Biodiversity and Ecosystem Function in the Gulf of Maine: Pattern and Role of Zooplankton and Pelagic Nekton

This paper forms part of a broader overview of biodiversity of marine life in the Gulf of Maine area (GoMA), facilitated by the GoMA Census of Marine Life program. It synthesizes current data on species diversity of zooplankton and pelagic nekton, including compilation of observed species and descriptions of seasonal, regional and cross-shelf diversity patterns. Zooplankton diversity in the GoMA is characterized by spatial differences in community composition among the neritic environment, the coastal shelf, and deep offshore waters. Copepod diversity increased with depth on the Scotian Shelf. On the coastal shelf of the western Gulf of Maine, the number of higher-level taxonomic groups declined with distance from shore, reflecting more nearshore meroplankton. Copepod diversity increased in late summer, and interdecadal diversity shifts were observed, including a period of higher diversity in the 1990s. Changes in species diversity were greatest on interannual scales, intermediate on seasonal scales, and smallest across regions, in contrast to abundance patterns, suggesting that zooplankton diversity may be a more sensitive indicator of ecosystem response to interannual climate variation than zooplankton abundance. Local factors such as bathymetry, proximity of the coast, and advection probably drive zooplankton and pelagic nekton diversity patterns in the GoMA, while ocean-basin-scale diversity patterns probably contribute to the increase in diversity at the Scotian Shelf break, a zone of mixing between the cold-temperate community of the shelf and the warm-water community offshore. Pressing research needs include establishment of a comprehensive system for observing change in zooplankton and pelagic nekton diversity, enhanced observations of "underknown'' but important functional components of the ecosystem, population and metapopulation studies, and development of analytical modeling tools to enhance understanding of diversity patterns and drivers. Ultimately, sustained observations and modeling analysis of biodiversity must be effectively communicated to managers and incorporated into ecosystem approaches for management of GoMA living marine resources.

Trace Elements in the Statoliths of Jumbo Flying Squid Off the Exclusive Economic Zones of Chile and Peru

Ontogenetic variation in 4 trace element ((88)Sr, (137)Ba, (24)Mg, (23)Na) concentrations and their ratios to Ca were measured in statoliths of the jumbo flying squid Dosidicus gigas off the Exclusive Economic Zone of Chilean and Peruvian waters using laser ablation inductively coupled plasma mass spectrometry (LA-ICP-MS). The element compositions of statoliths showed no significant differences between females and males. All of the elements in different growth zones showed significant variations, except for Mg. Sr:Ca and Mg:Ca were good indicators for distinguishing squid from autumn and winter spawning seasons. Sr:Ca and Ba:Ca distribution patterns in statoliths confirmed that paralarvae and juvenile squid inhabit surface waters, while subadult squid migrate into deeper waters. An increasing Sr: Ca ratio of subadult squid could be explained by declining temperature gradients from northern to southern sampling locations, although no significant Sr: Ca differences were observed (p > 0.05). Mg:Ca ratios decreased progressively from the nucleus to the peripheral zone, which might be correlated with statolith growth rates. Na:Ca ratios slightly declined from paralarvae to the subadult phase. Quantitative relationships between statolith trace elements and environmental conditions under different growth stages are needed to improve our understanding of life history of D. gigas.

