Environmental control of living symbiotic and asymbiotic

PALEOCEANOGRAPFIY, VOL. 10, 10,NO. NO.6, PALEOCEANOGRAPHY, VOL. 6, PAGES PAGES 987-1009, 987-1009, DECEMBER DECEMBER 1995 1995 Environmental control control of Environmental of living living symbiotic symbiotic and and asymbiotic asymbiotic foraminifera of the Current foraminifera of the California California Current J. J. D. D. Ortiz, Ortiz, A. A. C. C. Mix, Mix, and and R. R. W. W. Collier Collier College University, Corvallis Collegeof of Oceanic Oceanicand andAtmospheric AtmosphericSciences, Sciences,Oregon OregonState StateUniversity, Corvallis Abstract. tows from the northern California Current biological Abstract.Plankton Plankton tows from the northern California Currentconstrain constrain biologicaland andphysical physical influences influenceson on living living planktonic planktonicforaminifera. foraminifera.In Inthis thisregion, region,the thedominant dominantfactors factorscontrolling controllingthe the size offsizeand anddistribution distributionof of symbiotic symbioticand andasymbiotic asymbioticspecies speciesare arelight light and andfood. food.Food Fooddecreases decreases offshore. for photosynthesis, increases offshore as shore.Light, Light,needed needed forsymbiont symbiont photosynthesis, increases offshore aswater waterturbidity turbiditylessens. lessens. Asymbiotic right-coiling Neogloboquadrina Neogloboquadrina pachyderma, pachyderma, Globigerina Globigerina Asymbioticforaminifera foraminifera(e.g., (e.g.,right-coiling quinqueloba, and quinqueloba, andGlobigerina Globigerinabulloides), bulloides),which whichsurvive surviveby bygrazing, grazing,dominate dominatethe thecoastal coastalfauna. fauna. The these species, right-coiling Neogloboquadrina Neogloboquadrina pachyderma, did The most mostabundant abundantof of these species, right-coiling pachyderma, didnot notchange change in that in size sizein in response responseto to increasing increasingfood. food.Species Species thatbenefit benefitfrom fromsymbiont symbiontphotosynthesis photosynthesis (Orbulina dutertrei, ruber, glutinata) (OrbMinauniversa, universa,Neogloboquadrina Neogloboquadrina dutertrei,Globigerinoides Globigerinoides ruber,and andGlobigerinita Globigerinita glutinata) dominate are dominatethe the offshore offshorefauna. fauna.Individuals Individualsof ofthese thesespecies species arerare rareand andhave havesmaller smallershells shellsin in turbid turbid waters light is is limited. limited. G. of --14øC, "14°C, is waterswhere where light G. ruber, ruber,which whichis isnear nearits itsthermal thermaltolerance tolerancelimit limit of is the the only species to demonstrate a clear temperature response. Although temperature may control only speciesto demonstrate a cleartemperatureresponse.Althoughtemperaturemay controlaa foramimferal distribution near food foraminiferalspecies species' distribution nearthe thelimits limitsof ofits itsthermal thermaltolerance, tolerance, foodand andlight lightappear appearto to provide the primary primary control control under under more more favorable favorable thermal thermal conditions. conditions. We We infer infer that that gradients gradients in in providethe food sedimentary patterns foodand andlight lightcan canresult resultin in quantifiable quantifiable sedimentary patternsrelated relatedto tooceanic oceanicproductivity productivity through throughchanges changesin in plankton planktonbiomass biomassand andturbidity. turbidity. Introduction Introduction Approaches used to Approachesused to calibrate calibratethe therelationship relationshipbetween between environronmental conditions and environronmental conditions and planktonic planktonic foraminiferal foraminiferal distributions include sediment, and field distributionsinclude sediment,laboratory, laboratory,and field studies. studies. This field study assesses the relationships between This field study assessesthe relationshipsbetweenliving living planktonic foraminifera foraminifera and in the planktonic and environmental environmental conditions conditions in the California Specifically, we we investigate the relationCaliforniaCurrent. Current. Specifically, investigatethe relationships shipsamong among(1) (1) total totalforaminiferal foraminiferalstanding standingstock stockand andplankplank- that control the the distribution foraminifera. that control distribution of of living living foraminifera. Dissolution also from Dissolution also removes removes information information from the the sediments sediments and and obscures a species obscuresthe the primary primary relationship relationship between between a speciesand and its its environment. environment. Laboratory Laboratorystudies studiesculture cultureindividual individualforaminifera foraminiferaunder underconconTheir goal is to trolled environmental conditions. trolled environmental conditions. Their goal is to determine determine effects effects of of individual individual variables variableson on the thegrowth growthand andreproductive reproductive potential potential of of planktonic planktonic foraminifera foraminifera [e.g., [e.g., Be Bd et et al., al., 1981; 1981; Caron etetal., Caron al., 1987; 1987;Bijma Bijma et et al., al., 1990b]. 1990b]. However, However, it it can can be be ton biomass, ton biomass,(2) (2) species speciesstanding standingstock stockand andhydrographic hydrographic difficult difficult to to apply applyresults resultsfrom fromthese thesestudies studiesoutside outsidethe thelaboralaboravariability; and and (3) variability; (3) shell shellsize sizeand andenvironment. environment. tory tory where wheresuites suitesof of variables variablesmay mayproduce producesynergistic synergisticeffects effects Sediment calibration studies Sedimentcalibration studiescompare comparemodern modernenvironmenenvironmen- and by the and conditions conditions encountered encountered by the organism organism change change on on aa tal data of tal datato tocore coretop topmeasurements measurements offoraminiferal foraminiferalpercent percent variety variety of of timescales. timescales. For this reason, it remains necessary For this reason,it remainsnecessaryto to abundance, size, size, and/or and/or isotopic isotopic composition. composition. These abundance, Thesestudies studies augment laboratory studies with carefully conducted field augment laboratory studies with carefully conducted field include statistical estimates of a species' "optimum environinclude statisticalestimatesof a species'"optimumenviron- programs [e.g., Be et al., 1977]. Sampling tools used in field programs [e.g., Bd et al., 1977]. Samplingtools usedin field ment' [Hecht ment" [Hechtand andSavin, Savin,1972; 1972;Hecht, Hecht,1976; 1976;Molfino, Molfino, 1992] 1992] studies include sediment traps, plankton tow, and pumps. and statistical and statistical relationships relationships between between the the environment environment and and planktonic planktonic foraminiferal foraminiferal faunas faunas [Imbrie [Imbrie and and Kipp, Kipp, 1971; 1971; Loubere, Loubere, 1981; 1981; Molfino Molfino et et al., al., 1982; 1982; Ravelo Ravelo et et al., al., 1990; 1990; Dowsett, Transfer functions Dowsett, 1991]. 1991]. Transfer functions derived derived in in this this manner manner usually (SST) usuallyassume assumesea seasurface surfacetemperature temperature (SST) is isan anecological ecological variable or variablesignificant significantto to foraminiferal foraminiferalassemblages assemblages or is is linearlinearly variable signifily related relatedto tosome someunspecified unspecified variableofofecological ecological significance. aaspecies' growth cance.However, However,ititisisdifficult difficulttotodetermine determine species' growth environment environment (depth (depth and and season) season)from from its itssediment sedimentdistribution distribution (see Ctal. al. [1992] [1992] for for aa radiolarian radiolarianexample). example). It It is is thus (seeWelling Welling et thus difficult to determine objectively the environmental field(s) difficult to determine objectively the environmental field(s) studies include sediment traps, plankton tow, and pumps. Sediment traps traps measure integrated faunal faunal flux, flux, the Sediment measure integrated the link link between living living populations, and the between populations, and the fossil fossilforaminiferal foraminiferal record. Tow record. Tow and andpump pumpstudies studiesrelate relateforaminiferal foraminiferalstanding standing stock to stock to environmental environmental parameters. parameters. Using 200-Jim mesh nets, nets, systematic Using 200-gm mesh systematicplankton plankton tow tow surveys surveys of of the the Atlantic Atlantic and and Indian Indian Oceans Oceans were were conducted conducted from from the the Copyright Geophysical Union. Copyright1995 1995by bythe theAmerican American Geophysical Union. l950s to 1950s to the the 1970s 1970s [e.g., [e.g., Be, Bd, l959b; 1959b;Be Bd and andTolderlund, Tolderlund, 1971]. A 1971]. A primary primary goal goal of of these thesestudies studieswas wasto to determine determinethe the role of role of temperature temperatureon onforaminiferal foraminiferal species speciesdistribution. distribution. These These tows tows lack lack good good vertical vertical resolution resolution (generally (generally 00- to to 200-rn integrated integrated samples) samples) but but demonstrate that (1) 200-m demonstrate that (1) most most species of species of planktomc planktonic foraminifera foraminifera live live within within the theeuphotic euphotic zone [Be, zone [Bd, 1960] 1960] and and (2) (2) the the dominant dominantspecies speciescan can be be separated separated into zoogeographic into zoogeographicassemblages assemblages[Be [Bd and and Hamlin, Hamlin, 1967]. 1967]. Paper Papernumber number95PA02088. 95PA02088. 0883-8305/95/95PA-02088$l0.00 0883-8305/95/95PA-02088510.00 using nets nets with with meshes meshes ranging ranging from from 119 119to to 505 505 gm tm in using in size size [Bradshaw, Berger, 1969, [Bradshaw, 1959; 1959; Smith, Smith, 1963, !963, 1964; 1964; Berger, 1969, 1971; 197i; Similar data from Similar data from the Pacific Pacific and and Indian Indian oceans oceans were were obtained obtained 988 ORTIZ ET ET AL.: AL.: FORAMINIFERA OF ORTIZ FORAMINIFERA OFTUE THECALIFORNIA CALIFORNIACURRENT CURRENT spatial relationship relationship between between the the tows, tows, we we refer refer to spatial to them them by by their distance from the coast. Six single-net plankton their distance from the coast. Six single-net plankton tows tows were collected collected during during cruise using aa Puget were cruiseW8909A W8909A using PugetSound Soundnet. net. Four Four MOCNESS MOCNESS plankton plankton tows tows were were collected collectedduring duringcruise cruise Plankton W9009A. One Onesingle-net single-nettow towwas wascollected collectedduring duringNH5-19 NH5-19 Plankton tow tow studies studiesusing using complex complex samplers samplers like like the the W9009A. Deep conductivity-temperatureusing the Puget Sound net. Multiple Opening Closing Net Environmental Sensing Multiple Opening Closing Net Environmental Sensing using the Puget Sound net. Deep conductivity-temperatureSystem hydrographic depth (CTD) (CTD) stations stations (>1000 (>1000 m) m) were were sampled sampledacross acrossthe the System(MOCNESS) (MOCNESS) provide provideconcurrent concurrent hydrographic depth transect in in September These measurements and enhanced measurements and enhanced vertical vertical resolution. resolution. These transect September 1989 1989 (n (n== 14) 14) and and 1990 1990 (n (n = = 11), 11), respectively (Table (Table I). D measurements of improvements are critical critical to growth 1). CT CTD measurements of temperature temperatureand and improvements are to assessing assessing growthhabitats habitatsin in the the respectively salinity were collected during all three cruises. Studies of low-latitude field [Wiebe et al., 1976, 1985]. salinity were collected during all three cruises. Light Light transtransfield [Wiebe et al., 1976, 1985]. Studies of low-latitude mission was was measured planktonic using multiple measuredonly only during duringthe theW8909A W8909A and andW9009A W9009A planktonic foraminifera foraminifera using multiple net net plankton planktontows tows mission cruises. Advanced show that that many live below below the the sea Advancedvery veryhigh highresolution resolutionradiometry radiometry(AVHRR) (AVHRR) show manyforaminifera foraminiferalive seasurface surfaceat at the the cruises. sea surface deep maximum in of surfacetemperature temperatureimages imagesdocument documentthe the surface surfacethermal thermal deepchlorophyll chlorophyll maximum(DCM), (DCM), perhaps perhaps in search search offood food sea structure during cruise W9009A. [Fairbanks et al., 1979; Fairbanks and Wiebe, 1980; Be et al., structureduring cruiseW9009A. [Fairbankset al., 1979;Fairbanksand Wiebe,1980;Bd et al., Less 1985; Ravelo Ravelo et et al., al., 1990; Oberhänsli et The four tows were were collected collected west west of of the the shelf The four MOCNESS MOCNESS tows shelf 1985; 1990; Oberhiinsli et al., al., 1992]. 1992]. Less Although intercomparisons of the Miles, 1973]. 1973]. Although intercomparisons of the Atlantic, Atlantic, Indian, by Indian,and andPacific Pacificdata datasets setsare arecomplicated complicated bymesh meshsize sizevarivariation, the large-scale geographic patterns in each ation, the large-scalegeographicpatternsin each data data set set appear similar similar [Bradshaw, appear [Bradshaw,1959]. 1959]. work has work hasbeen beendone doneon onhigher higherlatitude latitudefaunas faunasusing usingmultiple multiple net towing net towingsystems. systems. Here Here we we analyze analyzeaa zonal zonaltransect transectof of MOCNESS plankton plankton tows tows across across the MOCNESS the California CaliforniaCurrent Currentto to break, which which occurs =95 km km offshore. break, occurs --95 offshore. Six Six or orseven sevenMOCNESS MOCNESS stocks stocks from from these these tows tows are areaccompanied accompaniedby bysupporting supporting hydrographic data including hydrographicdata includingtemperature, temperature,salinity, salinity,and and temperature, salinity, and temperature, salinity, and density density observations observationsto to sample sample mixed mixed layer, layer, seasonal seasonalthermocline, thermocline, and andsubtherrnocline subthermocline assess assessthe the controls controls on onmidlatitude midlatitude foraminifera. foraminifera. Standing Standing measures measuresof of small smalland andlarge largeplankton planktonbiomass. biomass. Because and provide Becausephytoplankton phytoplankton andzooplankton zooplankton providethe thebulk bulk of the of thenutritional nutritionalrequirements requirementsfor for adult adultheterotrophic heterotrophic planktonic foraminifera planktonic foraminifera [Hemleben [Hemleben et et al., al., 1988], 1988], we we hypothesize that changes in plankton biomass are mirrored by hypothesize thatchanges in planktonbiomass aremirrored by standing stock. changes in changes in asymbiotic asymbiotic foraminiferal foraminiferal standing stock. Likewise, it that provide Likewise, it has hasbeen beensuggested suggested thatphotosymbionts photosymbionts provide much much of of the thenutritional nutritionalneeds needsofofsymbiotic symbioticplanktonic planktonic foraminifera [Be et al., 1,1982; $pero Spero and and Parker, foraminifera [Bd et al., 198 1981,1982; Parker, 1985; 1985; Jørgensen JOrgensenet et al., al., 1985]. 1985]. This This implies impliesthat thatlight lightavailability availability should play play an of should an important importantrole rolein in the thedistribution distribution of symbiotic symbiotic Determining the planktonic foraminifera. planktonic foraminifera. Determining the spatial spatial and and temporal relationtemporalvariability variabilityof of the theforaminiferal-plankton foraminiferal-plankton relationship step shipprovides providesan animportant important steptoward towardultimately ultimatelycalibrating calibrating the the foraminiferal foraminiferal sedimentary sedimentary record record against against changes changesin in plankton [Mix, planktonbiomass biomassand andbiological biologicalproductivity productivity [Mix, 1989a, 1989a, b]. two b]. To Toachieve achievethis thisgoal, goal,we weaddress address twoquestions: questions:(1) (1) What What are are the the relative relativecontributions contributionsof of biological biologicalfactors factors(e.g., (e.g., food food and and and symbiont symbiontphotosynthesis) photosynthesis) and physical physicalfactors factors(e.g., (e.g., temperature, salinity, and and advection) on the temperature,salinity, advection)on the distribution distributionof of planktonic planktonic foraminiferal foraminiferal species speciesand andtotal totalforaminiferal foraminiferal planktonic foraminifera standing standingstock? stock? (2) (2) Do Do midlatitude midlatitude planktonic foraminifera exhibit the same affinity with the deep chlorophyll exhibit the sameaffinity with the deepchlorophyllmaximum maximum as astheir theirlow-latitude low-latitudecounterparts? counterparts? Materials and Materials and Methods Methods Experimental Design Design and and Field Experimental Field Methods Methods The The study studysites sitesare are part partof of the the'Multitracers" "Multitracers"program, program,aa study of of the 42°N. study theCalifornia CaliforniaCurrent CurrentSystem Systemat atapproximately approximately 42øN. These These locations locations are are excellent excellent test test sites sitesdue dueto tostrong stronghydrohydrographic gradients over a relatively small geographic region. graphicgradientsover a relatively small geographic region. Field work during Field work was wasconducted conducted duringcruises cruisesof of the theR/V R/V Wecoma Wecornaon on September 13-27, 17September 13-27, 1989 1989(cruise (cruiseW8909A), W8909A), and andSeptember September 17- 30, 1990 (cruise W9009A). W9009A). A A cruise of the the R/V R/V $acajawea Sacajawea 30, 1990 (cruise cruise of (cruise NH5-l9) on to (cruiseNH5-19) onJuly July23, 23,1991, 1991,provided providedthe theopportunity opportunityto samples were collected collected within within the the depth from 00- to sampleswere depth interval interval from to 1000 m at at each 1000 m each site. site. The Thedepth depthinterval intervalfor foreach eachMOCNESS MOCNESS net while towing net was was selected selected while towing by by monitoring monitoring CTD CTD regions. This paper regions.This paper discusses discussessamples samplesfrom from 00- to to 200 200 rn m only. only. Deep-dwelling planktonic planktonic foraminifera foraminifera observed observed in in the Deep-dwelling the samples >200 >200 m m are are discussed discussed by by Ortiz Ortiz [1995]. samples [1995]. The Thelocations locations of three of three of of the theMOCNESS MOCNESS tows towscorrespond correspondroughly roughly with with the the sites of sites of the the Multitracers Multitracerssediment-trap sediment-trapmoorings mooringsreferred referred to to as as "Nearshore" (120km), km), "Midway" "Midway (280 km), and and "Gyre" 'Gyre" (650 "Nearshore"(120 (280 km), (650 krn) Dymond et et al., et al., al., 1992; Ortiz and km) [e.g., [e.g., Dyrnond al., 1992; 1992; Lyle Lyle et 1992; Ortiz and Mix, Sancetta et et al., al., 1992]. Mix, 1992; 1992; $ancetta et al., al., 1992; 1992; Welling Welling et 1992]. The The Gyre (650 km Gyre mooring mooring (650 km offshore) offshore) is is farther farther west west than than the the location of the 572 km tow; however, hydrographic location of the 572 km tow; however, hydrographic conditions at at the the two conditions two sites sites are are similar. similar. The The seven seven Puget Puget Sound Sound net net tows tows were were collected collected with with aa simple towing towing system simple system consisting consistingof of aa conical, conical,63-j.tm 63-gm mesh mesh net (4:1 net (4:1 net net mesh mesh to to mouth mouth area) area) mounted mounted on on aa 70-cm 70-cm ring ring The W8909A (mouth (moutharea area0.385 0.385m2). m2). The W8909Atows towswere werecollected collected along the Multitracers transect at sites 67, 97, 129, alongthe Multitracerstransectat sites67, 97, 129, 298, 298, 649, 649, These tows and and 763 763 km km offshore offshore(Figure (Figure1). 1). These towswere werecollected collected over depth intervals intervals ranging rangingfrom from0-70 0-70mmto to 0-200 0-200m. m. The over depth The NH5-l9 tow tow (0-20 m) was was taken taken 10 km offshore offshorein in 50 50 m m of of NH5-19 (0-20 m) 10 km water. Sample processing for these seven single net tows was water. Sampleprocessing for thesesevensinglenet towswas identical to to the identical the MOCNESS MOCNESS tows, tows, with with the the exception exception that that standing stock stock and volume standing andplankton planktondisplacement displacement volume(see (seebelow) below) was not not estimated estimated from was from these thesesamples samplesdue dueto to the thevariable variabledepth depth intervals. intervals. The foraminiferal results from from these The foraminiferal results these tows tows are are presented as as percent percent abundance data. presented abundance data. To To calculate calculate aa comparable comparablepercent percent abundance abundancefor for the the MOCNESS data, we MOCNESS data, we integrated integratedthe the>125-j.tm >125-gm standing standingstocks stocks for for each each species speciesover over the the 00- to to200-rn 200-m depth depthinterval interval and and normalized by the normalized by the total total foraminiferal foraminiferalstanding standingstock stockat at each each site over over the the same interval. This all of of site same interval. This depth depthinterval interval includes includesall the the living living individuals individuals of of the thespecies speciesdiscussed discussedhere herebased basedon on of the the the September September1990 1990 MOCNESS MOCNESS data. data. Calculations Calculations of the September percent abundance for the September1990 1990 percent abundancefor the intervals intervals0-70, 0-70, 00100, and and 0-200 0-200 m m are are essentially essentially the the same. same. Comparison 100, Comparisonof of the the single-net tows to single-net tows to the theMOCNESS MOCNESS tows tows is is appropriate appropriate collect aa plankton collect planktontow towsample sampleduring duringan anactive activeupwelling upwelling assuming both sets assumingboth setsof of tows towssampled sampledthe thewater watercolumn columndown downto to event at at 44°N. event 44øN. Data Datafrom fromaatotal totalof of11 11plankton planktontows towsusing using the the depth depth of of the thedeepest deepestspecies speciesmaxima. maxima.This This depth depthwas was 63-lim mesh nets (Figure the 63-•m mesh netsare arepresented presented (Figure1). 1). To Toemphasize emphasize the approximately approximately70 70 m m based basedon onthe theMOCNESS MOCNESS data. data. ORTIZ Ef AL: ORTIZ ET AL.:FORAMINIFERA FORAMINIFERAOF OFTHE THE CAUFORNIA CALIFORNIACURRENT CURRENT 45 45 989 Multitracers Plankton Tow Tow Stations Stations Multitracers Plankton 44 44 3400 o• o o 43 43 Cape Cape Blanco Blanco 220 £42 42 0 41 41 0 Cape Mendocino 40 39 39 4600 38 134 134- 133 133 132 132 131 131 130 130 129 129 128 128 127 127 126 126 125 125 124 124 123 123 122 122 Longitude Longitude(°W) (øW) Figure Map of Figure 1. 1. Map of the theMultitracers Multitracersstudy studyregion. region. Open Opensquares squaresmark markthe theMultiple MultipleOpening OpeningClosing ClosingNet Net Environmental Sensing tows, while while solid solid circles tows. Environmental SensingSystem System(MOCNESS) (MOCNESS)tows, circlesmark markthe thesingle-net single-net tows. Sites Sitesare are labeled by offshore in Contours atat 200-rn intervals are bathymetry atat 15-mm labeled by distance distance offshore inkilometers. kilometers. Contours 200-m intervals are bathymetry 15-minresolution. resolution. use useof of hot hotbleach, bleach,peroxide peroxide[Sautter [Sautterand andThunell, Thunell,1991], 1991],or or Methods low-temperature ashing ashing [Oberhänsli et al., 1992]. low-temperature [Oberhansli et al., 1992]. Methods The MOCNESS samples were were preserved preserved at at sea The MOCNESS samples sea in in 10% 10% other than other thandensity densityseparation separationpreclude precludethe the possibility possibilityof of solutions of formalin solutionsof formalin (buffered (buffered to to pH pH =7.5 =7.5 using usingNaBO4) NaBO4) and and counting protoplasm-full shells, may damage the more fragile counting protoplasm-full shells, may damage the more fragile The samples placed in on placed in cold coldstorage storagefor forprocessing processing onland. land. The samples foraminifera, foraminifera, and and can can alter if altershell shellisotopic isotopiccomposition composition if were were later later split split to to within within±5% +_5%by by volume volumeusing usingaafour-way, four-way, partial dissolution occurs [Ganssen, 1981]. partial dissolution occurs [Ganssen, 1981]. Quarter splits OSU OSU sediment-trap sediment-trapsplitter splitter [Fischer, [Fischer,1984]. 