(Table 2) Metazoan meiofauna abundance in sediments obtained during POLARSTERN cruise ANT-XIX/3 (ANDEEP-1)

Meiofauna standing stocks and community structure are reported for the first time for abyssal soft-sediment samples in Antarctic waters. At seven stations within a depth range of 2274-5194 m a total of 128 sediment cores were retrieved with a multiple corer (MUC) on board of the R.V. Polarstern during the ANDEEP-1 cruise (ANT XIX/3). The metazoan meiofauna (defined by a lower size limit of 40 µm) was identified and counted, and one core per station was preserved for CPE, C/N, TOM and grain size analyses. Meiofauna densities are in the range of 2731 Ind./10 cm² at 2290 m depth and 75 Ind./10 cm² at 3597 m depth, with nematodes being the dominant group at all stations. Nematodes account for 84-94% followed by copepods with 2-8% of the total meiofauna. Other frequent taxa found at each station are kinorhynchs, loriciferans, tantulocarids, ostracods and tardigrades. There is a general tendency of decreasing abundances of metazoan meiofauna with increasing depth, but not all higher level taxa displayed this pattern. In addition, a tendency of decreasing higher taxon density with increasing depth was observed. Standing stocks are higher than the average found at similar depths in other oceans.

(Table 1) Ice station data during POLARSTERN expeditions ARK-XVIII/2 and ARK-XIX/1 to Fram Strait, the western Barents Sea and north of Svalbard

During two expeditions of the R.V. "Polarstern" to the Arctic Ocean, pack ice and under-ice water samples were collected during two different seasons: late summer (September 2002) and late winter (March/April 2003). Physical and biological properties of the ice were investigated to explain seasonal differences in species composition, abundance and distribution patterns of sympagic meiofauna (in this case: heterotrophs >20 µm). In winter, the ice near the surface was characterized by extreme physical conditions (minimum ice temperature: -22°C, maximum brine salinity: 223, brine volume: <=5%) and more moderate conditions in summer (minimum ice temperature: -5.6°C, maximum brine salinity: 94, most brine volumes: >=5%). Conditions in the lowermost part of the ice did not differ to a high degree between summer and winter. Chlorophyll a concentrations (chl a) showed significant differences between summer and winter: during winter, concentrations were mostly <1.0 µg chl a/l, while chl a concentrations of up to 67.4 µmol/l were measured during summer. The median of depth-integrated chl a concentration in summer was significantly higher than in winter. Integrated abundances of sympagic meiofauna were within the same range for both seasons and varied between 0.6 and 34.1×103 organisms /m**2 in summer and between 3.7 and 24.8×10**3 organisms /m**2 in winter. With regard to species composition, a comparison between the two seasons showed distinct differences: while copepods (42.7%) and rotifers (33.4%) were the most abundant sea-ice meiofaunal taxa during summer, copepod nauplii dominated the community, comprising 92.9% of the fauna, in winter. Low species abundances were found in the under-ice water, indicating that overwintering of the other sympagic organisms did not take place there, either. Therefore, their survival strategy over the polar winter remains unclear.

Spatial Relationships among Hydroacoustic, Hydrographic and Top Predator Patterns: Cetacean Distributions in the Mid-Atlantic Bight

Effective conservation and management of top predators requires a comprehensive understanding of their distributions and of the underlying biological and physical processes that affect these distributions. The Mid-Atlantic Bight shelf break system is a dynamic and productive region where at least 32 species of cetaceans have been recorded through various systematic and opportunistic marine mammal surveys from the 1970s through 2012. My dissertation characterizes the spatial distribution and habitat of cetaceans in the Mid-Atlantic Bight shelf break system by utilizing marine mammal line-transect survey data, synoptic multi-frequency active acoustic data, and fine-scale hydrographic data collected during the 2011 summer Atlantic Marine Assessment Program for Protected Species (AMAPPS) survey. Although studies describing cetacean habitat and distributions have been previously conducted in the Mid-Atlantic Bight, my research specifically focuses on the shelf break region to elucidate both the physical and biological processes that influence cetacean distribution patterns within this cetacean hotspot.

