Occurrence and Range-through database of functional diversity of marine ecosystems after the Late Permian mass extinction event

The Late Permian mass extinction event about 252 million years ago was the most severe biotic crisis of the past 500 million years and occurred during an episode of global warming. The loss of around two-thirds of marine genera is thought to have had substantial ecological effects, but the overall impacts on the functioning of marine ecosystems and the pattern of marine recovery are uncertain. Here we analyse the fossil occurrences of all known benthic marine invertebrate genera from the Permian and Triassic periods, and assign each to a functional group based on their inferred lifestyle. We show that despite the selective extinction of 62-74% of these genera, all but one functional group persisted through the crisis, indicating that there was no significant loss of functional diversity at the global scale. In addition, only one new mode of life originated in the extinction aftermath. We suggest that Early Triassic marine ecosystems were not as ecologically depauperate as widely assumed. Functional diversity was, however, reduced in particular regions and habitats, such as tropical reefs; at these smaller scales, recovery varied spatially and temporally, probably driven by migration of surviving groups. We find that marine ecosystems did not return to their pre-extinction state, and by the Middle Triassic greater functional evenness is recorded, resulting from the radiation of previously subordinate groups such as motile, epifaunal grazers.

Analyses of shocked quartz from the K-P boundary

The precise cause and timing of the Cretaceous-Paleocene (K-P) mass extinction 65 Ma ago remains a matter of debate. Many advocate that the extinction was caused by a meteorite impact at Chicxulub, Mexico, and a number of potential kill-mechanisms have been proposed for this. Although we now have good constraints on the size of this impact and chemistry of the target rocks, estimates of its environmental consequences are hindered by a lack of knowledge about the obliquity of this impact. An oblique impact is likely to have been far more catastrophic than a sub-vertical one, because greater volumes of volatiles would have been released into the atmosphere. The principal purpose of this study was to characterize shocked quartz within distal K-P ejecta, to investigate whether the quartz distribution carried a signature of the direction and angle of impact. Our analyses show that the total number, maximum and average size of shocked quartz grains all decrease gradually with paleodistance from Chicxulub. We do not find particularly high abundances in Pacific sites relative to Atlantic and European sites, as has been previously reported, and the size-distribution around Chicxulub is relatively symmetric. Ejecta samples at any one site display features that are indicative of a wide range of shock pressures, but the mean degree of shock increases with paleodistance. These shock- and size-distributions are both consistent with the K-P layer having been formed by a single impact at Chicxulub. One site in the South Atlantic contains quartz indicating an anomalously high average shock degree, that may be indicative of an oblique impact with an uprange direction to the southeast +/- 45°. The apparent continuous coverage of proximal ejecta in this quadrant of the crater, however, suggests a relatively high impact angle of >45°. We conclude that some of the more extreme predictions of the environmental consequences of a low-angle impact at Chicxulub are probably not applicable.

Eocene-Oligocene calcareous nannofossils from ODP sites 115-711 and 120-748 in the Indian Ocean

An Eocene-Oligocene calcareous nannofossil biostratigraphic framework for Ocean Drilling Program (ODP) Site 748 in the southern Indian Ocean is established, which provides a foundation for this and future quantitative biogeographic studies. This biostratigraphic analysis, together with quantitative nannofossil data, enables a reinterpretation of the preliminary magnetostratigraphy and a new placement for magnetic Subchron CBN in the lowermost Oligocene. Calcareous nannofossil species diversity is low at Site 748 relative to lower latitude sites, with about 13 taxa in the middle Eocene, gradually decreasing to about 6 in the late Oligocene. There is, however, no apparent mass extinction at any stratigraphic level. Similarly, no mass extinctions were recorded at or near the Eocene/Oligocene boundary at Site 711 in the equatorial Indian Ocean. Species diversity at the equatorial site is significantly higher than at Site 748, with a maximum of 39 species in the middle Eocene and a minimum of 14 species in the late Oligocene. The abundance patterns of nannofossil taxa are also quite different at the two sites, with chiasmoliths, Isthmolithus recurvus, and Reticulofenestra daviesii abundant and restricted to the high-latitude site and Coccolithus formosus, discoasters, and sphenoliths abundant at the equatorial site but impoverished at the high-latitude site. This indicates a significant latitudinal biogeographic gradient between the equatorial site and the high-latitude site in the Indian Ocean for the middle Eocene-Oligocene interval. The abundance change of warm-water taxa is similar to that of species diversity at Site 711. There is a general trend of decreasing abundance of warm-water taxa from the middle Eocene through the early Oligocene at Site 711, suggesting a gradual cooling of the surface waters in the equatorial Indian Ocean. The abundance of warm-water taxa increased in the late Oligocene, in association with an increase in species diversity, and this may reflect a warming of the surface waters in the late Oligocene. An abrupt increase in the abundance of cool-water taxa (from ~20% to over 90%) occurred from 36.3 to 35.9 Ma at high-latitude Site 748. Coincident with this event was a ~1.0 per mil positive shift in the delta18O value of planktonic foraminifers and the occurrence of ice-rafted debris. This abrupt change in the nannofossil population is a useful biostratigraphic event for locating the bottom of magnetic Subchron C13N in the Southern Ocean. The sharp increase in cool-water taxa coeval with a large positive shift in delta18O values suggests that the high-latitude surface waters drastically cooled around 36.3-35.9 Ma. The temperature drop is estimated to be 4°C or more at Site 748 based on the nannofossil population change relative to the latitudinal biogeographic gradient established in the South Atlantic Ocean during previous studies. Consequently, much of the delta18O increase at Site 748 appears to be due to a temperature drop in the high latitudes rather than an ice-volume signal. The ~0.1 per mil delta18O increase not accounted for by the temperature drop is attributed to an ice-volume increase of 4.6 * 10**3 km**3, or 20% the size of the present Antarctic ice sheet.