1984]. Quarter splits Settling in solution of (310 Settling inaasaturated saturated solution ofNaC1 NaC1 (310gIL, g/L,specific specific from from each each sample sample were were used usedto to determine determinethe the number numberof of gravity 1.20) separates the dense shelly plankters from gravity1.20)separates thedense shellyplankters fromless less planktonic foraminifera (125-150 and >150 tm) and the wet planktonicforaminifera(125-150 and >150 gm) and the wet dense, nonshelly plankters. The procedure entails pouring dense,nonshelly plankters.The procedure entailspouring volume plankton catch (>63 JLm) collected by each MOCNESS volumeplanktoncatch(>63 gm) collectedby eachMOCNESS aliquots of sample (=50 mL at a time) into a clear 1-L beaker of aliquots ofsample (=50mLata time)intoa clear1-Lbeaker of net. net. The The wet wet volume volume plankton plankton catch catch was was measured measuredin in aa Small sample aliquots help prevent MOCNESS Sample MOCNESS Sample Processing Processing saturated saturatedNaC1 NaC1solution. solution. Small samplealiquotshelp prevent graduated cylinder graduated cylinderafter afterallowing allowingthe theplankton planktonto to settle. settle. the foraminifera in the the foraminifera from from becoming becoming entangled entangled in the floating floating We distinguished visually between "living" (protoplasmWe distinguishedvisually between "living" (protoplasm- plankton. After the foraminifera settle, the floating material plankton. After the foraminifera settle, the floating material full) and 'dead" (protoplasm-empty) foraminiferal shells. full) and "dead" (protoplasm-empty)foraminiferal shells. is removed by suction, and the procedure is repeated until the is removedby suction,and the procedureis repeateduntil the Most samples Most sampleswere were analyzed analyzedwithout withoutprotoplasm protoplasmstaining. staining. However, Rose Rose Bengal Bengal was was added added to to several samples in in 11 g/L However, severalsamples g/L concentration prior prior to concentration to plankton planktoncatch catchdetermination determinationand and density separation, then allowed to soak density separation, then allowedto soakfor for 24 24 hours hours[Walton, [Walton, of the entire sample entire sample has has been beenprocessed. processed. Separation Separation of the planktonic foraminifera from the floating organic matter planktonicforaminiferafrom the floating organicmatterwas was not significantly not significantlysize sizedependent. dependent. Seven Seven randomly randomlyselected selected sample pairs indicate only 19 ± 14% of the samplepairs indicateonly 19 +- 14% of the125125-toto150-Rm 150-gm All foraminiferal 1952; Bernhard, 1988]. 1952; Bernhard, 1988]. All foraminiferal data data reported reportedhere here size size class class and and 10 10 ±+-6% 6%of ofthe the>l50-.Lm >150-gm size sizeclass classbecame became are individuals. are from fromprotoplasm-full protoplasm-full individuals. entangled. We inspected the residual floating material Forarninifera were separated from the less dense, nonshelly under Foraminiferawere separatedfrom the lessdense,nonshelly entangled. We inspectedthe residualfloating materialunder plankton catch by [Be, magnificationand and wet-picked wet-pickedany anyremaining remainingforaminifera. foraminifera. planktoncatch by density densityseparation separation [Bd,1959a]. 1959a].We Weprefer prefer magnification foraminifera to this foraminifera The separated separated foraminiferawere werethen thentransferred transferred to glass glasspetri petri thismethod methodof of separating separating foraminiferafrom fromplankton planktonover overthe the The ORTIZ ET AL.: FORAMTNTFERA OF THE THE CALIFORNIA CALIFORNIA CURRENT CURRENT FORAMINIFERA OF 990 990 Table Table 1. 1oW9009A W9009Aand andW8909A W8909A Deep DeepConductivityConductivityTemperature-Depth Temperature-Depth(CTD) (CTD) Stations Stations foraminiferal standing and plankton foraminiferal standingstock stock(F55) (Fss) and planktondisplacement displacement volume (Pdv). 1dv is aa measure volume (Pclv). P clvis measure of of the the>63-jim >63-gm plankton plankton biomass at at each biomass each site. site. Because Because the the MOCNESS MOCNESS flow flow meter meter failed failed Longitude, CTD Latitude, Longitude, CTD Cast CastDepth, Depth, Latitude, Station m m N N W W km Local km LocalTime, Time, PSI Offshore Offshore PST based on based on flow flow meter meterdata datafrom fromMOCNESS MOCNESS tows tows[Welling [Wellinget et al., al., 1991] collected at 1991] collected at the the same samelocations locationsin inSeptember September1987: 1987: W9009A Stations W9009A Stations 90-05 90-05 90-06 90-06 90-07 90-07 90-08 90-08 90-14 90-14 90-17 90-17 90-18 90-18 90-19 90-19 90-20 90-20 90-23 90-23 2652 2652 1502 2952 2952 2751 3400 3400 2502 2502 2951 1500 1500 1002 1002 42.375 42.375 42.082 42.082 42.080 42.080 42.188 42.188 41.592 41.592 41.666 41.666 41.751 41.751 41.832 41.832 41.831 41.831 42.084 42.084 89-04 89-04 89-14 89-14 89-23 89-23 89-32 89-32 89-39 89-39 89-49 89-49 89-56 89-56 89-58 89-58 89-59 89-59 89-60 89-60 89-65 89-65 89-74 89-74 89-75 89-75 89-76 89-76 976 976 42.160 42.160 42.006 42.006 42.086 42.086 126.375 126.375 126.001 126.001 126.997 126.997 127.615 127.615 131.982 131.982 131.233 131.233 130.002 130.002 128.964 128.964 128.200 128.200 125.365 125.365 184 184 149 231 282 282 646 583 481 394 394 331 97 00:05 00:05 22:53 22:53 04:26 04:26 09:49 09:49 13:36 07:11 13:36 21:50 21:50 03:29 03:29 22:34 22:34 67 129 129 00:25 00:25 23:49 00:30 00:30 00:09 23:58 23:57 23:50 23:50 00:19 00:19 00:35 23:51 00:37 00:37 23:48 23:55 00:05 00:05 W8909A Stations Stations W8909A 1000 2601 2700 2700 3600 3750 3450 3450 3100 3100 3300 3300 2870 3222 3222 1000 2693 3072 3072 42.091 41.331 41.151 41.349 41.349 41.420 41.420 41 .549 41.549 42.049 42.049 42.202 42.202 42.072 42.072 42.059 42.059 42.047 42.047 124.559 124.559 125.457 125.457 125.450 125.450 127.345 127.345 131.596 131.596 133.202 133.202 130.561 130.003 130.003 128.595 128.595 128.001 127.032 127.032 126.297 126.297 125.599 125.599 125.298 125.298 in September in September1990, 1990, we we estimated estimatedvolume volume filtered filtered from from tow tow duration using using aa linear (n 45, r2 duration linearregression regression (n = = 45, r2 = = 0.97, 0.97, p < 0.01) 0.01) 279 279 648 648 763 763 560 560 482 482 396 396 314 314 238 238 190 190 149 149 107 dishes, dishes,the the remaining remainingsaline salinesolution solutionwas wascarefully carefullyremoved removed by by pipette, pipette,and andthe thesample samplewas wasallowed allowedto toair airdry dryovernight. overnight. The were then then dry dry sieved The foraminifera foraminifera were sievedat at 125 125 and and150 150 .tm. [tm. We select We selectthese thesesize sizefractions fractionsto to facilitate facilitatecomparison comparisonwith with (1) (1) our trap study our 1987-1988 1987-1988 sediment sedimenttrap study[Ortiz [Ortiz and andMix, Mix, 1992], 1992], (2) (2) the >125-l.tm workfrom fromthe the Gulf Gulf of of Alaska the >125-[tm work Alaska [Sautter [Sautter and and Thunell, 1989], and (3) the >150-I.Lm sediment studies Thunell, 1989], and (3) the >150-gm sedimentstudiessuch suchas as Climate: Long-Range Climate: Long-RangeInvestigation, Investigation,Prediction, Prediction,and andMapping Mapping Foraminifera in in both (CLIMAP) (CLIMAP) [1976]. [1976]. Foraminifera both size size classes classes were were identified to to species species level level using using the the taxonomy taxonomyofofParker Parker identified [1962] and Be [1962] and Bd [1977]. [1977]. The The only only difference difference between between the the taxonomy used used here here and taxonomy and that that used usedin in our our1987-1988 1987-1988 sediment sediment trap study is with of trap study [Ortiz [Ortiz and and Mix, Mix, 1992] 1992] is with the the classification classificationof the Neogloboquadrina Neogloboquadrina pachyderma Neogloboquadrina the pachyderma -- Neogloboquadrina dutertrei dutertrei intergrade intergradecategory categoryof of Kipp Kipp [1976]. [1976]. After After careful careful analysis of the of Neogloboquadrinid analysisof the distribution distributionof Neogloboquadrinidplanktonic planktonic foraminifera in the foraminifera in the Multitracers Multitracerssediment sedimenttrap trap and andplankton plankton tows, tows, we we consider considerthe theP-D P-D intergrade intergradecategory categorytotobe beaamorphomorpho- logic subgroup subgroup of of N. N. dutertrei. logic dutertrei. We Wehave havegrouped groupedthese thesetwo two categories in this This grouping did not not affect categories in this paper. paper. This grouping did affect the the results results presented presentedhere; here; similar similar results resultswere were obtained obtainedwhen when N. N. dutertrei dutertreiwas wasanalyzed analyzedwithout withoutthe theaddition additionof ofindividuals individualsthat that some somestudies studieshave havereferred referredto to as asP-D P-D intergrade. intergrade. Foraminiferal Foraminiferal Standing Standing Stock Stock (F55) (Fss) and and Plankton Plankton Displacement Volume Displacement Volume (Pd,) (Ptlv) Calculations Calculations Using estimates of of the by the Using estimates the water water volume volume filtered filtered by the MOCNESS, we convert MOCNESS, we convert raw raw planktonic planktonic foraminiferal foraminiferal shell shell counts to counts (Ni) (Nf) and andwet wetvolume volumeplankton planktoncatch catch(Vu) (Vp)to V1= Vf= 16.2(t) 16.2(t)-- 4.7 4.7 (1) (1) (in cubicmeters). meters).InIn the the above (in cubic above relationship, relationship, tt is is the thetow tow duration in decimal duration in decimal minutes. minutes. Residual Residual errors errors associated associated with with (1) estimated (1) amount amount to to 12% <12% ofofthe the estimatedvolume volume filtered. filtered. Foraminiferal in shells Foraminiferal standing standingstock stock(F55) (Fss) in shellsper per cubic cubicmeter meter and and plankton plankton displacement displacementvolume volume (Pdv) (Pclv)in in milliliters milliliters per per were calculated and VP, cubic cubic meter meterwere calculatedby by dividing dividingNf Nfand Vp, respectively, by the respectively,by the sample samplesplit split size size (S) (S) and andvolume volumefiltered filtered (V1). The sample split size (5) was 1/4 in all cases. (Vf). Thesample splitsize(S) was1/4in all cases.Counting Counting errors errors based based on on the thespecies speciesmaximum maximum percent percentabundance abundanceat at each of each site site ranged ranged from from less less <1% <1% to to 3% 3% using using the the method method of Dryden Dryden [1931]. [1931]. Splitting, Splitting, counting, counting, and and towing towing errors errors produce aa total total measurement error of of 20% 20% for for the produce measurement error the foraminiferal foraminiferal standing stock dv are standing stock estimates estimates (F55). (Fss). Total Total errors errors in in Pdvare comparable. comparable. MOCNESS Tow Data MOCNESS Tow Data Analysis Analysis We present We presentthe the species' species'standing standingstocks stocksas asvertical verticalprofiles profiles factor analysis analysis [Kiovan across the across the transect. transect. Q-mode Q-mode factor [Klovan and and Imbrie, 1971] from 00- to Imbrie, 1971] based basedon on all all MOCNESS MOCNESS samples samples from to 200 taxa with 200 m m objectively objectivelygrouped grouped13 13 foraminiferal foraminiferaltaxa with similar similar distributions. Prior distributions. Prior to to the thefactor factoranalysis analysisaapercent-maximum percent-maximum transformation normalized normalized the the standing standing stock stock of transformation of each eachspecies species in each in each sample sampleby by its its maximum maximum standing standingstock stock in in the the entire entire the spatial structure of of the the data data set. data set. This This preserves preservesthe spatial structure data set set during the the factor but ranks ranks all all species equally. A during factor analysis analysis but speciesequally. A varimax rotation rotation optimized optimized the the factor varimax factor solution solution to to give give assemblages with with mostly assemblages mostly positive positive factor factorscores. scores. In the of In the absence absenceof of predation predationand andothers otherssources sources ofmortality, mortality, aa species optimum environment is identified by speciesoptimum environmentis identified by the the location location of its of its maximum maximum growth growth rate. rate. Our Our poisoned poisonedplankton plankton tow tow samples preclude the samples preclude the direct direct measurement measurementof of growth growthrates. rates. Therefore to to test Therefore test the the relationship relationshipbetween between a a species' species'standing standing stock and stock and environmental environmentalforcing forcing(e.g., (e.g.,light lightand andfood foodrequirerequirements, temperature temperature and and salinity etc.), we ments, salinity tolerances, tolerances,etc.), we assume assume that (1) that (1) the the preferred preferredhabitat habitatof of each eachspecies speciesat at each eachsite siteoccurs occurs at the and (2) at the depth depth of of its itsmaximum maximumconcentration concentration and (2) the the magnitude of of each magnitude eachspecies' species'maximum maximumreflects reflectsenvironmental environmental In conditions at at that conditions that site siteaveraged averagedover over the thetowing towinginterval. interval. In cases where where aa species cases speciesis is equally equallyabundant abundantwithin within error error bars barsin in two samples from the same site (Globigerinita glutinata two samplesfrom the samesite (Globigerinitaglutinataat at 220 220 km and and Globigerina Globigerina quinqueloba km quinqueloba121 121 km km offshore), offshore),we we plot plot both samples. both samples. By By focusing focusingon onthe thespecies' species'maxima maxima at at each each site we ratio and site we improve improve the the signal-to-noise signal-to-noiseratio and decrease decreasethe the effects of extraneous factors which might otherwise effects of extraneous factors which might otherwisemask mask relationships between relationships betweenspecies speciesdistribution distributionand andenvironmental environmental factors. The relationships isisassessed factors. Thestrength strengthof ofthe theobserved observed relationships assessed using the correlation coefficient using thesquared squared correlation coefficient(r2) (r2) which whichdetermines determines the fraction the fraction of of variance varianceexplained explained [Snedecor [Snedecor and and Cochran, Cochran, is assessed 1989]. Statistical 1989]. Statisticalsignificance significanceis assessedby by the theassociated associated "p-value" significant "p-value"which which is is reported reportedonly onlyfor forrelationships relationships significant levels. at the the<5% 5% or at or _<1% levels. ORTIZ ORTIZ FT ET AL.: FORAMINWERA FORAMINIFERA OF OF THE THE CALIFORNIA CALIFORNIA CURRENT CURRENT Foraminiferal Size Size Measurements Measurements Foraminiferal We supplement the standing We supplement the standing stock stock estimates estimates with with shell shell length length measurements measurementsto to obtain obtain aa second, second,independent independentmeasure measure of foraminiferal Shell length length was of foraminiferal success. success. Shell was measured measuredusing using an an ocular reticule (±6-tim precision). precision). On ocular reticule at at lOOx 100x magnification magnification (+6-gin On short timescales, short timescales, shell shell size size variations variations should should be be correlated correlated with We selected with aa species species'growth growth rate. rate. We selectedthree threeabundant abundant species on on which one species whichto to conduct conductthis thissize sizeanalysis, analysis, onewhich which harbors obligate (0. universa), harbors obligate endosymbionts endosymbionts(O. universa),one onewhich which harbors facultative harbors facultative endosymbionts endosymbionts(N. (N. dutertrei), dutertrei),and andone one which does not harbor endosymbionts (right-coiling which does not harbor endosymbionts (right-coilingN. N. 991 991 Results Results The The Physical Physical Environment Environment The The Multitracers Multitracers transect transectranges rangesfrom fromcool coolupwellingupwellinginfluenced waters near near the the coast influencedwaters coastto to waters watersinfluenced influencedby by the the oligotrophic subtropical subtropical gyre gyre offshore The oligotrophic offshore (Figure (Figure 2). 2). The prominent cool-water filament situated off Cape Blanco prominent cool-water filament situated off Cape Blanco (sampled by the anticyclonic (sampled by the 120 120 km km tow) tow) and andits itsassociated associated anticyclonic eddy to features eddy to the the north northare arepersistent persistent featuresduring duringthe theupwelling upwelling season Strub et et al., al., season [Laurs, [Laurs, 1967; 1967; Iketia Ikeda and and Emery, Emery, 1984; 1984; Strub 1991]. These mesoscale features (Figure 2) carry the influence 1991]. These mesoscalefeatures(Figure 2) carry the influence pachyderma) [Hemleben er al., al., 1988]. here pachyderma) [Hemleben et 1988].Our Ourobjectives objectives hereare are of of coastally 50 km coastallyupwelled upwelled water water farther farther offshore offshorethan than the the --50 km (1) to test whether changes in size can be linked to the degree (1) to testwhetherchanges in sizecanbe linkedto thedegree zone zone of of active active upwelling upwelling [Huyer, [Huyer, 1983; 1983; Strub Strub et et al., al., 1991]. 1991]. of of the the host and (2) of dependence dependenceof host on on its its symbionts symbiontsand (2) to to Similar Similar AVHRR AVHRR SSTs SSTs from from September 20, and September 20, and 29, 29, 1990 1990 determine if changes in size are correlated with changes in determineif changesin size are correlatedwith changes in suggest that that the suggest the surface surface thermal thermal structure structure was was essentially essentially standing stock. presented standingstock. The The size sizemeasurements measurements presentedhere hereare are unchanged unchanged throughout throughout cruise cruiseW9009A W9009A (Figure (Figure 2, 2, inset). inset). from shells at the standing from shellscollected collectedat the depth depthof of the thespecies species' standing The W9009A and W8909A deep CTD stations provide The W9009A and W8909A deep CTD stations provide stock maxima. stock maxima. transects of temperature, salinity, density (a,), geostrophic transectsof temperature, salinity, density ((Yt), geostrophic velocity (relative (relativeto to 500 500 m), m), and and C•, C, (Figures (Figures33and and4). 4). We velocity We Hydrographic Hydrographic Data Data Analysis Analysis discuss discuss the the September September 1990 1990 hydrography hydrography first first because because it it AVHRR 20, accompanies the the higher-resolution AVHRR images imagesof of September September 20, 1990 1990(22:13:30 (22:13:30GMT), GMT), accompanies higher-resolutionMOCNESS MOCNESS plankton planktontows. tows. and September 29, 1990 (22: 14:40 GMT), from the Scripps During the upwelling season, in response to During the upwelling season,in responsetowind-induced wind-induced and September 29, 1990 (22:14:40GMT), from the Scripps Satellite Center, were Satellite Oceanography OceanographyCenter, were converted convertedto to sea seasurface surface coastal coastalupwelling upwelling and and intense intensesouthward southwardflow, flow, the thetemperature, temperature, temperature using salinity, and and density salinity, density isolines isolines outcrop outcrop near near the the coast coast(Figure (Figure 22 temperature using the thecommercially commerciallyavailable availableX-Image X-Image software program. program. The Accordingly, the the surface and 3). software The SST SST values valuesat at 42.5°N 42.5øN across acrossthe the two two and 3). Accordingly, surface mixed mixed layer layer is is shallowest shallowest images were and images were then thenextracted extractedfor forintercomparison intercomparison andcloud cloud near near the the coast coast and and increases increases in in depth depth to to the the west. west. The The masked Columbia River plume, maskedvisually visually with with aa simple simplethreshold thresholdtest test[Stewart, [Stewart,1985]. 1985]. Columbia River plume, aa lens lensof of buoyant, buoyant,low-salinity low-salinity water water The CTD km from from the the coast, situated 250-450 250-450 km coast, is is evident evident in in the the salinity salinity The CTD used usedduring duringW8909A W8909A and andW9009A W9009A was wasequipped equipped situated with Seabird pressure, temperature, and conductivity sensors. (Figures with Seabirdpressure,temperature,and conductivity sensors. and and o rrttransects transects (Figures3b 3band and3c). 3c). Conductivity was converted converted to to salinity as defined The Conductivitywas salinityas definedby by the the1978 1978 The meridional meridional component component of of geostrophic geostrophic velocity velocity Practical Salinity (relative to to 500 500 m) m) indicates indicatessouthward southwardflow flow throughout throughoutmost most Practical SalinityScale Scale(PSS-78). (PSS-78). Temperature Temperatureand andsalinity salinity (relative were of were used used to to calculate calculate (1) (1) density density as as defined defined by by the the1980 1980 of the the transect, transect,with with strongest strongestsouthward southwardflow flow near near the the coast coast International Equation Equation of of State and weaker in the International State(IES-80) (IES-80) and and (2) (2) geostrophic geostrophic and weaker southward southwardtransport transportin the Columbia Columbia River River plume plume velocity relative to to an an assumed level of of no no motion at 500 (Figure 3d). The only velocity relative assumed level motion at 500 m m (Figure 3d). The only significant significant northward northward flow flow occurs occurs [Pond [Pond and and Pickard, Pickard,1983]. 1983]. The CTD The CTD was wasalso alsoequipped equippedwith with the thesame sameSea-Tech Sea-Techtranstransmissometer during during both both cruises. cruises. This the missometer This instrument instrumentmeasures measuresthe attenuation of monochromatic lightat at 670 670 nm nm along along aa path attenuationof monochromaticlight path between between 160 160 and and 220 220 km km offshore offshore in in an an eddy eddy between between the the coastal coastal upwelling upwelling front front and and the theColumbia Columbia River River plume. plume. data attenuation coefficient. C,, values values are dataas asCi,, Ct,,the theparticle particle attenuation coefficient.Ct, are influenced most heavily by <20-jim influenced most heavily by <20-gm microplankton microplanktonbiomass biomass [Pak et thus provides provides aa [Paket etal., al.,1988; 1988;Gardner Gardner etal., al.,1993]. 1993]. Ci,, C•,thus biomass measure biomass measureweighted weightedtoward toward phytoplankters phytoplanktersand andsmall small heterotrophs, which dv estimates estimates of of larger heterotrophs, which complements complementsthe the Pdv larger (>63 (>63 jim) gm) plankton planktonvolume. volume. The The attenuation attenuationof of ambient ambientlight light in in water wateris is proportional proportionalto to waters Particle waters near near the the coast coast and and subsurface subsurface waters waters offshore. offshore. Particle Similar flow flow patterns depths Similar patternsare are obtained obtainedfor for assumed assumedreference referencedepths of 750, 750, 800, and 1000 m. This to be be of 800, and 1000 m. Thisnorthward northwardflow flow appears appearsto length of of 25 25 cm cm [Spinrad, [Spinrad, 1986; Spinrad et er al., al., 1989]. the extension of the the length 1986; Spinrad 1989]. This This the extensionof the Cape Cape Blanco Blanco filament filament which which traverses traversesthe light 220 km Velocities 450-550 light attenuation attenuationis is linearly linearlyproportional proportionalto to the theparticle particleconcon- 220 km site site (Figure (Figure 2). 2). Velocities 450-550 km km offshore offshore centration volume in in the indicate slack water or weak northward transports. centration volume the water water [Zaneveld [Zaneveld er et al., al., 1979]. 1979]. indicate slack water or weak northwardtransports. Following Highest Following standard standardpractice, practice,we we present presentthe thetransmissometer transmissometer Highest particle particle attenuation attenuation coefficients coefficients occur occur in in surface surface the vertically of particles particles in in the the vertically integrated integrated concentration concentration of the water. To emphasize the spatial pattern of turbidity water. To emphasize the spatial pattern of turbidity that that attenuates ambient ambient light light with attenuates with depth, depth, we we calculate calculate the the relative relative particle load particle load along along the the transect transectin in September September1989 1989 and and 1990. 