Middle Miocene evolution and stable isotope ratios of Globorotalia (Fohsella) in the western equatorial Pacific

Evolution of the planktic foraminiferal lineage Globorotalia (Fohsella) occurred during the Miocene between 23.7 and 11.8 Ma and forms the basis for stratigraphic subdivision of the early middle Miocene (Zones N 10 through N 12). Important morphologic changes within the G. (Fohsella) lineage included a marked increase in test size, a transition from a rounded to an acute periphery, and the development of a keel in later forms. We found that the most rapid changes in morphology of G. (Fohsella) occurred between 13 and 12.7 Ma and coincided with an abrupt increase in the delta18O ratios of shell calcite. Comparison of isotopic results of G. (Fohsella) with other planktic foraminifers indicate that delta18O values of the lineage diverge from surface-dwelling species and approach deep-dwelling species after 13.0 Ma, indicating a change in depth habitat from the surface mixed layer to intermediate depth near the thermocline. Isotopic and faunal evidence suggests that this change in depth stratification was associated with an expansion of the thermocline in the western equatorial Pacific. After adapting to a deeper water habitat at 13.0 Ma, the G. (Fohsella) lineage became extinct abruptly at 11.8 Ma during a period when isotopic and faunal evidence suggest a shoaling of the thermocline. Following the extinction of G. (Fohsella), the ecologic niche of the lineage was filled by the Globorotalia (Menardella) group, which began as a deep-water form and later evolved to an intermediate-water habitat. We suggest that the evolution of G. (Fohsella) and G. (Menardella) were tightly linked to changes in the structure of the thermocline in the western equatorial Pacific.

Experiment: Adaptation of a globally important coccolithophore to ocean warming and acidification

Although oceanwarming and acidification are recognized as two major anthropogenic perturbations of today's oceanswe know very little about how marine phytoplankton may respond via evolutionary change.We tested for adaptation to ocean warming in combination with ocean acidification in the globally important phytoplankton species Emiliania huxleyi. Temperature adaptation occurred independently of ocean acidifcation levels. Exponential growth rates were were up to 16% higher in populations adapted for one year to warming when assayed at their upper thermal tolerance limit. Particulate inorganic (PIC) and organic (POC) carbon production was restored to values under present-day ocean conditions, owing to adaptive evolution, and were 101% and 55% higher under combined warming and acidification, respectively, than in non-adapted controls. Cells also evolved to a smaller size while they recovered their initial PIC:POC ratio even under elevated CO2. The observed changes in coccolithophore growth, calcite and biomass production, cell size and elemental composition demonstrate the importance of evolutionary processes for phytoplankton performance in a future ocean. At the end of a 1-yr temperature selection phase, we conducted a reciprocal assay experiment in which temperature-adapted asexual populations were compared to the respective non-adapted control populations under high temperature, and vice versa (1. Assay Data, Dataset #835336). Mean exponential growth rates ? in treatments subjected to high temperature increased rapidly under all high temperature-CO2 treatment combinations during the temperature selection phase (2. time series, Dataset #835339).

Seawater carbonate chemistry and combined mechanistic effects of CO2 and light on the N2-fixing cyanobacterium Trichodesmium IMS101, 2010

The marine diazotrophic cyanobacterium Trichodesmium responds to elevated atmospheric CO2 partial pressure (pCO2) with higher N2 fixation and growth rates. To unveil the underlying mechanisms, we examined the combined influence of pCO2(150 and 900 µatm) and light (50 and 200 µmol photons m-2 s-1) on TrichodesmiumIMS101. We expand on a complementary study that demonstrated that while elevated pCO2 enhanced N2 fixation and growth, oxygen evolution and carbon fixation increased mainly as a response to high light. Here, we investigated changes in the photosynthetic fluorescence parameters of photosystem II, in ratios of the photosynthetic units (photosystem I:photosystem II), and in the pool sizes of key proteins involved in the fixation of carbon and nitrogen as well as their subsequent assimilation. We show that the combined elevation in pCO2 and light controlled the operation of the CO2-concentrating mechanism and enhanced protein activity without increasing their pool size. Moreover, elevated pCO2 and high light decreased the amounts of several key proteins (NifH, PsbA, and PsaC), while amounts of AtpB and RbcL did not significantly change. Reduced investment in protein biosynthesis, without notably changing photosynthetic fluxes, could free up energy that can be reallocated to increase N2 fixation and growth at elevated pCO2 and light. We suggest that changes in the redox state of the photosynthetic electron transportchain and posttranslational regulation of key proteins mediate the high flexibility in resources and energy allocation in Trichodesmium. This strategy should enableTrichodesmium to flourish in future surface oceans characterized by elevated pCO2, higher temperatures, and high light.

