Q 11 U563 CRLSSI m\M 5- lllBf»- p: w jmr;'" •- •,-5 iMPiir SMITHSONIAN INSTITUTION MUSEUM OF NATURAL HISTORY For sale by the Superintendent of Documents, U.S. (Government Printing Office Washington, D.C. 20402 - Price $1.25 (paper cover) UNITED STATES NATIONAL MUSEUM BULLETIN 296 The Burrowing Barnacles (Cirripedia: Order Acrothoracica) JACK T. TOMLINSON Department of Marine Biology San Francisco State College SMITHSONIAN INSTITUTION PRESS CITY OF WASHINGTON 1969 Publications of the United States National Museum The scientific publications of the United States National Museum include two series, Proceedings of the United States National Museum and United States Na- tional Museum Bulletin. In these series are published original articles and monographs dealing with the collections and work of the Museum and setting forth newly acquired facts in the fields of anthropology, biology, geology, history, and technology'. Copies of each publication are distributed to libraries and scientific organizations and to specialists and others interested in the various subjects. The Proceedings, begun in 1878, are intended for the publication, in separate form, of shorter papers. These are gathered in volumes, octavo in size, with the publication date of each paper recorded in the table of contents of the volume. In the Bulletin series, the first of which was issued in 1875, appear longer, sep- arate publications consisting of monographs (occasionally in several parts) and volumes in which are collected works on related subjects. Bulletins are either octavo or quarto in size, depending on the needs of the presentation. Since 1902, papers relating to the botanical collections of the Museum have been published in the Bulletin series under the heading Contributions from the United States National Herbarium. This work forms number 296 of the Bulletin series. Frank A. Taylor Director, United States National Museum U.S. GOVERNMENT PRINTING OFFICE WASHINGTON: 1969 Contents Page History 1 Specimen Deposition 2 Acknowledgments 3 Abbreviations Used in the Figures 4 Methods 4 Orientation 7 Segmentation 8 Common Names 8 The Burrow 9 The Mantle 11 External Mantle Flap in Trypetesa lateralis ... 14 Digestive System and Feeding Mechanisms .... 17 The Cirri 19 FeedingHatoits of Trypetesa lateralis feraalea . 19 Excretory System 21 Muscular System 22 Circulatory and Respiratory Systems 24 Reproductive System 24 The Male 24 Ultrastructure of the Male and Its Sperm ... 25 Embryology and Mating Behavior 27 SexuaUty in the Cirripedia 28 Conjectures on Sexuality 29 Systematics and Characteristics 30 Fossils 134 Undescribed Fossil Species 141 Distribution 143 Key to the Order Acrothoracica 149 Phylogeny 151 Literature Cited 155 Index of Species and Figures 161 The Burrowing Barnacles (Cirripedia: Order Acrothoracica) History The study of the order Acrothoracica began in 1849, when Hancock discovered in shells of the gastropod Fusils and Buccinum a highly modified barnacle which he named Alcippe lampas. Darwin placed Alcippe in the Lepadidae very close to Alepas and Anelasma, although he recognized the great similarity to his Cryptophialus. For the latter he established a new order, the Abdominalia (1854). Noll (1872b) placed Alcippe with Cryptophialus in the Abdominalia. In 1905 Gruvel noticed the misconception upon which the name of Abdom- inalia was based, i.e., the cirri on the terminal segments of the body being of a thoracic nature rather than abdominal appendages, as Darwin had erroneously assumed. Gruvel changed the name Abdom- inalia to Acrothoracica and placed within this new order the genera Alcippe, Cryptophialus, Kochlorine Noll (1872a), and Lithoglyptes Aurivillius (1892). Norman (1903) noticed the synonymy of Alcippe with Blyth's genus of birds and changed the name to Trypetesa, although many subsequent workers failed to notice this. Berndt (1903a, 1903b, 1907b) and Kuhnert (1934) have reported on the anat- omy and development of these animals. Additional genera have been described and named IVeltneria Berndt 1907b, Berndtia Utinomi 1950a, Balanodytes Utinomi 1950b, Chytraea Utinomi 1950c, Rogeretla Saint-Seine 1951, Zapfella Saint-Seine 1954, Simonizapfes Codez 1957, and Brachyzapfes Codez 1957. The fossil form-genus Nygmites Magdefrau (1937) includes at least one species (A^. sacculus) which is considered to be an acrothoracican, anticipating Zapfella. 