The genus phymatolithon in the Gulf of Maine

Summary New information on anatomy, cytology and the development of reproductive structures is presented to show that Phymatolithon is a genus distinct from both Clathromorphum and the branching members of Lithothamnium. Also, a new species of Phymatolithon, Ph. rugulosum, is described. The reproductive cycles and geographic and bathymetric distributions of Ph. laevigatum and Ph. rugulosum in the Gulf of Maine are presented and discussed. There is strong indication that the geographic distribution of crustose corallines in the region is controlled primarily by maximum summer temperatures. The depth distributions are apparently controlled primarily by decrease of light with depth, though temperatures and substrate are also factors.Now at Colorado School of Mines, Golden, Colorado, U.S.A. The necessary field work for this study was made possible through the generous support of: The American Petroleum Institute, The Geological Society of America, the ONR (through the Department of Geology and Geophysics, Massachusetts Institute of Technology) and the Woods Hole Oceanographie Institution (Contr. No. 1500).     

Planktonic microbes in the Gulf of Maine area

In the Gulf of Maine area (GoMA), as elsewhere in the ocean, the organisms of greatest numerical abundance are microbes. Viruses in GoMA are largely cyanophages and bacteriophages, including podoviruses which lack tails. There is also evidence of Mimivirus and Chlorovirus in the metagenome. Bacteria in GoMA comprise the dominant SAR11 phylotype cluster, and other abundant phylotypes such as SAR86-like cluster, SAR116-like cluster, Roseobacter, Rhodospirillaceae, Acidomicrobidae, Flavobacteriales, Cytophaga, and unclassified Alphaproteobacteria and Gammaproteobacteria clusters. Bacterial epibionts of the dinoflagellate Alexandrium fundyense include Rhodobacteraceae, Flavobacteriaceae, Cytophaga spp., Sulfitobacter spp., Sphingomonas spp., and unclassified Bacteroidetes. Phototrophic prokaryotes in GoMA include cyanobacteria that contain chlorophyll (mainly Synechococcus), aerobic anoxygenic phototrophs that contain bacteriochlorophyll, and bacteria that contain proteorhodopsin. Eukaryotic microalgae in GoMA include Bacillariophyceae, Dinophyceae, Prymnesiophyceae, Prasinophyceae, Trebouxiophyceae, Cryptophyceae, Dictyochophyceae, Chrysophyceae, Eustigmatophyceae, Pelagophyceae, Synurophyceae, and Xanthophyceae. There are no records of Bolidophyceae, Aurearenophyceae, Raphidophyceae, and Synchromophyceae in GoMA. In total, there are records for 665 names and 229 genera of microalgae. Heterotrophic eukaryotic protists in GoMA include Dinophyceae, Alveolata, Apicomplexa, amoeboid organisms, Labrynthulida, and heterotrophic marine stramenopiles (MAST). Ciliates include Strombidium, Lohmaniella, Tontonia, Strobilidium, Strombidinopsis and the mixotrophs Laboea strobila and Myrionecta rubrum (ex Mesodinium rubra). An inventory of selected microbial groups in each of 14 physiographic regions in GoMA is made by combining information on the depth-dependent variation of cell density and the depth-dependent variation of water volume. Across the entire GoMA, an estimate for the minimum abundance of cell-based microbes is 1.7×10(25) organisms. By one account, this number of microbes implies a richness of 10(5) to 10(6) taxa in the entire water volume of GoMA. Morphological diversity in microplankton is well-described but the true extent of taxonomic diversity, especially in the femtoplankton, picoplankton and nanoplankton--whether autotrophic, heterotrophic, or mixotrophic, is unknown.     

The genus Clathromorphum (Corallinaceae) in the Gulf of Maine

Summary This paper is the second of a series covering the crustose corallines of the Gulf of Maine. Information on anatomy, cytology and the development of the reproductive structures is presented to show that Clathromorphum is a genus distinct from both Phymatolithon and the branching members of Lithothamnium. Also, for the two species of this genus present in the region, reproductive cycles and geographic and bathymetric distributions are presented and discussed.Part of a study undertaken while working on a Ph. D. program at the University of Michigan. The necessary field work was made possible by the generous support of: the American Petroleum Institute (Grant-in-aid No. 85), the Geological Society of America (Penrose Grant No. 912-62), the Office of Naval Research (Nonr 1841 (74), through the Department of Geology and Geophysics, Massachusetts Institute of Technology) and the Woods Hole Oceanographic Institution.     

