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Rcv. oras. occanogr., 44(1):W-80, 19C)6

Microalgae 01 the continental shelf off Paraná State, southeastern Brazil:a review of studies

Frederico P. BRANDINI1 & Luciano F. FERNANDES2

1 Universidade Federal do Paraná, Centro de Estudos do Mar(Av. Beira Mar, s/n, 83255-000, Pontal do Sul, Pontal do Paraná, PR, Brasil)

2 Universidade Federal do Paraná, Setor de Ciências BiológicasDepartamento de Botânica, Centro Politécnico

(Caixa Postal 1903, 81531-970, Curitiba, Paraná, PR, Brasil)

. Abstract: The paper reviews the taxonomic and ecological works on marinemicroalgae (phytoplankton and microphytobenthos) off Paraná State, SoutheasternBrazil. Various studies refer to the taxonomy of benthic diatoms. Few descriptivefield works in coastal and adjacent shelfwaters contribute to the understandingof phytoplankton dynamics in coastal and shc1f areas. Patterns of geographicdistribution and changes in the phytoplankton community are described in relationto water masses and seasonal variations in the hydrographic regime. Duringsummertime, when warm oligotrophic waters predominate over the shc1f, thephytoplankton is nummerically dominated by small phytoflagellates, dinoflagellates,coccolithophorids and filaments of cyanobacteria. During wintertime, theabundance of micro-size diatoms over the she1fincrease due to the dominance ofcold nutrient-rich waters from the South Atlantic Central Water and wind-drivenvertical circulation. The phytoplankton community off Paraná State may beclassified into two categorics: the diatom-dominated coastal assemblage, mainlycontrolled by nutrient inputs from land drainage and ressuspension of bottomsediments, and the flageIlate-dominated shelf assemblage, more affected by theseasonality of local hydrographic regime.

. Resumo: O trabalho é uma revisão dos estudos taxonômicos e ecológicos dasmicroalgas (planctônicas e bentônicas) realizados no Estado do Paraná. A maioriados trabalhos sobre microfitobentos são de caráter taxonômico e, basicamente,referem-se à diatomáceas bênticas sobre macroalgas ou fundos lodosos econsolidados. Poucos trabalhos contribuiram para o estudo da dinâmicaespaço-temporal do fitoplâncton em áreas costeiras e de plataforma. São descritosos padrões de distribuição geográfica e as mudanças sazonais na comunidadefitoplanctônica em relação ao regime hidrográfico. No verão, quando águasquentes oligotróficas predominam na superfície, o fitoplâncton é numericamentedominado por nanoflagelados, dinoflagelados, cocolitoforídeos e cianobactériasfilamentosas. No inverno, as diatomáceas do microplâncton são em geral maisabundantes devido ao domínio de águas frias ricas em nutrientes. A comunidadefitoplanctônica da plataforma paranaense pode ser classificada em duas categorias:as associações costeira, sujeitas ao aporte de nutrientes pela drenagem continentale ressuspensão do sedimento, e a comunidade da plataforma afastada da costa, maisafetada pela sasonalidade do regime hidrográfico.

. Descriptors:Phytoplankton, Microphytobenthos,Taxonomy,Paraná State,Southeastern Brazil.

. Descritores:Fitoplâncton, Microfitobentos, Taxonomia, Estado do Paraná, Regiãosueste do Brasil.

70 Rev. bras. oceanogr., 44(1),1996

The Paraná littoral and plane region

The coastline of Paraná State, Southeast Brazil, runsfrom north to south bctwecn 25°12' -58'S Lat. along the

48°W Long. The continental shelf is extensive, with theisobath of 200 meters starting at approximatcly 175-190kmfrom the coastline. The littoral plane is not extent(10-20 km, with a maximum of 50 km in the Bay ofParanaguá) due to its proximity to the Serra do Mar.Consequently, the continental drainage is small, despitethe high precipitation (Bigarella, 1978), beingconcentrated in the inner bays of Paranaguá andGuaratuba. Heavy rains gencrally occur in summcr, as theresult of the constant stationing of polar fronts. In winter,less solar radiation, couplcd with the frequent passage of apolar anticyclone, makes the atmosphere drier and colder(Bigarella, 1978;Nimer, 1990)

The littoral between the bays of Paranaguá andGuaratuba is basically formed by a single extensive sandybeach exposed to the open sea, but the inner bays arebordered by extensive mangrove forests producing organicdetritus which is then transported by continental drainageand tidal currents towards the adjacent sea.

In the coastal region, the pattems of circulation on aregional scale are govemed by tides, wind and precipitationpatterns, draining the continent and carrying nutrients tothe coastal waters. Meanwhile, it is impossible todisassociate the characteristics of the hydrographic systemof the mid-shelf and the contiguous oceanic arcas fromthe general oceanographic structure of the southem Brazil(see Emilsson, 1961) which is directly related to theseasonal dynamics of the Brazil Current and the proxirnityof the Subtropical Convergence.

Studies in shelf and oceanic areas

The first oceanographic studies of the SouthwestAtlantic Ocean were ma de in the 1920's, during theexpeditions of the RN "Meteor" (Wüst, 1932; Defant,1936)and RN "Discovery"(Deacon, 1933,1937).However,the data collected at that time permittcd only a general andsimplified vision of the water masses.