Regulations of SOX9 activity during chondrogenesis

Sox9 is a transcription factor required for chondrocyte differentiation and cartilage formation. In an effort to identify SOX9 interacting protein(s), we screened a chondrocyte cDNA library with a modified yeast two-hybrid method, Son of Sevenless (SOS) recruitment system (SRS). The catalytic subunit of cyclic AMP-dependent protein kinase A (PKA-Cα) and a new long form of c-Maf transcription factor (Lc-Maf) were found to interact specifically with SOX9. We showed here that two PKA phosphorylation consensus sites of SOX9 could be phosphorylated by PKA in vitro as well as in vivo. PKA phosphorylation of SOX9 increases its DNA binding and transcriptional activities on a Col2a1 chondrocyte-specific enhancer. Mutations of these two PKA phosphorylation sites markedly decreased the activation of SOX9 by PKA. ^ To test whether parathyroid hormone-related peptide (PTHrP) signaling results in SOX9 phosphorylation, we generated a phosphospecific antibody that specifically recognizes SOX9 that is phosphorylated at serine 181 (S 181) one of the two consensus PKA phosphorylation sites. Addition of PTHrP to COS7 cells cotransfected with SOX9 and PTH/PTHrP receptor strongly increased phosphorylation of SOX9 at S181; this phosphorylation was blocked by a PKA-specific inhibitor. In similar experiments we showed that PTHrP increased the activity of a SOX9-dependent Col2a1 enhancer. This increase in activity was abolished when a SOX9 mutant was used containing serine-to-alanine substitution in the two consensus PKA phosphorylation sites of SOX9. Using our phosphospecific SOX9 antibody we showed by immunohistochemistry of mouse embryos that Sox9 phosphorylated at S181 was localized almost exclusively in the pre-hypertrophic zone of the growth plate, an area corresponding to the major site of expression of PTH/PTHrP receptor. In contrast, no phosphorylation of Sox9 at S181 was detected in growth plates of PTH/PTHrP receptor null mutant mice. Sox9, regardless of phosphorylation state, was present in all chondrocytes of both genotypes except in hypertrophic chondrocytes. Thus, Sox9 is a target of PTHrP signaling and the PTHrP-dependent phosphorylation of SOX9 enhances its transcriptional activity. ^ In order to investigate the in vivo function of Sox9 phosphorylation by PKA, we are generating a mouse model of mutant Sox9 harboring point mutations in two PKA phosphorylation sites. Preliminary results indicated that heterozygous mice containing half amount of mutant Sox9 that can not be phosphorylated by PKA have normal skeletal phenotype and homozygous mice are being generated. ^ Lc-Maf encodes an extra ten amino acids at the carboxyl terminus of c-Maf and contains a completely different 3′ untranslated region. The interaction between SOX9 and Lc-Maf was further confirmed by co-immunoprecipitation and GST-pull down assays, which mapped the interacting domains of SOX9 to HMG DNA binding domain and that of Lc-Maf to basic leusine zipper motif. In situ hybridizations showed that RNA of Lc-Maf coexpressed with those of Sox9 and Col2a1 in areas of mesenchymal condensation during the early stages of mouse embryo development. A DNA binding site of Lc-Maf was identified at the 5′ part of a 48-bp Col2a1 enhancer element near the HMG binding site of SOX9. Lc-Maf and SOX9 synergistically activated a luciferase reporter plasmid containing a Col2al enhancer and increased the transcription of endogenous Col2a1 gene. In summary, Lc-Maf is the first identified SOX9-interating protein during chondrogenesis and may be an important activator of Col2a1 gene. ^

Foraminifera and microfossil occurence in Late Jurassic sediments of the North and South Atlantic

Biostratigraphical, taxonomical, and palaeocological results were obtained from Oxfordian to Tithonian foraminifers of the Northern and Southern Atlantic Ocean boreholes of the DSDP Legs 1, 11, 36, 41, 44, 50, and 79. An oversight on the cored Jurassic sections of the DSDP Legs 79 and the corresponding foraminiferal descriptions are given. The reddish brown, clayey and carbonaceous Cat Gap Formation (Oxfordian to Tithonian) of the Northern Atlantic Ocean, rich in radiolarians, yields less or more uniform, in most cases allochthonous foraminiferal faunas of Central European shelf character. No Callovian and Upper Tithonian foraminiferaI zones can be established. The zone of Pseudomarssonella durnortieri covers the Oxfordian/Kimmeridgian, the zone of Neobulimina atlantica the Kimmeridgian/Lower Tithonian interval. Characteristic foraminiferal faunas are missing since the Upper Tithonian to Valanginian for reason of a widely distributed regression which caused hiatuses observed all over the Northern Atlantic Ocean and in parts of Europe. The Upper Jurassic cannot be subdivided into single stages by foraminiferal biostratigraphy alone. The fovaminiferal zones established by Moullad (1984) covering a Callovian-Tithonian interval may be of some local importance in the Tethyan realm: It has too long-ranging foraminiferal species to be used as index marker in the word-wide DSDP boreholes. Some taxonomical confusion is caused because in former publications some foraminiferal species have got different names both in the Jurassic and Cretaceous. The foraminiferal biostratigraphy of drilled sections from DSDP boreholes is restricted by the drilling technique and for palaeo-oceanographical, biological, and geological reasons. Foraminiferal faunas from the DSDP originally described as ,,bathyal, or ,,abyssal,, have to be derived from shallower water. This contrasts the palaeo-water depths of 3000-4000 m which result from sedimentological and palaeo-geographical investigations.