The genus Pseudolithophyllum (Corallinaceae) in the Gulf of Maine

Summary This is the third of a series of papers covering the anatomy, morphology, systematics and ecology of the crustose corallines of the Gulf of Maine. Pseudolithophyllum orbiculatum, the only species of this genus in the region, is treated in this paper.Part of a study undertaken while working on a Ph. D. program at the University of Michigan. The necessary field work was made possible by the generous support of: the American Petroleum Institute (Grant-in-aid No. 85), the Geological Society of America (Penrose Grant No. 912-62), the Office of Naval Research (Nonr 1841-74), through the Department of Geology and Geophysics, Massachusetts Institute of Technology and the Woods Hole Oceanographic Institution.     

Use of Herring Bait to Farm Lobsters in the Gulf of Maine

Background

Ecologists, fisheries scientists, and coastal managers have all called for an ecosystem approach to fisheries management, yet many species such as the American lobster (Homarus americanus) are still largely managed individually. One hypothesis that has yet to be tested suggests that human augmentation of lobster diets via the use of Atlantic herring (Clupea harengus) as bait may contribute to recent increases in lobster landings. Currently 70% of Atlantic herring landings in the Gulf of Maine are used as bait to catch lobsters in traps throughout coastal New England.

Methodology/Principal Findings

We examined the effects of this herring bait on the diet composition and growth rate of lobsters at heavily baited vs. seasonally closed (i.e., bait free) sites in coastal Maine. Our results suggest that human use of herring bait may be subsidizing juvenile lobster diets, thereby enhancing lobster growth and the overall economic value and yield of one of the most valuable fisheries in the U.S.

Conclusions/Significance

Our study illustrates that shifting to an ecosystem approach to fisheries management should require consideration of cross-fishery interactions.     

Marine Foraging Birds As Bioindicators of Mercury in the Gulf of Maine

From existing databases, we compiled and evaluated 604 total mercury (Hg) levels in the eggs and blood of 17 species of marine foraging birds from 35 Gulf of Maine islands to provide baseline data and to determine the best tissue, age class, and species for future biomonitoring. While mean Hg levels in most species did not exceed adverse effects thresholds, levels in some individual eggs did; for all species arithmetic mean egg Hg levels ranged from 0.04 to 0.62 (μg/g, wet weight). Piscivorous birds had higher Hg levels than invertivores. Leach’s storm-petrel (Oceanodroma leucorhoa), razorbill (Alca torda), and black guillemot (Cepphus grylle) adult blood and egg Hg levels were higher than other species. Our results indicate that adult blood is preferable to chick blood for detecting long-term temporal trends because adult levels are higher and not confounded by metabolic effects. However, since we found that eggs and adult blood are comparable indicators of methylmercury bioavailability, we determined that eggs are the preferred tissue for long-term Hg monitoring because the relative ease in collecting eggs ensures consistent and robust datasets. We suggest specific sampling methods, and based on our results demonstrate that common eider (Somateria mollissima), Leach’s storm-petrel, double-crested cormorant, and black guillemot are the most effective bioindicators of Hg of the Gulf of Maine.     

Quantifying alosine prey in the diets of marine piscivores in the Gulf of Maine

The objectives of this work were to quantify the spatial and temporal distribution of the occurrence of anadromous fishes (alewife Alosa pseudoharengus, blueback herring Alosa aestivalis and American shad Alosa sapidissima) in the stomachs of demersal fishes in coastal waters of the north‐west Atlantic Ocean. Results show that anadromous fishes were detectable and quantifiable in the diets of common marine piscivores for every season sampled. Even though anadromous fishes were not the most abundant prey, they accounted for c. 5–10% of the diet by mass for several marine piscivores. Statistical comparisons of these data with fish diet data from a broad‐scale survey of the north‐west Atlantic Ocean indicate that the frequency of this trophic interaction was significantly higher within spatially and temporally focused sampling areas of this study than in the broad‐scale survey. Odds ratios of anadromous predation were as much as 460 times higher in the targeted sampling as compared with the broad‐scale sampling. Analyses indicate that anadromous prey consumption was more concentrated in the near‐coastal waters compared with consumption of a similar, but more widely distributed species, the Atlantic herring Clupea harengus. In the context of ecosystem‐based fisheries management, the results suggest that even low‐frequency feeding events may be locally important, and should be incorporated into ecosystem models.     

