Pt: ¥ TT The Natural History of ~ Enewetak Atoll a Volume Il Biogeography and Systematics DOE/EV/00703-T1-Vol. 2 - (DE87006111) United States Department of Energy Office of Energy Research Office of Health and Environmental Research Ecological Research Division atics raphy and System Biogeog © 2or 10HM/ Tay Top: Aerial view of Enewetak Atoll from an altitude of 10,000 ft looking north. The wide south passage to the lagoon is at the bottom of the pic- ture. The three islands to the right of the passage are Enewetak, Medren, and Japtan. The deep east pass is seen between Medren and Japtan. The five southwest islands are seen to the left of the wide south passage. Ikuren is the first one. North of these islands is the shallow southwest pass. The Atoll is elliptical in shape measuring about 41 km from north to south and 33 km from east to west. [Photography by P. L. Colin.] Bottom: Aerial view of the northern end of Enewetak Island showing the cluster of buildings. of the Mid-Pacific Research Laboratory. The quarry is visible on the reef flat. The small island immediately to the north is Bokandretak. [Photography by E. S. Reese.] DOE/EV/00703-T1-Vol. 2 (DE87006111) The Natural History of Enewetak Afoll Volume Il Biogeography and Systematics Beet Lee ool Edited by: Dennis M. Devaney Bernice P. Bishop Museum Honolulu, Hawaii United States Department of Energy Ernst S. Reese University of Hawaii Office of Energy Research Honolulu. Hawaii Office of Health and Environmental Research Beatrice L. Burch Bernice P. Bishop Museur Ecological Research Division Honolulu, Hawaii PSE) Philip Helfrich Office of Scientific and Technical Information University of Hawaii U.S. Department of Energy Honolulu, Hawaii B30 [Photograph by Annabelle Lyman.] Bok in, kon menninmour ko im menin eddok ko ion Enewetak, ej kein kememej im kautiej ri Enewetak. This volume on the natural history of Enewetak Atoll is dedicated to the people of Enewetak. International Copyright,©° U. S. Department of Energy, 1987, under the provisions of the Universal Copyright Convention. United States copyright 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. DOE/EV00703-T1-Vol. | (DE87006110). DOE/EV00703-T1-Vol. II (DE87006111). Includes indexes. 1. Natural history—Marshall Islands—Enewetak Atoll. I. Devaney, Dennis M. Il. United States. Dept. of Energy. Office of Scientific and Technical Information. QH198.E53N38 1987 508.96'83 87-24863 ISBN 0-87079-579-1 (set) ISBN 0-87079-580-5 (pbk. : set) ISBN 0-87079-581-3 (microfiche : set) Work performed under contract No. DE-AC08-76EV00703 The set of two volumes is available as DE87006110 (DOE/EV/0073-T1-Vol.1) and DE87006111 (DOE/EV/0073-T1-Vol.2) from NTIS Energy Distribution Center P. O. Box 1300 Oak Ridge, Tennessee 37831 Price Code: Paper Copy A99 Microfiche AO1 Printed in the United States of America 1987 Foreword As activity and funding at the Mid-Pacific Research Laboratory began to diminish in the early 1980s, it seemed fitting that a synthesis be prepared of the three decades of research that had been conducted at this Laboratory on Enewetak Atoll. For 30 years the Atoll served as a convenient, accessible location for studies of Mid-Pacific island ecosystems, and several hundred scientists utilized the facility. Primary fund- ing was provided by the Office of Health and Environmental Research, Ecological Research Division, U. S. Department of Energy (formerly the Atomic Energy Commission and the Energy Research and Development Administration). This is an attempt to synthesize in two 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 their research. Considerable time and effort were expended by many contributors to make this synthesis possible. Thanks are extended to all these authors for their manuscripts. Special appreciation is expressed for Dr. 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- tence of Dr. Ernst Reese in organizing and collecting the material. Deepest gratitude is acknowledged for his conscientious efforts. Helen M. McCammon, Director Ecological Research Division Office of Health and Environmental Research United States Department of Energy vii — —— )3=«6 i ee y= . ‘ ws y Cie : ay | ae Des Pua ©, : “ =< de vo ~~ fegirn! we ae = a = S Acknowledgments Many people have contributed in many ways to the production of these two volumes. Regardless of the nature of the contribution, everyone listed below has given thought and time, that most precious com- modity of thinking individuals, to bring The Natural History of Enewetak Atoll into publication. Authors of the chapters are not listed separately, even though, in most cases, they critically read other chapters. No doubt we have overlooked many who have contributed in important ways, and for these oversights we apologize. To all of you we wish to extend our deepest thanks. Donald P. Abbott Isabella A. Abbott Hazel K. Asher George H. Balazs Jerry L. Barnard Frederick M. Bayer Henry R. Bennett Richard A. Boolootian Thomas E. Bowman Harry U. Brown Fenner A. Chace, dr. G. Arthur Cooper Edward B. Cutler Mae G. De Rego Maxwell S. Doty lraneus Eibl-Eibesfeldt Robert E. Elbel William O. Forster Vicki S. Frey John T. Harrison Janet F. Heavenridge Derral Herbst Robert W. Hiatt Lipke B. Holthuis Richard Houbrick Arthur Humes Edwin Janss Robert E. Johannes Victor R. Johnson, dr. Irene D. Keller Phillip B. Lamberson Jere Lipps Francois Mautin Helen M. McCammon Ellen Moore William Newman Jeri-Lyn Palacio David Pawson William F. Perrin Marian Pettibone Colin S. Ramage Anita Savacool Hajo Schmidt Stephen V. Smith Sy Sohmer William J. Stanley Lyn Sweetapple Lori N. Yamamura =r Lae toe j ar ie 1 ae ; Prax lit 7 ray iy) il , ie Tv y arau i nt ite f if ; vil Hibippeietie i ai J >=. WR rw ee} i] = e iia - ay j iim ] a) pe a , " = my 1 J > Weipa & iz a She Fy { a4 oo Us i i. ‘ ai is Si f f i a : sph Lonel) iaaj oh ad ahd 't phat ie ' ' Si ' iia jar rT jeu BAe pieas)iG)* . dy i : y The inves? _$ + ay : ; Tre ’ = Vian) if j 1 pee + oak ‘ ; Ane 1" I - ‘ ecetf kX ipa y! j ( in Ae s r 7 a "7 > i , ’ \ wu, 6 Mahal | Te 2 a i gard) ! yi) ad om : Kah 7 te | ee ‘ 2) iv — Re all h Nerey’ ab a ib ; Lal a e j ic Goa | - Preface The two volumes of The Natural History of Enewetak Atoll summarize research done at the Mid-Pacific Research Laboratory from 1954 to 1984 under the auspices of the Department of Energy. The history of the laboratory and the reasons for its support by the United States Depart- ment of Energy are described in Chapter 1 of Volume 1. Over a thousand persons—established scientists, their assistants, and graduate students—conducted research at the laboratory during the 30-year period. Their efforts resulted in 223 publications. These have been collected in four volumes of reprints entitled Mid-Pacific Marine Labora- tory Contributions, 1955-1979, U. S. Department of Energy, Publication NVO 628-1. The laboratory has con- tinued operation on a limited scale to the present. A col- lection of papers recently appeared in the Bulletin of Marine Science, Volume 38, 1986. Much of the research conducted at the laboratory was on the marine environment. The reason was that the majority of scientists applying to work at Enewetak were marine biologists. For many, this was the first opportunity to study the biota of a coral atoll. Fewer studies were con- ducted in the terrestrial environment and its biota. Nevertheless, as these volumes attest, the coverage is amazingly 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 and intertidal environments and ecology, and those on reef processes and trophic relationships, summarize a great diversity of research carried out by many scientists for many years. In contrast, the chapters on meteorology and oceanography summarize research carried out under one integrated program involving fewer scientists working over a shorter period. Volume II of The Natural History of Enewetak Atoll provides information on the taxonomy of animals and plants known to occur at Enewetak Atoll. This taxonomy represents a fulfillment of one of the first assignments to the laboratory—to determine the scientific names of the biota of the atoll. The collections on which the checklists in each chapter are based are housed at the Bernice P. xi Bishop Museum in Honolulu and the U. S. National Museum of Natural History, Smithsonian Institution, Wash- ington, D. C. In addition to the species checklists, each chapter in Volume Il 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 expect to find on equivalent Indo-Pacific islands. This com- pendium of taxonomic 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 Eniwetok 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 were ob- literated by nuclear explosions. Currently there are 39 rec- ognizable islands, and these are shown on the map used throughout the book. These volumes do not report on the extensive radiolog- ical 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 auspices of the U. S. Department of Energy. Dennis M. Devaney, senior editor of this volume, disap- peared while collecting specimens off the Island of Hawaii on August 13, 1983. Dennis was doing what he loved best, collecting marine invertebrates, at the time of his death. He collected extensively at Enewetak, and he under- took the task of organizing the systematic chapters of Volume II. Beatrice L. Burch, Devaney’s assistant at the Bishop Museum, completed the task, and she has written the introduction 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 xii of human history. In a small way, this book stands as something good that has resulted from those years. Ernst S. Reese Professor of Zoology University of Hawaii, Honolulu Contributors A. Charles Arneson Scripps Institution of Oceanography, La Jolla, California Julie H. Bailey-Brock University of Hawaii, Honolulu, Hawaii Andrew J. Berger University of Hawaii, Honolulu, Hawaii A. J. Bruce Northern Territory Museum of Arts and Sciences Darwin, N. T., Australia Beatrice L. Burch B. P. Bishop Museum, Honolulu, Hawaii Elizabeth H. Chave University of Hawaii, Honolulu, Hawaii C. Allan Child Smithsonian Institution, Washington, D. C. Robert S. Cox University of Michigan, Ann Arbor, Michigan Roger J. Cuffey Pennsylvania State University, University Park, Pennsylvania Bertha M. Cutress University of Puerto Rico, Mayagiiez, Puerto Rico Charles E. Cutress University of Puerto Rico, Mayagiiez, Puerto Rico Dennis M. Devaney (Deceased) B. P. Bishop Museum, Honolulu, Hawaii Paul H. Dunn Riverside Fire Laboratory, Riverside, California Lucius G. Eldredge University of Guam, Mangilao, Guam John S. Garth University of Southern California, Los Angeles, California Ray P. Gerber St. Joseph’s College, North Windham, Maine Richard E. Grant Smithsonian Institution, Washington, D. C. Janet Haig University of Southern California, Los Angeles, California xili Scott Johnson University of Hawaii at Manoa, Honolulu, Hawaii E. Alison Kay University of Hawaii at Manoa, Honolulu, Hawaii Louis S. Kornicker Smithsonian Institution, Washington, D. C. Jens W. Knudsen Pacific Lutheran University, Tacoma, Washington Janet O. Lamberson U. S. Environmental Protection Agency, Newport, Oregon Judith C. Lang University of Texas at Austin, Austin, Texas Raymond B. Manning Smithsonian Institution, Washington, D. C. Gordon M. Nishida B. P. Bishop Museum, Honolulu, Hawaii Helen A. Randall B. P. Bishop Museum, Honolulu, Hawaii John E. Randall B. P. Bishop Museum, Honolulu, Hawaii Marjorie L. Reaka University of Maryland, College Park, Maryland Ernst S. Reese University of Hawaii at Manoa, Honolulu, Hawaii Don Reynolds Natural History Museum, Los Angeles, California Francis W. E. Rowe The Australian Museum, Sydney, Australia G. Allan Samuelson B. P. Bishop Museum, Honolulu, Hawaii Richard H. Titgen B. P. Bishop Museum, Honolulu, Hawaii Roy T. Tsuda University of Guam, Mangilao, Guam eo ie i ’ - - co =a — =i oe’ : ib T_ Aaa ee a ae A : Va 7 a = 9 ; 7 a vaya —— : rene, Pie tw eo AL a ae ie ae 7 . i . . 1 wes -_ We : > Pe lls) CLD ate cis” ‘av a ies aad <9 ~ in i & a - © ay~ - , } ee 7 , 78 Contents Chapter Introduction Beatrice L. Burch Marine Benthic Algae of Enewetak Atoll Roy T. Tsuda Fungi of Enewetak Atoll Paul H. Dunn and Don Reynolds Natural History of Terrestrial Vascular Plants of Enewetak Atoll Janet O. Lamberson Recent Foraminifera and Nonplanktonic Protozoans E. H. Chave and Dennis M. Devaney Porifera of Enewetak Atoll Dennis M. Devaney Sea Anemones of Enewetak Atoll C. E. Cutress and A. C. Arneson Octocorallia of Enewetak Atoll Dennis M. Devaney Scleractinia (Stony Corals) of Enewetak Atoll Dennis M. Devaney and Judith C. Lang Brachiopods of Enewetak Atoll Richard E. Grant Reef-Dwelling Bryozoans of Enewetak Atoll Roger J. Cuffey and Robert S. Cox Sipunculans and Echiurans of Enewetak Atoll Dennis M. Devaney Platyhelminthes, Nemertea, and Nematoda of Enewetak Atoll Dennis M. Devaney Polychaetes of Enewetak Atoll Dennis M. Devaney and Julie H. Bailey-Brock Mollusca of Enewetak Atoll E. Alison Kay and Scott Johnson Insects and Allies (Arthropoda) of Enewetak Atoll G. Allen Samuelson and Gordon M. Nishida Pycnogonida of Enewetak Atoll C. Allan Child Stomatopod Crustacea of Enewetak Atoll Marjorie L. Reaka and Raymond B. Manning Cirripedia of Enewetak Atoll Richard H.Titgen XV 105 147 179 181 191 Chapter 19 20 21 22 23 24 25 26 “ad, 28 29 30 31 Nonplanktonic Copepoda of Enewetak Atoll Dennis M. Devaney Lagoon Plankton of Enewetak Atoll Ray P. Gerber Ostracoda (Myodocopina) of Enewetak Atoll Louis S, Kornicker Crustacea Decapoda (Penaeidea, Stenopodidea, Caridea, and Palinura) of Enewetak Atoll Dennis M. Devaney and A. J. Bruce Crustacea Decapoda (Brachyura and Anomura) of Enewetak Atoll John S. Garth, Janet Haig, and Jens W. Knudsen Holothurians of Enewetak Atoll Bertha M. Cutress and Francis W. E. Rowe Echinodermata Other than Holothurians of Enewetak Atoll Dennis M. Devaney Protochordates of Enewetak Atoll Lucius G, Eldredge Annotated Checklist of the Fishes of Enewetak Atoll and Other Marshall Islands John E. Randall and Helen A. Randall Reptiles of Enewetak Atoll Janet O. Lamberson Birds of Enewetak Atoll Andrew J. Berger Mammals of Enewetak Atoll Ernst S. Reese Miscellaneous Species Records of Enewetak Atoll Richard H. Titgen and Beatrice L. Burch Author Index Subject Index xvi 345 347 Introduction Beatrice L. Burch B. P. Bishop Museum, Honolulu, Hawaii 96817 The organization and coordination of the taxonomic section of this volume was initiated by Dr. Dennis M. Devaney of the Bernice P. Bishop Museum and was com- pleted by Beatrice L. Burch after Dr. Devaney died in a tragic scuba-diving accident on August 13, 1983, as he was investigating shrimp offshore from the Big Island of Hawaii. His great interest in all invertebrates in the tropics was increased when the opportunity arose for him to work at the Mid-Pacific Research Laboratory at Enewetak Atoll. Devaney made his first collecting trip to the atoll in the early 1960s while he was still in graduate school. As the collection grew and taxonomy of the organisms became better known, it was soon apparent that the reference col- lection at Enewetak was becoming increasingly valuable. Devaney was pleased to participate in the Coral Reef Workshop held at the atoll in 1976, because he believed that the scleractinian coral collection was the key for the study of other organisms. The workshop brought together international coral specialists to establish species limits on this important and variable group. After the workshop was held, reference material from Enewetak was deposited in European and American museums for ready reference by a wider audience of scientists. Each year after the Coral Reef Workshop, Devaney went to Enewetak to curate the reference collection and to conduct his own research on echinoderms. At the same time, he encouraged the work of specialists to compile taxonomic and other research from Enewetak for this publication. The diversity of the organisms at Enewetak made it difficult to find specialists to study all groups, so Devaney prepared several chapters himself. Unfortunately, most groups were collected in the course of other work such as physiology, toxicity, etc., and were not extensively col- lected by specialists for a particular taxonomic group. The number of families, genera, and species reported in this volume either from the literature or from new records determined by the authors of this volume are presented in Table 1. References in this volume show that some or much work was done on a particular taxon. Many groups remain XVII TABLE 1 Taxonomic Groups at Enewetak Atoll No. of No. of No. of Taxon species genera _ families Algae 238 106 40 Fungi 112 58 18 Vascular plants 123 97 48 Forams