1990. This the C, Thiswas wasdone doneby bysumming summing the Cvdata datafrom from200 200m mto tothe the surface at each CTD site and dividing these values by surface at each CTD site and dividing these values by the the maximum value obtained obtained in in September September 1990. 1990. This maximum integrated integrated value This allows allows for for direct direct comparison comparison of of the theintegrated integratedparticle particle standing stocks stocks from from the standing the 2 2 years yearsby by referencing referencingchanges changesto to the the maximum value in September 1990. Higher surface values maximum value in September 1990. Higher surface values imply imply greater greatervertically vertically integrated integratedstanding standingstocks stocksof of particles particles and greater and greater light light attenuation. attenuation. attenuation rapidly decreases attenuation rapidly decreasesoffshore offshore and and with with increasing increasing depth depth (Figure (Figure 3e). 3e). This This pattern pattern indicates indicatesturbid, turbid, particle-rich particle-rich water water near near the the coast coastand andless lessturbid, turbid,particle-poor particle-poorwater waterfurther further offshore. offshore. Inshore Inshoreof of250 250km, km,the theparticle particlemaximum maximumis islargely largely a surface extending down downto to 50 50 m. m. Offshore a surface feature, feature, extending Offshore of of 250 250 km, the kin, the particle particle maximum maximum becomes becomesaa subsurface subsurfacefeature 'feature between 20 20 and and 60 between 60 m. m. The vertically closest to to The vertically integrated integratedparticle particle load load (Figure (Figure 31) 3f) closest the coast (--100 (l00 km) the coast km)isisroughly roughlytwice twiceas ashigh highas asthat thatfarthest farthest that the offshore (600 km). offshore (=600 km). Assuming Assuming that the light light scattering scattering properties of the propertiesof the particles particlesalong along the the transect transectare are similar, similar, the the high load near near the the coast result in in aa factor of 22 high particle particle load coastshould shouldresult factor of greater attenuationofof light light near greater attenuation near the the coast coast than than farther farther offshore. offshore. Measurements Measurements of of the the diffuse diffuse attenuation attenuation coefficient coefficient (Kd) active (Kd) of of ambient ambientphotosynthetically photosynthetically activeradiation radiation(PAR) (PAR) at at these same these same sites sites in in September September1991 1991 under undersimilar similar particle particle loads demonstrate demonstrate that that the the 1% PAR level is achieved at 35 loads 1% PAR level is achieved at 35 m m near the the coast coast (--100 ('l00 km) near kin) and andtwice twice as as deep deep at at 70 70 m m farther farther OF THE THE CALIFORNIA ORTIZ FT AL.: AL.: FORAMINIFERA ORTIZ ET FORAMINIFERA OF CALIFORNIA CURRENT CURRENT 992 992 44 44 - 43 43 'Cape .... Blanco 42 - 42 41ß''Cape : Mendocino ., 40 40 - 20 Seplember 20 September 29 September September : 20 18 16 ß .. ß 14 39 39 - 572 12 '° 8 700 700 .... I .... I .... 600 0O 500 I .... 400 I .... 300 300 I .... 200 200 I .... 100 100 I) 0 Distance Distance from Coast Coast (krn) (km) 38 m 134 I 133 132 I 131 I 130 1 I 128 127 I 126 125 PA 123 Longitude Longitude(°W) (øW) Figure 2. very high high resolution (AVHRR)image imageof of the the Multitracers Multitracers study study area area on Figure 2. Advanced Advanced very resolutionradiometer radiometer (AVHRR) on September 20, 1990 (22:13:30 GMT). GMT).Lighter Lightershades shadesnear nearthe thecoast coastindicate indicatecold coldupwelled upwelledwaters waters(--10øC), (l0°C), September 20, 1990 (22:13:30 while heavier shades shades indicate indicate warmer warmeroffshore offshorewaters waters(<18øC). (18°C). Large while heavier Largecrosses crossesdenote denotethe thelocations locationsof of the the four four MOCNESS plankton plankton tows: tows: 121,220, 121, 220, 289 289 and and 572 MOCNESS 572 km km offshore. offshore.The The small small crosses crossesdenote denotethe the locations locationsof of the the deep conductivity-temperature-depth CTD stations. stations. The The insert insert compares compares AVHRR AVHRR temperatures temperatures from from this this image image deep conductivity-temperature-depth CTD to temperatures to temperaturesfrom from September September29, 29, 1990. 1990. offshore (60O km) Oregon State values September offshore(=600 km)(C. (C.Roesler, Roesler, Oregon StateUniversity, University, Offshore Offshore valuesof of C,, Cpwere weretwice twiceas asgreat greatduring during September personal communication, 1989 personal communication,1994). 1994). 1989 (Figure (Figure 4e) 4e) than thanSeptember September1990 1990 (Figure (Figure 3e). 3e). The The Hydrographic conditions during during September Hydrographic conditions September1989 1989 were were integrated integratedoffshore offshorerelative relative particle particleload loadin inSeptember September1989 1989 qualitatively similar similar to to those was 30-40% 1990 qualitatively thoseof of September September1990. 1990. Offshore Offshore (Figure (Figure41) 4f) was 30-40% higher higherthan thanin in September September 1990(Figure (Figure mixed layer temperatures during September 1989 (Figure 4a) 31). In September 1989, the 80% particle load contour extends mixedlayertemperatures duringSeptember 1989(Figure4a) 3f). In September1989, the 80% particle load contourextends were '2°C cooler than September 1990 (Figure 3a). 3a). The In September were--2øC cooler thanduring during September 1990(Figure The across across the the entire entire transect. transect. In September 1990, 1990, the the 80% 80% Columbia River plume plume (=32.3 (32.3 practical salinity units contour extends extends less than than 250 250 km km offshore. offshore. ' Columbia River practical salinity units(psu)) (psu)) contour was Environmental data collected collected at at the the time time of tow Environmental data of the NH5-19 NH5-19 tow was closer closer to to the thecoast coastin inSeptember September1989 1989and andwas wasmuch much narrower than than in in September July 1991 indicate extremely high biomass and active narrower September1990. As Asaaresult, result,offshore offshorewaters waters in in July 1991 indicate extremely high biomass and active were saltier 1989 were saltierin in September September 1989(Figure (Figure4b) 4b) than thanin in September September 1990 (Figure in and 1990 (Figure3b). 3b). The The changes changes in temperature temperature andsalinity salinity resulted in in stronger density near the the coast resulted stronger densitygradients gradients near coast(Figure (Figure4c) 4c) and more flow 1989 and moreintense intensegeostrophic geostrophic flow during duringSeptember September 1989 (Figure 4d). regions (Figure 4d). The The locations locationsof of the thehigh-velocity high-velocity regionsdid did not change not changeappreciably appreciablyfrom from their theirSeptember September1990 1990locations. locations. upwelling to 20-m upwelling [Dickson, [Dickson, 1994]. 1994]. Temperature Temperatureover overthe the00-to 20-m towing interval ranged ranged from from 88 to towing interval to 10°C 10øCand andsalinity salinityranged ranged from to 34.2 from 33.8 33.8 to 34.2 psu. psu. These Thesevalues valuesare arecolder colderand andsaltier saltierthan than surface surface conditions conditions along along the the Multitracers Multitracers transect transectand and thus thus provide aa reasonable provide reasonable estimate estimate of of "end "endmember" member" seasonal seasonal upwelling conditions upwelling conditions in in the the northern northernCalifornia California Current. Current. 993 993 ORTIZ ORTIZ ET AL.: AL.: FORAMINIFERA FORAMINIFERA OF OF THE THE CALIFORNIA CALIFORNIA CURRENT CURRENT 00-• t• '• 2525-• =tg:t6-•t4•_• t2/" 100 150 , • , • , • , • 150 15o-' , 200 200 • 100 *BB ** ** B B * 2320 23•.20 --23.95 B T * *T B 23. 23.20 23.9 23.95 '"' / •- -"•'24.70 • 32.6 32.6%• 33.2 [33.2 f'•f ' I ' I ' I ' I ' I ' I 400 300 200 500 500 400 300 200 Distance from Distance from coast coast(km) (km) 600 600 100 0 * B B * ** * ' ' ' (1 -6 i 1 // B *T* *T* B I? / I \ / -3I -3 1100- tOO 125= . 125 150 150: 150 150= . 175 = 200 200- , 175 175: • , • , I ' I ' I ' 200 400 300 400 300 200 500 500 Distance from Distance from coast coast(km) 600 600 I , 0.16 0.16 . 0.08--i ! 7575 l 0.40 t, 0.40 e24 0.24 -- ID 200 200 0.32 0.32 _0.08..•,---- ,' I I I 200 500 400 300 500 400 200 Distance Distancefrom from coast coast(kin) (km) 600 100 tOO -September 1990: Attenuation Coefficient, September 1990:Particle Particle Attenuation Coefficient,42°N 42øN * ** * * *T** B *T B B 2550- • •-• 75 .l3\li3 - 100. tOO 125 -•-•--25.45"• 125= 0- . o 255050 75- • /N 25.45 .•"'...•5.45 .g i75 *T* *T * September 1990: Geostrophic Geostrophic Velocity Velocity (rd 500 m) September 1990: (re1500 m) September 1990: September 1990:Sigma-T, Sigma-T,42°N 42øN * B B 32.3 • 32.9- 8• 200 400 300 500 500 400 300 200 Distance Dist•ce from from coast coast(kin) (km) 600 600 2550- 25- • 25-' 505075-' 75l00tOO- . • ** * 3•.3 175175: , * ** B B 125= 125- 125 125: 150: 175 175: 00 * 0l 16 50- --•--•t2•'••.•'•: 504 7575 200 200 September 1990: September 1990:Salinity, Salinity,42°N 42øN September 1990: Temperature, 42°N September 1990:Temperature, 42øN * *T** B B * ** ** B B B *T ** 100 tOO September 1990: Relative Particle Particle Load, Load, 42øN 42°N September 1990:Relative 0- * 25 50 .g 75 B B * *BB BB * * *T * /2O-20 - tOO tOO 100. 125 125: 125150t50• 175- 1251 ß 150 150: 175 175: 200 200 I 600 6OO '' II ' I I ' I I ' I ' I 100 500 400 300 200 tOO 200 400 300 500 Distance from Distance fromcoast coast(kin) (km) 200 200• , I ', • I , ' ' ' 500 400 600 600 500 41•0300 360 200 260 100 t•)0 Distance from Distance fromcoast coast(kin) (km) properties (0-200 (0-200 m) m) across across the the Multitracers transect in in September 1990, based on Figure 3. Figure 3. Hydrographic Hydrographic properties Multitracers transect September 1990,based on across the the top top of of each nine located 97-646 ninedeep deepCTD CTDstations stations located 97-646km kmoffshore. offshore.Symbols Symbols across eachpanel paneldenote denoteCTD CTD locations (asterisks), plankton tow (T), where CTD tows locations (asterisks), plankton towlocations locations (T), or orlocations locations where CTDand andplankton plankton towswere werecoincident coincident data contoured contoured at at 2°C; data at 0.3 salinity unit (psu) (B). (B). (a) (a) Temperature Temperature data 2øC;(b) (b)salinity salinity datacontoured contoured at 0.3practical practical salinity unit (psu)interintercontoured at at 0.25 (d) component of geostrophic velocity vals; (cy vals;(c) (c)density density (ot)contoured 0.25kg kgm-3 m-3intervals; intervals; (d) meridional meridional component of geostrophic velocity particle attenuation coefficient relative to 500 in 33 cm relativeto 500m m contoured contoured cmss-1intervals; intervals;(e) (e) transmissometer transmissometer particle attenuation coefficient(Cr) (Cp) contoured intervals; (f) relative contouredin in 0.08 0.08 m-1 m-1 intervals; relativeparticle particleload loadcontoured contouredin in 20% 20% intervals. intervals. dv and and Ci,, Comparisons Comparisonsof of Pdv Cp various life life stages of dv is is dominated dominated by In these In thesesamples, samples,Pdv by various stagesof C1, is small calanoid copepods, euphausids, and chaetognaths. smallcalanoid copepods, euphausids, andchaetognaths. Cpis predominantly influenced influenced by by smaller predominantly smaller phytoplankton phytoplankton and and For each microheterotrophs (<20 microheterotrophs (<20 jim). gm). For eachMOCNESS MOCNESS plankton plankton tow sample, Table 22 lists tow sample, Table lists estimates estimates of of the the water water volume volume filtered, filtered, environmental environmentalvariables variables averaged averagedat at tow tow resolution, resolution, and stocks. The and total total foraminiferal foraminiferal standing standingstocks. Thetow towresolution resolutionat at the the Columbia ColumbiaRiver River plume plumesite site289 289 km km offshore offshoreis iscoarser coarser(0(0to to 60-m) 60-m) than than at at the theother otherthree threesites sites(0-30 (0-30 m mand and30-70 30-70 m). m). We We assess assessthe the potential potentialbias biasassociated associatedwith with this thissampling sampling problem at at the the averaging the I-rn C, problem averaging the 1-mresolution resolution Cpdata datainto into00- to to 60-m and and then then 00- to to 30-rn and 3030- to to 70-m 70-m bins. bins. The 60-m 30-m and The 0- to to 6060m at site m C, Cpaverage average atthe the289-km 289-km site(0.186 (0.186m-l) m-1)isis7% 7%smaller smaller than the the 0than 0- to to 30-rn 30-m average average(0.200 (0.200 m-l) m-1) and and38% 38% larger largerthan than the 30of (Table 2). 2). Because C,, the 30-to to70-rn 70-maverage average of0.054 0.054rn-1 m-1 (Table Because Cpis is all sites, dv and and F55 at all highly correlated with highly correlated with Pdv Fssat sites,we we infer inferthat thatthe the 0to 60-rn plankton tow at the 289-km site underestimated 00- to 60-m planktontow at the 289-kmsiteunderestimated 0to 30-rn "dv and and Fss F5 by the to 30-mPdv by10% 10%at atmost mostand andoverestimated overestimated the3030to values by by 40% 40% at at most. bias to 70-rn 70-m values most.The Thepotential potential biasof of+10% +10% from 0- to 30-rn from 030-m and -40% -40% from from 3030- to 70 m m is is similar similar to to the the dv and andFss. F5. Such measurernent error of of 20% for measurementerror for Pdv Sucherrors errorswould would not greatly values of of Cp, "dv' and and Fss F5 we not greatlyaffect affectthe themaximum maximum values Cp,Pdv, we ORTIZ ET EF AL.: AL.: FORAMINIFERA OF CURRENT ORTIZ FORAMINIFERA OFTHE THECALIFORNIA CALIFORNIA CURRENT 994 994 0 - 16'i6//I 16 16••••/•14•f•/ 50- ...•12• 75- •10'"' 75 100. lOO _.• '12.2// ' ._• ' I .... 200 100 600 600 500 500 400 460 300 360 260 160 Distance from coast Distance coast(km) September 1989: September 1989:Signia-T, Sigma-T,42°N 42øN ** ** ** ** ** B* B * ** 4.2 25-' *BTB *BIB 23../ 50 50-' ! 75 75: •32'6 ••'••2i•/?3•-'•'•••33. 5 150 15o: 26.0 • 175 175. 200 200 33.4 33.0 33.4 200 500 300 600 600 500 400 460 360 260 September 1989: Geostrophic Velocity 500m) September 1989:Geostrophic Velocity(rel (re1500m) ** ** B* ** *BTB B ** B * * * ** *BIB B -20 *********BTB B * * * * •02 0.2 2525- i 02 B * io--' I 2525 50so: I OT '_'10__ /I • / 1 200 2OO * * LLQ&, *BTB • 0.3 0.3 0.2_ 1 5050-' 7575-' too 100• 125-• 125 150• 150 '75• 175 [•'-t,5 Il0o /?l, -5 / 0 / .-5 x___/_i 175175 i D 200 100 600 500 400 300 600 500 400 360 260 Distance Distancefrom from coast coast(km) (km) 100 too Distance from Distance fromcoast coast(1cm) (km) ( 500 400 600 500 460 600 300 200 3OO 2;0 Distance from from coast Distance coast(km) 100 160 B ********* BTB September t989: September 1989:Relative RelativeParticle ParticleLoad, Load,42°N 42øN September 1989: Coefficient, 42°N September 1989:Particle ParticleAttenuation Attenuation Coefficient, 42øN B 0m * * * * B * * * *BIB '90 2525-' 5050- ---_•-•'50 ou 50..--• .g 75-' --•._•o•••--•4o • 3Ø .l00: 100• -----_.•30. -'"•30• 125'-20 125 • •""•20--'""20• "---..••80. '90 ••0._•_.0• 1501502 . 200 200 32.8 ,l00•100 1252 125: l50 1502 125 125- E 32'6•:•••33'6 32.••/•32.8 25k. 50- .32.650-' 752 is- .g 75 100 tOO- 00- B TB *BIB 175• I B 200 ** 32.4 p32.6 175 A 200 2oo 32.6 125: 150-' 150: 15015o 175 175 B* ** ** B * ** ** I328 •.100lOO-' 32.8.-. 125-' /8 l25 125 ! ** B 00- 16 25 - 0- September Salinity, 42øN 42°N September1989: 1989:Salinity, September 1989: Temperature, 42°N September 1989:Temperature, 42øN ** ** ** ** BB * ** ** *BTB *BIB B B ' I 600 600 ' I ' I ' I ' I 200 500 400 300 500 400 300 200 Distance Distancefrom from coast coast(km) (km) ' I 100 100 1751752 200 200 • 600 600 500 500 400 400 300 300 200 200 100 100 Distance from Distance fromcoast coast(1cm) (km) Hydrographic properties properties (0-200 (0-200 m) m) across across the the Multitracers Multitracers transect transect in in September September 1989, 1989, based based on on Figure 4. Figure 4. Hydrographic Plotting conventions as in Figure 3. 12 deep CTh stations located 107-648 km offshore. 12 deepCTD stationslocated107-648km offshore. Plottingconventions as in Figure3. Because our analyses report for report for the the 289-km 289-km site. site. Because our subsequent subsequent analyses are based are basedon on comparisons comparisonsof of maximal maximal values, values,we we conclude concludethe the or data, we the 15dv or C,, concentration orC,, Cpdata, weplotted plotted themaximum maximum Pdvor Cpconcentration from each site distance from from the the coast. coast. The from each site versus versusdistance Theexponential exponential fits that fits in in Figures Figures5a 5a and andSb 5bdemonstrate demonstrate thatdata datafrom fromthe thetow tow hindrance. hindrance. 220 220 km km offshore offshore do do not not fit fit the thenighttime nighttimetrend trenddescribed describedby by This tow the the other other three three tow tow site. site. This tow was was collected collected before before the the loss at site loss of of resolution resolution at the the 289-km 289-km site is is not not aa serious serious During cruise cruise W9009A, three of tows were During W9009A, three of the the MOCNESS MOCNESS tows were collected collected more more than than 1 1 hour hour after after sunset sunset at at sites sites 121, 121, 289, 289, and and 572 km 572 km offshore. offshore. The The fourth fourth MOCNESS MOCNESS tow, tow, 220 220 km km offoffafter sunset. This tow shore, shore,was wascollected collected13 13mm min after sunset. This tow may may still still show the show the influence influence of ofdaytime daytimephytoplankton phytoplankton growth. growth. Zooplankton Zooplanktonand and phytoplankton phytoplanktonoften oftenundergo undergodiel dielvariations variations in et in abundance abundance[Longhurst, [Longhurst, 1976; 1976; Dam Darn et et al., al., 1993; 1993; Cullen Cullen et al., al., 1992; 1992; Gardner Gardner et et al., al., 1993]. 1993]. Unless Unless these these diel diel variations variations in zooplankton in zooplankton depth depth distributions distributions and andnet netphytoplankton phytoplankton growth for, day day and and night night Pdv Pj and C,, growthare areaccounted accounted for, and Cpdata datacannot cannot be be compared compareddirectly. directly. To in either the Pdv dv To determine determineif if die! diel variations variationswere were evident evidentin eitherthe majority majority of of zooplankters zooplankters had had completed completed their their nighttime nighttime migration and prior prior to of the migration and to significant significant nighttime nighttime grazing grazing of the daily phytoplankton phytoplankton growth. growth. We daily We thus thusrefer refer to to the the220 220 km km offoffshore tow as the "daytime" tow. Based on the exponential shore tow as the "daytime" tow. Based on the exponential fits, dv at at 220 220 km km is fits, the the expected expectednighttime nighttime Pdv is 1.8 1.8 times times the the measured daytime daytime value, value, while nighttime C,, is measured whilethe theexpected expected nighttime Cpis 0.7 times the measured daytime value. We apply these scaling 0.7 times the measureddaytime value. We apply thesescaling factors to to the daytime P(J,, factors the00- to to30 30and and3030-toto70-rn 70-m daytime Pdvand andC, Cpdata data at 220 at 220 km km for for use usein in comparison comparisonto to date datefrom from the the other othersites. sites. data below below 70 70 m different A°dv Pdvand andC,, Cpdata mwere werenot notsignificantly significantly different between day day and and night between night tows tows so so no no corrections correctionswere were made. made. ORTIZ ET AL.: AL.: FORAMINIFERA OF ORTIZET FORAMINIFERA OFTHE THECALIFORNIA CALIFORNIACURRENT CURRENT 995 995 over are allindicate indicatehighest highest particle particle concentrations "dv dv' and Pdvand andC, Cpaveraged averaged overthe thetow towintervals intervals aresignificantly significantly Fss,Pdv, andCA! Cpall concentrations correlated both at at night 0.97, n n= in the seasonal 11, p in the cool, cool,subsurface subsurface seasonalthermocline thermocline(30-70 (30-70 m) m) at at the the correlatedboth night (21 (21 mL mL rn2, m-e,r2 r2 == 0.97, = 11, 0.01) and 0.0l) site _<0.01) andininthe theday day(9(9rnL mLrn-2, m-e,r2 r2 == 0.98, 0.98, nn == 4, 4, pp _<0.01) site 572 572 km km offshore offshoreand andin in the theproductive productivesurface surfacewater water(<30 (<30 (Figure 5c). The similarities between C, (predominately small m) at at the the other othersites sites(Figure (Figure6). 6). With With the the exception exceptionof of the the (Figure 5c). Thesimilarities between Cp(predominately small m) phytoplankton) and (predominately Columbia River plume site (289 km offshore), standing stocks large microdv phytoplankton) and P dr (predominately large micro- ColumbiaRiver plumesite (289 km offshore),standingstocks zooplankton) imply aa tight between trophic of zooplankton) imply tightcoupling coupling between trophiclevels levelsat at of 125125- to to150-jim 150-gmforaminifera foraminiferadecrease, decrease,while while>150-jim >150-gm these sites. correlations between C,, and large increase monotonically monotonically offshore. offshore. The foraminifera increase The Columbia Columbia these sites.Similar Similar correlations between Cpand largesize size foraminifera fraction River plume plume site to the fraction biomass biomass have have been been observed observed in in the the Joint Joint Global Global River site has hasvery very few few foraminifera foraminiferarelative relative to the other other Ocean OceanFlux FluxStudy Study(JGOFS) (JGOFS)North NorthAtlantic AtlanticBloom BloomExperiment Experiment [Gardner et at., 1993]. Because "dv and C,, are are so so tightly [Gardneret al., 1993]. Because PdvandCp tightly correlated at the correlatedat theMultitracers Multitracerssites, sites,we weplot plotthe theforaminiferal foraminiferal data of versus Ci,, dv only. dataagainst against Pdv only.Plots Plots offoraminifera foraminifera versus Cpwould would convey the convey the same sameinformation. information. three At each three sites, sites,particularly particularlyin in the thesmaller smallersize sizefraction. fraction. At each station, station, the the depth depth of of the theforaminiferal foraminiferalmaximum maximumcoincides coincides (within "dv (withinthe the limits limitsof of depth depthaveraging) averaging)with withthe thedepth depthof of the the Pdv and CA! maxima (Figure 6). However, the ratio of foraminifera andCpmaxima(Figure6). However,theratioof foraminifera to dvor orCp C, varies varies at at each each site site (Figure (Figure 7). 7). Relative dv' the the to P•tv Relativeto to P•tv, Total Total shells 0.97, nn = = 0.97, = 7, 7, pp _< <0.01). shellsmL-1 mL-l plankton plankton(r2 (r 2 = 0.01). In Incontrast, contrast, Planktonic Planktonic and Biomass Biomass Foraminiferal Foraminiferal 121- and and 289-km 121289-km "green" "green" water water sites sites have have 44 foraminifera foraminifera Distributions Distributions and farther offshore, offshore, the the 572-km "blue" water water site site has has 36 farther 572-km "blue" 36 Shells of of "living" are as Shells "living"individuals individuals areeasily easilyidentified identified ashaving having protoplasm with or without the aid of Rose Bengal staining. protoplasm with or withoutthe aid of RoseBengal staining. Virtually Virtually all foraminiferal foraminiferal shells shellscollected collectedin in tows towsshallower shallower than protoplasm-full. The than 200 200 m m are are protoplasm-full. The protoplasm protoplasmof of planktonic foraminifera collected collected in in the the day planktonic foraminifera daytow towis isgenerally generally green-brown. green-brown.In In the thenight nighttows, tows,ititisisyellow yellowto togolden-brown, golden-brown, presumably due to to nightly of presumablydue nightlydegradation degradation of chlorophyll chlorophyll"a" "a" in in their or prey prey [see Be and and Hamlin, theirsymbionts symbiontsor [seeBd Hamlin, 1967]. 