
The later part of the Devonian (the Frasnian and Famennian Stages) once marked the second largest marine mass extinction (~375 Ma) of the Phanerozoic Eon: it was one of the ‘Big Five’. For the last decade the drop in marine biodiversity in that interval has come under scrutiny: partly because it may have involved several events; no well-supported extinction mechanism has emerged; and extinctions seem have been concentrated on three animal groups – trilobites, brachiopods and reef corals. Something large did happen, as reef-building corals almost disappeared and coral reefs only returned with the rise of modern (scleractinian) corals in the Mesozoic. While the end of the Devonian still figures widely as having experienced a mass extinction event, more detailed palaeontological work at the genus and species level suggests another possibility.
‘Officially’ a mass extinction event must exceed the background extinction rate throughout the Phanerozoic and be above that in immediately preceding and following stages: statistically significant, that is. They always give rise to a marked reduction in biodiversity, but another mechanism can do that without extinctions suddenly increasing. The rate at which new species emerge can fall below that of species extinctions, when the overall number of living species falls. As far as ecosystems are concerned both processes are equally severe, but the causes may be very different.

Reviewing detailed records of Devonian species of two genera of brachiopods and one bivalve genus (50 species in all) in five North American stratigraphic sequences, Alycia Stigall of Ohio University, USA noted apparently significant variations in the local assemblages (Stigall, A. L. 2012. Speciation collapse and invasive species dynamics during the Late Devonian ‘Mass Extinction’. GSA Today, v. 22(1), p. 4-9). Speciation overall fell in the Frasnian and the preceding Givetian, while rate of extinction barely changed. For the three studied genera ,speciation reached low values in the Frasnian and Famennian, but that was accompanied by an equally large fall in extinctions. In this narrow sample we seem to be seeing not an extinction crisis but one of biodiversity. Why?
The Late Devonian saw repeated ups and downs in sea level, which repeatedly connected and disconnected shallow- to moderate-depth marine basins. The fossil record shows repeated cases of species from one basin colonising another, each invasion accompanying rapid marine transgression.. One means whereby species arise is through prolonged isolation of separate populations of the ancestral species through independent genetic drift and mutation. The episodic connection of basins may have prevented such allopatric speciation. Interestingly, the invading species were dominantly animals with a broad tolerance for environmental conditions.
Whether this mechanism applied to all three main animal groups whose diversity plummeted in Late Devonian times remains to be seen, and it begs the question ‘why didn’t it happen among other animal groups that were less affected by whatever the events were?’ One of the problems associated with decreasing biodiversity in modern marine (and terrestrial) settings is growth in the numbers of invasive species, so the work on 375 Ma fossils might help understand and mitigate current ecological issues. The only difference is that for many of the hyper-successful invader species the means of invasion has been provided by human activities. brachiopod brachioopod
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