and nonplanktonic protozoa 279 144 58 Porifera 40 33 26 Actiniaria 27 21 14 Octocorallia 31 17 12 Scleractinia 169 53 12 Brachiopoda 4 4 4 Bryozoa 84 61 39 Sipuncula 11 ve 3 Echiura 2 2 2 Platyhelminthes 31 11 10 Nemertea 1 1 1 Nematoda 1 1 1 Polychaeta 132 110 34 Mollusca (fossil, recent) 1240 453 151 Insects and related arthropods 190 157 9S Pycnogonida 5 4 4 Stomatopoda 12 4 4 Cirripedia 10 7 6 Lagoon plankton 285 177 82 Ostracoda 10 10 5 Natantia 145 56 14 Reptantia 4 3 3 Anomura 76 29 10 Brachyura 293 114 16 Holothuroidea 20 11 5 Echinodermata other than Holothuroidea 97 65 32 Fishes of the Marshall Islands 815 338 92 Reptilia 9 9 5 Aves 41 ii 12 Mammalia g) 7 6 Miscellaneous 124 87 40 Totals 4671 2284 902 to be worked on more thoroughly by specialists in particu- lar fields, such as Porifera or Tunicata, which, at the present time, seem to be represented so lightly at Enewetak Atoll. By having a named reference collection, the researchers there were able to identify organisms used in their studies on biochemistry, ecology, productivity, animal or plant associations, physiology, immunology, radiobiology, growth rates, and reproduction. They were also able to make broad interpretations of reef chronology, xviii geochemistry, stratigraphy, and biogeographic distribution. The checklist contained in each chapter has a coded entry symbol placed before the generic designation to indi- cate (1) if the organism represents a newly recorded species for Enewetak or for the Marshall Islands, (2) if it is a fossil record, or (3) if it has some other reason to be so marked. The explanations for these codes follow each species checklist. Chapter 1 Marine Benthic Algae of Enewetak Atoll ROY T. TSUDA Marine Laboratory, University of Guam UOG Station, Mangilao, Guam 96923 Our present knowledge of the floristics and ecology of the marine benthic algae on Pacific atolls is based primarily on studies conducted on Enewetak Atoll. There are more species of marine benthic algae known from this atoll than are known from any other Indo-Pacific atoll. The first published account of the marine benthic algae of Enewetak appeared in Taylor’s (1950b) treatise of the plants of Bikini, which was part of a comprehensive study of the flora and fauna of Bikini Atoll and other adjacent atolls conducted in 1946 prior to the atomic bomb tests of “Operation Crossroads.” The phycological study was not only the first for these atolls but also represented the pioneering study in the Marshall Islands. Descriptive and illustrative records of 57 species were included from Enewetak. Three new species of the green algal genus Halimeda were described based on Enewetak specimens as holotypes (H. fragilis Taylor, H. gigas Taylor, and H. lacunalis Taylor). Dawson (1957) made extensive collections of marine algae on Enewetak during the summer of 1955 under the auspices of the Eniwetok Marine Biological Laboratory (now called the Mid-Pacific Marine Laboratory). His objec- tive was to provide the laboratory with an algal reference collection. The extensive collections made by Ralph F. Palumbo of the Applied Fisheries Laboratory, University of Washington, in relation to his radiobiological survey in the Marshalls were also incorporated in the floristic account of the 211 species reported by Dawson (1957). A few of these species were mentioned in technical reports of the U. S. Atomic Energy Commission by Palumbo (1950, 1955, 1959) and a later paper (Lowman and Palumbo, 1962}. Based on the Enewetak specimens as holotypes, five new red algal species were described: Antithamnion percurrens Dawson, Callithamnion marshallensis Dawson, Ceramium marshallense Dawson, C. sympodiale Dawson, and C. vagabunde Dawson. The first ecological study on deep water algae in an atoll lagoon was conducted on Enewetak by Gilmartin (1960). During 1955 and 1956, he ran a transect across the lagoon in waters ranging from 20 to 65 m deep and recorded, as well as collected, algae he observed at the various depths. A total of 87 species was reported: 16 species represented new records not previously reported by Taylor (1950b) or Dawson (1957). Further algal studies which mention specimens from Enewetak were mainly monographic treatments of specific algae. Records of blue-green algae from Enewetak can be found in monographs by Drouet and Daily (1956) and Drouet (1968, 1973), who drastically revised the classifica- tion scheme. Drouet’s classification scheme is followed in this presentation, and it is coincidental that he identified all the blue-green algae reported by Taylor (1950b), Dawson (1957), and Gilmartin (1960). Other algal genera studied include records of Halimeda (Hillis, 1959), Turbinaria (Taylor, 1964), Tydemania (Gil- martin, 1966), Dawsoniella (Hollenberg, 1967), Polysi- phonia (Hollenberg, 1968a, 1968b), Herposiphonia (Hollen- berg, 1968c), Lophosiphonia (Hollenberg, 1968d), Fostliella on sea urchin spines (Lawrence and Dawes, 1969), and Caulerpa (Calvert et al., 1976). Polysiphonia pentamera Hollenberg (1968b) was described as a new species from Enewetak Atoll. A recent study on the distribution of Halimeda and Tydemania on Enewetak by Hillis-Colinvaux (1977) added two more species records of Halimeda to the flora. In the last chapter of the exhaustive review of Halimeda by Hillis-Colinvaux (1980), the habitats occupied by Halimeda at Enewetak are described in detail. In a paper presented at the 63 Annual Meeting of the Western Society of Naturalists held at the California State University of Long Beach, Hillis-Colinvaux (1982) de- scribed the large populations of Halimeda down to 100 m and growth to over 140 m on the windward and leeward outer slopes of Enewetak. The submersible “Makali’i” was used for observations. Aside from the above papers, several ecological studies mention algal species. Marsh (1970) carried out studies on the primary productivity of crustose coralline algae and later conducted studies on the productivity of the reef community as a whole (Smith and Marsh, 1973). Other studies which mention algae are those of Odum and Odum (1955), Bakus (1967), Johannes et al. (1972), and Gerber and Marshall (1974). Calothrix crustacea Schousboe and Thuret and Hormothamnion enteromorphoides B. and FI. 2 TSUDA were heterocystous blue-greens studied by Wiebe et al. (1975), Webb and Wiebe (1975), Webb et al. (1975), and Wiebe (1976) in their studies on the nitrogen cycle. On the basis of the papers cited above, 238 species (106 genera) of marine benthic algae are known from TABLE Enewetak Atoll: Cyanophyta (16 species), Chlorophyta (89 species), Phaeophyta (24 species), and Rhodophyta (109 species). Nine of the species (preceded by a dagger in Table 1) were described as new with the Enewetak speci- mens serving as holotypes. It is interesting to note that 40 1 Checklist of Marine Benthic Algae of Enewetak Division CYANOPHYTA (blue-green algae) Class CYANOPHYCEAE Order CHROOCOCCALES Family CHAMAESIPHONACEAE Entophysalis conferta (Kiitz.) Dr. and Daily: Dawson, 1957. Entophysalis deusta (Menegh.) Dr. and Daily: Drouet and Daily, 1956; Dawson, 1957. Family CHROOCOCCACEAE Anacystis dimidiata (Kiitz.) Dr. and Daily: Dawson, 1957. Coccochloris stagnina Sprengel: Dawson, 1957. Gomphosphaeria aponina Kitz.: Taylor, 1950b; Drouet Order OSCILLATORIALES Family NOSTOCACEAE and Daily, 1956; Dawson, 1957. Calothrix crustacea Schousboe and Thuret: Dawson, 1957; Bakus, 1967; Drouet, 1973; Webb et al., 1975; Webb and Wiebe, 1975; Wiebe et al., 1975; Wiebe, 1976. Rivularia atra Roth: Dawson, 1957. Rivularia polyotis (J. Ag.) B. and Fl.: Dawson, 1957. Hormothamnion enteromorphoides B. and FI.: Taylor, 1950b: Dawson, 1957; Gilmartin, 1960; Webb et al., 1975; Wiebe et al., 1975. Scytonema hofmanii Ag. Scytonema myocyrous (Dillw.) C. Ag.: Dawson, 1957. Scytonema polycystum B. and Fl.: Dawson, 1957; Gilmartin, 1960. Family OSCILLATORIACEAE Microcoleus lyngbyaceus (Kiitz.) Crouan. Hydrocoleum cantharidosmum (Mont.) Gomont: Taylor, 1950b. Hydrocoleum coccineum Gomont: Dawson, 1957. Hydrocoleum comoides (Harv.) Gomont: Dawson, 1957; Gilmartin, 1960. Hydrocoleum glutinosum (C. Ag.) Gomont: Dawson, 1957; Gilmartin, 1960. Hydrocoleum lyngbyaceum Kiitz.: Dawson, 1957. Lyngbya aestuarii (Mert.) Lyngb.: Dawson, 1957. Lyngbya confervoides C. Ag.: Taylor, 1950b; Dawson, 1957. Lyngbya majuscula (Dillw.) Harv.: Dawson, 1957; Gilmartin, 1960. Lyngbya meneghiniana (Kiitz.) Gomont: Taylor, 1950b; Dawson, 1957. Lyngbya semiplena (Ag.) J. Ag.: Dawson, 1957. Oscillatoria submembranacea Ard. and Straff: Drouet, 1968. Phormidium penicillatum Gomont: Dawson, 1957. Porphyrosiphon notarisii (Menegh.) Kiitz. Oscillatoria nigro-viridis Thw.: Dawson, 1957. Schizothrix arenaria (Berkeley) Gomont. Symploca laete-viridis Gomont: Dawson, 1957; Gilmartin, 1960. Schizothrix calcicola (C. Ag.) Gomont: Bakus, 1967; Drouet, 1968; Johannes et al., 1972; Webb and Wiebe, 1975. Phormidium crosbyanum Tilden: Taylor, 1950b; Dawson, 1957. Plectonema nostocorum Bornet: Taylor, 1950b; Dawson, 1957. Plectonema terebrans B. and Fl.: Dawson, 1957. Schizothrix lacustris A. Br.: Dawson, 1957. Schizothrix mexicana Gomont: Drouet, 1968. Lyngbya gracilis (Menegh.) Rabenh.: Taylor, 1950b; Dawson, 1957. Lyngbya sordida (Zanard.) Gomont: Dawson, 1957; Gilmartin, 1960. Symploca hydnoides (Harv.) Kiitz: Dawson, 1957; Gilmartin, 1960. Schizothrix tenerrima (Gomont) Dr.: Drouet, 1968. Microcoleus tenerrimus Gomont: Dawson, 1957. Spirulina subsalsa Oersted: Dawson, 1957. Spirulina major Kiitz.: Dawson, 1957. Spirulina tenerrima Kiitz.: Dawson, 1957. (This table continued on next page.) MARINE BENTHIC ALGAE TABLE 1 (cont’d) Division CHLOROPHYTA (green algae) Class CHLOROPHYCEAE Order CHAETOPHORALES Family CHAETOPHORACEAE *Entocladia viridis Reinke: Dawson, 1957. Order ULVALES Family ULVACEAE *Enteromorpha acanthophora Kiitz.: Dawson, 1957. Enteromorpha clathrata (Roth) J. Ag.: Dawson, 1957. Enteromorpha intestinalis (L.) Link: Dawson, 1957. Enteromorpha kylinii Bliding: Dawson, 1957. Enteromorpha lingulata J. Ag.: Taylor, 1950b. Enteromorpha ralfsii Harvey: Dawson, 1957. *Enteromorpha tubulosa (Kiitz.) Kiitz.: Dawson, 1957. Order CAULERPALES Family BRYOPSIDACEAE Bryopsis hypnoides Lamx.: Dawson, 1957. Bryopsis indica A. and E. S. Gepp: Taylor, 1950b; Dawson, 1957. Bryopsis pennata Lamx.: Dawson, 1957; Gilmartin, 1960. Family CAULERPACEAE Caulerpa ambigua Okamura Caulerpa vickersiae Boerg.: Dawson, 1957; Gilmartin, 1960. Caulerpa antoensis Yamada: Dawson, 1957. Caulerpa bikinensis Taylor: Dawson, 1957. Caulerpa brachypus Harv.: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960. Caulerpa elongata W. v. Bosse: Odum and Odum, 1955; Dawson, 1957; Calvert et al., 1976. Caulerpa filicoides Yamada: Dawson, 1957. Caulerpa acuta (Yamada) Yamada: Gilmartin, 1960. Caulerpa peltata Lamx.: Gilmartin, 1960. Caulerpa racemosa (Forsk.) J. Ag.: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960. Caulerpa serrulata (Forsk.) J. Ag.: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960; Calvert et al., 1976. Caulerpa sertularioides (Gmel.) Howe: Gilmartin, 1960. Caulerpa taxifolia (Vahl) C. Ag.: Dawson, 1957. Caulerpa urvilliana Montagne: Taylor, 1950b; Dawson, 1957; Calvert et al., 1976. Caulerpa verticillata J. Ag.: Gilmartin, 1960. Caulerpa webbiana Montagne: Dawson, 1957; Gilmartin, 1960; Calvert et al., 1976. Family CODIACEAE Aorainvillea lacerata Harv.: Taylor, 1950b; Dawson, 1957. Aorainvillea nigricans Decaisne: Gilmartin, 1960. Codium arabicum Kiitz.: Dawson, 1957. Codium edule Silva: Dawson, 1957. Codium geppii O. C. Schmidt: Dawson, 1957; Gilmartin, 1960. *Codium saccatum Okamura: Dawson, 1957. Codium tenue Kiitz.: Taylor, 1950b; Dawson, 1957. Halimeda copiosa Goreau and Graham: Hillis-Colinvaux, 1977. Halimeda opuntia f. hederacea Barton: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960. Halimeda cylindracea Decaisne: Hillis-Colinvaux, 1977. Halimeda monile (Solander) Lamx.: Dawson, 1957; Gilmartin, 1960, 1966. Halimeda discoidea Decaisne: Gilmartin, 1960. Halimeda distorta (Yamada) Colinvaux: Hillis-Colinvaux, 1977. Halimeda fragilis Taylor: Taylor, 1950b; Hillis, 1959; Gilmartin, 1960, 1966; Hillis-Colvinaux, 1977.+ +Halimeda gigas Taylor: Taylor, 1950b; Dawson, 1957; Hillis, 1959; Hillis-Colinvaux, 1977. Halimeda gracilis Harv.: Gilmartin, 1960; Hillis-Colinvaux, 1977. Halimeda incrassata (Ellis) Lamx.: Hillis-Colinvaux, 1977. Halimeda tridens (E. and S.) Lamx.: Gilmartin, 1960. Halimeda tridens f. lamourouxii (J. Ag.) W.v. Bosse: Taylor, 1950b; Dawson, 1957. Halimeda tridens f. tripartita (Barton) Collins: Taylor, 1950b. Halimeda tridens f. typica (Barton) Collins: Taylor, 1950b. “Only specimen recorded from Micronesia. +New species; holotype based on Enewetak specimens. (This table continued on next page.) TSUDA TABLE 1 (cont'd) Family CODIACEAE (cont’d) tHalimeda lacunalis Taylor: Taylor, 1950b; Dawson, 1957; Hillis, 1959; Gilmartin, 1960; Hillis-Colinvaux, 1977. Halimeda macrophysa Askenasy: Dawson, 1957; Hillis-Colinvaux, 1977. Halimeda micronesica Yamada: Hillis-Colinvaux, 1977. Halimeda minima (Taylor) Colinvaux: Hillis-Colinvaux, 1977. Halimeda opuntia f. minima Taylor: Dawson, 1957; Gilmartin, 1960. Halimeda opuntia (L.) Lamx.: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960; Hillis-Colinvaux, 1977. Halimeda stuposa Taylor: Taylor, 1950; Dawson, 1957; Hillis-Colinvaux, 1977. Halimeda taenicola Taylor: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960; Hillis-Colinvaux, 1977. Pseudochlorodesmis furcellata (Zanard.) Boerg.: Dawson, 1957; Gilmartin, 1960. Rhipilia diaphana Taylor: Taylor, 1950b; Dawson, 1957. Rhipilia geppii Taylor: Taylor, 1950b; Dawson, 1957. Rhipilia orientalis A. and E. S. Gepp: Dawson, 1957. Tydemania expeditionis W. v. Bosse: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960, 1966; Hillis-Colinvaux, 1977. Udotea indica A. and E. S. Gepp: Taylor, 1950b; Gilmartin, 1960. Udotea javensis (Mont.) A. and E. S. Gepp: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960. Udotea orientalis A. and E. S. Gepp: Gilmartin, 1960. Udotea palmetta Decaisne: Dawson, 1957. Family DERBESIACEAE Derbesia attenuata Dawson: Dawson, 1957. Derbesia marina (Lyngb.) Solier: Dawson, 1957. “Derbesia neglecta Reinbold: Gilmartin, 1960. Derbesia ryukiuensis Yamada and Tanaka: Dawson, 1957. Family PHYLLOSIPHONACEAE Ostreobium reineckii Bornet: Dawson, 1957. Order SIPHONOCLADALES Family BOODLEACEAE Boodlea composita (Harv.) Brand: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960. Boodlea siamensis Reinbold: Taylor, 1950b; Dawson, 1957. Struvea anastomosans (Harv.) Piccone: Dawson, 1957. Family SIPHONOCLADACEAE Cladophoropsis gracillima Dawson: Dawson, 1957; Gilmartin, 1960. Cladophoropsis sundanensis Reinbold: Dawson, 1957; Gilmartin, 1960. Siphonocladus rigidus Howe: Dawson, 1957. Family VALONIACEAE Boergesenia forbesii (Harv.) Feldmann: Dawson, 1957. Dictyosphaeria cavernosa (Forsk.) Boerg.: Taylor, 1950b; Dawson, 1957; Webb et al., 1975. Dictyosphaeria intermedia W. v. Bosse: Taylor, 1950b; Odum and Odum, 1955; Dawson, 1957. Dictyosphaeria versluysii W. v. Bosse: Dawson, 1957. Valonia aegagropila C. Ag.: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960. Valonia fastigiata Harv.: Dawson, 1957. Valonia utricularis (Roth) C. Ag.: Dawson, 1957; Gilmartin, 1960. Valonia ventricosa J. Ag.: Dawson, 1957; Gilmartin, 1960. Valoniopsis pachynema (Martens) Boerg.: Taylor, 1950b; Dawson, 1957. Order DASYCLADALES Family DASYCLADACEAE Acetabularia moebii Solms-Laubach: Dawson, 1957; Gilmartin, 1960. Neomeris bilimbata Koster: Dawson, 1957. Order CLADOPHORALES Family ANADYOMENACEAE Anadyomene wrightii Gray: Dawson, 1957. Microdictyon japonicum Setch.: Dawson, 1957. Microdiciyon okamurai Setch.: Taylor, 1950b; Dawson, 1957. Rhipidiphyllon reticulatum Askenasy: Taylor, 1950b; Dawson, 1957. Family CLADOPHORACEAE Chaetomorpha indica Kiitz.: Dawson, 1957. *Cladophora albida (Hudson) Kiitz.: Gilmartin, 1960. Cladophora crystallina (Roth) Kiitz.: Dawson, 1957. *Cladophora gracilis (Griffiths) Kiitz.: Gilmartin, 1960. *Only specimen recorded from Micronesia. +New species; holotype based on Enewetak specimens. (This table continued on next page.) MARINE BENTHIC ALGAE TABLE 1 (cont'd) Family CLADOPHORACEAE (cont'd) *Cladophora inserta Dickie: Dawson, 1957. Cladophora socialis Kiitz.: Dawson, 1957; Gilmartin, 1960. *Rhizoclonium implexum (Dillwyn) Kiitz.: Dawson, 1957. Division PHAEOPHYTA (brown algae) Class PHAEOPHYCEAE Order ECTOCARPALES Family ECTOCARPACEAE Ectocarpus breviarticulatus J. Ag.: Dawson, 1957. Ectocarpus mitchellae Harv.: Taylor, 1950b; Dawson, 1957. Feldmannia indica (Sonder) Womersley and Bailey. Ectocarpus indicus Sonder: Taylor, 1950b; Dawson, 1957. Feldmannia irregularis (Kiitz.) Hamel. Ectocarpus irregularis Kiitz.: Dawson, 1957. Order SPHACELARIALES Family SPHACELARIACEAE Sphacelaria furcigera Kiitz.: Dawson, 1957; Gilmartin, 1960. Sphacelaria novaehollandiae Sonder: Taylor, 1950b; Dawson, 1957. Sphacelaria tribuloides Menegh.: Dawson, 1957. Order DICTYOTALES Family DICTYOTACEAE Dictyopteris repens (Okam.) Boerg.: Dawson, 1957; Gilmartin, 1960, 1966. Dictyota cervicornis Kiitz.: Gilmartin, 1960. Dictyota dichotoma Lamx.: Gilmartin, 1960. Dictyota divaricata Lamx.: Dawson, 1957; Gilmartin, 1960; Lowman and Palumbo, 1962; Gilmartin, 1966. Dictyota patens J. Ag.: Dawson, 1957. Dictyota pinnatifida Kiitz.: Taylor, 1950b; Dawson, 1957. Lobophora papenfussii (Taylor) Womersley. Pocockiella papenfussii Taylor: Taylor, 1950b; Dawson, 1957. Lobophora variegata (Lamx.) Womersley: Smith and Marsh, 1973. Pocockiella variegata (Lamx.) Papenfuss: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960. Zonaria variegata (Lamx.) Mertens: Odum and Odum, 1955. Padina australis Hauck: Dawson, 1957. Padina tenuis Bory. Padina commersonii Bory: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960, 1966. * Spatoglossum schroederi (Mert.) J. Ag.: Gilmartin, 1960. Order SCYTOSIPHONALES Family SCY TOSIPHONACEAE Hydroclathrus clathratus (Bory) Howe: Taylor, 1950b; Dawson, 1957. * Rosenvingea fastigiata (Zanard.) Boerg.: Taylor, 1950b; Dawson, 1957. Rosenvingea intricata (J. Ag.) Boerg.: Taylor, 1950b; Dawson, 1957. *° Rosenvingea orientalis (J. Ag.) Boerg.: Gilmartin, 1960. Order DICTYOSIPHONALES Family CHNOOSPORACEAE Chnoospora implexa (Hering) J. Ag.: Dawson, 1957. Order FUCALES Family SARGASSACEAE Turbinaria ornata (Turn.) J. Ag.: Taylor, 1950b; Dawson, 1957, 1964. Division RHODOPHYTA (red algae) Class BANGIOPHYCEAE Order BANGIALES Family ERYTHROPELTIDACEAE Erythrotrichia carnea (Dillwyn) J. Ag.: Taylor, 1950b; Dawson, 1957. Erythrotrichia parietalis Tanaka: Dawson, 1957. Order PORPHYRIDIALES Family GONIOTRICHACEAE Asterocystis ornata (C. Ag.) Hamel: Taylor, 1950b; Dawson, 1957. Goniotrichum alsidii (Zanard.) Howe: Taylor, 1950b. Goniotrichum elegans (Chauvin) Zanard.: Dawson, 1957. *Only specimen recorded from Micronesia. (This table continued on next page.) TSUDA TABLE 1 (cont’d) Class FLORIDEOPHYCEAE Order NEMALIALES Family ACROCHAETIACEAE * Acrochaetium gracile Boerg.: Dawson, 1957. Acrochaetium robustum Boerg.: Dawson, 1957; Gilmartin, 1960. * Kylinia crassipes (Boerg.) Kylin: Dawson, 1957. * Kylinia secunda (Lyngb.) Papenfuss: Dawson, 1957. Family BONNEMAISONIACEAE Asparagopsis taxiformis (Delile) Collins and Hervey: Dawson, 1957; Gilmartin, 1960; Gerber and Marshall, 1974. Falkenbergia hillebrandii (Bornet) Falkenberg: Dawson, 1959; Gilmartin, 1960. Family CHAETANGIACEAE Galaxaura fastigiata Decaisne: Dawson, 1957. Galaxaura filamentosa Chou: Dawson, 1957. Family GELIDIACEAE * Gelidiella bornetii (W. v. Bosse) Feldmann and Hamel: Dawson, 1957. Gelidiella tenuissima Feldmann and Hamel: Dawson, 1957. * Gelidium crinale (Turn.) Lamx.: Dawson, 1957.* Gelidium pusillum (Stackh.) Ledolis: Dawson, 1957. Wurdemannia miniata (Lmk. and D. C.) Feldmann and Hamel: Dawson, 1957. Family HELMINTHOCLADIACEAE Liagora farinosa Lamx.: Dawson, 1957. * Liagora hawaiiana Butters: Dawson, 1957. * Liagora orientalis J. Ag.: Dawson, 1957. Liagora pinnata Harv.: Dawson, 1957. * Liagora robusta Yamada: Dawson, 1957. Order CRYPTONEMIALES Family CORALLINACEAE Fosliella farinosa (Lamx.) Howe: Dawson, 1957; Lawrence and Dawes, 1969. Heteroderma minutula Foslie: Dawson, 1957. Heteroderma subtilissima Foslie: Dawson, 1957. Jania capillacea Harv.: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960; Smith and Marsh, 1973. Jania decussato-dichotoma (Yendo) Yendo: Dawson, 1957; Gilmartin, 1960. Jania micranthrodia Lamx.: Dawson, 1957. Jania tenella Kiitz.: Dawson, 1957. Porolithon craspedium (Foslie) Foslie: Taylor, 1950b; Dawson, 1957. Porolithon gardineri (Foslie) Foslie: Taylor, 1950b; Dawson, 1957. Porolithon onkodes (Heyd.) Foslie: Taylor, 1950b; Marsh 1970; Smith and Marsh, 1973. Lithothamnion sp.: Taylor, 1950b. Family CRYPTONEMIACEAE Grateloupia filicina (Wulfen) C. Ag.: Dawson, 1957. Family PEYSSONELIACEAE Cruoriella dubyi (Crouan and Crouan) Schmidt: Dawson, 1957. Peyssonelia rubra (Crev.) J. Ag.: Dawson, 1957. Order GIGARTINALES Family DICRANEMACEAE * Dicranema rosaliae Setch. and Gard.: Dawson, 1957. Family HYPNEACEAE Hypnea esperi Bory: Dawson, 1957; Gilmartin, 1960. Hypnea nidulans Setch.: Dawson, 1957. Hypnea pannosa J. Ag.: Dawson, 1957. Hypnea spinella (C. Ag.) Kiitz.: Taylor, 1950b. Order RHODYMENIALES Family CHAMPIACEAE Champia parvula (Ag.) J. Ag.: Dawson, 1957; Gilmartin, 1960. Champia vieilardii Kiitz.: Dewson, 1957. Lomentaria hakodatensis Yendo: Dawson, 1957. Family RHODYMENIACEAE Botryocladia skottsbergii (Boerg.) Levring: Dawson, 1957; Gilmartin, 1960. Coelothrix ivregularis (Harv.) Boerg.: Dawson, 1957. Rhodymenia anastomosans W. v. Bosse: Dawson, 1957; Gilmartin, 1960. *Only specimen recorded from Micronesia. (This table continued on next page.) MARINE BENTHIC ALGAE TABLE 1 (cont'd) Order CERAMIALES Family CERAMIACEAE Antithamnion lherminieri (Crouan and Crouan) Nasr: Dawson, 1957; Gilmartin, 1960. *t+ Antithamnion percurrens Dawson: Dawson, 1957. Antithamnionella breviramosa (Dawson) Wollaston. Antithamnion breviramous Dawson: Dawson, 1957. *+ Callithamnion marshallensis Dawson: Dawson, 1957. Centroceras apiculatum Yamada: Dawson, 1957. Centroceras clavulatum (C. Ag.) Mont.: Dawson, 1957; Gilmartin, 1960. Centroceras minimum Yamada: Dawson, 1957. Ceramium clarionense Setch. and Gardn.: Dawson, 1957. Ceramium gracillimum Griff. and Harv.: Dawson, 1957; Gilmartin, 1960. *t+Ceramium marshallense Dawson: Dawson, 1957. Ceramium mazatlanense Dawson: Dawson, 1957; Gilmartin, 1960. Ceramium serpens Setch. and Gardn.: Dawson, 1957. +Ceramium sympodiale Dawson: Dawson, 1957. *Ceramium taylori Dawson: Dawson, 1957. +Ceramium vagabunde Dawson: Dawson, 1957. *Corynospora pedicellata (Smith) J. Ag. Neomonospora pedicellata v. tenuis Feldm.-Mazoyer: Dawson, 1957. Crouania attenuata (Bonnemaison) J. Ag.: Gilmartin, 1960. *Griffithsia ovalis Harvey: Dawson, 1957. Griffithsia tenuis C. Ag.: Dawson, 1957; Gilmartin, 1960. Haloplegma duperreyi Montagne: Gilmartin, 1960. * Herposiphonia nuda Hollenberg: Hollenberg, 1968c. Herposiphonia pacifica Hollenberg: Hollenberg, 1968c. Herposiphonia parca Setch.: Hollenberg, 1968c. Herposiphonia secunda (C. Ag.) Ambronn: Dawson, 1957. Herposiphonia tenella (C. Ag.) Naegeli: Dawson, 1957; Hollenberg, 1968c. Hetersiphonia wurdemanni (Bailey) Falkenberg: Dawson, 1957; Gilmartin, 1960. Lophosiphonia bermudensis Collins and Herv.: Dawson, 1957. *Lophosiphonia cristata Falkenberg: Hollenberg, 1968d. Lophosiphonia obscura (Ag.) Falkenberg: Dawson, 1957; Gilmartin, 1960. *Lophosiphonia villum (J. Ag.) Setch. and Gard.: Gilmartin, 1960. *Mesothamnion caribaeum Boerg.: Gilmartin, 1960. Polysiphonia coacta Tseng: Dawson, 1957. Polysiphonia exilis Harv.: Hollenberg, 1968b. Polysiphonia flaccidissima Hollenberg: Hollenberg, 1968a. + Polysiphonia pentamera Hollenberg: Hollenberg, 1968b. Polysiphonia poko Hollenberg: Hollenberg, 1968a. Polysiphonia setacea Hollenberg: Hollenberg, 1968a. Polysiphonia subtilissima Montagne: Dawson, 1957; Gilmartin, 1960. Polysiphonia tongatensis Harvey: Dawson, 1957; Gilmartin, 1960. Spermothamnion saccorhiza (Setch. and Gardn.) Feldmann-Mazoyer; Gilmartin, 1960. Spyridia filamentosa (Wulf.) Harv.: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960. * Taenioma perpusillum J. Ag.: Dawson, 1957. Tolypiocladia calodictyon (Harv.) Silva. Roschera calodictyon (Harv.) W. v. Bosse: Taylor, 1950b; Dawson, 1957; Gilmartin, 1960. Wrangelia anastomosans Yamada: Dawson, 1957. Wrangelia argus (Mont.) Montagne: Dawson, 1957. Wrangelia penicillata C. Ag.: Taylor, 1950b; Dawson, 1957. Family DASYACEAE Dasya adhaerens Yamada: Taylor, 1950b; Dawson, 1957. * Dasya corymbifera J. Ag.: Gilmartin, 1960. * Dasya iyengarii Boerg.: Dawson, 1957; Gilmartin, 1960. * Dasya mollis Harv.: Dawson, 1957; Gilmartin, 1960. * Dasyopsis geppii W. v. Bosse: Dawson, 1957. Dictyurus purpurascens Bory: Dawson, 1957. “Only specimen recorded from Micronesia. +New species; holotype based on Enewetak specimens. (This table continued on next page.) TSUDA TABLE 1 (cont’d) Family DELESSERIACEAE Hypoglossum minimum Yamada: Dawson, 1957; Gilmartin, 1960. Martensia fragilis Harv. Hemitrema fragilis {Harv.) Dawson: Dawson, 1957. Family RHODOMELACEAE *Chondria baileyana (Montagne) Harv.: Gilmartin, 1960. *Chondria minutula W. v. Bosse: Dawson, 1957. *Chondria polyrhiza Collins and Hervey: Dawson, 1957. Chondria repens Boerg.: Dawson, 1957; Gilmartin, 1960. * Dawsoniella bulborhiza Hollenberg: Hollenberg, 1967. Laurencia mariannensis Yamada: Taylor, 1950b; Dawson, 1957. Laurencia nana Howe: Dawson, 1957. Laurencia parvipapillata Tseng: Dawson, 1957. “Only specimen recorded from Micronesia. +Holotype based on Enewetak specimens. species (nine Chlorophyta, three Phaeophyta, and 28 Rho- dophyta) or 16% of the species reported from Enewetak represent the only collections known from the geographic region of Micronesia (preceded by an asterisk in Table 1). It is unlikely that these species are unique to Enewetak; further intensive collections from other areas in Micronesia will no doubt provide additional records. To date, there has been no record of a seagrass on Enewetak. Thalassia hemprichii (Ehrenb.) Aschers is the only species of seagrass reported from the Marshall Islands, and it has been found on Ujilang Atoll, Jaluit Atoll, and Ailinglapalap Atoll (Tsuda et al., 1977). Until further intensive collec- tions are made from other islands in Micronesia, it is diffi- cult to make a comparative analysis with the marine flora from other islands. Although it is hardly fair to compare the Enewetak marine flora with that of Guam, a high island located farther west, some inferences can be made. As can be seen in Table 2, there are slightly more genera known from Guam (118 genera) than Enewetak (106 gen- era), but there are more species reported from Enewetak. Taylor (1950a) described the dominant algae on the atoll and found it was difficult to compare the Marshall marine flora with those of other islands because of the paucity of TABLE 2 Comparison of the Marine Benthic Flora of Enewetak and Guam Genera Species Division ES Gt E° Gt Cyanophyta 1 10 16 13 Chlorophyta 29 30 89 69 Phaeophyta 12 15 24 27 Rhodophyta 53 63109 11 Total 106 118 238 120 “Enewetak. +Guam collections from other islands. He did mention that Sargassum was noticeably absent on Enewetak. This genus has yet to be found on any of the atolls in the Marshall Islands (Doty, 1954; Tsuda, 1976). Clearly, the phycologi- cal studies conducted on Enewetak Atoll have provided much of the information on the marine benthic algae within Micronesia. Locations of the early collection sites are available in the Departments of Botany and Invertebrate Zoology at the B. P. Bishop Museum. REFERENCES Bakus, G. J., 1967, The Feeding Habits of Fishes and Primary Production at Eniwetok, Marshall Islands, Micronesica, 3: 135-149. Calvert, H. E., C. J. Dawes, and M. A. Borowitzka, 1976, Phylo- genetic Relationships of Caulerpa (Chlorophyta) Based on Comparative Chloroplast Ultrastructure, J. Phycol., 12: 149-162. Dawson, E. Y., 1957, An Annotated List of Marine Algae from Eniwetok Atoll, Marshail Islands, Pac. Sci., 11: 92-132. Doty, M. S., 1954, Distribution of the Algal Genera Rhipilia and Sargassum in the Central Pacific, Pac. Sci., 8: 367-368. Drouet, F., 1968, Revision of the Classification of the Oscilla- toriaceae, Monogr. Nat. Acad. Sci. Phila., 15: 1-334. __, 1973, Revision of the Nostocaceae with Cylindrical Tri- chomes, Hafner Press, N. Y. ___, and W. A. Daily, 1956, Revision of the Coccoid Myxophy- ceae, Butler Univ. Bot. Stud., 12: 218. Gerber, R. P., and N. Marshall, 1974, Ingestion of Detritus by the Lagoon Pelagic Community of Eniwetok Atoll, Limnol. Oceanogr., 19: 815-824. Gilmartin, M., 1960, The Ecological Distribution of the Deep Water Algae of Eniwetok Atoll, Ecology, 41: 210-221. , 1966, Ecology and Morphology of Tydemania expeditionis, a Tropical Deep Water Siphonous Green Algae, J. Phycol., 2: 100-105. Hillis, L. W., 1959, A Revision of the Genus Halimeda (Order Siphonales), Publ. Inst. Mar. Sci. Univ. Texas, 6: 321-403. Hillis-Colinvaux, L., 1977, Halimeda and Tydemania: Distribution, Diversity and Productivity at Enewetak, Third International Coral Reef Symposium Proceedings (Miami), pp. 365-370. MARINE BENTHIC ALGAE 9 ——, 1980, Ecology and Taxonomy of Halimeda: Primary Pro- ducer of Coral Reefs, Adv. Mar. Biol., 17; 1-327. ——_, 1982, Deep Water Populations of Halimeda in the Economy of an Atoll, Western Society of Naturalists, 63rd Annual Meet- ing (Program/Abstract), p. 29. Hollenberg, G. J., 1967, New Genera in the Rhodomelaceae from the Central Pacific, Bull. South. Calif. Acad. Sci., 66: 201-221. ——, 1968a, An Account of the Species of Polysiphonia of the Central and Western Tropical Pacific Ocean, |. Oligosiphonia, Pac. Sci., 22: 56-98. —_, 1968b, An Account of the Species of the Red Alga Polysiphonia of the Central and Western Tropical Pacific Ocean, II. Polysiphonia, Pac. Sci., 22: 198-207. ——, 1968c, An Account of the Species of the Red Alga Herposiphonia Occurring in the Central and Western Tropical Pacific Ocean, Pac. Sci., 22: 536-559. ——, 1968d, Phycological Notes, III, New Records of Marine Algae from the Central Tropical Pacific Ocean, Brittonia, 20: 74-82. Johannes, R. E., J. Alberts, C. D’Elia, R. A. Kinzie, L. R. Pomeroy, W. Sottile, W. Wiebe, J. A. Marsh, Jr., P. Helfrich, J. Maragos, J. Meyer, S. Smith, D. Crabtree, A. Roth, L. R. McCloskey, S. Betzer, N. Marshall, M. E. Q. Pilson, G. Telek, R. I. Clutter, W. D. DuPaul, K. L. Webb, and J. M. Wells, Jr., 1972, The Metabolism of Some Coral Reef Communities: A Team Study of Nutrient and Energy Flux at Eniwetok, Bioscience, 22: 541-543. Lawrence, J. M., and C. J. Dawes, 1969, Algal Growth over the Epidermis of Sea Urchin Spines, J. Phycol., 5: 269. Lowman, F. G., and R. F. Palumbo, 1962, Occurrence of Bismuth-207 at Eniwetok Atoll, Nature (Lond.), 193: 796-797. Marsh, J. A., dr., 1970, Primary Productivity of Reef-Building Calcareous Red Algae, Ecology, 51: 255-263. Odum, H. T., and E. P. Odum, 1955, Trophic Structure and Pro- ductivity of a Windward Coral Reef Community on Eniwetok Atoll, Ecol. Monogr., 25: 291-320. Palumbo, R. F., 1950, Algae, Radiobiological Survey of Bikini, Eniwetok, and Likiep Atolls, July-August 1949, U. S. Atomic Energy Commission, AECD-3446, pp. 108-118. , 1955, Uptake of Iodine-131 by the Red Alga Asparagopsis taxiformis, U. S. Atomic Energy Commision, UWFL-44, 8 pp. ___, 1959, Gross Beta Radioactivity of the Algae at Enewetak Atoll, 1954-1956, U. S. Atomic Energy Commision, UWFL- 61, 32 pp. Smith, S. V., and J. A. Marsh, Jr., 1973, Organic Carbon Pro- duction on the Windward Reef Flat of Eniwetok Atoll, Limnol. Oceanogr., 18: 953-961. Taylor, W. R., 1950a, Marine Algal Vegetation of the Marshall Islands in Comparison with Other Tropical Areas, Seventh Intern. Bot. Congr. Proc. (Stockholm), pp. 826-827. —_, 1950b, Plants of Bikini and Other Northern Marshall Islands, University of Michigan Press, Ann Arbor. __, 1964, The Genus Turbinaria in Eastern Seas, J. Linn. Soc. Lond. Bot., 58: 475-490. Tsuda, R. T., 1976, Occurrence of the Genus Sargassum (Phaeo- phyta) on Two Pacific Atolls, Micronesica, 12: 279-282. __, F. R. Fosberg, and M. H.