Benthic foraminifera across the Cretaceous/Paleogene boundary in the Southern Ocean

The impact of an asteroid at the Cretaceous/Paleogene (K/Pg) boundary triggered dramatic biotic, biogeochemical and sedimentological changes in the oceans that have been intensively studied. Paleo-biogeographical differences in the biotic response to the impact and its environmental consequences, however, have been less well documented. We present a high-resolution analysis of benthic foraminiferal assemblages at Southern Ocean ODP Site 690 (Maud Rise, Weddell Sea, Antarctica). At this high latitude site, late Maastrichtian environmental variability was high, but benthic foraminiferal assemblages were not less diverse than at lower latitudes, in contrast to those of planktic calcifiers. Also in contrast to planktic calcifiers, benthic foraminifera did not suffer significant extinction at the K/Pg boundary, but show transient assemblage changes and decreased diversity. At Site 690, the extinction rate was even lower (~3%) than at other sites. The benthic foraminiferal accumulation rate varied little across the K/Pg boundary, indicating that food supply to the sea floor was affected to a lesser extent than at lower latitude sites. Compared to Maastrichtian assemblages, Danian assemblages have a lower diversity and greater relative abundance of heavily calcified taxa such as Stensioeina beccariiformis and Paralabamina lunata. This change in benthic foraminiferal assemblages could reflect post-extinction proliferation of different photosynthesizers (thus food for the benthos) than those dominant during the Late Cretaceous, therefore changes in the nature rather than in the amount of the organic matter supplied to the seafloor. However, severe extinction of pelagic calcifiers caused carbonate supersaturation in the oceans, thus might have given competitive advantage to species with large, heavily calcified tests. This indirect effect of the K/Pg impact thus may have influenced the deep-sea dwellers, documenting the complexity of the effects of major environmental disturbance.

Pollen profiles from three former lakes in the Lake Garda area

1. Late glacial and postglacial sediments from three former lakes in the Lake Garda area (Southern Alps) were investigated. 2. The pollen diagram from Bondone (1550 m) shows an older phase rich in NAP. A younger one corresponds with the Younger Dryas time according to two radiocarbon determinations. In the Preboreal no climatic deterioration could be found. 3. At first plants, which are nowadays typical for snow-ground, pioneer and dwarf shrub associations, immigrated into the surroundings of Bondone. In Alleröd times larch and pine appeared as the first trees. At the beginning of the Preboreal dense forest existed in that region. During the Alleröd timber line was at about 1500 m. 4. In the pollen diagrams from Saltarino (194 m) and Fiavè (654 m) an oldest period rich in NAP is followed by two stadial and two interstadial phases. Tree birches and larches immigrated during the oldest interstadial phase. 5. In the case of Saltarino and Fiavè only a preliminary dating could be made. A correlation seems to be possible with diagrams published by Zoller as well as with the diagram of Bondone. Discrepances in dating, which arise then, are discussed. According to the two possibilities of dating the youngest stadial is synchronous either with the so-called Piottino stadial or the Younger Dryas time. Consequently the oldest interstadial phase of Saltarino corresponds either with the Bölling or with a pre-Bölling interstadial. The last possibility seems to be more probable. 6. In the southern part of the Lake Garda area reforestation was preceded by a long shrub phase mainly with Juniperus. At about 650 m there was a period with Pinus mugo and only with a small amount of Juniperus before reforestation. A phase with Betula nana well known from areas north of the Alps could nowhere be found. 7. In the area under study larch appeared as the first tree. Lateron it has been the most important constituent of the forests near timber line. Birch, which plays an important role as a pioneer tree in Denmark - for instance at the transition of the pollen zones III/IV - as well as in Southern Germany during Bölling time, was of less importance at the southern border of the Alps. In that area the spreading of Pinus occurred very early causing dense forests. 8. During the last stadial phase (probably Younger Dryas time) dense forests with Pinus and Larix existed at 650 m. In the lower part of the Lake Garda area, however, both thermophilous trees as Quercus and herbs frequently occurred. This leads to the conclusion that during this time tree growth was limited by dryness in lower altitudes of the border of the Southern Alps. Pinus and Juniperus, however, do not show higher values in this period, a fact which cannot yet be explained. 9. A list of plants, which were found in the sediments, is compiled. Helodium lanatum, Dictamnus albus, Mercurialis cf. ovata, Buxus, Cerinthe cf. minor, Onosma, Anthericum and Asphodelus albus are findings, which are of special interest for the history of the flora of that region.