Oceanographic studies in shelf arcas off Brazil weresystematically begun only in the 1950's. Phytoplanktonsamples were collected in mid- and outcr-shclf off ParanáState within a larger sampling program covering large partsof the south and southeast regions of Brazil. Earlierinformations on the physical-chemical characteristics of thewater masses in this region were described by Emilsson(1959,1961), based on data obtained during expeditions ofthe Brazilian Navy, with the corvettes "Iguatemi","Solimões"and the RN "Almirante Saldanha".

The latitudinal displacements of the SubtropicalConvergence, as the consequence of seasonal changes inthe wind systems of the Southwest Atlantic, changecompletely the hydrography of the shelf. Northeasterlywinds predominate during summer transporting TropicalWater (TW) from the Brazil Current (Emilsson, 1%1;Miranda, 1982;Matsuura, 1986;Brandini 199Oa).In winter,the southeasterly winds predominate, allowing thepenetration and mixing of subantarctic waters along withshelf and tropical waters (Fig. 1). Strong gradients oftemperature were observed along the shelf off SantaCatarina and Paraná, indicating the penetration of coldwaters. The isohaline of 36.0 psu, which represents thelimits ofTropical Water from the Brazil Current, extendedalong the slope during winter 1982, but it was elose to thecoast during summer 1984. In the vertical plane, the mainhydrographic features affecting the physical-chernicalconditions of shelf waters and the planktonic communityare (i) the formation of a well-dcfined seasonal thermoclinein summer and (ü) the shelf-break upwellings (Fig 2).

Between May 1976and June 1978work was done on theshelfbetween Cabo Frio-RJ (22°S and 42OW)and Cabo deSanta Marta Grande - RS (28°S and 48°W) during variouscruises of the RN "Prof. W. Besnard" (IOUSP), supportedby the Financier of Studies and Projects (FINEP). Thesampling of phytoplankton material, restricted to thesurface layer, included some stations off Paraná State.Aidar-Aragão et ai. (1980) published the first informationon the potential primary production of phytoplankton in thesurface, using the C-14 technique of Steemann-Nielsen(1952) and analysis of chlorophyll-a. During the sameperiod, Vieira & Teixeira (1981) completed thephysiological studies by means of excretion of dissolvedorganic material, and Soares (1983) described thetaxonomic structure and the distribution of the dominant

phytoplankton groups. The diatoms were abundant inneritic stations and the more significant species wereSke/etonema costatum, Leptocy/indrus danicus, L.mediterranells, Tha/assionema nitzschioides, Bacteriastrom

sp, 17!a/assiosira sp, Nitzschia spp and Chaetoceros spp,followed by Proboscia ( = Rhizoso/enia) a/ata, R. ca/caravis,R.robusta, Gllinardia f/accida, Guinardia striata( = Rhizoso/enia sto/teifothii), Hemiau/lls membranaceusand H. hallckii. C/imacodillm frallenfe/dianllm wassignificant in one of the three cruises, but associated withtropical dinoflagellates, dominated by typical neritic andoceanic representatives of the Brazil Current (e.g.,Amphiso/enia bidentata, Ceratillm hexacanthum, C. fusus,C. karstenii, C. massi/iensis, C. trichoceros, C. tdpos,Dip/opsa/is sp., Goniodoma po/iedricum, Protoperidiniumgrande and Pyrophacus steinü).

29°

29'

2r' I. Temperature at 5 mFebruary-March 1984

\. Temperature at 5 mJuly-August 1982

Fig. 1. Geographical distributions of temperature and salinity at 5 meters in Southeast Brazil, inJuly-August 1982 and February-March 1984 (modifiedfrom Brandini,1990).

BRANDINT & FERNANDES: Microalgae of the continental shelf off Paraná State 71

lon W 48. 47. 46. 45' 44. long. W 4a' 47' 46' 45' '.4'

, I ',24. "'""V

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. Salinity at 5 m36 ;.February-March 1984

36 . :. . .

'" July-August 1982I

+"cI-'

long W "8. 4?O 46. 45. 44° long.W 48' 47° 46. 45° 44°

240L . .... ..24'

72 Rev. bras. oceanogr., 44(1), 1996

Summer

24

22

20

16~

16

14

Winter

51

Fig. 2. Crosss-shelf distributions of temperatrure, salinity and nutrients off the Bay of Paranaguá; ParanáState, inthe 5ummer1982and inthe winter1983 (modifiedfromBrandini,1990a).

.0

f

.'-''':.;.:.- .....-'36,5

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200 .---- 35.5--E..........300

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300-j N03 \1001 l N0312.0

100

200

30J Toe

BRANDINI & FERNANDES: Microalgae of the continental shelf off Paraná State

At the end af the 1970's, the expedition "Canversut I"af the Brazilian Navy studied the hydrographical andbialogical aspects of the Subtropica.l Convergence,comprising the shelves of southern Brazil, Uruguay andpart of Argentina (DHN, 1980;Hubold, 1980).

From July 1982 until November 1985, anotherimportant seasonal sampling program was carried out bythe Brazilian Navy; the expeditions "SUESTE"(DHN, 1985; 1986a; 1986b, 1987) were conducted by theRN "Almirante Saldanha", covering a sampling networkof 90 oceanographic stations in each cruise, organized in12 cross-shelf transects off São Paulo, Paraná andSanta Catarina states. During theses expeditions, theshelf-break upwellingof SACW (Mesquita etal., 1983) ..

were detected in different seasons of the year offthe Bay of Paranaguá (Brandini et ai., 1989;Brandini, 1990a). These upwellings fertilize the base ofthe euphotic zone of the oceanic oligotrophic regions,stimulating the production of phytoplankton and thedevelopment of the pelagic ecosystem.

