Deposit-feeding in benthic macrofauna : Tracer studies from the Baltic Sea

A low content of organic matter, which is largely refractory in nature, is characteristic of most sediments, meaning that aquatic deposit-feeders live on a very poor food source. The food is derived mainly from sedimenting phytodetritus, and in temperate waters like the Baltic Sea, from seasonal phytoplankton blooms. Deposit-feeders are either bulk-feeders, or selective feeders, which preferentially ingest the more organic-rich particles in the sediment, including phytodetritus, microbes and meiofauna. The soft-bottom benthos of the Baltic Sea has low species biodiversity and is dominated by a few macrobenthic species, among which the most numerous are the two deposit-feeding amphipods Monoporeia affinis and Pontoporeia femorata, and the bivalve Macoma balthica. This thesis is based on laboratory experiments on the feeding of these three species, and on the priapulid Halicryptus spinulosus. Feeding by benthic animals is often difficult to observe, but can be effectively studied by the use of tracers. Here we used the radioactive isotope 14C to label food items and to trace the organic matter uptake in the animals, while the stable isotopes 13C and 15N were used to follow feeding on aged organic matter in the sediment. The abundance of M. balthica and the amphipods tends to be negatively correlated, i.e., fewer bivalves are found at sites with dense populations of amphipods, with the known explanation that newly settled M. balthica spat are killed by the amphipods. Whether the postlarvae are just accidentally killed, or also ingested after being killed was tested by labelling the postlarvae with 14C and Rhodamine B. Both tracer techniques gave similar evidence for predation on and ingestion of postlarval bivalves. We calculated that this predation was likely to supply less than one percent of the daily carbon requirement for M. affinis, but might nevertheless be an important factor limiting recruitment of M. balthica. The two amphipods M. affinis and P. femorata are partly vertically segregated in the sediment, but whether they also feed at different depths was unknown. By adding fresh 14C-labelled algae either on the sediment surface or mixed into the sediment, we were able to distinguish surface from subsurface feeding. We found M. affinis and P. femorata to be surface and subsurface deposit-feeders, respectively. Whether the amphipods also feed on old organic matter, was studied by adding fresh 14C-labelled algae on the sediment surface, and using aged, one-year-old 13C- and 15N-labelled sediment as deep sediment. Ingestion of old organic matter, traced by the stable isotopes, differed between the two species, with a higher uptake for P. femorata, suggesting that P. femorata utilises the older, deeper-buried organic matter to a greater extent. Feeding studies with juveniles of both M. affinis and P. femorata had not been done previously. In an experiment with the same procedure and treatments as for the adults, juveniles of both amphipod species were found to have similar feeding strategies. They fed on both fresh and old sediment, with no partitioning of food resources, making them likely to be competitors for the same food resource. Oxygen deficiency has become more wide-spread in the Baltic Sea proper in the last half-century, and upwards of 70 000km2 are now devoid of macrofauna, even though part of that area does not have oxygen concentrations low enough to directly kill the macrofauna. We made week-long experiments on the rate of feeding on 14C-labelled diatoms spread on the sediment surface in different oxygen concentrations for both the amphipod species, M. balthica and H. spinulosus. The amphipods were the most sensitive to oxygen deficiency and showed reduced feeding and lower survival at low oxygen concentrations. M. balthica showed reduced feeding at the lowest oxygen concentration, but no mortality increase. The survival of H. spinulosus was unaffected, but it did not feed, showing that it is not a surface deposit-feeder. We conclude that low oxygen concentrations that are not directly lethal, but reduce food intake, may lead to starvation and death in the longer term.

