Estuarine, Coastal and Shelf Science (1994) 39, 287-302 Zooplankton Distribution in the Guarad River Estuary (South-eastern Brazil) Rubens M. Lopes Centro de Estudos do Mar, Universidade Federal do Parané, Travessa Alfredo Bufren, 140. Curitiba (PR), 80020-240 Brasil Received 4 January 1993 and in revised form 2 August 1993 Keywords: zooplankton; estuaries; community composition; abundance; seasonal variations; biological interactions; tropical coasts; Brazil The structure and seasonal variability of zooplanktonic associations were analysed over a L-year period in the Guarat River Estuary (south-east Brazil). ‘The spatial segregation of the zooplankton was mainly conditioned by the longitudinal salinity gradient, as revealed by multivariate analysis. The upper estuary was dominated by a large indigenous population of the estuarine copepod Pseudodiapromus richardi. The estuarine marine copepods Acartia Ulfeborgi and Oithona hebes, together with meroplanktonic organisms, co-occurred with P. richardi in the middle estuary. The maintenance of high standing-stocks of matine euryhaline species such as Paracalanus crassirostris and Preudodiapiomus acutus in the outer estuary, largely depended on recruit- ‘ment from the coastal water. Temperature fluctuations influenced the seasonal ‘occurrence of the zooplankton, with higher densities occurring in the warmer months. Increased river inflow caused by summer rainfalls had an evident but comparatively smaller effect on the zooplankton abundance. Predation on rotifers and grazing on nanoplankton are probably important factors regulating the local zooplankton dynamics. Introduction ‘The south-eastern coast of Brazil has been submitted to strong anthropogenic impacts, especially estuarine areas near large urban concentrations and important commercial ports (Tommasi, 1987). Recently an 82000-ha nature reserve (Juréia-ltatins Ecological Station; Figure 1) containing a preserved area of the Adantic Rainforest and a network of unpolluted streams and estuaries (Por & Imperatriz~ Fonseca, 1984), was established between the heavily impacted Santos Estuary and the well-investigated Cananéia estuarinelagunar system (Por et al. 19844; Shaeffer-Novelli et al., 1990). This has provided an opportunity to carry out hydrobiological studies in a conservation area. ‘The rivers of the Juréia-Itatins Reserve are relatively short and empty directly into the sea (Por et al., 19846; Lopes ef a/., 1986). Therefore, they show stronger alternation or superposition of tidal mixing and freshwater discharge than the larger and relatively stable estuarine-lagunar systems, such as Cananéia (Miyao er al., 1986). The estuarine 0272-7714/94/090287 + 16 $08.00/0 © 1994 Academic Press Limited288 RM, Lopes 47°30" 47°00" 46°30" 24°30" 25°00 ‘Figure 1, Southern coast of Sio Paulo State, Brazil, showing the limits of the Jurcia Ecological Station and the positioning of the sampling stations (1-3) in the Guarat_ River Estuary, The hatched area bordering the estuary represents the mangrove forest es, zooplankton dynamics are greatly influenced by these basic hydrographical features (Almeida Prado-Por & Lansac-Toha, 1984). Further investigations are required to improve knowledge of the plankton communi- ties of this region. In this paper an account is given of a I-year study of the zooplankton dynamics in the Guarat River Estuary (Figure 1). ‘Study area ‘The Guarati River is about 25 km long. It descends through a series of waterfalls and rapids from the north-eastern border of the Jtatins massif, forming two main branches which converge into the Guarait River proper (Figure 1). Like the nearby Verde River and numerous other clearwater streams of the Atlantic Rainforest (Por, 1992), the upper stretch of the Guarat River carries transparent, slightly acid waters with pH values of 5-65. This river type contrasts with the post-Pliocenic alluvional plain streams which have no defined spring-fed baseflow and carry black humic waters with pH of around 4 (Por, 1986; Lopes & Por, 1990). Such hydrochemical differences are evident in the freshwater streams but not along their estuarine reaches, where pH