Correlation of primary production as measured aboard ship in Southern California Coastal waters and as estimated from satellite chlorophyll images

Chlorophyll images suitable for the regional analysis of the Southern California Bight were obtained by satellite (Nimbus-7 Coastal Zone Color Scanner) on 27 February, 6 March and 11 March 1979. An algorithm to convert satellite chlorophyll data to primary production has been developed. In addition, primary production was measured along with other biological and physical factors from ships cruising in the Southern California Bight during late February and early March 1979. There was fair agreement between shipboard measurements and satellite estimations; for example, primary production (mg carbon m^-2 d^-1) averaged 554 mg from shipboard measurement and 403 mg from satellite estimation when averaged for the entire Bight for the course of the study. The high standard error associated with the variance in productivity index used in our algorithm prevented more accurate conversion of chlorophyll concentrations into primary productivity. However, the synoptic data provided by concurrent ship and satellite measurements can be used to quantify, and perhaps reduce, sampling errors associated with ship sampling alone and to provide estimates of the accuracy of primary productivity on a regional basis.     

Food availability as a selective factor on the chemical compositions of midwater fishes in the eastern North Pacific

In September and October 1980 we examined the relationships between food availability, depth, and chemical composition among 12 midwater fish species, from three adjacent areas of the eastern North Pacific: the eastern gyre, the California Current, and the transitional region between them. By comparing trends in chemical composition across a geographical productivity gradient, the influence of food availability could be examined both with and without depth as a dependent variable. In general, caloric density, lipids, and water content showed consistent trends along both vertical and horizontal gradients of food availability. Lipids and caloric contents were lowest among bathypelagic species and among fishes from the gyre. Water content was highest in the gyre and among deeper-living species. While protein content as a percentage of the total wet weight declined with depth, there was no clear trend among mesopelagic species between stations. Protein as percent of the total organic content showed no decrease with depth, but was significantly higher in the gyre among mesopelagic species. These findings suggest that compositional trends in water, lipids, and caloric content are correlated with food availability, regardless of any depth-related factors. In contrast, protein did not vary consistently with food availability and thus may be linked to other factors.     

Diurnal rhythms in netplankton and nannoplankton assimilation ratios

Surface nannoplankton and netplankton photosynthetic rates and chlorophyll a concentrations were measured 3 h before and 3 h after local apparent noon at 36 stations in the eastern Pacific. Morning and afternoon nannoplankton and netplankton photosynthetic capacities were plotted against chlorophyll a concentrations. Comparison of am (morning) and pm (afternoon) regression coefficients revealed that neither netplankton nor nannoplankton photosynthetic efficiencies varied diurnally in temperate California Current waters. However, in oligotrophic tropical surface waters where the photosynthetic rates of both fractions were nitrogen limited, netplankton assimilation ratios exhibited an afternoon maximum, while nannoplankton efficiencies exhibited a morning maximum. The netplankton followed the same pattern in eutrophic tropical surface waters with high nitrate concentrations, but nannoplankton assimilation ratios were highest in the afternoon. Assuming that midday light intensities inhibit surface photosynthesis, it was concluded that assimilation ratios will reach a mid-morning or mid-afternoon maximum depending upon the relative importance of netplankton and nannoplankton productivity and the degree to which nutrients are limiting.     

Relationships between nucleic acid concentrations and primary production in the Catalan Sea (Northwestern Mediterranean)

We explored the relationships between classical estimators of autotrophic biomass and primary production, such as chlorophyll a concentration and ¹⁴C-fixation rates, and biochemical indices based on DNA and RNA determinations, which have been proposed as indicators of physiological state in natural plankton populations. The measurements were made during two cruises across the Catalan Front, carried out in May 1989 and February 1990, corresponding respectively, to periods of stratification and moderate mixing. DNA and RNA concentrations (measured by a double-staining fluorimetric technique) were significantly correlated with chlorophyll a in February 1990, but not in May 1989, when a marked deep chlorophyll maximum was present. Significant positive correlations between RNA concentration and primary production and between RNA: DNA and primary production were found during both surveys, probably reflecting both higher RNA concentrations per cell and enhanced bacterial and microheterotrophic growth in high primary production situations. The results support the potential usefulness, in biological oceanography, of biochemical indicators based on DNA and RNA concentrations.     

Albatross species demonstrate regional differences in North Pacific marine contamination.