In Chapter One I review biologically important areas for cetaceans in the Atlantic waters of the United States. I describe the study area, the shelf break region of the Mid-Atlantic Bight, in terms of the general oceanography, productivity and biodiversity. According to recent habitat-based cetacean density models, the shelf break region is an area of high cetacean abundance and density, yet little research is directed at understanding the mechanisms that establish this region as a cetacean hotspot.

In Chapter Two I present the basic physical principles of sound in water and describe the methodology used to categorize opportunistically collected multi-frequency active acoustic data using frequency responses techniques. Frequency response classification methods are usually employed in conjunction with net-tow data, but the logistics of the 2011 AMAPPS survey did not allow for appropriate net-tow data to be collected. Biologically meaningful information can be extracted from acoustic scattering regions by comparing the frequency response curves of acoustic regions to theoretical curves of known scattering models. Using the five frequencies on the EK60 system (18, 38, 70, 120, and 200 kHz), three categories of scatterers were defined: fish-like (with swim bladder), nekton-like (e.g., euphausiids), and plankton-like (e.g., copepods). I also employed a multi-frequency acoustic categorization method using three frequencies (18, 38, and 120 kHz) that has been used in the Gulf of Maine and Georges Bank which is based the presence or absence of volume backscatter above a threshold. This method is more objective than the comparison of frequency response curves because it uses an established backscatter value for the threshold. By removing all data below the threshold, only strong scattering information is retained.

In Chapter Three I analyze the distribution of the categorized acoustic regions of interest during the daytime cross shelf transects. Over all transects, plankton-like acoustic regions of interest were detected most frequently, followed by fish-like acoustic regions and then nekton-like acoustic regions. Plankton-like detections were the only significantly different acoustic detections per kilometer, although nekton-like detections were only slightly not significant. Using the threshold categorization method by Jech and Michaels (2006) provides a more conservative and discrete detection of acoustic scatterers and allows me to retrieve backscatter values along transects in areas that have been categorized. This provides continuous data values that can be integrated at discrete spatial increments for wavelet analysis. Wavelet analysis indicates significant spatial scales of interest for fish-like and nekton-like acoustic backscatter range from one to four kilometers and vary among transects.

In Chapter Four I analyze the fine scale distribution of cetaceans in the shelf break system of the Mid-Atlantic Bight using corrected sightings per trackline region, classification trees, multidimensional scaling, and random forest analysis. I describe habitat for common dolphins, Risso’s dolphins and sperm whales. From the distribution of cetacean sightings, patterns of habitat start to emerge: within the shelf break region of the Mid-Atlantic Bight, common dolphins were sighted more prevalently over the shelf while sperm whales were more frequently found in the deep waters offshore and Risso’s dolphins were most prevalent at the shelf break. Multidimensional scaling presents clear environmental separation among common dolphins and Risso’s dolphins and sperm whales. The sperm whale random forest habitat model had the lowest misclassification error (0.30) and the Risso’s dolphin random forest habitat model had the greatest misclassification error (0.37). Shallow water depth (less than 148 meters) was the primary variable selected in the classification model for common dolphin habitat. Distance to surface density fronts and surface temperature fronts were the primary variables selected in the classification models to describe Risso’s dolphin habitat and sperm whale habitat respectively. When mapped back into geographic space, these three cetacean species occupy different fine-scale habitats within the dynamic Mid-Atlantic Bight shelf break system.

In Chapter Five I present a summary of the previous chapters and present potential analytical steps to address ecological questions pertaining the dynamic shelf break region. Taken together, the results of my dissertation demonstrate the use of opportunistically collected data in ecosystem studies; emphasize the need to incorporate middle trophic level data and oceanographic features into cetacean habitat models; and emphasize the importance of developing more mechanistic understanding of dynamic ecosystems.