is salinity-related and variable. ‘The tidal regime of the Guarait River is semi-diurnal and high tide levels vary between 0-25 and 1-5m. No data are available on freshwater inputs and tidal velocities. The mangrove forest is well developed, bordering the estuary from the mouth to the inner station (Figure 1)Zooplankton of the Guaratt River Estuary 289 Material and methods ‘Three stations along the upper 15 km from the river mouth (Figure 1) were visited monthly, during ebb and flow spring tides, from October 1986 to October 1987. Due to technical problems no collections were made during the low tide in October 1986, Zooplankton samples were obtained by filling a 9-1 Van Dorm bottle at the surface and just above the bottom, and pouring the water through a net of 75-um mesh aperture, ‘The volume of filtered water was 18-1801, depending on the abundance of the organisms, Semi-quantitative surface and bottom samples were also taken by means of 5-min horizontal rows, using a modified WP2 net (Fraser, 1968) and a sledge ‘D ’-net (Almeida Prado, 1973), respectively, both with a 75-ym mesh aperture. A comparison between each sample pair (bottle/net) revealed a maximum of 20% variation of the relative abundance (percentage) of each zooplankton species. Fast-swimming fish larvae were the only forms undersampled by the Van Dorn bottle (Lopes, 1989). Thus, bottle samples were considered representative of the micro- and mesozooplankton assemblages of Guaraa River. After the biological samplings, temperature, salinity and pH were measured in situ, with a Kahlsico RS5-3 thermo-salinometer and a portable Micronal B278 pH meter, respectively, both at the surface and near the bottom. Oxygen concentrations were determined for the same depths by a modified Winkler method (Grasshoff et al., 1983). Rainfall records were obtained from a climatological station from the Department of ‘Water and Energy of Sio Paulo State, situated c. 7 km from Guaraii River. Between one-quarter and the total zooplankton sample was counted for quantitative analysis (using the bottle samples), ensuring that at least 30 individuals of each taxa and developmental stage were enumerated. Aliquots of the semi-quantitative samples (net samples) were taken with a Stempel pipette or a Motoda splitter (Omori & Ikeda, 1984) for the calculation of the relative zooplanktonic abundance (percentage composition). Multivariate analysis was carried out with the whole quantitative data set (150 observations) after a log transformation. Principal component analysis was applied to a correlation matrix of selected standardized descriptors, computing separately the zooplankton taxa and physical/chemical variables. Cluster analysis of taxa was performed by the average linkage method, using Pearson’s correlation coefficient (Legendre & Legendre, 1984). Calculations were carried out by the BMDP statistical software (Dixon, etal., 1981). Results Envitonmental data Rainfall records were typical for the region, with a total annual precipitation of approximately 2400 mm. Late-spring and summer were the rainy periods, and the winter was dry (Figure 2). Salinity showed minor seasonal variation (Figure 3), weakly correlated with the rainfall record, The only exception occurred in December, when a strong freshwater outflow was observed as a consequence of the heavy summer rains. Within the range of the three stations, salinity fluctuated from near-zero values in the inner estuary to a maximum of 35-4, close to the river mouth. Vertical stratification was less pronounced during high tide, especially at station 3, but it occurred frequently during low tide, The pH was salinity-related, ranging from 5-3 in freshwater to 8-9 in seawater.290 RM, Lopes er) 160) 120 80 Rainfall (mm week") 40) oN DJF MAM Js J A SO 1986 1987 ‘Months Figure 2. Weekly rainfall records for the Guarati River Estuary. Arrows indicate days in which sampling took place. Water temperature and dissolved oxygen were inversely correlated throughout the year, showing a well-defined seasonal variation (Figure 4). However, both spatial (horizontal and vertical) and tidal differences were small. Zooplankton