Maastrichtian planktic foraminifera of the South Atlantic Ocean

Stratigraphic, faunal and isotopic analyses of the Maastrichtian at DSDP sites 525A and 21 in the South Atlantic reveal a planktic foraminiferal fauna characterized by two major events, an early late Maastrichtian diversification and end-Maastrichtian mass extinction. Both events are accompanied by major changes in climate and productivity. The diversification event which occurred in two steps between 70.5 and 69.1 Ma increased species richness by a total of 43% and coincided with the onset of major cooling in surface and bottom waters and increased surface productivity. The onset of the terminal decline in Maastrichtian species richness began at 67.5 Ma and the first significant decline in surface productivity occurred at 66.2 Ma, coincident maximum cooling to 13°C in surface waters and the reduction of the surface-to-deep temperature gradient to less than 5°C. Major climatic and moderate productivity changes mark the mass extinction and the last 500 kyr of the Maastrichtian. Between 200 and 400 kyr before the K-T boundary surface and deep waters warmed rapidly by 3-4°C and cooled again during the last 100 kyr of the Maastrichtian. Surface productivity decreased only moderately across the K-T boundary. Species richness began to decline during the late Maastrichtian cooling and by K-T boundary time, the mass extinction had claimed 66% of the species. Viewed within the context of Maastrichtian climate and productivity changes, the K-T mass extinction could have resulted from extreme environmental stress even without the addition of an extraterrestrial impact.

Marine diatoms in deep sea sediments

Long-term evolution is thought to take opportunities that arise as a consequence of mass extinction (as argued, for example, by Gould, 2002) and the following biotic recovery, but there is absolutely no evidence for this being the case. However, our study shows that eutrophication by oceanic mixing also played a part in the enhancement of several evolutionary events amongst marine organisms, and these results could indicate that the rates of oceanic biodiversification may be slowed if upwelling becomes weakened by future global warming. This paper defines three distinct evolutionary events of resting spores of the marine diatom genus Chaetoceros, to reconstruct past upwelling through the analysis of several DSDP, ODP and land-based successions from the North, South and equatorial Pacific as well as the Atlantic Ocean during the past 40 million years. The Atlantic Chaetoceros Explosion (ACE) event occurred across the E/O boundary in the North Atlantic, and is characterized by resting spore diversification that occurred as a consequence of the onset of upwelling following changes in thermohaline circulation through global cooling in the early Oligocene. Pacific Chaetoceros Explosion events-1 and -2 (PACE-1 and PACE-2) are characterized by relatively higher occurrences of iron input following the Himalayan uplift and aridification at 8.5 Ma and ca. 2.5 Ma in the North Pacific region. These events not only enhanced the diversification and increased abundance of primary producers, including that of Chaetoceros, other diatoms and seaweeds, but also stimulated the evolution of zooplankton and larger predators, such as copepods and marine mammals, which ate these phytoplankton and plants. Current thinking suggests new evolutionary niches open up after a mass extinction, but our study finds that eutrophication can also stimulate evolutionary diversification. Moreover, in the opposite fashion, our results show that as thermohaline circulation abates, global warming progresses and the ocean surface becomes warmer, many marine organisms will be affected by the environmental degradation.