Recent concern about negative effects on human health from elevated organochlorine and mercury concentrations in marine foods has highlighted the need to understand temporal and spatial patterns of marine pollution. Seabirds, long-lived pelagic predators with wide foraging ranges, can be used as indicators of regional contaminant patterns across large temporal and spatial scales. Here we evaluate contaminant levels, carbon and nitrogen stable isotope ratios, and satellite telemetry data from two sympatrically breeding North Pacific albatross species to demonstrate that (1) organochlorine and mercury contaminant levels are significantly higher in the California Current compared to levels in the high-latitude North Pacific and (2) levels of organochlorine contaminants in the North Pacific are increasing over time. Black-footed Albatrosses (Phoebastria nigripes) had 370-460% higher organochlorine (polychlorinated biphenyls [PCBs], dichlorodiphenyltrichloroethanes [DDTs]) and mercury body burdens than a closely related species, the Laysan Albatross (P. immutabilis), primarily due to regional segregation of their North Pacific foraging areas. PCBs (the sum of the individual PCB congeners analyzed) and DDE concentrations in both albatross species were 130-360% higher than concentrations measured a decade ago. Our results demonstrate dramatically high and increasing contaminant concentrations in the eastern North Pacific Ocean, a finding relevant to other marine predators, including humans.     

A comparison of zooplankton patterns in the California Current and North Pacific Central Gyre

This study compares the small-scale (100's to 1000's of meters) horizontal spatial distribution of zooplankton from stations in the California Current (29°N; 118°W) and in the North Pacific Central Gyre (31°N; 155°W). Patterns were inferred from the spacing of vertical tows and the variability in abundance of 15 taxonomic categories caught in these tows. In the California Current, 4 tows were taken at random positions within 2000 m of a drogue or fixed geographic position and 4 replicate tows were taken at the drougue or the fixed point. Four series of these 8 tows were taken around noon and midnight during two days. At the Central Gyre station, two similar series (one day and one night) were taken while following a drogue. Estimates of the scale of structure based on comparisons of replicate and random tow variability suggest that aggregated patterns in the Central Gyre are larger than a few hundred meters, while the California Current structures may be less than 100 m in size. Day-night changes in patch size were not apparent in either area. Evidence for multispecies patches was found only for the Central Gyre station. Replication samples in both areas gave more consistent measures of community structure (relative proportions of species) than the random tows. Similarity of community structure decreased with increasing distance between tows being compared. The decrease was much greater for the California Current station, a further indication of smaller patch size in this area. Diurnal changes in community structure occurred in both areas.Based in part on a dissertation submitted in partial fulfillment of the requirements of the Ph.D. degree at the University of California, San Diego. Portions of this research were supported by the Marine Life Research Program of the Scripps Institution of Oceanography.     

Copepod egg production in the Skagerrak during SKAGEX,May–June 1990

Egg production ofAcartia clausi andCentropages hamatus was measured along 6 transects in the Skagerrak every third day from 26 May to 20 June 1990. Egg production was highest in the shallow waters north of Denmark, with occasional peaks in frontal regions along the Swedish and the Danish west coasts. Linear regression analysis showed that the egg production was significantly (p<0.05) related to chlorophylla measured either as average surface concentration or integrated over the whole water column. When analysing each transect or each time period separately, the surface chlorophyll generally was a better predictor of egg production than the depth-integrated chlorophyll. Regressions improved when analysing the whole area for a short period of time rather than analysing a single transect for a month. The data suggest that the Skagerrak planktonic system functions more similarly over the whole area in a short period of time than over a month in a restricted area. Mixingwithin the system in frontal regions or in connection with eddies is more important for the secondary production thantransport to it by the Baltic Current or the Jutland Current.SKAGEX Contribution No. 4     

Dye tracing of a subsurface chlorophyll maximum of a red-tide dinoflagellate to surface frontal regions

Rhodamine dye (3701) was injected into a 22-m subsurface chlorophyll maximum of the red-tide forming dinoflagellate Prorocentrum mariae-lebouriae var. minimum in the northern Chesapeake Bay (USA) and traced for a sixday period as it spread over a 600 km² area. The precise physical mechanisms, which resulted in the transfer of dye and organisms to the surface, are documented. The major component of the dye and organisms was transported from the central bay into major tributary estuaries via net nontidal flow of bottom waters and surfaced upstream in frontal regions. Once in surface waters, the dye and organisms flowed downriver toward the bay. Due to the three-layer flow of the rivers at this time, the dinoflagellate and the rhodamine re-entered the bay proper at middepth below the fresher Susquehanna plume, thus forming a near-surface chlorophyll maximum (4–6 m) flowing in an opposite direction to the deep subsurface chlorophyll maximum (18–26 m). Current meter arrays verified the opposite flows of these two lenses. The near-surface, southward-flowing lens was followed downstream to an area where the influence of the Susquehanna begins to subside as indicated by isopycnals inclined to the surface. Here the near-surface lens is mixed upward to the surface forming massive red tides (25 000 cells ml^–1) delineated by a frontal region. In addition to the predominant along channel flow, major cross stream Ekman transport and upwelling of dye and organisms was detected in response to wind forcing resulting in localized surface patch formation along the western shore shoaling regions of the bay proper. Thus, annual variations in the locations of surface red tides can be correlated to streamflow and windinduced variations in the locations of frontal regions.     