The colIection of phytoplankton material produceddata on celI density, floristic composition, photosyntheticpigments, rates of photosynthesis, estimates of potentialand in situ primary production in the euphotic zone andphotosynthetic characteristics of phytoplankton. Thedynamics of the phytoplankton community wereassociated with the oceanographic structure of theregion (Brandini, 1988a;Brandini, 1988b;Brandin~ 199Oa;Brandini et ai., 1989). The expedition "Monitor IV"collected hydrographical and biological data duringsummer (DHN, 1989),off Rio de Janeiro, São Paulo andParaná States.

Brandini & Moraes (1986) and Brandini (1988b)described the patterns of geographic distribution ofdominant phytoplankton groups in relation to the watermasses. During winter, Brandini (1988b) showed thedifference between cold water assemblages of sub-antarcticorigin, dominated by diatoms, and the assemblages of theBrazil Current, dominated by cyanophyceans,coccolithophorids and dinoflagellates. A similar pattern inthe neritic region was observed by Fernandes (1992) duringa seasonal study in front of the Bay of Paranaguá.From these, it was deduced that seasonal changes inthe structure of the phytoplankton community off ParanáStafe, occurred as a result of changes in theoceanographic structure. With the predominance ofTropical Water in summer, prevails a regenerative typeproduction system (sensll Dugdale & Goering, 1967) inwhich nanoplanktonic phytoflagellates, coccolithophoridsand cyanophyceans are the dominant phytoplankters.During winter, the intrusion of cold and nutrient-rich

73

waters increases the relative contribution of

microplanktonic diatoms.ln the vertical plane, the structure of the

phytoplanktonic community in the euphotic zonechanges in the region of the continental slope due toshelf-break upwellings. The occurrence of such upwellingshas important implications on the primary production ofoceanic areas, where the contribution of nutrients to theeuphotic zone from deeper layers is generally limited bythe presence of the permanent thermocline. The upwardmotions of the SACW fertilizes the base of the euphoticzone, increasing the production of phytopIankton and,consequently, that of the higher trophic levels(Brandini et a/..,..1988).These authors observed that themost representative modifications in the phytoplanktoncommunity were the increases in concentr.ations of micro-and nanoplanktonic coccolithophorids (Umbilicosphaerasibogae, Emiliania huxleyii, Gephirocapsa oceanica),diatoms (Plellrosigmasp, Coscinosira sp, Eucampia sp,),cyanophyceans (Oscillatoria erythraea) and dinoflagellates.The cyanophycean Trichodesmium (=; Oscillatoria)occurred in the coastal areas and in the sub-surface

in oceanic areas (Fig. 3a). Diatoms predominate in coastalregions, mainly .. in sub-surface layers due tosedimentation (Fig. 3b). Dinoflagellates andcoccolithophorids were more abundant in offshore regions(Fig. 3c and d), associated with the shelf-break upwellingsof the South Atlantic Central Water (Fig. 3e).

The photosynthetic rates in the euphotic zone andits seasonal variation were attributed to changes in thehydrography and in the taxonomic structure ofphytoplankton (Brandini, 1988a, 1988b, 199Oa).Brandini(1990b) estimated the primary production rates in thecoast (0.04 -0.32 gC/m2/day) in the intermediate shelf (0.13

- 0.45 gC/m2/day), and in the oceanic (0.10 - 0.45gC/m2/day) regions, and compared the potential organicproduction in the southeastern Brazil with dataobtained in other seas. In the same work, thephotosynthetic characteristics of the phytoplanktoniccommunity were described during the stratificationperiods of summer, through the relations betweenthe rates of photosynthesis and light, and by observingphoto- adaptations occurring in the development ofcommunities physiologically distinct throughout theeuphotic zone.

Studies in the coastal region

Since the end of the 1950's, many works havecontributed to the knowledge of diatoms in the coastalregions between the Bays of Paranaguá and Guaratuba

74 Rev. bras. oceanogr., 44(1), 1996

D i s t a n c e (Km)Fig. 3. Cross-shelf distribution of the phytoplankton community and the vertical oceanographic

structure offthe Bay of Paranaguá, Param 8tate, in November of 1987. CW: coastal waters; TW:TropicalWater; SACW:South AtlanticCentral Water. Multiply numerical laOOlsof isolines in (A),(B)and (C) by 10, and in (D)by 100 (modifiedfrom Brandini,1989).

Stations

2 3 4 5 6O

50

100J :00 filame. tsl I

Cianophyceans """'\ -- A150

O

50I ''''''I'. .

1001->100 ceU,/1 ,<.1.

. .150 Diatoms> 20m

B

O

50

Ê IOOJ>50 cells/

-----. .<I

---.J:: IDinoflagellates> 20m

"- c..... 150a.Q)C

O

50

10</00. . <I

100-1.> 600 cells/l. .

ICoccolithophorids >20m " D150

O'CW \ \. . . ISW . . I . TW

50-1 ------ "'_------___ I1" . 4 .\

... ... ...

100-i CW-,

.- --...........Oceanographic Structure

.... ... .

150\ E

I I I , I

O 100 200 300

BRANDINI & FERNANDES: Microalgae of the continental shelf off Paraná State 75

(Moreira F1lho & Fernandes, 1958; Moreira-Filho, 1959;Moreira-Filho, 1961; Moreira-Filho & Kutner, 1962;Moreira Filho & Mômoli, 1962; Moreira-Filho, 1968;Aidar, 1970; Moreira-Filho et a/., 1975j Moreira-Filho &Valente-Moreira, 1979; Moreira-Filho &Valente-Moreira, 1980; Valente-Moreira &Moreira-Filho, 1981; Oliveira, 1983; Oliveira, 1984;Moreira-Filho & Valente-Moreira, 1984;Valente-Moreiraet a/., 1985; Valente-Moreira, 1987; Cunha, 1989;Moreira-Filho et a/., 1990).

