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A Manual for the Study of Meiofauna

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A Manual for the Study of Meiofauna NEIL C. HULIM A Manual and JOHN S. GKA' for the Study of Meiofauna SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 78 Neil c. Huiings A Manual and John S. Gray r •• tor the Study of Meiofauna SMITHSONIAN INSTITUTION PRESS CITY OF WASHINGTON 1971 SERIAL PUBLICATIONS OF THE SMITHSONIAN INSTITUTION The emphasis upon publications as a means of diffusing knowledge was expressed by the first Secretary of the Smithsonian Institution. In his formal plan for the Insti- tution, Joseph Henry articulated a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge not strictly professional." This keynote of basic research has been adhered to over the years in the issuance of thousands of titles in serial publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Annals of Flight Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to ^oology Smithsonian Studies in History and Technology In these series, the Institution publishes original articles and monographs dealing with the research and collections of its several museums and offices and of professional colleagues at other institutions of learning. These papers report newly acquired facts, synoptic interpretations of data, or original theory in specialized fields. Each publica- tion is distributed by mailing lists to libraries, laboratories, institutes, and interested specialists throughout the world. Individual copies may be obtained from the Smith- sonian Institution Press as long as stocks are available. S. DILLON RIPLEY Secretary Smithsonian Institution Official publication date w handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. UNITED STATES GOVERNMENT PRINTING OFFICE WASHINGTON : 1971 For tale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402 - Price $2.75 International Conference on Meiofauna 1-11 July 1969, Tunis, Tunisia ORGANIZATION N. C. Hulings, convener J. S. Gray, co-convener R. P. Higgins, co-convener EDITORIAL STAFF FOR MANUAL N. C. Hulings I. M. Newell J. S. Gray W. Sterrer A. D. Mclntyre J. B. J. Wells PARTICIPANTS Dr. C. Clausen Dr. W. D. Hummon Zoological Laboratory Department of Zoology University of Bergen Ohio University Bergen, Norway Athens, Ohio 45701 Dr. M. T. Gomoiu Dr. S. Husmann "Traian SaVelescu" Institute of Biology Limnologische Flusstation Department of Marine Biology Max-Planck-Instituts fiir Limnologie Constantza, Romania Schlitz, West Germany Dr. J. S. Gray Dr. B. O. Jansson Wellcome Marine Laboratory Askolaboratoriet Robin Hood's Bay Trosa, Sweden Yorkshire, England Dr. C. Jouin Laboratoire de Zoologie Dr. R. R. Hessler Faculty des Sciences Scripps Institution of Oceanography 7, Quai St. Bernard La Jolla, California 92037 Paris, France Dr. R. P. Higgins Dr. E. Kirsteuer Office of Oceanography and Limnology Department of Living Invertebrates Smithsonian Institution American Museum of Natural History Washington, D.C. 20560 New York, New York 10024 Dr. W. D. Hope Dr. H. Kiihl Department of Invertebrate Zoology Institute fiir Kiisten- und Binnenfischerei National Museum of Natural History Laboratorium Cuxhaven Washington, D.C. 20560 Cuxhaven, West Germany Dr. N. C. Hulings Dr. P. Lasserre Office of Oceanography and Limnology Institut de Biologie Marine Smithsonian Institution University de Bordeaux Washington, D.C. 20560 33 Arcachon, France in IV SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Mr. A. D. Mclntyre Dr. J. Renaud-Mornant Marine Laboratory Museum National d'Histoire Naturelle Aberdeen, Scotland 43, rue Cuvier Paris, France Dr. F. Monniot Museum National d'Histoire Naturelle Dr. F. Riemann Laboratoire de Biologie des Invert£bre"s Marins Institut fur Meeresforschung Paris, France Bremerhaven, West Germany Dr. W. Sterrer Dr. I. M. Newell Bermuda Biological Station Department of Life Sciences St. Georges West, Bermuda University of California Riverside, California 92502 Dr. B. Swedmark Kristineberg Zoological Station Dr. W. Noodt Fiskbackskil, Sweden Zoologisches Institut der Universitat Dr. G. Uhlig Hegewischstrasse 3 Biologisches Anstalt Helgoland Kiel, West Germany Helgoland, West Germany Dr. O. H. Pilkey Dr. J. B. J. Wells Department of Geology Department of Zoology Duke University University of Aberdeen Durham, North Carolina 27708 Aberdeen, Scotland Dr. L. W. Pollock Dr. W. Westheide Systematics-Ecology Program II Zoologisches Institut der Universitat Marine Biological Laboratory Gottingen, West Germany Woods Hole, Massachusetts 02543 Dr. R. Williams Oceanographic