Primary productivity in the Gulf of California

Primary productivity data from 6 cruises in the Gulf of California are summarized. A total of 31 stations with 133 samples was available. The Gulf of California represents a subtropical area with exceptionally high rates of primary productivity. This fact may be accounted for by temporary seasonal upwelling, a wind-mixed water column down to the shallow thermocline, especially in the northern part of the Gulf, and by tidal mixing, especially in the Ballenas Channel between Baja California and the island Angel de la Guardia. Rates of primary productivity in the Gulf of California are comparable to those in areas such as the Bay of Bengal, the upwelling areas off the west coast of Baja California, or North Africa. They are about 2 to 3 times greater than that in the open Atlantic or the open Pacific at similar latitudes.Contribution from the Scripps Institution of Oceanography, University of California, San Diego. This work was part of the Scripps Tuna Oceanography Research Program. It was supported by the Bureau of Commercial Fisheries under contract number 14-17-0007-963. Shiptime was provided from funds of the National Science Foundation.     

Population structure of the widely dispersing marine bryozoan Membranipora membranacea (Cheilostomata): implications for population history, biogeography, and taxonomy

Morphologically plastic, cryptic, or geographically widespread species pose similar challenges to the evolutionary biologist: their taxonomic status is often unclear yet must be known to study almost any aspect of their biology, ecology, evolution, or biogeography. The marine bryozoan Membranipora membranacea (L.) is morphologically plastic and geographically widespread in temperate oceans of the Northern and Southern Hemispheres, and its taxonomy is unclear. This study examined genetic relationships among allopatric populations and sympatric morphs of this species, or species complex. Colonies were collected from 1992 to 1995. Allozymes were used to elucidate the relationships among four widely separated populations, two in the North Atlantic and two in the North Pacific Ocean. Allozymes and mtDNA sequencing were used to clarify the genetic relationships among three sympatric morphs that might correspond to the species M. villosa Hincks and M. membranacea in the northeastern Pacific (Washington State). Populations in the North Atlantic and North Pacific had no fixed allelic differences at the loci tested but were separated by an average Nei's genetic distance of 0.581, suggesting their near-sibling species status. Populations from Friday Harbor (Washington) and Catalina Island (California) were not significantly differentiated, which was attributed to high gene flow. Populations on either side of the North Atlantic were genetically indistinguishable, which is most likely due to the recent establishment of the West Atlantic populations from European founders. At Friday Harbor, sympatric morphs varying in their spination and spine inducibility were genetically indistinguishable, supporting the hypothesis that M. villosa is an induced phenotype of M. membranacea and not a distinct species in the northeastern Pacific. Since such phenotypic plasticity is common in cheilostome bryozoans, the morphospecies concept must be used with caution. Received: 31 August 1998 / Accepted: 10 August 1999     

Abundance of picophytoplankton in the subsurface chlorophyll maximum layer in subtropical and tropical waters

A distinctive chlorophyll maximum was detected around 60-m depth in the western North Pacific Ocean and the South China Sea, and almost 55% of the total chlorophyll in the entire water column was found within 50 m around the subsurface chlorophyll maximum (SCM) layer. More than 70% of the chlorophyll was contained in picoplankton which passed through a 3-m Nuclepore but retained on 0.22-m Millipore filters at the SCM as well as the surface layers. By transmission electron microscopic observations, the picoplankton were identified as aChlorella-like coccoid green alga having a section size of 1.2 to 1.5 m and cyanobacteria of 0.5 to 2 m. No obvious difference in these two dominant groups was observed in the SCM and the surface samples except in numerous and heavily stacked thylakoids in the former samples.     