Long-term dynamics of adaptive evolution in a globally important coccolithophore to ocean acidification

Recent evolution experiments have revealed that marine phytoplankton may adapt to global change, for example to ocean warming or acidification. Long-term adaptation to novel environments is a dynamic process and phenotypic change can take place thousands of generations after exposure to novel conditions. Using the longest evolution experiment performed in any marine species to date (4 yrs, = 2100 generations), we show that in the coccolithophore Emiliania huxleyi, long-term adaptation to ocean acidification is complex and initial phenotypic responses may revert for important traits. While fitness increased continuously, calcification was restored within the first 500 generations but later reduced in response to selection, enhancing physiological declines of calcification in response to ocean acidification. Interestingly, calcification was not constitutively reduced but revealed rates similar to control treatments when transferred back to present-day CO2 conditions. Growth rate increased with time in controls and adaptation treatments, although the effect size of adaptation assessed through reciprocal assay experiments varied. Several trait changes were associated with selection for higher cell division rates under laboratory conditions, such as reduced cell size and lower particulate organic carbon content per cell. Our results show that phytoplankton may evolve phenotypic plasticity that can affect biogeochemically important traits, such as calcification, in an unforeseen way under future ocean conditions.

Raw-data to morphometric investigations about the Neogene planktonic foraminifer Globorotalia menardii and related forms from ODP Hole 154-925B (Céara Rise, western tropical Atlantic)

Evolutionary prospection is the study of morphological evolution and speciation in calcareous plankton from selected time-slices and key sites in the world oceans. In this context, the Neogene menardiform globorotalids serve as study objects for morphological speciation in planktic foraminifera. A downcore investigation of test morphology of the lineage of G. menardii-limbata-multicamerata during the past 8 million years was carried out in the western tropical Atlantic ODP Hole 925B. A total of 4669 specimens were measured and analyzed from 38 stratigraphic levels and compared to previous studies from DSDP Sites 502 and 503. Collection of digital images and morphometric measurements from digitized outlines were achieved using a microfossil orientation and imaging robot called AMOR and software, which was especially developed for this purpose. Most attention was given to the evolution of spiral height versus axial length of tests in keel view, but other parameters were investigated as well. The variability of morphological parameters in G. menardii, G. limbata, and G. multicamerata through time are visualized by volume density diagrams. At Hole 925B results show gradual test size increase in G. menardii until about 3.2 Ma. The combination of taxonomic determination in the light microscope with morphometric investigations shows strong morphological overlap and evolutionary continuity from ancestral to extant G. menardii (4-6 chambers in the final whorl) to the descendent but extinct G. limbata (seven chambers in the final whorl) and to G. multicamerata (>=8 chambers in the final whorl). In the morphospace defined by spiral height (dX) and axial length (dY) Globorotalia limbata and G. multicamerata strongly overlap with G. menardii. Distinction of G. limbata from G. menardii is only possible by slight differences in the number of chambers of the final whorl, nuances in spiral convexity, upper keel angles, radii of osculating circles, or by differences in reflectance of their tests. Globorotalia multicamerata can be distinguished from the other two forms by more than eight chambers in the final whorl. It appeared as two stratigraphically separate clusters during the Pliocene. Between 2.88 and 2.3 Ma G. menardii was severely restricted in size and abundance. Thereafter, it showed a rapid and prominent expansion of the upper test size extremes between 2.3 and 1.95 Ma persisting until present. The size-frequency distributions at Hole 925B are surprisingly similar to trends of menardiform globorotalids from Caribbean DSDP Site 502. There, the observations were explained as an adaptation to changes in the upper water column due to the emergence of the Isthmus of Panama. In light of more recent paleontological and geological investigations about the completion of the permanent land connection between North and South America since about 3 Ma the present study gives reason to suspect the sudden test size increase of G. menardii to reflect immigration of extra-large G. menardii from the Indian Ocean or the Pacific. It is hypothesized that during the Late Pliocene dispersal of large G. menardii into the southern to tropical Atlantic occurred during an intermittent episode of intense Agulhas Current leakage around the Cape of Good Hope and from there via warm eddy transport to the tropical Atlantic (Agulhas dispersal hypothesis).