Organic carbon in waters, suspended matter, and upper layer bottom sediments of the Kara Sea and Obskay Guba

Along three sections in the Kara Sea and Obskaya Guba concentrations of dissolved and particulate organic carbon (DOC and POC, respectively) in waters , as well as of organic carbon in bottom sediments (Corg) in September-October 2007 were determined. DOC varied from 6.3 to 2400 µg/l, POC - from 0.84 to 12.2 mg/l. For all samples the average DOC was 200 µg/l (n = 78; sigma = 368), the average POC - 2.7 mg/l (n = 92; sigma = 2.7). Concentrations of Corg in dried samples of upper layer bottom sediments varied from 0.13 to 2.10% (aver. = 0.9%; n = 21; sigma = 0.49%). It is shown that distribution of different forms of organic matter (OM) is an indicator of supply and scattering of particulate matter in the Kara Sea and that DOC and POC of the Kara Sea are formed under impact of runoff of the Ob and Yenisei Rivers. It is found that distribution of OM in bottom sediments is closely related to their grain size composition and to the structure of currents in the area. Variations in Corg concentration in bottom sediment cores from the zone of riverine and sea water mixing represent variability of OM burial.

(Table 1) Isotopic composition of Sm and Nd in mantle restites of the Central Atlantic and in associated plutonic and volcanic rocks

Ultrabasic rock samples collected from two areas of the crustal zone of the Mid-Atlantic Ridge (MAR): (1) 13-17°N (near the intersection of the ridge axis with the 15°20'N prime fracture zone), and (2) 33°40'N prime (the western intersection of the MAR crest with the Heis fracture zone) were objects of this study. Samples of peridotite and of plutonic and volcanic rocks associated with it were used to measure their Sm/Nd, 143Nd/144Nd, and 147Sm/144Nd ratios, which allowed to test time and genetic relationships between evolution of mantle material under the ridge crest and products of its magmatic activity. Results of this work proved ubiquitous discrepancy between melting degree values of extremely depleted mantle peridotites in the MAR area between 14°N and 16°N, obtained using petrologic and geochemical methods. This discrepancy suggests large-scale interaction between mantle material and magmatic melts and fluids enriched in incompatible elements or fluids. The results obtained suggest that repeated melting of the mantle under the axial MAR zone is an universal characteristic of magmatism in low-velocity spreading centers. The results of this study also proved the crestal MAR zone in the Central Atlantic region show distinct indications of isotope-geochemical segmentation of the mantle. It is suggested that the geochemically anomalous MAR mantle peridotite in the zone of the MAR intersection with the 15°20'N prime fracture zone can be interpreted as fragments of mantle substrate, foreign for the Atlantic mantle north of the equator.

Th-U ages of a stalagmite from the Kleegruben Cave (Table 1)