- Sachet, 1977, Distribution of Seagrasses in Micronesia, Micronesica, 13: 191-198. Webb, K. L., W. D. DuPaul, W. Wiebe, W. Sottile, and R. E. Johannes, 1975, Enewetak (Eniwetok) Atoll: Aspects of the Nitrogen Cycle on a Coral Reef, Limnol. Oceanogr., 20: 198-210. and W. J. Wiebe, 1975, Nitrification on a Coral Reef, Canadian J. Microbiol., 21: 1427-1431. Wiebe, W. J., 1976, Nitrogen Cycle on a Coral Reef, Micronesica, 12: 23-26. , R. E. Johannes, and K. L. Webb, 1975, Nitrogen Fixation in a Coral Reef Community, Science (Wash.), 188: 257-259. — Fungi of Enewetak Atoll PAUL H. DUNN* AND DON REYNOLDS} “Riverside Fire Laboratory, USDA Forest Service Riverside, California 92507 tNatural History Museum, Los Angeles County Los Angeles, California 90007 INTRODUCTION The microfungi of Enewetak Atoll are typical tropical forms and are not restricted to that atoll. Because of the extreme isolation of Enewetak, however, it seems likely that the fungi there are indigenous to the atoll and are not introductions by man. Whether the origin is terrestrial or from the beaches remains a question. The first study of microfungi on Enewetak (Sparrow, 1948) isolated unicellular chytrids of three different fami- lies from soil samples (Table 1), but no filamentous chy- Chapter 2 trids or Oomycetes were isolated. An extensive study (Dunn, 1973) isolated 101 specimens from the beaches of Enewetak. An additional eight newly recorded taxa are listed in Table 1. The specimens were isolated by the Warcup plating method (Warcup, 1950), which established only that viable propagules were present. Whether the fun- gal propagules of the Enewetak beaches are dormant or active was not defined. Fungal propagules have been preserved for long periods when stored in distilled water (Campbell and Stewart, 1980). It can be assumed, there- fore, that nutrient-poor seawater could provide a habitat in which those propagules that tolerate salinity could remain viable. Reports from the atoll beaches (Dunn, 1973; Steele, 1967) are based on isolations from enrichment culture. Marine fungi have been defined as those observed growing on a substrate in a marine habitat (Kohlmeyer and TABLE 1 Checklist of Enewetak Atoll Fungi Kingdom MYCETEAE Class CHYTRIDIOMYCETINA Order CHYTRIDIALES Family OLPIDIACEAE Olpidium rhizophlyctidis Sparrow: Sparrow, 1948. Family PHLYCTIDIACEAE Rhizophydium marshallense Sparrow: Sparrow, 1948. Family RHIZIDIACEAE Rhizophlyctis rosea (DeBary and Woronin) Fischer: Sparrow, 1948. Class ZYGOMYCOTINA Order MUCORALES Family PIPTOCEPHALIDACEAE *Syncephalastrun racemosum Cohn ex Schroter, 1886. Family THAMNIDIACEAE *Helicostylum glomeratum (van Tieghem and Le Monnier) van Tieghem, 1876. Family MUCORACEAE Absidia corymbifera (Cohn) Saccardo and Trotter: Dunn, 1973. Mucor sp.: Dunn, 1973. Rhizopus nigricans Ehrenberg: Dunn, 1973. Order ENTOMOPHTHORALES Family ENTOMOPHTHORACEAE Conidiobolus sp.: Dunn, 1973. *New Enewetak Atoll record. 11 (This table continued on next page.) DUNN AND REYNOLDS TABLE 1 (cont'd) Class ASCOMYCOTINA Order SPHAERIALES Family CHAETOMIACEAE Ascotricha guamensis Ames: Dunn, 1973. Chaetomium globosum Kunze ex Fries: Dunn, 1973. Family MELANOSPORACEAE *Melanospora sp. Order PLEOSPORALES Family SPORORMIACEAE Sporormia sp.: Dunn, 1973. Order EUROTIALES Family GYMNOASCACEAE Emericellopsis minima Stolk: Dunn, 1973. Gymnoascus reesii Baranetsky: Dunn, 1973. Order MICROASCALES Family MICROASCACEAE Microascus cinereus (Emile-Weil and Gaudin) Curzi: Dunn, 1973. Microascus trigonosporus Emmons and Dodge: Dunn, 1973. Class DEUTEROMYCOTINA Order SPHAEROPSIDALES Family SPHAERIOIDACEAE Lasiodiplodia theobromae (Pat.) Griff. and Maubl.: Dunn, 1973. Order MELANCONIALES Family MELANCONACEAE Pestalotia sp.: Dunn, 1973. Order MONILIALES Family MONILIACEAE ; Acremonium bacillisporum (Onions and Barron) W. Gams (Teleomorph: Chaetomium sp.): Dunn, 1973. *Acremonium charticola: (Lindau) W. Gams, 1971. Acremonium pteridii W. Gams and Frankland: Dunn, 1973. Acremonium terricola (Miller et al.) W. Gams: Dunn, 1973. Aspergillus aculeatus lizuka: Dunn, 1973. Aspergillus carneus (v. Tiegh) Blochwitz: Dunn, 1973. Aspergillus conjunctus Kwon and Fennell: Dunn, 1973. Aspergillus flavipes (Bain. and Sart.) Thom and Church: Dunn, 1973. Aspergillus flavus Link: Dunn, 1973. Aspergillus ochraceus Wilhelm; Dunn, 1973. Aspergillus oryzae (Ahlburg) Chon: Dunn, 1973. Aspergillus recurvatus Raper and Fennell: Dunn, 1973. Aspergillus terreus Thom: Dunn, 1973. Aspergillus terricola? Marchal: Dunn, 1973. Aspergillus tubingensis (Schober) Mosseray: Dunn, 1973. Aspergillus ustus (Bainier) Thom and Church: Dunn, 1973. Aspergillus variecolor (Berk. and Br.) Thom and Raper: Dunn, 1973. Aspergillus versicolor (Vuill.) Tiraboschi: Dunn, 1973. Aspergillus wentii group: Dunn, 1973. Emmonsia sp.: Dunn, 1973. Geotrichum sp.: Dunn, 1973. Hansfordia ovalispora Hughes: Dunn, 1973. Nodulisporium sp.: Dunn, 1973. Oidiodendron sp.: Dunn, 1973. Paecilomyces sp.: Dunn, 1973. Paecilomyces lilacinum (Thom) Samson: Dunn, 1973. Penicillium sp. (monoverticillate, with sclerotia): Dunn, 1973. Penicillium sp. (polyverticillate): Dunn, 1973. Penicillium canescens Sopp.: Dunn, 1973. Penicillium citreo-viride Biourge: Dunn, 1973. Penicillium corylophilum Dierckx: Dunn, 1973. Penicillium frequentans Westling: Dunn, 1973. “New Enewetak Atoll record. (This table continued on next page.) FUNGI TABLE 1 (cont'd) Family MONILIACEAE (cont'd) Penicillium funicolosum Thom: Dunn, 1973. Penicillium janthinellum Biourge: Dunn, 1973. Penicillium verruculosum Peyronel: Dunn, 1973. Phialophora sp.: Dunn, 1973. Sporothrix sp.: Dunn, 1973. Sporotrichum sp.: Dunn, 1973. *Trichoderma aureoviride Rifai, 1969. *Trichoderma harzianum Rifai, 1969. *Trichoderma pseudokongii Rifai, 1969. Tritirachium sp.: Dunn, 1973. Verticillium lecanii (Zimm.) Viegas: Dunn, 1973. Yeast: Dunn, 1973. Family DEMATIACEAE Acrophialophora fusispora (Saksena) M. B. Ellis: Dunn, 1973. Aureobasidium pullulans (DeBary) Arnaud: Dunn, 1973. Cercospora sp.: Dunn, 1973. Cladosporium cladosporioides (Fresen.) de Vries: Dunn, 1973. Cladosporium herbarum (Pers.) Link ex S. F. Gray: Dunn, 1973. Cladosporium sphaerospermum Penz.: Dunn, 1973. Curvularia sp.: Dunn, 1973. Curvularia clavata Jain: Dunn, 1973. Curvularia lunata (Wakker) Boedijn: Dunn, 1973. Curvularia oryzae Bugnicourt: Dunn, 1973. Curvularia senegalensis (Speg.) Subram.: Dunn, 1973. Curvularia tuberculata Jain: Dunn, 1973. Drechslera australiensis (Bugnicourt) Subram. and Jain Ex M. B. Ellis: Dunn, 1973. Drechslera halodes (Drechsler) Subram. and Jain: Dunn, 1973. Drechsiera hawaiiensis (Bugnicourt) Subram. and Jain Ex M. B. Ellis: Dunn, 1973. Drechslera papendorfii (van der Aa) M. B. Ellis: Dunn, 1973. Gliomastix murorum (Corda) Hughes: Dunn, 1973. Gliomastix murorum (Corda) Hughes var. felina (Marchal) Hughes: Dunn, 1973. Gliomastix murorum (Corda) Hughes var. polychroma (van Beyma) Dickinson: Dunn, 1973. Humicola fuscoatra Traaen: Dunn, 1973. Humicola sp.: Dunn, 1973. Leptographium sp.: Dunn, i973. Mammaria sp.: Dunn, 1973. Memnoniella echinata (Riv.) Galloway: Dunn, 1973. Nigrospora sphaerica (Sacc.) Mason: Dunn, 1973. Nigrospora state of Khuskia oryzae Hudson: Dunn, 1973. Rhinocladiella cellaris (Pers. ex S. F. Gray) M. B. Ellis: Dunn, 1973. Scolecobasidium humicola Barron and Busch: Dunn, 1973. Scolecobasidium variabile Barron and Busch: Dunn,1973. Scopulariopsis baarnensis Morton and Smith: Dunn, 1973. Scopulariopsis brumptii: Dunn, 1973. Scopulariopsis brevicaulis (Sacc.) Bainier: Dunn, 1973. Scopulariopsis chartarum (Smith) Morton and Smith: Dunn, 1973. Scopulariopsis fimicola (Cost. and Matr.) Vuill.: Dunn, 1973. Scopulariopsis sphaerospora Zach: Dunn, 1973. Scopulariopsis sp.: Dunn, 1973. Stachybotrys atra Corda: Dunn, 1973. Stachybotrys state of Melanopsamma pomiformis (Pers. ex Fr.) Sacc.: Dunn, 1973. Stemphylium vesicarium (Wallr.) Simmons: Dunn, 1973. Trichocladium canadense Hughes: Dunn, 1973. *Ulocladium chartarum (Preuss) Simmons, 1967. Family STILBACEAE Graphium putredinis (Corda) Hughes: Dunn, 1973. Isaria sp.: Dunn, 1973. *New Enewetak Atoll record. (This table continued on next page.) 14 DUNN AND REYNOLDS TABLE 1 (cont'd) Family TUBERCULARIACEAE Cylindrocarpon sp.: Dunn, 1973. Fusarium episphaeria (Tode) Fr.: Dunn, 1973. Fusarium solani (Mart.) Sacc.: Dunn, 1973. Fusarium sp.: Dunn, 1973. Metarrhizium anisopliae (Metschn.) Sorok.: Dunn, 1973. Myrothecium roridum Tode ex Steudel: Dunn, 1973. Myrothecium striatisporum Preston: Dunn, 1973. Class BASIDIOMYCOTINA Unidentified air isolate. *New Enewetak Atoll record. Kohlmeyer, 1979). None of the fungi isolated from Enewetak, therefore, fits the definition of a marine fungus. Inhibitors in seawater (Kirk, 1980) and in soil (Lockwood, 1977) may prevent fungal spores from germinating until changes occur in the endogenous or exogenous nutrient status or until the exogenous spore inhibitors are removed. The exogenous inhibitors of nonmarine fungi can be partly removed or deactivated by autoclaving seawater; most marine fungi can germinate in raw seawater. Addition of glucose can also counteract the inhibition (Kirk, 1980). Few studies have been done on decomposition of sub- tidal substrate. In a study of seagrass, Thalassia testudinum Konig, Newell and Fell (1980) showed that although non- marine fungi were present, they played only a minor role in decomposition of T. testudinum leaves until the leaves were deposited in the intertidal. The terrestrial fungi are active in the intertidal zone, but their significance in rela- tion to marine fungi is unknown (Newell and Fell, 1980, 1982). On the basis of the present discussion, it can be suggested that the fungal propagules in Enewetak Atoll beaches are also dormant until they come in contact with a proper substrate. Fungi from the beaches of Enewetak Atoll have been compared with those from other locations. When the specimens isolated by Dunn (1973) were compared with fungi plated from sediment of a coastal estuary of North Carolina with salinities from 0 to 36°/,,, (Borut and John- son, 1962), a Dice similarity quotient of 17% was found, indicating a distinct biota for these two very different marine habitats. In a comparison of the fungi isolated by Warcup plat- ing from beaches at Enewetak Island and the Hawaiian Islands, a Dice similarity quotient of 43% was found (Dunn, 1973). The beaches in the Hawaiian Islands are both vol- canic and carbonate, whereas the beaches at Enewetak Island are carbonate. Similarity is greater between the mycobiota of the carbonate beaches in Enewetak and Hawaii (48%) than between the carbonate and volcanic beaches in Hawaii (41%). The principal difference between the mycobiota of the carbonate beaches sampled in Enewetak and those in Hawaii was the abundance of species present in both beaches relative to the total iso- lates. Typical of the pattern are Gymnoascus reesii Baranetsky and two species in the genus Microascus, which were common at Enewetak Island beach but were infrequently isolated from Kahala Beach, Oahu, Hawaii. CHYTRIDIOMYCOTINA The isolates of this class were all unicellular. Rhizophyctis rosea (DeBary and Woronin) Fischer was the most common isolate, and it is the most widespread of all chytrids (Sparrow, 1960). Unicellular chytrids were found in the beach areas by Dunn (1973) but were not identified. Sparrow (1948) consistently found only the same three species of this class on Enewetak and on three other Marshall Island atolls. The filamentous species of the class were conspicuously absent. OOMYCOTINA Although many species in this class are common plant pathogens and would be expected to be associated with the vegetation, none has been reported from this atoll. They may be too sensitive to salinity to do well in beach habitats (TeStrake, 1959; Lee and Baker, 1972), although some have been isolated from other beach areas. ZYGOMYCOTINA Conidiobolus sp. was isolated from subtidal areas on Enewetak. Species in this genus are normally associated with dung of insect-eating animals or decaying vegetation. This species was isolated only from seawater sources and seems to require salinity. It does not resemble any described species and does not last many generations in culture. Members of the order Mucorales are common iso- lates from beaches at Enewetak but have salinity optima more appropriate for terrestrial species (Dunn, 1973). ASCOMYCOTINA Many ascomycetes are known to produce two types of spores: the teleomorph, or sexual state, and the anamorphs, or asexual states. The teleomorph may be associated with one or more anamorphs. The teleomorph genus regularly associated with the isolated Scopulariopsis FUNGI anamorph, for example, is well documented but was not isolated in the studies. Ascotricha guamensis Ames previ- ously was isolated from Guam, New Guinea, India, and more recently, from soil at Enewetak. Microascus cinereus (Emele-Weil and Gaudin) Gurzi, the teleomorph, and its associated anamorph were among the most frequently iso- lated fungi in the beach study. These were rarely isolated from carbonate beaches in Hawaii. Gymnoascus reesii is typical of the saline habitats all through the Pacific basin (Baker and Meeker, 1972). DEUTEROMYCOTINA At Enewetak Atoll, the majority of the Deutero- mycotina genera isolated from beaches was comprised of members of the order Moniliales (94 taxa), with the majority of species evenly divided between the families Dematiaceae and Moniliaceae. Acremonium bacillisporum (Onions and Barron) W. Gams was a prominent member of the carbonate sand community at both Enewetak and Kahala Beach, Oahu, Hawaii (Dunn, 1973). The aspergilli were common and the penicillia were rare in beaches at both localities (Dunn, 1973). These patterns coincide with other data indicating that the aspergilli are more common in the subtropics, and the penicillia are more common in temperate climates. Graphium putredinis (Corda) Hughes was found commonly in the anaerobic zone in deeper sedi- ments off Enewetak Atoll. Fusarium solani (Mart.) Sacc. was common in beaches at Enewetak but mostly was found near the land zone. The moniliaceous genus Dreschlera is represented on Enewetak Atoll by four species (Table 1). All are known from the tropics and southernmost Africa. Two of the species’ of Cladosporium, C. cladosporioides, and C. sphaerosper- mum are found in tropical and subtropical climates in saline habitats. A third species, C. herbarum, was not found frequently at Enewetak and is cosmopolitan in distribution. BASIDIOMYCOTINA The most common isolate from air at Enewetak Atoll is a basidiomycete identifiable by large clamp connections on the hyphae (Dunn, 1973). It did not sporulate in culture. Other basidiomycetes in the Marshall Islands, mostly asso- ciated with decaying wood, were collected by Rogers (1947) and Taylor (1950). They did not collect fungi on Enewetak Atoll, and fungi from decaying wood have not been studied on Enewetak. CONCLUSION The mycological data from Enewetak Atoll reflect two principles of fungal biogeography: similar regions have similar biotas, and distributional data are often due to bias collection activity of mycologists (Baker and Meeker, 1972). For example, the genus Myrothecium was initially reported in the Pacific from New Zealand. This genus was 15 then reported from Moorea, Society Island (Peterson, 1960). As data from other Pacific islands became available, the distribution was enlarged to include Tonga, Hawaii, New Caledonia (Baker and Meeker, 1972), and Enewetak (Baker and Meeker, 1972; Dunn, 1973). A similar expan- sion through the tropical Pacific may be expected for most of the species reported here. ACKNOWLEDGMENTS The authors thank G. E. Baker for help during the beach studies and acknowledge the financial support of the U. S. Nuclear Regulatory Commission Grant AT-(29-2)-229 and the use of its Mid-Pacific Research Laboratory. REFERENCES Baker, G. E., and J. A. Meeker, 1972, Ecosystems, Mycologist and the Geological Distribution of Fungi in the Central Pacific, Pac. Sci., 26: 418-432. Borut, S. Y., and T. W. Johnson, Jr., 1962, Some Biological Observations on Fungi in Estuarine Sediments, Mycologia, 54: 181-193. Campbell, M. C., and J. L. Stewart, 1980, The Medical Mycology Handbook, John Wiley and Sons, New York. Dunn, P. H., 1973, The Ecology of Filamentous Heterotrophic Microorganisms in Subtropical and Tropical Marine Psammen Habitats, Ph.D. dissertation, University of Hawaii. Kirk, P. W., Jr., 1980, The Mycostatic Effect of Seawater on Spores of Terrestrial and Marine Higher Fungi, Bot. Mar., 23: 233-238. Kohlmeyer, J., and E. Kohlmeyer, 1979, Marine Mycology, The Higher Fungi, Academic Press, New York. Lee, B. K. H., and G. E. Baker, 1972, An Ecological Study of the Soil Microfungi in a Hawaiian Mangrove Swamp, Pac. Sci., 26: 1-10. Lockwood, J. L., 1977, Fungistasis in Soils, Biol. Rev. (Camb. Philos. Soc.), 52: 1-43. Newell, S. Y., and J. W. Fell, 1980, Mycoflora of Turtlegrass (Thalassia testudinum Konig) Is Recorded After Seawater Incu- bation., Bot. Mar., 23: 265-275. , 1982, Surface Sterilization and the Active Mycoflora of Leaves of a Seagrass, Bot. Mar., 25: 339-346. Peterson, R. H., 1960, Some Soil and Coprophilous Fungi from the South Pacific Area, Mycologia, 52: 552-556. Rogers, D. P., 1947, Fungi of the Marshall Islands, Central Pacific Ocean, Pac. Sci., 1: 92-107. Sparrow, F. K., 1948, Soil Phycomycetes from Bikini, Eniwetok, Rongerik, and Rongelap Atolls, Mycologia, 40: 445-453. ——, 1960, Aquatic Phycomycetes, 2nd edition, University of Michigan Press, Ann Arbor. Steele, C. W., 1967, Fungus Populations in Marine Waters and Coastal Sands of Hawaiian, Line, and Phoenix Islands, Pac. Sci, 21: 317-331. Taylor, W. R., 1950, Plants of Bikini and other Northern Marshall Islands, University of Michigan Press, Ann Arbor. TeStrake, D., 1959, Estuarine Distribution and Saline Tolerance of Some Saprolegniaceae, Phyton, 