Total number of benthic foraminifera in sediments below the K/T boundary in the Atlantic and Pacific Ocean

Most species of Late Cretaceous deep-sea benthic foraminifera are believed to be cosmopolitan and therefore to exhibit only minor biogeographical differences. In this preliminary report, six Deep Sea Drilling Project (DSDP) sites from different oceans, paleolatitudes, and paleodepths were analyzed for terminal Cretaceous abyssal-bathyal benthic foraminifera in order to investigate their assumed cosmopolitan distribution and the question of whether different faunal compositions are related to time, different paleolatitudes, and/or different paleodepths. The material studied was obtained from the low-latitude Site 465 (Pacific Ocean), and the intermediate-latitude Sites 384 (North Atlantic) and 356, 516, 525, and 527 (South Atlantic). The material analyzed represents a time slice encompassing the last 20-50 k.y. of the Cretaceous. The faunas contain numerous "Velasco-type" species, such as Gavelinella beccariiformis (White), Cibicidoides velascoensis (Cushman), Nuttallides truempyi (Nuttall), Gaudryina pyramidata Cushman, and various gyroidinoids and buliminids. The results contradict the general assumption of the cosmopolitan nature of Late Cretaceous deep-sea benthic foraminifera advocated in the literature. Only about 9% of the taxa identified were found to be truly "cosmopolitan" through their occurrence at all the sites analyzed. On the basis of correspondence analysis and relative abundance data, three assemblages and three subassemblages were recognized: (1) a bathyal-abyssal assemblage [Nuttallinella sp. A, Cibicidoides hyphalus (Fisher), Valvalabamina sp. evolute form, and Gyroidinoides spp.] at the South Atlantic Sites 356, 516, 525, and 527, divided into three subassemblages, namely (a) a middle bathyal subassemblage [Eouvigerina subsculptura McNeil and Caldwell, Truaxia aspera (Cushman), and G. pyramidata] at Sites 516 and 525, (b) a lower bathyal subassemblage [Osangularia? sp., Pyramidina rudita (Cushman and Parker), and Quadrimorphina camerata (Brotzen)] at Site 356, and (c) an abyssal subassemblage [Gyroidinoides sp. C, Hyperammina-Bathysiphon, Gyroidinoides beisseli (White), and Globorotalites sp. B] at Site 527; (2) an abyssal assemblage [Buliminella cf. plana (Cushman and Parker) and Bulimina incisa Cushman] at the North Atlantic Site 384; and (3) a middle bathyal assemblage [Vulvulina sp. A, Osangularia navarroana (Cushman), Alabamina? sp., Bulimina velascoensis (Cushman), Spiroplectammina spp. calcareous forms, and Bulimina trinitatensis Cushman and Jarvis] at the Pacific Site 465.

Eocene benthic foraminiferal community DSDP of Holes 95-612, 95-613 and 11-108

Benthic foraminiferal biofacies may vary independently of water depth and water mass; however, calibration of biofacies and stratigraphic ranges with independent paleodepth estimates allows reconstruction of age-depth patterns applicable throughout the deep Atlantic (Tjalsma and Lohmann, 1983). We have attempted to test these faunal calibrations in a continental margin setting, reconstructing Eocene benthic foraminiferal distributions along a dip section afforded by the New Jersey Transect (DSDP Sites 612, 108, 613). The following independent estimates of Eocene depths for the transect were obtained by "backtracking," "backstripping," and by assuming increasing depth downdip ("paleoslope"): Site 612, near the middle/lower bathyal boundary (about 1000 m); Site 108, in the middle bathyal zone (about 1600 m); and Site 613, near the lower bathyal/upper abyssal boundary (about 2000 m). Within uncertainties of backtracking (hundreds of meters), these estimates agree with estimates of paleodepth based on comparison of the New Jersey margin biofacies with other backtracked faunas. The stratigraphic ranges of many benthic taxa correspond to those found at other Atlantic DSDP sites. The major biofacies patterns show: (1) a depth dichotomy between an early to middle Eocene Nuttallides truempyidominated biofacies (greater than 2000 m) and a Lenticulina-Osangularia-Alabamina cf. dissonata biofacies (1000- 2000 m); and (2) a difference between a middle and a late Eocene biofacies at Site 612. The faunal boundary at about 2000 m, between bathyal and abyssal zones, occurs not only on the margin, but also throughout the deep Atlantic. The faunal change between the middle and late Eocene at Site 612 was due to a decrease of Lenticulina spp., the local disappearance of N. truempyi, and establishment of a Bulimina alazanensis-Gyroidinoides spp. biofacies. Although this change could be attributed to local paleoceanographic or water-depth changes, we argue that it is the bathyal expression of a global deep-sea benthic foraminiferal change which occurred across the middle/late Eocene boundary.