Ecology of larval herring in relation to the oceanography of the Gulf of Maine

The dynamics of larval herring in the north-eastern Gulf ofMaine are reviewed with reference to the important physicaland biological oceanographic processes of the region. Particularattention is given to the apparently conflicting hypothesesof larval drift from the tidally well-mixed spawning areas tonursery areas, and larval retention in spawning areas for aperiod of several months. Both processes have been reportedfor the eastern Maine-Grand Manan spawning area of the Gulfof Maine, but the relative importance of each to larval survivalthrough the winter and recruitment to the juvenile stage isnot clear. Both transport and retention are interpreted in lightof oceanographic processes that might impart variability, especiallybetween years, in the proportion of larvae transported awayversus that retained. Results of recent survey cruises in theGulf of Maine show both hypotheses to have merit. It is suggestedthat (i) interannual differences in slope water intrusions intothe Gulf of Maine as they affect the residual circulation, (ii)the lunar periodicity in the intensity of tidal mixing in relationto hatching times, and (iii) the potential for variable spawninglocations relative to the tidal fronts, may affect the distributionsof herring larvae immediately after hatching in the fall, andmay control the proportion of larvae that are advected awaywith the residual currents versus that retained in the vicinityof spawning. It is also suggested that those processes thataffect larval distributions and survival in the fall are importantin determining the overwintering distributions of larvae inthe Gulf of Maine, where the subtle influences of variable foodregimes and water temperatures could potentially exact largeinterannual differences in winter survival and recruitment tothe juvenile stage.     

Subsurface chlorophyll maxima and vertical distribution of zooplankton in the Gulf of Maine

The vertical disthbution of chlorophyll, zooplankton and physicalstructure were measured using a pumping system and CTD on twocruises in the Gulf of Maine during June and September 1982.The vertical distribution of chlorophyll was closely relatedto the density structure of the water column. In waters witha pronounced pycnocline subsurface chiorphyll maxima (SCM) werelocated at or just above the pycnocline. Chlorophyll concentrationswere maximal in the surface waters at those stations sampledin June where the pycnocline was not well defined. The relationshipbetween the zooplankton and chlorophyll distribution differedbetween cruises. In June, the zooplankton, particularly post-naupliarcopepods, were associated with the depth of the chlorophyllmaxirnum, while in September the post-naupliar copepods weremost abundant in the surface waters above the SCM at the stratifiedstations. During the September cruise we observed that the copepodnauplii were most abundant at the depth of the SCM, and thatthe larger protozoans (>35 µm) were most abundant atdepths of 55–85 m, which were well below the SCM and pycnocline. ^*Bigelow Laboratory for Ocean Sciences Contribution No. 83025     

A Regional Assessment of Salt Marsh Restoration and Monitoring in the Gulf of Maine