The main objectives of these works were to performfloristic and taxonomic surveys of diatoms in the Paranácoast. Moreira-Filho (1959) began by publishing the flJ'stinformation about epiphytic microalgae on Sargassumcymosum collected from the beach of Caiobá. In this work,31 species of diatoms and two species of silicoflagellateswere identified. Moreira-Filho & Kutner (1962) studied thecommunity of diatoms in sediments of Alexandra mangrove(Bay of Paranaguá), identifying various planktonic species.A series of works folIowed, surveying the floristic ofepiphytic microalgae on macroalgae (Moreira-Filho et al.,1977; Moreira-Filho & Valente-Moreira, 1980;Valente-Moreira et a/., 1980; Moreira-Filho &Valente-Moreira, 1981; Paula, 1990), in shelI deposits(Hurt & Blasi, 1960;Moreira- Filho & Mômoli, 1962) andin the digestive tract of invertebrates (Moreira-Filho, 1960;Valente-Moreira et a/., 1994) and in flsh (Valente-Moreiraet al., op.cit.). The quantitatively most signiflcant specieswere Achnanthes brevipes, A./ongipes, Biddu/phiapu/che//a, Cocconeis heteroidea, C. pe//ucida, C. scute//um,Grammatophora marina, G. oceanica, Licmophoraabreviata, Mastog/oia binotata, M. crucico/a, M. sp/endida,Para/ia su/cata and Synedra tabu/ata. Cunha (1989)described the temporaly variations in the structure ofdiatom community in the Galheta Channel, Bay ofParanaguá, on a seasonal scale. Moreira-Filho et a/., (1990)performed a survey of diatoms identilled in the whole Stateof Paraná, noting the occurrence of 636 species distributedin 115 classes, surpassing alI the other groups ofphytoplankton. In relation to benthic microalgae in theneritic are a, it is difflcult to establish any pattem ofdominance and seasonal variation, since few works havebeen performed in Paraná. Meanwhile, recentinvestigations of Brandini et alo (in preparation) andPellizzaril (1994) on the seasonal dynamics of theperiphitic community in artificial substrates in the Bay ofParanaguá supply the first indications that they could

(1) Pellizzari, F. M. 1994. Ecologia do perifíton da Baía de Paranaguá.

Paraná. In: EVENTO DE INICIAÇÃO CIENfÍFICA da Universidade

Federal do Paraná (EVINCI - 94/resumos). Anais. Curitiba.

UFI'RlPRPPG. p. 121.

assume a major role in the local ecosystem, by-passingphytoplanktonic biomass in certain sections of the bay,besides becoming available for the planktonic community(e.g. Cy/indrotheca c/osterium and P/eurosigma sp). Theseauthors noted the dominance of the diatom Navicu/a spduring rainy periods of summer, foUowed by Me/osira sp,Coscinodiscus sp, Thalassiosira sp, Cy/indrothecac/osterium, Licmophora sp, Nitzsehia aff. /ongissima andP/eurosigma sp (Fig.4 ), suggesting that the production ofthe community is controlIed mainly by temperature.Concentrations of chlorophylI-a were in accordance withdiatom densities. Femandes (1992) studied the annualvariation of phytoplankton on the continental shelf off theBay of Paranaguá and recorded the presence of variousbenthic species in ~e water column, possibly due toresuspension by tidal and wind-induced turbulence. Themost signiflcants were Cyc/ote//a sty/orum, Paralia su/eata(main/y), Anorthoneis eurystoma, De/phineis surire//a,Rhaphoneis amphieeros and Thalassiosira spp (3 species ofvalvar diameter inferior to 20 p.m, coated with grains ofsediment). There also occurred the epiphytes Biddulphiasp, Liemophora ehrenbergii, L. flabe//ata and Podocystisadriatica, and the planktonic genus Chaetoceros, whichforms benthic resting-spores.

Since the 1980's works have been initiated in the Bay ofParanaguá with the purpose of studying the patterns ofspatial and temporal variation of phytoplankton in relationto hydrographic and climatological parameters. Thepreliminary works concentrated on the analysis of dailyvariations of phytoplankton concentration andhydrographic parameters in relation to tides (Oliveira2 eta/., 1983; Freitas, 1989). Brandini (1985a) described thechanges in the phytoplanktonic community structure in aflxed station, in relation to pluviosity. Brandini (1985b) andBrandiniet ai. (1988) studied the seasonal variation and thespatial distribution af chlorophyll-a and basic hydrographicparameters within the Bay of Paranaguá.

Recently, besides the local climatology, the annualvariation studies described the effects of the hydrographicregime in the adjacent shelf on the seasonal dynamics ofphytoplankton in the bay (Rezende & Brandini3, 1988;Rebello & Brandini, 1990; Femandes, 1992; Brandini &

(2) Oliveira, c.; Dortas, E. M.; Sert, M. A.; Soares, F. S. & Brandini, F.

P. 1983. Variação nictemeral do fitoplâncton da Baía de Paranaguá. In:

SIMPÓSIO DE ESTAGIÁROS, 3. Resumos. Londrina, 1983. Fund.

