Pollen records from Elbe-Weser Dreieck

A long-running interdisciplinary research project on the development of landscape, prehistoric habitation and the history of vegetation within a "siedlungskammer" (limited habitation areal from neolithic to modern times has been carried out in the NW German lowlands, The siedlungskammer Flögeln is situated between the rivers Weser and EIbe and comprises about 23.5 km^2. It is an isolated pleistocene area surrounded by bogs, the soils consisting mainly of poor sands. In this siedlungskammer large-seale archaeological excavations and mappings have been performed, parallel to pedological, historical and above all pollen analytical investigations. The aim of the project is to record the individual phases in time, to delimit the respective settlement areas and to reconstruct the conditions of life and economy for each time period. A dense network of 10 pollen diagrams has been constructed. Several of them derive from the marginal area and from the centre of the large raised bog north of the siedlungskammer. These diagrams reflect the history of vegetation and habitation of a large region; due to the large pollen source area the habitation phases in the diagrams are poorly defined. Even in the utmost marginal diagram of this woodless bog, a great village with adjoining fields, situated only 100 m away from it, is registered with only low values of anthropogenic indicators. In contrast to this, the numerous pollen diagrams from kettle-hole bogs inside the siedlungskammer yield an exact picture of the habitation of the siedlungskammer and their individual parts. Early traces of habitation can be identified in the pollen diagram soon after the elm decline (around 5190 BP). Some time later in the middle neolithic period there follows a marked habitation phase, which starts between 4500 and 4400 BP and reflects the immigration of the trichterbecher culture. It corresponds to the landnam phase of Iversen in Denmark and begins with a sharp decline of the pollen curves of lime and oak, followed by the increase of anthropogenic indicators pointing to arable and pastural farming. High values of wild grasses and Calluna witness extensive forest grazing. This middle to late neolithic habitation is also registered archaeologically by settlements and numerous graves. After low human activity during Bronze Age and Older Iron Age times the archaeological and pollen analytical records of Roman and Migration periods is again very strong. This is followed by a gap in habitation during the 6th and 7th centuries and afterwards in the western part of the siedlungskammer from about 700 AD until the 14th century by the activity of the medieval village of Dalem, that was also excavated and whose fields were recorded by phosphate mapping to a size of 117 hectares. This medieval settlement phase is marked by much cereal cultivation (mainly rye). The dense network of pollen diagrams offers an opportunity to register the dispersion of the anthropogenic indicators from the areas of settlement to different distances and thus to obtain quantitative clues for the assessment of these anthropogenic indicators in pollen diagrams. In fig. 4 the reflection of the neolithic culture in the kettle-hole bogs and the large raised bog is shown in 3 phases: a) pre landnam, b) TRB-landnam, c) post landnam. Among arboreal pollen the reaction of Quercus is sharp close to the settlement but is not found at more distant profiles, whilst in contrast to this Tilia shows a significant decline even far away from the settlements. The record of most anthropogenic indicators outside the habitation area is very low, in particular cereal pollen is poorly dispersed; much more certain as an indicator for habitation (also for arable farming!) is Plantago lanceolata. A strong increase of wild grasses (partly Calluna aswell) some distance from the habitation areas indicates far reaching forest grazing. Fig. 5 illustrates the reflection of the anthropogenie indicators from the medieval village Dalem. In this instance the field area could be mapped exactly using phosphate investigations, and it has been possible to indicate the precise distances of the profile sites from the medieval fields. Here also, there is a clear correlation between decreasing anthropogenic indicators and increasing distance. In a kettle-hole bog (FLH) a distance of 3000 m away this marked settlement phase is not registered. The contrast between the pollen diagrams SWK and FLH (fig. 2 + 3, enclosure), illustrates the strong differences between diagrams from kettlehole bogs close to and distant from the settlements, for the neolithic as well as for the medieval period. On the basis of the examples presented here, implications concerning the interpretation of pollen diagrams with respect to habitation phases are discussed.

Biological traits, and egg and fecal pellet production of Calanus finmarchicus and Calanus glacialis in situ and during experiments

The effects of temperature and food was examined for Calanus finmarchicus and C. glacialis during 3 phases of the phytoplankton spring bloom in Disko Bay, western Greenland. The 2 species were collected during pre-bloom, bloom, and post-bloom and exposed to temperatures from 0 to 10°C, combined with deficient or excess food. Fecal pellet and egg production were measured as indices for grazing and secondary production, respectively. Furthermore, changes in body carbon, nitrogen, and lipid content were measured. C. glacialis sampled before the bloom and incubated with excess food exhibited high specific egg production at temperatures between 0 and 2.5°C. Higher temperatures did not increase egg production considerably, whereas egg production for C. finmarchicus more than tripled between 2.5 and 5°C. Starved C. glacialis produced eggs at all temperatures stimulated by increasing temperatures, whereas starved C. finmarchicus needed temperatures above 5°C to produce eggs fueled by their lipid stores. Few C. finmarchicus had mature gonads at the initiation of the pre-bloom and bloom experiment, and egg production of C. finmarchicus therefore only increased as the ratio of individuals with mature gonads increased. During the bloom, both C. glacialis and C. finmarchicus used the high food availability for egg production, while refueling or exhausting their lipid stores, respectively. Finally, during the post-bloom experiment, production was low by C. finmarchicus, whereas C. glacialis had terminated production. Our results suggest that a future warmer ocean will reduce the advantage of early spawning by C. glacialis and that C. finmarchicus will become increasingly prevalent.