We compiled salt marsh monitoring datasets from 36 complete or imminent restoration projects in the Gulf of Maine to assess regional monitoring and restoration practices. Data were organized by functional indicators and restoration project types (culvert replacement, excavation works, or ditch plugging) then pooled to generate mean values for indicators before restoration, after restoration, and at reference sites. Monitoring data were checked against the regional standards of a voluntary protocol for the Gulf of Maine. Data inventories showed that vegetation and salinity indicators were most frequently collected (89 and 78% of sites, respectively), whereas nekton, bird, and hydrologic measures were collected at only about half of the sites. Reference conditions were monitored at 72% of sites. Indicators were analyzed to see if project sites were degraded relative to reference areas and to detect ecological responses to restoration activities. Results showed that compared to reference areas, prerestoration sites had smaller tidal ranges, reduced salinity levels, greater cover of brackish plants species, and lower cover of halophyte plants. Following restoration, physical factors rebounded rapidly with increased flood and salinity levels after about one year, especially for culvert projects. Biological responses were less definitive and occurred over longer time frames. Plant communities trended toward recovered halophytes and reduced brackish species at 3+ years following restoration. Nekton and avian indicators were indistinguishable among reference, impacted, and restored areas. The protocol was successful in demonstrating restoration response for the region, but results were limited by regional inconsistencies in field practices and relatively few multiyear datasets. To improve future assessment capabilities, we encourage greater adherence to the standard protocol throughout the Gulf of Maine salt marsh restoration community.     

Regional variation in fish predation intensity: a historical perspective in the Gulf of Maine

Summary Regional variation in the intensity of fish predation on tethered brittle stars and crabs was measured at 30–33 m depths in the rocky subtidal zone at seven sites representing coastal and offshore regions of the Gulf of Maine, USA. Analysis of covariance comparing the slopes of brittle star survivorship curves followed by multiple comparisons tests revealed five groupings of sites, with significantly greater predation rates in the two offshore than in the three coastal groups. Brittle stars tethered at the three offshore sites were consumed primarily by cod, Gadus morhua, with 60–100% prey mortality occuring in 2.5 h. In striking contrast, only 6–28% of brittle star prey was consumed in the same amount of time at the four coastal sites, which were dominated by cunner, Tautogolabrus adspersus. In several coastal trials, a majority of brittle star prey remained after 24 h. The pattern of higher predation offshore held for rock crabs as well with only 2.7% of tethered crabs consumed (n=36) at coastal sites versus 57.8% of crabs (n=64) consumed at offshore sites. Another important predatory fish, the wolffish, Anarhichas lupus, consumed more tethered crabs than brittle stars. Videos and time-lapse movies indicated that cod and wolffish were significantly more abundant at offshore than at coastal sites. Three hundred years of fishing pressure in New England has severely depleted stocks of at least one important benthic predator, the cod, in coastal waters. We speculate that this human-induced predator removal has lowered predation pressure on crabs and other large mobile epibenthos in deep coastal communities. Transect data indicate that coastal sites with few cod support significantly higher densities of crabs than offshore sites with abundant cod.     

A mechanism for offshore initiation of harmful algal blooms in the coastal Gulf of Maine

A combination of observations and model results suggest a mechanismby which coastal blooms of the toxic dinoflagellate Alexandriumfundyense can be initiated from dormant cysts located in offshoresediments. The mechanism arises from the joint effects of organismbehavior and the wind-driven response of a surface-trapped plumeof fresh water originating from riverine sources. During upwelling-favorablewinds, the plume thins vertically and extends offshore; downwellingwinds thicken the plume and confine it to the nearshore region.In the western Gulf of Maine, the offshore extent of the riverplume during upwelling conditions is sufficient to entrain upward-swimmingA. fundyense cells germinated from offshore cyst beds. Subsequentdownwelling conditions then transport those populations towardsthe coast.     

Zooplankton community grazing impact on a bloom of Alexandrium fundyense in the Gulf of Maine

