ANTARCTIC RESEARCH SERIES. American Geophysical Union

Transcription

1 ANTARCTIC RESEARCH SERIES American Geophysical Union

2 A N T A R C T I C R E S E A R C H S E R I E S A m e r i c a n G e o p h y s i c a l V n i o n V o l u m e 1 B I O L O G Y O F T H E A N T A R C T I C S E A S M i l t o n O. L e e, E d i t o r V o l u m e 2 A N T A R C T I C S N O W A N D I C E S T U D I E S M a l c o m M e l l o r, E d i t o r V o l u m e 3 P O L Y C H A E T A E R R A N T I A O F A N T A R C T I C A O l g a H a r t m a n V o l u m e 4 G E O M A G N E T I S M A N D A E R O N O M Y A. H. W a y n i c k, E d i t o r V o l u m e 5 B I O L O G Y O F T H E A N T A R C T I C S E A S I I G e o r g e A. L l a n o, E d i t o r V o l u m e 6 G E O L O G Y A N D P A L E O N T O L O G Y O F T H E A N T A R C T I C J a r v i s B. H a d l e y, E d i t o r V o l u m e 7 P O L Y C H A E T A M Y Z O S T O M I D A E A N D S E D E N T A R I A O F A N T A R C T I C A O l g a H a r t m a n V o l u m e 8 A N T A R C T I C S O I L S A N D S O I L F O R M I N G P R O C E S S E S 1. C. F. T e d r o w, E d i t o r V o l u m e 9 S T U D I E S I N A N T A R C T I C M E T E O R O L O G Y M o r t o n 1. R u b i n, E d i t o r V o l u m e 1 0 E N T O M O L O G Y O F A N T A R C T I C A 1. L i n s l e y G r e s s i t t, E d i t o r V o l u m e 1 1 B I O L O G Y O F T H E A N T A R C T I C S E A S I I I W a l d o L. S c h m i t t a n d G e o r g e A. L l a n o, E d i t o r s V o l u m e 1 2 A N T A R C T I C B I R D S T U D I E S O l i v e r L. A u s t i n, J r., E d i t o r V o l u m e 1 3 A N T A R C T I C P a t r i c i a K o t t V o l u m e 1 4 A S C I D I A C E A A N T A R C T I C C I R R I P E D I A W i l l i a m A. N e w m a n a n d A r n o l d R o s s

3 Volume 14 ANTARCTIC RESEARCH SERIES Antarctic Cirripedia Monographic account based on specimens collected chiefly under the United States Antarctic Research Program, William A. Newman and Arnold Ross Published with the aid of a grant from the National Science Foundation PUBLISHER AMERICAN GEOPHYSICAL UNION OF THE National Academy of Science-National Research Council January 29, 1971

4 A N T A R C T I C l~olume 14 R E s E A R c H ( S E R I E S ANTARCTIC CIRRIPEDIA 1970 bj- the American Geophysical Union Suite 435, 2100 Pennsylvania Avenue, N.W. Washington, D. C Library of Congress Catalogue Card No Standard Book No X List Price, $25.00 Printed by THE HORN-SHAFER COMPANY DIVISION OF Geo. W. King Printing Co. Baltimore, Maryland

5 THE ANTARCTIC RESE14RCH SERIES THE A\T-~RCTIL RESEARCH SERIES is designed to provide a medium for presenting authoritati~e reports on the extensile and detailed scientific research work being carried out in Antarctica. The series lias been successful in eliciting contributions from leading research scientists engaged in antarctic investigations; it seeks to maintain high scientific and publication standards. The scientific editor for each volume is chosen from among recognized authorities in the discipline or theme it represents, as are the reviewers on whom the editor relies for advice. Eeginning uith the scientific investigations carried out during the International Geophysical Year, reports of research results appearing in this series represent original contributions too lengthy or other~+ise inappropriate for publication in the standard journals. In some cases an entire volume is devoted to a monograph. The material published is directed not only to specialists actively engaged in the ~+ork but to graduate students, to scientists in closely related fields, and to interested laymen versed in the biological and the physical sciences. Many of the earlier volumes are cohesive collections of papers grouped around a central theme. Future volumes may concern themselves with regional as well as disciplinary aspects. or 1% ith a comparison of antarctic phenomena ~+ith those of other regions of the globe. But the central theme of Antarctica nil1 dominate. In a sense, the series continues the tradition dating from the earliest days of geographic exploration and scientific expeditions-the tradition of the expeditionary volumes which set forth in detail everything that Nas seen and studied. This tradition is not necessarily outmoded, but in much of the present scientific mork one expedition blends into the next, and it is no longer scientifically meaningful to separate them arbitrarily. Antarctic research has a large degree of coherence; it deserves the modern counterpart of the expeditionary volumes of past decades and centuries which the Antarctic Research Series provides. With the aid of a grant from the Eational Science Foundation in 1962, the American Geophysical Union initiated the Antarctic Research Series and appointed a Board of Associate Editors to implement it. A supplemental grant received in 1966, the income from the sale of volumes in the series, and income from reprints and other sources have enabled the AGU to continue this series. The response of the scientific community and the favorable comments of reviewers cause the Board to look for^+-ard with optimism to the continued success of this endeavor. To represent the broad scientific nature of the series, the members of the Board were chosen from all fields of antarctic research. At the present time they include: Eugene L. Boudette, representing geology and solid Earth geophysics; A. P. Crary, seismology and glaciology; George A. Llano, botany and zoology; Martin A. Pomerantz, aeronomy and geomagnetism; Morton J. Rubin. meteorology; Waldo L. Schmitt, marine biology and oceanography; and Laurence M. Gould, honorary chairman. Fred G. Alberts, secretary to the U. S. Advisory Committee on Antarctic Names, gives valuable assistance in verifying place names, locations, and maps. AGU staff members responsible for the series are: Judith S. McCombs, managing editor. and Jane Bruce, style editor. MORTON J. RUBIN Chairman, Board of Associate Editors Antarctic Research Series