Characteristics of eggs and female adults of Calanus finmarchicus, C. glacialis and C. hyperboreus from Disko Bay, western Greenland

This is the first high temporal-resolution study in Disko Bay covering population dynamics, grazing, reproduction, and biochemical composition of 3 dominating copepod species (Calanus finmarchicus, C. glacialis and C. hyperboreus) from late winter to midsummer in 2008. C. finmarchicus and C. glacialis ascended to the surface layer at the onset of the spring phytoplankton bloom, followed by C. hyperboreus 2 wk later. C. finmarchicus spawning occurred during the bloom and postbloom period, partially fueled by wax esters. C. glacialis commenced spawning before the bloom, yet it was greatly stimulated when food became available. However, feeding and reproduction was terminated after the main bloom despite the presence of food. In terms of feeding, this was also the strategy for C. hyperboreus. Between pre-bloom and post-bloom, C. finmarchicus showed an increase in carbon, nitrogen, and phospholipid content but a decrease in total lipid content. This was likely the result of protein synthesis, oocyte maturation, and spawning fueled by wax esters and by feeding. C. glacialis showed a similar pattern, although with an increasing total lipid content from pre-bloom to post-bloom, and an increasing wax ester and decreasing phospholipid content after reproduction was terminated. C. hyperboreus showed greatly increased content of carbon, nitrogen, and all lipid classes between the pre- and post-bloom periods. Hence, C. finmarchicus commenced feeding and spawning at the onset of the bloom and continued throughout the remaining study period. Both C. glacialis and C. hyperboreus females refueled their storage lipids (wax esters) during the bloom and post-bloom period, suggesting that they may spawn in an additional year.

Nannofossil assemblages of mid-Pliocene sediments from the equatorial Atlantic

Downcore cyclic variation in high-resolution nannofossil abundance records from mid-Pliocene equatorial Atlantic ODP Sites 662 and 926 demonstrate the direct response by several Pliocene taxa (notably Discoaster, Sphenolithus and Florisphaera profunda) to orbitally forced climatic variation. In particular, these records display strong obliquity and precessional signals reflecting primarily high latitude, Southern hemisphere changes influencing upwelling intensity and local low-latitude, insolation-driven climatic changes (via the productivity and/or turbidity influence of Amazon-sourced terrigenous material) at Sites 622 and 926 respectively. In seasonal studies of coccolithophorid assemblages, only part of the variation observed can be explained by abiotic processes, so it is perhaps not surprising that in this study few Pliocene nannofossil taxa demonstrate significant correlations with each other or with physical environmental parameters. Only some variance in nannofossil abundances can be explained by the primary controls of temperature and productivity. The rest is attributed to nonlinear responses to climatic changes; biotic processes such as grazing, predation, viral infection and competition, and/or, abiotic factors for which there is no readily available proxy (e.g. salinity). The lack of strong, consistent intra- and inter-relationships of the nannoflora and the environment reflects an ecologically complex, differentiated original community producing a complex integrated signal transmitted into the fossil record.

Biomass of nanoprotozooplankton in the Atlantic sector of the Southern Ocean

The dynamic of early spring nanoprotozoa was investigated in three characteristic water masses of the Southern Ocean: the Marginal Ice Zone, the intermediate waters of the Antarctic Circumpolar Current and the Polar Frontal Zone. Biomass and feeding activities of nanoprotozoa were measured, as well as the biomass of their potential prey-bacteria and phototrophic flagellates-on the 6°W meridian in the Southern Ocean along three repetitive transects between 47 and 60° South from October to November 1992. On average, nanoprotozooplankton biomass accounted for 77% of the combined biomass of bacteria and phototrophic flagellates, and was dominated by dinoflagellates and flagellates smaller than 5 µm. As a general trend, low protozoan biomass of 2 mg C/m**3 was typical of the ice covered area, while significantly higher biomasses culminating at 15 mg C/m**3 were recorded at the Polar Front. Biomasses of bacteria and total phytoplankton were distributed accordingly, with larger values at the Polar Front. Phototrophic flagellates did not show any geographical trend. No seasonal trend could be identified in the Marginal Ice Zone and in the intermediate waters of the Antarctic Circumpolar Current. On the other hand, at the Polar Front region a three-fold increase was observed within a 2-month period for nanoprotozooplankton biomass. Such a biomass increase was also detected for bacterioplankton and total phytoplankton biomass. Half-saturation constants and maximum specific ingestion of nanoprotozoan taxons feeding on bacteria and phototrophic flagellates were determined using the technique of fluorescent labelled bacteria (FLB) and algae (FLA) over a large range of prey concentrations. Maximum ingestion rates ranged between 0.002 and 0.015/h for bactivorous nanoprotozoa and heterotrophic flagellates larger than 5 µm feeding on phototrophic flagellates. The markedly high maximum ingestion rates of 0.4/h characterising nanophytoplankton ingestion by dinoflagellates evidenced the strong ability of dinoflagellates for feeding on nanophytoplankton. Daily ingestion rates were calculated from nanoprotozoan grazing parameters and carbon biomass of prey and predators. This indicated that nanoprotozoa ingestion of daily bacterioplankton and phytoplankton production in early spring ranged from 32 to 40%.