Distribution of meiobenthos in sediment samples of the northeastern Atlantic

The meiofauna of the deep sea areas (800 - 5500 m) between Madeira and Lisbon was quantitatively investigated during "Meteor" cruises in 1970 and 1971. With respect to numbers and biomass the meiofauna (especially nematodes and harpacticoid copepods) of the investigated areas is relatively poor averaging about 66,000 individuals per m**2 and 34 mg per m**2 wet weight biomass (polychaetes and foraminifera excluded). Regional differences are more pronounced in the investigated areas than differences due to depth. A comparison with the results of other authors from other areas confirms the regional variations in the meiofauna abundance of the deep sea.

Results from a stable isotope tracer experiment in Potter Cove, King George Island, Antarctica

Antarctic meiofauna is still strongly understud- ied, and so is its trophic position in the food web. Primary producers, such as phytoplankton, and bacteria may repre- sent important food sources for shallow water metazoans, and the role of meiobenthos in the benthic-pelagic coupling represents an important brick for food web understanding. In a laboratory, feeding experiment 13C-labeled freeze- dried diatoms (Thalassiosira weissflogii) and bacteria were added to retrieved cores from Potter Cove (15-m depth, November 2007) in order to investigate the uptake of 3 main meiofauna taxa: nematodes, copepods and cumaceans. In the surface sediment layers, nematodes showed no real difference in uptake of both food sources. This outcome was supported by the natural delta 13C values and the community genus composition. In the first centimeter layer, the dominant genus was Daptonema which is known to be opportunistic, feeding on both bacteria and diatoms. Copepods and cumaceans on the other hand appeared to feed more on diatoms than on bacteria. This may point at a better adaptation to input of primary production from the water column. On the other hand, the overall carbon uptake of the given food sources was quite low for all taxa, indicating that likely other food sources might be of relevance for these meiobenthic organisms. Further studies are needed in order to better quantify the carbon requirements of these organisms.

Ocean acidification affects the phyto-zoo plankton trophic transfer efficiency

The critical role played by copepods in ocean ecology and biogeochemistry warrants an understanding of how these animals may respond to ocean acidification (OA). Whilst an appreciation of the potential direct effects of OA, due to elevated pCO2, on copepods is improving, little is known about the indirect impacts acting via bottom-up(food quality) effects. We assessed, for the first time, the chronic effects of direct and/or indirect exposures to elevated pCO2 on the behaviour, vital rates, chemical and biochemical stoichiometry of the calanoid copepod Acartia tonsa. Bottom-up effects of elevated pCO2 caused species-specific biochemical changes to the phytoplanktonic feed, which adversely affected copepod population structure and decreased recruitment by 30 %. The direct impact of elevated pCO2 caused gender-specific respiratory responses in A.tonsa adults, stimulating an enhanced respiration rate in males (> 2-fold), and a suppressed respiratory response in females when coupled with indirect elevated pCO2 exposures. Under the combined indirect+direct exposure, carbon trophic transfer efficiency from phytoplankton-to-zooplankton declined to < 50 % of control populations, with a commensurate decrease in recruitment. For the first time an explicit role was demonstrated for biochemical stoichiometry in shaping copepod trophic dynamics. The altered biochemical composition of the CO2-exposed prey affected the biochemical stoichiometry of the copepods, which could have ramifications for production of higher tropic levels, notably fisheries. Our work indicates that the control of phytoplankton and the support of higher trophic levels involving copepods have clear potential to be adversely affected under future OA scenarios.