Shipboard grazing experiments were conducted in the Gulf of Maine and on Georges Bank during of June 2006 to estimate zooplankton community grazing impact on a natural bloom of the toxic dinoflagellate Alexandrium fundyense. Surface seawater samples containing natural populations of grazers and A. fundyense from 23 stations were incubated at ambient temperatures. Concentrations of A. fundyense after incubations were compared to those at the start of each experiment to determine net increases due to population growth, or decreases presumed to be primarily due to grazing losses. Abundances of both microzooplankton (tintinnids, oligotrich ciliates, rotifers, copepod nauplii and heterotrophic dinoflagellates) and mesozooplankton (copepod nauplii, copepodites and adult copepods, rotifers, marine cladocerans, and meroplankton) grazers in experimental aliquots were also determined. The total zooplankton community had minimal grazing impact on natural populations of A. fundyense at most stations. At 70% of the stations where grazing experiments were performed, there were no significant differences in initial and final concentrations of A. fundyense. This indicated that growth of, and grazing on A. fundyense were in approximate balance. At 2 stations, which had the highest A. fundyense abundances of the cruise (>10⁴ cells l⁻¹), % of the A. fundyense population grazed per day was significantly negative, indicating that net population growth of A. fundyense exceeded grazing losses. At 5 stations, which had low concentrations of A. fundyense (10²–10³ cells l⁻¹), % of the A. fundyense population grazed per day was significantly positive, indicating that losses of A. fundyense due to grazing exceeded net population growth. For stations with significant differences between Initial and Grazed concentrations of A. fundyense, grazing had the greatest impact at lower concentrations of A. fundyense, and grazing impact by the larger mesozooplankton was inversely related to zooplankton abundance. There was no relationship between microzooplankton abundance and grazing impact on A. fundyense. Grazing exceeded growth only where A. fundyense abundance was low, and growth exceeded grazing only where A. fundyense abundance was high. The inverse relationship between grazing impact and A. fundyense abundance implies that grazing may be capable of retarding bloom development at low concentrations typical of the early stages of a bloom, but at higher concentrations once a bloom becomes established, either grazing maintains a balance with A. fundyense growth, or growth exceeds grazing losses at highest concentrations.     

Distribution and long-term temporal patterns of four invasive colonial ascidians in the Gulf of Maine

Invasive ascidians are a growing concern for ecologists and natural resource managers, yet few studies have documented their short- and long-term temporal patterns of abundance. This study focuses on the invasion of the Gulf of Maine by the colonial ascidians Botryllus schlosseri, Botrylloides violaceus, Diplosoma listerianum and Didemnum sp. A. We examined the time of arrival and potential vectors for these four invasive ascidians using survey data (collected from 1969 onwards) and literature documentation. We also compared the dominance and seasonal patterns of abundance of these species using data from two identical panel studies; one conducted from 1979 to 1980, the other from 2003 to 2005. Didemnum and Botrylloides were most likely first introduced into the Damariscotta River, Maine in the early 1970's through oyster aquaculture while Botryllus and Diplosoma were probably transported by commercial and recreational vessels. The overall abundance of colonial ascidians has increased since 1979 and 1980. Botryllus was the only invasive colonial ascidian present during the 1979 to 1980 study and accounted for an average of 6.16% cover on panels. From 2003 to 2005, the more recently arrived colonial ascidians Botrylloides and Didemnum accounted for 7.38% and 2.08% cover respectively, while Botryllus covered only 1.16%. Our results reveal a shift in seasonal abundance between 1979 to 1980 and 2003 to 2004. In 1979 and 1980, colonial ascidians had the highest percent cover in fall and winter while in 2003 and 2005 they had highest percent cover in summer and fall. Seasonal patterns of space occupation by colonial ascidians were correlated with seasonal changes in seawater temperature.     

Metagenomic analysis of a complex marine planktonic thaumarchaeal community from the Gulf of Maine

Thaumarchaea, which represent as much as 20% of prokaryotic biomass in the open ocean, have been linked to environmentally relevant biogeochemical processes, such as ammonia oxidation (nitrification) and inorganic carbon fixation. We have used culture-independent methods to study this group because current cultivation limitations have proved a hindrance in studying these organisms. From a metagenomic data set obtained from surface waters from the Gulf of Maine, we have identified 36,111 sequence reads (containing 30 Mbp) likely derived from environmental planktonic Thaumarchaea. Metabolic analysis of the raw sequences and assemblies identified copies of the catalytic subunit required in aerobic ammonia oxidation. In addition, genes that comprise a nearly complete carbon assimilation pathway in the form of the 3-hyroxypropionate/4-hydroxybutyrate cycle were identified. Comparative genomics contrasting the putative environmental thaumarchaeal sequences and 'Candidatus Nitrosopumilus maritimus SCM1' revealed a number of genomic islands absent in the Gulf of Maine population. Analysis of these genomic islands revealed an integrase-associated island also found in distantly related microbial species, variations in the abundance of genes predicted to be important in thaumarchaeal respiratory chain, and the absence of a high-affinity phosphate uptake operon. Analysis of the underlying sequence diversity suggests the presence of at least two dominant environmental populations. Attempts to assemble complete environmental genomes were unsuccessful, but analysis of scaffolds revealed two diverging populations, including a thaumarchaeal-related scaffold with the full urease operon. Ultimately, the analysis revealed a number of insights into the metabolic potential of a predominantly uncultivated lineage of organisms. The predicted functions in the thaumarchaeal metagenomic sequences are directly supported by historic measurements of nutrient concentrations and provide new avenues of research in regards to understanding the role Thaumarchaea play in the environment.     