Diatom assemblages, HBIs and TEXL86 in sediment core NBP99-03_10

The West Antarctic ice sheet is particularly sensitive to global warming and its evolution and impact on global climate over the next few decades remains difficult to predict. In this context, investigating past sea ice conditions around Antarctica is of primary importance. Here, we document changes in sea ice presence, upper water column temperatures (0-200 m) and primary productivity over the last 9000 yr BP (before present) in the western Antarctic Peninsula (WAP) margin from a sedimentary core collected in the Palmer Deep Basin. Employing a multi-proxy approach, based on the combination of two biomarkers proxies (highly branched isoprenoid (HBI) alkenes for sea ice and TEXL86 for temperature) and micropaleontological data (diatom assemblages), we derived new Holocene records of sea ice conditions and upper water column temperatures. The early Holocene (9000-7000 yr BP) was characterized by a cooling phase with a short sea ice season. During the mid-Holocene (~7000-3800 yr BP), local climate evolved towards slightly colder conditions and a prominent extension of the sea ice season occurred, promoting a favorable environment for intensive diatom growth. The late Holocene (the last ~2100 yr) was characterized by warmer temperatures and increased sea ice presence, accompanied by reduced local primary productivity, likely in response to a shorter growing season compared to the early or mid-Holocene. The gradual increase in annual sea ice duration over the last 7000 yr might have been influenced by decreasing mean annual and spring insolation, despite increasing summer insolation. We postulate that, in addition to precessional changes in insolation, seasonal variability, via changes in the strength of the circumpolar Westerlies and upwelling activity, was further amplified by the increasing frequency/amplitude of the El Nino-Southern Oscillation (ENSO). However, between 3800 and 2100 yr BP, the lack of correlation between ENSO and climate variability in the WAP suggests that other climatic factors might have been more important in controlling WAP climate at this time.

Revised age models and foraminiferal records obtained for the Last Interglacial period in six marine sediment cores

The dataset contains the revised age models and foraminiferal records obtained for the Last Interglacial period in six marine sediment cores: - the Southern Ocean core MD02-2488 (age model, sea surface temperatures, benthic d18O and d13C for the period 136-108 ka), - the North Atlantic core MD95-2042 (age model, planktic d18O, benthic d18O and d13C for the period 135-110 ka), - the North Atlantic core ODP 980 (age model, planktic d18O, sea surface temperatures, seawater d18O, benthic d18O and d13C, ice-rafted detritus for the period 135-110 ka), - the North Atlantic core CH69-K09 (age model, planktic d18O, sea surface temperatures, seawater d18O, benthic d18O and d13C, ice-rafted detritus for the period 135-110 ka), - the Norwegian Sea core MD95-2010 (age model, percentage of Neogloboquadrina pachyderma sinistral, sea surface temperatures, benthic d18O, ice-rafted detritus for the period 134-110 ka), - the Labrador Sea core EW9302-JPC2 (age model, percentage of Neogloboquadrina pachyderma sinistral, sea surface temperatures, benthic d18O for the period 134-110 ka).

Chemical and Sr isotopic compositions of 3 step leachates and residues of bulk samples, zeolites, and smectites from Lower Miocene sediments at ODP Site 166-1007

