Was the Late Ordovician mass extinction truly exceptional?

The Late Ordovician mass extinction event is the oldest of the five great extinction events in the fossil record. It has long been regarded as an outlier among mass extinctions, primarily due to its association with a cooling climate. However, recent temporally better resolved fossil biodiversity estimates complicate this view, providing growing evidence for a prolonged but punctuated biodiversity decline modulated by changes in atmospheric composition, ocean chemistry, and viable habitat area. This evolving view invokes extinction drivers similar to those that occurred during other major extinctions; some are even factors in the current human-induced biodiversity crisis. Even this very ancient and, at first glance, exceptional event conveys important lessons about the intensifying 'sixth mass extinction'.

Synchronizing rock clocks in the late Cambrian.

The Cambrian is the most poorly dated period of the past 541 million years. This hampers analysis of profound environmental and biological changes that took place during this period. Astronomically forced climate cycles recognized in sediments and anchored to radioisotopic ages provide a powerful geochronometer that has fundamentally refined Mesozoic-Cenozoic time scales but not yet the Palaeozoic. Here we report a continuous astronomical signal detected as geochemical variations (1 mm resolution) in the late Cambrian Alum Shale Formation that is used to establish a 16-Myr-long astronomical time scale, anchored by radioisotopic dates. The resulting time scale is biostratigraphically well-constrained, allowing correlation of the late Cambrian global stage boundaries with the 405-kyr astrochronological framework. This enables a first assessment, in numerical time, of the evolution of major biotic and abiotic changes, including the end-Marjuman extinctions and the Steptoean Positive Carbon Isotope Excursion, that characterized the late Cambrian Earth.

Building mountain biodiversity: Geological and evolutionary processes.

Mountain regions are unusually biodiverse, with rich aggregations of small-ranged species that form centers of endemism. Mountains play an array of roles for Earth's biodiversity and affect neighboring lowlands through biotic interchange, changes in regional climate, and nutrient runoff. The high biodiversity of certain mountains reflects the interplay of multiple evolutionary mechanisms: enhanced speciation rates with distinct opportunities for coexistence and persistence of lineages, shaped by long-term climatic changes interacting with topographically dynamic landscapes. High diversity in most tropical mountains is tightly linked to bedrock geology-notably, areas comprising mafic and ultramafic lithologies, rock types rich in magnesium and poor in phosphate that present special requirements for plant physiology. Mountain biodiversity bears the signature of deep-time evolutionary and ecological processes, a history well worth preserving.

The evolutionary dynamics of the early Palaeozoic marine biodiversity accumulation.

The early Palaeozoic Era records the initial biodiversification of the Phanerozoic. The increase in biodiversity involved drastic changes in taxon longevity, and in rates of origination and extinction. Here, we calculate these variables in unprecedented temporal resolution. We find that highly volatile origination and extinction rates are associated with short genus longevities during the Cambrian Period. During the Ordovician and Silurian periods, evolutionary rates were less volatile and genera persisted for increasingly longer intervals. The 90%-genus life expectancy doubled from 5 Myr in the late Cambrian to more than 10 Myr in the Ordovician-Silurian periods. Intervals with widespread ecosystem disruption are associated with short genus longevities during the Cambrian and with exceptionally high longevities during the Ordovician and Silurian periods. The post-Cambrian increase in persistence of genera, therefore, indicates an elevated ability of the changing early Palaeozoic marine ecosystems to sustainably maintain existing genera. This is evidence of a new level of ecosystem resilience which evolved during the Ordovician Period.

Cascading trend of Early Paleozoic marine radiations paused by Late Ordovician extinctions.