Oxygen isotope values from biogenic apatie (conodont elements and fish teeth) from the Lower Triassic Mianwali Formation (Salt Range, Pakistan)

Recovery from the end-Permian mass extinction is frequently described as delayed, with complex ecological communities typically not found in the fossil record until the Middle Triassic epoch. However, the taxonomic diversity of a number of marine groups, ranging from ammonoids to benthic foraminifera, peaked rapidly in the Early Triassic. These variations in biodiversity occur amidst pronounced excursions in the carbon isotope record, which are compatible with episodes of massive CO2 outgassing from the Siberian Large Igneous Province. Here we present a high-resolution Early Triassic temperature record based on the oxygen isotope composition of pristine apatite from fossil conodonts. Our reconstruction shows that the beginning of the Smithian substage of the Early Triassic was marked by a cooler climate, followed by an interval of warmth lasting until the Spathian substage boundary. Cooler conditions resumed in the Spathian. We find the greatest increases in taxonomic diversity during the cooler phases of the early Smithian and early Spathian. In contrast, a period of extreme warmth in the middle and late Smithian was associated with floral ecological change and high faunal taxonomic turnover in the ocean. We suggest that climate upheaval and carbon-cycle perturbations due to volcanic outgassing were important drivers of Early Triassic biotic recovery.

The distributions, C- and O-isotopic compositions, and frequency of giant-ooids

Early Triassic oceans were characterized by deposition of a number of "anachronistic facies", including microbialites, seafloor carbonate cement fans, and giant ooids. Giant ooids were particularly prevalent in Lower Triassic sections across South China and exhibit unusual features that may provide insights into marine environmental conditions following the end-Permian mass extinction. The section at Moyang (Guizhou Province) contains abundant giant ooids ranging in size between 2 and 6 mm (maximum 12 mm) and exhibiting various cortical structures, including regular, deformed, compound, regenerated and "domed". Preservation of ooid cortical structure is generally good as indicated by petrographic observations, and trace element and carbon isotope analyses suggest that diagenesis occurred in a closed diagenetic system. All ooids exhibit fine concentric laminae, frequently alternating between light-colored coarsely crystalline and dark-colored finely crystalline layers probably reflecting variation in organic content or original mineralogy. Under scanning electron microscope, biomineralized filaments or biofilms and tiny carbonate fluorapatite (CFA) crystals are commonly found in the finely crystalline layers. We infer that the precipitation of CFA was related to adsorption of P via microbial activity on the surfaces of ooids following episodic incursions of deep waters rich in carbon dioxide, hydrogen sulfide and phosphate into shallow-marine environments. Giant ooid precipitation may have been promoted in shallow ramp settings during these events by increased watermass agitation and supersaturation with respect to calcium carbonate, as well as reduced carbonate removal rates through biotic skeletal formation. Spatio-temporal distribution data reveal that giant ooids were widespread in the Tethyan region during the Early Triassic, and that they were most abundant immediately after the end-Permian crisis and disappeared gradually as metazoans repopulated marine environments.

Benthic foraminifera across the Cretaceous/Paleogene boundary in ODP Hole 198-1210A

Deep-sea benthic foraminifera show important but transient assemblage changes at the Cretaceous/Paleogene (K/Pg) boundary, when many biota suffered severe extinction. We quantitatively analyzed benthic foraminiferal assemblages from lower bathyal-upper abyssal (1500-2000 m) northwest Pacific ODP Site 1210 (Shatsky Rise) and compared the results with published data on assemblages at lower bathyal (~ 1500 m) Pacific DSDP Site 465 (Hess Rise) to gain insight in paleoecological and paleoenvironmental changes at that time. At both sites, diversity and heterogeneity rapidly decreased across the K/Pg boundary, then recovered. Species assemblages at both sites show a similar pattern of turnover from the uppermost Maastrichtian into the lowermost Danian: 1) The relative abundance of buliminids (indicative of a generally high food supply) increases towards the uppermost Cretaceous, and peaks rapidly just above the K/Pg boundary, coeval with a peak in benthic foraminiferal accumulation rate (BFAR), a proxy for food supply. 2) A peak in relative abundance of Stensioeina beccariiformis, a cosmopolitan form generally more common at the middle than at the lower bathyal sites, occurs just above the buliminid peak. 3) The relative abundance of Nuttallides truempyi, a more oligotrophic form, decreases at the boundary, then increases above the peak in Stensioeina beccariiformis. The food supply to the deep sea in the Pacific Ocean thus apparently increased rather than decreased in the earliest Danian. The low benthic diversity during a time of high food supply indicates a stressed environment. This stress might have been caused by reorganization of the planktic ecosystem: primary producer niches vacated by the mass extinction of calcifying nannoplankton may have been rapidly (<10 kyr) filled by other, possibly opportunistic, primary producers, leading to delivery of another type of food, and/or irregular food delivery through a succession of opportunistic blooms. The deep-sea benthic foraminiferal data thus are in strong disagreement with the widely accepted hypothesis that the global deep-sea floor became severely food-depleted following the K/Pg extinction due to the mass extinction of primary producers ("Strangelove Ocean Model") or to the collapse of the biotic pump ("Living Ocean Model").