1 2 UNITED STATES NATIONAL MUSEUM BULLETIN 296 Specimen Deposition The following museums have been chosen as major depositories of acrothoracican material. Under each species description the depositories for that species will be abbreviated as indicated in the following schedule. The name of the individual most instrumental in assisting me at each institution and the one to whom the material was sent also is included. In addition to these major depositories, about one hundred in- stitutions were sent a standard set of three species of acrothoracican cirripeds: Weltneria spinosa, Kochlorine Jloridana, and ''Crypto- phialus" melampygos (see page 118). These institutions are too nu- merous to mention individually. The sets were bulk-labeled, so local fractioning into five sets and redistributing is expected. Aust Australian Museum, College Street, Sydney, N.S.W., Australia; Elizabeth C. Pope. BA Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Avda. Angel Gallardo 470, Buenos Aires, Argentina; Elena Martinez Pontes. Belg Institut Royal des Sciences Naturelles de Belgique, Rue Vautier, 31, Brussels 4, Belgium; W. Adam. BPB Bernice P. Bishop Museum, 1355 Kalihi St., Honolulu, Hawaii 96819; Yoshio Kondo. Accession number 7246, and catalog numbers B460 to B465. Brit British Museum (Natural History), Cromwell Road, London, S.W. 7, England; J. P. Harding, Keeper of Zoology. CA California Academy of Sciences, Golden Gate Park, San Francisco, California 94118; Department of Invertebrate Zoology; AUyn G. Smith. Dublin National Museum of Ireland, Kildare Street, Dublin, 2, Ireland; Colm O'Riordan, Natural History Division. Catalog numbers NMI.11.1967/ 1 to 17. Mex National University of Mexico, Cuidad Universitaria, Mexico City, Mexico; Instituto de Biologia; Llamas Flores. Catalog numbers: IBUNAM 022468—1 to 20; 090468—1 to 6. Paris Museum National d'Histoire Naturelle, 61, rue de Buffon, Paris Ve, France; Laboratoire de Zoologie; J. Forest. Seto Seto Marine Biological Laboratories (of the University of Kyoto), Sirahama, Wakayama-ken, Japan; Huzio Utinomi. SFSC San Francisco State College, San Francisco, California 94132; De- partment of Marine Biology. SIO Scripps Institution of Oceanography, University of California, La JoUa, California 92038; William A. Newman. UCT University of Cape Town, Cape Town, Cape Province, South Africa; Zoology Department; John Field. USNM United States National Museum, Washington 25, D.C.; Marine In- vertebrates; Fenner A. Chace, Jr. Catalog numbers: 122603 to 122629 and 123331 to 123334. : ) THE BURROWING BARNACLES (CIRRIPEDIA: ACROTHORACICA 3 Vict National Museum of Victoria, 284-321 Russell Street, Melbourne, Victoria, Australia; Edmund D. Gill. Catalog numbers: J- 186 to J-192. Acknowledgments It is a pleasure to acknowledge the kind consideration and assistance offered to me by the institutions and individuals listed above under the subject of Specimen Distribution. In addition, the following individuals and institutions have been most helpful Australia: John Yaldwyn and Donald McMichael, Australian Museum; Robert Endean and the Great Barrier Reef Committee. France: Dr. B. Salvat and Dr. F. Salvat, Museum National d'Histoire Naturelle, Paris. Great Britain: R. Bassindale of Bristol; D. J. Crisp of Menai Bridge Labora- tories; H. Barnes of Millport, Scotland. Hong Kong: Dennis Hill, University of Hong Kong. Italy: Stazione Zoologica di Napoli (NSF Table Award). Japan: Tadeshige Habe, National Science Museum, Tokyo; S. M. Shiino, Mie Prefectural University, Tsu. Liberia: Frank Tichy and Dr. Joshna of the University; Dr. and Mrs. William Winnett; Russell Riley. New Zealand : Elizabeth Batham, and the staff of the Universities of Canterbury, Otago, and Auckland. Nigeria: Alice O' Grady. Panama: Stewart and Claire Jadis, Canal Zone Police; Joe Becker, U.S. Army. Philippines: The National Museum. Singapore: D. S. Johnson of the University, and Eric Alfred of the National Museum. South Africa: Mr. and Mrs. Lawrence Meiring of Cape Town; Mrs. C. M. Connolly of Fishoek. Trust Territories of the Pacific Islands: The Commissioners and their staffs; Peter and Ann Wilson; the Robert Owens, George Hagiwaras, and Peter Hills. United States: Ensign and Mrs. Gaylord Galiher, USN; Mr. and Mrs. Henry Scott at Addis Ababa, Ethiopia; Leo Hertlein at the California Academy of Sciences; C. G. Bookhout and Lawrence McCloskey of Duke University Marine Laboratories; Tom Hopkins of New College and Cape Haze Marine Labora- tories, Sarasota, Florida; University of California at Santa Barbara; University of Hawaii Marine Laboratories; Dr. Willard D. Hartman of the Peabody Museum, Yale University. This work was begun as a thesis for the degree of Doctor of Philos- ophy in Zoology at the University of Cahfornia at Berkeley under the supervision of Dr. Cadet Hand. Substantially furthered by a sabbatical leave for one year from San Francisco State College, it was completed under a five-year grant for a study on the reproduction 4 UNITED STATES NATIONAL MUSEUM BULLETIN 29 6 of barnacles from the National Institutes of Health, United States Public Health Service, number GM-09953. Special thanks