Univ. f';stadual de Londrina, p. 5.

(3) Rezende, K R. V. & Brandini, F. P. 1988. Variação temporal do

fitoplâncton em um ponto fIXoda praia de Pontal do Sul (Paraná). In:

ENCONTRO BRASILEIRO DE PlÂNCfON, 3. Resumos.

Universidade Federal do Paraná. Centro de Biologia Marinha, 1988.p.45.

76 Rev. bras. oceanogr., 44(1),1996

400000 0.04

NE()

1::Q)..cE:JC

ã)()

300000~...,I II II II III. I

fi II I II I II I'. I .

I' II "I )IIII

200000

100000

o

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N

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- - - - - - - chlorophyll-aCIJ

...!.>..s::c.o~o1:()

0.02

II Ifi II I II "I .

f

0.01

0.00

1993- 1994

Fig. 4. Seasonal variation in the abundance of benthic diatoms in external sections of the Bay of Paranaguá,Paraná State. Abundance is expressed interms of accumulation of cellsand chlorophyll-aper area, after 7 daysof submersion of artificialsubstrates (frem Brandiniet aI. 1985).

Thamm, 1994).As a basis for these works, it was noted thatthe structure of the microalgae community is dominated bydiatoms (see below) in summer, when occurs an enrichmentof the water column with inorganic nutrients due to higherprecipitation (Brandini, 1985a) (Fig. 5). Meanwhile, theturbulence generated by the stronger winds of winter,resuspending nutrients from the bottom, could aIsoincrease the density of diatoms in the coastaI areas adjacentto the Bay of Paranaguá, with a significant contribution ofbenthic species such as Paralia !nUcata,Delphineis surirellaand Cyclotella stylomm (Fcrnandes, 1992). During winter,the presence of cold water species, such as Bacteriastmmhyalinum, B. varians,Eucampia comuta, Lauderia annulata,Detonula pumila (= Schroederella delicatula),Thalassiosira punctigera, Thalassiosira tumida,Thalassiothrix mediterranea, Dinophysis rotundata,Oxyphysis oxytoxoides and Protoperidinium oceanicum,indicate the influence of sub-antarctic waters. Fernandes

(1992) also observcd a bloom of Phaeocystis sp immediateIyafter the diatoms bIoom, similar to what occurs in coastalregions of the North Sca (Lancc1ot et ai., 1991).

Phytoplankton assemblages in the shelf offParaná State

The accomplished works in the coastaI area and in theintermediate she1f allow for the distinguishing of thefollowing phytoplanktonic associations:

a) The coastal assemblage, dominated by diatoms,mainly Pseudonitzschia seriata, Pseudonitzscllia

delicatissima, Leptocylindrus danicus, Leptocylindmsminimus, Chaetoceros affinis, Chaetoceros debilis,Ozaetoceros didymus, Chaetoceros curvisetum, Chaetoceroscompressum, Guinardia (=Rhizosolenia) striata, G.(= Rhizosolenia) delicatllla, Dactyliosolen fragilissimus(= Rhizosolenia fragilissima), Cerataulina bergonii,Hemiaulus hauckii, H. sinensis, Asterionellopsis glacialis,Thalassionema nitzschioides and Skeletonema costatum.

Dinoflagellates are less abundant, and numericallydominated by the genus Gymnodinium, Amphidinium,Cochlodinium, Gyrodinium and Torodinium robustum,followed by Ceratium furca, Cera ti um fusus,Protoperidinium depressum, Protoperidinium spp,Prorocentrum balticum, Prorocentrum compres um,Prorocentrum micans, Prorocentrum minimum,Podolampas palmipes, Podolampas spinifera, DinophysisaCllminata,Dinophysis caudata, Dinophysis rotundata andNoctiluca miliaris. Occasionally, some of these species maycause bIooms. The predominant siIicoflageUate isDictyocha fibula. The coccoIithophorids are normaUyrepresentcd by Gephyrocapsa oceanica and Emilianiahu.rleyii, followed by Anoplosolenia sp, Antosphaera sp,Calciopapus sp, Calciosolenia sp, Corisphaera sp,Crycosphaera sp and Helichosphaera carterae, whichoccurred in low concentrations.

b) The shelf assemblage, greatly influenced byoligotrophic waters of the Brazil Current, wheredinoflagellates are dominated by Ceratium azoricllm, C.gravidum, C. massiliensis, C. trichoceros, C. vllltur,Protoperidinium elegans, Amphisolenia bidentata,

BRANDINJ & FERNANDES: Microalgae of the continental shelf off Paraná State 77

120mm

100% o Phytoflagellates

O Diatoms. Dinoflagellates

50

A s o N o

(8)

J F M A M J

Fig. 5. Precipitation.air temperature (A)and reiative abundance of the phytoplanktonic groups (B)atthe surface of the Bay of Paranaguá. The figureshows the dornain ofdiatoms in the warmer andrainierperiods. and of 1Iagellateforms during winterwith less continental drainage (modifiedfromBrandini,1985a).

Goniodoma poliedricllm, PyrophaCllssteinii, Gonyalllaxspp and Ceratillm spp (see review of Balech, 1988). Thecoccolithophorids are essentially represented byGephyrocapsa oceanica, Emiliania lmxleyii andUmbilicosphaera sibogae. Secondly, occurred the diatomsClimacodillm frallenfeldianum, CoscinodiscllS gigas, C.centralis, Proboscia (=Rhizosolenia) alata, Hemialllllsmembranacells and Fragilariopsis (= Pselldoellnotia)doliolus. The species Gossleriela tropica and Planktoniellasol are rare but always present in net samples. Thecyanobaeteria Oscillatoria erythraea is common, formingsometimes extensive patches. During summer, thesespecies also occur in inner-shelf regions, being mixed withthe coastal assemblage.

