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     

Biodiversity and ecosystem function: the consumer connection

Proposed links between biodiversity and ecosystem processes have generated intense interest and controversy in recent years. With few exceptions, however, empirical studies have focused on grassland plants and laboratory aquatic microbial systems, whereas there has been little attention to how changing animal diversity may influence ecosystem processes. Meanwhile, a separate research tradition has demonstrated strong top‐down forcing in many systems, but has considered the role of diversity in these processes only tangentially. Integration of these research directions is necessary for more complete understanding in both areas. Several considerations suggest that changing diversity in multi‐level food webs can have important ecosystem effects that can be qualitatively different than those mediated by plants. First, extinctions tend to be biased by trophic level: higher‐level consumers are less diverse, less abundant, and under stronger anthropogenic pressure on average than wild plants, and thus face greater risk of extinction. Second, unlike plants, consumers often have impacts on ecosystems disproportionate to their abundance. Thus, an early consequence of declining diversity will often be skewed trophic structure, potentially reducing top‐down influence. Third, where predators remain abundant, declining diversity at lower trophic levels may change effectiveness of predation and penetrance of trophic cascades by reducing trait diversity and the potential for compensation among species within a level. The mostly indirect evidence available provides some support for this prediction. Yet effects of changing animal diversity on functional processes have rarely been tested experimentally. Evaluating impacts of biodiversity loss on ecosystem function requires expanding the scope of current experimental research to multi‐level food webs. A central challenge to doing so, and to evaluating the importance of trophic cascades specifically, is understanding the distribution of interaction strengths within natural communities and how they change with community composition. Although topology of most real food webs is extremely complex, it is not at all clear how much of this complexity translates to strong dynamic linkages that influence aggregate biomass and community composition. Finally, there is a need for more detailed data on patterns of species loss from real ecosystems (community “disassembly” rules).     

The role of zooplankton in the functioning of the Taganrog Gulf ecosystem in the Sea of Azov

The taxonomic composition, distribution of biomass and seasonal variability of zooplankton were studied in various areas of the Taganrog Gulf of the Sea of Azov. The structural-functional transformation of zooplankton was analyzed for ecosystems of various levels of organization (the hypereutrophic central and eutrophic western parts). Unusually strong water blooming caused by cyanobacteria and the pressure of predatory comb jelly results in the elimination of zooplankton, the simplification of the trophic web, and the loss of the ecosystem’s capacity for biotic self-purification. The considerable participation of microzooplankton, especially that of tintiniids, in the metabolism of the Taganrog Gulf ecosystem is one of the important features of its sustainable functioning.     

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.     

Spatial and temporal patterns of zooplankton on baleen whale feeding grounds in the southern Gulf of Maine

This study addresses the regional variation of zooplankton inthe Great South Channel area in the southern Gulf of Maine betweenCape Cod, Massachusetts and Georges Bank. This is a region ofparticular interest because of the intense concentrations ofthe copepod Calanus finmarchicus in the spring, along with theco-occurrence of right whales that feed upon these copepod aggregations.Zooplankton in the Great South Channel were sampled with theMOCNESS plankton sampler during spring 1988 and 1989 as partof the SCOPEX (South Channel Ocean Productivity Experiment)project. Zooplankton variation was addressed through comparisonsof taxonomic composition and water column abundances among towswithin and between years, and between locations with and withoutright whales. Results showed that zooplankton community compositionwas highly similar between tows within each year average percentsimilarity index (PSI) for pairwise comparisons of tows = 82.2and 88.8% in 1988 and 1989, respectively] as well as betweenyears (mean PSI = 84.4). The copepods C.finmarchicus and Pseudocalanusspp. dominated the zooplankton in terms of total water columnabundance (>94% of all zooplankton), with C.finmarchicuscomprising an average of 84% of the copepods. Highest abundancesof these copepods (particularly the younger life stages) coincidedwith a region of low-salinity transport from the north. In addition,these copepods had higher abundances in 1989, which may be relatedto the fact that low-salinity transport was approximately twiceas large in 1989 as in 1988. Given the physical dynamics ofthe region, it is possible that developing copepod populationswere adverted into the Great South Channel from the northwesternGulf of Maine via the low-salinity plume. Each year, whaleswere located in areas of both high and low copepod abundance,and tended to concentrate near the leading edge of the low-salinityplume. In 1988, there were no significant differences in zooplanktonabundance between right whale areas and non-whale areas forany taxon. In 1989, whale areas had a greater proportion ofC.finmarchicus relative to other zooplankton in 1989, whichsuggests that whales preferred regions enriched in C.finmarchicus.     