Massive clinoptilolite authigenesis was observed at about 1105 meters below sea floor (mbsf) in lower Miocene wellcompacted carbonate periplatform sediments from the Great Bahama Bank [Ocean Drilling Program, ODP Leg 166, Site 1007]. The diagenetic assemblage comprises abundant zeolite crystallized within foraminifer tests and sedimentary matrix, as well as Mg smectites. In carbonate-rich deposits, the formation of the zeolite requires a supply of silica. Thus, the objective of the study is to determine the origin of the silica supply, its diagenetic evolution, and consequently the related implications on interpretation of the sedimentary record, in terms of local or global paleoceanographic change. For lack of evidence for any volcaniclastic input or traces of Si-enriched deep fluids circulation, an in situ biogenic source of silica is validated by isotopic data and chemical modeling for the formation of such secondary minerals in shallow-water carbonate sequences. Geochemical and strontium isotopic data clearly establish the marine signature of the diagenetic zeolite, as well as its contemporaneous formation with the carbonate deposition (Sr model ages of 19.6-23.2 Ma). The test of saturation for the pore fluids specifies the equilibrium state of the present mineralogical assemblage. Seawater-rock modeling specifies that clinoptilolite precipitates from the dissolution of biogenic silica, which reacts with clay minerals. The amount of silica (opal-A) involved in the reaction has to be significant enough, at least 10 wt.%, to account for the observed content of clinoptilolite occurring at the most zeolite-rich level. Modeling also shows that the observed amount of clinoptilolite (~19%) reflects an in situ and short-term reaction due to the high reactivity of primary biogenic silica (opal-A) until its complete depletion. The episodic occurrence of these well-lithified zeolite-rich levels is consistent with the occurrence of seismic reflectors, particularly the P2 seismic sequence boundary located at 1115 mbsf depth and dated as 23.2 Ma. The age range of most zeolitic sedimentary levels (biostratigraphic ages of 21.5-22 Ma) correlates well with that of the early Miocene glaciation Mi-1 and Mi-1a global events. Thus, the clinoptilolite occurrence in the shallow carbonate platform environment far from volcanogenic supply, or in other sensitive marine areas, is potentially a significant new proxy for paleoproductivity and oceanic global events, such as the Miocene events, which are usually recognized in deep-sea pelagic sediments and high latitude deposits.

Age determination, sedimentological, biogeochemical, and plant macrofossils analyses of outcrop PG2038-1

Thermokarst lakes are a widespread feature of the Arctic tundra, in which highly dynamic processes are closely connected with current and past climate changes. We investigated late Quaternary sediment dynamics, basin and shoreline evolution, and environmental interrelations of Lake El'gene-Kyuele in the NE Siberian Arctic (latitude 71°17'N, longitude 125°34'E). The water-body displays thaw-lake characteristics cutting into both Pleistocene Ice Complex and Holocene alas sediments. Our methods are based on grain size distribution, mineralogical composition, TOC/N ratio, stable carbon isotopes and the analysis of plant macrofossils from a 3.5-m sediment profile at the modern eastern lake shore. Our results show two main sources for sediments in the lake basin: terrigenous diamicton supplied from thermokarst slopes and the lake shore, and lacustrine detritus that has mainly settled in the deep lake basin. The lake and its adjacent thermokarst basin rapidly expanded during the early Holocene. This climatically warmer than today period was characterized by forest or forest tundra vegetation composed of larches, birch trees and shrubs. Woodlands of both the HTM and the Late Pleistocene were affected by fire, which potentially triggered the initiation of thermokarst processes resulting later in lake formation and expansion. The maximum lake depth at the study site and the lowest limnic bioproductivity occurred during the longest time interval of ~7 ka starting in the Holocene Thermal Maximum and lasting throughout the progressively cooler Neoglacial, whereas partial drainage and an extensive shift of the lake shoreline occurred ~0.9 cal. ka BP. Correspondingly, this study discusses different climatic and environmental drivers for the dynamics of a thermokarst basin.

Electron spin resonance (ESR) and 14C dating in sediments of Bonaire, Leeward Antilles

The coastal deposits of Bonaire, Leeward Antilles, are among the most studied archives for extreme-wave events (EWEs) in the Caribbean. Here we present more than 400 electron spin resonance (ESR) and radiocarbon data on coarse-clast deposits from Bonaire's eastern and western coasts. The chronological data are compared to the occurrence and age of fine-grained extreme-wave deposits detected in lagoons and floodplains. Both approaches are aimed at the identification of EWEs, the differentiation between extraordinary storms and tsunamis, improving reconstructions of the coastal evolution, and establishing a geochronological framework for the events. Although the combination of different methods and archives contributes to a better understanding of the interplay of coastal and archive-related processes, insufficient separation, superimposition or burying of coarse-clast deposits and restricted dating accuracy limit the use of both fine-grained and coarse-clast geoarchives to unravel decadal- to centennial-scale events. At several locations, distinct landforms are attributed to different coastal flooding events interpreted to be of tsunamigenic origin. Coastal landforms on the western coast have significantly been influenced by (sub)-recent hurricanes, indicating that formation of the coarse-clast deposits on the eastern coast is likely to be related to past events of higher energy.