Laboratory (S.M.B.A.) Edinburgh, Scotland Foreword The term meiofauna has been in use for only a relatively brief period. It was coined by Mare in 1942 to describe metazoans (and Foraminifera) too small to be retained on the 1-mm sieves used by marine biologists for the extraction of bottom fauna from samples of sediment when analyzing benthic communities. Thus, meiofauna refers only to a size class between macrofauna and microfauna. The term is now used for benthos and phytal fauna, but not for planktonic organisms, which are treated traditionally as a separate category and by special methods. The variation in form and the biological complexity of the meiofauna make it difficult to determine upper and lower limits in terms of body size which can be applied to all taxa. For the time being, however, the lack of a definitive and gen- erally accepted definition is less important than standardized methods of work for the taxonomists and ecologists studying these organisms. Practically all classes of the Metazoa, including the smaller species of macro- fauna, are represented in the meiofauna, while the Protozoa are represented by their largest forms, e.g., Foraminifera and Ciliata. The smaller protozoans are generally considered as microfauna. Gnathostomulida, Gastrotricha, Kinorhyn- cha, and Tardigrada are some groups of metazoans that are known exclusively from the meiofauna, while Mollusca, Brachiopoda, Echinodennata, and Ascidia- cea are examples of groups represented only by a very few species. Mclntyre (1969) uses the term "permanent members" for species belonging to the meiofauna during the whole of their life cycle. The meiofauna also includes a large number of "temporary members," i.e., larvae and juvenile stages of species that, as adults, belong to the macrofauna. If we refrain from an exact division according to type of biotope but are satis- fied to compare the three main groups—mud, interstitial sand, and phytal meio- fauna—we find certain morphological and biological features for each of these categories, reflecting the adaptation of the organisms to particular environmental conditions. These adaptations undoubtedly are most marked in the interstitial fauna of sand. Since Remane's pioneering work in the 1920s the interest of psam- monologists has been focused on the study of such adaptations. Within several systematic groups a greater proportion of elongated forms is found dian in types other dian the interstitial environment. The development of various types of adhesive organs is another adaptation found in many groups belonging to the interstitial fauna. Other adaptations are common for the meiofauna as a whole— for example, reproductive adaptations, which are related to the relatively small number of gametes that small animals produce. Copulation, hermaphroditism, and the formation of egg cocoons, as well as brood protection, improve the pros- pects of development and ensure the survival of the populations. The discovery of new groups of animals, particularly in the interstitial sand fauna—often of high taxonomic rank (Gnathostomulida, Mystacocarida, Actin- ulida, Macrodasyoidea, Acochlidiacea)—undoubtedly has been one of the major contributions to systematic zoology during the present century. Several research workers regard the many aberrant forms, morphological spe- cializations, and appearance of new organs in the meiofauna as having evolved vi SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY under the influence of the unique environment. Neoteny seems often to have been one of the reflections of evolution; it has resulted in a simplified organization compared with the presumed ancestral form and may be manifested in the loss of organs. In addition, neoteny has led to diminution of body size. Specializations such as the appearance of new organs, loss of organs, and reduction in body size have emphasized the systematic isolation of the aberrant species. It is difficult, for example, to demonstrate unequivocally the relationship between the Gnatho- stomulida and any definite class of the other Metazoa, or to show the affinity of the Archiannelida with a definite systematic group within the Polychaeta. The phylogenetic relationships and the taxonomic problems resulting from regressive evolution must remain unsolved until we discover new links in the often rather fragile morphological series. The definition of phylogenetic relationships is today one of the motives behind the work of many meiobenthologists in the fields of systematics and morphology. The discovery during recent years
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