A mass-spectrometric uranium-series dated stalagmite from the Central Alps of Austria provides unprecedented new insights into high-altitude climate change during the peak of isotope stage 3. The stalagmite formed continuously between 57 and 46 kyr before present. A series of 'Hendy tests' demonstrates that the outer parts of the sample show a progressive increase of both stable C and O isotope values. No such covariant increase was detected within the axial zone. This in conjunction with other observations suggests that the continuous stable oxygen isotope profile obtained from the axial zone of the stalagmite largely reflects the unaltered isotopic composition of the cave drip water. The delta18O record shows events of high delta18O values that correlate remarkably with Interstadials 15 (a and b), 14 and 12 identified in the Greenland ice cores. Interstadial 15b started rapidly at 55.6 kyr and lasted ~300 yr only, Interstadial 15a peaked 54.9 kyr ago and was even of shorter duration (~100 yr), and Interstadial 14 commenced 54.2 kyr ago and lasted ~3000 yr. This stalagmite thus represents one of the first terrestrial archives outside the high latitudes which record precisely dated Dansgaard-Oeschger (D/O) events during isotope stage 3. Provided that rapid D/O warmings occurred synchronously in Greenland and the European Alps, the new data provide an independent tool to improve the GRIP and GISP2 chronologies.

Grain size distribution in surface sediments samples of the continental shelf and slope of southeast Greenland

During "Meteor" Cruise 6/1966 in the northwest Atlantic a systematic survey of the bottom topography of the southeast Greenland continental margin was undertaken. Eighty-seven profiles transverse to the shelf edge at distances of 3-4 nautical miles and two longitudinal profiles parallel to the coast were carried out with the ELAC Narrow Beam Echo-Sounder giving a reliable record of even steep slopes. On the basis of the echo soundings the topography and morphology of the continental shelf and slope are evaluated. A detailed bathymetric chart and a serial profile chart were designed as working material for the morphological research. These maps along with the original echograms are morphometrically evaluated. The analysis of the sea bottom features is the basis of a subsequent morphogenetical interpretation, verified and extended by means of interpretation of magnetic data and sediment analysis (grain size, roundness, lithology). The results of the research are expressed in a geomorphological map. The primary findings can be summarized as follows: 1) The southeast Greenland shelf by its bottom topography can be clearly designated as a glacially formed area. The glacial features of the shelf can be classified into two zones nearly parallel to the coast: glacial erosion forms on the inner shelf and glacial accumulation forms on the outer shelf. The inner shelf is characterized by the rugged and hummocky topography of ice scoured plains with clear west/east slope asymmetry. On the outer shelf three types of glacial accumulation forms can be recognized: ice margin deposits with clearly expressed terminal moraines, glacial till plains and glaciomarine outwash fans. Both zones of the shelf can be subdivided into two levels of relief. The ice scoured plains, with average depths of 240 meters (m), are dissected to a maximum depth of 1060 m (Gyldenloves Trough) by trough valleys, which are the prolongations of the Greenland fjords. The banks of the outer shelf, with an average depth of 180 m, surround glacial basins with a maximum depth of 670 meters. 2) The sediments of the continental shelf can be classified as glacial due to their grain size distribution and the degree of roundness of the gravel particles. The ice margin deposits on the outer shelf can be recognized by their high percentage of gravels. On the inner shelf a rock surface is suggested, intermittently covered by glacial deposits. In the shelf troughs fine-grained sediments occur mixed with gravels. 3) Topography and sediments show that the southeast Greenland shelf was covered by an ice sheet resting on the sea floor during the Pleistocene ice-age. The large end moraines along the shelf edge probably indicate the maximum extent of the Wurm shelf ice resting on the sea floor. The breakthroughs of the end moraines in front of the glacial basins suggest that the shelf ice has floated further seaward over the increasing depths. 4) Petrographically the shelf sediments consist of gneisses, granites and basalts. While gneisses and granites occire on the nearby coast, basalt is not known to exist here. Either this material has been drifted by icebergs from the basalt province to the north or exists on the southeast Greenland shelf itself. The last interpretation is supported bythe high portion of basalt contained in the sediment samples taken and the strong magnetic anomalies probably caused by basaltic intrusions. 5) A magnetic profile allows the recognition of two magnetically differing areas which approximately coincide with the glacial erosion and accumulation zones. The inner shelf shows a strong and variable magnetic field because the glacially eroded basement forms the sea floor. The outer shelf is characterized by a weak and homogenous magnetic field, as the magnetized basement lies at greater depthy, buried by a thick cover of glacial sediments. The strong magnetic anomalies of the inner shelf are probably caused by dike swarms, similar to those observed further to the north in the Kangerdlugssuaq Fjord region. This interpretation is supported by the high basalt content of the sediment samples and the rough topography of the ice scoured plains which correlates in general with the magnetic fluctuations. The dike structures of the basement have been differentially eroded by the shelf ice. 6) The continental slope, extending from the shelf break at 313 m to a depth of 1270 m with an average slope of 11°, is characterized by delta-shaped projections in front of the shelf basins, by marginal plateaus, ridges and hills, by canyons and slumping features. The projections could be identified as glaciomarine sediment fans. This conclusion is supported by the strong decrease of magnetic field intensity. The deep sea hills and ridges with their greater magnetic intensities have to be regarded as basement outcrops projecting through the glaciomarine sediment cover. The upper continental rise, sloping seaward at about 2°, is composed of wide sediment fans and slump material. A marginal depression on the continental rise running parallel to the shelf edge has been identified. In this depression bottom currents capable of erosion have been recorded. South of Cape Farvel the depression extends to the accumulation zone of the "Eirik" sedimentary ridge. 7) By means of a study of the recent marine processes, postglacial modification of the ice-formed relief can be postulated. The retention effect of the fjord troughs and the high velocity of the East Greenland stream prevents the glacial features from being buried by sediments. Bottom currents capable of active erosion have only been found in the marginal depression on the continental rise. In addition, at the time of the lowest glacio-eustatic sea level, the shelf bottom was not situated in the zone of wave erosion. Only on the continental slope and rise bottom currents, sediment slumps and turbidity currents have led to significant recent modifications. Considering these results, the geomorphological development of the southeast Greenland continental terrace can be suggested as follows: 1. initial formation of a "peneplain", 2. fluvial incision, 3. submergence, and finally 4. glacial modification.