Identification of distinct movement patterns in Pacific leatherback turtle populations influenced by ocean conditions

Interactions with fisheries are believed to be a major cause of mortality for adult leatherback turtles (Dermochelys coriacea), which is of particular concern in the Pacific Ocean, where they have been rapidly declining. In order to identify where these interactions are occurring and how they may be reduced, it is essential first to understand the movements and behavior of leatherback turtles. There are two regional nesting populations in the East Pacific (EP) and West Pacific (WP), comprising multiple nesting sites. We synthesized tracking data from the two populations and compared their movement patterns. A switching state-space model was applied to 135 Argos satellite tracks to account for observation error, and to distinguish between migratory and area-restricted search behaviors. The tracking data, from the largest leatherback data set ever assembled, indicated that there was a high degree of spatial segregation between EP and WP leatherbacks. Area-restricted search behavior mainly occurred in the southeast Pacific for the EP leatherbacks, whereas the WP leatherbacks had several different search areas in the California Current, central North Pacific, South China Sea, off eastern Indonesia, and off southeastern Australia. We also extracted remotely sensed oceanographic data and applied a generalized linear mixed model to determine if leatherbacks exhibited different behavior in relation to environmental variables. For the WP population, the probability of area-restricted search behavior was positively correlated with chlorophyll-a concentration. This response was less strong in the EP population, but these turtles had a higher probability of search behavior where there was greater Ekman upwelling, which may increase the transport of nutrients and consequently prey availability. These divergent responses to oceanographic conditions have implications for leatherback vulnerability to fisheries interactions and to the effects of climate change. The occurrence of leatherback turtles within both coastal and pelagic areas means they have a high risk of exposure to many different fisheries, which may be very distant from their nesting sites. The EP leatherbacks have more limited foraging grounds than the WP leatherbacks, which could make them more susceptible to any temperature or prey changes that occur in response to climate change.     

Temporal separation of two fin whale call types across the eastern North Pacific

Fin whales (Balaenoptera physalus) produce a variety of low-frequency, short-duration, frequency-modulated calls. The differences in temporal patterns between two fin whale call types are described from long-term passive acoustic data collected intermittently between 2005 and 2011 at three locations across the eastern North Pacific: the Bering Sea, off Southern California, and in Canal de Ballenas in the northern Gulf of California. Fin whale calls were detected at all sites year-round, during all periods with recordings. At all three locations, 40-Hz calls peaked in June, preceding a peak in 20-Hz calls by 3–5 months. Monitoring both call types may provide a more accurate insight into the seasonal presence of fin whales across the eastern North Pacific than can be obtained from a single call type. The 40-Hz call may be associated with a foraging function, and temporal separation between 40- and 20-Hz calls may indicate the separation between predominately feeding behavior and other social interactions.     

Deep chlorophyll maximum distribution in the central Tyrrhenian Sea described by a towed undulating vehicle

The towed undulating vehicle (TUV), named SARAGO, was used for two fine-scale surveys between the Italian and the Sardinian coasts during the Astraea 2 cruise (6-7 and 26-27 September 1995), studying the deep chlorophyll maximum distribution. SARAGO sections identify a sub-surface doming with higher chlorophyll a and primary production concentrations in the upwelling area of a cyclonic gyre region, detected by sea-surface temperature images. In the first section, the cyclone presents a double doming, in density and salinity, with shallower and concentrated patches of chlorophyll a for about 2 miles. Twenty days later, the second section shows that the gyre changes shape and extension, showing a single doming with higher primary production and chlorophyll a concentrations, distributed over a large area of about 40 nautical miles. SARAGO allows analysis of this high-variability phenomenon (cyclonic gyre) and allows concentrated patches (2 nm) to be identified, thus proving the importance of TUVs in the study of mesoscale processes.     

Abundance and production rates of floating diatom mats (Rhizosolenia castracanei and R. imbricata var. shrubsolei) in the Eastern Pacific Ocean

Abundance and production rates were measured on freefloating mats composed of the diatoms Rhizosolenia castracanei and R. imbricata var. shrubsolei in the California Current and boundary waters of the central North Pacific during October, 1980. Mats ranged from 3.0 to 10.6 cm in maximum length and had a mean volume of 6.4 ml. Production rates of the diatom mats averaged 4.0 g C colony^-1 h^-1, more than 10³ times higher than that of an equal volume of surrounding water. However, because of their low density at these sites, about 1 mat m^-3, diatom mats contributed only about 1% of the total primary production. The two large Rhizosolenia species comprised almost all of the phytoplankton biomass within the mats. Rhizosolenia species rarely occurred in seawater between mats, where the phytoplankton community was dominated by a diverse array of nannoplankton. The Rhizosolenia species in the mats appeared to be in healthy condition and contained intracellular bacteria. The very high production rates of the colonies indicate high nutrient demand and, since these particular diatoms lack the symbiotic cyanobacterium Richelia intracellularis, which is implicated in N-fixation in other species of Rhizosolenia, our results indicate that other nutrient sources must be present. We discuss the potential role of the intracellular bacteria in nitrogen fixation.     

