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Feeding by Shallow Water Holothuroids (Echinodermata) and Its Effect on the Environment

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Feeding by Shallow Water Holothuroids (Echinodermata) and Its Effect on the Environment ББК 28.691 Л34 УДК 591.53:593.96 Левин В. С. Л34 Питание мелководных голотурий и его влияние на донные осадки. — СПб.: Политехника, 1999. — 254 с: ил. ISBN 5-7325-0484-2 В монографии описаны новые методические приемы исследования, наиболее полно отвечающие биологической специфике шельфовых голотурий, выяснены основные за- кономерности использования этими животными пищевых частиц, приведены простран- ственные характеристики потребления ими пищевого материала, выявлены наиболее значимые для питания морфоэкологические особенности голотурий, оценено влияние трофической деятельности голотурий на среду обитания. Материал представлен на английском и русском языках. Книга адресована зоологам, экологам, специалистам, интересующимся измене- ниями в окружающей среде, вызываемыми жизнедеятельностью морских организ- мов. Levin У. S. Feeding by shallow water holothuroids (Echinodermata) and its effect on the environment. The aim of this book was to use new methodological approaches to comprehensively describe feeding by shallow-water holothurians, to quantitati"ely estimate the impact of feeding on the environment, especially on the bottom sediments, and to study latitudinal regularities in the modification of the environment by feeding activity. Л 1907000000-287 Без объявл. ББК 28.691 045 (01)-99 На переплете: Кукумария Eupentacta fraudatrix. Фото В. Даркина, Владивосток Cucumaria Eupentacta fraudatrix. Photograph of V. Darkin.Vladivostoc ©В. С. Левин, 1999 ISBN 5-7325-0484-2 © Издательство «Политехника», 1999 CONTENT Preface 7 Acknowledgements 8 Chapter 1. MATERIAL AND METHODS 9 1.1. Study areas and periods 9 1.2. Studied species 10 1.3. Basic methods 13 Chapter 2. BASIC FEEDING PECULIARITIES OF SHALLOW-WATER HOLOTHURIANS 15 2.1. Ecological classification 15 2.2. Structure of food-catching organs 20 2.3. Trophic-ecological features 23 2.4. Evolution of trophically important morpho-functional characteristics 28 Chapter 3. BOTTOM SEDIMENTS AS SOURCES OF FOOD MATERIAL 35 3.1. Sediment structure and composition 35 3.1.1. Methods 35 3.1.2. Sediment composition 37 3.2. Composition of food particles 46 3.2.1. Methods 47 3.2.2. The Japanese sea cucumber Apostichopus japonicus 47 3.2.3. Tropical Aspidochirotida 49 3.3. Hydrodynamic characteristics of food particles 54 Chapter 4. FEEDING PATTERNS OF DEPOSIT-FEEDING HOLOTHURIANS 57 4.1. Foraging behaviour 57 4.1.1. Methods 57 4.1.2. Food capture 59 4.1.3. Feeding movements 60 4.2. Feeding intensity and dynamics 65 4.2.1. Methods 65 4.2.2. Relationship between feeding intensity and food distribution 66 4.3. Feeding conditions and distribution patterns of deposit-feeding holothurians . 67 Chapter 5. FEEDING SELECTIVITY AND TROPHIC SPECIALIZATION OF SHALLOW- WATER HOLOTHURIANS 72 5.1. Feeding selectivity of deposit-feeding holothurians 72 5.1.1. Selectivity assessment methods 72 5.1.2. Feeding selectivity of Apostichopus japonicus and tropical holothurians. 74 5.1.3. Mechanism for selectivity 76 5.1.4. Are holothurians «optimal foragers»? 85 5.2. Trophic specialization of holothurians 87 5.3. Utilization of food by holothurians: Comparative aspects 89 Chapter 6. HABITAT MODIFICATION BY GRAZING BY SHALLOW-WATER HOLOTHURIANS 91 6.1. Basic types of impact on bottom sediments 91 6.2. Transport of food material 92 6.3. Alteration of physical and mechanical properties of sediments 97 6.3.1. Methods 98 6.3.2. Results of experiments 100 6.4. Latitudinal regularities of environment-modifying activity 103 в.4.1. Distribution of trophic groups of shallow-water holothurians 103 в.4.2. Latitudinal zonation of pattern and intensity of influence on bottom sediments Ill Conclusions 114 References 116 Index of Latin names 129 ПЕГАСЕ Holothuroids are distributed throughout the World Ocean, inhabiting various types of bottom from the intertidal zone to abyssal depths. They are dominant ia abundance, biomass and ecological importance in numerous bottom commu- aities. Holothuroids inhabiting the productive coastal marine zone deserve special attention since their role in structuring the habitat is most significant there. Despite rather extensive information on the feeding of particular species of holothuroids in particular areas, standard techniques for studying their trophic importance in coastal ecosystems are still lacking. This may largely be explained by the biological peculiarities of holothuroids, which often are not amenable to conventional methods of investigation. Future progress in under- standing the fundamental feeding characteristics of this important group of animals will require overcoming these methodological challenges. My aim in this book was to use new methodological approaches