Final remarks

The climatology and the seasonal variation of theoceanographic structure of the region, renders even morecomplex the space- temporal dynamics of the microalgaecommunities. Although the studies on the composition andbiogeography of microalgal species are more or lessadvanced, there is missing physioeeological data tocomplement the interpretation of patterns of distribution

and development in relation to environmental parameters.From that point on, it will be possible to evalulatequantitatively the role of micfoalgae in the marineecosystem on the Paraná shelf, and to trace ecologicalmodels which will allow to foresee regularities andvariations in the community over a longer periQd of time.

References

Aidar, E. 1970.O fitoplâncton da Baía de Paranaguá.Relatórioapresentado à SUDEPE. (não publicado)

Aidar.Aragão, E.; Tcixeira, C. & Vieira, A. 1980.Produção primária e concentração de clorofila-a nacosta brasileira (Lat. 22"31'S - Long. 41052'W a Lat.28°43'S . Long 47°57'W). Bolm Inst. oceanogr., SPaulo, 29(2):9-14.

Balech, E. 1988. Los dinoflagctados deI AtlânticoSudoccidental. Publ. Esp. Inst. Esp. Oceanogr.,Madrid, 1:310p.

78 Rev. bras. oceanogr., 44(1),1996

Bigarella,J. J. 1978.A Serra do Mar c a porção orientaldo Estado do Paraná. Um problema de segurançaambiental e nacional. (Contribuições à geografia,geologiae ecologia regional). Curitiba, SEP/ADEA.249p.

Brandini, F. P. 1985a. Seasonal succession of thephytoplanktonin the Bay of Paranaguá (Paraná State- Brazil).Revtabras. Biol.'45(4):687-694.

Brandini, F. P. 1985b.Ecological studies in the Bay ofParanaguá. I. Horizontal distributions and seasonaldynamicsof the phytoplankton.BolmInst. oceanogr.,S Paulo,33(2):139-147.

Brandini, F. P. 1988a. Hydrography, phytoplanktonbiomass and photosynthesis in shelf and oceanic watersoff Southeastern Brazil during autumn (May/June1983). Bolm Inst. oceanogr., S Paulo, 36(1/2):63-72.

Brandini, F. P. 1988b. Composição e distribuição dofitoplâncton na Região Sueste e suas relações com asmassas d'água (Operação Sueste I - Invemo/1982).Ciênc. Cult., S Paulo, 40(4):33+341.

Brandini, F. P. 199Oa.Hydrography and characteristics ofthe phytoplankton in shelf and oceanic waters offSoutheastern Bra7il during winter (Ju!y/August 1982)and summer (February/March 1984). Hydrobiology,1%(2): 111~148.

Brandini, F. P. 199Ob.Produção primária e característicasfotossintéticas. do fitoplâncton na Região Sueste doBrasil. Bolm Inst. oceanogr., S Paulo, 38(2):147-159.

Brandini, F. P. & Moraes, C. L. B. 1986. Composição edistribuição do fitoplâncton em áreas costeiras eoceânicas da Região Sueste do Brasil. Nerítica, Pontaldo Sul, 1(3):9-21.

Brandini, F. P. & Thamm, C. A. C. 1994.Variações diáriase sazonais do fitoplâncton e parâmetros ambientais naBaía de Paranaguá. Nerítica, Pontal do Sul,8(1/2):55-72.

Brandini, F. P.; Moraes, C. L. B. & Thamm, C. A. C. 1989.Shelf-break upwelling, subsurface maxima ofchlorophyll and nitrite, and vertical distribution of asubtropical nano-microplankton community offsoutheastern Brazil. In: Brandini, F. P. ed.Memóriasdo 111 Encontro Brasileiro de Plâncton. Caiobá,Paraná. p.47-55.

Brandini, F. P.; Thamm, C. A. H. & Ventura, I. 1988.Ecological studies in the Bay of Paranaguá. m.Seasonal and spatial variations of nutrients andchlorophyll-a.Nerítica, Pontal do Su~ 3(1):1-30.

Cunha, J. A. 1989. Relações entre composição e variaçãoqualitativa relativa genérico/específica, dasdiatomáceas (Chrysophyta-Bacillariophyceae), noCanal da Galheta, município de Paranaguá, Estado doParaná, Brasil. Dissertação de mestrado.Universidade Federal do Paraná, Depto de Botânica.598p.

Deacon, G. E. R. 1933.A general account ofthe hydrologyof the Southern. Atlantic Oceano Discovery Rep.7:171-238.

Deacon, G. E. R. 1937. The hydrology of the SouthernOceano Discovery Rep., 15:1-124.

Defant, A. 1936. Die troposphãre des AtlantischenOzeans. Wiss. Ergebn. dto atlant. Exped. "Meteor",6(1):289-411.

D.H.N. 1980. 718 Comissão Oceanográfica/OperaçãoConversut I, Costa Sul - Oceanografia Física eBiológica. N/Oc."Almirante Saldanha". Diretoria deHidrografia e Navegação da Marinha do Brasil.21p.

D.H.N. 1985. 1018 Comissão Oceanográfica/OperaçãoSueste 11,Costa Sueste -Oceanografia Física, Químicae Biológica. N/Oc."Almirante Saldanha". Diretoria deHidrografia e Navegação da Marinha do Brasil.31p.

