Tidal and Diel Dynamics in a Nursery Area: Patterns in Fish Migration in a Mangrove in North Brazil

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Tidal and Diel Dynamics in a Nursery Area: Patterns in Fish Migration in a Mangrove in North Brazil Center for Tropical Marine Ecology Tidal and Diel Dynamics in a Nursery Area: Patterns in Fish Migration in a Mangrove in North Brazil Uwe Krumme Thesis submitted in partial fulfilment of the requirements for the degree of Doctor of Natural Sciences Faculty 2 (Biology/Chemistry) University of Bremen August, 2003 Center for Tropical Marine Ecology Tidal and Diel Dynamics in a Nursery Area: Patterns in Fish Migration in a Mangrove in North Brazil Uwe Krumme Thesis submitted in partial fulfilment of the requirements for the degree of Doctor of Natural Sciences Faculty 2 (Biology/Chemistry) University of Bremen August, 2003 ADVISORY COMMITTEE 1. Referee: Prof. Dr. Ulrich Saint-Paul (University Bremen, Center for Topical Marine Ecology, Germany) 2. Referee: Prof. Dr. Reiner Eckmann (University Konstanz, Limnological Institute, Germany) 1. Examiner: Prof. Dr. Wilhelm Hagen (University Bremen, Germany) 2. Examiner: Dr. Werner Ekau (Center for Topical Marine Ecology, Germany) 1. Member: Dipl.-Biol. Inga Nordhaus (Center for Topical Marine Ecology, Germany) 2. Member: Cand. rer. nat. Marion Kohn (University Bremen, Germany) To Reciane, Andreas and Chico 1 TABLE OF CONTENT List of Papers……………………………………………………………………………. 2 Summary.................................................................................................................…....... 3 Resumo...………............................……..............................................................…...….... 4 Zusammenfassung..............................................................................................….....….. 6 Scope of the Thesis ...........................................................................................…..…..…. 8 Sampling methods in tidal environments………….....….................................…....… 11 Temporal resolution, sampling effort and significance of results...................……….. 13 Interactions of tidal and diel cycle………............….......................................……...... 15 Importance of high-resolution sampling for identification of key patterns in tidal ecosystems...…..............……………………………………………...............…......... 16 Results................................................................................................................…............. 18 Patterns in the tidal migration of mangrove fish..........................................…..…....... 18 Tide and diel patterns..........................................................................…...…...... 18 Within-tidal cycle patterns....................................................................…...….... 19 Concluding Remarks and Research Perspectives...........................................….....….. 22 Acknowledgements..............................................................................................….......... 25 Literature cited.....................................................................................................…......... 27 Paper I Krumme et al., Tidal and diel changes in fish assemblage structure….…….…... 34 Paper II Krumme & Saint-Paul, Horizontal beaming…………………………….……... 77 Paper III Krumme, Vertical beaming…………………………………………………… 100 Paper IV Krumme & Liang, Tidal induced changes in a zooplankton community…….. 133 Paper V Brenner & Krumme, Migration and patterns in feeding of A. anableps.……..... 152 2 LIST OF PAPERS This thesis is based on the following papers, which are referred to by their Roman numerals. All papers are reproduced with the kind permission of the copyright holder. Paper I Krumme U., Saint-Paul U., Rosenthal H., (Submitted). Tidal and diel changes in the structure of a nekton assemblage in small intertidal creeks in Northern Brazil. Aquatic Living Resources. Paper II Krumme U. and Saint-Paul U., 2003. Observations of fish migration in a macrotidal mangrove channel in Northern Brazil using a 200-kHz split-beam sonar. Aquatic Living Resources, 16(3): 175-184. Paper III Krumme U., (Submitted) Patterns in the tidal migration of fish in a North Brazilian mangrove channel as revealed by vertical beaming using a 200 kHz split-beam sonar. Fisheries Research. Paper IV Krumme U. and Liang, T.H., (In Press) Tidal induced changes in a copepod-dominated zooplankton community in a macrotidal mangrove channel in northern Brazil. Zoological Studies. Paper V Brenner M. and Krumme U., (Submitted) Tidal migration and patterns in feeding of the four-eyed fish Anableps anableps L. in a North Brazilian mangrove. Marine Ecology Progress Series. 