12: 147-152. Warcup, J. H., 1950, The Soil Plate Method for the Isolation of Fungi from the Soil, Nature (Lond.), 166: 117-118. . Chapter 3 Natural History of Terrestrial Vascular Plants of Enewetak Atoll JANET O. LAMBERSON U. S. Environmental Protection Agency Hatfield Marine Science Center, Newport, Oregon 97365 INTRODUCTION Enewetak Atoll has evolved over geological time from a raised limestone island environment supporting mangrove swamps and an upland mixed forest to a low coral sand island habitat with typical strand vegetation. The 128 species of plants recorded from the atoll include many introduced weeds and ornamental or food plants, although 43% of the flora is considered to be indigenous to Micronesia. The dry, windy season from November through April causes many of the plants to drop some or all of their leaves or to die back completely. Summer—Jdune through September—is the season of maximum plant growth and more flower and seed production. The effects of World War II and post-war military activity, the nuclear test program from 1948 to 1958, and the radiological cleanup and rehabilitation for the return of the people of Enewetak have been significant with regard to the vegetation. Many species were accidentally or inten- tionally introduced to the atoll, and some species have disappeared. The vegetation was greatly altered because of destruction of habitat with removal of soil and nutrients and exposure of the plants to salt spray and drying winds. Several collections and studies of the flora have been reported. This chapter describes stages in the succession of the vegetation following disturbance. The vegetation of the islands during the period 1975 to 1977 is also described in some detail. ENVIRONMENT AND BIOGEOGRAPHY The islands and shallow reefs of Enewetak Atoll are the top of a limestone cap approximately 1 km thick, rest- ing on the basalt remains of an extinct volcano which was active during the pre-Tertiary geological period. Core sam- 17 ples from drilling on the island of Medren and on the former island of Elugelab include basalt encountered at depths of 1271 and 1405 m, respectively. The samples also show that during the reef’s geological history it was exposed and submerged several times as a result of fluc- tuations in sea level (Emery, Tracey, and Ladd, 1954; Tracy and Ladd, 1974). Analysis of pollen grains in the cores (Leopold, 1969) showed that during Miocene times Enewetak was a raised limestone island supporting an intertidal mangrove swamp, mangrove depressions on beach ridges, an upland forest of salt-intolerant plants, and a typical strand vegetation (Fos- berg, 1960) similar to that presently found at Enewetak. Leopold (1969) listed 17 angiosperm genera among 54 species of plants found as fossil remains in the cores, including Pandanus, Pisonia, Terminalia, Tournefortia, Cordia, Morinda, and Guettarda. The islands of Enewetak Atoll have apparently reached their present form within the Holocene. The reef around the atoll grew to keep pace with the rising sea level, then was eroded by wave action during a more recent drop in sea leve! within the past 2000 to 4000 years (Buddemeier, Smith, and Kinzie, 1975). The terrestrial plants that occur now at Enewetak have been introduced by chance or inten- tionally as seeds and cuttings brought to the atoll by ocean currents, wind, birds, and people. In comparison with other atolls in the Marshalls, Enewetak has fewer species of plants, and to those familiar with the dense undergrowth and luxuriant ferns of the southern atolls, Enewetak may seem somewhat barren. Only 79 out of 155 species of plants, including those cul- tivated in gardens, listed by Fosberg (1955) in the northern Marshall Islands have been reported from Enewetak. Species indigenous to Micronesia comprise 43% of the total flora (55 species). (See Tables 1 through 4.) Speci- mens of all species in Table 1, except Cocos nucifera and others noted in Table 4 were deposited in the reference collection at the Mid-Pacific Laboratory at Enewetak. Addi- tional specimens from Enewetak, including the new species listed, are deposited at the Bishop Museum (BPBM) and at the U. S. National Museum of Natural History (USNM). The taxonomic nomenclature for the Dicotyledonae and 18 LAMBERSON TABLE 1 Checklist of Terrestrial Vascular Plants of Enewetak Atoll* Division TRACHEOPHYTA Class PSILOPSIDA Order PSILOTALES Family PSILOTACEAE +Psilotum nudum (L.) Beauvais: Lamberson, 1982; Fosberg et al., 1982. Class PTEROPSIDA Order FILICALES Family POLYPODIACEAE +Polypodium scolopendria Burmann f. Phymatodes scolopendria (Burmann f.) Ching: St. John, 1960. Polypodium phymatodes Linn: Bryan, 1944. Class ANGIOSPERMAE Subclass MONOCOT YLEDONEAE Family PANDANACEAE +Pandanus sp.: Bryan, 1944; Taylor, 1950; Woodbury, 1962; Fall et al., 1971; Koranda et al., 1973; Lamberson, 1982. +Pandanus tectorius Park: Fosberg, 1955; Welander et al., 1966. +Pandunus brachypodus Kanehira: Kanehira, 1935; St. John, 1960. +Pandanus enchabiensis Kanehira: Kanehira, 1935; St. John, 1960. +Pandanus korrorensis Kanehira: St. John, 1960. +Pandanus odoratissimus L. f. var. novocaledonicus (Martelli) St. John: St. John, 1960. +Pandanus odoratissimus L. f. var. novoguineensis (Martelli) St. John: St. John, 1960. +Pandanus pulposus (Warb.) Martelli: St. John, 1960. +Pandanus rectanulatus Kanehira: Kanehira, 1935; St. John, 1960. +Pandanus rhombocarpus Kanehira: Kanehira, 1935; St. John, 1960. +Pandanus utiyamai Kanehira: St. John, 1960. Family GRAMINAE Cenchrus brownii Roemer and Schultes: Taylor, 1950; St. John, 1960. Cenchrus echinatus L.: Bryan, 1944; St. John, 1950; Taylor, 1950; Fosberg, 1955; St. John, 1960; Welander et al., 1966; Lamberson, 1982. Cenchrus sp.: Fall et al., 1971. Chloris inflata Link: St. John, 1950; Fosberg, 1955; St. John, 1960; Welander et al., 1966; Lamberson, 1982. Cynodon dactylon (L.) Persoon: Bryan, 1944; Fosberg, 1955; St. John, 1960; Welander et al., 1966; Koranda et al., 1973; Lamberson, 1982. Dactyloctenium aegypticum (L.) Willdenow: St. John, 1960; Welander et al., 1966; Lamberson, 1982. +Digitaria setigera Roemer and Schultes: St. John, 1960; Lamberson, 1982. Digitaria pruriens (Trinius) Buese: Taylor, 1950. Digitara microbachne (Presl) Henrard: Fosberg, 1955. Eleusine indica (L.) Gaertner: Bryan, 1944; Taylor, 1950; Fosberg, 1955; St. John, 1960; Welander et al., 1966; Lamberson, 1982. Eragrostis scabrifolia Swallen: Lamberson, 1982. Eragrostis tenella (L.) Roemer and Schultes: Lamberson, 1982. Eragrostis ambilis (L.) Wight and Arnott: Taylor, 1950, Fosberg, 1955; St. John, 1960; Welander et al., 1966. Eragrostis sp.: Lane, 1960. Lepturopetium marshallense Fosberg and Sachet, 1982. Lepturus repens (Forster f.) R. Brown var. occidentalis: St. John, 1960. Lepturus repens (Forst. f.) R. Brown: Bryan, 1944; Taylor, 1950; Fosberg, 1955; Palumbo, 1962; Woodbury, 1962; Welander et al., 1966; Fall et al., 1971; Koranda et al., 1973; Lamberson, 1982. Lepturus sp.: Lane, 1960; Carpenter et al., 1968. {Lepturus repens (Forster f.) R. Brown var. subulatus Fosberg. Brachiaria mutica (Forsskal) Stapf.: Lamberson, 1982. Setaria verticillata (L.) Beauvais: Bryan, 1944; Taylor, 1950; Fosberg, 1955; St. John, 1960. +Thuarea involuta (Forster f.) Roemer and Schultes: Taylor, 1950; Fosberg, 1955; St. John, 1960; Lamberson, 1982. Tricachne insularis (L.) Nees: Fosberg, 1955; St. John, 1960; Welander et al., 1966; Lamberson, 1982. Tricholaena rosea Nees Tricholaena repens (Willdenow) Hitchcock: Taylor, 1950; Lane, 1960; St. John, 1960. *Excludes plant species reported only from gardens, which are considered to be transient components of the flora. +Species considered indigenous to Micronesia. +The taxonomy of the genus Pandanus is disputed. Some taxonomists would consider most of the species listed here as forms of the species Pandanus tectorius. New Enewetak record; see Table 4. (This table continued on next page.) HISTORY OF TERRESTRIAL VASCULAR PLANTS 19 TABLE 1 (cont’d) Family CYPERACEAE Cyperus compressus L.: Lamberson, 1982. +Cyperus javanicus Houttuyn: St. John, 1960. Cyperus ferax L. C. Richard Cyperus odoratus L.: St. John, 1960. Cyperus rotundus L.: Lamberson, 1982. +Fimbristylus cymosa R. Brown ssp. spathacea (Roth) Koyama, 1964. Fimbristylus annua Roemer and Schultes forma diphyllia (Retz) Kukenth: Bryan, 1944. Fimbristylus cymosa R. Brown: Taylor, 1950. Fimbristylus atollensis St. John: Lane, 1960; St. John, 1960; Welander et al., 1966; Fall et al., 1971; Koranda et al., 1973; Koranda et al., 1978; Lamberson, 1982. Fimbristylus sp.: Woodbury, 1962. Family PALMAE Cocos nucifera L.: Bryan, 1944; St. John, 1950; Taylor, 1950; Fosberg, 1955; St. John, 1960; Palumbo, 1962; Woodbury, 1962: Welander et al., 1966; Carpenter et al., 1968; Fall et al., 1971; Koranda et al., 1973; Lamberson, 1982. Family AMARYLLIDACEAE +Crinum asiaticum L.: Bryan, 1944; St. John, 1960; Lamberson, 1982. Pancratium littorale Jacquin: Lamberson, 1982. Family TACCACEAE +Tacca leontopetaloides (L.) O. Kuntze: Taylor, 1950; Fosberg, 1955; St. John, 1960; Welander, et al., 1966; Koranda et al., 1973; Lamberson, 1982. Tacca pinnatifida Forster: Bryan, 1944. Subclass DICOTYLEDONEAE Family MORACEAE Artocarpus altilis (Parkinson) Fosberg: Fosberg et al., 1979. Artocarpus incisus (Thunberg) L. f.: St. John, 1960. +Artocarpus mariannensis Trecul: Fosberg et al., 1979. Family URTICACEAE +Laportea ruderalis (Forster f.) W. L. Chew: Fosberg et al., 1979; Lamberson, 1982. Fleurya ruderalis (Forster f.) Gaudichaud: St. John, 1950; Taylor, 1950; Fosberg, 1955; Lane, 1960; St. John, 1960; Welander et al., 1966. Family OLACACEAE +Ximenia americana L.: Taylor, 1950; Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. Family POLYGANACEAE Coccoloba uvifera (L.) L.: Lamberson, 1982. Family AMARANTHACEAE +Achyranthes aspera L.: Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. +Achyranthes canescens R. Brown: St. John, 1960. +Amaranthus dubius Martell: Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. Amaranthus viridis L.: Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. Family NYCTAGINACEAE +Boerhavia albiflora var. powelliae Fosberg: Fosberg et al., 1979. +Boerhavia repens L. Boerhavia diffusa L. var. eudiffusa: St. John, 1950. Boerhavia diffusa L.: Taylor, 1950; Fosberg, 1955; Lane, 1960; Welander et al., 1966; Fall et al., 1971. Boerhavia diffusa L. var. diffusa: St. John, 1960. +Boerhavia tetrandra Forster: Taylor, 1950; Fosberg, 1955; Palumbo, 1962; Woodbury, 1962; Welander et al., 1966. Boerhavia diffusa var. tetrandra (Forster f.) Heimerl: Bryan, 1944; St. John, 1960. Boerhavia spp.: Lamberson, 1982. +Pisonia grandis R. Brown: Bryan, 1944; Taylor, 1950; Fosberg, 1955; Lane, 1960; St. John, 1960; Welander et al., 1966; Carpenter et al., 1960; Fall et al, 1971; Koranda et al., 1973; Koranda et al., 1978; Fosberg et al., 1979; Lamberson, 1982. Pisonia sp.: Woodbury, 1962; Lee and Lee, 1978. Family PORTULACACEAE +Portulaca australis Endl.: Fosberg et al., 1979. Portulaca quadrifida L.: Bryan, 1944; Taylor, 1950; Welander et al., 1966. Portulaca samoensis v. Poelln.: Fosberg, 1955: Lane, 1960; St. John, 1960; Lamberson, 1982. +Portulaca lutea Solander ex Forster f.: Taylor, 1950; Fosberg, 1955; St. John, 1960; Welander et al., 1966; Fosberg et al.. 1979; Lamberson, 1982. +Species considered indigenous to Micronesia. (This table continued on next page.) 20 LAMBERSON TABLE 1 (cont'd) Family PORTULACACEAE (cont'd) Portulaca oleracea L.: Bryan, 1944; St. John, 1950; Biddulph and Cory, 1952; Fosberg, 1955; Lane, 1960; St. John, 1960; Palumbo, 1962; Welander et al., 1966; Fosberg et al., 1979; Lamberson, 1982. Family LAURACEAE Cassytha filiformis L.: Bryan, 1944; Taylor, 1950; Fosberg, 1955; St. John, 1960; Woodbury, 1962; Welander et al., 1966; Koranda et al., 1973; Fosberg et al., 1979; Lamberson, 1982. +Cythya filiformis: Koranda et al., 1978. Family FABACEAE +Canavalia cathartica Thouars: Fosberg et al., 1979; Lamberson, 1982. Canavalia microcarpa (DeCandolle) Piper: Fosberg, 1955; St. John, 1960; Welander et al., 1966. Desmodium incanum DeCandolle. Desmodium adscendens (Sw.) DeCandolle: Lamberson, 1982. Leucaena leucocephala (Lamarck) DeWit: Lamberson, 1982. +Vigna marina (Burmann) Merrill: Bryan, 1944; St. John, 1960. Family ZYGOPHYLLACEAE Tribulus cistoides L.: Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. Family RUTACEAE {Citrus aurantifolia (Christm.) Swingle. Family SURIANACEAE Suriana maritima L.: Taylor, 1950; Fosberg, 1955; Lane, 1960; St. John, 1960; Woodbury, 1962; Welander et al., 1966; Koranda et al., 1973; Fosberg et al., 1979; Lamberson, 1982. Family SIMAROUBACEAE +Soulamea amara Lamarck: Welander et al., 1966. Family EUPHORBIACEAE +Euphorbia chamissonis (Klotsch and Garcke) Boissier: Fosberg, 1955; St. John, 1960; Fosberg et al., 1979; Lamberson, 1982. Euphorbia atoto Forster: Welander et al., 1966. Euphorbia hirta L.: Fosberg, 1955; St. John, 1960; Welander et al., 1966; Fosberg et al., 1979; Lamberson, 1982. Euphorbia thymifolia L.: St. John, 1960; Welander et al., 1966; Lamberson, 1982. Phyllanthus amarus Schum. and Thonn.: St. John, 1960; Fosberg et al., 1979; Lamberson, 1982. Ricinus communis L.: Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. Family TILIACEAE +Triumfetta procumbens Forster: Bryan, 1944; Taylor, 1950; Fosberg, 1955; Lane, 1960; St. John, 1960; Palumbo, 1962; Woodbury, 1962; Welander et al., 1966; Carpenter et al., 1968; Fall et al., 1971; Koranda et al., 1973; Fosberg et al., 1979; Lamberson, 1982. Family MALVACEAE +Hibiscus tiliaceus L.: Lamberson, 1982. Malvastrum coromandelianum (L.) Garcke: St. John, 1960. +Sida fallax Walpers: Bryan, 1944; Taylor, 1950; Fosberg, 1955; St. John, 1960; Welander et al., 1966; Koranda et al., 1973; Fosberg et al., 1979; Lamberson, 1982. Family STERCULIACEAE Waltheria indica L.: Lamberson, 1982. Family LYTHRACEAE +Pemphis acidula Forster: Taylor, 1950; Fosberg, 1955; Lane, i960; St. John, 1960; Welander et al., 1966; Koranda et al., 1973; Fosberg et al., 1979; Lamberson, 1982. Pemphis sp.: Carpenter et al., 1968. Family COMBRETACEAE +Terminalia samoensis Rechinger: Fosberg, 1955; St. John, 1960; Welander et al., 1966; Fall et al., 1971; Fosberg et al., 1979. Terminalia litoralis Seeman: Taylor, 1950; Lane, 1960; Lamberson, 1982. Terminalia sp.: Woodbury, 1962. Family LOGANIACEAE Polypremum procumbens L.: Lamberson, 1982. Family APOCY ANACEAE +Neisosperma oppositifolia (Lamarck) Fosberg and Sachet: Fosberg et al., 1979; Lamberson, 1982. Ochrosia parviflora (Forster) Henslow: Bryan, 1944; Taylor, 1950. Ochrosia oppositifolia (Lamarck) K. Schumann: Fosberg, 1955; Lane, 1960; St. John, 1960; Welander et al., 1966; Fall et al., 1971; Koranda et al., 1973. Ochrosia sp.: Woodbury, 1962. +Species considered indigenous to Micronesia. ‘New Enewetak record; see Table 4. (This table continued on next page.) HISTORY OF TERRESTRIAL VASCULAR PLANTS 21 TABLE 1 (cont'd) Family APOCYANACEAE (cont'd) {Plumeria obtusa L., 1753. {Plumeria rubra L., 1753. Family CONVOLVULACEAE +Ipomoea macrantha Roemer and Schultes: Fosberg et al., 1979; Lamberson, 1982. Ipomoea grandiflora (Choisy) Hallier: Bryan, 1944. Ipomoea tuba (Schlechtendahl) G. Don: St. John, 1950; Biddulph and Biddulph, 1953; Fosberg, 1955; Lane, 1960; St. John, 1960; Woodbury, 1962; Welander et al., 1966. Ipomoea alba L.: Taylor, 1950. +Ipomoea pes-caprae ssp. brasiliensis (L.) v. Ooststroom: St. John, 1950; Fosberg, 1955; Lane, 1960; St. John, 1960; Welander et al., 1966; Koranda et al., 1973; Koranda et al., 1978; Fosberg et al., 1979. Ipomoea brasiliensis (L.) Sweet: Lamberson, 1982. Ipomoec sp.: Carpenter et al., 1968; Fall et al., 1971. Family BORAGINACEAE +Cordia subcordata Lamarck: Taylor, 1950; Fosberg, 1955; Lane, 1960; St. John, 1960; Welander et al., 1966; Fall et al., 1971; Koranda et al., 1973; Fosberg et al., 1979; Lamberson, 1982. Cordia sp.: Woodbury, 1962; Carpenter et al., 1968. +Heliotropium procumbens var. depressum (Chamisso) Fosberg and Sachet. Heliotropium anomalum Hooker and Arnott: Welander et al., 1966; Lamberson, 1982. Tournefortia argentea L. f.: Taylor, 1950: Carpenter et al., 1968; Fall et al., 1971; Fosberg et al., 1979. Messerschmidia argentea (L.f.) Johnston: Bryan, 1944; St. John, 1950; Fosberg, 1955; Lane, 1960; St. John, 1960; Woodbury, 1962; Palumbo, 1962; Welander et al., 1966; Jackson and Carpenter, 1967; Koranda et al., 1973; Koranda et al., 1978; Lamberson, 1982. Messerschmidia sp.: Lee and Lee, 1978. Family VERBENACEAE +Clerodendrum inerme var. oceanicum A. Gray: Fosberg et al., 1979. Lippia nodiflora (L.) Rich: Lamberson, 1982. +Premna obtusifolia R. Brown: Lamberson, 1982. Stachytarpheta jamaicensis (L.) Vahl. Stachytarpheta