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Aus Dem Institut Für Polarökologie Der Christian-Albrechts-Universität Zu Kiel Aus dem Institut für Polarökologie der Christian-Albrechts-Universität zu Kiel ECOLOGICAL STUDIES ON THE TROPICAL LIMPET Cellana testudinaria (Linnaeus, 1758): INFLUENCE OF ENVIRONMENTAL FACTORS ON THE ROCKY SHORE BENTHOS OF THE BIG KAI ISLAND, SOUTHEAST MOLLUCCAS, INDONESIA Dissertation zur Erlangung des Doktorgrades der Mathematisch-Naturwissenschaftlichen Fakultät der Christian-Albrechts-Universität zu Kiel Vorgelegt von Abraham Seumel Khouw KIEL 2003 „Gedruckt mit Unterstützung des Deutschen Akademischen Austauschdienstes“ Referent: Priv. –Doz. Dr. D. Piepenburg Korreferent: Prof. Dr. M. Spindler Tag der mündlichen Prüfung: 4 December 2003 Zum Druck genehmigt: Kiel, den ................................ Der Dekan ECOLOGICAL STUDIES ON THE TROPICAL LIMPET Cellana testudinaria (LINNAEUS, 1758) : INFLUENCE OF ENVIRONMENTAL FACTORS ON THE ROCKY SHORE BENTHOS OF THE BIG KAI ISLAND, SOUTHEAST MOLLUCCAS, INDONESIA Dedication This thesis is dedicated to : my beloved wife Costavina Litamahuputty and our son Winfield Bierhoff Khouw Da ist das Meer – so unendlich groß und weit, unzählbar sind die Tiere darin, große wie kleine Psalm 104, 25 Es lobe IHN Himmel und Erde, die Meere mit allem, was sich darin regt. Psalm 69, 35 © A.S.Khouw ACKNOWLEDGEMENTS ACKNOWLEDGEMENTS So many people have worked so hard and supported me so much to make this thesis possible. Words alone cannot express my appreciation. I am truly grateful for all their help and encouragement. First and foremost, I would like to thank Priv. Doz. Dr. D. Piepenburg who supervised me and to whom I will be eternally indebted for the surveillance, guidance, support and opportunity that he has given me. I hope that one day I will be able to return all his efforts to my students in Indonesia. I would also like to convey my thanks to Prof. Dr. M. Spindler for his tireless efforts in providing opinion, criticism, and guidance on the manuscript and especially for providing me with all the equipments needed during the field experiments. This study was done while I have been awarded a postgraduate scholarship by the German Academic Exchange Service (DAAD), which made my stay in Germany possible. In this regard I am most grateful to Dr. G. Gad and Dr. U. Toyka-Fuong. I am exceptionally gratefull to Dr. M. Schmid and Prof. Dr. K. Bischof, for their constructive criticism and helpful comments on the manuscript. I appreciate the cooperation and excellent assistance of several people in Institute for Polar Ecology of Kiel University. In particular, M. Steffens, Dr. I. Werner, M. Bartz, A. Lorenz, H. Schünemann, and my roommate Dr. E. Renjaan, Ms. L. Puturuhu, and Mr. I. J. Zakaria. I wish to thank all people in IPÖ: Dr. I. Hempel, Dr. K. Meiners, Prof. Dr. M. Bötler, S. Lischka, R. Krapp, J. Kotzerka, R. Möller, A. Scheltz, F.P. Rapp, and J. Boyens-Seegert, for the tremendous and interest atmosphere that they have performanced during my study. I am very gratefull and particularly indebted to Piepenburg’s family for their excellent accomodation during my first day in Kiel, especially for staying overnight. Valuable field assistance came from J. Notanubun. The same can be given to Renjaan’s family, H. Suikeno, J. Pattikawa, Ongkers’s family, and Masrikat’s family. Special thanks goes to Manusama’s family (Pak Kris and Tante Joti), Andreas Harms, Seth Xeflide, Andreas Beck, Tomasz Barganowski, and other friends for their big support and prayers. Many thanks and deep gratitude to my dear friends Corra Pattiselanno, Gino & Rita Limmon, and Julius Masrikat not only for their financial support during my stay in Indonesia, but also for their tremendous prayers. My final acknowledgement goes to my parents and my families who have been open and honest in sharing their heartwarming greatest love, support and prayers to make my study possible. Ecological studies on the tropical limpet … A. S. Khouw SUMMARY / ZUSAMMENFASSUNG SUMMARY / ZUSAMMENFASSUNG SUMMARY The present study addresses the effects of environmental factors (seawater temperature, salinity, humidity, precipitation, insolation, air temperature, wave action, and tidal cycle) on the ecology of the patellid limpet Cellana testudinaria at the intertidal rocky shore of Ohoiwait (Big Kai Island, Southeast Molluccas, Indonesia). Sampling was conducted once each month at three different shore levels from October 2001 to September 2002. Average ambient seawater temperature ranged from 29.3 0C in August to 32.2 0C in October (31.2 ± 1.0 0C), and salinity fluctuated from 30.3 in August to 34.0 in October (31.9 ± 1.1). The highest insolation (76 %) occurred in September, the lowest (28 %) during December (54.3 ± 18.0 %). The mean monthly air temperature varied