Palaeoenvironment in North-Western Romania during the last 15 000 years

The objectives of this thesis are to establish a chronological framework for environmental changes during the last 15,000 years in northwest Romania, to reconstruct the vegetation development, and to evaluate the underlying processes for forest dynamics. Furthermore, an overview of earlier and ongoing pollenstratigraphic work in Romania is provided. Sediments from two former crater lakes, Preluca Tiganului and Steregoiu, situated in the Gutaiului Mountains, on the western extremity of the Eastern Carpathians at 730 m and 790 m a.s.l., respectively were obtained and analysed for high-resolution pollen, macrofossils, charcoal, mineral magnetic parameters and organic matter. The chronostratigraphic framework was provided by dense AMS 14C measurements. Cold and dry climatic conditions are indicated by the occurrence of open vegetation with shrubs and herbs, and cold lake water prior to 14,700 cal. yr BP. The climatic improvement at the beginning of the Lateglacial interstadial (around 14,700 cal. yr BP) is seen by the development of open forests. These were dominated by Pinus and Betula, but contained also new arriving tree taxa, such as Populus, Alnus and Prunus. The gradual establishment of forests may have led to a stabilization of the soils in the catchment. Between ca. 14,100 and 13,800 cal. yr BP the forest density became reduced to stands of Pinus, Betula, Alnus, Larix and Populus trees and grassland expanded, suggesting colder climatic conditions. Picea arrived as a new taxon at around 13,800 cal. yr BP, and between 13,800 and 12,900 cal. yr BP, the surroundings of the sites were predominantly covered by Picea forest. This forest included Betula, Pinus, Alnus, Larix and Populus and, from 13,200 cal. yr BP onwards also Ulmus. At ca. 12,900 cal. yr BP, the forest became significantly reduced and at 12,600 cal. yr BP, a recurrence of open vegetation with stands of Larix, Pinus, Betula, Salix and Alnus is documented, lasting until 11,500 cal. yr BP. This distinct change in vegetation may by taken as a strong decline in temperature and moisture availability. At the transition to the Holocene, at ca. 11,500 cal. yr BP, Pinus, Betula and Larix quickly expanded (from small local stands) and formed open forests, probably as a response to warmer and more humid climatic conditions. At 11,250 cal. yr BP Ulmus and Picea expanded and the landscape became completely forested. The rapid increase of Ulmus and Picea after 11,500 cal. yr BP may suggest the existence of small residual populations close to the study sites during the preceding cold interval. Ulmus was the first and most prominent deciduous taxa in the early Holocene in the Gutaiului Mountains. From ca. 10,750 cal. yr BP onwards Quercus, Tilia, Fraxinus and Acer expanded and Corylus arrived. A highly diverse, predominantly deciduous forest with Ulmus, Quercus, Tilia, Fraxinus, Acer, Corylus and Picea developed between 10,700 and 8200 cal. yr BP, which possibly signifies more continental climatic conditions. The development of a Picea-Corylus dominated forest between 8200 and 5700 cal. yr BP is likely connected to a more humid and cooler climate. The establishment of Carpinus and Fagus was dated to 5750 cal. yr BP and 5200 cal. yr BP, respectively. The dominance of Fagus during the late Holocene, from 4000 cal. yr BP onwards, may have been related to cooler and more humid climatic conditions. First signs of human activities are recorded around 2300 cal. yr BP, but only during the last 300 years did local human impact become significant. The vegetation development recorded in the Gutaiului Mountains during the Lateglacial is very similar to reconstructions based on lowland sites, whereas higher elevation sites seem not to have always experienced visible vegetation changes. The time of tree arrival and expansion during the past 11,500 cal. yr BP seems to have occurred almost synchronously across Romania. The composition of the forests during the Holocene in the Gutaiului Mountains is consistent with that reconstructed at mid-elevation sites, but differs from the forest composition at higher elevations. Important differences between the Gutaiului Mountains and other studied sites in Romania are a low representation of Carpinus and a late and weak human impact. The available data sets for Romania give evidence for the presence of coniferous and cold-tolerant deciduous trees before 14,700 cal. yr BP. Glacial refugia for Ulmus may have occurred in different parts of Romania, whereas the existence of Quercus, Tilia, Corylus and Fraxinus has not been corroborated.