1967]. The Thecolor color of of N. N. dutertrei, dutertrei,Globigerinoides Globigerinoidesruber, ruber,and andGloboquadrina Globoquadrina hexagona is particularly striking. Their protoplasm in hexagona is particularly striking.Theirprotoplasm inthe theday day tow from of tow is is bright brightgreen greenin in color, color,presumably presumably fromthe thepresence presence of symbionts or or the and/or symbionts theingestion ingestionof ofphytoplankters phytoplankters and/orphytophytodetritus. Dead detritus. Dead specimens specimensof of all all species speciesare areclearly clearlyidentified identified by their by theirbone-white, bone-white,protoplasm-empty protoplasm-emptyappearance. appearance. All standing stocks stocks reported reported here here are standing are based basedon on living living specimens. spocimens. foraminifera shells shells mL mL-1 almost an an order foraminifera -1 plankton, plankton, almost order of of magnitude greater greater (r2 0.84, nn == 4). magnitude (r 2 = = 0.84, 4). The The220-km 220-kmsite sitefalls falls between these these extremes between extremes with with 99 foraminifera foraminifera shells shells mL-1 mL-1 plankton (r2 = 0.99, 0.99, nn == 4, <0.01). plankton (r2 = 4, pp _< 0.01). Species Distributions Species Standing Standing Stock Stock Distributions We observe a total of 13 foraminiferal We observe a total of 13 foraminiferal taxa taxa in in the the <200-rn <200-m samples at the four MOCNESS tow stations (Table 3). We samplesat the four MOCNESS tow stations(Table 3). We limit to species with limit our ourdiscussion discussion toseven sevenfelatively relativelyabundant abundant species with clear patterns: N. N. dutertrei, dutertrei, O. 0. universa, clear distribution distributionpatterns: universa,G. G. ruber, ruber, G. N. pachyderma, G. quinqueloba, and G. glutinata, glutinata,right-coiling right-coiling N. pachyderma, quinqueloba, and Globigerina here Globigerina bulloides. bulloides. The The seven sevenspecies speciesdiscussed discussed here account for 93 93 +± 9% 95 +± 8% accountfor 9% of of the the>125-jim >125-gm and and 95 8% of of the the>150>150l.Lmforaminiferal foraminiferalstanding standingstock stockatat the the four gm four sites. sites. Total Table Plankton Tow Tow Table 2. 2. Multiple MultipleOpening OpeningClosing ClosingNet NetEnvironmental EnvironmentalSensing SensingSystem System(MOCNESS) (MOCNESS) Plankton Environmental and Foraminifera Data for Small (125-150 jim) and Large (>150 jim) Size Classes EnvironmentalandForaminiferaData for Small (125-150 gm) andLarge(>150 gm) Size Classes Tow TowSite, Site,Date, Date, and andTimea Timea 121 km 42.060°N 42.060 øN 125.665 øW °W 125.665 Sept. Sept.21, 21, 1990 1990 00:23:00 00:23:00 220km 220 km 42.167°N 42.167 øN Depth, Depth, m m 0-30 0-30 30-70 30-70 70-100 70-100 100-200 100-200 Volume, Volume, Temp. Temp, rn3 m3 C C 33 14.8 14.8 10.9 39 8.5 2.3 0.4 0.4 153 153 105 105 84 14 0.5 25 40 40 20 20 21 21 23.8 23.8 24.9 24.9 25.7 25.7 26.3 26.3 0.230 0.230 0.128 0.128 0.058 0.058 0.064 6.3 6.3 3.3 0.8 0.8 8.1 8.1 32.6 32.9 33.2 33.8 33.8 105 37 2 10 10 14.0 14.0 8.4 8.4 7.7 32.4 32.4 32.8 33.8 24.1 24.1 0.186 0.186 0.054 0.054 0.044 0.044 5.3 5.3 1.1 0.5 0.5 12 12 3 9 17.9 11.9 8.9 8.9 7.5 7.5 32.6 32.7 32.6 33.0 33.0 23.5 23.5 24.8 24.8 25.3 25.3 25.8 25.8 0.070 0.070 0.084 0.084 0.055 0.055 1.1 1.7 1.1 0.031 0.031 0.5 0.5 16.3 12.2 289km 289km 060b,c 0-60b,c 57 57 42.169 42.169 °N øN 127.694 øW °W 127.694 60-100 60-100 100-200 100-200 40 40 0-30 0-30 30-70 30-70 70-100 70-100 100-200 100-200 46 46 72 32 18 :27:00 18:27:00 99 Small Large Large Small, Small, Small Count Count Count shells shells rn-3 Count m-3 0.307 0.307 0.119 0.119 0.061 0.054 34 34 19 113 113 126.858 °w 126.858 øW Sept.28, Sept.28, 1990 1990 dv, Pdv mL rn-3 mL •-3 24.3 25.2 25.2 25.8 25.8 26.3 26.3 8.9 8.9 20 Density, Density, Cp Cp, kg rn-' kgrn-3 m-• m-1 32.8 33.0 33.3 33.8 33.8 114 147 147 0-30 0-30 30-70 30-70 70-100 70-100 100-200 100-200 Salt, Salt, psu psu 8.0 8.0 8.9 8.9 25.5 25.5 26.4 26.4 0.3 Large, Large, shells shellsrn-3 m-3 18.8 8.5 0.5 12.9 0.7 0.6 361 81 14 12.4 12.4 7.4 7.4 0.4 0.4 42.8 42.8 16.1 2.9 2.9 34 0.3 1.2 1.2 205 205 0.8 0.8 26 14 0.3 0.3 0.4 0.4 14.4 14.4 2.6 2.6 38 56 18 101 805 93 93 3.3 3.3 3.1 2.2 2.2 8.9 44.7 44.7 11.6 44 44 87 87 1.4 1.4 2.8 2.8 4.1 0.7 0.6 0.6 Sept. 23, 1990 Sept.23, 1990 19:24:00 19:24:00 572km 572 km 41.493 °N 41.493 øN 131.070°W 131.070 øW Sept. Sept.26, 26, 1990 1990 125 125 23 :30:00 23:30:00 apacific apacific Standard StandardTime Time at at start startof of the the200200- to to100-m 100-mtow tow interval. interval. bAverage temperature, salinity, density, and for 0-30 mare 16.8°C, 32.3 PSU, 23.5 kg 0.200 rn-1. bAyerage temperature, salinity, density, andCC for 0-30 mare 16.8øC, 32.3 PSU, 23.5 kgrn-3, m-3,and and 0.200 m-1. CAverage temperature, salinity, density, and •pp for30-70 mare 10.4øC, 32.5 PSU, 25.0 kgm-3,and 0.143 m-1. cAverage temperature, salinity, density, and C for 30-70 m are 10.4°C, 32.5 PSU, 25.0 kg rn-3, and 0.143 rn-'. ORTIZ ET FJ AL: CURRENT ORTIZ AL.'FORAMINIFERA FORAMINIFERAOF OFTHE THECALIFORNIA CALIFORNIACURRENT 996 10 10 ,-' E 'O I 8-1 6420 700 700 E c) I1 _.-IJ ---- --I' .- II II I I 600 600 500 500 400 400 300 300 200 200 100 100 .1 I 0.30.30.2- S.) 0.1- 0 0.0 0.0 I Night Night 1dv Pdv 0!"l Day Day 1dv Pdv o Night C Night Cp DayC Day Cp A A I 0.4 0.4 S.) as I. i 0 I B B 700 700 I I II I I I 600 600 500 500 400 400 300 300 200 200 100 100 0 Distance from Coast Distance from Coast (Km) (Km) 0.4 0.4 Day ,' 0.30.3 I Night, 121 Night, 121 kin km site site o Day, 220 220 km km site Day, site Night 0.20.2 Night, 289 289 km km site Night, site C 0.1.0.1 £& Night. 572 Night, 572km kmSite site ri Model Night, Night, 220 220 km km site Model site c 0.0 0.0 . . 0 2 I 5 8 10 (ml m' m3) dv (ml Pdv 3) Figure dv and and (b) (b) maximum Figure 5. 5. Spatial Spatial relationships relationshipsbetween between(a) (a) maximum maximumnighttime nighttimeand anddaytime daytime Pdv maximum nightnighttime C, over intervals) at tow sites. lines timeand anddaytime daytime Cp(averaged (averaged overthe thetow towdepth depth intervals) atthe thefour fourMOCNESS MOCNESS tow sites.The Thecurved curved lines represent best functions to dv and and C1, data. (c) C1, and Pdv 'dv data represent bestfit fitexponential exponential functions tothe thenighttime nighttime Pdv Cpdata. (c) The Thenighttime nighttime Cpand data follow data follow follow aa trend trendwith withslope slopeof of 0.03 0.03mL mL m-2. m-2.The Thedaytime daytime datafrom fromthe thesite site220 220km kmoffshore offshore followaarelationrelationship shipwith withslope slopeof of0.08 0.08mL mLm-2. m-2. planktonic planktonicforaminiferal foraminiferalstanding standingstock stock>125 >125 .tm gm ranges rangesfrom from 15 to 55 (Table 2). 2). Species diversity 15 to 55 shells shellsrn-3 m-3(Table Species diversityat atthe thefour fourtow tow sites sitesis is typical typicalof ofmidlatitude midlatitudesites sitesnear nearoceanic oceanicmargins margins[Be, [Bd, 1977]. 1977]. A varimax A varimax rotated, rotated, Q-mode Q-mode factor factor model model (Table (Table 4) 4) based basedon on 13 species species in in all 13 all of of the theMOCNESS MOCNESS samples samplesfrom from 00- to to 200-rn 200-m grouped groupedspecies specieswith with similar similar distributions. distributions.These Thesefour four factors factors account for for 82% 25-tm percentaccount 82% of of the theinformation information in in the the>1 >125-gm percent- The seven into three of the The sevenspecies specieslisted listed above abovegroup groupinto threeof the four four faunal of faunal factors. factors.The Thefourth fourthfactor factorisiscomposed composed ofdeep-dwelling deep-dwelling are Individuals of of these asymbiotic asymbiotic species. species. Individuals these species species are maximum normalized data data set. maximum normalized set. Because Because of of the the percentpercentmaximum normalization, all all species maximum normalization, speciesare are equally equally weighted weightedin in this analysis. this analysis. The The resulting resultingfactors factorsare areuseful usefulfor forgrouping grouping and (Figure (Figure10). 10). The The terms terms"shallow," "shallow,""subsurface," "subsurface," and"deep" "deep" species specieswith with similar similardistribution distributionpatterns, patterns,but butthe theinformation information explained by by each each factor explained factor (Table (Table 4) 4) is is not notproportional proportionalto to the the 2). Thus species' species' maximum maximum standing standing stocks stocks (Table (Table 2). Thus comparison of the by each comparisonof the information informationexplained explainedby eachfactor factoris is not not relevant relevant to to aa discussion discussionof of standing standingstock stockdistributions. distributions. "asymbiotic" derives derives from from the of "asymbiotic" the designations designations of Hemleben Hernlebenet et al. al. [1988]. We of [1988]. We assume assumethat thatthese thesegroups groupsdefine defineassemblages assemblages of sometimes present in in the sometimespresent the 00- to to200-m 200-msamples samplesbut butare aremore more common in the the >200-m samples [Ortiz, [Ortiz, 1995]. commonin >200-m samples 1995]. We We refer referto to the the species speciesassociated associatedwith with these thesethree threefactors factorsas asthe the"subsurface "subsurface symbiotic" symbiotic" species species (Figure (Figure 8), 8), the the "shallow "shallowsymbiotic" symbiotic" species species(Figure (Figure 9), 9), and andthe the"shallow "shallowasymbiotic" asymbiotic"species species derive derive from from the the relative relative depth depthof of the thespecies speciesstanding standingstock stock maxima at these these locations. locations. Use maxima at Useof ofthe theterms terms"symbiotic" "symbiotic"and and ecologically ecologically similar similar species. species. The species (N. and O. 0. The "subsurface "subsurfacesymbiotic" symbiotic" species (N. durertrei dutertrei and universa) maximum universa)both bothhave haveoffshore offshoresubsurface subsurface maximumat at the the572572- ORTIZ Ef AL.: OF CURRENT ORTIZET AL.:FORAMINIFERA FORAMINIFERA OFTHE THECALIFORNIA CALIFORNIA CURRENT Total Total Foraminifers Foraminifers (shells (shellsrn-3) m-3) 20 40 60 00 20 40 0- I _ I I Total Total Forarninifers Foraminifers (shells rn-3) (shells m-3) 40 60 0 20 20 40 60 Total Foraminifers Total Foraminifers (shells (shellsrn-3) m-3) 20 40 60 20 40 60 0 00 • • • • -:. :3.4 .':;•:• ............................................................... 50 5050 5o - ..-::.•::.......:½ 0. . Total Forarninifers Total Foraminifers 0. 0 • Size fractions fractions of of Size total plasiktic foraminifers: total plankticforaminifers: ............ .......... 50 5o 505o i!-:::::::::::::::::::::::::::::::::::::::::: ..................... .' iiiiiiiiiiiii •$.... ......... E 100100 (shells rn-3) (shells m-3) 20 40 60 20 40 • 0 0 'i:?.-...,.........•j!,:,,•:i [' ' :,, l00 100 125-150 urn 125-150 um ............... ß ß • 100100 S 997 997 >150 urn >150 um 100100 Cp Cp ;. (. •s0iI 200 -! I I 0.2 0.2 0.3 0.3 0.1 ,50] 200-i 200 I 0.4 0.4 I I 00 0.1 0.1 0.2 0.2 0.3 0.3 0.4 0.4 0 0.4 0.1 0.2 0.3 0.4 0.1 0.2 0.3 0.1 0.1 0.2 0.3 0.3 0.4 0.4 I I I 0 0.1 0.1 0.2 0.2 0.3 0.3 0 0 0.3 0.4 0.1 0.2 0.3 0.1 0.4 i o 50 - 121 km 121 km 0.4 0.4 I I I , U 0 0.1 0.1 0.2 0.2 0.3 0.3 0.4 0.4 0o 0.2 0.3 0.4 0.1 0.2 0.1 0.3 0.4 Cp Cp(rn-') (m-l) Cp (rn-1) Cp (m-1) I o ,50]Night, 200-. 200 I Cp Cp(rn-1) (m-l) Cp Cp(rn-1) (m-l) 0 200 200 I Night. Day, Day, 220 km 220 km [ | 289 289 km km 572 572 km km 200 150- Night. Night, Night, Night, 0 l50 150 150 - i I i o ,,..:.., ,: •:•""i "•" 0 .::..::..:.', ?•'..'..'.:. '.':'.' l 50 50 100 100 100 100 IOOJ 150 150 150 150 150 150 D '[2• 50 -'--: Pdv(rnl Pdv (mlrn-3) m-3) Cp Cp 100 150 - Night, 572 km 200 200 0 I I I I 2 4 66 8 Pdv (ml m-3) or3) Pdv(ml Night, Night, 121 km • [ Mod.night 200 10 10 100 02468 10 0 2 4 6 8 10 200 I 0 ]I eeo I I I 8 i 10 0246810 2 Pdv(ml Pdv (mlrn-3) m-3) 4 6 Pdv (ml (ml rn-3) Pdv m-3) 200 t 0246810 0 2 I I I 4 6 8 10 Pdv (ml (ml rn-3) Pdv m-3) Figure 6. of foraminiferal volume Figure 6. Cumulative Cumulativehistograms histogramsof foraminiferalstanding standingstock, stock,plankton planktondisplacement displacement volume(Pd,,), (Pdv),and and transmissometer particle particle attenuation coefficient (C,,) tow Light shading (upper transmissometer attenuation coefficient (Cp)at atthe thefour fourMOCNESS MOCNESS towsites. sites.Light shading (upper panels) denotes denotes 125to 150-jim panels) 125- to 150-proplanktonic planktonicforaminifera, foraminifera,while while heavy heavy shading shading(upper (upperpanels) panels)indicates indicates>150>150jim planktonic foraminifera. Dashed pm planktonicforaminifera. Dashedlines linesat at the thedaytime daytimesite site(220 (220 km km offshore) offshore)indicate indicatethe the inferred inferredmagmagnitude of dvand and Ct, C,,(see (see Figure Figure 33 and data are are plotted nitude of the thenighttime nighttime Pav andtext textfor fordetails). details).The TheC,, Cpdata plottedover overthe the foraminiferal data data to to demonstrate that the the depth stock maxima maxima at at each foraminiferal demonstratethat depth of of the the foraminiferal foraminiferal standing standing stock each site site is is coincident with with the the particle coincident particleconcentration concentrationmaxima maxima at at each eachsite. site. km site km site and andsurface surfacemaxima maximaat at the theother otherlocations locations(Figure (Figure8). 8). Together, these two species account for most Together, these two species account for most of of the the foraminiferal fauna fauna at at the the four tow sites foraminiferal four tow sitesand andthe the majority majorityof of the the >150-jim foraminifera foramimfera (Table (Table 3). 3). The >150-pm Themaximum maximumabundance abundanceof of these species toward the these speciesdecreases decreasesmonotonically monotonicallytoward the coast, coast,with with the exception of the Columbia River plume site (289 the exceptionof the ColumbiaRiver plume site (289 km km offoffshore), which which has has too of either shore), too few few individuals individualsof either species speciesto to fit fit this general N. duterrrei this generaltrend. trend. N. dutertreiis is the themost mostabundant abundantspecies species collected atat these these sites; sites; O. 0. universa universa was collected was the the third third most most abundant. abundant. The consist of of G. The "shallow "shallowsymbiotic" symbiotic"species speciesconsist G. ruber ruber and andG. G. glutinata. both species rare in glutinata. Although Althoughboth speciesare are relatively relatively rare in these these samples, their their distributions are similar 9). G. samples, distributions are similar (Figure (Figure 9). G. ruber ruber reaches maximum abundance in the 0to 30-m sample reachesmaximum abundancein the 0- to 30-m sampleat atthe the 572- and 572and220-km 220-kin sites. sites. G. G. glurinara glutinata is is most mostabundant abundantin in the the 30- to to 70-rn at 220 3070-m sample sampleat 220 km km and and in in the the00- to to30-rn 30-m sample sample 572 km The 572 km offshore. offshore. 'The two two sites sites with with lowest lowest abundance abundance for for these thesespecies species(121 (121 and and289 289 km) kin) experience experiencesouthward southwardflow flow of of relatively cool, plankton rich waters relatively cool, plankton rich waters (Figure (Figure 3) 3) which which may may have excluded have excludedthese thesesubtropical subtropicalspecies. species. The "shallow asymbiotic" The "shallow asymbiotic" species species (right-coiling (right-coiling N. N. pachyderma, pachyderma, G. G. quinqueloba, quinqueloba,and andG. G. bulloides) bulloides) are aremost most common in in the common the cool cool surface surface waters waters of of the the coastal coastal sites sites where where total biomass biomass is is highest highest (Figure (Figure 10). 10). These total Thesespecies speciesare are rare rare in in the low-salinity waters of the Columbia River plume the low-salinity waters of the Columbia River plume (289 (289 km). At kin). At the the572-km 572-km site sitethey theyare arefound foundfrom from 3030- to to 100 100 m m in in the seasonal the seasonaland andpermanent permanentthermocline. thermocline. Right-coiling Right-coiling N. N. pachyderma and pachyderma andG. G. quinqueloba quinquelobawere werethe thesecond secondand andfourth fourth most common species in these samples. They accounted for most common speciesin these samples. They accountedfor the majority the majority of of the the 125125- to to 150-jim 150-pm foraininifera. foraminifera. Percent Abundance Percent Abundance Comparison Comparison -Between Between MOCNESS MOCNESS and Single-Net and Single-Net Tows Tows Species percent percent abundance Species abundancedata datafrom from the theSeptember September1989 1989 and July and July 1991 1991 single-net single-net tows tows extend extend the the temporal temporal and and 998 998 ORTIZ ET ET AL.: AL: FORAMINIFERA CURRENT ORTIZ FORAMINIFERA OF OF THE THE CALIFORNIA CALIFORNIA CURRENT 60 60 • 505o- .= 40 L • • 0 121 kmsite 121 km site o 220 km km site 220 (using dv) (usingmodel modelnight nightPdv) 3030 289 km site 289 site 20 "Green' "Green" • [] £ 572 572 km km site site ß Water 10 I 2 0 10 8 5 "dv m3) Pdv(ml (mlm' 3) Figure Figure 7. 7. The The relationship relationshipbetween betweenplanktonic planktonicforaminiferal foraminiferalstanding standingstock stockand and Pdv dv. The The 220-km 220-km dv values. values. The The "green" 'green' water foraminifer data are foraminiferdata areplotted plottedagainst againstthe themodel modelnighttime nighttimePdv watersites sites121 121and and289 289km km while the offshore plot along - Pdv dv slope, offshoreplot alongaa single singleline linewith withaa shallow shallowforaminifer foraminiferslope,while the"blue" "blue"water watersite site572 572km kmsite site plots Pd slope. 220 lies these plotsalong alongaa steep steepforaminiferforaminiferPdv slope.The Thesite site 220km kmoffshore offshore liesbetween between thesetwo twoextremes. extremes. Table Species Shells Per >1 50-jim Size Table 3. 3.Foraminiferal Foraminiferal Species Shells PerCubic CubicMeter Meterfor forthe the125125-toto150-jim 150-gmSize SizeClass Classand and >150-gm SizeClass ClassFrom From September 1990 September 1990 63-jim 63-gm Mesh Mesh MOCNESS MOCNESS Samples Samples Site Site Depth,m Depth, rn O.un.a O. un.a G.ru.a G. ru.a G.aq. G. aq. G.ca. G. ca. G.bu.a G. bu.a N.pa.Ra Gd,. N.pa.L G. di. G.qu.a G. qu.a N. pa.L N. pa. R a N.du.a N. du.a G.sc. G. sc. G.me. G. me. G.gl.a G. gl.a 125- to to 150-gm 150-jim Size Class 125Size Class 121 km 0-30 121km 0-30 30-70 30-70 70-100 70-100 100-200 100-200 220 km 0-30 220km 0-30 30-70 30-70 70-100 70-100 100-200 100-200 289 km 0-60 289km 0-60 60-100 60-100 100-200 100-200 572 km 0-30 572km 0-30 30-70 30-70 70-100 70-100 100-200 100-200 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.1 1.1 0.4 0.4 0.1 0.1 0.1 13.5 13.5 1.8 0.8 0.1 7.7 0.8 0.4 0.4 1.0 15.3 15.3 1.1 1.1 0.2 0.2 0.1 0.1 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.9 0.9 0.3 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 1.8 0.8 0.8 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.1 0.1 3.3 3.3 1.7 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.2 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.9 0.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.3 6.3 0.5 0.3 9.0 3.0 0.0 0.2 0.0 0.1 0.2 0.2 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 9.8 9.8 1.4 1.4 0.0 0.0 0.1 3.2 3.2 4.4 4.4 0.0 0.0 0.0 0.0 0.6 0.2 0.0 0.0 0.3 0.2 0.2 0.4 0.2 0.1 0.1 0.0 0.0 0.1 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 6.6 6.6 1.6 1.6 0.1 0.4 0.4 0.0 0.0 0.0 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.0 0.0 1.9 1.9 2.6 2.6 1.6 1.6 0.7 0.7 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.2 0.1 0.0 0.0 >150-jim >150-gm Size Size Class Class 121 121kmkm 0-30 0-30 30-70 30-70 70-100 70-100 100-200 100-200 220km 0-30 220km 0-30 30-70 30-70 70-100 70-100 100-200 100-2• 289 289kmkm 0-60 0-60 60-100 60-100 100-200 100-200 572 km 0-30 572km 0-30 30-70 30-70 70-100 70-100 100-200 100-200 0.0 0.0 0.1 0.1 0.1 0.1 0.1 0.2 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.2 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 00 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.3 0.3 0.0 0.0 0.0 0.0 0.7 0.7 1.8 1.8 0.2 0.2 0.1 0.1 1.3 1.3 4.0 4.0 0.2 0.2 0.7 0.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.5 0.5 0.0 0.0 0.0 0.0 1.4 0.1 0.1 10.6 10.6 4.6 4.6 0.6 0.6 0.1 0.3 0.3 0.0 0.0 0.1 0.4 0.4 1.3 3.5 0.8 0.8 2.9 2.9 0.4 0.4 0.0 0.0 0.1 14.9 4.2 4.2 0.6 0.6 0.1 6.0 6.0 1.8 0.0 0.0 3.4 3.4 25.3 25.3 6.7 6.7 1.5 0.0 0.0 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.4 0.4 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.3 0.2 0.2 0.0 0.0 0.5 0.6 0.6 0.2 0.2 0.0 0.0 0.1 Abbreviations are O. 0. un., ru.. Globigerinoides niber (White); aq.. Globigerinella Globigerinella aequilateralis; aequilateralis; G. Globigerina Abbreviations are un.,Orbulina Orbulinauniversa; universa;G. G. ru., Globigerinoides ruber (White);G. G. aq., G. bu., bu., Globigerina bulloides; G. di., Globigerinella digitata; quinqueloba; N. pa. L., L, Neogloboquadrinapachyderma (Left); bulloides; G. di., Globigerinella digitata;G. G.qu., qu.,Globigerina Globigerina quinqueloba; N. pa. Neogloboquadrina pachyderma (Left);N. N.pa. pa.R., R.,N. N. pachyderma (Right); N. N. du., du., N. dutertrei; G. Globorotalia scitula; scitula; G. me.,Globorotalia Globorotaliamenardii; menardii; and G. gl., gi.. Globigerinita Globigerinira glutinata. glutinata. pachyderma (Right); N. dutertrei; G. sc., sc.,Globorotalia G. me., andG. aa Species discussed in Species discussed in the thetext. text. 999 AL: FORAMINIFERA ORTIZ ET AL.: FORAMINIFERAOF OFTHE THE CALIFORNIA CALIFORNIA CURRENT CURRENT Table for Table 4. 4. Q-mode Q-modeFactor FactorScores Scores forPercent-Maximum Percent-Maximum Transformed TransformedMOCNESS MOCNESS Tow Tow Data Data (0-200 (0-200 m) m) Taxonornic Taxonomic Deep Shallow Shallow Shallow Shallow Deep Asymbiotic Asymbiotic Symbiotic Symbiotic Symbiotic SymbioticAsymbiotic Asymbiotic Species Species Species Species Species Species Species Species Subsurface Subsurface Grouping Grouping 061a N. N. dutertreia dutertrei a 0.61a 053a 0. universaa O. universa a 0.53a 0.47 G. calida G. calida 0.47 0.25 G. G. menardii menardii 0.25 0.12 G. Gorubera ruber a 0.12 0.03 G. glutinataaa G.glutinata 0.03 -0.07 G. aequilateralis G.aequilateralis -0.07 0.11 G. G.digitata digitata 0.11 -0.07 G. G.quinquelobaa quinqueloba a -0.07 N. pachyderma (R)a N.pachyderma (R)a 0.17 0.17 -0.04 G. bulloides bulloidesaa G. -0.04 0.01 G. scitula scitula G. 0.01 N. pachyderma (L) N.pachyderma (L) -0.03 -0.03 Information (in Information (in%) %) 17.9% 17.9% 003 0.03 0.04 0.04 -0.02 -0.02 -0.14 -0.14 -0.17 -0.17 -0.01 -0.01 0.02 0.02 0.00 0.00 0.01 0.01 0.06 0.06 0.05 0.05 -0.10 -0.10 -0.06 -0.06 067a 0.67a 050a 0.11 0.11 0.23 0.23 -0.06 -0.06 0.02 0.02 -0.09 -0.09 0.02 -0.01 003 -0.01 0.03 075a 0.75a 065a -0.11 -0.11 0.09 0.09 10.3% 10.3% 0.09 0.09 18.6% 18.6% within km of within 100 100 km of the the coast, coast, G. G. bulloides bulloides was wasthe thedominant dominant The la). planktonic planktonicforaminifera foraminifera(Figure (Figure 111a). The subsurface subsurface symbiotic species (O. (0. universa symbioticspecies universaand andN. N. dutertrei) dutertrei)were wereless less abundant offshore in in September September 1989 1989(Figure (Figure11c) llc) than abundantoffshore than in in September 1990 (Figure (Figure 1lid). symbiotic species September1990 ld). The Theshallow shallow symbiotic species (G. patterns (G. ruber ruberand andG. G. glutinata) glutinata)had hadsimilar similardistribution distribution patternsin in the two two years years (Figures (Figures 1lie the le and and lit), 1lf),but butthe therelative relativedominance dominance -0.00 0.02 080a 0.80 a 045a 0.45 a 030a 0.30a -0.11 -0.11 accounted accountedfor for =20% --20% of of the theoffshore offshorefauna faunain inSeptember September1989 1989 (Figure 1 la), and =10% of the fauna during September 1990 (Figure11a), and --10% of the faunaduringSeptember 1990 coastal (Figure (Figure1l ib). lb). In In the thehigh-biomass high-biomass coastalupwelling upwellingzone zone -0.00 -0.00 -0.01 -0.01 -0.00 -0.00 -0.00 -0.07 -0.07 -0.08 -0.08 0.50 a 0.42 0.42 0.65 a of the of the two two species speciesswitched switchedbetween betweenyears. years. 35.5% 35.5% Species Species Maxima, Maxima, Environmental Environmental Variability, Variability, and and Size Changes Size Changes The The four-factor four-factor model model accounts accountsfor 82.3% 82.3% of the the transformed transformed information. information. aaThese which the are Thesespecies, species, whichdominate dominate thefactor factorscores, scores, arediscussed discussed in the the text. text. in 0 N. dutertrei dutertrei N. (shells rn-3) (shells m-3) 10 20 30 30 I II 0 0 nI.____ 0 G. G. ruber ruber was wasthe theonly onlyspecies speciesto todemonstrate demonstrateaa clear clear distribution trend trend as as aa function distribution functionof of temperature. temperature. This This species species N. N. duterrrei dutertrei (shells (shellsrn3) m-3) 10 20 30 20 I I 1 Ic, and and 111e; le; geographic range range of of our la, 11c, geographic ourstudy study(Figures (Figures111a, Table 5). These tows show patterns similar to the September Table 5). These tows show patternssimilar to the September 1990tows tows(Figures (Figures1lb, lib,1lid, and 11 lit). 