Living benthic foraminiferal assemblages in surface samples of the Arabian Sea

The distribution, biomass, and diversity of living (Rose Bengal stained) deep-sea benthic foraminifera (>30 µm) were investigated with multicorer samples from seven stations in the Arabian Sea during the intermonsoonal periods in March and in September/October, 1995. Water depths of the stations ranged between 1916 and 4425 m. The distribution of benthic foraminifera was compared with dissolved oxygen, % organic carbon, % calcium carbonate, ammonium, % silica, chloroplastic pigment equivalents, sand content, pore water content of the sediment, and organic carbon flux to explain the foraminiferal patterns and depositional environments. A total of six species-communities comprising 178 living species were identified by principal component analysis. The seasonal comparison shows that at the western stations foraminiferal abundance and biomass were higher during the Spring Intermonsoon than during the Fall Intermonsoon. The regional comparison indicates a distinct gradient in abundance, biomass, and diversity from west to east, and for biomass from north to south. Highest values are recorded in the western part of the Arabian Sea, where the influence of coastal and offshore upwelling are responsible for high carbon fluxes. Estimated total biomass of living benthic foraminifera integrated for the upper 5 cm of the sediment ranged between 11 mg Corg m**-2 at the southern station and 420 mg Corg m**-2 at the western station. Foraminifera in the size range from 30 to 125 ?m, the so-called microforaminifera, contributed between 20 and 65% to the abundance, but only 3% to 28% to the biomass of the fauna. Highest values were found in the central and southern Arabian Sea, indicating their importance in oligotrophic deep-sea areas. The overall abundance of benthic foraminifera is positively correlated with oxygen content and pore volume, and partly with carbon content and chloroplastic pigment equivalents of the sediment. The distributional patterns of the communities seem to be controlled by sand fraction, dissolved oxygen, calcium carbonate and organic carbon content of the sediment, but the critical variables are of different significance for each community.

Quaternary through early Eocene benthic foraminifera off Peru

Stratigraphic assemblages of Quaternary through early Eocene benthic foraminifers were recovered among 10 Peru margin drill sites. Various hiatuses and intervals barren in foraminifers characterize the sections, but numerous samples contain abundant, well-preserved benthic foraminifers. Bathymetry of the extant species and California-based estimates of the paleobathymetry of the extinct species permit recognition of Quaternary sea-level fluctuations between shelf and upper bathyal depths that produced vertical migrations of oxygenated and low-oxygen habitats at the six shallow sites. Assemblages from lower-slope sites at about 9° and 11°S indicate a general subsidence of the continental margin from shelf or upper bathyal depths in Eocene time to the present lower bathyal depths. Data from 11°S suggest a major part of this subsidence occurred in late Oligocene to early Miocene time. Downslope-transported shelf specimens, particularly the small biserial species, Bolivina costata and B. vaughani, are major contributors to these lower bathyal assemblages from the middle Miocene through Quaternary time.

Abundance and biomass of in- and epibenthic invertebrate macrofauna in the German Bight from 1969 to 2000

In order to examine the long-term development of offshore macrozoobenthic soft-bottom communities of the German Bight, four representative permanent stations (MZB-SSd, -FSd, -Slt, -WB) have been sampled continuously since 1969. Inter-annual variability and possible long-term trends were analysed based on spring-time samples from 1969 until 2000. This is part of the ecological long-term series of the AWI and is supplemented by periodic large-scale mapping of the benthos. The main factors influencing the development of the benthic communities are biological interactions, climate, food supply (eutrophication) and the disturbance regime. The most frequent disturbances are sediment relocations during strong storms or by bottom trawling, while occasional oxygen deficiencies and extremely cold winters are important disturbance events working on a much larger scale. Benthic communities at the sampling stations show a large inter-annual variability combined with a variation on a roughly decadal scale. In accordance with large-scale system shifts reported for the North Sea, benthic community transitions occurred between roughly the 1970ies, 80ies and 90ies. The transitions between periods are not distinctly marked by strong changes but rather reflected in gradual changes of the species composition and dominance structure.