composition Holo- and meroplanktonic taxa and their respective relative frequencies are shown in ‘Table 1, In general, the zooplankton community was dominated by estuarine and estuarine/marine copepods, including Pxeudodiapromus richardi, Acartia lilljeborgi, Para~ calanus crassirostris and Oithona hebes. They belong to the ‘ central group ’ of species which appears in the Brazilian estuaries from the Amazon up to 25°C (Almeida Prado-Por & Lansac-Téha, 1984). Ergasilid copepods (non-parasitic males and juveniles) were also frequent, although much less abundant. Marine stenohaline species of copepods seldom appeared. Among other holoplanktonic taxa, the most important in terms of relative or absolute abundance were tintinnid ciliates (nine species), the appendicularian Oikopleura divica {at station 3), and the rotifer Synchacia bicornis (at stations 1 and 2). The latter species is recorded here for the first time in the Neotropical region (following Koste & Paggi, 1982). The occurrence of an unidentified acoelous turbellarian belonging to the family Convolutidae, which appeared in population blooms, also merits attention. Meroplanktonic forms were dominated by polychaetes (mainly spionids), cirripede larvae and, to a lesser extent, mollusc veligers and brachyuran zoeze, which sometimes reached high densities. Meiobenthic organisms, such as cyclopoid and harpacticoid copepods, nematodes, water mites, chydorids and insect larvae present in the samples, were probably washed out from decaying leaves, macroscopic algae or from the river bottom itself, due to the turbulent water flow. There were no truly freshwater plankton species, Zooplankton abundance and distribution Overall, lower zooplankton densities were found during the colder period from April to September and in the rainy month of December, while higher densities were found during the warmer period (January-March and October; Figure 5). Total abundance varied almost 4-fold throughout the year.Zooplankton of the Guarati River Estuary 291 wo ‘Eo T\ ° td) oO fe) 8 8 : % 35} A, A Satinity | 3 J J | M A M M, F F, J J Z Figure 3. Salinity values (practical salinity scale) registered at the three stations of Guarad River Estuary, Station 1 during low tide was not figured because the highest salinity recorded was 0-5 (=freshwater). (a) Station 1, high tide; (b) station 2, low tide; (© station 2, high tide; (d) station 3, low tide; and (¢) station 3, high tide. The percentage composition of selected zooplankton taxa at key periods and their numerical abundance along the year are shown in Figures 6 and 7, respectively. There were distinct patterns of zooplankton distribution in different sections of the estuary. ‘The copepod Pseudodiaptomus richardi dominated at the inner estuary (Figure 6). In this area, total zooplankton densities closely followed P. richardi fluctuations, a species which peaked in summer (January-March) and in October (Figure 7). Young copepodites and nauplii always outnumbered adults, the latter being most abundant292 RM, Lopes Station 1 400 25 80 60 20 40 20 = Spit og 8 3 rd 100 2 2 25) 80 FE [ oo 8 £ 20 ao 8 é 20 3 "Seaton 3 a a 100 25 80 60 20 40 20 6 0 ONDJFMAMJJASO ONDJFMAMJJASO 1986 1987 Months Figure 4. Average temperature and dissolved oxygen values registered during the sampling period in the Guarza River Estuary. Bars indicate the overall standard deviation between surface, bottom, high tide and low tide values. near the bottom. Pseudodiaptomus richardi was also found in the higher salinities of station 3, but here it never exceeded 3000 individuals m~*. ‘The estuarine marine copepods Acartia lilljeborgi and Oithona hebes co-occurred with P. richardi in the middle estuary, but were generally more abundant at station 3 (Figure 6). Both species appeared throughout the year, usually in salinities higher than 15. Oithona hebes had poorly defined annual maxima, while A. [ilijeborgi showed peak numbers in January and October (Figure 7). Paracalanus crassirostris and Pseudodiaptomus acutus were virtually restricted to the outer estuary and occurred in relatively low densities (Figures 6 and 7) Polychaete larvae (dominated by spionids) and cirripede nauplii occurred mainly in the middle and outer estuary (Figure 6), the former reaching high abundance in January and October 1987, and the latter in January and August (Figure 7). Table 2 shows the peak densities recorded for some other important taxa which occurred in the Guarati River. The marine euryhaline copepods Euterpina acutifons and Oncaea media appeared only in winter at station 3. The large number of acoels observed in October 1986 rapidly decreased until March 1987, when a new population developed in the outer estuary for a short period (see Figure 6). Tintinnopsis mortenseni and Oikopleura dioica reached maximum densities in February at station 3; both species were carried into the estuary from the nearby coastal area. The rotifer Synchaeta bicornis was found at stations 1 and 2 in small densities of less than 3000 individuals m~*, except forZooplankzon of the Guavait River Estuary 293 ‘TaBLs 1. List of the zooplankton taxa recorded from October 1986 to October 1987 in the Guarad River Estuary Foraminifera Tintinaina “Amphorellopsis acura Sche Contilla longa Brande* Eutintinrus lusus-tndae Entz* Facella ehrenbergi Claparede & Lachmann Leprotintinnus nordgoisti Brandt Tinsinnopsis directa Hada Tintinnopsis morcenseni Schmidt (TB) Tintinnopsis parvula Jorgensen" Tintinnopsis tacantinensis Kofoid & Campbell Medusae (ME) ‘Turbellatia (Acocla) (AC) Rotifera Brachionus spt Synchaeta bicontis Smith (SB) Non-identified” Nematoda Gastropoda (larvae) Bivalvia (larvae) (BI) Polychaeta (larvae) (PO) Hydracarina Cladocera Evade tergestina Claus" Penilia avivostis Dana® Chydoridae” Copepoda Acarta liljeborgé Giesbrecht (AL) Conycacus amazonicus F. Dahl” Coryeacus giesbrechti F. Dahl" Eucalanus pileatus Giesbrecht” Euerpina acutifrons Dana (EA) Halicyctops crassicornis Herbst Halizyctops exiguus Kiefer” Healicyclops sp.a ‘Microsetella rosea Dana ie Oithona hebes Giesbrecht (OH) Oithona ostealdocrust Oliveira Oithona plumifera Baied* Oithona simplex Farran* Oncaea curea Sars" ‘Oncaea media Giesbrecht (OM) Paracalanus erassirostis F. Dabl (PC) Preudodiaptomus aeutus F. Dahl (PA) Preudodiaptomus richard F. Dabl (PR) ‘Saphirella sp. Guveniles) Temora styljera Dana Ergasilidae (ER) Benthic Cyclopoida (CY) Benthic Harpacticoida (HR) Ostracoda ‘Mysidacea “Metamysidopsis clongata Bacescu® Isopoda Epicarides) Amphipoda* Decapoda Lucifer faxoni Borradsile* Brachyura (zoese) Other larvae* Cirripedia (nauplii) (CD) Insecta (larvae) CChironomidae (CH) Culicidae* Ephemeroptere* Homoptera* Trichoptera® Echinodermata (larvae) Chactognatha Sagitta friderici Ritter-Zahoni (SF) Appendiculari Oikoplewra divica Fol (OD) PPiscis (cggs and larvae) Occurred in less than 5% of the samples. Species symbols used in Figures 8-10 are indicated in parentheses. the cases included in Table 2. During October 1987 5. bicormis was the numerically dominant species in the inner estuary (Figure 6). The tintinnid Favella ehrenbergi peaked in the same month, but in higher salinity water. Mollusc larvae and brachyuran zoeae were frequent in the samples but seldom appeared in high densities (Table 2). Brachyuran zoeae were important in terms of the percentage in December (Figure 6); but low zooplankton densities were recorded on that occasion, The taxa selected for the cluster analysis were split into five groups, as follows (Figure 8) Group A: Paracalanus crassivostris, Oithona hebes, Acartia lilljeborgi, cirripede nauplii and polychaete larvae Group B: Tintinnopsis montenseni, Oikopleura dioica and bivalve larvae Group C: Oncaea media, Euterpina acutifrons and Pseudodiaptomus acutus Group D: Preudodiaptomus richardi, ergasilids and chironomid larvae Group E: Benthic cyclopoid and harpacticoid copepods294 RM. Lopes High tide Low tide (Staton) 38.2850 100 [Station 1 THEI ‘80 60 sesoya o8 Abundance (individuals m~* x 103) & s BS8 o8 ONDJFMAMJJASO NDJFMAMJIJASO 1986 1987 Months Figure 5. Total abundance of 2ooplankton in the Guarati River Estuary. (-. Surface samples; (——) bottom samples. The positioning of these groups relative to the first two axes of the principal component analysis is shown in