1990 ld, and f). Shallow Shallowasymbiotic asymbiotic species are abundant near the speciesare abundantnear the coast, coast,while while shallow shallowand andsubsubsurface offshore. surfacesymbiotic symbioticspecies speciesare aremore moreabundant abundant offshore. The The offshore offshoretransition transitionfrom from asymbiotic asymbioticto tosymbiotic symbioticdominance dominance appeared sharper 1989 1990. appeared sharperin in September September 1989than thanin inSeptember September 1990. Right-coiling Right-coiling N. N. pachyderma pachydermaand andG. G. quinqueloba quinquelobaeach each N. dutertrei N. dutertrei (shells (shellsrn-3) m-3) I I 0 10 20 I I I 30 30 0 I_...__ 50 50 50 50 50 50 100 100 100 100 100 100 150 150 150 150 150 150 572 289 289 km km km 0 0 10 10 20 20 00 30 30 I 50 .z 100 150 10 20 20 10 20 30 30 0 0 0 I 50 50 50 50 5O 100 100 100 100 100 150 150 150 150 150 572 km 572 km 289 289 km km 200 200 220 km 220 km 200 l 125-150 j.trn 125-150 gm l >150 gm (shells (shellsrn-3) m-3) (shells (shellsrn-3) m-3) 0 II 0. universa O. universa 0. universa O. universa 30 30 I 121 121 km km km 200 200 0. O. universa universa (shells m3) (shellsm-3) 0. universa O. universa (shells m-3) rn3) (shells 200 220 200 200 200 200 N. dutertrei N. dutertrei (shells (shellsrn-3) m-3) 20 30 10 10 10 I[ 20 30 30 II 121 km 121 km 200 200 Cumulative histograms histograms of of the the subsurface symbiotic planktonic foraminifera foraminifera N. N. dutertrei dutertrei and 0. Figure 8. Figure 8. Cumulative subsurface symbiotic planktonic andO. universa denotes small foraminifera (125-150 universaat atthe thefour fourMOCNESS MOCNESStow towsites. sites.Heavy Heavyshading shading denotes smallplanktonic planktonic foraminifera (125-150lim), gm), while light shading indicates larger foraminifera (>150 Rm). These species are most common in whilelightshading indicates largerforaminifera (>150gm). Thesespecies aremostcommon inthe theoffshore offshore thermocline thermocline and and exhibit exhibit surface surfacemaxima maxima at at the the other othersites. sites. ORTIZ E AL.: ORTIZ ET AL.:FORAMINIFERA FORAMINIFERA OF OFTHE THE CALIFORNIA CALIFORNIA CURRENT CURRENT 1000 1000 G. nther G. ruber (shells (shellsrn-3) m-3) G. ruber tuber (shells (shellsrn-3) m-3) 012345 012345 1 2 3 4 5 0 2 3 4 I I I I G. ruber G. tuber (shells rn-3) (shells m-3) 012345 5 0 1 2 3 4 I I I 012345 5 0 50 100 100 100 100 150 150 150 150 150 572 289 km 289 km km 00 0.5 0.5 0.5 0.5 0 0 0 , I G. glutinata G. glutinata (shells rn-3) (shells m-3) 0 0 1 0.5 0.5 1 , 50 50- - 100 100 100 100 100 100 150 150 150 150 150 150 289 km 289 km 200 2oo 220 km 200 4 5 II 0.5 0.5 0 , 50 km 3 II l 125-150 125-150 gm l > 150 gm G. G. glutinata glutinata (shells rn-3) (shells m-3) 50 5o 572 II 200 200 50 200 2 [I 121 km 121 km 200 G. glutinata glutinata G. (shells rn-3) (shells m-3) 1 1 I 220 220 km km 200 G. G. glutinata glutinata (shells rn-3) (shells m-3) 0 5050 100 2OO • 1 50 50 G. ruber ruber G. (shells rn-3) (shells m-3) I 1 , 121 121 km km 200 200 Figure ofofthe G. ruber ruber and and G. G. glutinata glutinata Figure 9. 9. Cumulative Cumulativehistograms histograms theshallow shallowsymbiotic symbioticplanktonic planktonicforaminifera foraminifera G. at tow sites. sites. Shading are at the the four four MOCNESS MOCNESS tow Shadingis isas asin in Figure Figure6. 6. These Thesespecies species aremost mostcommon commonin inthe thewarm warmsurface surface waters waters572 572 km km offshore offshoreand andin in the thenorthward northwardflowing flowingwaters waters220 220km kmoffshore. offshore. decreased decreasedin in abundance abundancewith with decreasing decreasingternperature temperatureand and is is very very rare of less rare in in waters waters of lessthat that 15°C 15øC (Figure (Figure 12b). 12b). None None of of the theother other species speciesdemonstrated demonstratedany any statistically statisticallysignificant significant or or physiophysiologically meaningful standing stock variations with logically meaningful standingstock variations with respect respectto to temperatureor or salinity salinity in in these (Tables 22 and temperature these samples samples (Tables and 3; 3; Figures 12a, Figures 12a, 12b, 12b, and and12c). 12c). 0. O. universa, universa,N. N. dutertrei dutertrei (Figure (Figure 12a), and G. (Figure 12b) 12a), and G. glutinata glutinata (Figure 12b) have have minimum minimum standing standing stock stock near near 15°C 15øC and and higher higher standing standing stocks stockstoward toward both both temperature extremes. N. pachyderma, temperature extremes. Right-coiling Right-coiling N. pachyderma,G. G. quinqueloba, 2c) had quinqueloba,and andG. G. bulloides bulloides(Figure (Figure112c) had patterns patternswhich which were equally equally complex. complex. The were The coarse coarseresolution resolution(0-60 (0-60 m) m) sample sample at the at the site site 289 289 km km offshore offshore does doesnot not greatly greatly affect affect these these results. results.The Theaverage averagetemperature temperaturefrom from 00- to to 60 60 m m at atthis thissite siteis is 14.0°C. The from 00- to 14.0øC. The average averagetemperature temperaturefrom to 30 30 m m is is 16.8°C. 16.8øC. Assuming foraminiferal foraminiferal standing standing stock stock is to Assuming is proportional proportional toC,,, Cp, the 00the 0- to to 60-rn 60-m average averageunderestimates underestimates 0- to to30-rn 30-m forarniniferal foraminiferal standing standingstock stockby by "°lO%, =10%, which which is is less lessthan thanthe the20% 20%measuremeasurement of dv and and C,,). menterror error(see: (see:Comparisons Comparisons of Pttv Cp). Plotting Plottingthe the foraminiferal data data at at 16.8°C, foraminiferal 16.8øC, rather rather than than 14.0°C, 14.0øC, would would not not alter the alter the basic basic structure structure of of these these results. results. The shape The shapeof of the theobserved observeddistribution distributionpatterns patternswith with respect respect to temperature to temperature is is unlike unlike the thegaussian gaussiandistribution distributionpatterns patterns expected based based on expected on each each species' species'optimum optimum temperature temperature as as defined by large-scale Be and and definedby large-scaleplankton planktontow tow studies studies[Be, [Bd,1977; 1977;Bd Hutson, [Bijma Hutson,1977] 1977]and andlaboratory laboratoryculture cultureexperiments experiments [Bijma et et al., al., 1990b]. 1990b]. It It thus thusseems seemslikely likely that thatfactors factorsother otherthan than temperature are are heavily •emperature heavily influencing influencing the the distributions distributionsof of these these species speciesat at these thesesites. sites. In In contrast, contrast,the the relationships relationshipsbetween betweenthe thestanding standingstock stock maxima biomass maxima for for each eachspecies speciesat ateach eachsite siteand andzooplankton zooplankton biomass are physiologically physiologicallyplausible plausible (Figures (Pd) are (Pdv) (Figures 12d,12d, 12e,12e, and and 12f). species (Figures (Figures 12d 12d and and 12e) l2e) 12f). The The shallow shallow symbiotic symbiotic species have have highest highest concentration concentration in in low-biomass, low-biomass, less less turbid turbid offshore offshore waters. waters. These These species speciesdecrease decreasein in abundance abundanceas as biornass biomass and and water water turbidity turbidity increase. increase. For For comparison, comparison,the the shallow asymbiotic asyrnbiotic species species (Figure (Figure 12f) 120 are shallow aremost mostabundant abundantin in the the high-biomass, high-biomass, high-turbidity high-turbidity water water near near the the coast. coast. G. G. bulloides was bulloides was the the least least common common of of the the three three shallow shallow asymbiotic species. species. This rapidly between 99 asymbiotic This species speciesdecreases decreases rapidly between and 55 mL and is dv values values of of 55 and mL rn-3 m-3 and is essentially essentiallyabsent absentbelow below Pdv rnL N. pachyderrna pachyderma decreased rapidly mL rn-3. m-3. Right-coiling Right-coilingN. decreased rapidlybelow below of 6 6 mL had 'dv Pdvvalues valuesof mL rn-3. m-3. G. G. quinqueloba quinqueloba hadroughly roughlyconstant constant dv values standing stock over a large range of standingstock over a large range of Pdv values then then dropped dropped dvfell fell below below 22 rnL rapidly stock rapidlyin in standing standing stockwhen whenPdv mLrn-3. m-3. Shell size size inin O. 0. universa Shell universa and and N. N. dutertrei dutertrei increases increases offshore offshore (Table 6). (Table 6). Increasing Increasingmean mean shell shellsize sizeisispositively positivelycorrelated correlated ORTIZ ORTIZ ET AL.: AL.: FORAMINIFERA FORAMINIFERA OF OF THE THE CALIFORNIA CALIFORNIA CURRENT CURRENT N. pachyderma pachyderma (R) N. (R) N. pachyderma (R) N.pachyderma (R) (shells rn-3) (shells m-3) 00 5 5 10 t0 15 15 20 20 till (shells (shellsrn3) m-3) 0 5 10 10 15 15 20 20 I I N. pachyderma pachyder,na (R) N. (R) N. pachyderma (R) N.pachyderma (R) (shells m3) (shells m-3) (shells rn3) (shells m-3) 00 55 t0 10 15 15 20 20 I 1001 00 55 t0 10 15 15 20 20 0 0 / 50 50 5, 50 50 5o 50 5o 100 too t100 O• 100 100 100 150 15o 150 t5• 150 150 150 150 125-150 125-150 jim gm E 572 572 km km 200 2oo >t50gm 289 289 km km 200 20• G. quinqueloba G.quinqueloba (shells rn-3) (shells m-3) 0 0 5 10 t0 I I I [ G. G.quinqueloba quinqueloba (shells rn-3) (shells m-3) 0 55 10 15 t0 15 I I I I i [ 15 15 I 121 km 220 km 200 200 o 200 200 G. quinqueloba quinqueloba G. (shells rn-3) (shells m-3) G. quinqueloba G.quinqueloba (shells rn-3) (shells m-3) 00 5 10 t0 0 15 15 50 50- 5o 50 50 5o 54 50 100 too- 100 lOO 100 lOO 100 t0• 150 15o 150 15o - 150 15o 572 572 km km 0 I t 2 Ii [ 200 200 G. G. bulloides bulloides (shells rn-3) (shells m-3) 0123 0123 o 3 0 t 2 I II 3 200 200 G. G. bulloides bulloides G. G. bulloides bulloides (shells rn-3) (shells m-3) 0123 0123 (shells rn-3) (shells m-3) 0 0 ti t 2 I I 3 0 50 - 50 50 50 5o 100 too- 100 - 100 100 100 100 150 15o 150 - 150 150 150 200 200 572 km 572 km 200 289 289 km km 200 200 220 220 km km ' t 2 0 50 - 15 220 km 200 200 G. G. bulloides bulloides (shells rn-3) (shells m-3) 10 t0 150 151 289 km 289 km 200 200 5 o I 121 km 200 200 Figure 10. Cumulative histograms histograms of of the asymbiotic planktonic foraminifera right-coiling N. Figure 10. Cumulative theshallow shallow asymbiotic planktonic foraminifera right-coiling N. pachyderma, G. quinqueloba, and G. bulloides at the four MOCNESS tow sites. Shading is as in Figure pachyderma, G. quinqueloba, andG. bulloides at thefourMOCNESStowsites.Shading is asin Figure6. 6. Asymbiotic species species are are most waters near near the offshore. Asymbiotic mostcommon commonin in high-biomass high-biomass waters the coast coastand anddecrease decrease offshore. 0.65, nn = with 0. universa (r2 = with standing standingstock stock(Figure (Figure13a) 13a) for O. universa (r2 = = 0.65, 4) = 0.44, = 4). Size changes 4) and andN. N. dutertrei dutertrei(r2 (r2 = 0.44, nn = 4). Size changesin these lhese species species(Figure (Figure13b) 13b)are areinversely inverselycorrelated correlatedwith withincreasing increasing plankton biomass as by planktonbiomass asmeasured measured by Pdv dv(r2 (r2== 0.96, 0.96,nn == 4, p 0.05 and = 4). right-coiling N. _<0.05 andr2 r2== 0.86, 0.86, nn = 4). In In contrast, contrast, right-coiling N. pachyderma shells do not vary significantly in size across the pachydermashellsdo not vary significantly in sizeacrossthe transect (Table in the transect (Table6). 6). Changes Changesin thesize sizeof ofright-coiling right-coilingN. N. pachyderma in its pachyderma are areuncorrelated uncorrelatedwith with either either changes changesin its standing stock stock (Figure (Figure 13a) or changes in dv (Figure (Figure 13b). standing 13a)or changes in Pdv 13b). Discussion Discussion control control under under favorable favorable thermal thermal conditions. conditions. Because Because the the MOCNESS plankton tows MOCNESS plankton towsused usedfor for this thispurpose purposewere werecollected collected at different sites over a span of only 7 days, we cannot at different sitesover a spanof only 7 days,we cannotaddress address the the possible possiblerole role of of lunar lunarcyclicity cyclicity on on the thestanding standingstock stockand and size sizechanges changeswe we observe observe[Hemleben [Hemlebenet et al., al., 1988; 1988;Bijma Bijma et et al., al., 1990a; Erez Erez et et al., al., 1991]. 1990a; 1991]. However, However,minimum minimum and andmaximum maximum standing stock and standing stock and size size in in these thesesamples samplesare arenot notcorrelated correlated with Longer with the the timing timing of of the thefull full or ornew newmoon moonat atour oursites. sites. Longer records recordsfrom from midlatitude midlatitudesites siteswith withbetter bettertemporal temporalresolution resolution are are necessary necessaryto to address addressthis this topic. topic. With the exception of does With the exceptionof G. G. ruber, ruber,temperature temperature doesnot notplay playaa dominant role in dominant role in determining determining the the mesoscale mesoscaledistribution distribution of of We have tested for relationships between planktonic foraminifera at at the the sites We have tested for relationships between living living planktonic foraminifera siteswe we studied. studied. During During foraminiferal standing stock and size against light availSeptember 1990, G. ruber decreases in abundance foraminiferal standing stock and size against light availSeptember 1990, G. ruber decreasesin abundancewith with dedeability, planktonicbiomass biomass(Pay (d and Ce). and temperature and and is is essentially absent from from waters waters of of creasingtemperature essentiallyabsent ability,planktonic and Cp),temperature, temperature, and creasing salinity. These is the <15°C. G. salinity. These data data suggest suggestthat that while while temperature temperature is the <15øC. G. ruber ruberwas wasabsent absentin inSeptember September1989 1989 when whenmixed mixed dominant on aa species dominant control control on speciesdistribution distribution near near the the limits limits of of its thermal food and its thermaltolerance, tolerance,food and light light may may provide providethe the primary primary layer layer temperatures temperaturesaveraged averaged=2°C --2øC cooler. cooler. The The 14° 14øisotherm isotherm marks the northern limit of its distribution in plankton marks the northern limit of its distribution in planktontows tows 1002 1002 ORTIZ ORTIZ ET AL.: FORAMINIFERA FORAMINIFERA OF OFTHE THECALIFORNTA CALIFORNIA CURRENT CURRENT 100 1 •0 100 100 1989 1989and and1991 1991 80- 80 A I 6060 4040 2020 TEI o....o ..................... 0:::: .... 0........... 0 ...............................I o 00 700 700 800 800 600 600 0 ........FI ........ 500 500 400 400 300 300 60 •0 .0 40 ;0 20 a 0 I 200 200 100 100 700 700 800 800 0 1989 and 1991 100 1989 and 1991 80- iC 80 6060 200 200 1990 1990 80 80- 100 100 0 0 200 200 D :• D ß &.... 20 0 300 300 300 300 40 .... 500 500 400 400 400 400 100 100 404040 [] ......... •. .................................... {•'-. ,...'"'"" \ i 202020 •.............................. '•l'"" .... 0 - I I I I I I I •'a • 600 600 500 500 60 60- A 700 700 600 600 00 ........ N. quinqueloba pachyderma (R) ....... • ....... G. G.quinqueloba ........ N.pachyderma (R) G. bulloides bulloides 100 800 800 B .• B 1990 1990 80 ;0 100 100 800 800 0 •li•: ] I I I I II II I 700 700 600 600 500 500 400 400 300 300 200 200 100 100 0 dutertrei ........• ........ 0. O.universa universa ........I• ........ N. dutertrei 20 20 20 20 iE E 1989 1989and and1991 1991 1515- 1515 *.... 1010- 10 ß 10V... 5 5 5I 00- V1 V 800 700 •F F 1990 1990 600 0 500 300 500 400 400 300 200 200 100 100 00 I 800 800 700 600 700 600 500 300 500 400 400 300 200 200 100 0 Distance Distance offshore offshore(km) (km) cc ruber ........ • ........ G. G.ruber inata ........ •lIE ........ G. G.glut glutinata percent abundance abundance (>125 (>125 gm) Rm) across across the the Multitracers Multitracers transect transect during during the the upwelling Figure Figure 11. 11. Species Speciespercent upwelling season in in 1989, 1989, 1990, 1990,and and1991. 1991. The The 1989 1989 and and 1991 1991data data(Figures (Figures11a, lla, 11c, llc, and season and lie) lie) are arefrom fromsingle-net single-net plankton tows: all all sites were collected collected in in September September1989. 1989. The The tow planktontows: sites>67 >67 km km offshore offshorewere tow data data10 10 km km offshore offshore (solid symbols symbolsin inFigures Figures1lla, were collected ininJuly of G. G. bulloides bulloides (solid l a, lic, 1lc,and andlie) 11e) were collected July1991. 1991.Note Notehigh highabundance abundance of within 100 100 km km of of coast. coast. The 110 areare 0-0to to 200-rn integrated MOCNESS within The 1990 1990data data(Figures (Figureslib, lib, ild, lid,and and 11f) 200-m integrated MOCNESS samples. Vertical lines at 100 km mark the most coastal station from 1990, for comparison to 1989 samples.Verticallinesat 100km markthemostcoastalstationfrom 1990,for comparison to 1989and and1991 1991 tow Species groupings are asymbiotic species (Figures 11 symbiotic towlocations. locations. Species groupings areshallow shallow asymbiotic species (Figures 1l aa and andI1ib), lb),subsurface subsurface symbiotic species, (Figures 1 ic and 1 ld), and shallow symbiotic species (Figures 1 le and I if). species, (Figures1lc and1ld), andshallowsymbiotic species (Figures1le and1If). [Be and Tolderlund, [Bg and Tolderlund, 1971; 1971; Be Be•and and Hamlin, Hamlin, 1967] 1967] and and its its lower lower thermal et al., al., 1990a]. thermal limit limit in in the the laboratory laboratory[Bijma [Bijma et 1990a]. Our Ourfield field results agree agree that that G. G. ruber results ruber does doesnot notsucceed succeedat attemperatures temperatures <14°C. <14øC. Effect Effect of of Food Food and and Light Light on onSpecies SpeciesDistributions Distributions Comparison of total to the dv Comparisonof total foraminiferal foraminiferalstanding standingstock stockto the Pdv data data (Figure (Figure 6) 6) indicates indicatestwo two biological biological provinces provinces in in the the waters waterswe we study, study,coastal coastalupwelling upwellinginfluenced influenced"green" "green"waters waters that that have have low low foraminifera foraminifera to to biomass biomass ratios ratios and and offshore offshore "blue' waters "blue" watersthat thathave havehigher higherforaminifera foraminiferato to biomass biomassratios. ratios. The The foraminiferal foraminiferalfaunas faunasin in the the "green" "green"sites sitesare aredominated dominatedby by asymbiotic asymbiotic foraminifera, foraminifera, while while foraminifera foraminifera with with algal algal symbionts flourish in in the symbiontsflourish the'blue" "blue"waters watersoffshore. offshore. These Thesetwo two faunal provinces reflect the physical and biological faunal provincesreflect the physicaland biologicalchanges changes that occur occur as as one onetraverses traversesthe thecoastal coastaltransition transitionzone zone(CTZ). (CTZ). During increasing particle Duringthe theupwelling upwellingseason, season, increasing particleload loadtoward toward the the coast coastis is associated associatedwith with aa factor factorof of 22 decrease decreasein in the thedepth depth of the 1% PAR light level from =70 m offshore to =35 of the 1% PAR light level from --70 m offshoreto --35 m m near near the shallow dwelling the coast. coast.As Asa aresult, result, shallow dwellingforaminifera foraminiferanear nearthe the coast coastcan canexperience experiencelower lowerlight lightlevels levelsthan thandeeper deeperdwelling dwelling individuals individuals in the the offshore offshore thermocline. thermocline. ORTIZ ET ET AL.: AL.: FORAMINIFERA ORTIZ FORAMINIFERA OF OF THE THE CALIFORNIA CALIFORNIA CURRENT 0) 0 C) r.' C) - - C) C) \O - C) en C) q eq C) C) C) qC) qC)0o ----,,--, c•. 0 o - Q C C..' Ct 0) z 0) b1 C.'1 'I, 0) 2 C) 0 ,•' C) 0 en The in standing stock The depth depthof of the themaximum maximum inforaminiferal foraminiferal standing stock and plankton biomass (F5, Pj,, and C; Figure 6) also depends andplankton biomass (Fss, Pdv,andCp;Figure 6) alsodepends on on the thebiophysical biophysicalenvironment. environment. At the the 121-km 121-km site, site,where where the the Cape CapeBlanco Blancofilament filamentsupplies suppliesnutrient-rich nutrient-richturbid turbidwaters waters from from coastal coastal upwelling upwelling sites, sites, plankton plankton biomass biomass and and foraminiferal standing foraminiferal standingstock stockare are concentrated concentratednear nearthe thesurface surface (0In contrast, (0- to to 30-m 30-m net; net;Figure Figure6). 6)ø In contrast,plankton planktonbiomass biomassand and q 000 q - C)N d d C) C0 o C) C) NC) dr--:. oc•d 0 C) o C) en d 00 - en foraminiferal standing stock stock at at the foraminiferal standing the relatively relatively clear clear water, water, oligotrophic site 572-km offshore is is concentrated in the oligotrophic site 572-km offshore concentrated in the thermocline (30(30- to to 70-rn net), near near the the major of new thermocline 70-m net), major source sourceof new nutrients at nutrients at this this location. location. We infer We infer that that the the foraminiferal foraminiferal concentrations concentrations and and size size changes in in these of changes thesewaters watersarise arisefrom fromthe thedifferent differentresponses responses of asymbiotic and symbiotic foraminifera to changes in food and asymbioticand symbioticforaminiferato changesin food and light. At foraminiferal light. At these thesestudy studysites, sites,asymbiotic asymbiotic foraminiferalstanding standing stock is stock is positively positively correlated correlatedwith with increasing increasingbiomass, biomass, aa measure measureof of greater greaterpotential potentialfood. food. This This implies impliesasymbiotic asymbiotic foraminifera foraminifera become becomeincreasingly increasinglyfood-limited food-limited as asbiomass biomass decreases. decreases. In In the thelaboratory, laboratory,all all species speciesof ofplanktonic planktonicforaminifera foraminifera will accept prey of various types [Hemleben will acceptprey of varioustypes [Hemlebenet et al., al., 1988, 1988,and and references However, prey prey ingestion referencestherein]. therein]. However, ingestionalone alonecannot cannot en 00-N- q'fi C.1 C) en C'l 00 C..' '.0 0 .0 00 C' a0) 0) C/) 2 2 '.0 N C) en N 000 N C..' 00 en 00 C') '.0 fl a en 0 q 0 0 0 C) o C) o eq C) c5 qC)0 C '.0 '. en q explain the standing explain the standing stock stock and and size size distribution distribution of of the the symbiotic symbioticspecies. species. If If food foodwere werelimiting limiting their theirstanding standingstocks stocks and size at these sites, we would observe increasing standing and size at thesesites, we would observeincreasing standing en - - N- C-.' C) c5 F.It Ct C) e N dd c5 C) 0 C) 0 c5 c5 • •o -o CO qC) U 0) qC) 2 I1C 0). C..' C). C). 0 0 C) o A C C) C) o C) C) C) C) C) 0 Q 0 0 0 0 C) c5 C) o C) c5 C) o C) c5 ,•- o C) o C.'1 c',! o o C) c5 asymbiotic asymbiotic species species can can outcompete outcompetesymbiotic symbiotic species speciesin in .0 '.0 •o C) o .0 C) o do C) c5 qQ NO - o C) c5 o C) c5 •. '.0 •o E. C) C) C) ch. 0. o,, o •C') 0-o biomass coastal sites biomasscoastal sites than than in in the theopen openocean. ocean. Because Becauseour our study studydemonstrates demonstratesasymbiotic asymbioticspecies speciesincrease increasein inabundance abundance with we infer with increasing increasingcoastal coastalbiomass, biomass,we infer light light limitation limitationof of C') 2.. foraminiferal standing stocks stocksin in coastal coastal waters. waters. Where foraminiferal standing Wherelight light symbiotic speciesmay may be be the the cause symbiotic species cause of of the the lower lower total total is is abundant, abundant,at at lower lowerlatitude latitudeand andwhere wherewaters watersare areless lessturbid, turbid, symbiotic standing stock stock and symbiotic foraminiferal foraminiferal standing and size sizeincrease, increase, C) while while asymbiotic asymbioticforaminiferal foraminiferalstanding standingstock stockdecreases. decreases. At plankton At plankton biomass biomass levels levels below below those those measured measured at at the the 0) C-) N en en en '.0 en Cfl fl .00 C'. C) en en 000 - - C..' C-.' C') C') C..' 0)0) - C'. C'. C'. C'. Ct that are less that planktonic planktonic foraminifera foraminifera are less abundant abundantin in highhigh- en • •. 