urticaefolia (Salisb.) Sims: Lamberson, 1982. (Vitex trifolia L. Family SOLANACEAE Nicotiana glauca Graham: St. John, 1960. Physalis angulata L. var. angulata St. John, 1960; Welander et al., 1966; Lamberson, 1982. Family ACANTHACEAE Pseuderanthemum carruthersii (Seemann) Guillaumin var. carruthersii: Lamberson, 1982. Pseuderanthemum carruthersii (Seemann) Guillaumin var. atropurpureum (Bull.) Fosberg; Lamberson, 1982. Family RUBIACEAE +Guettarda speciosa L.: Bryan, 1944; St. John, 1950; Taylor, 1950; Lane, 1960; St. John, 1960; Palumbo, 1962; Welander et al., 1966; Fall et al., 1971; Koranda et al., 1973; Fosberg et al., 1979; Lamberson, 1982. Guettarda sp.: Woodbury, 1962. Morinda citrifolia L.: Bryan, 1944; St. John, 1950; Taylor, 1950; Fosberg, 1955; St. John, 1960; Welander et al., 1966; Fall et al., 1971; Koranda et al., 1973; Fosberg et al., 1979; Lamberson, 1982. Morinda sp.: Woodbury, 1962. Family GOODENIACEAE Scaevola taccada (Gaertner) Roxburgh: Lane, 1960; Jackson and Carpenter, 1967; Carpenter et al., 1968; Fall et al., 1971; Fosberg et al., 1979; Lamberson, 1982. Scaevola frutescens (Miller) Krause: Bryan, 1944; St. John, 1950; Taylor, 1950; Fosberg, 1955; St. John, 1960; Woodbury, 1962; Welander et al., 1966; Koranda et al., 1973; Koranda et al., 1978. Scaevola seriacea Vahl: Palumbo, 1962. Scaevola sp.: Lee and Lee, 1978. Family COMPOSITAE Bidens pilosa L. var. minor (BI.) Sherff: Lamberson, 1982. Conyza bonariensis (L.) Cronquist: Fosberg et al., 1979; Lamberson, 1982. Erigeron bonariensis L.: St. John, 1960. Eclipta alba (L.) Hasskarl: Lamberson, 1982. +Pluchea indica (L.) Lessing: St. John, 1960; Fosberg et al., 1979; Lamberson, 1982. +Species considered indigenous to Micronesia. {New Enewetak record; see Table 4. (This table continued on next page.) 22 LAMBERSON TABLE 1 (cont’d) Family COMPOSITAE (cont’d) Pluchea symphytifolia (Miller) Gillis: Lamberson, 1982. Pluchea odorata (L.) Cassini: Fosberg, 1955; St. John, 1960; Welander et al., 1966; Koranda et al., 1973; Koranda et al., 1978. Tridax procumbens L.: Lamberson, 1982. Veronia cinerea (L.) Lessing: Fosberg, 1955; St. John, 1960; Welander et al., 1966; Fosberg, 1979; Lamberson, 1982. Wedelia trilobata (L.) Hitchcock: Fosberg et al., 1979; Lamberson, 1982. +Wollastonia biflora (L.) DeCandolle. Wedelia biflora (L.) DeCandolle: Bryan, 1944; Fosberg, 1955; St. John, 1960; Welander et al., 1966; Lamberson, 1982. +Species considered indigenous to Micronesia. TABLE 2 Garden Plants Reported from Enewetak Atoll, Considered to Be Transient Members of the Flora Division TRACHEOPHYTA Class ANGIOSPERMAE Subclass MONOCOT YLEDONEAE Family GRAMINAE Sorghum bicolor (L.) Moench var. technicum (Koern.) Fiori and Paoli, “broom corn”: Fosberg, 1955. Sorghum vulgare (L.) Persoon: Taylor, 1950. Zea mays L., “corn”: St. John, 1960. Family AGAVACEAE *Cordyline terminalis (L.) Kunth, “ti.” Family LILIACEAE Allium cepa L., “onion”: Bryan, 1944; St. John, 1960. Family MUSACEAE *Musa sapientum L., “banana.” Family ARACEAE *Epipremnum pinnatum (L.) Engler cv. “Aureum.” Subclass DICOT YLEDONEAE Family NYCTAGINACEAE Mirabilis jalapa L., “four-o’clock”: Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. Family CRUCIFERAE Brassica oleracea L. capitata, “cabbage”: Bryan, 1944; St. John, 1960. Brassica pekinensis (Lour.) Rupr., “Chinese cabbage”: Bryan, 1944; St. John, 1960. Brassica sp.: Bryan, 1944; Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. Raphanus sativus L., “radish”: Bryan, 1944; St. John, 1960. Family FABACEAE Phaseolus sp., “beans”: Bryan, 1944; St. John, 1960. Family MALVACEAE *Hibiscus hybrid (H. rosa-sinensis x schizopetalus?), “hibiscus.” Family CARICACEAE Carica papaya L., “papaya”: St. John, 1960. Family CONVOLVULACEAE Ipomoea purpurea (L.) Roth: St. John, 1960. Family SOLANACEAE Solanum lycopersicum L., “tomato”: Bryan, 1944; Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. Family CUCURBITACEAE Citrullus lanatus var. caffrorum (Alef.) Fosberg, “watermelon”: Fosberg, et al., 1979. Citrullus vulgaris Schrader, “watermelon”: Bryan, 1944; Fosberg, 1955; St. John, 1960. Cucumis melo L., “canteloupe”: Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. Cucurbita maxima Duchesne, “squash”: Bryan, 1944; Fosberg, 1955; St. John, 1960; Fosberg et al., 1979. Family COMPOSITAE Lactuca sativa L., “lettuce”: Bryan, 1944; St. John, 1960. *New Enewetak record; see Table 4. HISTORY OF TERRESTRIAL VASCULAR PLANTS 23 TABLE 3 Plants Reported from Enewetak Atoll but Found Only as Drift Seeds on Beaches Division TRACHEOPHYTA Class ANGIOSPERMAE Subclass DICOTYLEDONEAE Family HERNANDIACEAE Hernandia sonora L.: St. John, 1960 Family FABACEAE Caesalpina bonduc (L.) Roxbury: St. John, 1960. Dioclea reflexa Hook. f.: St. John, 1960 Entada phaseoloides (L.) Merrill: St. John, 1960. Mucuna urens (L.) Medicus: St. John, 1960. Family EDPHORBIACEAE Aleurites moluccana (L.) Willdenow: St. John, 1960. Family SAPINDACEAE Sapindus saponaria L.: St. John, 1960. TABLE 4 Collection Data of Newly Recorded Terrestrial Vascular Plants from Enewetak* Family GRAMINAE Lepturus repens (Forst. f.) R. Brown var. subulatus Fosberg. Enewetak Islet: Mar. 1976; Enjebi Islet: s.n.s.d., coll. by R. Warner (MPRL). Family MUSACEAE Musa sapientum L. Enjebi Islet: Planted in Lawrence Livermore Laboratory experimental garden. Japtan Islet: Planted near homes. Not collected. Family ARACEAE Epipremnum pinnatum (L.) Engler cv. “Aureum” Enewetak Islet: Mar. 3 1975, under air conditioner (MPRL). Family AGAVACEAE Cordyline terminalis (L.) Kunth. Enewetak Islet: Feb. 12, 1975, near buildings, coll. by E. H. Bryan, Jr. (BPBM). Family RUTACEAE Citrus aurantifolia (Christm.) Swingle. Enjebi Islet: Planted in Lawrence Livermore Laboratory experimental garden. Not collected. Family MALVACEAE Hibiscus sp. (hybrid, H. rosa-sinensis x schizopetalus?). Enewetak Islet: Mar. 1975, near chapel (MPRL). Family APOCYANACEAE Plumeria obtusa L. Enewetak Islet: Feb. 1975, near buildings (MPRL). Plumeria rubra L. Enewetak Islet: Feb. 1975, near buildings, coll. by E. H. Bryan, Jr. (BPBM). Family VERBENACEAE Vitex trifolia L. Enewetak Islet: Feb. 1975, near buildings at southwest end of runway; also collected Feb. 12, 1975 by E. H. Bryan, Jr. (MPRL, BPBM). *Taxa are listed in the same order as in Tables 1 through 3. Specimens were collected by J. O. Lamberson, 1975 through 1977, except as noted, were determined by H. St. John and F. R. Fosberg, and are deposited in the U. S. National Museum Herbarium (USNM), the Bishop Museum Herbarium (BPBM), and the Mid-Pacific Research Laboratory Reference Collection (MPRL). the Pteridophyta correlates with that given by Fosberg et al. (1979, 1982). Thirty-six species (28.1% of the total) are introduced weeds, and 30 species (23.4%) are introduced ornamentals or food plants. Nineteen of these species occur only under cultivation and are considered transient members of the flora (Table 2). Seven species (5.5%) have been reported only as drift seeds found on beaches (Table 3). New records for the atoll reported in this chapter (Table 4) bring the total number of species recorded from the atoll to 128. Enewetak Atoll averages 125 to 150 cm of rain annu- ally and is much drier than atolls farther south, which are closer to the tropical convergence zone just north of the equator. Observations during the period 1975 to 1977 and meteorological records show that there is a definite seasonality on Enewetak: dry and windy from November through April, with more rain and less wind the other half of the year. The plants of Enewetak reflect this seasonality, produc- ing more flowers and fruit in summer and shedding some or most of their leaves in winter, depending on the species. Pisonia forests become open and light in the winter because Pisonia trees shed most of their leaves. This species flowered abundantly in January and February 1976, and the flowers were followed by the sticky fruits and new growth of leaves in March and April. Terminalia trees shed almost all of their leaves, which turn yellow and drop in the dry season. Cordia, Tournefortia, Guettarda, and Scaevola shed some of their leaves during the dry sea- son and produce more flowers and fruit during the summer months. Premna and Hibiscus tiliaceus shed many of their leaves in winter, and new growth is produced in April or May. Pemphis, Suriana, Cocos, and Morinda produce flowers and fruit year round, though the plants may appear more wind-blown and dry in winter, and the seeds usually germinate in early summer. Many of the ground plants and weedy species also reflect the seasonality. Annuals such as Conyza, Physalis, and some of the grasses are seen mostly in summer, as are Laportea and Portulaca. Ipomoea species are present year round but flower in summer. Tacca dies down in October or November and grows from the underground tubers in April or May, producing flowers in June and seeds in July. Cyperus species flower in summer, although Fimbristylus is in bloom year round. Euphorbia species are present year round, but Phyllanthus is much more abundant in summer. Waltheria also dies down in winter, and new growth is seen in April or May. Tropical storms, though uncommon in the Marshall Islands, do occasionally occur in the vicinity of Enewetak, and storm waves sometimes wash over the islands. The dense shrub growth at the top of most of the island beaches affords some protection to plants farther inland, but storms may wash away parts of the islands or damage the larger trees, especially Pisonia, which is soft and easily broken. A typhoon approached the atoll in the summer of 1977, causing some damage to the Pisonia on the leeward 24 LAMBERSON islands and sending storm waves over the northeast end of Enewetak Island to a depth of more than 1 m. HISTORY AND PREVIOUS OBSERVATIONS During the late 1800s and early 1900s, the people of Enewetak Atoll cultivated Tacca, breadfruit, coconut, Pandanus, and other plants on the atoll and traded copra to the Germans. Some of the coconut trees planted in neat rows during German times still survive and produce coconuts. German soil used as ballast in ships was de- posited on the island of Japtan to promote coconut growth. Plants and plant parts were utilized extensively in the Marshall Islands for food, drink, building materials (for houses and canoes), utensils, clothing, toys, medicine, and other uses (Anon., 1951; Bryan, 1972). In 1914, Japan acquired Enewetak as a result of a World War I mandate, and a Japanese trader and his assis- tants resided on the atoll. The first known botanical collec- tion from Enewetak was one of Pandanus made by Ohba in 1934 and published by Kanehira (1935). In the early 1940s, the Japanese cleared parts of some of the islands for construction of military fortifications, including an air- field on Enjebi. In 1944, the atoll was attacked and captured by United States forces. The heavy shelling during the battle and subsequent clearing and grading resulted in large-scale destruction of the vegetation on Enjebi, Medren (then called Parry), and Enewetak Islands. Bryan (1944) described the atoll’s vegetation at that time and noted that Medren resembled a plowed field with almost no plants at all. Bryan’s report and 1954 Army Map Service maps based on aerial photographs taken in 1944 indicate, how- ever, that some coconut trees were still present on several islands. In 1946, Fosberg visited the atoll and reported the results of his botanical survey and collection of Enewetak plants with those from other atolls in the Marshall Islands (Fosberg, 1955). Taylor (1950) described the marine and terrestrial vegetation and the plant species he encountered when he visited the atoll in 1946. At that time, Pisonia had been cut on Ikuren, Jedrol, and Alembel; military activity had also destroyed some of the vegetation on Lujor, Alembel, Runit, Japtan, and Jedrol; and the islands of Enewetak, Medren, and Enjebi had been reduced to “wastes of white sand” between buildings with a few weeds and scrubby coconuts. Runit, however, was com- pletely occupied by a formal coconut grove, except for a military installation at the south end. Aerial photographs taken in 1956 show that the flora of almost all the islands was affected by military activity during the nuclear test program conducted from 1948 to 1958. The islands of Louj through Boken (Irene) were almost bare, as were Runit, Boko, and Bokandretok. Enewetak, Medren, and Enjebi were covered with build- ings, and only a few plants grew on these islands among the structures and roadways. Most of the other islands sup- ported scrub vegetation or were partially cleared, and only the leeward islands from Ikuren through Biken were rela- tively undisturbed. The effects of the nuclear test program on the biota and the environment at Enewetak and Bikini were extensively studied by the Applied Fisheries Laboratory of the University of Washington, later renamed the Labora- tory of Radiation Biology. Their results were reported in UWEL series, in published papers, and in journalistic form in the book Proving Ground (Hines, 1962). An article by St. John (1960) provides a comprehen- sive record of the land plants found on the atoll up to that time. It gives a systematic account of species recorded, with notes on islands, collectors, and dates. Also included are a key to species, index to genera, and other data help- ful to an understanding of the flora of the atoll. The long-term effects on the vegetation because of the nuclear test program were primarily those resulting from destruction of habitats and the removal of soil and vegeta- tion from the islands (St. John, 1950; Biddulph and Cory, 1952: Palumbo, 1962). Some abnormalities, including tumors, calcium deficiency, growth irregularities, abnormal fruits and flowers, color abnormalities, and sterility, were found in plants near test sites (St. John, 1950; Biddulph, 1950; Biddulph and Cory, 1952; Biddulph and Biddulph, 1953). However, habitat disturbance and removal of the soil with loss of organic matter, inorganic nutrients, and water retention capability apparently had more lasting effects on the vegetation than the radioactivity. With the destruction of protective habitats, exposure to adverse environmental factors such as salt spray and drying winds was Greater. Following the conclusion of nuclear testing on Enewetak in 1958, the atoll was used for various other military and semi-military purposes during the 1960s and early 1970s. The Pacific Cratering Experiment (PACE) pro- gram bulldozed Aomon to the water table level and detonated nonnuclear explosives which left several small craters on the island. The Enewetak Marine Biological Laboratory (EMBL) was established in 1954. An extensive reference collection of atoll biota, including marine and terrestrial plants, has been gradually gathered, and the laboratory has provided facilities for a large number of visiting scientists. Specimens for the reference collection were contributed by G. R. Baker, E. H. Bryan, P. Colinvaux, V. Frey, M. Gilmartin, J. Lamberson, P. Lamberson, I. E. Lane, M. A. Lee, H. St. John, and R. Warner. Some botanical studies have been made through the laboratory and are described in the laboratory annual reports (Bryan and Lee, 1975; Lee and Lee, 1978). In 1957 to 1958, the University of Hawaii, the U. S. Weather Bureau, and the Joint Task Force Seven con- ducted microclimatic weather observations at Enewetak. During that study, Lane made botanical collections and notes and contributed a detailed section, which is included in that report, on the vegetation of some of the islands (Lane, 1960). HISTORY OF TERRESTRIAL VASCULAR PLANTS 25 Woodbury (1962) gathered ecological data on the atoll for the University of Utah from February to May 1962. His report includes very general descriptions of the vegeta- tion on the various islands at that time. In 1964, the Laboratory of Radiation Biology of the University of Washington conducted a radiological resurvey of Bikini and Enewetak Atolls (Welander et al., 1966). This report includes many photographs and descriptions of the vegetation on various islands, including notes on vege- tation recovery since the nuclear test days. Observations on vegetation have also been recorded in reports of studies on terrestrial radioisotope cycling (Jackson and Carpenter, 1967; Koranda et al., 1978), rats (Jackson, 1969; Fall et al., 1971), and birds (Carpenter et al., 1968). The Atomic Energy Commission conducted another radiological survey of the biota and environment of Enewetak in 1972 and 1973. The three-volume report includes color aerial photographs of each of the islands, as well as radiological data on the water, sediments, soil, and biota. Terrestrial biota