Figure 9, Factor 1, explaining 34-65% of the variance, is a component of spatial heterogeneity, which in the Guarait River was salinity-related, Groups A, B, and C were positively associated with this axis, in contrast to groups D and E. The second factor, which accounted for 11-02% of the variance, reflects the influence of seasonal variations of temperature, dissolved oxygen and rainfall, on the zooplankton distribution. Taxa belonging to group B together with polychaetes, the rotifer S. bicornis and the copepod P. richardi, reached higher densities duting the warmer months, and were associated with the positive side of axis 2. Copepod species belonging to group C, along with Sagitra friderici and hydromedusae, were more closely associated with the colder months, as shown by the negative side of this axis. The only exception was P. acutus, whose abundance maxima occurred during the warmer period (see below). ‘The quadrant containing the negative values of both axes had no associated taxon, since it aggregated the point-objects of December, when the lowest zooplankton abundance and diversity were recorded, ‘An additional cluster analysis was carried out in an attempt to correlate separately females, males, young copepodites and nauplii of the most abundant copepod species (Figure 10). In general, the same groups shown in Figure 8 were formed. Pseudodiapromus acutus had a significant correlation with P. crassirostris, O. hebes and A, lilieborgi, instead of having an association with O. media and E. acurifrons, as indicated by the initial analysis. This was due to the weak correlation between P. acutus and other non-copepod taxons belonging to groups A and B, mentioned above.Zooplankton of the Guarati River Estuary 295 ol eT | iL 1H 2b 2H) BL SH o1L) 1H L2H” SL oH Stations Figure 6, Overall percentage composition of the zooplankton in the Guarad River Estuary over the different stations and tidal periods at four quarterly intervals: H,, high tide; L, low tide. O, Preudodiaptomus richardis ©, Psoudodiapiomus acutus; 2, Paracala- us crassivostnss W, Acartia lilleborgis ll, Oithona hebes; B8, Synchacta dicomis; FA, Acocle; #4, Polychacta; , Cirripedia; 4, Btachyura; @, others. (a) December, (b) March, (c) June and (4) October. Concerning the different developmental stages and sexes, the dendrogram (Figure 10) shows that adults of O. hebes, A. lilfeborgi, P. acutus and P. richardi, grouped apart from juveniles and nauplii. A similar pattern was found for E, acutifons, although males and females of this species were not as strongly correlated. Males of P. crassirostris occurred in very low numbers. Accordingly, they were weakly associated with other representa- tives of the species. The same is true regarding the parasitic females of ergasilid copepods. Discussion The environment Salinity profiles at the head and mouth of the Guaraii River Estuary were vertically homogeneous during most of the yeat, whereas the intermediate reaches were almost permanently stratified, suggesting a partially mixed circulation pattern (type B of Pritchard, 1955). Similar salinity profiles were recorded by Por et al. (19848) in the estuary of Una River but a contrasting picture is apparent in the small and shallow Verde River Estuary where a nearly permanent halocline is observed (Lopes er al., 1986). ‘Temperature and dissolved oxygen values were within the range reported for other estuaries of the region (Tundisi et al., 1978; Por et al., 1984b; Lopes er al., 1986). Zooplankton composition For unknown reasons, species richness was much lower in the Guarati River than in other adjacent estuaries, where several marine stenohaline species occur during high296 RM. Lopes High tide Oithona hebes Low tide Station 2 LA ch we Pseudodiaptomus richardi Abundance (individuals m”? x 103) s-inatane“i] ONDJ FMAMJJASOONDJFMAMJJASO 1986 1987 Months Figure 7. Abundance of the dominant copepod species and mefoplankton taxa during. the sampling period in the Guarati River Estuary. (--—) Surface samples; (———-) bottom samples. tides, including the copepods Paracalanus quasimodo, Clausocalanus furcatus, Centropages elificatus and Oithona