The greatest relative changes in marine biodiversity accumulation occurred during the Early Paleozoic. The precision of temporal constraints on these changes is crude, hampering our understanding of their timing, duration, and links to causal mechanisms. We match fossil occurrence data to their lithostratigraphical ranges in the Paleobiology Database and correlate this inferred taxon range to a constructed set of biostratigraphically defined high-resolution time slices. In addition, we apply capture-recapture modeling approaches to calculate a biodiversity curve that also considers taphonomy and sampling biases with four times better resolution of previous estimates. Our method reveals a stepwise biodiversity increase with distinct Cambrian and Ordovician radiation events that are clearly separated by a 50-million-year-long period of slow biodiversity accumulation. The Ordovician Radiation is confined to a 15-million-year phase after which the Late Ordovician extinctions lowered generic richness and further delayed a biodiversity rebound by at least 35 million years. Based on a first-differences approach on potential abiotic drivers controlling richness, we find an overall correlation with oxygen levels, with temperature also exhibiting a coordinated trend once equatorial sea surface temperatures fell to present-day levels during the Middle Ordovician Darriwilian Age. Contrary to the traditional view of the Late Ordovician extinctions, our study suggests a protracted crisis interval linked to intense volcanism during the middle Late Ordovician Katian Age. As richness levels did not return to prior levels during the Silurian-a time of continental amalgamation-we further argue that plate tectonics exerted an overarching control on biodiversity accumulation.

Identifying the most surprising victims of mass extinction events: an example using Late Ordovician brachiopods.

Mass extinction events are recognized by increases in extinction rate and magnitude and, often, by changes in the selectivity of extinction. When considering the selective fingerprint of a particular event, not all taxon extinctions are equally informative: some would be expected even under a 'background' selectivity regime, whereas others would not and thus require special explanation. When evaluating possible drivers for the extinction event, the latter group is of particular interest. Here, we introduce a simple method for identifying these most surprising victims of extinction events by training models on background extinction intervals and using these models to make per-taxon assessments of 'expected' risk during the extinction interval. As an example, we examine brachiopod genus extinctions during the Late Ordovician Mass Extinction and show that extinction of genera in the deep-water 'Foliomena fauna' was particularly unexpected given preceding Late Ordovician extinction patterns.

Biogeographic and bathymetric determinants of brachiopod extinction and survival during the Late Ordovician mass extinction.

The Late Ordovician mass extinction (LOME) coincided with dramatic climate changes, but there are numerous ways in which these changes could have driven marine extinctions. We use a palaeobiogeographic database of rhynchonelliform brachiopods to examine the selectivity of Late Ordovician-Early Silurian genus extinctions and evaluate which extinction drivers are best supported by the data. The first (latest Katian) pulse of the LOME preferentially affected genera restricted to deeper waters or to relatively narrow (less than 35°) palaeolatitudinal ranges. This pattern is only observed in the latest Katian, suggesting that it reflects drivers unique to this interval. Extinction of exclusively deeper-water genera implies that changes in water mass properties such as dissolved oxygen content played an important role. Extinction of genera with narrow latitudinal ranges suggests that interactions between shifting climate zones and palaeobiogeography may also have been important. We test the latter hypothesis by estimating whether each genus would have been able to track habitats within its thermal tolerance range during the greenhouse-icehouse climate transition. Models including these estimates are favoured over alternative models. We argue that the LOME, long regarded as non-selective, is highly selective along biogeographic and bathymetric axes that are not closely correlated with taxonomic identity.

Onset of main Phanerozoic marine radiation sparked by emerging Mid Ordovician icehouse.

The Great Ordovician Biodiversification Event (GOBE) was the most rapid and sustained increase in marine Phanerozoic biodiversity. What generated this biotic response across Palaeozoic seascapes is a matter of debate; several intrinsic and extrinsic drivers have been suggested. One is Ordovician climate, which in recent years has undergone a paradigm shift from a text-book example of an extended greenhouse to an interval with transient cooling intervals - at least during the Late Ordovician. Here, we show the first unambiguous evidence for a sudden Mid Ordovician icehouse, comparable in magnitude to the Quaternary glaciations. We further demonstrate the initiation of this icehouse to coincide with the onset of the GOBE. This finding is based on both abiotic and biotic proxies obtained from the most comprehensive geochemical and palaeobiological dataset yet collected through this interval. We argue that the icehouse conditions increased latitudinal and bathymetrical temperature and oxygen gradients initiating an Early Palaeozoic Great Ocean Conveyor Belt. This fuelled the GOBE, as upwelling zones created new ecospace for the primary producers. A subsequent rise in δ(13)C ratios known as the Middle Darriwilian Isotopic Carbon Excursion (MDICE) may reflect a global response to increased bioproductivity encouraged by the onset of the GOBE.