletal content is a general pattern and not unique to the two basins studied. The long interval (~40 myr) between the initial Cambrian appearance of carbonate skeletons and the subsequent Ordovician diversification of heavily skeletonized organisms provides an important perspective on the Ordovician radiation. Geochemical data increasingly support the hypothesis that later Cambrian oceans were warm and, in subsurface water masses, commonly dysoxic to anoxic. We suggest that surface waters in such oceans would have been characterized by relatively low saturation states for calcite and aragonite. Mid-Ordovician cooling would have raised oxygen concentrations in subsurface water masses, establishing more highly oversaturated surface waters. If correct, these links could provide a proximal trigger for the renewed radiation of heavily skeletonized invertebrates and algae

Lipid biomarker and stable isotopic profiles through Early-Middle Ordovician carbonates from Spitsbergen, Norway

One of the most dramatic episodes of sustained diversification of marine ecosystems in Earth history took place during the Early to Middle Ordovician Period. Changes in climate, oceanographic conditions, and trophic structure are hypothesised to have been major drivers of these biotic events, but relatively little is known about the composition and stability of marine microbial communities controlling biogeochemical cycles at the base of the food chain. This study examines well-preserved, carbonate-rich strata spanning the Tremadocian through Upper Dapingian stages from the Oslobreen Group in Spitsbergen, Norway. Abundant bacterial lipid markers (elevated hopane/sterane ratios, average = 4.8; maximum of 13.1), detection of Chlorobi markers in organic-rich strata, and bulk nitrogen isotopes (delta N-15(total)) averaging 0 to -1 parts per thousand for the open marine facies, suggest episodes of water column redox-stratification and that primary production was likely limited by fixed nitrogen availability in the photic zone. Near absence of the C-30 sterane marine algal biomarker, 24-n-propylcholestane (24-npc), in most samples supports and extends the previously observed hiatus of 24-npc in Early Paleozoic (Late Cambrian to Early Silurian) marine environments. Very high abundances of 3 beta-methylhopanes (average = 9.9%; maximum of 16.8%), extends this biomarker characteristic to Early Ordovician strata for the first time and may reflect enhanced and sustained marine methane cycling during this interval of fluctuating climatic and low sulfate marine conditions. Olenid trilobite fossils are prominent in strata deposited during an interval of marine transgression with biomarker evidence for episodic euxinia/anoxia extending into the photic zone of the water column. (C) 2019 Elsevier Ltd. All rights reserved.Peer reviewe

The Ordovician succession adjacent to Hinlopenstretet, Ny Friesland, Spitsbergen

Peer reviewe

Records of carbon and sulfur cycling during the Silurian Ireviken Event in Gotland, Sweden