Web-based Biodiversity Citizen Science Database (assembled 2012)

The collective impact of humans on biodiversity rivals mass extinction events defining Earth's history, but does our large population also present opportunities to document and contend with this crisis? We provide the first quantitative review of biodiversity-related citizen science to determine whether data collected by these projects can be, and are currently being, effectively used in biodiversity research. We find strong evidence of the potential of citizen science: within projects we sampled (n = 388), ~1.3 million volunteers participate, contributing up to US Dollar 2.5 billion in-kind annually. These projects exceed most federally-funded studies in spatial and temporal extent, and collectively they sample a breadth of taxonomic diversity. However, only 12% of the 388 projects surveyed obviously provide data to peer-reviewed scientific articles, despite the fact that a third of these projects have verifiable, standardized data that are accessible online. Factors influencing publication included project spatial scale and longevity and having publically available data, as well as one measure of scientific rigor (taxonomic identification training). Because of the low rate at which citizen science data reach publication, the large and growing citizen science movement is likely only realizing a small portion of its potential impact on the scientific research community. Strengthening connections between professional and non-professional participants in the scientific process will enable this large data resource to be better harnessed to understand and address global change impacts on biodiversity.

Planktic foraminiferal turnover across the Paleocene-Eocene transition in the Bay of Biscay, North Atlantic

Planktic foraminifera across the Paleocene-Eocene transition at DSDP Site 401 indicate that the benthic foraminiferal mass extinction occurred within Subzone P 6a of Berggren and Miller (1988), or PS of Berggren et al. (1995) and coincident with a sudden 2.0? excursion in 6r3C values. The benthic foraminiferal extinction event (BFEE) and Sr3C excursion was accompanied by a planktic foraminiferal turnover marked by an influx of warm water species (Morozovella and Acarinina), a decrease in cooler water species (Subbotina), a sudden short-term increase in low oxygen tolerant taxa (Chiloguembelina), and no significant species extinctions. These faunal changes suggest climatic warming, expansion of the oxygen minimum zone, and a well stratified ocean water column. Oxygen isotope data of the surface dweller M. subbotina suggest climate warming beginning with a gradual 0.5? decrease in delta180 in the 175 cm preceding the benthic foraminiferal extinction event followed by a sudden decrease of 1? (4°C) at the BFEE. The delta13C excursion occurred over 27 cm of sediment and, assuming constant sediment accumulation rates, represents a maximum of 23 ka. Recovery to pre-excursion delta13C values occurs within 172 cm, or about 144 ka. Climate cooling begins in Subzone P 6c as indicated by an increase in cooler water subbotinids and acarininids with rounded chambers and a decrease in warm water morozovellids.

Maastrichtian calcareous nannofossils of the Indian Ocean and Shatsky Rise

The biotic effects of volcanism have long been the unknown factors in creating biotic stress, and the contribution of the Deccan volcanism to the K-T mass extinction remains largely unknown. Detailed studies of the volcanic-rich sediments of Indian Ocean Ninetyeast Ridge Sites 216 and 217 and Wharton Basin Site 212 reveal that the biotic effects of late Maastrichtian volcanism on planktic foraminifera and calcareous nannofossils are locally as severe as those of the K-T mass extinction. The biotic expressions of these high stress environments are characterized by the Lilliput effect, which includes reduced diversity by eliminating most K-strategy species, and reduction in specimen size (dwarfing), frequently to less than half their normal adult size of both r-strategy and surviving K-strategy species. In planktic foraminifera, the most extreme biotic stress results are nearly monospecific assemblages dominated by the disaster opportunist Guembelitria, similar to the aftermath of the K-T mass extinction. The first stage of improving environmental conditions results in dominance of dwarfed low oxygen tolerant Heterohelix species and the presence of a few small r-strategy species (Hedbergella, Globigerinelloides). Calcareous nannofossil assemblages show similar biotic stress signals with the dominance of Micula decussata, the disaster opportunist, and size reduction in the mean length of subordinate r-strategy species particularly in Arkhangelskiella cymbiformis and Watznaueria barnesiae. These impoverished and dwarfed late Maastrichtian assemblages appear to be the direct consequences of mantle plume volcanism and associated environmental changes, including high nutrient influx leading to eutrophic and mesotrophic waters, low oxygen in the water column and decreased watermass stratification.