3 SUMMARY Anthropogenic degradation of mangroves - the intertidal forested wetlands on tropical and subtropical coasts - has increased dramatically in recent years. This thesis is part of a long-term project in the comparably undisturbed mangrove estuary of the Caeté River near Bragança (Pará, North Brazil), exposed to semidiurnal macrotides. The ecological investigations of the project focus on factors determining characteristic spatio-temporal patterns and the development of new methodological approaches to study such patterns. Mangroves are considered important as fish nurseries worldwide. The tide induces large-scale exchange processes of fish between the submerged mangrove at high water (HW) and the subtidal areas at low water (LW) on a short-time scale. These short-time dynamics in the tidal migrations of the fish are yet poorly understood, albeit they likely represent one of the most direct ecological links in the system because they include the everyday survival strategies and use patterns of the fish in this amphibious environment. Block netting in intertidal mangrove creeks at HW revealed the overwhelming significance of the factors tide and time of day and their combinations (spring tide at dusk and dawn, neap tide at midday and midnight) in structuring fish assemblages. In contrast to neap tide and daylight when nekton catches were poorest, spring tide and darkness produced the most complex fish assemblage structure when abundant species occurred with increased proportions and more rare species entered. Extreme spring tides produced extraordinary high nekton catches. Fish apparently respond with increased immigration to flood features that indicate increased HW levels. A stronger flood tide provides earlier habitat accessibility and increased resource availability for intertidal fish. Intertidal fish were juveniles or adults of small species, or juveniles of larger species. The almost complete absence of larger piscivores in the intertidal creek network suggests a low risk of predation for the juveniles, emphasizing the mangroves’ great nursery function. Stomach analyses of fish showed that the intertidal movements are usually feeding migrations. Fish were concentrated in the subtidal sections at LW. However, not all intertidal species and age groups reside at LW near the areas where they feed at HW. Hence, the fish may split their migration routes intra- and interspecifically at ebb tide to return to a specific LW habitat. Pioneer studies with a 200 kHz split-beam sonar beaming horizontally and vertically in a large mangrove channel, revealed that the entire fish community rode the first flood rise to achieve upstream transport and to enter the intertidal zone. At slack HW the fish milled around, indicating active foraging behavior. Despite low Secchi depths, diel changes in vertical distribution were apparent when fish exploited the water column only at night. Tracking parameters like direction of travel, swimming speed, track tortuosity and target strength showed tidal-related changes. The sonar provided unique high-resolution, non-invasive insights into the natural movement patterns of the fish. It has a high potential to advance the methodological approach in studying short-time patterns in fish migration. The surface-swimming four-eyed fish Anableps anableps entered the intertidal creeks rapidly with the first flood rise. They fed in the submerged mangrove at HW and returned gradually after the ebb current maximum to concentrate again in the subtidal parts of the main channel at LW. The combination of high inundation at daylight (spring tide-dawn) provided optimum foraging conditions for A. anableps while darkness and low inundation was linked to poorest foraging conditions (neap tide-night). The tidal migration of A. anableps was controlled by the water level, not by time. Fish that move into the intertidal zone early and leave late in the tidal cycle most likely optimize their foraging time in this shallow water environment. The mangrove zooplankton was dominated by copepods, an important food for juvenile fish. Zooplankton densities were higher at spring than at neap tide. In contrast to the number of zooplankton species, the densities were lowest at HW and highest at LW. Obviously, the temporal variations in abundance of fish and zooplankton are synchronized by the tide. This points to the need for simultaneous multi-disciplinary short-time sampling to comprehensively cover the range of possible functional interrelations between system compartments in areas exposed to extreme environmental conditions. 4 RESUMO As degradações antropogênicas de manguezais (florestas inter-tidais de litorais tropicais e subtropicais) aumentaram dramaticamente nos útimos anos. Esta dissertação faz parte de um projeto de longo prazo, realizado em uma área de manguezal no estuário do rio Caeté, em Bragança (Pará - Norte do Brasil). Este estuário é exposto a macro-marés semi-diúrnas, e é relativamente preservado se comparado com outros estuários. As investigações ecológicas do projeto
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