(Table 1) Organic geochemistry of ODP Hole 134-832B and Site 134-833

Ocean Drilling Program (ODP) Leg 134 was located in the central part of the New Hebrides Island Arc, in the Southwest Pacific. Here the d'Entrecasteaux Zone of ridges, the North d'Entrecasteaux Ridge and South d'Entrecasteaux Chain, is colliding with the arc. The region has a Neogene history of subduction polarity reversal, ridge-arc collision, and back-arc spreading. The reasons for drilling in this region included the following: (1) to determine the differences in the style and time scale of deformation associated with the two ridge-like features (a fairly continuous ridge and an irregularly topographic seamount chain) that are colliding with the central New Hebrides Island Arc; (2) to document the evolution of the magmatic arc in relation to the collision process and possible Neogene reversal of subduction; and (3) to understand the process of dewatering of a small accretionary wedge associated with ridge collision and subduction. Seven sites were occupied during the leg, five (Sites 827-831) were located in the d'Entrecasteaux Zone where collision is active. Three sites (Sites 827, 828, and 829) were located where the North d'Entrecasteaux Ridge is colliding, whereas two sites (Sites 830 and 831) were located in the South d'Entrecasteaux Chain collision zone. Sites 828 (on North d'Entrecasteaux Ridge) and 831 (on Bougainville Guyot) were located on the Pacific Plate, whereas all other sites were located on a microplate of the North Fiji Basin. Two sites (Sites 832 and 831) were located in the intra-arc North Aoba Basin. Results of Leg 134 drilling showed that forearc deformation associated with the North d'Entrecasteaux Ridge and South d'Entrecasteaux Chain collision is distinct and different. The d'Entrecasteaux Zone is an Eocene subduction/obduction complex with a distinct submerged island arc. Collision and subduction of the North d'Entrecasteaux Ridge results in off scraping of ridge material and plating of the forearc with thrust sheets (flakes) as well as distinct forearc uplift. Some offscraped sedimentary rocks and surficial volcanic basement rocks of the North d'Entrecasteaux Ridge are being underplated to the New Hebrides Island forearc. In contrast, the South d'Entrecasteaux Chain is a serrated feature resulting in intermittent collision and subduction of seamounts. The collision of the Bougainville Guyot has indented the forearc and appears to be causing shortening through thrust faulting. In addition, we found that the Quaternary relative convergence rate between the New Hebrides Island Arc at the latitude of Espiritu Santo Island is as high as 14 to 16 cm/yr. The northward migration rate of the d'Entrecasteaux Zone was found the be ~2 to 4 cm/yr based on the newly determined Quaternary relative convergence rate. Using these rates we established the timing of initial d'Entrecasteaux Zone collision with the arc at ~3 Ma at the latitude of Epi Island and fixed the impact of the North d'Entrecasteaux Ridge upon Espiritu Santo Island at early Pleistocene (between 1.89 and 1.58 Ma). Dewatering is occurring in the North d'Entrecasteaux Ridge accretionary wedge, and the wedge is dryer than other previously studied accretionary wedges, such as Barbados. This could be the result of less sediment being subducted at the New Hebrides compared to the Barbados.