Meiofauna of surface sediments in the Cretan Sea

Quantitative information on metazoan meiofaunal abundance and biomass was obtained from three continental shelf (at 40, 100 and 200 m depth) and four deep-sea stations (at 540, 700, 940 and 1540 m depth) in the Cretan Sea (South Aegean Sea, NE Mediterranean). Samples were collected on a seasonal basis (from August 1994 to September 1995) with the use of a multiple corer. Meiofaunal abundance and biomass on the continental shelf of the Cretan Sea were high, in contrast to the extremely low values reported for the bathyal sediments that showed values comparable to those reported for abyssal and hadal environments. In order to explain the spatial and seasonal changes in metazoan meiofauna these data were compared with: (1) the concentrations of 'food indicators' (such as proteins, lipids, soluble carbohydrates and CPE) (2) the bacterial biomass (3) the flux of labile organic compounds to the sea floor at a fixed station (D7, 1540 m depth). Highly significant relationships between meiofaunal parameters and CPE, protein and lipid concentrations and bacterial biomass were found. Most of the indicators of food quality and quantity (such as CPE, proteins and carbohydrates) showed a clear seasonality with highest values in February and lowest in September. Such changes were more evident on the continental shelf rather than at deeper depths. On the continental shelf, significant seasonal changes in meiofaunal density were related to changes in the input of labile organic carbon whereas meiofaunal assemblages on the deep-sea stations showed time-lagged changes in response to the food input recorded in February 95. At all deep-sea stations meiofaunal density increased with a time lag of 2 months. Indications for a time-lagged meiofaunal response to the food inputs were also provided by the increase in nauplii densities during May 95 and the increase in individual biomass of nematodes, copepods and polychaetes between February and May 1995. The lack of strong seasonal changes in deep sea meiofaunal density suggests that the supply of organic matter below 500 m is not strong enough to support a significant meiofaunal development. Below 700 m depth >92% of the total biomass in the sediment was represented by bacteria. The ratio of bacterial to meiofaunal biomass increased with increasing water depth indicating that bacteria are probably more effective than meiofauna in exploiting refractory organic compounds. These data lead us to hypothesise that the deep-sea sediments of the Cretan Sea are largely dependent upon a benthic microbial loop.

Relative abundance of euphausiids (Crustacea) in water samples of the Arabian Sea (Table 2)

In the present paper, the ecology and feeding habits of euphausiids are described. The samples were taken at the time of the NE-monsoon (1964/65) by R. V. "Meteor" in the Arabian Sea and adjacent waters. 24 species were determined. According to distribution of the species, the following marine areas can be distinguished: Arabian Sea: 24 species, dominant are Euphausia diomedeae, E. tenera, E. distinguenda, Stylocheiron carinatum. Gulf of Aden: 10 species, dominant are Euphausia diomedeae, E. distinguenda. Red Sea: 6 species, dominant are Euphausia diomedeae, E. distinguenda. Gulf of Oman : 5 Species, dominant are Euphausia distinguenda, Pseudeupbaufia latifrons. Persian Gulf: 1 species - Pseudeuphausia latifrons. The total number of euphausiids indicate the biomass of this group. High densities of euphausiids (200-299 and > 300 individuals/100 m**3) occur in the innermost part of the Gulf cf Aden, in the area south of the equator near the African east coast, near Karachi (Indian west coast) and in the Persian Gulf. Comparison with data relating to production biology confirms that these are eutrophic zones which coincide with areas in which upwelling occurs at the time of the NE-monsoon. The central part of the Arabian Sea differs from adjacent waters by virtue of less dense euphausiid populations (> 199 individuals/100 m**3). Measurements relating to production biology demonstrate a relatively low concentration of primary food sources. Food material was ascertained by analysis of stomach content. The following omnivorous species were examined: Euphausia diomedeae, E. distinguenda, E. tenera, Pseudeuphausia latifrons and Thysanopoda tricuspidata. Apart from crustacean remains large numbers of Foraminifera, Radiolaria, tintinnids, dinoflagellates were found in the stomachs. Quantitatively crustaceans form the most important item in the diet. Food selection on the basis of size and form appears to be restricted to certain genera of tintinnids. The genera Stylocheiron and Nematoscelis are predators. Only crustacean remains were found in the stomachs of Stylocheiron abbreviatum, whereas Radiolaria, Foraminifera and tintinnids occurred to some extent in Nematasceli sp. Different euphausiids in the food chain in the Arabian Sea. In omnivorous species the position is variable, since they not only feed by filtering autotrophic and heterotrophic Protista, but also by predation on zooplankton. Carnivorous species without filtering apparatus feed exclusively on zooplankton of the size of copepods. Only these species are well established as occupying a higher position in the food chain. The parasitic protozoan Tbalassomyces fagei was found on Euphausia diomedeae, E. fenera, E. distinguenda and E. sanzoi.