Biodiversity change and ecosystem function in tropical forests

Recent debate about the fate of tropical forests has focused attention on the consequences of forest degradation and fragmentation for their diversity and composition, and the likely functional consequences of these changes. Existing data suggest that the responses of tropical forest plant and animal communities to habitat change are idiosyncratic, although a few consistent patterns are emerging. In particular, it is apparent that conventional diversity and richness metrics may not adequately represent anthropogenic changes to community structure and organisation. A widespread trend is towards ‘biotic homogenisation’: while disturbed forests may often have an equal or even a greater number of species than undisturbed forests, these species are typically drawn from a restricted pool; and endemic, restricted-range or habitat-specialist species are most likely to decline or go extinct. Similarly, studies have documented marked changes in the structure of food webs, even where the richness and diversity of component species remains little altered. What are the likely consequences of such changes for the important ecosystem functions performed by biodiversity, such as pollination and decomposition? Much of the extensive literature on the relationship between biodiversity and ecosystem function is of limited utility for answering this question, because experimental designs do not consider species-specific contributions to ecosystem function, abundance, degree of redundancy, or extinction-proneness; and few such studies have been carried out under realistic levels of diversity under field conditions, particularly in high-diversity ecosystems such as tropical forests. Furthermore, the focus has almost always been on richness as the explanatory variable, rather than the composition or structural attributes of communities. I briefly review recent papers that have begun to tackle these important issues, and consider how future research might help us understand the functional consequences of realistic changes to species composition and food-web ‘biostructure’ in tropical forests.     

Biodiversity of Collembola and their functional role in the ecosystem

More than 6500 species of Collembola are known from throughout the world and these are only a small part of the still undescribed species. There are many checklists and catalogues of Collembola for smaller territories and entire continents. Biogeographical analyses have been made for some genera and smaller territories. The most serious problems for a global biogeographical analysis is the lack of enough records from immense territories of all continents. Local biodiversity of Collembola can be very high, reaching over 100 species in small mountain ranges. Sampling methods do not impede documenting biodiversity on a global scale. Collembola have well differentiated ecomorphological life-forms and feeding guilds which enable the functional role that Collembola play in ecosystems to be recognised in some degree. Collembola play an important role in plant litter decomposition processes and in forming soil microstructure. They are hosts of many parasitic Protozoa, Nematoda, Trematoda and pathogenic bacteria and in turn are attacked by different predators. They utilise as food Protozoa, Nematoda, Rotatoria, Enchytraeidae, invertebrate carrion, bacteria, fungi, algae, plant litter, live plant tissues, and some plant pathogens. Soil acidification, nitrogen supply, global climate change and intensive farming have greatly impacted collembolan diversity.     

Organisation in the pelagic ecosystem

A continuous, steady-state theory has been developed for the abundance of organisms in the pelagic ecosystem as a function of their body weight. It is based on accepted relationships for the weight-dependence of metabolism and growth, in a context where individual organisms are assigned to one of a series of size classes for which the nominal weights increase in a geometric progression. Analysis of the biomass flow in such a representation leads to the conclusion that, in the steady state, the total biomass in any given size class decreases in a regular manner with increasing size. Explicitly,b(w ₂)/b(w ₁)~(w ₂/w ₁)^0.22, whereb(w ₂) andb(w ₁) are the total biomasses in the size classes characterised by weightsw ₂ andw ₁, respectively. The exponent (–0.22) represents a balance between catabolism and anabolism, based on published reviews concerning the revelant parameters. This result agrees favourably with data collected by other workers in the subtropical oceans. The theory can be used to draw conclusions about the functional dynamics of the pelagic ecosystem, such as community respiration and rate of biomass flow.     

