Report of the 2005 Workshop on Ocean Ecodynamics Comparison in the Subarctic Pacific

I. Scientific Issues Posed by OECOS II. Participant Contributions to the OECOS Workshop A. ASPECTS OF PHYTOPLANKTON ECOLOGY IN THE SUBARCTIC PACIFIC Microbial community compositions by Karen E. Selph Subarctic Pacific lower trophic interactions: Production-based grazing rates and grazing-corrected production rates by Nicholas Welschmeyer Phytoplankton bloom dynamics and their physiological status in the western subarctic Pacific by Ken Furuya Temporal and spatial variability of phytoplankton biomass and productivity in the northwestern Pacific by Sei-ichi Saitoh, Suguru Okamoto, Hiroki Takemura and Kosei Sasaoka The use of molecular indicators of phytoplankton iron limitation by Deana Erdner B. IRON CONCENTRATION AND CHEMICAL SPECIATION Iron measurements during OECOS by Zanna Chase and Jay Cullen 25 The measurement of iron, nutrients and other chemical components in the northwestern North Pacific Ocean by Kenshi Kuma The measurement of iron, nutrients and other chemical components in the northwestern North Pacific Ocean by Kenshi Kuma C. PHYSICAL OCEANOGRAPHY, FINE-SCALE DISTRIBUTION PATTERNS AND AUTONOMOUS DRIFTERS The use of drifters in Lagrangian experiments: Positives, negatives and what can really be measured by Peter Strutton The interaction between plankton distribution patterns and vertical and horizontal physical processes in the eastern subarctic North Pacific by Timothy J. Cowles D. MICROZOOPLANKTON Microzooplankton processes in oceanic waters of the eastern subarctic Pacific: Project OECOS by Suzanne Strom Functional role of microzooplankton in the pelagic marine ecosystem during phytoplankton blooms in the western subarctic Pacific by Takashi Ota and Akiyoshi Shinada E. MESOZOOPLANKTON Vertical zonation of mesozooplankton, and its variability in response to food availability, density stratification, and turbulence by David L. Mackas and Moira Galbraith Marine ecosystem characteristics and seasonal abundance of dominant calanoid copepods in the Oyashio region by Atsushi Yamaguchi, Tsutomu Ikeda and Naonobu Shiga OECOS: Proposed mesozooplankton research in the Oyashio region, western subarctic Pacific by Tsutomu Ikeda Some background on Neocalanus feeding by Michael Dagg Size and growth of interzonally migrating copepods by Charles B. Miller Growth of large interzonal migrating copepods by Toru Kobari F. MODELING Ecosystem and population dynamics modeling by Harold P. Batchelder III. Reports from Workshop Breakout Groups A. PHYSICAL AND CHEMICAL ASPECTS WITH EMPHASIS ON IRON AND IRON SPECIATION B. PHYTOPLANKTON/MICROZOOPLANKTON STUDIES C. MESOZOOPLANKTON STUDIES IV. Issues arising during the workshop A. PHYTOPLANKTON STOCK VARIATIONS IN HNLC SYSTEMS AND TROPHIC CASCADES IN THE NANO AND MICRO REGIMES B. DIFFERENCES BETWEEN EAST AND WEST IN SITE SELECTION FOR OECOS TIME SERIES C. TIMING OF OECOS EXPEDITIONS D. CHARACTERIZATION OF PHYSICAL OCEANOGRAPHY V. Concluding Remarks VI. References (109 page document)

Considerations for Assessing Maximum Critical Temperatures in Small Ectothermic Animals: Insights from Leaf-Cutting Ants