D.H.N. 1986a. 988 Comissão Oceanográfica/OperaçãoSuesí:eI, Costa Sueste -Oceanografia Física, Químicae Biológica. N/Oc."Almirante Saldanha". Diretoria deHidrografia e Navegação da Marinha do Brasil.33p.

D.H.N. 1986b. 2008 Comissão Oceanográfica/OperaçãoSueste 111, Costa Sueste - Oceanografia Física,Química e Biológica. N/Oc."Almirante Saldanha".31p.

D.H.N. 1987. 988 Comissão Oceanográfica/OperaçãoSueste IV, Costa Sueste - Oceanografia Física,Química e Biológica. N/Oc. "Almirante Saldanha".Diretoria de Hidrografia e Navegação da Marinhado Brasil. 31p.

BRANDTNT& FERNANDES: Microalgae of the continental shelf off Paraná State 79

D.H.N. 1989. 1411 Comissão Oceanográfica/OperaçãoMonitor IV. Costa Sul: Oceanografia Química, Físicae Biológica. N/Oc. "Almirante Saldanha". Diretoriade Hidrografia e Navegação da Marinha do BrasiL26p.

Dugdale, R. C. & Goering, J. J. 1%7. Uptake of new andregenerated forms of nitrogen in primary productivity.Limnol. Oceanogr., 12:196-206.

Emilsson, I. 1959.Alguns aspectos físicos e químicos daságuas marinhas brasileiras. Ciênc. Cult., S Paulo,11(2):44-54.

Emilsson, I. 1961 The shelf and coas tal waters offSouthern Brazil. Bolm Inst. oceanogr., S Paulo,11(2):101-112.

Fernandes, L. F. 1992. Variação sazonal do fitoplânctone parâmetros hidrográficos em uma estação costeirade Paranaguá - Paraná. Dissertação de mestrado.Universidade Federal do Paraná, Depto de Botânica.93p.

Freitas, C. A. F. 1989. Parâmetros hidrográficos efitoplâncton. In: Almeida, C. ed. Estudo biológicointegrado da foz da Gamboa do Maciel (Paranaguá-Paraná) durante dois ciclos de maré. UniversidadeFederal do Paraná, Centro de Biologia Marinha,UFPR, p.13-36.

Hubold, G. 1980.Hydrography and plankton off SouthernBrazil and Rio de La Plata, August -November 1977.Atlântica, Rio Grande, 4(1):1-22.

Hurt, R. W. & Blasi, O. 1960. O sambaqui do Macedo.Universidade do Paraná, FFDAA, A52.

Lancelot, c.; Billen, G. & Barth, H. 1991. The dynamicsof Phaeocystis blooms in nutrient enriched coastalzones. Wat. Pollut. Res. Rep., 23:116p.

Matsuura, Y. 1986. Contribuição ao estudo da estruturaoceanográfica da região sudeste entre Cabo Frio (RJ)e Cabo de Santa Marta Grande (Se). Ciênc. Cult., SPaulo, 38:1439-1450.

Mesquita, A. R. de; Leite, J. B. A. & Rizzo, R. 1983.Noteon the shelf break upwelling off the southeast coast ofBrazil (Lat. 26"30'S). Bolm Inst. oceanogr., S Paulo,32(2):193-198.

Miranda, L. B. de 1982. Análise de massas de água daplataforma continental e da região oceânica adjacente:Cabo de São Tomé (RJ) a Tlhade São Sebastião (SP).Dissertação de livre- docência. Universidade de SãoPaulo, Instituto Oceanográfico. 193p.

Moreira-Filho, H. 1959.Diatomáceas do Paraná: I. A floradiatomológica no Sargassum. Bolm Inst. Hist. nat.,curitiba, 2:1-18.

Moreira-Filho, H. 1960. Diatomáceas no trato digestivode Tegula viridllla Gmelin. BoI. Univ. Paraná, 1:1-24.

Moreira-Filho, H. 1961. Diatomáceas da Baía deGuaratuba. BoI. Univ. Paraná, 3:1-33.

Moreira-Filho, H. 1968. Margaritllm (Podosira) tenebro(Leudger- Fortmorel) nov.genlts et nov.comb.(Chrysophyta- Bacilllariophyceae). BoI. Univ. Fed.Paraná, 20:1-2.

Moreira-Filho, H. & Fernandes, E. C. L. 1958. Notapreliminar sobre as Bacillariophyceae da Baía deGuaratuba (Paraná). Trib. farm.,Curitiba, (6):81-87.

Moreira-Filho, H. & Kutner, M. B. B. 1962. Contribuiçãopara o conhecimento das diatomáceas do manguezalde Alexandra. BoI. Univ. Paraná, 4:1-24.

Moreira-Filho, H. & Mômoli, D. M. M. 1962.Sobre apresença de diatomáceas em alguns sambaquis dolitoral paranaense. BoI.Univ.Paraná, 5:1-9.

Moreira-Filho, H.; Valcnte-Moreira, I. M. &Trippia-Cecy, I. I. 1975. Diatomáceas da Baía deParanaguá. Bolm Mus. Bot. Munic., 20:1-23.

Moreira-Filho, H.; Valente-Moreira, I. M. & Matos, A.1977. Diatomáceas epífitas em Codium decorticatum(Wood.) Howe. Trib. farm., Curitiba, 44(6):3-17

Moreira-Filho, H. & Valente-More ira, I. M. 1979.Diatomáceas da enseada da Prainha, Matinhos,Paraná. BolmMus. Bot.Munic.,35:1-12.