Salinity and fish effects on Salton Sea microecosystems: zooplankton and nekton

The Salton Sea is the largest inland lake in California. Currently (1997) the salinity of the lake is about 44 g l-1 and is increasing gradually as a result of continued agricultural wastewater inflows, high evaporation rates, and lack of an outlet. A microcosm experiment was carried out to determine the effects of salinity (30, 39, 48, 57, and 65 g l-1) on Salton Sea algae and invertebrates in outdoor aquatic microcosms. The experiment was also designed to assess the effects of tilapia ( Oreochromis mossambicus) on this community at two of these salinities (39 and 57 g l-1). Fiberglass tanks containing Salton Sea water were adjusted to the appropriate salinity by the addition of salts, identically inoculated with organisms from the Salton Sea and other saline water bodies in the region, and monitored for 15 months. Planktonic and nektonic invertebrates were sampled monthly at night from the upper part of the water column. The dominant invertebrates present were Gammarus mucronatus, Artemia franciscana, Trichocorixa reticulata, and an assemblage of ciliate protozoans. Gammarus decreased and Trichocorixa increased with increasing salinity. Artemia was present only at the two highest salinities. Rotifers, harpacticoid and cyclopoid copepods, barnacle larvae, and protozoans all showed marked and varied responses. During the latter half of the experiment, the invertebrate assemblage was dominated by Gammarus at 30 and 39 g l-1, by protozoans at 48 g l-1, and by protozoans and Trichocorixa at 57 and 65 g l-1. The presence of tilapia caused a 99 percent reduction in Gammarus at 39 g l-1 and a 70–90 percent decrease in Trichocorixa at 57 g l-1. These were accompanied by substantial increases in rotifers, copepods, and certain protozoans, and decreases in other protozoans. As the salinity of the Salton Sea continues to increase, large changes in the invertebrate populations are expected. This study suggests that the principal change would be an increase in Trichocorixa densities, the loss of Gammarus, and the appearance of Artemia at about 60–70 g l-1, when both fish and invertebrate predators are likely to be scarce or absent. Protozooplankton abundance is likely to increase when tilapia declines and later decrease when and if large Artemia populations develop. This revised version was published online in July 2006 with corrections to the Cover Date.     

The correlation of downwelling irradiance and staggered vertical igration patterns of zooplankton in Wilkinson Basin, Gulf of Maine

Field studies on the characteristics of light that influencevertical migrations in the mesopelagic realm are sparse, dueto the difficulty in simultaneously monitoring changes in speciesdistributions with changes in downwelling irradiance. Usingthe Johnson-Sea-Link submersible as a platform, in situ measurementsof the changes in downwelling irradiance at sunset were madesimultaneously with observations on changes in animal distributionpatterns in Wilkinson Basin, Gulf of Maine. The results indicatethat the vertical migrations for several species of large zooplanktonare staggered, with euphausiids (Meganyctiphanes norvegica)migrating first, cydippid ctenophores (Euplokamus) migratingnext, and two species of caridean shrimp (Dichelopandalus lepiocerusand Pasiphaea multidentata) migrating last Data collected ondaytime dives indicate that the daytime depth distribution isnot solely responsible for the migration order, and that differentspecies may be responding to different cues, or have differentthresholds for the same cue.     

Feeding ecology of pelagic fish species in the Gulf of Riga (Baltic Sea): the importance of changes in the zooplankton community

The feeding ecology of four pelagic fish species was studied in relation to their prey availability in the Gulf of Riga (Baltic Sea) during the summer 1999-2006. The zooplankton community was dominated by the cladoceran Bosmina longispina, rotifers Keratella cochlearis and K. quadrata and the copepod Eurytemora affinis, with the highest interannual variability in abundance recorded for B. longispina. The last influenced the diet of adult sprat Sprattus sprattus, juvenile smelt Osmerus eperlanus and three-spined stickleback Gasterosteus aculeatus as these were strongly selecting for B. longispina. The fish feeding activity did not match the abundance dynamics of their preferred prey, suggesting that fishes may switch to consume other prey in case the preferred diet was limited. A considerable dietary overlap indicated high potential competition between pelagic fish species. While herring Clupea harengus membras and G. aculeatus were relying on very different food, the diets of young O. eperlanus and G. aculeatus were very similar. Interannual variability in zooplankton composition and abundance significantly affected the diet composition of fishes, but those changes were insufficient to exert a consistent influence upon fish feeding activity and total amounts of zooplankton consumed.     

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 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).     