Sedimentological, petrographical and biostratigraphical characteristics of ODP Leg 190 volcanic ashes and siliceous claystones

Leg 190 was the first of a two-leg program across the Nankai accretionary prism and Trough, offshore Japan, aiming to evaluate existing models for prism evolution and to constrain syntectonic sedimentation, deformation styles, mechanical properties, and prism hydrology (Moore, Taira, Klaus, et al., 2001; Moore et al., 2001). More than 400 volcanic ash and siliceous claystone (altered ash) layers were penetrated and sampled during drilling of the six sites from two transects across the accretionary prism (Sites 1173-1178). In sites from the subducting Shikoku Basin (Sites 1173 and 1177) and in the trench axis (Site 1174), recognition of ash layers and diagenetically altered ashes was initially important in defining major lithostratigraphic units. However, it is clear that understanding the diagenesis of the volcanic ashes has considerable implications for prism evolution, mechanical properties, prism hydrology, geochemistry, and fluid flow in the accretionary prism and associated subducting sediments (cf. Masuda et al., 1996, doi 10.1346/CCMN.1996.0440402). Particle size, chemical composition, temperature, depth of burial, and time are all thought to be factors that may affect volcanic ash diagenesis and preservation (Kuramoto et al., 1992, doi:10.2973/odp.proc.sr.127128-2.235.1992; Underwood et al., 1993, doi:10.2973/odp.proc.sr.131.137.1993). The overall aim of this research is to evaluate factors influencing volcanic ash diagenesis in the Nankai Trough area. This data report presents just the results of the sedimentological and petrographic analysis of the volcanic ashes and siliceous claystones from Sites 1173, 1174, and 1177. It is anticipated that when the results of additional geochemical analysis of these lithologies is available a more meaningful evaluation of factors influencing volcanic ash alteration will be possible.

Documentation, profile descriptions and analytical results of loes profiles from Bavaria, southern Germany

This study investigates the landscape evolution and soil development in the loess area near Regensburg between approximately 6000-2000 yr BP (radiocarbon years), Eastern Bavaria. The focus is on the question how man and climate influenced landscape evolution and what their relative significance was. The theoretical background concerning the factors that controlled prehistoric soil erosion in Middle Europe is summarized with respect to rainfall intensity and distribution, pedogenesis, Pleistocene relief, and prehistoric farming. Colluvial deposits , flood loams, and soils were studied at ten different and representative sites that served as archives of their respective palaeoenvironments. Geomorphological, sedimentological, and pedological methods were applied. According to the findings presented here, there was a high asynchronity of landscape evolution in the investigation area, which was due to prehistoric land-use patterns. Prehistoric land use and settlement caused highly difIerenciated phases of morphodynamic activity and stability in time and space. These are documented at the single catenas ofeach site. In general, Pleistocene relief was substantially lowered. At the same time smaller landforms such as dells and minor asymmetric valleys filled up and strongly transformed. However, there were short phases at many sites, forming short lived linear erosion features ('Runsen'), resulting from exceptional rainfalls. These forms are results of single events without showing regional trends. Generally, the onset of the sedimentation of colluvial deposits took place much earlier (usually 3500 yr BP (radiocarbon) and younger) than the formation of flood loams. Thus, the deposition of flood loams in the Kleine Laaber river valley started mainly as a consequence of iron age farming only at around 2500 yr BP (radiocarbon). A cascade system explains the different ages of colluvial deposits and flood loams: as a result of prehistoric land use, dells and other minor Pleistocene landforms were filled with colluvial sediments. After the filling of these primary sediment traps , eroded material was transported into flood plains, thus forming flood loams. But at the moment we cannot quantify the extent ofprehistoric soil erosion in the investigation area. The three factors that controlled the prehistoric Iandscapc evolution in the Ioess area near Regensburg are as follows: 1. The transformation from a natural to a prehistoric cultural landscape was the most important factor: A landscape with stable relief was changed into a highly morphodynamic one with soil erosion as the dominant process of this change. 2. The sediment traps of the pre-anthropogenic relief determined where the material originated from soil erosion was deposited: either sedimentation took place on the slopes or the filled sediment traps of the slopes rendered flood loam formation possible. Climatic influence of any importance can only be documented as the result of land use in connection with singular and/or statistic events of heavy rainfalls. Without human impact, no significant change in the Holocene landscape would have been possible.