is reported by Koranda et al. (1973). By 1975, the vegetation was gradually recovering on all of the islands and was growing over the remains of buildings and equipment left from the war and nuclear test days. Many of the plants which dominated the vegetation on the islands were introduced weeds, and a few ornamen- tal and crop plants were growing in private gardens. A photographic field guide to plants observed on the atoll from 1975 to 1977 has been compiled (Lamberson, 1982). In July 1975, a 12-acre plot of land was cleared on Enjebi for an experimental garden project to monitor the uptake of radionuclides from the soil into food plants. This long-term study is supported by the Department of Energy and is contracted to the Lawrence Livermore Laboratory (LLL) of Livermore, Calif. Radiologically “clean” soil from Jedrol was moved to Enjebi in 1975 for control studies. Breadfruit, pandanus, coconut, banana, sweet potato, lime, watermelon, and papaya were planted. On September 16, 1976, legal control of Enewetak Atoll was formally transferred from the U. S. government back to the people of Enewetak, with the agreement that radioactive debris and soil and nonradioactive scrap would be removed from inhabitable islands. In late 1976, federal funds were approved for the cleanup, and in 1977 the project was well under way. Some of the people of Enewetak returned from Ujelang Atoll to live on Japtan in March 1977. The program of rehabilitation and resettle- ment of the people included building of houses on Enewetak, Medren, and Japtan and replanting of coconut, banana, breadfruit, pandanus, and other crop plants. The effect of the cleanup on the vegetation of radiolog- ically contaminated islands has been very significant. Radioactive soil and debris were removed from many of the northern islands and deposited in a crater on Runit, which will be a permanently radiologically quarantined island. Reconstruction of housing on Enewetak and Lojwa to support cleanup personnel resulted in fairly extensive clearing of the vegetation, and the rehabilitation program involved extensive clearing and replanting of crops and food trees on the southeast islands from Ananij to Enewetak. Homes were built for the returning people of Enewetak on the islands of Japtan, Medren, and Enewetak, and the people returned to the atoll between 1977 and 1979. The radiological cleanup and the rehabilitation pro- grams were completed in 1980, but it will be years before the atoll again supports mature groves of coconut, bread- fruit, and pandanus. SUCCESSION AND OBSERVATION, 1975 TO 1977 After nuclear testing ceased at Enewetak Atoll in 1958, the vegetation was relatively undisturbed for almost 20 years, permitting the observation of plant distribution and succession under “natural,” if unusual, conditions. The scrap and radiological cleanup operations beginning in 1977 and the rehabilitation and replanting for the return of the people of Enewetak have resulted in large-scale altera- tions to the vegetation. It seems appropriate, therefore, to report on the condition of the vegetation before the cleanup, 1975 to 1977. Plant succession on the leeward islands, Ikuren through Biken, was followed through literature reports, aerial pho- tographs, personal communication, and observation. For convenience, several stages of succession were defined. Stage I, early pioneers, was seen on sand bars, spits, and small islands that were subject to storm damage and washover. Plants had to survive harsh conditions of intense sun, drying winds, and salt spray. They also had to over- come high salt concentration in the sand and the lack of water and nutrients. This stage was seen on narrow portions of Ikuren; on the small island between Elle and Bokenelab; on Bokinwotme; and on the small islands of Boko, Munjor, and Jinedrol. Plant genera present were Tournefortia, Scaevola, Lepturus, and Triumfetta and see- dlings of Guettarda, Morinda, Suriana, and Cocos (Fig. 1). As these plants become established, there is more pro- tection from wind and sun, and organic matter accumulates in the soil to provide nutrients and to hold moisture. In Stage Il, a thick scrub growth of mixed genera occurs, impenetrable without a sharp machete and _ pa- tience. This stage was seen on the eastern ends of Ikuren, the southeast end of Boken (Irwin), and on Bokandretok and formed the beach vegetation on many of the islands. It forms a protective barrier against salt spray and allows for development of vegetation inland on the larger islands. In addition to genera found in Stage I, vines such as Ipomoea, Canavalia, and Cassytha are present. Terminalia, Cordia, Pemphis, and Pandanus may also appear, and ground plants such as Lepturus and Triumfetta are con- fined to the margins (Fig. 2). In Stage III, the soil is more fertile, the trees are larger, and birds nesting in the branches add to the soil nutrients with their droppings and an occasional regurgitated fish. An open grassland develops under the trees, and ground 26 LAMBERSON as Ss ae ae on Fig. 1 Stage I, early pioneer plants. Plants seen in this photograph are Scaevola, Tournefortia, Suriana, Lepturus, and Triumfetta. A Na PE Fig. 2 Stage Il, scrub growth. Plants in this photograph are Scaevola and Tournefortia. HISTORY OF TERRESTRIAL VASCULAR PLANTS 27 plants such as Portulaca, Boerhavia, Laportea, and Sida appear. Pisonia may become a component of the young mixed forest, and vines are abundant. Cassytha may help to open up the forest by killing some trees. Larger plants present include Tournefortia, Scaevola, Cocos, Pandanus, Cordia, Terminalia, Morinda, Guettarda, Pisonia, Suriana, and Pemphis. This stage was found on the middle section of Ikuren, on Mut, on Kidrenen, and on the ocean side of Biken. It was also seen on Ananij, Van, the central portion of Alembel, and Bokenelab (Fig. 3). Fig. 3. Stage Ill, young mixed forest. Plants shown include Scaevola, Tournefortia, Guettarda, Pemphis, and Lepturus. 28 LAMBERSON In Stage IV, Pisonia and Cocos dominate the older mixed forest, and Scaevola is nearly shaded out. Morinda, Tournefortia, Guettarda, and Terminalia reach for sun, and Neisosperma may appear. The whole forest is more open, and the undergrowth consists of small Cocos and Pisonia with ground plants of Laportea, Boerhavia, Lepturus, and Portulaca. Sida and Scaevola are restricted to open areas, Suriana and Pemphis occur mostly at the forest margins, and vines become fewer, usually growing only in the sun in clearings and at the outer margins. This stage predom- inated on the main portions of Ikuren and Ananij (Fig.4). In Stage V, Pisonia takes over. Cocos trees become unproductive and gradually die out; other species of trees are confined to the edges of the forest. There is little Fig. 4 Stage IV, older mixed forest. Plants illustrated are predominantly Cocos and Pisonia. HISTORY OF TERRESTRIAL VASCULAR PLANTS 29 ground cover other than Pisonia seedlings and regenerated Stage Ill and common in Stage IV, become less abundant. branches. Nesting noddy terns fill the branches of the Stage V was seen on Biken and Kidrenen (Fig. 5). Pisonia trees, and the forest is open and easy to walk Portions of Japtan were between Stages IV and V. through. Coconut crabs, which are often present in Fig. 5 Stage V, Pisonia forest. Only Pisonia is seen in this photograph. 30 LAMBERSON THE ISLANDS, 1975 TO 1977 Between January 1975 and May 1977, the author visited most of the atoll islands to record vegetative data. An island-by-island summary of plant species collected or observed is presented in Table 5. The northern islands from Bokoluo to Boken were extensively disturbed by the nuclear test program. These islands had an open vegetation of scattered Scaevola and Tournefortia to 6 m in height with some Cordia, Guet- tarda, Cocos, and Pandanus with vines; a ground cover of Lepturus and Fimbristylis; and some Boerhavia, Cassytha, Triumfetta, Sida, Ipomoea, and Euphorbia (Stages | through Ill). Thirty-four plant species, other than those planted in the LLL garden project, were found on Enjebi. The gen- eral vegetation (was open to dense scrub of Tournefortia and Scaevola to.6 m in height, with occasional Pandanus, Pemphis, Morinda, and Cocos (modified Stage II to III). Cenchrus (reported extinct on this island in 1964), Lepturus, Fimbristylis, Cassytha, Euphorbia spp., Phyl- lanthus, Ipomoea spp., Pluchea spp., Stachytarpheta, Heliotropium, and Tridax were common, especially in the LLL garden. This island was once planted with coconuts and was the home of the people of Enjebi, but the coconut plantation was destroyed by Japanese and American mili- tary activities. Ten nuclear tests were conducted on or near the island, and during the test days the vegetation consisted of only grass, vines, and low shrubs. By 1964, however, Tournefortia trees were up to 6 m in height, and Scaevola almost to 5 m (Welander et al., 1966). Some Pisonia was reported in 1946 (Fosberg, 1955) and 1964 (Welander et al., 1966) but was not seen in 1975 to 1977. The islands of Mijikadrek through Lujor supported only a scrub vegetation, consisting predominately of Scaevola and Tournefortia (Stage II to Ill). Coconut trees were present on Bokenelab; a few sprouts had appeared on Lujor; and a grove of coconut trees with Pisonia occurred on Elle. Pisonia was also present on Mijikadrek, and its sticky fruits may eventually be transported by birds from there to other northern islands. Lepturus, Cassytha, Fim- bristylus, Triumfetta, Sida, and Ipomoea were fairly com- mon, and there was some Guettarda. Lujor had much Morinda, and Elle had a grove of Terminalia on the south end, ocean side. The island of Taiwel or Percy, between Kidrenen and Bokenelab, has been barren at least since 1944, and storm waves sometimes wash over it. It has a border of beachrock and may once have been vegetated, but the only evidence of plant life seen there from 1975 to 1977 was a sprouted coconut which had died. The islands of Eleleron through Bijile were once joined by causeways, though Eleleron was divided by a nuclear test in 1956, and part of it is now separate, although part is still joined to Aomon by a causeway. Aomon was bulldozed and pockmarked with small craters during the PACE program. There was an airstrip on Bijile, which was joined by a bridge to Lojwa, the site of the northern camp during the 1977 and 1978 cleanup operations. All of these islands had a mixed scrub vegetation of Tournefortia and Scaevola, with some Cocos, Pandanus, Pisonia (on Bijile), Suriana, Morinda, and Terminalia (on Aomon), Cordia, and Guettarda (Stage II to III). Pluchea was common, as were Lepturus, Fimbristylus, Euphorbia, Triumfetta, Ipomoea, and Heliotropium. Stachytarpheta, Lippia, Eragrostis, Portu- laca, Cassytha, and Sida were present but uncommon. Aomon was occupied by the Marshallese people during the war and postwar period, and in 1977 Tacca still grew on the ocean side, near stands of Hibiscus tiliaceus and Wollastonia_ biflora. In 1977, wedge-tailed shearwaters were found nesting in burrows near Pisonia trees on Bijile, and Laportea and Tacca were seen near the nesting area. Alembel was relatively undisturbed and had a central mixed forest of Pisonia, Cocos, Scaevola, Tournefortia, Premna, Morinda, and Guettarda (Stage III). Cordia Was present on the lagoon side, and two large Pandanus trees—one bearing fruit—were at the northernmost end. The outer margin was a thick scrub growth, primarily of Tournefortia, Guettarda, and Scaevola, with scrub Pisonia and Pemphis on the ocean side, and Suriana on the lagoon side (Stage Il). Runit, which was extensively disturbed during the nuclear test program and is now a depository for radioac- tive debris from other islands, had an open vegetation of scattered large Tournefortia and Scaevola bushes and a few coconut trees (modified Stage II). There was a ground cover of Cassytha, Conyza, Heliotropium, Lepturus, Fim- bristylis, Euphorbia, Triumfetta, and Ipomoea pescaprae. A few individuals of Suriana, Pemphis, Morinda, Guettarda, and Pluchea were also seen. This island was bulldozed during the nuclear test days and was the site of 17 tests, including Lacrosse (1956) and Cactus (1958) which left fairly large craters at the north end. The small islands of Boko, Munjor, and Inedrol sup- ported a limited vegetation (Stage 1). Inedrol had only Lepturus and Tournefortia, and the others had those species plus Scaevola. Inedral and Van had more species, including Pisonia, Boerhavia, Ipomoea macrantha, and Guettarda. Inedral also had Suriana, and Van had a young mixed forest (Stage III) of coconut, Triumfetta, Cordia, and Canavalia. Ananij had a Cocos—Pisonia forest (Stage IV) in the middle of the wide portion of the island, with Lepturus and Boerhavia beneath the trees and Scaevola and Tournefortia around the edges of the old runway (Stage II). There were several Terminalia trees, plus a few Cordia, Suriana, and Guettarda. The consolidated rock bar extending oceanward perpendicular to the island was covered with Pemphis. Jinimi is an unusual little island which was covered with a dense mat of Boerhavia with scattered scrub Pisonia, Tournefortia, and Scaevola bushes. Portulaca and Fimbristylis were also common. The island was sometimes the site of a large nesting colony of sooty terns, which were also found nesting on Aomon. Japtan was partially cleared during the war and test days and in 1977 became the home of about 80 TABLE 5 Island-by-Island Summary of Plant Species Observed or Collected on Enewetak Atoll, 1975-1977, Excluding Garden Species HISTORY OF TERRESTRIAL VASCULAR PLANTS 31 pjjauay sysosbD1sz x x x x x % D pyOfiuqoos sijso418p1g x x & 3 ° Te) poIpul auisna|/z x x x x 3 fo) 3 Dq|D 0yd12z x 2 E i= fe) piabias DI4DpI5IG x x x Ss re) © wnup9ou! Wun|powsaq x a winoydAbap wniuajI0/4490q x x snpunjoi sniadaa x x x snssaidwos sniadid x uo]4j2pp uopoudd x winoybIsD WnulZ x x DJDp4s09gns vDIP4OD x x x x x x x x x x x x x x x x x sisualinuog DzAuod x x x x x Diafionu so90D x x x x x x x x x x xxx x x x x x x x x Peles atin ep x Diafian pgojo29075, x x pjDjful suojyD, x x numoig sniysuad x snyoulyda sniy2uaz x x x x x s1uofiylf DYyAssDD x x x x x x x x x x x x x x x x x x x DOIEDYJDI DI|DADUDZ) x x x Don DIuDIYIDIG x x ‘dds pianyi20g x x x x x x x x x x x x x x x x x * x x psojid suapig x 3 =x = ° 2s 2 < «= ED a te) v os CF Se 8D = = 2 ~ es = = wo S ees Os f= G5) ox o 2 2&2 23558255 ~£S ston o SBEQteseee?s§ §25c SSSSESSSEESE SSEMSERELQSSSE SERS SE FL Les TOMO! Sees SS SS Ceo see ese rEEESSS ®ssGgeU=x Persia JON, SSL GR8ARGSS Sut tnt¢mhicmemMes S¢S8 8S 802 S8eRH TABLE 5 (cont’d) 32 LAMBERSON pyofydyduids vayonjq x x x x x x DoIpul DaYyIN|q x x Ea a ed sIpupi6 pluosig x x x cs ES x x Dee DenDG x x x x x x pypjnBup sijosayq x ie sued snapwp snyjup]|44q x x x x x pinpiop siydwad | x x x x x x x eg me mee ‘dds snuppuog | x x x Mee 6 2 Be flue x x ajD4ayyl] WintyD4s2UDg x pyofizisoddo puuadsosiayy x x DYyOfiayl9 DpulLopy x x x * x x x x x x x x x x x piojfipou viddi7 x x x pjpydaz092na] DuaponaT cs suadas sninjdaq x x a £4 x x x x x x xxx Kx x x a ee a ed S!