6 PREFACE THIS IS THE FIRST ATTEMPT to bring together what is known of the antarctic cirriped fauna in one comprehensive treatise. It covers the systematics of the barnacles of subantarctic and antarctic regions and their distribution, both recent and paleontological. A second part already in preparation will report on planktonic larvae. Antarctic Cirripedia, the second monographic contribution in the Antarctic Research Series, generally follows the pattern set by Kott7s Antarctic Ascidiacea (1969). The study, as far as possible, covers all records made by previous expeditions to Antarctic waters and culminates in this volume with the cruises of USNS Eltanin. The list of the principal vessels which have brought back cirriped materials from antarctic regions is a notable one: Aurora, Belgica, Challenger, Discovery, Endeavour, Ob', Pourquoi-Pas?, SGya, Terra Nova, Umitaka-Maru and William Scoresby. Yet the number of benthic samplings of cirripeds made by Eltanin under the United States Antarctic Research Program from south of the Antarctic Convergence exceeds the total of those returned by all earlier expeditions. In accordance with procedures established under the U.S. Antarctic Research Program, biological materials from terrestrial and marine activities forwarded to the Smithsonian Oceanographic Sorting Center are made available to qualified individuals recommended by the Advisory Committees for distribution of specimens to specialists. William A. Newman of the University of California, San Diego, was the specialist entrusted with the Eltanin cirriped collections in He was joined by Arnold Ross, Natural History Museum of San Diego, California, as junior author. Altogether, Newman and Ross have examined 85 species, 29 genera, and 9 families of cirripeds. Of these 20 species, 9 genera and 1 family are described as new. The text is illustrated by 48 plates, 90 text-figures, 11 charts, and 3 tables. All the Cirripedia from the Antarctic are figured and described with keys to each; charts are used to show their present geographical distribution. Paleogeographical and biogeographical aspects of the antarctic cirriped fauna are discussed in context with related forms throughout the world. The report includes the extraordinary discoveries of a new gigantic, sessile barnacle, the deepest balanomorphan known, and the first deep-water acrothoracican; the remarkably complex and uniquely constructed shell walls of the former are illustrated in three color plates. GEORGE A. LLANO Member, Board of Associate Editors Antarctic Research Series