Phylogeography of the diamond turbot (Hypsopsetta guttulata) across the Baja California Peninsula

We compared morphology and sequenced nuclear and mitochondrial genes from 11 populations of a previously genetically unstudied “Baja California disjunct” species, the diamond turbot (Hypsopsetta guttulata). This species exhibits very limited adult movement and restriction to soft-bottom habitats but has a moderately long pelagic larval duration. Therefore, if pelagic larval duration is correlated with gene flow between Gulf of California and Pacific populations, we expect a reduced level of genetic and morphological differentiation. However, if adult habitat and ecology have more effect on gene flow, we expect the populations in the two bodies of water to be more highly differentiated. We used logistic regression to compare morphological features and phylogenetic and population genetic analyses to compare nucleotide sequence data. Gulf of California H. guttulata are different from Pacific populations in morphology and both mitochondrial and nuclear gene sequences. MtDNA shows reciprocal monophyly, and nuclear sequences from the Gulf of California formed a monophyletic group. Population genetic analyses also suggest further population subdivision within the Pacific and within the Gulf of California. We argue that adult ecology has a significant effect on migration rates among populations in the Pacific Ocean and the Gulf of California.     

Standing crops and vertical distribution of four groups of marine planktonic ciliates in relation to phytoplankton chlorophyll a

Standing crops and the vertical distribution of four groups of ciliates, autotrophic naked ciliates (ANC), heterotrophic naked ciliates (HNC ), mixotrophic naked ciliates (MNC) and loricated ciliates (LC ), were analysed in relation to phytoplankton chlorophyll a in the western Pacific. Data were gathered from the upper 60 to 200 m of water in the subarctic North Pacific in spring, in the subtropical North Pacific in spring, in Toyama Bay in summer, off eastern Australia in spring and off Sanriku in spring and fall. Of the four groups, the standing crop of HNC (cells l⁻¹) showed the highest correlation to phytoplankton chloro- phyll a [CHL, μg l⁻¹; r=0.66, n=365 (HNC=490 ×CHL ^0.653)]. Depths of the maximum layers of HNC and MNC were usually shallower than that of chlorophyll a, while those of ANC and LC occurred frequently beneath the chlorophyll maximum layer. This indicates that these four ciliate groups are substantially different from each other, and that each group holds a different ecological position in the marine microbial food web. Received: 27 October 1997 / Accepted: 27 June 1998     

Deep chlorophyll maximum distribution in the central Tyrrhenian Sea described by a towed undulating vehicle

The towed undulating vehicle (TUV), named SARAGO, was used for two fine-scale surveys between the Italian and the Sardinian coasts during the Astraea 2 cruise (6–7 and 26–27 September 1995), studying the deep chlorophyll maximum distribution. SARAGO sections identify a sub-surface doming with higher chlorophyll a and primary production concentrations in the upwelling area of a cyclonic gyre region, detected by sea-surface temperature images. In the first section, the cyclone presents a double doming, in density and salinity, with shallower and concentrated patches of chlorophyll a for about 2 miles. Twenty days later, the second section shows that the gyre changes shape and extension, showing a single doming with higher primary production and chlorophyll a concentrations, distributed over a large area of about 40 nautical miles. SARAGO allows analysis of this high-variability phenomenon (cyclonic gyre) and allows concentrated patches (2 nm) to be identified, thus proving the importance of TUVs in the study of mesoscale processes.     

Bioassay of biotin and its distribution in the sea

A simple, rapid and sensitive method of microbiological assay for the determination of biotin in seawater has been developed by using a marine bacterium, Achromobacter sp. (Strain yH-51) as an assay organism. When the organism was incubated at 25°C for 48 h in a synthetic medium with constant shaking, a standard reference curve of bioassay was obtained. The organism is sensitive in the range 0.1 to 8.0 ng/l biotin. The distribution of biotin has been determined in the East China Sea, the North Pacific Ocean, and the bays and inlets along the Pacific coast of Japan. Biotin concentrations in surface waters were in the range from less than 0.1 ng/l up to 57.9 ng/l; generally high in coastal waters and low in deep open waters. The concentrations were generally high in surface waters, and rapidly decreased with depth down to 50 m, below which values were generally low. The vertical distribution of biotin followed closely the concentration of chlorophyll a, with high concentrations of biotin restricted to the euphotic layer of high chlorophyll a contents at most stations observed. Biotin concentrations in surface waters were generally high during summer, but in autumn and winter the concentrations decreased markedly.