ATP content in surface sediments of the Northeast Atlantic (Table 1)

1. ATP in deep-sea sediments can be determined after it is adsorbed on a mixture of the sediment and calcium carbonate by measuring the luminescence of the reaction of the mixture and luciferin-luciferase. 2. ATP contents of the toplayer of northeastern Atlantic sediments (Josephine Bank and northern Canary Basin) decrease with increasing depths of 252, 408, 1445, 1769, 2149, 4897, 5510m: 0.96, 0.61, 0.13, 0.10, 0.21, 0.05, 0.07 µg ATP/ml wet sediment. The decreasing values are in accordance with the decrease of macrobenthos and meiobenthos biomass in the deep-sea. 3. The ATP content of deep-sea nematodes is about 1 ? of their wet weight. 4. At the two deepest stations, less than 50% of the ATP measured in the sediment is represented by nematodes, copepods, other "hard" meiofauna groups and bacteria.

Seawater carbonate chemistry and biological processes of zooplankton Amphiascoides atopus and Schizopera knabeni during experiments, 2010

Increased atmospheric CO2 concentrations are causing greater dissolution of CO2 into seawater, and are ultimately responsible for today's ongoing ocean acidification. We manipulated seawater acidity by addition of HCl and by increasing CO2 concentration and observed that two coastal harpacticoid copepods, Amphiascoides atopus and Schizopera knabeni were both more sensitive to increased acidity when generated by CO2. The present study indicates that copepods living in environments more prone to hypercapnia, such as mudflats where S. knabeni lives, may be less sensitive to future acidification. Ocean acidification is also expected to alter the toxicity of waterborne metals by influencing their speciation in seawater. CO2 enrichment did not affect the free-ion concentration of Cd but did increase the free-ion concentration of Cu. Antagonistic toxicities were observed between CO2 with Cd, Cu and Cu free-ion in A. atopus. This interaction could be due to a competition for H+ and metals for binding sites.

Under-ice fauna, sea-ice concentration, sea-ice thickness and under-ice water parameters from the Eurasian central Arctic Ocean during POLARSTERN cruise ARK-XXVII/3 (PS80) in August-September 2012

Arctic sea-ice decline is expected to have a significant impact on Arctic marine ecosystems. Ice-associated fauna play a key role in this context because they constitute a unique part of Arctic biodiversity and transmit carbon from sea-ice algae into pelagic and benthic food webs. Our study presents the first regional-scale record of under-ice faunal distribution and the environmental characteristics of under-ice habitats throughout the Eurasian Basin. Sampling was conducted with a Surface and Under-Ice Trawl, equipped with a sensor array recording ice thickness and other physical parameters during trawling. We identified 2 environmental regimes, broadly coherent with the Nansen and Amundsen Basins. The Nansen Basin regime was distinguished from the Amundsen Basin regime by heavier sea-ice conditions, higher surface salinities and higher nitrate + nitrite concentrations. We found a diverse (28 species) under-ice community throughout the Eurasian Basin. Change in community structure reflected differences in the relative contribution of abundant species. Copepods (Calanus hyperboreus and C. glacialis) dominated in the Nansen Basin regime. In the Amundsen Basin regime, amphipods (Apherusa glacialis, Themisto libellula) dominated. Polar cod Boreogadus saida was present throughout the sampling area. Abrupt changes from a dominance of ice-associated amphipods at ice-covered stations to a dominance of pelagic amphipods (T. libellula) at nearby ice-free stations emphasised the decisive influence of sea ice on small-scale patterns in the surface-layer community. The observed response in community composition to different environmental regimes indicates potential long-term alterations in Arctic marine ecosystems as the Arctic Ocean continues to change.