Plio-Pleistocene benthic foraminifera of the Tyrrhenian Sea

Benthic foraminifers from Site 652, Site 653 (Hole 653A), and Site 654 of Leg 107 (Tyrrhenian Sea, Western Mediterranean), which penetrated with more or less good recovery the Plio-Pleistocene stratigraphic interval, were studied in a total of 699 close-spaced samples. A total number of 269 species have been classified and their quantitative distribution in each sample is reported. The benthic foraminifers assemblage is more diversified in Site 654, less diversified in Site 652. Less than a half of the benthic foraminifers species listed from Plio-Pleistocene Italian land sections are present in the coeval deep-sea Tyrrhenian record, in which shallow water species are missing and Nodosarids are poorly represented. A very few species have comparable stratigraphic distribution in the three deep-sea sequences and in Italian land sections when compared against calcareous plankton biostratigraphy. In the same three sites, the first appearance levels of several species are younger and younger, and last appearance levels are earlier and earlier from Site 654 to Site 653 and Site 652. Five biostratigraphic events, biochronologically evaluated and occurring at the same level in the deepsea Tyrrhenian record and in several land sections, have been selected as zonal boundaries of the proposed benthic foraminifers biostratigraphic scheme. The Plio-Pleistocene interval has been subdivided into four biozones and one subzone, recognizable both in the deep-sea and land-based sequences. The Cibicidoides (?) italicus assemblage zone stretches from the base of the Pliocene to the extinction level of the zonal marker, biochronologically evaluated at 2.9 Ma. The Cibicidoides robertsonianus interval zone stretches from the Cibicidoides (?) italicus extinction level to the Pliocene Mediterranean FO of Gyroidinoides altiformis, evaluated at 2.4 Ma. The Gyroidinoides altiformis interval zone stretches from the Mediterranean Pliocene FO of the zonal marker to the appearance level of Articulina tubulosa, evaluated at 1.62 Ma. The Articulina tubulosa assemblage zone stretches from the appearance level of the zonal marker to the Recent. In the Articulina tubulosa biozone, the Hyalinea baltica subzone is proposed. The appearance level of Hyalinea baltica is evaluated at 1.35 Ma, well above the Plio-Pleistocene boundary as defined in the Vrica stratotype section.

Upper Maestrichtian to Eocene benthic foraminifera in ODP Leg 121 holes

Late Maestrichtian to late Eocene bathyal benthic foraminiferal faunas at Sites 752,753, and 754 on Broken Ridge in the eastern Indian Ocean were analyzed as to their stratigraphic distribution of species to clarify the relation between faunal turnovers and paleoceanographic changes. Based on Q-mode factor analysis, eight varimax assemblages were distinguished: the Stensioina beccariiformis assemblage in the upper Maestrichtian to upper Paleocene; the Cibicidoides hyphalus assemblage in the upper Maestrichtian; the Cibicidoides cf. pseudoperlucidus assemblage in the upper Paleocene; the Anomalinoides capitatusldanicus assemblage in the uppermost Paleocene to lower Eocene; the Cibicidoides subspiratus assemblage in the lower Eocene; the Nuttallides truempyi assemblage in the lower and middle Eocene; the Osangularia sp. 1 - Hanzawaia ammophila assemblage in the upper Eocene; and the Lenticulina spp. assemblage in the uppermost Eocene, Oligocene, and lower Miocene. The presence of the Osangularia sp. 1 - Hanzawaia ammophila assemblage is related to the shallowing episode on Broken Ridge (upper bathyal), as a result of the rifting event that occurred in the middle Eocene. The most distinct faunal change (the disappearance of about 37% of the species) occurred between the S. beccariiformis assemblage and the A. capitatusldanicus assemblage, at the end of the upper Paleocene. A. capitatusldanicus, Lenticulina spp., and varied forms of Cibicidoides replaced the Velasco-type fauna at this time. The timing of this event is well correlated with the known age at South Atlantic sites (Thomas, 1990 doi:10.2973/odp.proc.sr.113.123.1990; Kennett and Stott, 1990 doi:10.2973/odp.proc.sr.113.188.1990; Katz and Miller, 1990 doi:10.2973/odp.proc.sr.114.147.1991). The primary cause of the extinction of the Stensioina beccariiformis assemblage is elusive, but may have resulted from the cessation of deep-water formation in the Antarctic (Katz and Miller, 1990), and subsequent arrival of warm saline deep water (Thomas, 1990; Kennett and Stott, 1990). Another possibility may be a weakened influence of high-salinity water formed at the low latitudes such as the Tethys Sea. The extinction event corresponds to the change from higher delta13C values in benthic foraminifers to lower ones. An interpretation of delta13C values is that the eastern Indian deep water, characterized by young and nutrient-depleted water, became old water which was devoid of a supply of new water during the latest Paleocene to early Eocene. Prior to this benthic event, signals of related faunal change were detected in the following short periods: early and late Paleocene, near the boundary of nannofossil Zone CP4, and Zone CP5 of the late Paleocene at Site 752. Among common taxa in the upper Maestrichtian, only seven species disappeared or became extinct at the Cretaceous/ Tertiary boundary at Site 752. The benthic foraminiferal population did not change for up to 2 m above the boundary, in contrast to the rapid decrease of the plankt onic foraminiferal population at the boundary. A decrease in the number of benthic foraminifers occurs after that level, corresponding to an interval of decreased numbers of planktonic foraminifers and higher abundance of volcanic ash. Reduced species diversity (H') suggests a secondary effect attributable to the dissolution of foraminiferal tests. The different responses of planktonic and benthic foraminifers to the event just above the boundary suggest that the Cretaceous/Tertiary event was a surface event as also suggested by Thomas (1990). In addition, a positive shift of delta13C in benthic foraminifers after the event indicates nutrient-depleted bottom water at Site 752.