000 C) C) 0) - B high-biomass waters and and vice vice versa. versa. Berger high-biomasswaters Berger[1969] [1969]observed observed C) o o 00 C) C) 0 C) C..' C) - C) C) C) C) C) C) fl C) C') 0. d stock In fact, stock and and size sizewith withincreasing increasingplankton planktonbiomass. biomass. In fact, their standing their standing stocks stocksdecrease decreasewith with increasing increasingplankton plankton biomass ambient light. light. We biomassand and increase increasewith with increasing increasingambient We thus thus infer these symbiotic species gain additional nutrition infer these symbiotic speciesgain additional nutritionfrom from their symbionts their symbionts as as light light levels levelsincrease increaseoffshore offshoreinto into lower lower biomass waters. Consistent with with our biomass waters. Consistent our field field observations, observations, published can publishedculture culturework work demonstrate demonstratethat thatendosymbionts endosymbionts can provide provide an an important importantsource sourceof of nutrition nutritionto toforaminiferal foraminiferalhosts hosts [Beet et [Bd et al., al., 1981, 1981,1982; 1982;Spero Speroand andParker, Parker,1985; 1985;JØrgensen JOrgensen et al. al. 1985]. 1985]. Our explanation Our findings findingssuggest suggestaaphysiological physiological explanationfor for why why eq 0 eq e eq N 0)) 1003 -: 00 'd -: - N N '.0 fl C) '.0 en C..' C..' C') - en - en .- - - . - C') 00 -. C'. 00 C'. C'. 00 C'. C'. 00 0'. C'. 00 '. - 0'. C'. en a 0) C) N 0) N 1)0.0'. C') 0) 00 0) '.0 0) - - '.0 Cd) 0'. C/) - Cd) C') 0/) '.0 C/) N Cd) 572-km symbiotic foraminifera may 572-kmsite sitein inthis thisstudy, study, symbiotic foraminifera maybecome become limited by by some If food limited somevariable variableother otherthan thanlight. light. If food were were the the limiting ininsymbiotic limiting agent, agent,this thiswould wouldresult resultin inaadecrease decrease symbiotic species high speciessize sizeand andstanding standingstocks stocksat at very verylow lowbiomass, biomass, high light lightsites. sites. This This seems seemsto to be bethe thecase casein in the thecentral centralequatorial equatorial Pacific than Pacific along alongl40°W 140øWat atsites sitesthat thatare aremore moreoligotrophic oligotrophic than the Multitracers sites [Watkins et al., During the Multitracers sites [Watkins et al., 1995]. 1995]. During sampling, temperature temperature at at these these low low latitude sampling, latitudesites siteswas wasnearly nearly constant, constant,the the foraminiferal foraminiferal community community was was dominated dominatedby by symbiotic species, species, and symbiotic and both both symbiotic symbiotic and andasymbiotic asymbiotic foraminifera were associated with increasing foraminifera were associated with increasingplankton plankton biomass. biomass. Another Another recent recent foraminiferal foraminiferal study study in in the theeastern eastern ORTIZ OF ORTIZEl ETAL.: AL.:FORAMINIFERA FORAMINIFERA OFTHE THECALIFORNTA CALIFORNIACURRENT CURRENT 1004 1004 30 30 t• iE 20 • E 20 10 • 0 -ri- - ---ra--- ' 20 0 i 2 ' i 4 ' i ' 6 i 8 _ 10 10 1.0 -0.8 0 ' 5 -0.6 0.6 . -0.6 0.4 -0.4 0.4 -0.4 0.0 0.0 0 20 15 30 3,• 20- -3 •'I, -22 -1 l 20 •0--- N. pachyderma N.pachyderma (Right) (Right) ----0---.... • .... G. G.quinqueloba quinqueloba • 0(5 ----El--- -- - - G. G.bulloides bulloides 0 0 15 G. G. glutinata glutinata .5 -44 10 ---•--- G. ruber 5 2 E 20 rl 10 0 30 V G. ruber 0.2 -0.2 _ 0.0 0.0 I 10 0.8 -0.8 _ 0.2 -0.2 _ N. dutertrei dutertrei N. ' 1.0 B fr--iX 0. universa O. universa 10 0 I 5 D ,, 0 Temperature (°C) Temperature (øC) 2 4 6 8 10 Pcjv(m1m3) Pdv (mlm-3) dy (a-c) Planktonic foraminiferal foraminiferal standing Figure Figure 12. 12. Planktonic standingstock stockversus versustemperature temperatureand and Pdv. (a-c) G. G. ruber ruberis is the theonly only species to demonstrate a clear relationship to temperature. (d-e) Symbiotic species decrease with increasing speciesto demonstrate a clear relationshipto temperature. (d-e) Symbioticspeciesdecreasewith increasing Measurement errors errors of of 20% 20% apply foraminiferal (t) while species increase with dv "dv' Pdv,(f) whileasymbiotic asymbiotic species increase with Pdv. Measurement applyfor for foraminiferal standing stock and standingstock and Pdv. dy equatorial Atlantic equatorial Atlantic linked linked high highforaminiferal foraminiferalstanding standingstocks stocks to intermediate levels of phytoplankton biomass [Oberhansli to intermediatelevels of phytoplanktonbiomass[OberhZinsli et al., et al., 1992]. 1992]. These These researchers researchersfound found foraminiferal foraminiferal concenconcentration decreased both toward high-biomass, coastal upwelling trationdecreased both towardhigh-biomass, coastalupwelling sites and open-ocean, oligotrophic sites andtoward towardlower-biomass, lower-biomass, open-ocean, oligotrophic sites. that decrease toward sites.We Wesuggest suggest thatthe theoffshore offshore decrease towardoligooligotrophic by Oberhansli trophicconditions conditionsobserved observedby OberhZinsliet etal. al. [1992] [1992]arises arises Accounting differences between Accounting for for differences between symbiotic symbiotic and and asymbiotic species may asymbiotic species may resolve resolve the the apparent apparentconflict conflict discussed by Thunell and Sautter discussedby Thunell and Sautter [1992] [1992] as as to to whether whether N. N. dutertrei dutertrei should should reach reach maximum maximum seasonal seasonal abundance abundance in in upwelling zones zones earlier earlier or or later later than than G. G. bulloides. upwelling bulloides. Kroon Kroon and and Ganssen Ganssen[1989], [1989], working workingon onpump pumpsamples samplesfrom from the thenorthern northern Indian Ocean, stock Indian Ocean,suggest suggestthat thatthe thestanding standing stockmaximum maximumof ofN. N. from from food food limitation limitation of of both bothsymbiotic symbioticand andasymbiotic asymbiotic dutertrei dutertreishould shouldlead leadthat thatof of G. G. bulloides, bulloides,because becauseN. N. dutertrei dutertrei species, toward feeds primarily that peak species,while while the theonshore onshoredecrease decrease towardhigh-biomass high-biomass feeds primarily on on phytoplankters phytoplanktersthat peakin in abundance abundance coastal coastalsites sitesarises arisesfrom fromlight lightlimitation limitationof ofsymbiotic symbioticspecies. species. If this interpretation is correct, our inference If this interpretation is correct, our inference of of light light limitation at at the sites may to limitation the midlatitude midlatitudeMultitracers Multitracerssites may extend extendto turbid in the turbid continental continentalmargins marginsin the tropics tropicsas aswell. well. Table 6. Individual Size Species From Table 6.Mean Mean Individual SizeofofSelected Selected Species From the MOCNESS MOCNESS Tows Tows the Right-coiling Right-coiling Distance before the the zooplankters zooplankters on on which which G. G. bulloides bulloides feeds. feeds. Thunell before Thunell and the pattern andSautter Sautter[1992] [1992]observed observed theopposite opposite patternin in the theSan San Pedro Basin off California. They explain this apparent Pedro Basin off California. They explain this apparent conflict of variability betweer the conflictin in terms terms ofhydrographic hydrographic variability betweer• thetwo two regions and suggest that N. regions and suggestthat N. dutertrei dutertrei prefers prefersaa thermally thermally stratified chlorophyll stratified water water column column with with aapronounced pronounced chlorophyll maximum. maximum. We We argue arguethat thatasymbiotic asymbioticG. G. bulloides bulloidesreaches reaches maximum maximum abundance abundancein in turbid turbid waters watersat at times timesof of greatest greatestfood food availability, availability, while while symbiotic symbioticN. N. dutertrei dutertreican canonly onlyreach reachhigh high N. dutertrei N. Distance 0. O.univera univera N. dutertrei N.pachyderma pachydermastanding standing stocks stocks and and largest largest size sizein inlower-turbidity, lower-turbidity, lowerlowerSize Size Offshore, km Offshore, km Size Size Size Size 343±83 ± 26(168) 343+83 (12) (12) 236±31 236+31 (41) (41) 202 202+26(168) biomass waters. Our biomasswaters. Ourexplanation explanationof of this thispattern patterndiffers differs from from that of Thunell and Sautter or Kroon that of Thunell and Sautter [1992] [1992] or Kroon and and Ganssen Ganssen 403±78 403_+78 (79) (79) 213±38 213_+38 (12) (12) distribution of N. distribution of N. dutertrei dutertrei rather ratherthan than the the light light needs needsof of its its Mean deviation) Meansize size(± (-+one onestandard standard deviation)is is given givenin in microns. microns.Nunber Nunber of is of shells shellsmeasured measured islisted listedin inparenthesis. parenthesis. endosymbionts endosymbiontsas as we we suggest. suggest. Our Our conceptual conceptualmodel model of of food foodand andlight lightlimitation limitationexpands expands upon upon the the current currentparadigm paradigmof of planktonic planktonicforaminiferal foraminiferal dietary dietary 121 121 220 220 289 289 572 572 441 ± 53(107) 441+53(107) 493±89(110) 493_+89 (110) 576±53(153) 576_+53 (153) 342 ± 79(121) 218± 30 (121) 342__+79(121) 218_+30(121) 434±63(201) 434_+63 (201) 231±30 231+ 30 (32) (32) [1989] who who stress of herbivory [1989] stress the the importance importance of herbivory on on the the ORTIZ ET AL.: FORAMINIFERA CALIFORNIA CURRENT ORTIZ FORAMINIFERA OF OFTI-LB THE CALIFORNIA CURRENT 1005 1005 800 800 700 ...... '& ....... 700- 600 • ...._•. .............................. I-i 600- - 500 500 400 400I,4$I. 300-4-. 300 .1. .I .. I I. 200 200 100 100 I 0 10 5 15 15 20 20 25 30 35 Shells Shellsrn-3 m -3 700 B 500 400 D [] 0. O. universa universa O N. dutertrei N. dutertrei o Right-coiling N. N. pachyderma pachyderma Right-coiling ... z 100 . 0 22I ' 4I ' 6I ' 8I ' 110I• ' 12 12 Pth (ml Pdv (mlrn-3) m-3) Figure 13. between dv Two species (O. (0. Figure 13. Relationships Relationships betweenforaminiferal foraminiferalsize, size,standing standingstock, stock,and and Pdv. Two symbiotic symbiotic species universa size and universaand andN. N. dutertrei) dutertrei)demonstrate demonstrate(a) (a) positive positivecorrelation correlationbetween betweensize and standing standingstock stock and and (b) (b) sigsignificant negative between size valuesofofPdv Pa imply turbid water. water. The dv Increasing nificant negative correlations correlationsbetween size and and Pdv. Increasingvalues imply more more turbid The asymbiotic asymbioticspecies, species,right-coiling right-coiling N. N. pachyderma, pachyderma,does doesnot notdemonstrate demonstratesignificant significantcorrelation correlationbetween betweensize size and dy andstanding standingstock stockor or size sizeand and Pdv. needs by synthesizing field and and laboratory laboratory results. results. Much needs by synthesizingfield Much of of the the nutritional nutritional needs needs of of these these species speciesare are met metthrough through omnivorous omnivorousfeeding feeding habits habits[Hemleben [Hemlebenet et al., al., 19881. 1988]. Previous Previous MOCNESS studies from from open-ocean open-ocean sites sites using using 333-gm 333-tm mesh MOCNESS studies mesh nets nets stress stressthe the linkage linkagebetween betweenspecies speciesdistribution distributionand andfood, food, demonstrated by the demonstrated by the presence presenceof of foraminifera foraminifera at at the theDCM DCM [Fairbanks et [Fairbanks et al., al., 1979; 1979; Fairbanks Fairbanks and andWiebe, Wiebe,1980; 1980; Ravelo Ravelo et et active upwelling upwelling (Figure (Figure 1ha). active l a). What What prevents preventsG. G. bulloides bulloides from thriving thriving farther farther offshore? offshore? We from We suggest suggestthat that the the low low offoffshore of G. shore concentrations concentrations of G. bulloides bulloides arise arise from from food food limilimiof this tation. The tation. The distribution distribution of this species speciesserves servesas as an an extreme extreme example of of aa process example processcommon common to to all all three threeof of the theasymbiotic asymbiotic species specieswe we studied. studied. We propose We proposethat that asymbiotic asymbioticspecies specieshave have different different critical critical is to al., al., 1990; 1990; Be Bd et et al., al., 1985]. 1985]. Our Our added added contribution contribution is to evaluate interactionofof food food and factors evaluate the the interaction and light light as as factors contributing to the and to contributing to the nutrition nutrition of of foraminiferal foraminiferal species species and to food Based on on the food thresholds thresholds below below which which each each is is absent. absent. Based the MOCNESS standing stock stock data MOCNESS standing data (Figure (Figure 13), 13), these thesepresumed presumed assess assesstheir their impact impacton onspecies speciesdistribution distributionpatterns patternsin in the themidmidlatitudes. latitudes. the the three three asymbiotic asymbiotic species), species),followed followed by by right-coiling right-coiling N. N. pachyderma, leben et et al. pachyderma, and andfinally finally G. G. quinqueloba. quinqueloba.Hem Hemleben al. [1988] report that that G. [1988] report G. quinqueloba quinquelobaharbors harborscrysophycophyte crysophycophyte Food Food Threshold Threshold for for Asymbiotic Asymbiotic Species Species The of G. The low low concentrations concentrations of G. bulloides bulloides 121 121 km km offshore offshore in in the the California California Current Current are are somewhat somewhat surprising surprising given given the the affinity affinity of of this this species speciesfor for cold, cold,productive productivecoastal coastalupwelling upwelling zones zones [e.g., [e.g., Cullen Cullen and and Prell, Prell, 1984; 1984; Curry Curry et et al. al. ,, 1992; 1992; Thunell and Sautter, Thiede and and J•inger, Junger, 1992]. Thunell and Sautter, 1992; 1992; Thiede 1992]. Low Low concentrations concentrationsin in the theMOCNESS MOCNESS plankton planktontows tows>121 >121 km km offoffshore are are consistent with this this species' rarity in shore consistent with species' rarity in aa yearlong yearlong sediment trap trap time sediment time series seriesfrom from the the same samesites sites[Ortiz [Ortiz and andMix, Mix, 1992]. 1992]. However, However, in in the the September September1989 1989 and and 1991 1991 tows, tows, G. G. bulloides ininhigh inshore bulloides are areencountered encountered highpercent percentabundance abundance inshore of to the of 100 100 km km at at the thehigh-biomass high-biomass sites sites closest closest to the zone zone of of food for G. of food needs needswere were greatest greatestfor G. bulloides bulloides(the (the least leastabundant abundantof symbionts. are similar to those symbionts. These These symbionts symbionts are similar to those of of Globigerinella Globigerinellaaequilateralis aequilateralisand anddo donot notappear appearto tocontribute contributeas as much to host much to hostnutrition nutrition as asdo dodinoflagellate dinoflagellatesymbionts symbionts[Faber [Faber et et al., al., 1988; 1988; 1989]. 1989]. This Thisexplains explainswhy whythe thedistribution distributionof of G. G. quinqueloba quinqueloba in inthis thisstudy studybears bearsgreatest greatestresemblance resemblanceto to asymbiotic The small asymbioticspecies. species. The small contribution contributionto to host hostnutrition nutrition provided symbionts provided by by the thecrysophycophyte crysophycophyte symbiontscould, could,however, however, explain survives atatlower explain why why G. G. quinqueloba quinqueloba survives lowerfood foodconcenconcentrations than right-coiling N. pachyderma and G. bulloides. trationsthan right-coilingN. pachydermaand G. bulloides. Effects of Food Effects of Food and and Light Light on on Shell Shell Size Size The The observed observed correlations correlations between between shell shell size, size,standing standing stock, dv for for O. 0. universa stock,and and Pdv universaand andN. N. dutertrei dutertrei(Figure (Figure13) 13) are are 1006 ORTIZ ET ORTIZ ET AL.: AL.: FORAMINIFERA FORAMINIFERA OF OFTHE THECALIFORNTA CALIFORNIA CURRENT CURRENT consistent plume consistentwith with laboratory laboratory culture culture work work on on symbiont symbiontphotophotoplume and and aa deep deep(70-100 (70-100 m), m),potentially potentially subducted subducted synthesis Bijma et et al., al., 1992]. distribution 572 572 km km offshore offshore than than at at sites sites inshore inshore of of 220 220 km. km. synthesis[Spero, [Spero, 1992; 1992; Bijma 1992]. These Theselaboratory laboratory distribution studies Another studiesdemonstrate demonstratethat that symbiotic symbioticforaminifera foraminifera grown grown under under Another California California Current Current plankton plankton tow tow study studyfound foundsimilar similar high species patterns among high light light conditions conditionsreach reach larger larger size size than thanindividuals individualsgrown grown speciesdistribution distributionpatterns among a a doliodid, doliodid, aa calanid, calanid,and and under et al., under low low light light conditions. conditions. Additional Additional laboratory laboratory studies studies juvenile juvenile euphausids euphausids[Mackas [Mackas et al., 1991]. 1991]. They Theyattribute attributethe the show of these show that that the the symbiotic symbiotic foraminifer foraminifer G. G. sacullifer sacullifer reaches reaches offshore offshore deepening deepening of these species' species'maxima maxima to to filament filament larger larger adult adult size sizewhen whenwell well fed fedand andunder underhigh highlight lightconditions conditions [Be et et al., al., 1981; Caron et [Bd 1981; 1982; 1982; Caron et al., al., 1981; 1981; Caron Caron and and Be, Bd, subduction subduction[Hofinann [Hofmann et et al., al., 1991]. 1991]. While the physical exclusion While the physical exclusionargument argumentmay may explain explainboth both 19841. 1984]. Our Our field field data dataimply implythat thatlarger largerindividuals individualsof of the thesubsub- the the low low abundance abundanceof of asymbiotic asymbioticforaminifera foraminifera in in the thehighhigh- surface surface symbiotic symbiotic species species are are associated associatedwith with higher higher light light conditions d) and conditions (as (as inferred inferred from from lower lower Pdv) andwith withabundant, abundant, thriving as measured thriving populations populations as measured by by their their species' species'standing standing stock. stock. In In contrast, contrast,the the standing standingstock stockand andspecies speciespercentages percentagesof of asymbiotic species increase with food availability. asymbiotic species increase with food availability. Despite Despite this this positive positive correlation, correlation, right-coiling right-coiling N. N. pachyderma, pachyderma, the the most most abundant abundant of of the the asymbiotic asymbiotic species, species, maintains maintains aa constant that constantsize size as asfood food increases increases(Figure (Figure 12). 12). This This suggests suggests that feeding rate does not provide a strong control on calcification feeding rate does not provide a strong control on calcification rate Similarly, other rate in in asymbiotic asymbioticforaminifera. foraminifera. Similarly, other asymbiotic asymbiotic calcifying calcifying organisms organisms (e.g., (e.g., mollusca, mollusca,benthic benthicforaminifera, foraminifera, and and cnidaria) cnidaria) generally generally calcify calcify more more slowly slowly than than related, related, symbiotic counterparts 1988, and symbiotic counterparts [Jones [Jones et et al., al., 1988, and references references therein]. therein]. Comparing Comparing symbiotic symbiotic and and asymbiotic asymbiotic species, species,we we infer availability does does not that food infer that food availability not radically radically alter alter calcification rate rate but but that calcification that symbiont symbiont activity activity does does through through modification of the chemistry at at the the site modification of the carbonate carbonatechemistry site of of calcite calcite deposition. deposition. Asymbiotic Site: Asymbiotic Species Species at at the the Low-Salinity Low-Salinity Site: Salinity Salinity Limits Limits or or Advection? Advection? Despite Despite the the lack lack of of aaclear clearpattern patternbetween betweenasymbiotic asymbiotic species standing standing stock stock and species and salinity, salinity, their their conspicuous conspicuousabsence absence from the the low-salinity River plume from low-salinity Columbia Columbia River plume site site 289 289 km km offoffshore shore(Figure (Figure 10) 10) suggests suggestseither either (1) (1) secondary secondarycontrol control by by aa low-salinity threshold threshold or or (2) (2) physical low-salinity physical exclusion exclusion from from the the plume by the hydrographic front associated with the plume by the hydrographic front associatedwith theCape Cape Blanco filament (Figures (Figures 22 and and 3d). 3d). We Blanco filament We compare comparethe theobserved observed salinity salinity of of the theplume plume(=32.3 (--32.3psu, psu,Figure Figure3) 3)with withlow-salinity low-salinity thresholds The low-salinity thresholds for for each eachspecies. species. The low-salinity threshold thresholdwas was determined determined from from the the species' species'salinity salinity optimum optimum and andstandard standard deviation reported by by Be deviation reported Bg [1977]. [1977]. Be Bg [1977] [1977] defined defined the the salinity optimum for for each each species based on on the salinity optimum species based the mean mean sea sea surface salinity at at the surface salinity the location location of of the thespecies' species'maximum maximum standing standingstock stock using using data data from from over over 150 150plankton planktontows tows(0-200 (0-200 m). We m). We assume assumethat that aaspecies specieslow-salinity low-salinity tolerance toleranceis is 11 standard deviation below below its its optimum. optimum. All standarddeviation All three threespecies specieshave have salinity thresholds thresholds lower lower than than the salinity the salinity salinity of of the theColumbia Columbia River plume: plume: N. N. pachyderma pachyderma (34.1 2.9 -lcy == 31.2 River (34.1±_+ 2.9psu, psu,-1(5 31.2 psu), psu), G. quinqueloba G. quinqueloba (34.5 (34.5 ± + 3.6 3.6 psu, psu, -1 -1(5 = = 30.9 30.9 psu), psu), and and G. G. bulloides (34.8 = 29.7 that bulloides (34.8 ±_+5.1 5.1 psu, psu,-1 -1(5 = 29.7 psu). psu). This Thissuggests suggests that low salinity low salinity was was not not responsible responsible for for their their absence absencefrom from the the plume. plume. Lagrangian drifter studies Lagrangian drifter studies of of filaments filaments in in the theCalifornia California Current that these Current demonstrate demonstrate that these dense, dense, cold, cold, and and saline saline features features subduct subduct under under less less dense dense offshore offshore surface surface waters, waters, or or entrain entrain them, them, rather rather than than diffuse diffuse into into warmer, warmer, fresher, fresher, surface surface waters waters [Paduan Our [Paduan and and Niiler, Niiler, 1990; 1990; Swenson Swenson et etal., al.,19921. 