Mesozooplankton in Bornholm Basin, Baltic Sea

The seasonal vertical distribution of mesozooplankton was investigated in the Bornholm Basin in October 1988, July 1991 and April 1992 by means of horizontal tows using a small net with a mesh size of 47 micrometers. This study shows that the vertical distribution of multivoltine copepods is linked with ontogeny and may also change seasonally, even in shallow areas such as the Baltic. Trends between copepod species, stages and seasons were discovered. Secondly, the correlation was investigated between the vertical mesozooplankton distribution on the one hand and chlorophyll a concentration and the density of medusae and fish larvae on the other hand. In several cases the comparison of the shape of the profiles led to greater differentiation than the coefficient of correlation. The abundances found allowed only scant statements about non-copepod taxa.

Sampling information, length measurements, DNA alignments, models and phylogenetics trees for Oithona similis s.l.

Traditionally, many small-sized copepod species are considered to be widespread, bipolar or cosmopolitan. However, these large-scale distribution patterns need to be re-examined in view of increasing evidence of cryptic and pseudo-cryptic speciation in pelagic copepods. Here, we present a phylogeographic study of Oithona similis s.l. populations from the Arctic Ocean, the Southern Ocean and its northern boundaries, the North Atlantic and the Mediterrranean Sea. O. similis s.l. is considered as one of the most abundant species in temperate to polar oceans and acts as an important link in the trophic network between the microbial loop and higher trophic levels such as fish larvae. Two gene fragments were analysed: the mitochondrial cytochrome oxidase c subunit I (COI), and the nuclear ribosomal 28S genetic marker. Seven distinct, geographically delimitated, mitochondrial lineages could be identified, with divergences among the lineages ranging from 8 to 24 %, thus representing most likely cryptic or pseudocryptic species within O. similis s.l. Four lineages were identified within or close to the borders of the Southern Ocean, one lineage in the Arctic Ocean and two lineages in the temperate Northern hemisphere. Surprisingly the Arctic lineage was more closely related to lineages from the Southern hemisphere than to the other lineages from the Northern hemisphere, suggesting that geographic proximity is a rather poor predictor of how closely related the clades are on a genetic level. Molecular clock application revealed that the evolutionary history of O. similis s.l. is possibly closely associated with the reorganization of the ocean circulation in the mid Miocene and may be an example of allopatric speciation in the pelagic zone.

Feeding of larval sardine Sardina pilchardus (Walbaum) off the north coast of Spain

In April and May 1991 and between March and June 1992 data regarding the diet of larval S. pilchardus in relation to food availability was gathered. Interpretation of results is compromised by the tendency of sardine larvae to defecate their gut contents during sampling. The most common food organisms in the guts (78-89%) were the developmental stages of copepods (eggs, nauplii and copepodites). Percentage composition of copepod nauplii in the diet decreased with increasing larval size, while copepodites increased. There was no consistent relationship between food availability and feeding success, probably because feeding conditions were generally adequate.

Warmer, deeper and greener mixed layers in the north Atlantic subpolar gyre over the last 50 years.

Shifts in global climate resonate in plankton dynamics, biogeochemical cycles, and marine food webs. We studied these linkages in the North Atlantic subpolar gyre (NASG), which hosts extensive phytoplankton blooms. We show that phytoplankton abundance increased since the 1960s in parallel to a deepening of the mixed layer and a strengthening of winds and heat losses from the ocean, as driven by the low frequency of the North Atlantic Oscillation (NAO). In parallel to these bottom-up processes, the top-down control of phytoplankton by copepods decreased over the same time period in the western NASG, following sea surface temperature changes typical of the Atlantic Multi-decadal Oscillation (AMO). While previous studies have hypothesized that climate-driven warming would facilitate seasonal stratification of surface waters and long-term phytoplankton increase in subpolar regions, here we show that deeper mixed layers in the NASG can be warmer and host a higher phytoplankton biomass. These results emphasize that different modes of climate variability regulate bottom-up (NAO control) and top-down (AMO control) forcing on phytoplankton at decadal timescales. As a consequence, different relationships between phytoplankton, zooplankton, and their physical environment appear subject to the disparate temporal scale of the observations (seasonal, interannual, or decadal). The prediction of phytoplankton response to climate change should be built upon what is learnt from observations at the longest timescales.