nana, the chaetognath Sagitta enflata and the appendicularian Oikopleura longicauda (Teixeira et al., 1965; Lopes et al., 1986; Monti: & Cordeiro, 1988;Zooplankton of the Guarait River Estuary 297 High tide Cirripedia (nauplii) Low tide Station 2 a (larvae) Abundance (individuals m™ x 10°) Pseudodiaptomus acutus 20 oF ONDJ FMAMJJASOONDJFMAMJJASO 1986 1987 Months Figure 7 (continued) Almeida & Prado-Por et al., 1989; Lansac-Téha & Lima, in press). Similarly, mysids were scarce and represented by only one species, Metamysidopsis elongata, contrasting with the higher diversity reported for the Cananéia estuarine-lagunar system (Por et al.; 1984a). ‘Among the microzooplankton forms, tintinnids were relatively diversified in the Guarai River, with nine species listed. An equivalent figure was obtained by Lansac- ‘Téha and Lima (in press) in the Una River Estuary and by Sassi and Melo (1982) in a larger estuary in north-eastern Brazil. ‘The rotifer Synchaeta bicomis was an important component of the microzooplankton assemblage at the inner estuary as indicated by its jarge standing-stock at certain periods. Although Synchaea is a widespread genus in brackish waters around the world (sce Egloff, 1986, and references therein) it has not been observed in the Brazilian estuaries, including the Suape estuarine area where rotifers were studied in detail (Neumann-Leitao et al., 1992).298 RM, Lopes ‘Taste 2. Annual peak abundances of selected zooplankton taxa recorded in the Guarait River Estuary Annual maxima Date, station. Texon (Gndividuals m=?) and tidal phase Oncea media 2600 August 87, 3H Euterpina acutfrons 4900 August 87, 3H Oikopleura dioica 8000 January 87, 34 9000 February 87, 3H ‘Synchaeta bicornis 22000 July 87, 2L 35.000 October 87, 2L 525.000 October 87, 1H Tinsinnopsis 34.000 February 87, 3H 4000 October 87, 3L Favella 5000 October 87, 3HU31. Acoel (Convolutidze) 22000 October 86, 3H. 90.000 March 87, 3H Gastropoda 12.000 September 87, 3H Bivalvia 5000 January 87, 3H 10.000 March 87, 3H Brachyura (Zoeae) 5000 January 87, 2HV/IL. H, High tide; L, low tide. co on aH ct PO or BI | é Similarity Figure 8. Similarity dendogram of zooplenksonic associations (A-E) at the Guarati River Estuary. Species symbols are identified in Table 1. Zooplankton distribution Pseudodiapromus richardi was the only planktonic copepod to develop a large indigenous population in the estuary, reaching very high densities in salinities from 0-5 to 20. Almeida Prado-Por and Lansac-Toha (1984) and Walter (1989) showed that P. richardi is one of the most common estuarine calanoids in tropical and subtropical areas of theZooplankton of the Guarati River Estuary 299 Fy (11-02%) {Temperature Fy 84-65%) } Salinity A pluviosity ~ \ 03 to, | Temperature 08 FFigure 9. Principal componenc analysis applied to correlation matrix of selected biotic and abiotic variables, The equilibrium eycle of descriptors (Legendre & Legendre, 1984) and the species association obrained through cluster analysis are indicated, Species symbols are identified in Table 1. south-western Atlantic, The cyclopoid Qithona oswaldocruzi, frequently one of the dominant copepods in low-salinity waters of Brazilian estuaries (Rocha, 1986), was found in low numbers and only during winter. ‘The retention of the P. richardi population within the inner estuary might be explained by the higher densities of adults and late-stage copepodites near the bottom (data not shown), thus avoiding the stronger surface currents during the ebb tide. A similar mechanism was proposed for Pseudodiapromus hessei in a South African estuary (Wooldridge & Erasmus, 1980), Severe flooding decreased the abundance of Pseudo- diaptomus and other zooplankters in the Guarai River Estuary in December. Flushing may occur for longer periods in other tropical and subtropical estuaries (Chua, 1973; Qasim, 1977; Nordi, 1982; Wooldridge & Bailey, 19825 Por er al., 1984; Lopes et al., 1986). ‘The maintenance of relatively high standing-stocks of marine-curyhaline copepods such as Pseudodiaptomus acurus, Oncaea media, Euterpina acutifrons and Paracalanus crassirostris at station 3 depended on