Biodiversity in the Gulf of Guinea: an overview

Patterns of species-richness and endemism in the Gulf of Guinea reflect the region's biogeographic history. Bioko is a continental-shelf island that was recently connected to the African mainland, whereas Príncipe, São Tomé and Annobón are truly oceanic and have never been connected with each other or with the mainland. As a result, Bioko supports a much more diverse flora and fauna but with relatively low levels of endemism at the species level, whereas the oceanic islands are relatively depauperate because of their isolation but rich in endemic taxa. Species endemism is 0–3% on Bioko for angiosperms, bats, birds, reptiles and amphibians, compared with much higher values on Principe for these same taxa of 8% (plants) to 100% (amphibians), on São Tomé between 14% (plants) and 100% (amphibians), and on Annobón 0% (bats) to 71% (reptiles). On a global scale, for their size both Príncipe and São Tomé support unusually high numbers of single-island endemic species of birds, reptiles and amphibia. For its tiny size, Annobón is also notable for its endemic birds and reptiles. Among terrestrial molluscs the rates of endemism are in general higher than for plants and vertebrates, from ca 50% on Bioko to ca 80% on the oceanic islands. In contrast and as might be expected, only Bioko supports a rich freshwater fish fauna and it contains many endemic taxa, whereas the oceanic islands support only a few salt-tolerant species. The Gulf of Guinea islands are also important for their marine organisms, amongst which coral reef fish and marginellid molluscs show high levels of endemism, though they are not especially species-rich. The Gulf of Guinea islands are of great interest to conservationists and evolutionary biologists. Each island, of greatly differing size and degree of isolation, has acquired its unique sub-set of plants and animals separately from the neighbouring mainland, followed by adaptive radiations in situ. For this reason the conservation value of the archipelago as a whole is greater than the sum of the biodiversity contained in its individual islands. Conservation initiatives in the Gulf of Guinea should therefore ensure that representative terrestrial, freshwater and marine habitats and groups of organisms are targeted in a co-ordinated manner among the islands.     

Biodiversity of the deep-sea continental margin bordering the Gulf of Maine (NW Atlantic): relationships among sub-regions and to shelf systems

Background

In contrast to the well-studied continental shelf region of the Gulf of Maine, fundamental questions regarding the diversity, distribution, and abundance of species living in deep-sea habitats along the adjacent continental margin remain unanswered. Lack of such knowledge precludes a greater understanding of the Gulf of Maine ecosystem and limits development of alternatives for conservation and management.

Methodology/Principal Findings

We use data from the published literature, unpublished studies, museum records and online sources, to: (1) assess the current state of knowledge of species diversity in the deep-sea habitats adjacent to the Gulf of Maine (39–43°N, 63–71°W, 150–3000 m depth); (2) compare patterns of taxonomic diversity and distribution of megafaunal and macrofaunal species among six distinct sub-regions and to the continental shelf; and (3) estimate the amount of unknown diversity in the region. Known diversity for the deep-sea region is 1,671 species; most are narrowly distributed and known to occur within only one sub-region. The number of species varies by sub-region and is directly related to sampling effort occurring within each. Fishes, corals, decapod crustaceans, molluscs, and echinoderms are relatively well known, while most other taxonomic groups are poorly known. Taxonomic diversity decreases with increasing distance from the continental shelf and with changes in benthic topography. Low similarity in faunal composition suggests the deep-sea region harbours faunal communities distinct from those of the continental shelf. Non-parametric estimators of species richness suggest a minimum of 50% of the deep-sea species inventory remains to be discovered.

Conclusions/Significance

The current state of knowledge of biodiversity in this deep-sea region is rudimentary. Our ability to answer questions is hampered by a lack of sufficient data for many taxonomic groups, which is constrained by sampling biases, life-history characteristics of target species, and the lack of trained taxonomists.     

Bioluminescence of sound-scattering layers in the Gulf of Maine

Submersible-based investigations of bioluminescence were conductedin sound-scattering layers (SSLs) in the Gulf of Maine, usingintensified video and dual-beam acoustic methods. Stimulatedbioluminescence in the SSLs was high (3–41 µW sr^–1m^–3 while spontaneous bioluminescence was not detected.The average intensity of individual bioluminescent sources inthe SSLs was 30–200 times greater than the intensity oflight emitters outside the SSLs. The two brightest sources ofbioluminescence were identified as the euphausiid, Meganyctiphanesnorvegica and the cydippid ctenophore, Euplokamis sp. Meganyctiphanesnorvegica formed a SSL within 50 m of the bottom during theday and migrated to the uppermost 30 m of the water column atnight, forming a near-surface SSL. Euplokamis sp., which producedexceptionally intense and long-lasting bioluminescent secretions,occurred within the near-bottom SSL in concentrations up to7 m^–3. Our findings indicate that traditional methodsof identifying the primary light emitters in a region, basedon light measurements from net- or pump-captured organisms,may have underestimated the significant in situ bioluminescencepotential of euphausiids and gelatinous zooplankton. ³ Present address: NOAA/NURP/R-OR2, Silver Spring, MD 20910,USA