1992]. Our plankton tows tows suggest plankton suggest that that the theasymbiotic asymbiotic planktonic planktonic foraminifera inhabiting inhabiting these these filaments foraminifera filaments record recordmuch much the the same same pattern, with pattern, with exclusion exclusion from from the the less lessdense denseColumbia Columbia River River biomass biomasswaters watersof of the theColumbia ColumbiaRiver River plume plumeand andtheir theirdeeper deeper distribution the distribution at at the the 572-km 572-km site, site, we we cannot cannotentirely entirely exclude exclude the direct affects of low salinity on the foraminiferal distributions direct affects of low salinity on the foraminiferal distributions in in the the plume. plume. This This is is because becausethe theabundant abundantand andshallow shallow symbiotic species symbiotic specieswhich which presumably presumably become become entrained entrainedinto into the the plume plume from from the the offshore offshoreside sideand andthe thesouth southalso alsodecrease decreasein in abundance within 0. universa abundance within it. O. universa and and N. N. dutertrei dutertrei either either are are not as adversely affected by by low low salinity as other as adversely affected salinity as other species species of of planktonic or are planktonic foraminifera foraminifera or are not not excluded excluded as aseffectively, effectively, because the the dense, because dense, cold, cold, and and salty saltyfilaments filamentsassociated associatedwith with the upwelling front are are not not found found on on the side of of the the the upwelling front the offshore offshore side plume. plume. for for Paleoceanographic Paleoceanographic Studies Studies Mix [l989a, bi suggested that temperature Mix [1989a, b] suggestedthat temperatureand andoceanic oceanic productivity (through their influence on respiration productivity (through their influence on respirationrate rate and and available food) food) were available were the the dominant dominant variables variables recorded recorded in in the the Implications Implications foraminiferal fossil fossil record. record. He foraminiferal He further furthersuggested suggestedthat that because because these two were these two processes processes were uncorrelated uncorrelatedat at large largespatial spatialscales, scales,it it was was possible possibleto to reconstruct reconstructthe thebasin-scale basin-scalefeatures featuresof of both both environmental factors through environmental factors through statistical statistical transfer transferfunctions functions using calibration data data set. set. We using the the same sameforaminiferal foraminiferal calibration We have havenot not compared compared our our foraminiferal foraminiferal data data directly directly to to primary primary productivity productivity rate rate measurements, measurements,but but the the Multitracers Multitracers sites sitesare are located located in in aa region regionwhere wherechanges changesin inbiomass biomassand andambient ambient light light are are often often tightly tightly linked linked to to variations variationsin in oceanic oceanicprimary primary productivity. productivity. The The results resultsof ofour ourstudy studyindicate indicatephysiological physiological linkages linkages between between foraminiferal foraminiferal species speciescomposition composition and and variations in ambient variations in ambient light light and and available available food food driven driven by by changes changesin in oceanic oceanicproductivity. productivity. While While the the statistical statisticalmodel model employed by Mix employed by Mix [1989a, [1989a, b] b] did did not notspecifically specificallyincorporate incorporate any any biological biological processes, processes, our our study study supports supports its its basic basic assumptions and suggests suggests ways ways to to improve assumptionsand improve such suchmodels models by by considering symbiotic considering symbiotic and and asymbiotic asymbioticforaminifera foraminifera separately separately and and by by including includingshell shellsize sizeinformation. information. It It is is useful usefulto to consider considerthe theMOCNESS MOCNESS plankton planktontow tow data datain in terms of of percent because this this is is the the form terms percent abundance abundancebecause form of of data data most most easily easily accessible accessible from from the the sediments sediments(Figure (Figure 11). 11). Comparison of the 1989 and 1991 data with the 1990 Comparisonof the 1989 and 1991 data with the 1990 percent percent data yields data yields essentially essentiallythe thesame sameforaminiferal foraminiferaldistributions: distributions: shallow shallow asymbiotic asymbioticspecies speciesare are most mostabundant abundantnear nearthe thecoast, coast, while species are are most most common common offshore. offshore. We while symbiotic symbioticspecies We argue argue that that the the same sameenvironmental environmental factors factors (food (food and and light) light) are are responsible for for the responsible the structure structureof of the theforaminiferal foraminiferalcommunity community during the 1989 upwelling seasons. seasons. The in during the 1989 and and 1990 1990 upwelling The increase increasein percent abundance percent abundanceof of asymbiotic asymbioticspecies species(and (andthus thusdecrease decreaseof of symbiotic symbiotic species) species)in in the the offshore offshorefauna faunaduring during1989 1989(Figure (Figure 11 was accompanied accompaniedby by aa doubling doubling of of the 1 l a)a)was the offshore offshoreparticle particle standing stock stock as by C,, (Figure 4). 4). The in standing asmeasured measured by Cp(Figure Theincrease increase in particle was with particleconcentration concentration wasalso alsoassociated associated withaa2°C 2øCcooling. cooling. This This cooling cooling was was sufficient sufficient to to eliminate eliminate G. G. ruber ruberfrom fromthe the 1989 fauna because because this this species is near 1989 fauna speciesis nearthe the limit limit of of its itsthermal thermal ORTLZET ETAL.: AL.: FORAMINIFERA FORAMINIFERA OF OF THE THE CALIFORNIA CALIFORNIACURRENT CURRENT ORTIZ tolerance at at these these sites. sites. Based tolerance Based on on the the MOCNESS MOCNESS results results from from September 1990 1990 (Figure (Figure 13), 13), it it is September is unlikely unlikely the the2°C 2øCcooling cooling was responsible was responsible for for the the asymbiotic asymbioticspecies speciesincrease increaseand and symbiotic species symbiotic speciesdecrease decreaseduring duringSeptember September1989. 1989. If If these these distribution patterns patterns are are recurrent features of of the distribution recurrent features the high-flux high-flux upwelling season, season, the the factors factors that that give upwelling give rise rise to to them themshould should leave traces traces in in the the sedimentary record. We leave sedimentaryrecord. We would would expect expectto to see see these patterns in shell accumulation rates, community shifts these patternsin shell accumulationrates, communityshifts as measured by percent as measuredby percentabundance, abundance,and and size sizechanges changeswithin within aa species. species. To be applied in this To explore explorehow how size sizeinformation informationmight might be appliedin this context, context, we we reconsider reconsiderthe the results resultsof of sediment sedimentstudies studiesof of shell shell size variations variationsinin O. 0. universa. [1973] assumed size universa. Be B• et et al. al. [1973] assumed temperature controlled 0. temperaturecontrolled O. universa universashell shellsize sizeand andwere wereable ableto to correlated correlated shell shell size size from from Indian Indian Ocean Ocean core core top topsediments sediments with SST with SST as asaapaleo-SST paleo-SSTproxy. proxy. Be Bd and andDuplessy Duplessy[1976] [1976] assumed this this correlation assumed correlation applied applied through through time time and andstudied studied downcore shell size at two downcore shell size variations variations at two sites sites near near the the presentpresentday Subtropical Convergence (STC): RC17-69, =500 km off off day SubtropicalConvergence(STC): RC17-69, =500 km the the SE SE African African coast coast(31.50°S (31.50øS32.60°E) 32.60øE)and andRC9-150, RC9-150, =125 =125 km km off off the the SW SW Australian Australian coast coast(31.28°S (31.28øS 114.5°E). 114.5øE). They They found 0. universa were found O. universashells shells were=200 =200 jim gm larger largerduring during warm warm interglacial than colder interglacial than colder glacial glacial stages. stages. They They suggested suggestedthat that 450-jim sized shells 450-gm sized shells track track the the location location of of the the STC STC and and concluded concluded the the STC STC moved moved as asfar far north north as as331l°S øS(from (from its its modem modem location of 38-40°S) during glacial stages. location of 38-40øS) during glacial stages. Hutson of Hutson [1980] [ 1980] disagreed. disagreed.He He concluded concluded(1) (1) the the location location of the the STC STC was was best bestdescribed describedby by aatransition transitionfrom from aadominant dominant subtropical to aa subantarctic subtropicalto subantarcticforaminiferal foraminiferal fauna fauna and and that that the the STC remained south of the location of RC17-69 during STC remained south of the location of RC17-69 during glacial glacial stages and (2) stagesand (2) size size in in 0. O.universa universawas wasnot notdiagnostic diagnosticof of the the STC STC location. location.Hutson Hutson[1980] [ 1980]provided providedno noexplanation explanationof ofwhat what 1007 1007 biomass waters waters with biomass with abundant abundantfood. food. These These species speciesexhibit exhibit aa range of range of critical critical food food thresholds thresholdsbelow below which whicheach eachwas wasabsent. absent. For example, For example, G. G. bulloides, bulloides, which which has hasthe thegreatest greatestapparent apparent food need need of of these to the food thesespecies, species,is is largely largely confined confinedto the very very high high biomass upwelling upwelling waters waterswithin within 100 100km km of of the the coast coast in in the the biomass study studyarea. area. In contrast, In contrast, ambient ambient light light levels levels seemed seemedto to determine determinethe the distribution of symbiont-bearing species, which distribution of symbiont-bearingspecies,which increased increasedin in Meager standing abundance as as water abundance water turbidity turbidity decreased. decreased. Meager standing stocks and small such as as 0. stocks and small shell shell sizes sizes of of symbiotic symbiotic species speciessuch O. universa and universa and N. N. dutertrei dutertrei are areassociated associatedwith withhigh-biomass high-biomass waters, waters, while while large large individuals individuals with with greater greaterstanding standingstocks stocks occur in in low-biomass waters. These occur low-biomass waters. These results results suggest suggestthat that the the small small individuals individuals of of symbiotic symbioticspecies speciespreserved preservedin in sediments sediments of high-biomass of high-biomass (high-turbidity) (high-turbidity) regions regions reflect reflect stressed stressed growth under under light light limitation reflect growth limitation and and that that larger larger specimens specimensreflect less These field less turbid turbid waters watersand andhigh highsymbiont symbiontactivity. activity. These field observations are consistent observations are consistent with with laboratory laboratory studies studies of of the the foraminiferal foraminiferal host-symbiont host-symbiontcomplex. complex. This plankton This plankton tow tow study study indicates indicates mechanistic mechanistic links links between foraminiferal species composition composition and and variations variations in in between foraminiferal species light driven light availability availabilityand andfood foodconcentration concentration drivenby by changes changesin in oceanic productivity. Accordingly, these findings oceanic productivity. Accordingly, these findingsidentify identify biological processes processes that that support biological support the the reconstruction reconstruction of of oceanic productivity oceanic productivity using usingmultivariate multivariatetransfer transferfunctions functionsand and foraminiferal faunal faunal data data [Mix, foraminiferal [Mix, 1989a, 1989a, b]. b]. We We suggest suggest strategies for reconstructing paleoproductivity from strategiesfor reconstructingpaleoproductivityfrom shell shell size size and the and the relative relative abundance abundanceof of symbiotic symbioticand andasymbiotic asymbiotic foraminifera. Future foraminifera. Future research research on on the the reconstruction reconstruction of of oceanic oceanic paleotemperature and paleoproductivity paleotemperature and paleoproductivity should should focus focus on on might might be be driving driving the theobserved observedglacial-interglacial glacial-interglacial size size developing developing strategies strategiesto to determine determinewhen whenforaminiferal foraminiferalfaunas faunas are are biologically biologically (e.g., (e.g., food food and andlight) light) as asopposed opposedto tothermally thermally controlled. controlled. changes in O. 0. universa. changesin universa. Our Our results resultssuggest suggestan analternative alternativeinterpretation interpretationfor for these these size of 0. size changes changesof O. universa universain inIndian IndianOcean Oceansediments. sediments.Given Given Wecoma, the Multitracers trap group, group, and and M. Wecoma, the Multitracers sediment sediment trap M. Hill Hill and and M. M. the these two the near-coastal near-coastal environments environments ofof these two cores, cores, the the observed observed size size differences differences are are consistent consistent with with changes changes in in ambient light of of the the waters in which 0. universa ambient light waters in which O. universa grew. grew. Smaller Smaller 0. O. universa universashells shellsduring duringglacial glacialperiods periodscould couldimply imply either higher biomass (due to higher productivity) either higher biomass(due to higher productivity)or or cloudier cloudier conditions conditionsalong along the the glacial glacial Indian Indian Ocean Oceanmargin marginthan thantoday. today. Our Our study studyargues arguesthat thatproductivity productivitydriven driven variations variationsin in water water turbidity turbidity are are more more important important than than small smallchanges changes in in temperature as aa control on the the size temperatureas control on size of of 0. O.universa universaand andother other symbiotic symbioticspecies speciesat at midlatitude midlatitudeand andhigh-latitude high-latitudesites. sites. Conclusions Conclusions Temperature does does not not provide Temperature provide the the sole soleor oreven evendominant dominant control on local foraminiferal size and distribution control on local foraminiferal size and distributionpatterns patternsin in the California the California Current Current off off southern southernOregon. Oregon. Temperature Temperature changes are are uncorrelated with changes changes in in standing or changes uncorrelated with standing stock stock or size size of of foraminiferal foraminiferalspecies speciesin in this thisstudy studyregion regionexcept exceptfor for G. G. ruber, ruber, which whichwas wasnear nearits itsthermal thermaltolerance tolerancelimit. limit. Although Although temperature controls controls aa foraminiferal species distribution temperature foraminiferal species' distributionnear near the limits the limits of of its its thermal thermaltolerance, tolerance,food food and and light light appear appear to to provide the provide the primary primary control control under under more more favorable favorable temperatures. temperatures. Shallow-dwelling asymbiotic asymbiotic species Shallow-dwelling species(e.g., (e.g., right-coiling right-coiling N. N. pachyderma, pachyderma,G. G. quinqueloba, quinqueloba,and and G. G. bulloides), bulloides), which which survive by by grazing, survive grazing, were were most mostcommon commonoff off Oregon Oregonin in highhigh- Acknowledgments. the Acknowledgments. We Wethank thank thecaptain captainand andcrew crewof ofR/V R/V Willis Willis for for help help with with the the MOCNESS. MOCNESS. S. S.Richard Richardhelped helpedprocess processsome someof of the as part part of of an project. C. the MOCNESS MOCNESS samples samplesas an REU REU summer summerproject. C. Roesler Roesler provided for toto CA! M. Abbott Abbott and and T T. Strub Strub provided1991 1991PAR PARdata data forcomparison comparison C M provided providedcomputer computerfacilities facilitiesfor forAVHRR AVHRR analysis. analysis. J. J. Huyer Huyer and andB. B. Smith gave gave helpful helpful insights insights on on hydrography. hydrography. The Smith Thetext textwas wasimproved improvedby by comments from N. N. Pisias, commentsfrom Pisias, P. P. Wheeler, Wheeler, M. M. Abbott, Abbott, L. L. Welling, Welling, and and J. J. Watkins from C. C. Ravelo and H. H. Spero. Watkins and and insightful insightfulreviews reviews from Ravelo and Spero. Funding Funding for for this thisproject projectwas wasprovided providedby by aaNASA NASA Graduate Graduatestudent studentfellowship fellowshipto to the project. Curation thefirst firstauthor authorand andby by NSF NSF funding fundingto to the the Multitracers Multitracersproject. Curation of of the the plankton planktontow tow samples samplesatatthe theNORCOR NORCOR Marine Marine Geological Geological Repository Repositoryat at OSU OSU was wasprovided providedby by aagrant grantfrom fromthe theNSF. NSF. References References Be, A. A. W. for of from B6, W. H., H., A A method method forrapid rapidsorting sorting offoraminifera foraminifera frommarine marine plankton J. 33, planktonsamples, samples, J.Paleontol., Paleontol., 33,846-848, 846-848,1959a. 1959a. Be, planktonic foraininifera; I,1,Areal B6,A. A. W. W. H., H.,Ecology Ecologyof ofrecent recent planktonic foraminifera; Arealdisdistribution North 5, tributionin inthe thewestern western NorthAtlantic, Atlantic,Micropaleontology, Micropaleontology, 5, 77-100, 77-100, 1959b. 1959b. Be, A. B6, A. W. W. H., H., Ecology Ecologyof ofrecent recentplanktonic planktonicforaminifera, foraminifera, 2, 2, Bathymetric and seasonal distributionsin in the the Sargasso Sea off off Bathymetric and seasonaldistributions SargassoSea Bermuda, 6, Bermuda,Micropaleontology, Micropaleontology, 6, 373-392, 373-392,1960. 1960. Be, A. A. W. and B6, W. H., H., An An ecological, ecological,zoogeographic, zoogeographic, andtaxonomic taxonomicreview reviewof of recent vol. recentplanktonic planktonicforaminifera, foraminifera,in in Oceanic OceanicMicropaleontology, Micropaleontology, vol. 1, Ramsey, pp. San 1, edited editedby by A. A. T. T.S.So Ramsey, pp.1-100, 1-100,Academic, Academic, SanDiego, Diego, Calif., Calif., 1977. 1977. B6, Subtropical convergence fluctuations B6, A. A. W. W. H., H., and andJ.J.C. C.Duplessy, Duplessy, Subtropical convergence fluctuations and andquaternary quaternaryclimates climatesin inthe themiddle middlelatitudes latitudesof of the theIndian IndianOcean, Ocean, Science, Science,194, 194, 419-422, 419-422, 1976. 1976. BC, A. W. planktonic B6, A. W. H., H., and andW. W. H. H.Hamlin, Hamlin,Ecology Ecologyofofrecent recent planktonic 1008 ORTLZ FT AL.: AL.: FORAMINIFERA OF THE CALIFORNIA CURRENT CURRENT ORTIZ ET foraminifera, foraminifera,3, 3, Distribution Distributionin in the theNorth NorthAtlantic Atlanticduring duringthe thesummer summer of of 1962, 1962,Micropaleontology, Micropaleontology,13, 13, 87-106, 87-106,1967. 1967. Be, B6, A. A. W. W. H., H., and andW. W. H. H.Hutson, Hutson,Ecology Ecologyof ofplanktonic planktonicforaminifera foraminifera and patterns of of life life and in the and biogeographic biogeographicpatterns and fossil fossil assemblages assemblagesin the Indian 23, IndianOcean, Ocean,Micropaleontology, Micropaleontology, 23,369-414, 369-414,1977. 1977. Be, B6, A. A. W. W. H., H., and andD. D. S. S.Tolderlund, Toldefiund,Distribution Distributionand andecology ecologyof of living living planktonic in the planktonic foraminifera foraminifera in the surface surfacewaters watersof of the theAtlantic Atlantic and and Indian by B. Indian Oceans, Oceans,in in The The Micropaleontology Micropaleontologyof of Oceans, Oceans,edited editedby B. M. M. Funnel Funnel and and W. W. R. R. Riedel, Riedel, pp. pp.105-149, 105-149, Cambridge CambridgeUniv. Univ. Press, Press,New New York, York, 1971. 1971. BC, A. W. W. H., H., S. S. M. M. Harrison, Harrison, and and L. L. Lott, LoU, Orbulina Orbulina universa universa d'Orbigny d'Orbigny B6, A. in in the theIndian IndianOcean, Ocean,Micropaleontology, Micropaleontology,19, 19,150-192, 150-192,1973. 1973. BC, A. W. W. H., H., C. C. Hemleben, 0. R. B6, A. Hemleben, O. R. Anderson, Anderson,M. M. Spindler, Spindler,J. J. Hacunda, Hacunda, and of and S. S. Tuntivate-Choy, Tuntivate-Choy,Laboratory Laboratoryand andfield field observations observations of living living planktonic Micropaleontology, 23, planktonicforaminifera, foraminifera,Micropaleontology, 23,155-179, 155-179,1977. 1977. BC, A. A. W. W. H., H., D. D. A. A. Caron, Effects freB6, Caron,and and0. O.R. R.Anderson, Anderson, Effectsofoffeeding feeding frequency quency on on life life processes processes of of the the planktonic planktonic foraminifer foraminifer Globigerinoides sacculifer in culture, J.J.Mar. Globigerinoides sacculifer inlaboratory laboratory culture, Mar.Biol. Biol.Assoc. Assoc. U. K., 61,257-277, 61,257-277, 1981. U. K., 1981. BC, A. A. W. W. H., H., H. H. J. J. Spero, and 0. Effects B6, Spero,and O.R. R.Anderson, Anderson, Effectsof of symbiont symbiont Dickson, M.-L. M.-L. Nitrogen Nitrogen dynamics dynamics in in aa coastal Dickson, coastalupwelling upwellingregime, regime,Ph.D. Ph.D. thesis, 214 pp., Oreg. thesis, 214 pp., Oreg.State StateUniv., Univ., Corvallis, Corvallis,1994. 1994. Dowsett, H. H. J., of foraminiferal transfer Dowsett, J., The Thedevelopment development of aalong-range long-range foraminiferal transfer function and and application North function applicationto to the thePleistocene Pleistocene NorthAtlantic Atlanticclimatic climatic extremes, Paleoceanography, Paleoceanography, 6, extremes, 6, 259-273, 259-273, 1991. 1991. Dryden, A. representation of Dryden, A. L., L., Accuracy Accuracyin in percentage percentage representation of heavy heavymineral mineral frequencies,Proc. Nail. frequencies,Proc. Natl.Acad. Acad.Sci. Sci.U.S.A., U.S.A.,17, 17,233-238, 233-238,1931. 1931. Dymond, J., E. sediments: A Dymond,J., E. Suess, Suess,and andM. M. Lyle, Lyle,Barium Bariumin indeep-sea deep-sea sediments: A geochemical proxy for Paleoceanography, 7, geochemical proxy for paleoproductivity, paleoproductivity, Paleoceanography, 7,163163183, 1992. 183, 1992. Erez, and S. Erez, J., J., A. A. Almogi-Labin, Almogi-Labin, and S. Avraham, Avraham, On On the the life life history history of of planktonic planktonic foraminifera: foraminifera: Lunar Lunar reproduction reproduction cycle cycle in Globigerinoides 6, Globigerinoidessacculifer sacculifer(Brady), (Brady), Paleoceanography, Paleoceanography, 6,295-306, 295-306, 1991. 1991. Faber, W. W. W., W., 0. Faber, O.R. R.Anderson, Anderson,J.J.L. L.Lindsey, Lindsey,and andD. D.A. A.Caron, Caron,AlgalAlgalforaminiferal in the foraminiferalsymbiosis symbiosisin the planktonic planktonicforaminifer foraminiferGlobigerinella Globigerinella aequilateralis, and aequilateralis,I,I,Occurence Occurence andstability stabilityofoftwo twomutually mutuallyexclusive exclusive chrysophyte endosymbionts and and their their ultrastructure, ultrastructure, J. J. Foraminiferal Foraminiferal chrysophyteendosymbionts Res., Res., 18, 18, 334-343, 334-343, 1988. 1988. Faber, and Algal-foraminiferal Faber, W. W. W., W., 0. O.R. R.Anderson, Anderson, andD. D.A.A.Caron, Caron, Algal-foraminiferal symbiosis foraminifer Globigerinella Globigerinella aequilateralis, aequilateralis, elimination symbiosisin in the the planktonic planktonicforaminifer elimination and and reinfection reinfection on on the the life life process processof ofthe theplanktonic planktonic II, foraininifer Globigerionoides sacculifer, Mar. II, Effect Effect of of two twosymbiont symbiontspecies specieson onforaminiferal foraminiferalgrowth growthand and foraminifer Globigerionoides sacculifer, Mar.Biol. Biol.Berlin, Berlin,70, 70,73-86, 73-86, longevity, 1982. 1982. longevity,J. J. Foraminiferal ForaminiferalRes., Res.,19, 19, 185-193, 185-193,1989. 1989. BC, A. A. W. W. H., H., J. J. Bishop, Bishop, M. M. Sverdlove, and Standing stock, FairbanksR., R., and andP. P.Wiebe, Wiebe,Foraminiferan Foraminiferanand andchlorophyll chlorophyllmaximum: maximum: B6, Sverdlove, andW. W. Gardner, Gardner, Standing stock, Fairbanks Vertical and and flux vertical distribution Vertical distribution, distribution,seasonal seasonalsuccession, succession, andpaleoceanographic paleoceanographic vertical distribution and flux of of planktonic planktonicforaminifera foraminiferain in the the significance, Panama Basin, Basin, Mar. Mar. Micropaleontol., Micropaleontol., 9, significance,Science, Science,209, 209, 1524-1526, 1524-1526,1980. 1980. Panama 9, 307-333, 307-333,1985. 1985. Berger, W. ofofliving foraminifera, Fairbanks R. G., G., P. P. H. H. Wiebe, Wiebe, and andA. A. W. W. H. H.BC, B6, Vertical Vertical distribution distributionand and Berger, W. H., H., Ecologic Ecologicpatterns patterns livingplanktonic planktonic foraminifera, Fairbanks isotopic Deep Sea isotopiccomposition compositionof of planktonic planktonicforaminifera foraminiferain in the thewestern westernNorth North Deep SeaRes., Res.,16, 16, 1-24, 1-24, 1969. 1969. Berger, W. foraminifera: Sediment production in Berger, W. H., H., Planktonic Planktonic foraminifera: Sediment production in an an oceanic J. Foraminiferal oceanicfront, front,J. ForaminiferalRes., Res.,1, 1, 95-118, 95-118,1971. 1971. Bernhard, J. M., Bernhard, J. M., Postmortem Postmortemvital vital staining stainingin inbenthic benthicforaminifera: foraminifera: Duration studies, Durationand andimportance importancein in population populationand anddistributional distributional studies,J. J. Foraminiferal ForaminiferalRes., Res.,18, 18,143-146, 143-146,1988. 