between 26.2 0C in August and 28 0C in October (27.1 ± 0.6 0C). Air humidity and precipitation ranged from 81 to 90 % (85.9 ± 3.3 %) and from 51 to 547 mm (251.8 ± 159.1 mm), respectively. The tides were of the 'mixed prevailing diurnal' type. The average spring range was 1.5 m, the neap range was 1 m. The monthly tidal excursion ranged from 80 hrs in June to 145 hrs in October and November (117.1 ± 25.2 hrs), while the wave heights varied between 0.5 m in September and October and 3 m in March and April (1.6 ± 0.9 m). Population structure and dynamics The distribution, abundance and biomass of the intertidal limpet C. testudinaria are described separately for the high, middle and low shore levels. A total of 2402 limpets were obtained in 12 monthly collections. The mean density over the whole period was 11.12 ± 4.51 ind.m-2. The density tended to be higher at the high (15.79 ± 7.54 ind.m-2) and the middle (14.67 ± 13.99 ind.m-2) than at the low shore level (2.90 ± 2.44 ind.m-2). Analysis of variance showed significant density differences among shore levels, months, as well as a significant interaction between shore levels and months. The small-scale dispersion patterns did not show any seasonal variability. They were strongly clumped throughout the year and at each shore level. The conversion factor between body wet weight (WW) and ash-free dry weight (AFDW) was 0.19. The average energy content of the limpets was 21.59 kJ.g-1 AFDW. The power function describing the relationship between body size (shell length L, in mm) and body weight (B, in mg AFDW) was B = 0.03236 L 2.7703. The mean annual population biomass was estimated to be 1013 ± 748 mg AFDW.m-2 (21.8 kJ.m-2). The highest biomass (3236 mg AFDW.m-2 or 69.9 kJ.m-2) occurred in September 2002, whilst the lowest (544 mg AFDW.m-2 or 11.7 kJ.m-2) was in March 2002. The monthly mean biomass (619 mg AFDW.m-2 or 13.4 kJ.m-2) of the rainy season was lower than that (899 mg AFDW.m-2 or 19.4 kJ.m-2) of the dry season. Ecological studies on the tropical limpet … A. S. Khouw SUMMARY / ZUSAMMENFASSUNG The average relative condition factor (CF) was higher (1.27) for limpets at the middle shore level than for those at the high (1.25) and the low shore (1.22) levels. Monthly shell-length frequency distributions were used to analyse the size and age structure of the population. The lengths of the collected specimens ranged from 8.0 to 31.8 mm. The analysis of the successive frequency distributions suggested that the population consisted of 4 to 5 distinct age groups (cohorts) at any given time, and that two new cohorts recruited during the one-year investigation period. An analysis to determine the growth pattern using the FiSAT software showed that the longevity of C. testudinaria extended up to 2 years. The values of von Bertalanffy growth -1 parameters (L∞, K and t0), estimated from size-frequency distributions, were 33.1 mm, 1.4 yr , and 0.09, respectively. The highest growth increments were 25 % and 18 % of the asymptotic length during the first 3 and 6 months, respectively. The effects of environmental variables on the growth rates showed important seasonal variations, with the highest increment of 2.6 mm/month during the dry season. The present study also highlights the fact that the recruitment of C. testudinaria was highly variable between months and shore levels. Both methods used to determine recruitment (direct counting and FiSAT) yielded similar results, which indicated that recruitment was highest during the transition season (March/April). The mean mortality of C. testudinaria was estimated, using three different methods, to be 94 % per year (Z = 2.81). Highest monthly mortality rates, determined with the Ault-Ehrhardt approach, were observed in October 2001 and April 2002, when desiccation stress and wave action, respectively, were highest. However, there was no significant linear correlation between mortality and environmental variables. A study on the reproduction of C. testudinaria indicated that the population consisted of 50 % neuters (incl. juveniles as well as adults with resting gonads and, hence, doubtful sex), 30 % adult males and 20 % adult females. Spawning took place in July and August. Males spawned earlier than females. Both sexes had mature gonads from May to August. The mean gonad index (GI) and the gonad condition (MGC) of males were higher than that of females. Male gonads accounted for a maximum of 23 % of the total body weight in July while females invested a maximum of 22 % of their body weight in gonad tissue in August. Ecological studies on the tropical limpet ... A. S. Khouw SUMMARY / ZUSAMMENFASSUNG The annual population production of C. testudinaria was estimated by the mass specific growth rate method to be 2.81 g.m-2 or 60.7 kJ.m-2.yr-1.
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