Late Quaternary carbonate and isotope stratigraphy of DSDP Site 90-594 east of South Island, New Zealand

The late Quaternary sequence off eastern South Island, New Zealand, consists of ~100 m of alternating bluish gray pelagic oozes and greenish gray hemipelagic oozes that extend uninterruptedly back to the Brunhes/Matuyama boundary (0.73 m.y.). A very high resolution (~2400 yr.) record of sediment texture, calcium carbonate content, and planktonic and benthic foraminiferal oxygen and carbon isotope composition demonstrates an in-phase cyclical fluctuation between the sedimentary parameters that closely correspond to the pelagic-hemipelagic sedimentation cycles and the isotope composition. Pelagic oozes, formed during interglacial periods of high eustatic sea level, are characterized by calcareous microfossils, relative enrichment in sand and clay sizes, high carbonate contents, reduced delta18O values, and increased delta13C values. Hemipelagic oozes, associated with glacial episodes and lowered eustatic sea level, include common terrigenous material and siliceous microfossils, are enriched in silt sizes, have low carbonate contents, high delta18O values, and low delta13C values. The history of alpine glaciations and associated erosion of the South Island of New Zealand, as expressed by the appearance of hemipelagic oozes, can be correlated directly with the major fluctuations of Northern Hemisphere ice sheets as expressed by the influence of eustatic sea-level changes on the oxygen isotope composition of both planktonic and benthic foraminifers. This high-accumulation-rate record contains conspicuous intervals of highfrequency, high-amplitude isotope variability including the presence of multiple glacial/interglacial intervals within single isotope stages, and offers one of the best sections cored to date for detailed study of the evolution and history of climate change over the last 0.75 m.y.

Proportion of bryozoa, isopoda and ostracoda, and bryozoan species richness on the Antarctic continental shelf, slope and abyss

The Antarctic continental slope spans the depths from the shelf break (usually between 500 and 1000 m) to ~3000 m, is very steep, overlain by 'warm' (2-2.5 °C) Circumpolar Deep Water (CDW), and life there is poorly studied. This study investigates whether life on Antarctica's continental slope is essentially an extension of the shelf or the abyssal fauna, a transition zone between these or clearly distinct in its own right. Using data from several cruises to the Weddell Sea and Scotia Sea, including the ANDEEP (ANtarctic benthic DEEP-sea biodiversity, colonisation history and recent community patterns) I-III, BIOPEARL (Biodiversity, Phylogeny, Evolution and Adaptive Radiation of Life in Antarctica) 1 and EASIZ (Ecology of the Antarctic Sea Ice Zone) II cruises as well as current databases (SOMBASE, SCAR-MarBIN), four different taxa were selected (i.e. cheilostome bryozoans, isopod and ostracod crustaceans and echinoid echinoderms) and two areas, the Weddell Sea and the Scotia Sea, to examine faunal composition, richness and affinities. The answer has important ramifications to the link between physical oceanography and ecology, and the potential of the slope to act as a refuge and resupply zone to the shelf during glaciations. Benthic samples were collected using Agassiz trawl, epibenthic sledge and Rauschert sled. By bathymetric definition, these data suggest that despite eurybathy in some of the groups examined and apparent similarity of physical conditions in the Antarctic, the shelf, slope and abyssal faunas were clearly separated in the Weddell Sea. However, no such separation of faunas was apparent in the Scotia Sea (except in echinoids). Using a geomorphological definition of the slope, shelf-slope-abyss similarity only changed significantly in the bryozoans. Our results did not support the presence of a homogenous and unique Antarctic slope fauna despite a high number of species being restricted to the slope. However, it remains the case that there may be a unique Antarctic slope fauna, but the paucity of our samples could not demonstrate this in the Scotia Sea. It is very likely that various ecological and evolutionary factors (such as topography, water-mass and sediment characteristics, input of particulate organic carbon (POC) and glaciological history) drive slope distinctness. Isopods showed greatest species richness at slope depths, whereas bryozoans and ostracods were more speciose at shelf depths; however, significance varied across Weddell Sea and Scotia Sea and depending on bathymetric vs. geomorphological definitions. Whilst the slope may harbour some source populations for localised shelf recolonisation, the absence of many shelf species, genera and even families (in a poorly dispersing taxon) from the continental slope indicate that it was not a universal refuge for Antarctic shelf fauna.