1988. Bijma, Lunar reproductive Bijma,J., J.,J. J.Erez, Erez,and andC. C.Hemleben, Hemleben, Lunarand andsemi-lunar semi-lunar reproductive cycles planktonic J. Foraminiferal cyclesin in some somespinose spinose planktonicforaminifers, foraminifers,J. Foraminiferal Res., 20, 20, 117-127, Res., 117-127, 1990a. 1990a. Bijma, J., J., W. W. W. Temperature and Bijma, W. Faber FaberJr., Jr.,and andC. C.Hemleben, Hemleben, Temperature andsalinity salinity limits foraminifers in limitsfor for growth growthand andsurvival survivalof ofplanktonic planktonic foraminifers in laboralaboratory J. Res., torycultures, cultures, J.Foraminiferal Foraminiferal Res.,20, 20,128-148, 128-148,1990b. 1990b. Bijma, H. and The Bijma,J., J.,H. H. Hemleben, Hemleben, H. Oberhänsli, Oberhfinsli, andM. M. Spindler, Spindler, Theeffects effectsof of increased planktonic foraminifers increasedwater waterfertility fertilityon ontropical tropicalspinose spinose planktonic foraminifers in laboratory cultures, J. Foraminiferal Res., 22, 242-256, l92. in laboratory cultures, J. Foraminiferal Res.,22,242-256, 19•}2. Bradshaw, J., foraminifera in Bradshaw, J.,Ecology Ecologyof of living livingplanktonic planktonic foraminifera in the thenorth northand and equatorial Pacific Oceans, Oceans, Contrib. Contrib. Cushman Found. Foraminiferal Foraminiferal equatorial Pacific Cushman Found. Res., 10, 25-64, 1959. 1959. Caron, feeding Caron,D. A., A., and andA. A. W. W. H. H.BC, B6, Predicted Predictedand andobserved observed feedingrates ratesof of the planktonic foraminifer Globigerinoides sacculifer, Bull. thespinose spinose planktonic foraminifer Globigerinoides sacculifer, Bull. Mar. Mar. Sd., Sci.,35, 35, 1-10, 1-10, 1984. 1984. Caron, Effects of in Caron,D. A., A., A. A. W. W. H. H. BC, B6, and and0. O.R. R.Anderson, Anderson, Effects ofvariations variations in light ofofthe planktonic foraminifer light intensity intensityon onlife lifeprocesses processes the planktonic foraminifer Globigerinoides sacculifer ininlaboratory culture, J.J.Mar. Globigerinoides sacculifer laboratory culture, Mar.Biol. Biol.Assoc. Assoc. U. K., K., 62, 62,435-452, U. 435-452, 1981. 1981. Caron, D. Caron, D. A., A., W. W. W. W. Faber FaberJr., Jr.,and andA. A. W. W. H. H. BC, B6, Effects Effectsof of temperature temperature and salinity and salinityon on the thegrowth growthand andsurvival survivalof of the theplanktonic planktonicforaminifer foraminifer Globigerinoides .cacculifer, J. Mar. 67, 323-341, Globigerinoides sacculifer,J. Mar. Biol. Biol. Assoc. Assoc.U. U. K., K., 67, 323-341, Atlantic, Science, Science,207, 61-63, 61-63, 1979. 1979. Fischer, Fischer, K. K. M., M., Particle Particlefluxes fluxes in in the theeastern easterntropical tropicalPacific PacificOcean Ocean-Sources Sourcesand and processes, processes,Ph.D. Ph.D. thesis, thesis,225 225 pp., pp.,Oreg. Oreg.State StateUniv., Univ., Corvallis, Corvallis, 1984. 1984. Ganssen, G., G., Isotopic Ganssen, Isotopicanalysis analysisof of foraminifera foraminiferashells: shells:interference interferencefrom from chemical treatment, Palaeogeogr. Palaeogeogr. Palaeoclimatol., chemical treatment, Palaeoclimatol.,Palaeoecol., Palaeoecol.,33, 33, 271-276, 271-276, 1981. 1981. Gardner, D. Walsh, Biophysical Gardner,W. W. D., D., I.I.D. Walsh,and andM. M. J.J.Richardson, Richardson, Biophysicalforcing forcing of of particle particleproduction productionand anddistribution distributionduring duringaa spring springbloom bloomin in the the North NorthAtlantic, Atlantic,Deep DeepSea SeaRes., Res.,Part Part11, II, 40, 40,171-195, 171-195,1993. 1993. Hecht, A.D., A. D., An Hecht, An ecologic ecologicmodel model for for test testsize sizevariation variationin in recent recentplankplanktonic to the tonicforaminifera: foraminifera:Applications Applicationsto the fossil fossilrecord, record,J. J. Foraminiferal Foraminiferal Res., Res., 6, 6, 295-311, 295-311, 1976. 1976. Hecht, D., and Hecht,A. A.D., andS. S.M. M. Savin, Savin,Phenotypic Phenotypicvariation variationand andoxygen oxygenisotope isotope ratios J. Foraminiferal ratiosin in recent recentplanktonic planktonicforaminifera, foraminifera,J. ForaminiferalRes., Res.,2, 2,555557, 1972. 1972. Hemleben, C., M. M. Spindler, and O. 0. R. Hemleben, C., Spindler, and R. Anderson, Anderson,(Eds.), (Eds.), Modern Modern Planktonic New PlanktonicForamini/'era, Foraminifera,363 363pp., pp.,Springer-Verlag, Springer-Verlag, NewYork, York,1988. 1988. Hofmann, Hofmann, E. E. E., E., K. K. S. S. Hedstrdm, Hedstr6m,J. J. R. R. Moisan, Moisan,D. D. B. B. Haidvogel, Haidvogel,and andD. D. L. L. Mackas, Mackas, Use Use of of simulated simulateddrifter drifter tracks tracksto toinvestigate investigategeneral general transport times Transition Zone, transportpatterns patternsand andresidence residence timesin inthe theCoastal Coastal Transition Zone, J. J. Geophys. Geophys.Res., Res.,96, 96, 15,041-15,052, 15,041-15,052,1991. 1991. Hutson, Hutson, W. W. H., H., The The Agulhas AgulhasCurrent Currentduring duringthe theLate LatePleistocene: Pleistocene: Analysis Science, 207, Analysisof of modern modemfaunal faunalanalogs, analogs, Science, 207,64-66, 64-66,1980. 1980. Huyer, Huyer, A., A., Coastal Coastalupwelling upwellingin in the theCalifornia CaliforniaCurrent CurrentSystem, System,Prog. Prog. Oceanogr., Oceanogr.,12, 12,1434-1450, 1434-1450,1983. 1983. Ikeda, of Ikeda, M., M., and and W. W. J. J.Emery, Emery,Satellite Satelliteobservations observationsand and modeling modelingof meanders meandersin in the the California CaliforniaCurrent CurrentSystem Systemoff off Oregon Oregonand andnorthern northern California. J. Phys. California.J. Phys.Oceanogr., Oceanogr.,14, 14, 1434-1450, 1434-1450,1984. 1984. Imbrie, J. and method Imbrie, J. andN. N. G. G.Kipp, Kipp,AAnew newmicropaleontological micropaleontological methodfor for quantitative paleoclimatology: Application to to aa Late quantitative paleoclimatology:Application LatePleistocene Pleistocene Caribbean core, core, in in The by K. K. Cullen J. J. L., Caribbean The Late Late Cenozoic CenozoicGlacial Glacial Ages, Ages, edited edited by Cullen L., and and W. W. L. L. Prell, Prell, Planktonic Planktonicforaminifera foraminifera of of the the northern northern Turekian, pp. pp. 71-181, Indian Turekian, 71-181, Yale Yale Univ. Univ. Press, Press,New New Haven, Haven,Conn., Conn.,1971. 1971. Indian Ocean: Ocean: Distribution Distribution and and preservation preservationin in surface surfacesediments, sediments, Jones, D. D. S., Mar. Micropaleontol., Jones, S., D. D. F. F. Williams, Williams, and and H. H. J.J.Spero. Spero,More More light lighton onphotophotoMar. Micropaleontol.,9, 9, 1-52, 1-52,1984. 1984. symbiosis in in fossil fossil mollusks: mollusks:The Thecase caseof ofMercenaria Mercenaria 'tridacnoides", Cullen J. J. J., J., M. symbiosis "tridacnoides", Cullen M. R. R. Lewis, Lewis,C. C. 0. O.Davis, Davis,and andR. R.1. T.Barber, Barber,Photosynthetic Photosynthetic Palaeogeogr., Palaeoclimatol., Palaeoecol., 64, characteristics and estimated estimated growth growth rates rates indicate is the Palaeogeogr., Palaeoclimatol., Palaeoecol., 64,141-152, 141-152,1988. 1988. characteristicsand indicate grazing grazing is the Jørgensen, B. B. B., B., J. proximate control Pacific, J0rgensen, J. Erez, Erez, N. N. P. P.Revsbech, Revsbech,and andY. Y. Cohen, Cohen,Symbiotic Symbiotic proximate controlof of primary primaryproduction productionininthe theequatorial equatorial Pacific,.1. J. photosynthesis in aa planktonic Geophys. Res., 97, 97, 639-654, 639-654, 1992. photosynthesisin planktonicforaminiferan, foraminiferan, Globigerinoides Globigerinoides Geophys.Res., 1992. sacculifer Limnol. Curry, B., B., D. producsacculifer(Brady), (Brady),studied studiedwith withmicroelectrodes, microelectrodes, Limnol.Oceanogr., Oceanogr., Curry, D. R. R. Ostermann, Ostermann,and and M. M. V. V. S. S. Gupta, Gupta,Foraminiferal Foraminiferalproduc30, tion and 30, 1253-1267, 1253-1267, 1985. 1985. tion and monsoonal monsoonalupwelling upwelling in in the theArabian ArabianSea: Sea:Evidence Evidencefrom from Kipp, N. sediment traps, traps, in Kipp, N. G., G., New New transfer transferfunction functionfor forestimating estimatingpast pastsea-surface sea-surface sediment in Evolution Evolutionof ofUpwelling UpwellingSystems: Systems: Miocene Miocene to to conditions Present, edited W. Prell, conditionsfrom from sea-bed sea-beddistributions distributionsof ofplanktonic planktonicforaminiferal foraminiferal Present, edited by by C. C. P. P. Sunanerhayes, Summerhayes,W. Prell,and andK. K. C. C. Emeis, Emeis, assemblagesinin the the north north Atlantic, in investigations of Late Geol. assemblages Atlantic, in Investigations of Late Geol. Soc. Soc.Spec. Spec.PubI. Publ. London, London,64, 64, 93-106, 93-106, 1992. 1992. Quaternary and edited Dam, H. H. G., QuaternaryPaleoceanography Paleoceanography andPaleontology, Paleontology, editedby by K. K. K. K. Dam, G., C. C. A. A. Miller, Miller, and andS. S.H. H. Jonasdottir, Jonasdottir,The The trophic trophicrole role of of Turekian, Geol. Mem. 145, 3-41, 1976. mesozooplankton at at 47°N, Turekian, Geol. Soc. Soc.Am. Mem. 1976. mesozooplankton 47øN, 20°W 20øW during during the the North North Atlantic Atlantic Bloom Bloom Experiment, Deep 1-195, 1993. Kiovan, Experiment, Deep Sea SeaRes., Res.,Part Part ii, II, 40, 40,17 171-195, 1993. Klovan,J. J. B., E., and andJ. J.Imbrie, Imbrie,An Analgorithm algorithmand andFORTRAN FORTRAN IV IV program program 1987. 1987. Climate: Long-Range Long-Range Investigation, Prediction, Climate: Investigation, Prediction,and andMapping Mapping(CLIMAP) (CLIMAP) Project Members, Members, The The surface surface of of the Project the ice ice age ageEarth, Earth,Science, Science,191, 191, 1131-1137, 1976. 1976. ORTIZ FT AL: OF CURRENT ORTIZET AL.:FORAMINIFERA FORAMINIFERA OFTHE THECALIFORNIA CALIFORNIA CURRENT for for large largescale scaleQ-mode Q-modefactor factoranalysis, analysis,J. J.In:. lnt.Assoc. Assoc.Math. Math. Geol., Geol.,3, 3, 61-77, 61-77, 1971. 1971. Kroon D., and cells and and Kroon D., and G. G. Ganssen, Ganssen,Northern NorthernIndian Indian Ocean Oceanupwelling upwellingcells Smith, A. A. B., B., Distribution of foraminifera in the the northSmith, Distribution ofliving livingplanktonic planktonic foraminifera in north- eastern Found. Res., easternPacific, Pacific,Contrib. Contrib.Cushman Cushman Found.Foraminiferal Foraminiferal Res.,14, 14,1115, 15, 1963. 1963. the stable the stableisotope isotopecomposition compositionof of living livingplanktonic planktonicforaminifera, foraminifera, Deep Sea Part A, Deep Sea Res., Res., Part A, 36, 36, 1219-1236, 1219-1236, 1989. 1989. Laurs, R. R. M., levels, Laurs, M., Coastal Coastalupwelling upwellingand andthe the ecology ecologyof of lower lower trophic trophiclevels, Ph.D. thesis, thesis, 121 Ph.D. 121 pp., pp., Oreg. Oreg.State StateUniv., Univ., Corvallis, Corvallis,1967. 1967. Smith, A. A. B., along an an eastSmith, B., Living Livingplanktonic planktonicforaminifera foraminiferacollected collectedalong east- west in the west traverse traverse in the north north Pacific, Pacific, Contrib. Contrib. Cushman Cushman Found. Found. Foraminiferal 1-134, 1964. ForaminiferalRes., Res.,15, 15,13 131-134, 1964. Longhurst, A. in of Longhurst, A. R., R.,Vertical Verticalmigration, migration, inThe TheEcology Ecology ofthe theSeas, Seas,edited edited by J. Walsh, by D. D. H. H. Cushing Cushingand and J. J. J. Walsh,pp. pp. 116-137, 116-137,Saunders, Saunders, Philadelphia, Philadelphia,1976. 1976. Loubere, parameters reflected distriLoubere,P., P., Oceanographic Oceanographic parameters reflectedin in the theseabed seabed distribution of bution of planktic planktic foraminifera foraminifera from from the the North North Atlantic Atlantic and and Mediterranean Sea, J. Fora,niniferal Res., Mediterranean Sea,J. Foraminiferal Res.,11, 11,137-158, 137-158,1981. 1981. Lyle, M., M., R. R. Zahn, N. Pisias, and E. E. Lyle, Zahn,F. F. Prahl, Prahl,J. J.Dymond, Dymond,R. R. Collier, Collier,N. Pisias,and Suess, and carbon Suess,Paleoproductivity Paleoproductivityand carbonburial burialacross acrossthe theCalifornia California Current: The The Multitracers Multitracers transect, transect, 42øN, 42°N, Paleoceanography, Paleoceanography, 7, 251Current: 7, 251272, 1992. 272, 1992. Mackas, D. and Zooplankton community Mackas, D. L., L.,L. L.Washburn, Washburn, andS. S.L. L.Smith, Smith, Zooplankton community pattern with aa California J. pattern associated associatedwith California Current Current cold cold filament, filament, J. Geophys. Res., 96, 14,781-14,798, 1991. Geophys.Res.,96, 14,781-14,798, 1991. Miles, G. G. A., in the Miles, A., Living Living planklonic planktonicforaminifera foraminiferain the northeast northeastPacific Pacific Ocean, Ocean,M.A. M.A. thesis, thesis,131 131pp., pp.,Univ. Univ.of of Oreg., Oreg.,Univ. Univ.of of Oreg. Oreg.Press, Press, Eugene, Eugene,1973. 1973. Mix, paleoproductivity: Evidence from Mix, A. A. C., C.,Pleistocene Pleistocene paleoproductivity: Evidence fromorganic organic carbon species, in Productivity Productivity of of the the Oceans: Oceans: Past Past carbonand andforaminiferal foraminiferal species, in and V. and and Present, Present,edited editedby by W. W.H.H.Berger, Berger, V.S.S.Smetacek, Smetacek, andG. G. Wefer, Wefer, pp. pp.313-340, 313-340,John JohnWiley, Wiley,New NewYork, York,1989a. 1989a. Mix, of variations on atmoMix, A. A. C., C.,Influence Influence ofproductivity productivity variations onlong-term long-term atmospheric Nature, 337, sphericCO2. CO2,Nature, 337, 541-544, 541-544,1989b. 1989b. Molfino, B., B., Statistics Molfino, Statisticsdefine defineecologic ecologiccontrols controlsof of fossil fossilplanktonic planktonic forams in Volume, GEOMAR forams(Abstract), (Abstract), inJCP ICPIV IVAbstract Abstract Volume, GEOMARRep. Rep.15, 15, edited editedby by M. M. Sarnthein, Samthein,J. J.Thiede, Thiede,and andR. R. Zahn, Zahn,p. p. 204, 204,GEOMAR, GEOMAR, Kid, Kiel,Germany, Germany,p. p.204, 204, 1992. 1992. Molfino, of Molfino,B., B., N. N. G. G. Kipp, Kipp,and andJ.J.Morley, Morley,Comparison Comparison offoraminiferal, foraminiferal, Coccolithophorid, and paleotemperature equations: Coccolithophorid, andRadiolarial Radiolarial paleotemperature equations: Assemblage coherency Quat. Assemblage coherencyand andestimate estimateconcordancy, concordancy, Quat.Res., Res.,17, 17, 279-313, 279-313, 1982. 1982. Oberhansli, H., C. Bénier, and Oberh•insli, H., B6nier,G. G. Meinecke, Meinecke,H. H. Schmidt, Schmidt,R. R. Schneider, Schneider, and G. Wefer, G. Wefer, Planktonic Planktonicforaminifers foraminifersas astracers tracersof of oceanic oceaniccurrents currentsin in the eastern eastern south south Atlantic, Atlantic, Paleoceanography, Paleoceanography, 7, the 7,607-632, 607-632,1992. 1992. Ortiz, foraniinifers of Current: Last Ortiz,J. J.D., D.,Planktic Planktic foraminifers ofthe theCalifornia California Current: Lastglacial glacial maximum and present, maximumand present,Ph.D. Ph.D. thesis, thesis,211 211 pp., pp.,Oreg. Oreg.State StateUniv., Univ., Corvallis, 1995. 1995. Ortiz, and sucOrtiz, J. J. D., D., and andA. A. C. C.Mix, Mix, The Thespatial spatialdistribution distribution andseasonal seasonal succession in the cession of of planktonic planktonic foraminifera foraminifera in the California California Current Current off off Oregon, Evolution Oregon,September September1987-1988, 1987-1988,in inUpwelling UpwellingSystems: Systems: Evolution Since W. L. Sincethe theEarly EarlyMiocene, Miocene,edited editedby byC. C. P. P.Summerhayes, Summerhayes, W. L. Prell, Prell, and 197-2 13, 1992. andK. C. C. Emeis, Emeis,Geol. Geol.Soc. Soc.Spec. Spec.PubI. Publ.64 64London, London, 197-213, 1992. Paduan, J. description of in Paduan, J.D., D.,and andP. P.P.P.Niiler, Niiler,AAlagrangian lagrangian description ofmotion motion in northern California coastal transition filaments, .1. Res., 95, northern California coastal transition filaments, J.Geophys. Geophys. Res.,95, 18,095-18,109, 18,095-18,109, 1990. 1990. Pak, Meridional in Pak,H., H., D. D. A. A. Kiefer, Kiefer,and andK. K.C. C.Kitchen, Kitchen, Meridionalvariations variations in the the concentration of and in concentration of chlorophyll chlorophyll andmicroparticles microparticles inthe theNorth NorthPacific Pacific Ocean,DeepSeaRes.,partA,35, Ocean,DeepSeaRes.,Part A, 35,1151-1171, 1151-1171,1988. 1988. 1009 1009 Snedecor, G. G. W., W., and and W. Statistical Methods, 8th ed., 503 Snedecor, W.G. G.Cochran, Cochran, Statistical Methods, 8thed., 503 pp., pp., Iowa Iowa State StateUniv. Univ. Press, Press,Ames, Ames,1989. 1989. Spero, foraminifera accurately record Spero,H. H. J., J.,Do Doplanktic planktic foraminifera accurately recordshifts shiftsin inthe the carbon isotopic composition of seawater Mar. carbonisotopic composition of seawaterECO2?, •CO27 , Mar. Micropaleontol., 19, Micropaleontol., 19, 275-285, 275-285, 1992. 1992. Spero, H. Photosynthesis in planktonic Spero, H. J., J.,and andS. S.L. L. Parker, Parker, Photosynthesis inthe thesymbiotic symbiotic planktonic foraminifer and contribution to foraminiferOrbulina Orbulinauniversa, universa, andits itspotential potential contribution to oceanic primary J. Res., 15, 1, oceanic primaryproductivity, productivity, J.Foraminiferal Foraminiferal Res., 15,273-28 273-281, 1985. 1985. Spinrad, R. R. W., W., A diagram of specific beam attenuation, attenuation, J. J. Spinrad, A calibration calibration diagram of specific beam Geophys. Res., Res., 91, 7761-7764, Geophys. 7761-7764,1986. 1986. Spinrad, R. W., and Spinrad,R. W., H. H. Glover, Glover,B. B. B. B.Ward, Ward,L. L.A. A.Codispoti, Codispoti, andG. G. Kullenberg, Suspended Suspended particle particle and Kullenberg, and bacterial bacterialmaxima maximain in Peruvian Peruvian coastal waters Deep coastal watersduring duringaa cold coldwater wateranomaly, anomaly, DeepSea SeaRes., Res.,Part PartA, A, 36, 715-733, 36, 715-733, 1989. 1989. Stewart, R. H., of Oceanography, 360 Stewart, R. H., Methods Methods ofSatellite Satellite Oceanography, 360pp., pp.,Univ. Univ.of of Calif. Press, Press, Berkeley, Calif. Berkeley,1985. 1985. Strub, P. P. T., T., P. The Strub, P. M. M. Kosro, Kosro,A. A. Huyer, Huyer,and andCTZ CTZCollaborators, Collaborators, Thenature nature of the the cold in Current System. J. Geophys. of coldfilaments filaments in the theCalifornia California Current System. J. Geophys. Res., 96, 14,743-14,768, Res., 96, 14,743-14,768, 1991. 1991. Swenson, M. S., Swenson, M. S., P. P. P. P. Niiler, Niiler,K. K. H. H.Brink, Brink,and andM. M. R. R.Abbott, Abbott,Drifter Drifter observations of a a cold off in July July observations of coldfilament filament off Point PointArena, Arena,California, California, in 1988, J. .1. Geophys. Geophys. Res., Res., 97, 97, 3593-36 10, 1992. 1988, 3593-3610, 1992. Thiede, J., J., and Thiede, and B. B. Jtinger, Jtinger,Faunal Faunal and and floral floral indicators indicatorsof of coastal coastal upwelling, in Systems: Evolution Since Since the the Early Early Miocene, Miocene, upwelling, inUpwelling Upwelling Systems: Evolution edited by by C. W. Geol. edited C.P.P.Summerhayes, Summerhayes, W.L. L.Prell, Prell,and andK. K.C. C.Emeis, Emeis, Geol. Soc. Spec. Spec. Publ. Publ. London, London, 64, 47-76, Soc. 47-76, 1992. 1992. Thunell, R., and and L. R. foraminiferal faunal Thunell, R. Sautter, Sautter,Planktonic Planktonic foraminiferal faunaland andstable stable isotopic indices of A trap isotopic indices of upwelling: upwelling: A sediment sediment trapstudy studyin inthe theSan SanPedro Pedro Basin, California Systems: Evolution Basin,Southern Southern CaliforniaBight, Bight,in inUpwelling Upwelling Systems: Evolution Since the Early Early Miocene, Miocene, edited W. Sincethe editedby byC. C.P.P.Summerhayes, Summerhayes, W. L. L. Prell, Prell, and 64,, 1992. andK. K. C. C. Emeis, Emeis,Geol. Geol.Soc. Soc.Spec. Spec.Pubi. Publ.London, London, 64,,197-213, 197-213, 1992. Walton, W. R., Techniques for recognition of living foraminifera, Walton,W. R., Techniques for recognition of living foraminifera, Contrib. Found. Res., Contrib.Cushman Cushman Found.Foraminiferal Foraminiferal Res.,3, 3,56-60, 56-60,1952. 1952. Watkins, J., J., A. Living foraminifera: Watkins, A. C. C. Mix, Mix,and andJ.J.Wilson, Wilson, Livingplanktonic planktonic foraminifera: Tracers water and regimes Tracersof of surface surface watermasses masses andproductivity productivity regimesin in the the central Equatorial Pacific, Res., Part Part II, 1995. central Equatorial Pacific,Deep DeepSea SeaRes., II, in in press, press, 1995. L. A., A., N. N. G. 0. Pisias, as tracers tracers of of Welling, L. Pisias,and andA. A. K. K. Roelofs, Roelofs,Radiolaria Radiolariaas currents and in California Current System: currents andwater watermasses masses inthe thenorthern northern California Current System: A study, Eos A MOCNESS MOCNESSstudy, Eos Trans. Trans.AGU, AGU, 72(17), 72(17), Spring SpringMeet. Meet. Supplement, Supplement,156, 156,1991. 1991. Welling, Radiolarian microfauna Welling,L. A., A., N. N. G. G. Pisias, Pisias,and andA. A.K. K.Roelofs, Roelofs, Radiolarian microfauna in California Current System: Indicators of in the thenorthern northern California Current System: Indicators of multiple multiple processes controlling productivity, in Evolution processes controlling productivity, inUpwelling UpwellingSystems: Systems: Evolution Since the Early Early Miocene, edited by W. Sincethe Miocene, edited byC. C.P. P.Summerhayes, Summerhayes, W. L. L. Prell, Prell, and 64, andK. K. C. C. Emeis, Emeis,Geol. Geol.Soc. Soc.Spec. Spec.PubI. Publ.London, London, 64,177-197, 177-197,1992. 1992. Wiebe, P. P. H., Wiebe, H., K. K. H. H. Burt, Burt,S. S.H. H.Boyd, Boyd,and andA. A.W. W.Morton, Morton,AAmultiple multiple opening/closing net net and and environmental sensing system system for for sampling opening/closing environmental sensing sampling Parker, F., Planktonic foraminiferal species in Pacific sediments, zooplankton, J. Parker,F., Planktonic foraminiferal speciesin Pacificsediments, zooplankton, J. Mar. Res., Res.,34, 313-326, 313-326, 1976. 1976. Micropaleontology, 8,8,2219-254, 19-254, 1962. Micropaleontology, 1962. Wiebe, P. P. H., H., A. M. Bradley, Wiebe, A. W. W. Morton, Morton, A. A.M. Bradley, R. R. H. H. Backus, Backus,J. J. E. E. Pond, S., and G. Pickard, Introductory Dynamical Oceanography, 2nd Craddock, V. Pond, S.,andG. Pickard, Introductory Dynamical Oceanography, 2nd Craddock, V. Barber, Barber,T. T. J. J.Cowles, Cowles,and andG. G.R. R.Filerl, Filed,New Newdevelopdeveloped., Tarrytown, N.Y., ed.,329 329pp., pp.,Pergamon, Pergamon, Tarrytown, N.Y.,1983. 1983. ments in for zooplankton and ments in the theMOCNESS, MOCNESS,an anapparatus apparatus forsampling sampling zooplankton and Ravelo, A. C., R. G. Fairbanks, and S. G. H. Philander, Reconstructing micronekton, 13-323, 1985. micronekton,Mar. Mar. Biol. Biol.Berlin, Berlin,87, 87,3313-323, 1985. Ravelo, A. C., R. G. Fairbanks, and S. G. H. Philander, Reconstructing tropical atlantic hydrography using planktonic foraminifera Zaneveld, J. J. R., and an tropical atlantic hydrography using planktonic foraminifera and an Zaneveld, R., R. R. W. W. Spinrad, Spinrad,and andR. R.Bartz, Bartz,Optical Opticalproperties propertiesof of ocean model, Paleoceanography, 5, 1990. turbidity Ocean ocean model,Paleoceanography, 5,409-431, 409-431, 1990. turbiditystandards, standards, OceanOptics OpticsVI, VI, Proc. Proc.SPIE SPIEmt. Int.Soc. Soc.Opt. Opt.Eng., Eng., Sancetta, C., M. M. Lyle, Huesser, R. and J. P. Bradbury, 208, 208, 159-168, 159-168, 1979. 1979. Bradbury, LateSancetta, C., Lyle,L. L. Huesser, R.Zahn, Zahn, and J.P. LateGlacial to to Holocene changes in the upwelling and Glacial Holocene changes in thewinds, winds, upwelling andseasonal seasonal production of California Current System, Quat. Res., 38, production ofthe thenorthern northern California Current System, Ouat. Res., 38, 359-370, 1992. 1992. J. D. A. C. C. Mix, Mix, and J. D. Ortiz, Ortiz, A. andR. R.W. W. Collier, Collier,College Collegeof of Oceanic Oceanicand and Sautter, L., and R. Thunell, Seasonal succession of the planktonic Atmospheric Sciences, Sciences, Oregon Oregon State University, Corvallis, OR Sautter, L., and R. Thunell, Seasonal succession of the planktonic Atmospheric State University, Corvallis, OR 97331. 97331. foraminifera: Results from a four year rime-series sediment trap (e-mail: jortiz jortiz@oce.orst.edu; amix@oce.orst.edu; rcollier@oce.orst.edu) foraminifera: Results from a fouryear time-series sediment trap (e-mail: @oce.orst.edu; amix@oce.orst.edu; rcollier@oce.orst.edu) experiment in in the the northern northern Pacific, Pacific, J. J. Foraminiferal Foran,iniferal Res., 19, experiment Res., 19,253253267, 1989. Sautter, L., R. Thunell, Planktonic foraminiferal response to Sautter, L.,and and R. Thunell, Planktonic foraminiferal response to upwelling and seasonal hydrographic conditions: Sediment trap upwelling and seasonal hydrographic conditions: Sediment trap (Received September 19, July results from San Pedro basin, southern California, J. (Received September 19,1994; 1994;revised revised July5, 5,1995; 1995; results fromSanPedro basin, southern California, J.Foraminferal Foraminiferal Res., 21, 347-363, 1991. accepted July 5, 1995.) Res.,21, 34%363, 1991. acceptedJuly 5, 1995.)

Environmental control of living symbiotic and asymbiotic

Related documents

Products

Support

Environmental control of living symbiotic and asymbiotic

Related documents

Add this document to collection(s)

Add this document to saved

Suggest us how to improve StudyLib