The thermal limits of individual animals were originally proposed as a link between animal physiology and thermal ecology. Although this link is valid in theory, the evaluation of physiological tolerances involves some problems that are the focus of this study. One rationale was that heating rates shall influence upper critical limits, so that ecological thermal limits need to consider experimental heating rates. In addition, if thermal limits are not surpassed in experiments, subsequent tests of the same individual should yield similar results or produce evidence of hardening. Finally, several non-controlled variables such as time under experimental conditions and procedures may affect results. To analyze these issues we conducted an integrative study of upper critical temperatures in a single species, the ant Atta sexdens rubropiosa, an animal model providing large numbers of individuals of diverse sizes but similar genetic makeup. Our specific aims were to test the 1) influence of heating rates in the experimental evaluation of upper critical temperature, 2) assumptions of absence of physical damage and reproducibility, and 3) sources of variance often overlooked in the thermal-limits literature; and 4) to introduce some experimental approaches that may help researchers to separate physiological and methodological issues. The upper thermal limits were influenced by both heating rates and body mass. In the latter case, the effect was physiological rather than methodological. The critical temperature decreased during subsequent tests performed on the same individual ants, even one week after the initial test. Accordingly, upper thermal limits may have been overestimated by our (and typical) protocols. Heating rates, body mass, procedures independent of temperature and other variables may affect the estimation of upper critical temperatures. Therefore, based on our data, we offer suggestions to enhance the quality of measurements, and offer recommendations to authors aiming to compile and analyze databases from the literature.

Factors structuring Fucus communities at open and complex coastlines in the Baltic Sea

This thesis deals with physical factors and biological interactions affecting the distribution of two fucoid species, Fucus vesiculosus and F. serratus, in the Baltic Sea. Studies have been carried out in two quite different environments: an archipelago, and an open rocky coast. The archipelago has an extremely long coastline with a heterogeneous submerged landscape of different substrate types, slopes, water qualities, and degrees of wave exposure. The factors influencing F. vesiculosus distribution, morphology and epiphyte composition were studied in the Stockholm archipelago using field surveys and spatial modelling in Geographic information systems (GIS). A GIS-method to estimate wave exposure was developed and validated by comparing the result to an index based on vertical zonation of lichens. Wave exposure was considered an important factor for predicting the distribution of F. vesiculosus by its ability to clean hard surfaces from silt, and a predictive model was constructed based on the information of wave exposure and slope of the shore. It is suggested that the lower distribution boundary of attached F. vesiculosus is set by sediment in sheltered parts of the archipelago, and by light availability in highly wave exposed parts. The morphology of F. vesiculosus was studied over a wave exposure gradient, and several characters responded in accordance with earlier studies. However, when separating effects of wave exposure from effects of other confounding water property parameters, only thallus width was significantly different. Several water property parameters were shown to be correlated with wave exposure in the Stockholm archipelago, and the mechanism responsible for the effects on F. vesiculosus morphology is discussed. The composition of epiphytes on F. vesiculosus varied over a wave exposure gradient with a positive correlation to Elachista fucicola, and a negative to Chorda filum. At an open coast the physical environment is much less heterogeneous compared to an archipelago. The distributions of F. vesiculosus, F. serratus, turf-forming algae, and the seafloor substrate, were surveyed along the open coasts of Öland and Gotland. Turf-forming algae dominated all hard substrates in the area, and Polysiphonia fucoides was most abundant. At the Gotland coast F. vesiculosus was less abundant than at the Öland coast, and F. serratus occurred only in the southern-most part. Fucus serratus was increasingly more common towards south which was interpreted as an effect mainly of the Baltic salinity gradient, or the variation of salinity that has occurred in the past. The effects of turf-forming algae and sediment on F. serratus recruitment at 7 m depth off the Öland east coast were studied in the field, and by laboratory experiments. Almost no recruits were found in the algal turf outside the F. serratus patches. More fine sediment was found in the turf than in the F. serratus patches, suggesting that the turf accumulates sediment by decreasing resuspension. Both filamentous algae and sediment decreased the attachment ability of F. serratus zygotes and survival of recruits, and sediment had the strongest effect. It is therefore suggested that F. serratus has difficulties recruiting outside its patches, and that these difficulties are enforced by the eutrophication of the Baltic Sea, which has favoured growth of filamentous algae and increased sedimentation. An overall conclusion is that Fucus distribution is affected by large-scale-factors, such as the eutrophication and salinity changes of the Baltic Sea, as well as by small-scale variation in wave exposure, substrate and slope, and by surface competition with neighbouring species.