One size fits all: stability of metabolic scaling under warming and ocean acidification in echinoderms

Responses by marine species to ocean acidification (OA) have recently been shown to be modulated by external factors including temperature, food supply and salinity. However the role of a fundamental biological parameter relevant to all organisms, that of body size, in governing responses to multiple stressors has been almost entirely overlooked. Recent consensus suggests allometric scaling of metabolism with body size differs between species, the commonly cited 'universal' mass scaling exponent (b) of ¾ representing an average of exponents that naturally vary. One model, the Metabolic-Level Boundaries hypothesis, provides a testable prediction: that b will decrease within species under increasing temperature. However, no previous studies have examined how metabolic scaling may be directly affected by OA. We acclimated a wide body-mass range of three common NE Atlantic echinoderms (the sea star Asterias rubens, the brittlestars Ophiothrix fragilis and Amphiura filiformis) to two levels of pCO2 and three temperatures, and metabolic rates were determined using closed-chamber respirometry. The results show that contrary to some models these echinoderm species possess a notable degree of stability in metabolic scaling under different abiotic conditions; the mass scaling exponent (b) varied in value between species, but not within species under different conditions. Additionally, we found no effect of OA on metabolic rates in any species. These data suggest responses to abiotic stressors are not modulated by body size in these species, as reflected in the stability of the metabolic scaling relationship. Such equivalence in response across ontogenetic size ranges has important implications for the stability of ecological food webs.

Cenozoic benthic foraminifers from ODP Leg 134 holes

This paper discusses the Paleobathymetric and paleoenvironmental history of the New Hebrides Island Arc and North d'Entrecasteaux Ridge during Cenozoic time based on benthic foraminiferal and sedimentological data. Oligocene and Pliocene to Pleistocene benthic foraminiferal assemblages from Sites 827, 828, 829, and 832 of Ocean Drilling Program (ODP) Leg 134 (Vanuatu) are examined by means of Q-mode factor analysis. The results of this analysis recognize the following bathymetrically significant benthic foraminiferal biofacies: (1) Globocassidulina subglobosa biofacies and Bulimina aculeata-Bolivinita quadrilatera biofacies representing the upper bathyal zone (600-1500 m); (2) Gavelinopsis praegeri-Cibicides wuellerstorfi biofacies, indicating the Pacific Intermediate Water (water depth between 1500 and 2400 m); (3) Tosaia hanzawai-Globocassidulina muloccensis biofacies, Valvulineria gunjii biofacies, and the Melonis barleeanus-Melonis sphaeroides biofacies, which characterize the lower bathyal zone; (4) the Nuttallides umbonifera biofacies, which characterizes the interval between the lysocline (approximately 3500 m) and the carbonate compensation depth (approximately 4500 m); and (5) the Rhabdammina abyssorum biofacies representing the abyssal zone below the carbonate compensation depth. Benthic foraminiferal patterns are used to construct Paleobathymetric and paleogeographic profiles of the New Hebrides Island Arc and North d'Entrecasteaux Ridge for the following age boundaries: late Miocene/Pliocene, early/late Pliocene, Pliocene/Pleistocene, and Pleistocene/Holocene.

Benthic foraminifera in Northwest Indian Ocean surface sediments

The relationship between the distribution of benthic foraminifera and sediment type and depositional environment in the Arabian Sea is discussed. The benthic foraminiferal fauna were sampled in nineteen Recent surface sediment samples, and geochemical variables of the sediment of the same samples were measured. The water depths for the box core samples varies from 440 to 4040 m. A total of 103 species and six species-complexes were identified. The geochemical properties were found to correspond well to the sediment type and depositional environment and six different sediment/depositional environment types could be distinguished. Analysis of the benthic foraminiferal fauna reveals specific faunal assemblages that are closely related to these sediment/depositional environment types.