The Natural History of Enewetak Atoll, Vol. I the Ecosystem: Environments, Biotas, & Processes, DOE/Ev/00703-T1-Vol. 1 (DE87006110)

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The Natural History of G78 !‘3Iurne I The Ecosystem: Environments, Biotas, and Processes Edited by: Dennis M. Devaney Bernice P. Bishop Museum xpartment of Energy Office of Energy Research Office of Health and Ecological Research Division Office of Scientific and Technical Information U.S. Department of Energy University of Hawaii Bernice P. Bishop Museum University of Hawaii

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International Copyright,” U. S. Department of Energy. 1987. under the provisions of the Universal Copyright Convention. United States copyrtght IS not asserted under the United States Copyright Law, Title 17. United States Code. Library of Congress Cataloging-in-Publication Data The natural history of Enewetak Atoll. k!Gli 1. Devaney. Dennis M II Umred States Dept of Energy. Office of Scientific and Jschnlcal Work performed under contract No. DE-ACO8-76EV00703 The set of two volumes is available as DE870061 10 (D@E/EV;0073 T1-Vol 1) and DEq:i0061 11 NTIS Energy Distrlbutton Center Oak Ridge, Tennessee 37831 Price Code. Paper Copy A99 Printed in the United States of America

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As activity and funding at the Mrd-Pacific Research Laboratory began to diminrsh in the early 19805, j,Tt?medfltting that a syntheses be prepared of the three decades of research that had been conducted at -.~ . Laboratory on Enewetak Atoll For 30 years the Atoll served as a convenient. accessible location for : .:..?s of Mid-Pacific island ecosystems. and several hundred scientists utilized the facility. Primary fund -; icas pro\,ided by the Office of Health and Environmental Research, Ecologrcal Research Divisron. .’ S Department of Energy iformerly the Atomic Energy Commission and the Energy Research andi This is an attempt to synthesize in twp volumes the results of the Mid-Pacific Research Laboratory studies that have been published in hundreds of widely dispersed publications. It is hoped that present and future scientists involved in studies of Mid-Pacific islands will find this synthesis a convenient resource for Considerable time and effort were expended by many contributors to make this synthesis possible. -hanks are extended to all these authors for their manuscripts. Special appreciation is expressed for 3 Dennis Devaney’s dedication in filling gaps in the taxonomic descriptions of several invertebrate groups. This publication would not have been possible, however, without the determination and persis- :ence of Dr. Ernst Reese in organizing and collecting the material. Deepest gratitude is acknowledged for his conscientious efforts. Ecological Research Division Office of Health and Environmental Research United States Department of Energy

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..,,rk. from 1954 to 1984 under the auspices of the ; ,,.;,,,rlment of Energy. The history of the laboratory and ,j,,. ,easons for its support by the United States Departr,,,.nt of Energy are described in Chapter 1 of Volume Lover a thousand persons-established scientists, their !!I<’ laboratory during the 30.year period. Their efforts ::!~,:tiL! operation on a limited scale to the present A coli,,ctlon of papers recently appeared in the Bul/er!n of Much of the research conducted at the laboratory was on the marine environment. The reason was that the majority of scientists applying to wfork at Enewetak were “i.!rlr:e biologists. For many, this was the first opportunity :he biota of a coral atoll. Fewer studies were con- .n the terrestrial environment and its biota. dmaztng!y complete and thorough, and there are few, if any, studies of an equivalent ecosystem that equal the total research effort reported in these volumes. Volume I provides a synthesis of the research carried out under the subject headings of the respective chapters. Certain of the chapters, e.g., those on geology, subtidal --d intertidal environments and ecology, and those on reef .‘-scs and trophic relationshtps, summarize a great .>rty of research carrred out by many scientists for ‘aany years. In contrast, the chapters on meteorology and oceanography summarize research carried out under one integrated program involving fewer scientists working over Volume II of The Natural H~stoq of Eneuetok ;I!oll provides information on the taxonomy of animals and plants known to occur at Enewetak Atoll Thus taxonomy ’ ?!r+!sents a fulfillment of one of the first assignments to laboratory-to determine the screntifrc names of the ~:a of the atoll The collections on which the checkhsts 1: each chapter are based are housed at rhe Bernlctl F Bishop Museum in Honolulu and the U S National Museum of Natural History, Smithsonian Institutton, WashIn addition to the species checklists. each chapter in Volume II provides a succinct summary of the biota with respect to endemism, range extensions, and other features that set the Enewetak biota apart from those one might pendium of taxonomrc information for an atoll should prove of immense value to scientists interested in biogeog raphy and evolutionary biology of island ecosystems for years to come. One of the problems of editing these volumes has been the correct use of place names In some cases authors used the military code names for islands while others used the native names. Even the native names have changed from early phonetic spellings to the spellings currently in use and preferred by the Enewetak people For example, the name of the atoll has changed from Eniusetok to Enewetak. and, although the correct current spelling is used throughout, the old spelling occurs in older references and maps which appear in these volumes Maps giving the military code names and the native names preferred by the Enewetak people are located in Chapter 1 of Volume I. Surprisingly. it is difficult to determine the exact number of islands. Due to the effects of storms. small islands are ephemeral, and two islands and part of a third u’ere ob kterated by nuclear explosions Currently there are 39 recognizable islands, and these are shown on the map used throughout the book These volumes do not report on the exrensrve radiolog ~cal surveys and studies which have been conducted by the Lawrence Livermore Laboratory, University of California, and the Radiation Laboratory, University of Washington, also under the ausptces of the U. S Department of Dennis M Devaney, senior edrtor of thus \,olume. disap peared while coliectmg specimens off the island of Hawaii on August 13, 1983 Dennis was dorng what he loved best. collecting marine invertebrates. at the ttme of his dt!ath He collected extensively at Enewetak. and he undertook. the task of orgamzing the systematic chapters of L’olome II batrice 1. Burch. Dcvaney’5 assistan: at the