Moreira-Filho, H. & Valente-Moreira, I .M. 1980.Diatomáceas epífitas em Uiva fasciata Delile. BolmMus. Bot. Munic., 41:1-10.

80

Morcira-Filho, H. & Valentc-Morcira~I. M. 1981.Avaliação taxonômica e ecoÜSgicadas diatomáceas

(Bacillariophyceae) epífitas em alga~ pluricelularesobtidas nos litorais dos Estados do Par~ná, SantaCatarina e São Paulo. Bolm Mus. Bot. Munic., 47:1-17.

Moreira-Filho, H. & Valente-Moreira, I. M. 1984.Catálogo das diatomáceas (Chrysophyta-BacilIariophyceae) marinhas e estuarinas do Estadodo Paraná, Brasil. Acta Biol. Par., Curitiba,13(1,2,3,4):3-49.

Moreira-Filho, H.; Valente-More ira, I. M.;Souza-Mosimann, R. M. & Cunha, J. A. 1990.Avaliação florística e ecológica das diatomáceas(Chrysophyt a-Bacill ariophyceae) marinhas eestuarinas nos Estados do Paraná, Santa Catarina eRio Grande do Sul. Est. Biologia, PUCIPR, 25:5-48. .

Nimer, E.I990. Clima. In: Geografia do Brasil. Região Sul.Publ. Inst. bras. Geogr. Estatíst., 2:151-187.

Oliveira, R. J. M. 1983. Um levantamento quantitativorelativo do gênero ActinopthychllS Ehrenberg(Chrysophyta- Bacillariophyceae) no estuário do rioItiberê -Paraná, Brasil. Est. Biologia, PUCIPR, 8:1-27.

Oliveira, R. J. M.1984. Contribuição ao conhecimento dasdiatomáceas (Bacillariophyceae) no plânctonestuarino do rio Itiberê, município de Paranaguá,Estado do Paraná, Brasil. Dissertação de mestrado.Universidade Federal do Paraná. 441p.

Paula, J. L. T. M. 1990. Diatomáceas (Chrysophyta-Bacillariophyceae) epífitas em Centrocerasclavlllatllm(CAgardh) Montagne (Rhodophyta-Ceramiales) dailha do Faro~ balneário de Caiobá, Paraná, Brasil.Dissertação de mestrado. Universidade Federal doParaná, Depto de Botânica. 129p.

RebelIo, J. & Brandini, F. P. 1990. Variação temporal deparâmetros hidrográficos e material particulado emsuspensão em dois pontos fixos da Baía de Paranaguá,Paraná Gunho/87 - fevereiro/88).Nerítica,Pontal doSu~ 5(1):95-111.

Soares, L. F. S. 1983. Estudo do fitoplâncton de águascosteiras e oceânicas da região de Cabo Frio - RJ(23°31'S; 41°52'W) até o Cabo de Santa MartaGrande-SC (28°43'S 47°57'W). Dissertação demestrado. Universidade de São Paulo, InstitutoOceanográfico. "118p.

Rev. bras. oceanogr., 44(1), 1996

Stecmann-Niclsen, E. 1952. The use of the radioactivecarbon (C- 14)for measuring organic production in thesea. Rapp. P.-v."Réun. Cons. perm. int. Explor. Mer.,144:38-45.

Valente-Morcira, I. M. 1987. A fI6rula diatomol6gicamadr, ~ e estuarina nos balneários de Canoas e

Ipane.. Estado do Paraná, Brasil. Est. Biologia,PUCIPR,17:26-48.

Valente-Me ~ira, I. M. & Moreira Filho, H. 1981.Diatomàceas de Pontal do Sul, Município deParanaguá, Estado do Paraná, Brasil.(Chrysophyta-Bacillariophyceae). Phycol. Lat. Amer.1:156-185.

Valente-More ira, I. M.; Moreira Filho, H. & Cunha, J. A.1985. Diatomáceas (BacilIariophyceae) marinhas eestuarinas nas praias de Atami e Shangri-Iá, Municípiode Paranaguá, Estado do Paraná, Brasil. Est.Biologia, PUCIPR, 12:1-47.

Valente-Moreira, J. M.; Moreira Filho, H.; Cunha, J. A.& Shirata, M. T. 1987. A fIórula diatomol6gicamarinha e estuarina nos balneários de Canoas e

Ipanema, Estado do Paraná, Brasil. I. Estudoqualitativo e ecológico. Est. Biologia, PUC/PR,17:23-48.

Valente-Moreira, I. M.; Moreira Filho, H.; Cunha, J. A.& Nakamura, I. T. 1994. Diatomáceas(Chrysophyta-Bacillariophyceae) no conteúdoestomacal de peixes e crustáceos do manguezal do rioPerequê, Pontal do Sul, Estado do Paraná, Brasil. Est.Biologia, PUCIPR, 3(39):99-114.

Vieira, A. A. H. & Teixeira, C. 1981.Excreção de matériaorgânica dissolvida por populações fitoplanctônicasda costa leste e sudeste do Brasil. Bolm Inst. oceanogr.,S Paulo, 30(1):9-25.

Wüst, G. 1932. Das ozeanographischebeobachtungsmaterial. Wiss. Ergebn. dt. atlant.Exped. "Meteor", 4(2):290p.

(Manllscript received 17lll/Y 1995; revised 04December 1995; accepted 20 Marsh 1996)