(Table 2) Biomass and size characteristics of Aurelia aurita in the Black Sea as determined from the Argus manned submersible

Based on estimate of Aurelia aurita concentration in the Black Sea from the Argus manned submersible in April-May 1984, as well as on author's data and published information on metabolic rate and feeding of medusa, biomass of medusa Aurelia aurita in the epipelagic zone of the Black Sea is estimated to be about 400 million tons of wet weight, and its mean annual production to be 400-900 million tons wet weight or about 1.1-2.5 million tons of organic carbon, equivalent to approximately 1-3% of primary production.

Studies of the microbial diversity on analyzed samples from from Peru Margin, Equatorial Pacific, Hydrate Ridge, and Juan de Fuca Ridge

Marine sediments harbor an enormous quantity of microorganisms, including a multitude of novel species. The habitable zone of the marine sediment column begins at the sediment-water interface and probably extends to depths of several thousands of meters. Studies of the microbial diversity in this ecosystem have mostly relied on molecular biological techniques. We used a complementary method - analysis of intact polar membrane lipids - to characterize the in-situ microbial community in sediments covering a wide range of environmental conditions from Peru Margin, Equatorial Pacific, Hydrate Ridge, and Juan de Fuca Ridge. Bacterial and eukaryotic phospholipids were only detected in surface sediments from the Peru Margin. In contrast, deeply buried sediments, independent of their geographic location, were dominated by archaeal diether and tetraether lipids with various polar head groups and core lipids. We compared ring distributions of archaeal tetraether lipids derived from polar glycosidic precursors with those that are present as core lipids. The distributions of these related compound pools were distinct, suggestive of different archaeal sources, i.e., the polar compounds derive from sedimentary communities and the core lipids are fossil remnants from planktonic communities with possible admixtures of decayed sedimentary archaea. This in-situ production of distinct archaeal lipid populations potentially affects applications of the TEX86 paleotemperature proxy as demonstrated by offsets in reconstructed temperatures between both pools. We evaluated how varying cell and lipid stabilities will influence the sedimentary pool by using a box-model. The results are consistent with (i) a requirement of continuous inputs of freshly synthesized lipids in subsurface sediments for explaining the observed distribution of intact polar lipids, and (ii) decreasing lipid inputs with increasing burial depth.

Biology community and bottom sediments in the West Bay of the Furugelm Island, Sea of Japan

An experiment was carried out on the soft bottom in the sublitoral zone of the Furugelm Island (Peter the Great Bay, Sea of Japan) to study formation of benthic communities. Boxes with defauned sediments were placed on depths of 4, 6 and 13 m and exposed during 60 days in the summer period. Half of them were covered with a net with mesh size 2 cm to prevent effect of large predators. It was found that spatial pattern of invertebrates' sinking in the bay conforms to distribution of benthic communities. Larvae of benthic invertebrates sinks in general in places inhabited by their adult species. The main factors responsible for recolonzation are: sediment type and local hydrodynamic conditions. Heart-shaped sea urchin Echinocardium cordatum is numerically dominated in the bay on depth 3-4.5 m, but its larvae sinks in the deeper area. Community structure is supported by mature specimen migration to places inhabited by species. Predators affect largely on the species.