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Estuary of Mundaka, Bay of Biscay) SCI. MAR., 61 (2): 173-188 SCIENTIA MARINA 1997 Tidal influence on zonation and occurrence of resident and temporary zooplankton in a shallow system (Estuary of Mundaka, Bay of Biscay) F. VILLATE Landare-Biologia eta Ekologia Saila, Ekologi Laborategia, Zientzi Fakultatea, Euskal Herriko Unibertsitatea, 644 P.K. E-48080 Bilbao, Spain. SUMMARY: Tide-induced variability in the zooplankton community was analysed during 2 tidal series of 14 h during spring tides and neap tides in the shallow estuary of Mundaka by relating short-term changes in composition and spatial dis- tribution of populations to tide-associated parameters (water level, current velocity and direction, mixing and stratification, and salinity). A strong tidal influence was found in the zooplankton composition and distribution on a day-week time scale (spring-neap tidal cycles) and at hour time scale (high-low tidal cycle). During spring tides, suspended meiofauna shows high densities at flood and ebb tides; the neritic zooplankton can reach the inner estuary at high water, and the autochtho- nous zooplankton move from the inner zone, at high water, to the outer zone, at low water, over distances of more than one third of the estuary length. During neap tides, autochthonous populations remain around the low middle estuary, the pene- tration of neritic zooplankton is feeble, and the occurrence of benthic forms in the water column is negligible. The observed effects on the abundance and zonation of resident and temporary zooplankton indicate that tides are a key factor in the estu- ary of Mundaka by regulating the stock of neritic and autochthonous zooplankton within the estuary, the reproductive suc- cess of benthic populations which have planktonic larval development, and meiofauna dispersion and availability as food in the water column. Key words: Zooplankton, tychoplankton, tidal cycle, transport, estuaries. RESUMEN: INFLUENCIA DE LAS MAREAS EN LA ZONACIÓN Y PRESENCIA DEL ZOOPLANCTON EN UN SISTEMA COSTERO (ESTUARIO DE MUNDAKA, BAHIA DE VIZCAYA). – Los cambios en la comunidad zooplanctónica por efecto de las mareas fueron analiza- dos durante 2 series mareales de 14 h en mareas vivas y muertas en el estuario somero de Mundaka, relacionando los cam- bios a pequeña escala temporal de la composición y distribución espacial de las poblaciones con parámetros ligados a las mareas (nivel del agua, velocidad y dirección de corriente, mezcla y estratificación y salinidad). El efecto de la marea sobre la composición y la distribución del zooplancton fue muy notorio tanto a una escala temporal de dias-semanas (ciclos de marea vivas-muertas) como a una escala de tiempo horaria (ciclos pleamar-bajamar). En mareas vivas, la densidad de meio- fauna resuspendida es elevada durante el ascenso y el descenso de la marea, el zooplancton nerítico alcanza el tramo interi- or del estuario en pleamar y el zooplancton autóctono es transportado desde el tramo alto del estuario, en pleamar, hasta el tramo exterior, en bajamar, sobre distancias superiores a un tercio de la longitud total del estuario. En mareas muertas, las poblaciones autóctonas permanecen alrededor del tramo medio bajo del estuario, la penetración del zooplancton nerítico es escasa y la aparición de formas bentónicas en la columna de agua es irrelevante. El efecto observado sobre la abundancia y distribución del zooplancton residente y temporal indica que en el estuario de Mundaka las mareas son un factor determi- nante en la regulación de los efectivos de zooplancton nerítico y autóctono en el estuario, los sucesos reproductivos de las poblaciones bentónicas con desarrollo larvario planctónico, y la dispersión y disponibilidad como alimento de la meiofau- na en la columna de agua. Palabras clave: Zooplancton, ticoplancton, ciclos de marea, transporte, estuarios. *Received August 23, 1996. Accepted February 10, 1997. TIDAL EFFECTS ON ESTUARINE ZOOPLANKTON 173 INTRODUCTION (Whitfield, 1988); and tidal influence on sediment transport and settling (Uncles et al., 1992; Ten Tides play a major role in the functioning of Brinke & Dronkers, 1993; Ten Brinke, 1994; Smith many coastal systems, being responsible for notice- and FitzGerald, 1994). able mid-term (spring-neap cycles) and short-term Regarding zooplankton, several papers have (low-high water cycles) variations in the abiotic and pointed to the importance of tidal dynamics induc- biotic characteristics of these systems. This role has ing spatial variability in zooplankton distribution inspired several authors to analyse tidal control of within estuaries (e.g. Lee and McAlice, 1979; All- communities and processes in different systems. dredge and Hammer, 1980; Gagnon and Lacroix, Examples include: tide-induced phytoplankton and 1981, 1982). However, tides are not only responsi- microphytobenthos exchanges (Riaux-Gobin, ble for distributional patterns, but also for dynamic 1987); tidal influence on bacteria, microphytoplank- events and survival of endemic populations by ton and microzooplankton abundance (Morales- exporting and reducing their standing stocks. This Zamorano et al., 1991); tidal control of seston quan- effect of tidal exchange in regulating estuarine zoo- tity and quality (Flegey et al., 1992); tidal export of plankton may be especially relevant in small sys- particulate organic matter (Boto and Bunt, 1981); tems where strong river discharges can occur (Ken- tidal redistribution of macrodetrital aggregates nish, 1990). FIG. 1. – Map of the estuary of Mundaka showing the sampling sites, and the boundaries of the three parts mentioned in the text. 174 F. VILLATE This report examines the effects of tides on mid- around 13 km long, and is characterized by three term and short-term variability of suspended living clearly distinguishable physiographic parts. The organisms larger than 200 µm in a shallow tidal sys- outer part is a broad area open to the sea and filled tem (the estuary of Mundaka). The aim of the study by large intertidal flats, the middle part is occupied was to evaluate the importance of changes in zoo- mainly by emerging salt-marshes, and the inner part plankton composition, abundance and zonation comprises a narrow man-made channel that meets induced by tides in this type of estuary. the meandering channel of the river in the upper estuary and its surrounding wetlands. The river inflow is normally low (monthly mean MATERIAL AND METHODS about 1 m3 s-1 from the main tributary) in relation to the volume of the estuarine basin (mean volume of Study area 32.9 105 m3), such that the estuary is mainly polyha- line and euhaline. However, strong increases in The estuary of Mundaka is a shallow meso-tidal freshwater inputs are frequent due to the torrential system located in the south-east of the Bay of Bis- regime of tributary streams. As a result, the resi- cay, on the Basque Coast (Fig. 1). It forms the cen- dence time is highly variable, and calculated flush- tral axis of the Urdaibai Biosphere Reserve, is ing times (21 to 581 days) at moderate and low river FIG. 2. – Time series of water height, salinity and current velocity during spring tides atB and G sites of the estuary of Mundaka. Circles: time series on 12 August 1991. Squares: time series on 13 August 1991. For salinity and current velocity plots, open symbols: surface data, filled symbols: bottom data, semi-filled symbols: mid-depth data. TIDAL EFFECTS ON ESTUARINE ZOOPLANKTON 175 discharges differ largely (Villate et al., 1989). Tides Sampling and data are semidiurnal, and at spring tide the high water is not significantly delayed between the outer estuary Data presented here were obtained from analy- and the upper reaches. The tidal exchange is high, ses of samples collected during a study on the tidal with the mean tidal prism-mean basin volume ratio effects on seston quantity and quality in the estuary greater than 1 (Villate et al., 1989). of Mundaka. The study was carried out in summer Previous research on mesozooplankton (Vil- (August 1991) when disturbances induced by river- late, 1989-90; 1991; Villate et al., 1993) showed runoff fluctuations are smaller and the develop- that true estuarine holoplankters such as Eury- ment of endemic zooplankton reaches the annual temora species cannot develop in the estuary of maximum. Mundaka. The endemic mesozooplankton is dom- Sampling was performed at two sites in the inated by the estuarine-marine species Acartia main channel during two spring tidal cycles on 12- bifilosa, which reach the highest density in sum- 13 August and two neap tidal cycles on 18-19 mer. The meroplankton contributes largely to the August. Sampling sites were located at the lower total zooplankton, and is dominated by barnacle part (station B) and at the upper part (station G) of and polychaete spionid larvae in spring and by the middle estuary (Fig. 1). These sites were gastropod larvae in summer. selected to obtain the most accurate estimation of FIG. 3. – Time series of water height, salinity and current velocity during neap tides at B and G sites of the estuary of Mundaka. Circles: time series on 18 August 1991. Squares: time series on 19 August 1991. For salinity and cur- rent velocity plots, open symbols: surface data, filled symbols: bottom data, semi-filled symbols: mid-depth data. 176 F. VILLATE populations moving throughout the estuary during using a hydraulic flow meter General Oceanics. the tidal cycle. Seaward to the B site, data record- The tidal height was determined by recording the ed at a single point may be less suitable because water level on graduated stakes located at each the basin is wider and the water circulation sampling site. Measurements were made just becomes more complex. Upward to the G site, before and after sampling. although most the water volume moves into the For zooplankton analysis, the entire sample was artificial channel, estuarine waters and popula- examined under a stereoscopic microscope. All indi- tions are also derived from the original channel. viduals were counted, and keyed to different taxo- When tides fall, waters coming from both chan- nomic categories.
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