Physiological advantages of dwarfing in surviving extinctions in high-CO2 oceans

Excessive CO2 in the present-day ocean-atmosphere system is causing ocean acidification, and is likely to cause a severe biodiversity decline in the future, mirroring effects in many past mass extinctions. Fossil records demonstrate that organisms surviving such events were often smaller than those before, a phenomenon called the Lilliput effect. Here, we show that two gastropod species adapted to acidified seawater at shallow-water CO2 seeps were smaller than those found in normal pH conditions and had higher mass-specific energy consumption but significantly lower whole-animal metabolic energy demand. These physiological changes allowed the animals to maintain calcification and to partially repair shell dissolution. These observations of the long-term chronic effects of increased CO2 levels forewarn of changes we can expect in marine ecosystems as CO2 emissions continue to rise unchecked, and support the hypothesis that ocean acidification contributed to past extinction events. The ability to adapt through dwarfing can confer physiological advantages as the rate of CO2 emissions continues to increase.

(Table 1) Element analyses of DSDP Hole 15-149

An iridium anomaly has been found in coincidence with the known microtektite level in cores from Deep Sea Drilling Project site 149 in the Caribbean Sea. The iridium was probably not in the microtektites but deposited simultaneously with them; this could occur if the iridium was deposited from a dust cloud resulting from a bolide impact, as suggested for the anomaly associated with the Cretaceous-Tertiary boundary. Other workers have deduced that the microtektites are part of the North American strewn tektite field, which is dated at about 34 million years before present, and that the microtektite horizon in deep-sea cores is synchronous with the extinction of five radiolarian species. Mass extinctions also occur in terrestrial mammals within 4 million years of this time. The iridium anomaly and the tektites and microtektites are supportive of a major bolide impact about 34 million years ago.

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.

Calcareous dinocysts and grain size distribution along profile Stevns-Klint

The distribution patterns of calcareous dinoflagellate cysts were studied in the classic Cretaceous Tertiary (K-T) boundary section of Stevns Klint, Denmark, focusing mainly on the response of the cyst association to an abrupt environmental catastrophe. A major part of the Fish Clay, which covers the K-T boundary at its base and is exposed in the investigated section, contains fallout produced by an asteroid impact. Calcareous dinoflagenate cysts are the best preserved remains of carbonate-producing phytoplankton in this layer. The potential of this group of microfossils for the analysis of survival strategies and extinction patterns has been underestimated. The cyst species of the investigated section can be grouped into four assemblages that represent victims, survivors, opportunists, and specially adapted forms. The victims (Pithonelloideae) were an extremely successful group throughout the Upper Cretaceous, but were restricted to the narrow outer shelf. This restriction minimized their spatial distribution, which generally should be large to facilitate escape from unfavorable conditions. Spatial restriction optimized the population decrease by mass mortality, disabling a successful recovery. In contrast, the survivors that became the dominating group in the Danian had a wide spatial range from the shelf environment to the oceanic realm. A unique calcareous dinocyst assemblage in the Fish Clay shows that even under the stressed conditions immediately following the impact event, some species flourished due to special adaptation or high ecological tolerance. The ability of these dinoflagellate species to form calcareous resting cysts in combination with their generally wide spatial distribution in a variety of environments appears to be the main reason for a low extinction rate at the K-T boundary as opposed to the high extinction rate of other phytoplankton groups, such as the coccolithophorids.

(Table 1) Lipid yields for samples across the Cretaceous/Tertiary boundary in DSDP Hole 86-577

Core samples of calcareous sediments taken from above and below the proposed Cretaceous/Tertiary boundary (Sample 577-12-5, 130 cm) were examined for geochemical evidence of the mass extinctions and faunal successions that marked this period. The lipid compositions of the six core samples examined were virtually identical and were characterized by a large component of unresolved naphthenic hydrocarbons and a homologous series of an/mo-alkanes, both presumably of bacterial origin. The results of this preliminary study suggest that the lipids of sediments deposited over a several million year period encompassing the Cretaceous-Tertiary extinctions have been almost completely recycled by bacterial metabolism, which occurred under oxic depositional and/or diagenetic conditions and which left a unique bacterial signature with only minor traces of the original sedimentary lipids.

(Table 1) Sample descriptions, carbon contents, and lipid yields across the K/T boundary at DSDP Hole 93-605 and at Stevns Klint, Denmark

This reconnaissance study was undertaken to determine whether the mass extinctions and faunal successions that mark the Cretaceous/Tertiary (K/T) boundary left a discernible molecular fossil record in the sediments of this period. Lipid signatures of sediments taken from above and below the K/T boundary were compared in core and outcrop samples taken from two locations: the U.S. east coast continental margin (western Atlantic Ocean, DSDP Site 605) and Stevns Klint, Denmark. Four calcareous sediments taken from above and below the K/T boundary in DSDP Hole 605, Section 605-66-1, revealed changing lipid signatures between above and below that are characterized by a large component of unresolved naphthenic hydrocarbons and a homologous series of n-alkanes ranging from Ci6 to C33. These lipid signatures are attributed to an influx of a terrestrial higher plant component and to bacterial reworking of the sediments under partially anoxic depositional and/or diagenetic conditions. The outcrop samples from Stevns Klint had extremely low concentrations of indigenous lipids. The fish clay at the K/T boundary contained traces of microbial hydrocarbons and fatty acids, whereas the carbonates above and below had only microbial fatty acids and additional terrestrial resin acids. The data from both sites indicate a perturbation in the deposition of lipid compound classes across the K/T boundary.

Benthic foraminifera extinctions in the Cenozoic record

In the late Pliocene-middle Pleistocene a group of 95 species of elongate, cylindrical, deep-sea (lower bathyal-abyssal) benthic foraminifera became extinct. This Extinction Group (Ext. Gp), belonging to three families (all the Stilostomellidae and Pleurostomellidae, some of the Nodosariidae), was a major component (20-70%) of deep-sea foraminiferal assemblages in the middle Cenozoic and subsequently declined in abundance and species richness before finally disappearing almost completely during the mid-Pleistocene Climatic Transition (MPT). So what caused these declines and extinction? In this study 127 Ext. Gp species are identified from eight Cenozoic bathyal and abyssal sequences in the North Atlantic and equatorial Pacific Oceans. Most species are long-ranging with 80% originating in the Eocene or earlier. The greatest abundance and diversity of the Ext. Gp was in the warm oceanic conditions of the middle Eocene-early Oligocene. The group was subjected to significant changes in the composition of the faunal dominants and slightly enhanced species turnover during and soon after the rapid Eocene-Oligocene cooling event. Declines in the relative abundance and flux of the Ext. Gp, together with enhanced species loss, occurred during middle-late Miocene cooling, particularly at abyssal sites. The overall number of Ext. Gp species present began declining earlier at mid abyssal depths (in middle Miocene) than at upper abyssal (in late Pliocene-early Pleistocene) and then lower bathyal depths (in MPT). By far the most significant Ext. Gp declines in abundance and species loss occurred during the more severe glacial stages of the late Pliocene-middle Pleistocene. Clearly, the decline and extinction of this group of deep-sea foraminifera was related to the function of their specialized apertures and the stepwise cooling of global climate and deep water. We infer that the apertural modifications may be related to the method of food collection or processing, and that the extinctions may have resulted from the decline or loss of their specific phytoplankton or prokaryote food source, that was more directly impacted than the foraminifera by the cooling temperatures.