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Wisllop Museum. completed the task, and she has, written tfle lntroductlon to Volume II. It is fitting that the two volumes of this book are dedi cated to the people of Enewetak Atoll. They, like so many other human beings, were caught up by forces beyond their control and understanding in an immense cataclysm of human history 1~ a small way. this book stands as somethlng good that ha> rc>sulted from those years Professor of Zoology University of Hawaii, Honolulu

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Untverstty of Hawati. Honolulu. Hawart Untversity of Papua New Guinaa. Port Moresby. Papua New Guinea ’ ..~,I.G:s:iv of Rhode Island. Ktnqston. Rhode island 5: Jos2ph’s College. North Wrndham. Maine Unrversrty of Hawati. Honolulu. Hawati Bowltng Green State Untversity. Bowling Green, Ohio ‘.‘rxrerstty of Hawaii. Honoluiu. Hawaii University of Washtngton. Seattle. Washington University of Guam, Mangrlao. Guam University of Rhode Island. Kingston. Rhode Island University of Rhode Island. Kingston. Rho& Island University of Hawaii at Kanoa. Honolulu. Hawait Albuquerque, New Mexico Bowling Green State University. Bowling Green. Ohio

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Research at Enewetak Atoll: A Historical Perspective Philip Helfrlch and Roger Ray History of the People of Enewetak Atoll Physiography of Enewetak Atoll Geology and Geohydrology of Enewetak Atoll Oceanography of Enewetak Atoll Meteorology and Atmospheric Chemistry of Enewetak Atoii John T. Merrill and Robert A. Duce Subtidal Environments and Ecology of Enewetak Atoll intertidal Ecology of Enewetak Atoll Reef Processes: Energy and Materials Flux Trophic Relationships at Enewetak Atoll Nelson Marshall and Ray P. Gerber Terrestrial Environments and Ecology of Enewetak Atoll Bjology of the Rodents of Enewetak Atoll Avifauna of Enewetak Atoll

