The Late Ordovician mass extinction (LOME), sometimes referred to as the end-Ordovician mass extinction or the Ordovician–Silurian extinction, is the first of the "big five" major mass extinction events in Earth's history, occurring roughly 445 million years ago (Ma). It is often considered to be the second-largest-known extinction event, behind only the end-Permian mass extinction, in terms of the percentage of genera that became extinct. Extinction was global during this interval, eliminating 49–60% of marine genera and nearly 85% of marine species. Under most tabulations, only the Permian–Triassic mass extinction exceeds the Late Ordovician mass extinction in biodiversity loss. The extinction event had a profound impact on all major contemporary taxonomic groups; it caused the disappearance of one third of all brachiopod and bryozoan families, as well as numerous groups of conodonts, trilobites, echinoderms, corals, bivalves and graptolites. Despite its taxonomic severity, the Late Ordovician mass extinction did not produce major changes to ecosystem structures compared to other mass extinctions, nor did it lead to any particular morphological innovations. Diversity gradually recovered to pre-extinction levels over the first 5 million years of the Silurian period. The Late Ordovician mass extinction is traditionally considered to occur in two distinct pulses. The first pulse (interval), known as LOMEI-1, began at the boundary between the Katian and Hirnantian stages of the Late Ordovician epoch. This extinction pulse is typically attributed to the Late Ordovician glaciation, which abruptly expanded over Gondwana at the beginning of the Hirnantian and shifted the Earth from a greenhouse to icehouse climate. Cooling and a falling sea level brought on by the glaciation led to habitat loss for many organisms along the continental shelves, especially endemic taxa with restricted temperature tolerance and latitudinal range. During this extinction pulse, there were also several marked changes in biologically responsive carbon and oxygen isotopes. Marine life partially rediversified during the cold period and a new cold-water ecosystem, the "Hirnantia fauna", was established. The second pulse (interval) of extinction, referred to as LOMEI-2, occurred in the later half of the Hirnantian as the glaciation abruptly receded and warm conditions returned. The second pulse was associated with intense worldwide anoxia (oxygen depletion) and euxinia (toxic sulfide production), which persisted into the subsequent Rhuddanian stage of the Silurian period. Some researchers have proposed the existence of a third distinct pulse of the mass extinction during the early Rhuddanian, evidenced by a negative carbon isotope excursion and a pulse of anoxia into shelf environments amidst already low background oxygen levels. Others, however, have argued that Rhuddanian anoxia was simply part of the second pulse, which according to this view was longer and more drawn out than most authors suggest.
Impact on life
Ecological impacts The Late Ordovician mass extinction followed the Great Ordovician Biodiversification Event (GOBE), one of the largest surges of increasing biodiversity in the geological and biological history of the Earth. At the time of the extinction, most complex multicellular organisms lived in the sea, and the only evidence of life on land are rare spores from small early land plants. At the time of the extinction, around 100 marine families became extinct, covering about 49% of genera (a more reliable estimate than species). The brachiopods and bryozoans were strongly impacted, along with many of the trilobite, conodont and graptolite families. The extinction was divided into two major extinction pulses. The first pulse occurred at the base of the global Metabolograptus extraordinarius graptolite biozone, which marks the end of the Katian stage and the start of the Hirnantian stage. The second pulse of extinction occurred in the later part of the Hirnantian stage, coinciding with the Metabolograptus persculptus zone. Each extinction pulse affected different groups of animals and was followed by a rediversification event. Statistical analysis of marine losses at this time suggests that the decrease in diversity was mainly caused by a sharp increase in extinctions, rather than a decrease in speciation. Following such a major loss of diversity, Silurian communities were initially less complex and broader niched. Nonetheless, in South China, warm-water benthic communities with complex trophic webs thrived immediately following LOME. Highly endemic faunas, which characterized the Late Ordovician, were replaced by faunas that were amongst the most cosmopolitan in the Phanerozoic, biogeographic patterns that persisted throughout most of the Silurian. LOME had few of the long-term ecological impacts associated with the Permian–Triassic and Cretaceous–Paleogene extinction events. Furthermore, biotic recovery from LOME proceeded at a much faster rate than it did after the Permian–Triassic extinction. Nevertheless, a large number of taxa disappeared from the Earth over a short time interval, eliminating and altering the relative diversity and abundance of certain groups. The Cambrian-type evolutionary fauna nearly died out, and was unable to rediversify after the extinction.
Biodiversity changes in marine invertebrates
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