Fatty acids and intact polar lipids of laminated microbial sediments from St. Peter Ording, German Wadden Sea

Hidden for the untrained eye through a thin layer of sand, laminated microbial sediments occur in supratidal beaches along the North Sea coast. The inhabiting microbial communities organize themselves in response to vertical gradients of light, oxygen or sulfur compounds. We performed a fine-scale investigation on the vertical zonation of the microbial communities using a lipid biomarker approach, and assessed the biogeochemical processes using a combination of microsensor measurements and a 13C-labeling experiment. Lipid biomarker fingerprinting showed the overarching importance of cyanobacteria and diatoms in these systems, and heterocyst glycolipids revealed the presence of diazotrophic cyanobacteria even in 9 to 20 mm depth. High abundance of ornithine lipids (OL) throughout the system may derive from sulfate reducing bacteria, while a characteristic OL profile between 5 and 8 mm may indicate presence of purple non-sulfur bacteria. The fate of 13C-labeled bicarbonate was followed by experimentally investigating the uptake into microbial lipids, revealing an overarching importance of cyanobacteria for carbon fixation. However, in deeper layers, uptake into purple sulfur bacteria was evident, and a close microbial coupling could be shown by uptake of label into lipids of sulfate reducing bacteria in the deepest layer. Microsensor measurements in sediment cores collected at a later time point revealed the same general pattern as the biomarker analysis and the labeling experiments. Oxygen and pH-microsensor profiles showed active photosynthesis in the top layer. The sulfide that diffuses from deeper down and decreases just below the layer of active oxygenic photosynthesis indicates the presence of sulfur bacteria, like anoxygenic phototrophs that use sulfide instead of water for photosynthesis.

Geochemistry of rare earth elements in basalts from Walvis Ridge

Selected basalts from a suite of dredged and drilled samples (IPOD sites 525, 527, 528 and 530) from the Walvis Ridge have been analysed to determine their rare earth element (REE) contents in order to investigate the origin and evolution of this major structural feature in the South Atlantic Ocean. All of the samples show a high degree of light rare earth element (LREE) enrichment, quite unlike the flat or depleted patterns normally observed for normal mid-ocean ridge basalts (MORBs). Basalts from Sites 527, 528 and 530 show REE patterns characterised by an arcuate shape and relatively low (Ce/Yb)N ratios (1.46-5.22), and the ratios show a positive linear relationship to Nb content. A different trend is exhibited by the dredged basalts and the basalts from Site 525, and their REE patterns have a fairly constant slope, and higher (Ce/Yb)N ratios (4.31-8.50). These differences are further reflected in the ratios of incompatible trace elements, which also indicate considerable variations within the groups. Mixing hyperbolae for these ratios suggest that simple magma mixing between a 'hot spot' type of magma, similar to present-day volcanics of Tristan da Cunha, and a depleted source, possibly similar to that for magmas being erupted at the Mid-Atlantic Ridge, was an important process in the origin of parts of the Walvis Ridge, as exemplified by Sites 527, 528 and 530. Site 525 and dredged basalts cannot be explained by this mixing process, and their incompatible element ratios suggest either a mantle source of a different composition or some complexity to the mixing process. In addition, the occurrence of different types of basalt at the same location suggests there is vertical zonation within the volcanic pile, with the later erupted basalts becoming more alkaline arid more enriched in incompatible elements. The model proposed for the origin and evolution of the Walvis Ridge involves an initial stage of eruption in which the magma was essentially a mixture of enriched and depleted end-member sources, with the N-MORB component being small. The dredged basalts and Site 525, which represent either later-stage eruptives or those close to the hot spot plume, probably result from mixing of the enriched mantle source with variable amounts and variable low degrees of partial melting of the depleted mantle source. As the volcano leaves the hot spot, these late-stage eruptives continue for some time. The change from tholeiitic to alkalic volcanism is probably related either to evolution in the plumbing system and magma chamber of the individual volcano, or to changes in the depth of origin of the enriched mantle source melt, similar to processes in Hawaiian volcanoes.