to compre- hensively describe feeding by shallow-water holothurians, to quantitatively estimate the impact of feeding on the environment, especially on the bottom sediments, and to study latitudinal regularities in the modification of the environment by feeding activity. At all stages of work I aspired to follow the remarkable definition of a famous Russian zoologist D.N.Kashkarov (1933): «The projection on the habi- tat is the basic characteristic of ecological research». Valery LEVIN ACKNOWLEDGEMENTS The author is happy to have this opportunity to acknowledge everyone who have helped to make this book possible. It is a pleasure to thank my colleagues from the Pacific Institute of Bioor- ganic Chemistry, the Institute of Marine Biology, the Zoological Institute, the Institute of Oceanology, the Institute of Biology of Southern Seas, the Institute of Evolutionary Morphology and Ecology of Animals, and the Pacific and All- Union Institutes of Fisheries and Oceanography for attention and help in the research efforts, valuable advice, and exacting but always well-wishing criti- cism. In collection of material I was assisted by professional and amateur SCUBA divers from various cities in Russia. Their efforts resulted in the collection of a vast amount of «background» material which helped to widen the scien- tific scope of the author. They also partici pated in complicated experiments requiring perfect div- ing skill, accuracy and exactness. I am deeply grateful to all of them. Samples of some species of holothurians from the Bahamas and the USA were kindly placed at my disposal by Carol Mosher (unfortunately now de- ceased) and Scott Smiley. Some printed and manuscript material not present- ed in the libraries of the former Soviet Union were made available thanks to Gustave Cherbonnier, Chantal Conand, Howard Feder, Jean-Pierre Feral, John Lawrence, Madelon Mottet, Ann Muscat, David Pawson, Francis Rowe and oth- er foreign colleagues. Finally, I wish to acknowledge the assistance of the most qualified editor I know, John M. Lawrence, who gave liberally of his time for this book. It was only by his great work with the English text of the manuscript and his most valuable remarks and advice that I could prepare it for publication. Chapter 1. MATERIAL AND METHODS 1Л- STUDY AREAS AND PERIODS Tfce basic studies were made between 1971 and 1995 with the use of some results of earlier observations (since 1967). The locations of the biological stations are presented in Fig. 1.1. Observations and field experiments on North Pacific holothurians were performed in the Peter-the-Great Bay of the Sea of Japan. The laboratory experiments were made at the Marine Experimental Stations of the Institute Motl». Fig. 1.1. Location of research and collection areas. Sites of stationary underwater observations and experiments are shown by squares, sites of personal observa- tions and samplings by dark circles, and sampling of collection material by light circles. Insets A-C show areas of most intensive research 10 Material and methods of Bioorganic Chemistry (Troitsa Inlet, 1967-1975), of the Pacific Institute of Fisheries and Oceanography (Popov Island, 1975-1978) and of the Pacific Oceanological Institute (Vityaz Inlet, 1978-1986). Tropical holothurians were studied in the cruises of research vessel «Dmi- triy Mendeleev» (1971, south-western part of the Pacific Ocean), of R/V «Izum- rud» (1974, the Indian Ocean), the cruise by the Pacific Institute of Bioorganic Chemistry to Cuba (1972-1973), the cruises by the Institute of Marine Biology (Far East Branch of the Russian Academy of Sciences) to Vietnam (1985 and 1987-1988). The following collection materials were analyzed: collections from expedi- tions of the Institute of Marine Biology to the Sea of Japan, the Okhotsk and Bering Seas (1976-1993) and to Vietnam (1980-1985), Pacific Institute of Fisheries and Oceanography to the Sea of Japan (1970-1974), the Pacific In- stitute of Bioorganic Chemistry to various Indo-Pacific areas (1978—1987) and North Kuril Islands (1981-1982), collections of the Institute of Oceanol- ogy of Cuba (1962-1972), and the Institute of Marine Research of the Republic of Vietnam (1980-1985). Samples of five species of holothurians from Bahamas and their gut con- tents were made available through the courtesy of C.Mosher (Eleuthera, Baha- mas). Samples of Parastichopus (=Stichopus) californicus from Friday Har- bour Bay and their tissue extract for chemical analysis were kindly prepared by S.Smiley (University of California, Los Angeles, USA). 1.2. STUDIED SPECIES The publication represents the results of the author's study of 90 species of Holothurioidea (57 — Aspidochirotida, 21 — Dendrochirotida, 10 — Apodida, 2 — Molpadiida) (Table 1.1). All species' identification was made by the author. In this work I used the conventional holothurian system (Clark, 1922; Heding,
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