Field and stable isotope work was supported by an Agouron Institute grant to DAF and WWF as well as a Packard Fellowship and a Hanse-wissenschaftskolleg Fellowship awarded to DAF.Early Silurian (∼431 Ma) carbonate rocks record a ca. 4.5‰ positive excursion in the stable isotopic composition of carbonate carbon (δ13Ccarb). Associated with this isotopic shift is a macroevolutionary turnover pulse known as the ‘Ireviken Event’. The onset of this carbon isotope excursion is commonly associated with a shallowing-upward facies transition that may have been accompanied by climatic change, as indicated by a parallel positive shift (∼0.6‰) in the stable isotopic composition of carbonate oxygen (δ18Ocarb). However, the relationships among carbon cycle perturbations, faunal turnover, and environmental changes remain enigmatic. Here we present a suite of new isotopic data across the Ireviken Event from multiple sections in Gotland, Sweden. These samples preserve no systematic change in δ18Ocarb but show positive excursions of equal magnitude in both carbonate (δ13Ccarb) and organic (δ13Corg) carbon. In addition, the data reveal a synchronous perturbation in sulfur isotope ratios, manifest as a ca. 7‰ positive excursion in carbonate-associated sulfate (δ34SCAS) and a ca. 30‰ positive excursion in pyrite (δ34Spyr). The increase in δ34Spyr values is accompanied by a substantial, concomitant increase in stratigraphic variability of δ34Spyr. The relatively constant offset between the δ13Ccarb and δ13Corg excursions throughout the Ireviken Event could be attributed to increased organic carbon burial, or possibly a change in the isotopic composition of CO2 sources from weathering. However, a positive correlation between carbonate abundance and δ13Ccarb suggests that local to regional changes in dissolved inorganic carbon (DIC) during the shallowing-upward sequence may have been at least partly responsible for the observed excursion. The positive excursion recorded in δ34SCAS suggests a perturbation of sufficient magnitude and duration to have impacted the marine sulfate reservoir. An inverse correlation between CAS abundance and δ34SCAS supports the notion of decreased sulfate concentrations, at least locally, consistent with a concomitant increase in pyrite burial. A decrease in the offset between δ34SCAS and δ34Spyr values during the Ireviken Event suggests a substantial reduction in the isotopic fractionations (εpyr) expressed during microbial sulfur cycling and pyrite precipitation through this interval. Decreased εpyr and the concomitant increase in stratigraphic variation in δ34Spyr are typical of isotope systematics observed in modern shallow-water environments, associated with increased closed-system behavior and/or oxidative sedimentary reworking during early sediment diagenesis. While the isotopic trends associated with the Ireviken Event have been observed in multiple locations around the globe, many sections display different magnitudes of isotopic change, and moreover, are typically associated with local facies changes. Due to the stratigraphic coherence of the carbon and sulfur isotopic and abundance records across the Ireviken Event, and their relationship to changes in local depositional environment, we surmise that these patterns more closely reflect biogeochemical processes related to deposition and lithification of sediment than global changes in carbon and sulfur burial fluxes.PostprintPeer reviewe

A high-resolution record of early Paleozoic climate

The spatial coverage and temporal resolution of the Early Paleozoic paleoclimate record are limited, primarily due to the paucity of well-preserved skeletal material commonly used for oxygen-isotope paleothermometry. Bulk-rock δ¹⁸O datasets can provide broader coverage and higher resolution, but are prone to burial alteration. We assess the diagenetic character of two thick Cambro–Ordovician carbonate platforms with minimal to moderate burial by pairing clumped and bulk isotope analyses of micritic carbonates. Despite resetting of the clumped-isotope thermometer at both sites, our samples indicate relatively little change to their bulk δ¹⁸O due to low fluid exchange. Consequently, both sequences preserve temporal trends in δ¹⁸O. Motivated by this result, we compile a global suite of bulk rock δ¹⁸O data, stacking overlapping regional records to minimize diagenetic influences on overall trends. We find good agreement of bulk rock δ¹⁸O with brachiopod and conodont δ¹⁸O trends through time. Given evidence that the δ¹⁸O value of seawater has not evolved substantially through the Phanerozoic, we interpret this record as primarily reflecting changes in tropical, nearshore seawater temperatures and only moderately modified by diagenesis. Focusing on the samples with the most enriched, and thus likely least-altered, δ¹⁸O values, we reconstruct Late Cambrian warming, Early Ordovician extreme warmth, and cooling around the Early–Middle Ordovician boundary. Our record is consistent with models linking the Great Ordovician Biodiversification Event to cooling of previously very warm tropical oceans. In addition, our high-temporal-resolution record suggests previously unresolved transient warming and climate instability potentially associated with Late Ordovician tectonic events

The Geozoic Supereon

Geological time units are the lingua franca of earth sciences: they are a terminological convenience, a vernacular of any geological conversation, and a prerequisite of geo-scientific writing found throughout in earth science dictionaries and textbooks. Time units include terms formalized by stratigraphic committees as well as informal constructs erected ad hoc to communicate more efficiently. With these time terms we partition Earth’s history into utilitarian and intuitively understandable time segments that vary in length over seven orders of magnitude: from the 225-year-long Anthropocene (Crutzen and Stoermer, 2000) to the ,4-billion-year-long Precambrian (e.g., Hicks, 1885; Ball, 1906; formalized by De Villiers, 1969)