7 ACKNOWLEDGMENTS THE RESOIJRCES OF MA\Y INSTITLTIOAS AND COLLEAGLES have been drar+n upon during the course of this studj. Initial arrangements for the cirripeds collected by the LSYS Eltanin were made ~ ith Dr. R. B. Tibbj. Universit) of Southern California. under the auspices of the United States Antarctic Research Program (USARP). The Eltanzn collections mere processed largely bj the Smithsonian Oceanographic Sorting Center. Washington, D. C.. under the directicn of Dr. H. Adair Fehlman. ~ith the assistance of Mrs. Beatrice Burch and her successor. Miss P. A. McLaughlin. The efficient and enthusiastic services of the SOSC have been much appreciated. We are especiallj indebted to Dr. Thomas E. Bouman. Smithsonian Institution, for providing tjpe specimens and for critical selection of comparative materials housed in the collections of the United States Kational Museum. Many thanks are due Miss Elizabeth Pope of the Australian Museum for the loan of specimens collected by the Aurora; Mrs. Patricia Lofthouse-Barker of the British Museum (Aatural History), for the loan of specimens taken by the Challenger; Dr. C. A. Fleming of the hew Zealand Geological Survey. uho made available the fossil specimens of Hexelasma az~cklandicum upon ~jhich the late T. H. Withers based his descriptions; Dr. I. G. Speden for providing specimens described in his ~ork on antarctic Pleistocene fossil deposits. presently housed in the New Zealand Geological Survey collections; Dr. Jan Stock, Zoological NIuseum, Amsterdam. for the loan of paratypes of Bathybalanus Hoek; and Dr. Huzio Utinomi. Seto Marine Biological Laboratory. Japan, for making arrangements with Dr. Jiro S&no. Tokyo University of Fisheries, for us to borrow the JARE specimens he described. Special thanks are due Dr. A. J. Southward, The Laboratory. Pljmouth, England. for the remarkable photographs of living and preserved specimens of Hexelasma hirsutum, and for the kindness of comparing this species with type material of H. aucklandicum and H. corolliforme housed in the British Museum (Natural History). For the loan or donation of other specimens that materially aided this study. we thank Dr. H. G. Stubbings. Admiralty Materials Laboratory, England; Dr. Brian Foster, Lniversity of Auckland, New- Zealand; Dr. J. T. Tomlinson. San Francisco State College. California; Dr. Victor A. Zullo, Marine Biological Laboratories, Woods Hole; Dr. J. A. Bullivant, New Zealand Oceanographic Institute; Norman E. Weisbord, Florida State University, Florida; Dr. Gilbert Rowe, Duke University, North Carolina; and Dr. Torben Wolff, Universitetets Zoologiske Museum, Denmark. The able bibliographic work by Mrs. Carol Kourtz-Platt, the inking of drawings by Mrs. Sally Beer, drafting by Howard Shirley, portions of the photographic work by Ronald Lam and Miss Nancy Ford and the stenographic assistance by Mrs. Cecelia Ross during the initial phase of the work were much appreciated. Support under a grant entitled Cooperative Systematic Studies in Antarctic Biology (GA-1374) from the National Science Foundation through Dr. I. E. Wallen, Director, Office of Environmental Sciences, for systematic studies of biological materials collected under USARP made this work possible and is gratefully acknowledged. WILLIAM A. NEWMAN ARNOLD ROSS Scripps Institution of Oceanography Natural History Museum La Jolla, California San Diego, California 92112

8 TABLE OF CONTENTS The Antarctic Research Series... Preface Ackno~~ledpments Introduction Historical Account Biogeographical Account... Paleogeography.... Contemporary Biogeography... Materials and Methods... Systematic Account... Key to Recent Antarctic Cirripeds... Order Ascothoracica Lacaze-Duthiers... Family Synagogidae Gruvel... Genus Ascothorax Djakanov... Order Acrothoracica Gruvel... Suborder Pygophora Berndt... Family Crypt~~hialidae Gerstaecker Genus Cryptophialus Darwin Key to the Austral Group of Cryptophialus... Family Zapfellidae Saint-Seine... Genus Brachyzapfes Codez Order Thoracica Darwin Suborder Lepadomorpha Pilsbry Family Lepadidae Darwin Key to Antarctic Lepadidae...., Genus Lepas Linne... Subgenus Lepas s.s.... Subgenus Dosima Gray... Genus Conclzoderma Olfers Family Scalpellidae Pilsbry Key to Recent Antarctic Genera of Scalpellidae... Genus Smilium Leach Genus Euscalpellum Hoek Genus Cretiscalpellum Withers Genus Zeugmatolepas Withers Genus Arcoscalpellum Hoek Key to Antarctic Species of Arcoscalpellum xi... v vii

9 Key to Genera Allied to Neoscalpellum Genus Neoscalpellum Pilsbry Genus Abathe~cal~ellum nov Genus Gymnoscalpellurn nov Genus Litoscalpellurn nov Genus Mesoscalpellum Hoek Genus Annandaleum nov Genus Scalpellum Leach Key to Species-Groups of the Genus Scalpellum Genus Australscalpellurn nov GenusBrochianov Suborder Verrucomorpha Pilsbry Family Verrucidae Darwin Genus Verruca Schumacher Key to Subgenera of Verruca Subgenus Altiverruca Pilsbry Suborder Balanomorpha Pilsbry Key to Families of Balanomorpha Family Bathylasmatidae nov Families of the Balanomorpha: Comparative Aspects Key to Genera of Bathylssmatidae Genus Bathylasma nov Key to Species of Bathylasma Genus Tetrachaelasma nov Genus Tessarelasma Withers Genus Hexelasma Hoek Key to Species of Hexelasma Genus Aaptolasma nov Key to Species of Aaptolasma Family Balanidae Leach Genus Balanus DaCosta Subgenus Austrobalanus Pilsbry Subgenus Chirona Gray Subgenus Bathybalanus Hoek Subgenus Balanus DaCosta Subgenus Megabalanus Hoek Genus Coronula Lamarck Genus Xenobalanus Steenstrup Order Rhizocephala Miiller Suborder Kentrogonida Delage Family Peltogastridae Lilljeborg Genus Briarosaccus Boschma xii

10 Appendices 1. Localities arid Bathjmetry of Previously Known Antarctic Cirripedia 2. Summary of Stations Yielding Specimens Covered in This Study 3. Eltaniri Cirripedia Listed According to Station 4. Eltanin Cirripedia Listed According to Depth 5. Rathymetric Distribution for Sampling Devices llsed during El~anzn Expeditions Covered in This Repor t References Glossary Plates Plate Legends Indices Systematic Subject