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I ::rsr volume of The Natural History of ‘!Znewetak \ : ,,T~~~~desa summary of the research carried out over !I,,’ :j()-year period from 1954 to 1984. The frontispiece ,jjustrates the dramatic contrasts between the immensity of ,jlc lagoon and the seemingly fragile necklace of small ,,innds which surrounds it, and also between the sea condi- .,,)i, on the windward, seaward side of the reef and the i b48.2first chapter discusses the history of research at .?k Atoll. The reasons behind the establishment of :_;!<,\Letak Marine Biological Laboratory are described. authors, Philip Helfrich and Roger Ray, have been1< dssociated with activities at Enewetak from the very early days They conferred with Robert W. Hiatt, the first direcror of the laboratory. In Chapter 2, Robert C. Krste, a foremost authority on the people of Micronesia, provides a hrstory of the Enewetak people to whom these volumes T& next four chapters deal with the physical environ- > oi Enewetak Atoll. In Chapter 3, Patrick L Colin -,‘r~oes the physiography of Enewetak. Colin served as_. rzsrdent scientist in charge of the laboratory from 1979 to The end of 1983 when all resident scientific staff left the atoll. Following the description of the atoll, Byron L. Ristvet, a frequent scientific visitor to Enewetak, provides a summary of the geology and geohydrology in Chapter 4. Next, in Chapter 5, Marlin J. Atkinson describes the oceanography. Under the direction of Stephen \r Smith. “Ainson participated in an important study of the lagoon -cularron Chapter 6 on the meteorology and atmos- -‘Ieric chemistry is the final chapter in the group of :napters dealing with the physical environment Of Enewetak Atoll. Written by John T. Merrill and Ro’bert A Duce. the chapter is based on the results of the SEAREX Project. Duce served as the director and principa! :nvesti3ator of the project. The next four chapters are devoted to the marine ecosystem and its biota. They summarize the large amount .ory in the marine environment. All of the authors were frequent visitors to the laboratory, and they have done a splendid job of reviewing the research carried out in their area of interest. In Chapter 7, Patrick L. Colin describes the subtidal environments of Enewetak and reports on the research done on the subtidal biota. This is followed in Chapter 8 by Alan J. KohE’s masterful summary of research in the intertidal environment. Kohn has been a student of tropical intertidal ecology for 30 years. He tackled a particularly difficult task because of the extensive study of the intertidal environment and its biota by many scientists over the years. Chapters 9 and 10 deal with processes and relation. ships in the marine environment. In Chapter 9. James A. Marsh, another frequent visitor to the laboratory and a recognized authority on coral reef processes, reviews the extensive work which was carried out at Enewetak on the community metabolism of coral reefs and related topics such as calcification processes, nitrogen and phosphorus cycles, and the role of detritus in the ecosystem. Nelson Marshall and Ray P. Gerber extend the ecosystem approach in Chapter 10 to include the entire atoll. They discuss the trophic relationship between the shallow reefs and the lagoon. Both Gerber and Marshall conducted research at Enewetak. The final three chapters are devoted to the terrestrial environment. Because fewer scientists applied to conduct research in the terrestrial environment, less work was accomplished, and an integrated overview is not possible. In Chapter 11, I report on the life history, behavior, and ecology of land crabs, review what is known about atoll soils, and conjecture on the carrying capacity of an atoll such as Enewetak. For a description of the vegetation, the reader is referred to Chapter 3 in Volume II by Janet 0. Lamberson. William B. Jackson, a frequent visitor to Enewetak over the years, and his co-workers Stephen H. Vessey and Robert K. Bastian report on their long-term study of the rodents in Chapter 12, and Andrew J. Berger summarizes our knowledge of the bird life of the atoll in Chapter 13 Berger, a noted ornithologist and the foremost authority on Hawaiian birds, made a number of trips to Enewetak. 1 suspect that few readers will read this volume from cover to cover, but those who do will gain an appreciation for the complexity of the atoll ecosystem and a better

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understanding of the intimate relationships between the seemingly fragile components of the ecosystem the lagoon, the reefs, the islands and their biotas, all perched on a volcanic and coral pinnacle in the vastness of the Pacific Ocean. In the final analysis, however, the book will serve its purpose best if the reader comes away wtth more questtons than answers and a desert to fmd the answers to these questions in future research on the natural history of 1 coral reefs and islands

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Research at Enewetak Atoll: A Historical Perspective i ,:c.r31.L~S Vegas. Nevada 89114. current -i:i-c theater of operations In World War Ii and their activities on the Pacific Islands afforded (>C contact and awareness of the physiography and . ....zr~l history of these small dots of land scattered in the .es: zkpanse of ocean. This enhanced awa’reness, coupled A!lh 2 recognized need by the military establishment for :xreased knowledge of Pacific Island areas, led to _._ ‘-,f many individual scientists whose interest had .Lred by wartime visits to these islands. In the . : ; further interest on the coral atoll of the tropical ;‘cc~i:c and Influenced the future of research at Enewetak Aio!l (Figs. 1 and 2). The origin of the spelling “Eniwetok” :s ~OS: but would appear to be a phonetic rendering of ‘~.i.lti: :he people called their atoll. In 1973 it gave way to :hG :urrent spelling, consistent with written Marshallese. ‘ yrld War II demonstrated the importance of these --ryered land masses to any milirary confrontation_.< ‘ACIDIC basin. After the war, the U S. Navy moved :<‘.elop a series of permanent bases from among the 32:‘:. temporary wartime bases and outposts which had 3e-5-. established across the Pacific. With the prominent iol+ of the Navy in developing and maintaining these baj+- it is not surprising that the Navy’s research arm !!f:: Irogram In the late 1940s aimed at a better under . ~3 of atoll morphology and of all aspects of island :Y mlcroorganisms lo human inhabitants The ONR I d series of expcdltions In conjunctlon with the 2 ; Science Association, many of u;hlch were to atolls In the central and western tropical Pacific. Arno Atoll in the southern Marshall Islands and Onotoa Atoll in the Gilbert Islands (now Kiribati) were subjects of intensive investigation in 1950 and 1953, respectively. Scientists involved in these atoll studies contributed to the establishment of the Eniwetok Marine BiologIcal Laboratory (EMBL) on Medren Island, Enewetak Atoll, in 1954. The second postwar activity which served to focus attention on the mid-Pacific area was the atomic weapons testing program in the northern Marshall Islands. Two atomic weapons had inflicted mortal damage upon Japan and had brought a precipitous end to the war in the Pacific. Military planners and strategists knew very little about this new and awesome strategic resource. Thus, an area was sought which might accommodate full-scale testing of atomic weapons. Neil Hines …