Life History and Population Structure of the Exploited Mangrove Crab Ucides cordatus cordatus (Linnaeus, 1763) (Decapoda: Brachyura) in the Caeté Estuary, North Brazil vorgelegt dem Fachbereich 2 (Biologie / Chemie) der Universität Bremen als Dissertation zur Erlangung des Grades eines Doktors der Naturwissenschaften - Dr. rer. nat. - von Karen Diele Diplom Biologin Bremen 2000 Tag des öffentlichen Kolloquiums: 3. April 2000 Gutachter der Dissertation: 1. Prof. Dr. Ulrich Saint-Paul, Universität Bremen, Zentrum für Marine Tro- penökologie (ZMT) 2. Prof. Dr. Wilhelm Hagen, Universität Bremen ACKNOWLEDGEMENTS I would like to thank Prof. Dr. Ulrich Saint-Paul for having given me the opportunity to conduct this thesis, for all his support and for patiently awaiting the first megalopa find. I am also grateful to Prof. Dr. Wilhelm Hagen for kindly accepting the second supervision. Many persons were involved in one or the other way and I cannot thank them enough for their advice and help: Prof. Dr. Klaus Anger, Domingos, Raimunda and Simei Araújo, Tatjana Barros, Dr. Uta Berger, Dr. Matthias Birkicht, Dr. John Christy, Joo Deusdete, Dr. Victoria Isaac, Katrin Franke, Dr. Marion Glaser, Olaf Gundermann, Sabine Kadler, Dr. Volker Koch, Gesche Krause, Dr. Ruben Lara, Anke Machetzky, Ingrid Meeßen, Ulf Mehlig, Aldo and Katio Melo, Antonio Peixoto, Olegario Pinheiro, Marco Ortiz, José Nas- cimento and Christina Rosario, Rosa Saraiva, Prof. Dr. Horacio Schneider, Dr. Dirk Schories, Antonio Silveira, Dr. Iracilda Sampaio, Daniela Torres, Dr. Michael Türkay, Ursula Werner, Prof. Dr. Matthias Wolff and many other unnamed ZMT collegues. Rosa, Volker, Ulf and Dirk - again so many thanks for your friendship, support, sugges- tions, and corrections. Joo, Aldo and Senhor Domingos - data collection would not have been possible without your experience and help. Your sense of humour and friendship was invaluable for keeping me going. I am also very grateful to Helene Heider, Hans-Dieter Heider and Thea and Manfred Weyer for their hospitality and support. I warmly thank my parents and my brother for their loving encouragement and being there for me. Uli- thanks for all your love and help. This work was conducted in the frame of the bilateral research project MADAM, which is supported by the BMBF (code 03F0154A) and CNPq. Contents I CONTENTS List of Figures ................................................................................................................... IV List of Tables..................................................................................................................... VI List of Appendices............................................................................................................. VI SUMMARY .....................................................................................................................VII RESUMO ..........................................................................................................................IX 1 GENERAL INTRODUCTION................................................................................... 1 1.1 FRAME AND OBJECTIVE OF THE STUDY ..................................................................... 1 1.2 SHORT NOTE ON THE SYSTEMATIC POSITION AND DISTRIBUTION OF THE GENUS UCIDES ..................................................................................................................... 4 2 ENVIRONMENTAL SETTING................................................................................. 5 3 REPRODUCTION ...................................................................................................... 9 3.1 INTRODUCTION......................................................................................................... 9 3.2 METHODS ............................................................................................................... 10 3.2.1 Determination of breeding cycles and abundance of ovigerous females ............. 10 3.2.2 Plankton sampling................................................................................................ 12 3.2.2.1 Net-sampling........................................................................................................ 12 3.2.2.2 Passive collector-sampling................................................................................... 15 3.2.2.3 Processing of samples .......................................................................................... 16 3.2.3 Larvae rearing in the laboratory........................................................................... 17 3.3 RESULTS................................................................................................................. 18 3.3.1 Breeding cycle, abundance and size-frequency distribution of ovigerous females ............................................................................................ 18 3.3.2 Larvae findings in the estuary .............................................................................. 22 3.3.2.1 Water temperature and salinity ............................................................................ 22 3.3.2.2 Seasonal and lunar abundance of zoea larvae ...................................................... 23 3.3.2.3 Tidal and diel pattern of zoeal abundance............................................................ 24 3.3.2.4 Development time, seasonal and daily abundance of megalopae ........................ 27 3.3.2.5 Tidal pattern of megalopal abundance ................................................................. 30 3.3.3 Larvae findings in offshore waters....................................................................... 30 Contents II 3.4 DISCUSSION............................................................................................................ 32 3.4.1 Timing of reproduction and its implication for the dispersal of zoea larvae........ 32 3.4.2 Within- and between-year differences in breeding activities and larval output... 35 3.4.3 Size-frequency distribution of ovigerous females................................................ 38 3.4.4 Timing of estuarine re-invasion by megalopae .................................................... 39 4 GROWTH ................................................................................................................. 43 4.1 INTRODUCTION....................................................................................................... 43 4.2 METHODS ............................................................................................................... 44 4.2.1 Laboratory studies on early juvenile growth........................................................ 44 4.2.2 Field studies on later juvenile and adult growth................................................... 44 4.2.3 Age at size estimates ............................................................................................ 47 4.3 RESULTS................................................................................................................. 48 4.3.1 Early juvenile growth........................................................................................... 48 4.3.2 Later juvenile and adult growth ........................................................................... 51 4.4 DISCUSSION............................................................................................................ 54 4.4.1 Early juvenile growth........................................................................................... 54 4.4.2 Later juvenile and adult growth and age at size estimates ................................... 56 5 POPULATION STRUCTURE AND COMMERCIAL EXPLOITATION.............. 59 5.1 INTRODUCTION....................................................................................................... 59 5.1.1 Frame of the study................................................................................................ 59 5.1.2 Commercial crab capture and production systems............................................... 60 5.2 METHODS ............................................................................................................... 62 5.2.1 Field sampling...................................................................................................... 62 5.2.2 Size monitoring of commercial landings ............................................................. 63 5.2.3 Statistical analyses ............................................................................................... 64 5.3 RESULTS................................................................................................................. 65 5.3.1 Population structure............................................................................................. 65 5.3.1.1 Rhizophora mangle forest habitat ........................................................................ 65 5.3.1.2 Peripheral habitats................................................................................................ 72 5.3.2 Size of commercially landed crabs....................................................................... 73 5.4 DISCUSSION............................................................................................................ 76 5.4.1 Population structure............................................................................................. 76 5.4.2 Size of commercially landed crabs....................................................................... 79 Contents III CONCLUSIONS AND PERSPECTIVES