Larval settlement of the green sea urchin, Strongylocentrotus droebachiensis, in the southern Gulf of Maine

Abstract. Settlement timing and differential settlement for the larval stage of the green sea urchin, Strongylocentrotus droebachiensis , in the southern Gulf of Maine was studied during the summer of 1996. Settlement densities on astroturf panels were highest in June and early July (13 to 37 m ⁻² d⁻¹), and peaked in mid-June (199 m⁻² d⁻¹). Settlement was low to nonexistent from mid-July through August (0 to 2 m⁻² d⁻¹). During the peak in settlement, no selection for substrate type was observed. In the remainder of the settlement period, differential settlement occurred, with a preference for substrate covered with live coralline algae. Test diameter of newly settled urchins varied among the substrates, with urchins settling on live coralline algae having the largest test diameter (0.43 ± 0.01 mm). There were no differences in test diameter among the different weeks in which sampling was done. Sustained onshore winds occurred only during peak settlement, suggesting that wind drift currents may concentrate larvae and influence patterns of larval settlement.     

The relations between larval fishes and zooplankton in two inshore areas of the Gulf of Maine

Larval fishes and zooplankton were sampled in two hydrographicallydifferent areas on the coast of the Gulf of Maine: SullivanHarbor, an embayment in eastern Maine, and the DamariscottaRiver estuary in western Maine. Sampling was conducted at weeklyintervals from late winter to early summer in each area in 1979and 1980. Phytoplankton chlorophyll concentrations were determinedin each area in 1979. The time of peak catch rates of the dominantlarval fish species occurred one to three weeks earlier in thewestern sample area, the Damariscotta estuary, than in SullivanHarbor in the east. The phytoplankton and zooplankton bloomsalso occurred one to three weeks earlier in the Damariscottaestuary than in Sullivan Harbor. These timing trends are believedto result from the differences in the seasonal hydrographicchanges of the inshore and coastal source waters. Analyses ofthe feeding, length-frequencies, and condition factors of thedominant larval fish species, Pholis gunnellus, are used torelate the apparent survival of the larvae to the timing ofappearance of their forage organisms, the dynamics of whichare determined by the local hydrography and resultant phytoplanktondynamics.     

Molecular quantification of toxic Alexandrium fundyense in the Gulf of Maine using real-time PCR

The toxic dinoflagellate Alexandrium fundyense is widespread in the northeastern part of North America, including the Gulf of Maine, and is responsible for seasonal harmful algal blooms in these regions. Even at low cell densities, A. fundyense toxins can accumulate in shellfish and result in paralytic shellfish poisoning (PSP). PSP can be debilitating or lethal to humans and other shellfish consumers and is a public health concern. As a result, accurate measurements of A. fundyense distributions, particularly at low cell density, are critical to continued PSP monitoring and mitigation efforts. Towards this end we have developed a real-time quantitative PCR (qPCR) method to monitor A. fundyense. Laboratory validation indicates that the qPCR assay is sensitive enough to detect 10 cells per sample, and that it does not detect co-occurring dinoflagellates such as Alexandrium ostenfeldii. The qPCR methodology was used to quantify A. fundyense cell densities in samples collected during a spring 2003 transect in the Gulf of Maine, and the data were compared to those obtained in parallel from light microscope and DNA hybridization-based methods. Results show that A. fundyense cell density was low during this period relative to typical cell densities required for PSP contamination of local shellfish, and that qPCR values were comparable to numbers determined by independent methods.