recruitment from the adjacent coastal water. Acartia liljeborgi and Oithona hebes reproduced in the middle and outer estuary, as suggested by the high densities of ovigerous females of these species recorded at stations 2 and 3. Juveniles and nauplii of the coastal copepod species penetrated farther up the estuary than did adults. Conversely, nauplii and young copepodites of Pseudodiaptomus richardi occurred more frequently in more saline areas than the adults. These results were confirmed by the cluster analysis (Figure 10). Bjémberg and Moreira (in press) interpretated this differential ontogenetic distribution as a consequence of the higher tolerance of larval and young stages to the fluctuation of environmental variables.300 RM, Lopes 10 08 06 04 02 0 02 Similarity Figure 10. Similarity dendrogram of the associations among females (F), mates (M), young copepodites (C} and nauplii (N) of the most frequent copepod species collected jn the Guarati River Estuary. Other species symbols are identified in Table 1. Large simultaneous peaks of Acartia lilljeborgi, Pseudodiapromus richardi, larval spio- nid polychaetes and the rotifer Synchaera bicornis were recorded at the inner estuary in October 1987. There is evidence of high consumption rates of Synchaeta spp. on small phytoplankton cells in brackish waters (Egloff, 1988). In addition, a negative relation- ship between the abundance of rotifers (mainly Synchaeta) and microflagellates was reported by Dolan and Gallegos (1991). The peak densities of Synchaeta bicornis in the Guarai River were probably linked to the same kind of trophic interaction. The unusually high numbers of polychaete larvae and copepods were apparently associated with exploitation of the lange rotifer standing-stock. Predation of mesozooplankton on estuarine Synchacia spp. has been documemed, especially for the copepod Acartia tonsa (Allan et al., 1976; Stoecker & Egloff, 1987; Dolan & Gallegos, 1992). The positive correlations between Oikopleura dioica, Tintinnopsis mortenseni and bivalve larvae are also suggestive of trophic relationships. Bivalves were reported to feed on tintinnids in estuarine enclosures (Horsted et al., 1988; Riemann et al., 1990). Appen- dicularians and ciliates consume mainly pico- and nanoplanktonic particles, tespectively (King et al., 1980; Rassoulzadegan & Etienne, 1981; Stoecker et ai., 1983), which were thus possibly co-occurring with Oikoplewa dioica and tintinnids during the warmerZooplankton of the Guarait River Estuary 301 months in the Guarai River. This supposition is also corroborated by the high densities of pelagic acoclous turbellarians in the October-March period, since these animals are potential hosts of nanopianktonic photobionts (Stoecker et al., 1989). Acknowledgements I thank Prof. T. K. , Bjémberg for her advisorship during this work. Comments and suggestions by Prof. F, D. Por, S. A. Gaeta, F. P. Brandini, P. C. Lana and two anonymous referees are fully acknowledged. I am indebted to R. Sassi, C. E. F, Rocha and A. L. Oliveira for their taxonomical assistance and to T. A. Cordeiro for drawing Figure 6. The Department of General Ecology of the University of So Paulo provided the academic support for this work. Financial support was provided by FAPESP grant 85/2925-0 to R.M.L. and by FAPESP grant 86/0004-7 to T. K. S. Bjémberg. References Allan, J. D., Kinsey, T. G. & James, M. C. 1975 Abundance and production of copepods in the Rhode River sub-estuary of Chesapeake Bay. Chesapeake Science 17, 86-89. Almeida Prado, M. S. 1973 Distribution of Mysidacea (Crustucae) in the Cananéia region. Boletim de Zoologia ¢ Biologia Marinha, Sao Paulo 30, 395-417 Almeida Prado-Por, M. S. & Lansac-Taha, F. A. 1994 The distribution of brackish water Calanoide (Copepoda) along the Coast of Brazil. Hydrobiologia 113, 147-150. Almeida Prado-Por, M. S., Pompeu, M. & Por, F. D. 1989 ‘The impact of the Valo-Grande canal on the planktonic copepod populations of the Mar Pequeno Seaway (Sdo Paulo, Brazil). in Environmental Quality and Ecosystem Stability (Spanier, E., Steinberger, Y. & Luria, H., eds). 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