|D4apni Day.odpT x x x x x x apidp2-sad paowody] x x x x x x Ax xe x x DAYJUDIIDU D2OWOM] x x x x El et ke eS gg SJ EP ES 5 Eb Ed eg ce py ce snaonljiq snosiqipy x x suaquinsoid winido.jolazy x x x x x x x x psolzads Dp4pyan) x x x x x x x Pd x xxx x x x x x x xx x x x x DsOWAD snjAjsiuquiy | x x x x x -“x x x Be ob pyofiudy} viqsoydny x x x x x x pa fd puy piqsoydny | x x x >< x i ei sluossiuipy) viqioydny | x x x x x ———— SEE a ae = ce) Qe 2 ) ° Fr: 5 = 3 (= s BS) c = — iS s i“ 5 Sobeee Geese 4 8oe Seer ose ete race = 25 gece See PSSSSESS Eo ge SSS SSS SLSGSSSERERE SSS SSS pasion, | BOL IAARHSS ASH FHCaItHemS eS FSESSISGGzSaerva (Table continued on following page.) TABLE 5 (cont’d) DIO/fIq DIUOJSD/|O/, bjDqo}}44 Dijapa/yy, DOIpUul DLaYI]D/ DaJau|d DJUOLa/\ suaquinsoid pyafwiniy suaquinsoid xpp}4]. §j4bjNsu} aUYIDII4 pajuabip p}jofauino [, Dynjoaul paiony [, SISUBOLUDS DI/DUIWIa | Sap|ojpjadojuoa] D290 | DWI DUDING sjsuaajblup! pjaydupjAyonjg xo)|0f DpIs Dpb290} vjonaDIS winpnu winjojisg 1S4ayjnsibd WnWayjudJapnasd DIOfisnjgo DUWAadId DaaD1ajO DIDINUOd Dan] vaDjnj4og SI/D4ISND DIDjNJOg suaquinsoid winwaidéjod PeysiA JON, HISTORY OF TERRESTRIAL VASCULAR PLANTS 33 * x < rd x * tad x x x x x x x eK * x x x x * x bad Cr ee te cai teat iCall etait * * bd * * x ~~ MRK RK KK x x ~~ 6 Ke MK KK a et od * * * ta tad ad * x x Ke * x * x * * * * x x x bad x x MK * x x * bd rad * x x x x x * x * * * ad xx x x xX RK KK ad x tet peta hikeviteciataPetesetat tal x x 36 SS oe OS OS OM: OS bad * * * tad * * * x x ~*~ xX cad x x x * * * * IF = ) ve 2 rs ae 32 Bose = ra) sop TS Beoes 5 5 SEEWSSESEESE pSPoSERE OSS ER TESES ESE S EES Bee S888 SSe8 Snes eae RSs Sef S8eRisedern 34 LAMBERSON Marshallese people. It had a productive coconut plantation (established during the time of the German occupation) which was somewhat overgrown and a field of Tacca. The ocean side of the island was overgrown with scrub Scaevola and Tournefortia (Stage II), and the cleared area had a ground cover of Cenchrus and other grasses, sedges, Euphorbia spp., Physalis, Triumfetta, Portulaca, Phyl- lanthus, Conyza, Ipomoea, Stachytarpheta, Heliotropium, and Tridax. Pluchea was notably absent. The residents cleared the underbrush from the area around the buildings and planted Pandanus and bananas. There was also a mixed forest on the southwest third of the island dom- inated by Pisonia, with Neisosperma, Cocos, Morinda, Guettarda, Cordia, and Pandanus, with Boerhavia near the margins (Stage IV to V). Jedrol, a small island inside the deep pass, set back from the atoll rim, had a mature Pisonia forest with no Cocos but with Scaevola, Tournefortia, Terminalia, and Guettarda (Stage IV to V). Fimbristylis, Lepturus, and Ipomoea macrantha were common, and Portulaca and Boerhavia were present but less common. Some soil was removed from this island in 1975 for the Enjebi garden. Medren and Enewetak were completely cleared during the war and nuclear test days for construction of housing and other buildings. Medren was abandoned in 1969, and the buildings and equipment left there became overgrown with plants (modified Stage II). Large shrubs and trees of Tournefortia, Scaevola, and Cocos were growing, and Pluchea spp. were abundant. There were remains of gar- dens containing Coccoloba, Crinum, and Leucaena. There was a ground cover of various grasses (not including Cenchrus), sedges, Portulaca, Cassytha, Euphorbia spp., Phyllanthus, Triumfetta, and Ipomoea and introduced spe- cies such as Tridax, Lippia, Stachytarpheta, Conyza, Ver- nonia, Heliotropium, and Waltheria. Enewetak was less overgrown because it had been continually occupied but had even more introduced species, including the weeds Polypremum, Desmodium, and Bidens. Ornamentals in- cluded Crinum, Wedelia, Pseuderanthemum, Coccoloba, Plumeria, and Hibiscus, and gardens contained many more transient species of food plants and flowers not recorded for this report. Bokandretok is a small islet just north of Enewetak which may be reached from there on foot at low tide. It was barren and occupied by troops in the 1950s but in 1977 supported a dense cover of Scaevola and Tour- nefortia, with Lepturus and Triumfetta around the edges (Stage Il). The leeward islands, Ikuren through Ribewon, were lit- tle affected by the war and test days and were not involved in the cleanup and rehabilitation. There was a weather station on Kidrenen, and there were a few derelict barges and boats left along the shores, but the vegetation in 1977 was Stage Ill to IV mixed forest. Coconuts were abundant on Ikuren, Mut, and Kidrenen, but those on Mut were old and seemed to be dying out (Stage IV to V). Pisonia trees were present on all of the leeward islands, and there were some nearly pure stands of this species (Stage V) and of Pemphis on Ribewon and Boken. The successional stages described earlier were well illustrated on these islands. Biken, isolated 20 km along the reef southwest of Bokoluo, in 1977 had a forest dominated by Pisonia with some old coconut trees and open underneath (Stage V). There was a scrub Scaevola—Tournefortia border along the south side, with a few Guettarda and Morinda. Cenchrus, Sida, and Ipomoea were present but uncommon on the lagoon side, and on the north side there were Pandanus and a group of Terminalia trees. A grove ot old Cordia trees was found at the entrance to an overgrown road on the northwest side. The vegetation on this island was dam- aged by Oak test to the north in 1958. It was described in 1964 as thick and overgrown with Ipomoea macrantha making transit difficult (Welander et al., 1966), but in 1975 through 1977 it was open and easy to walk through. Ipomoea and Cassytha were still present but not abundant. The ground cover outside the Pisonia area included Boerhavia, Lepturus, and some Portulaca and Triumfetta. There were a number of small craters on the island and reef flat dating from World War II when the island was used as a target for bombing practice. Coconut crabs were abundant, and roosting immature frigate birds were often seen, as were many other species of birds. ACKNOWLEDGMENTS This report is based in part on observations made by the author during employment at the Mid-Pacific Marine Laboratory from January 1975 through May 1977. Species of plants reported as new records for Enewetak were identified by Harold St. John of the Bernice P. Bishop Museum, Honolulu, Hawaii, and by F. R. Fosberg of the National Museum of Natural History. The Mid- Pacific Research Laboratory is supported by the Division of Biomedical and Environmental Research of the Depart- ment of Energy under Contract No. AT(26-1)-628 to the University of Hawaii. The author’s sincere thanks are extended to Edwin H. Bryan, Jr., Harold St. John, and F. R. Fosberg for botanical information and help in identifi- cation, for their interest and support during the period of observation, and in the writing of this report. Appreciation also is extended to S. H. Sohmer and Darrel Herbst for many helpful comments in their review of the manuscript and to Peggy Gaines for typing it. REFERENCES Anonymous, 1951, The Agriculture of Arno Atoll, Marshall Islands, Atoll Res. Bull. No. 6, Pacific Science Board, Wash- ington, D. C. Biddulph, O., 1950, Section VI of Radiobiological Survey of Bikini, Eniwetok and Likiep Atolls, July-August, 1949, UWFL-23, Appl. Fish. Lab. Univ. Wash. __, and R. 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Oliver, 1982, Geographical Checklist of the Micronesian Pteridophyta and Gymnospermae, Micronesica, 18: 23-82. Hines, N. O., 1962, Proving Ground. An Account of the Radiobiological Studies in the Pacific, 1946-1961, University of Washington Press, Seattle. Jackson, W. B., 1969, Survival of Rats at Eniwetok Atoll, Pac. Sci., 23: 265-275. __, and M. L. Carpenter, 1967, Radioisotope Cycling in Terres- trial Communities at Eniwetok Atoll, in Proceedings of the Second National Symposium on Radioecology, Ann Arbor, May 15-17, 1967, Atomic Energy Commission, Oak Ridge, Tennessee, pp. 644-651. Kanehira, R., 1935, An Enumeration of Micronesian Plants, Kyushu Univ. Dep. Agric. J., 4: 237-464. Koranda, J. J., J. R. Martin, S. E. Thompson, Jr., M. L. Stuart, D. R. Mcintyre, and G. Potter, 1973, Enewetak Radiological Survey, NVO-140, U. S. Atomic Energy Commission, Las Vegas, Nevada, pp. 225-235. __, W. L. Robison, S. E. Thompson, and M. L. Stuart, 1978, Enewetak Radioecology Research Program: I. Ecological Studies on Enjebi Island 1975-1976, Lawrence Livermore Laboratory Report UCRL-52409-1, University of California, Livermore. Koyama, T., 1964, The Cyperaceae of Micronesia, Micronesica, 1: 59-117. Lamberson, J. O., 1982, A Guide to Terrestrial Plants of Enewetak Atoll, Pacific Scientific Information Center, Bernice P. Bishop Museum, Honolulu. Lane, I. E., 1960, in Microclimatic Observations at Eniwetok, D. I. Blumenstock and D. F. Rex, Atoll Research Bull., No. 71, Pacific Science Board, Washington, D. C., pp. 15-19. Lee, M. A., and J. S. Lee, 1978, Terrestrial Nutrient Cycling: Analysis of Compartments, Mid-Pacific Marine Laboratory Annual Report, 1975-1976, NVO 0628-2, University of Hawaii, Honolulu. Leopold, E. B., 1969, Miocene Pollen and Spore Flora of Eniwetok Atoll, Marshall Islands, U. S. Geol. Surv. Prof. Pap. 260-II. Morat, P., 1981, Note sur les Graminées de las Nouvelle- Caledonie. VI: Lepturopetium, genre nouveau endemique, Adansonia, 20: 377-381. Palumbo, R. F., 1962, Recovery of the Land Plants at Eniwetok Atoll Following a Nuclear Detonation, Radiation Biol., 1: 182-189. St. John, H., 1950, Flora of Engebi, Aomon-Biijire, and Runit Islands, Rep. UWFL-23, Sect. V, Appl. Fish. Lab. Univ. Wash. , 1960, Flora of Eniwetok Atoll, Pac. Sci., 14: 313-335. Taylor, W. R., 1950, Plants of Bikini and Other Northern Marshall Islands, University of Michigan Press, Ann Arbor. Tracey, J. I., dr., and H. S. Ladd, 1974, Quaternary History of Eniwetok and Bikini Atolls, Marshall Islands, in Proceedings of Second International Symposium on Coral Reefs, Great Bar- rier Reef Committee, Brisbane, Australia, pp. 537-550. Welander, A. D., K. Bonham, L. R. Donaldson, R. F. Palumbo, S. P. Gessel, F. G. Lowman, and W. B. Jackson, 1966, Bikini-Eniwetok Studies, 1964, Rep. UWFL-93, Lab. Radiation Biol. Univ. of Wash. Woodbury, A. M., 1962, A Review of the Ecology of Eniwetok Atoll, Pacific Ocean, Contr. No. DA-42-007-403-CML-427, Univ. Utah Inst. of Env. Biol. Res., U. S. Army Chem. Corps, Dugway Proving Ground. Chapter 4 Recent Foraminifera and Nonplanktonic Protozoans* E. H. CHAVE7F and DENNIS M. DEVANEY (deceased)+ tUniversity of Hawaii, Hawaii Undersea Research Laboratory, Honolulu, Hawaii 96822 {Bernice P. Bishop Museum, Honolulu, Hawaii 96817 FORAMINIFERA In 1946, during Operation Crossroads, 45 samples containing Recent Foraminifera were obtained in the Enewetak Lagoon from the shore to depths of 62 m (204 feet) and on the outer slopes at depths from 44 to 914 m (144 to 3000 feet) (Cushman et al., 1954). Of tne 204 species of foraminifera recorded at Enewetak, approxi- mately 60% were also collected from the reef flats, lagoons, and outer slopes of other Marshall Island atolls (Rongerik, Rongelap, Bikini) and Sylvania Guyot. Fourteen of the 18 new species and two new varieties described at that time from the Marshall Islands were found at Enewetak. In the 1954 study, the Marshall Island foraminiferal fauna was composed of nearly 85% species characteristic of the tropical Pacific, 6% new to science, and the balance undetermined. No significant differences in the foraminif- eral composition were evident between the four atolls sam- pled during the study. Compared with other unspecified Pacific islands, the absence of Baculogypsina spp., Calcarina calcar, and medium-to-large Rotalia, as well as the presence of a greater variety of miliolid species, was apparent (Cushman et al., 1954). General features of specific biotopes (reef flat, lagoon, outer slope, and deep waters) with their dominant or re- stricted foraminiferal components were given, together with specific station localities (op. cit.). Foraminifera from Enewetak were also recorded from borings at two drill sites on the atoll (Cole, 1957; Todd and Low, 1960). These sites include a large number of extant species. In fact, Cole (op. cit.) reported that samples ‘With the exception of planktonic foraminiferans, which are included in this chapter, other planktonic protozoans (dinoflagel- lates, radiolarians, and tintinnids) are presented in Chapter 20 (this volume). 37 from the surface to about 186 m (610 feet) contained only Recent species. The present list of Recent Enewetak Foraminifera recorded from these two papers includes 23 additional species that were listed among Recent Marshall Island species by Cushman et al. (1954). Todd (1960) included Enewetak among localities dis- cussed in the distribution of Calcarina spengleri and absence of members of the genus Baculogypsina. Lacking a representative identified collection of Fora- minifera and other protozoans at Enewetak, Mid-Pacific Research Laboratory (MPRL) contracted for such a collec- tion to be made in 1956. Approximately 140 foraminiferal species were determined from the 142 samples obtained from shallow water throughout the atoll. The seaward reef and reef flat genera were compared with those found in beach sands and plankton tows (Hirshfield et al., 1968). This work also compared the foraminiferal genera of Enewetak with that from smaller samples taken in Hawaii, Kwajalein, and the Solomon and Caroline Islands. The species list (unpublished report to MPRL by Hirshfield, Helson, and Charmatz, 1957) included several additions to the Enewetak fauna, six of which were also found by Chave in 1981. Subsequent work on Foraminifera at Enewetak included studies on the fine structure and morphology of Carterina spiculotesta (Lipps and Enrico, 1973; Deutsch and Lipps, 1976; Conger et al., 1977). Foraminiferan pre- dation by fishes and the methods of feeding by foraminif- erans were also investigated at Enewetak (Lipps and Delaca, 1980). The species, Amphistegina lessonii and A. obifera, reviewed by Halloch and Larsen (1979) included material from Enewetak. Showers and Atkinson (1979) worked at the atoll on the different forms of Rosalina globularis. The checklist of Recent Foraminifera and nonplank- tonic protozoans from Enewetak is presented in Table 1 and contains 280 species. The above-mentioned publica- tions list 233 of these species. Forty-seven new Enewetak records have been added to this list by Chave (Table 2). Fourteen of these species were listed by Cushman et al. (1954) from other Marshall Island atolls; 33 species are new to Enewetak and the Marshall Islands. CHAVE AND DEVANEY TABLE 1 Checklist of Recent Foraminifera and Nonplanktonic Protozoans* Phylum SARCODINA Class RETICULAREA Subclass GRANULORETICULOSIA Order FORAMINIFERIDA Family SACCAMMINIDAE Saccammina sp. Proteonina sp.: Cushman et al., 1954. Family HYPERAMMINIDAE Sagenina frondescens (Brady): Todd and Low, 1960. Family NOURIIDAE +Nouria polymorphinoides Heron-Allen and Earland: Cushman et al., 1954. Family LITUOLIDAE *Placopsilina bradyi Cushman and McCullouch, 1939. Family TEXTULARIIDAE Bigenerina nodosaria d’Orbigny: Todd and Low, 1960. +Bigenerina sp.: Cushman et al., 1954. Clavulina angularis d’Orbigny: Todd and Low, 1960. Clavulina pacifica Cushman: Cushman et al., 1954. Textularia agglutinans d’Orbigny: Cushman et al., 1954; Todd and Low, 1960. Textularia candeiana d’Orbigny: Cushman et al., 1954; Todd and Low, 1960. Textularia conica d’Orbigny: Cushman et al., 1954; Todd and Low, 1960. Textularia foliacea Heron-Allen and Earland: Cushman et al., 1954. Textularia foliacea var. oceanica Cushman: Cushman et al., 1954; Todd and Low, 1960. Textularia kerimbaensis Said: Todd and Low, 1960. Textularia milletti Cushman: Cushman et al., 1954. *Textularia sagitula var. fistulosa Brady, 1884. Textularia semialata Cushman: Cushman et al., 1954; Todd and Low, 1960. Valvulina davidiana Chapman: Cushman et al., 1954; Todd and Low, 1960. Family TROCHAMMINIDAE *Trochammina nitida Brady, 1884. +Trochammina cf. rotaliformia Wright: Cushman et al., 1954. Family ATAXOPHRAGMIIDAE +Gaudryina cf. pauperata Earland: Cushman et al., 1954. Gaudryina triangularis angulata Cushman: Todd and Low, 1960. G. triangularis var. angulata Cushman: Cushman et al., 1954. Gaudryina trullissata Todd: Cushman et al., 1954. Gaudryina (Siphogaudryina) rugulosa Chapman: Cushman et al., 1954: Todd and Low, 1960. Gaudryina (Siphogaudryina) siphonifera Brady: Todd and Low, 1960. Karreriella bradyi (Cushman): Cushman et al., 1954. Family FISCHERINIDAE Cornuspira planorbis Schultze: Cushman et al., 1954; Todd and Low, 1960. Planispirinella exigua (Brady). Planispirinia exigua (Brady): Cushman et al., 1954. Family NUBECULARIIDAE Nebecularia lacunensis Chapman: Cushman et al., 1954. *Nubeculina divaricata (Brady, 1879). *Ptychomiliola separans Eimer and Fickert, 1921. +Spiroloculina acescata Cushman: Cushman et al., 1954. Spiroloculina angulata Cushman: Cushman et al., 1954; Todd and Low, 1960. Spiroloculina clara Cushman: Cushman et al., 1954; Todd and Low, 1960. Spiroloculina clara var. lirata Cushman: Cushman et al., 1954; Todd and Low, 1960. Spiroloculina communis Cushman and Todd: Cushman et al., 1954; Todd and Low, 1960. Spiroloculina corrugata Cushman and Todd: Cushman et al., 1954; Todd and Low, 1960. Spiroloculina folium Todd: Todd and Low, 1960. Spiroloculina sp. A: Cushman et al., 1954. *Spiroloculina grateloupi d’Orbigny, 1839. Spiroloculina marshallana Todd: Cushman et al., 1954; Todd and Low, 1960. *New Enewetak record. +Not found in 1981 Enewetak collections. (This table continued on next page.) FORAMINIFERA AND NONPLANKTONIC PROTOZOANS TABLE 1 (cont'd) Family NUBECULARIIDAE (cont’d) Spiroloculina mayori Cushman: Cushman et al., 1954; Todd and Low, 1960. *Spiroloculina sp. B (of Cushman et al., 1954). Vertebralina striata d’Orbigny: Todd and Low, 1960. Wiesnerella auriculata (Egger): Cushman et al., 1954. Family MILIOLIDAE tArticulina elongata Cushman: Cushman et al., 1954. Articulina pacifica Cushman: Cushman et al., 1954; Todd and Low, 1960. * Articulina sagra d’Orbigny, 1839. Hauerina bradyi Cushman: Cushman et al., 1954. Hauerina diversa Cushman: Cushman et al., 1954; Todd and Low, 1960. Hauerina involuta Cushman: Cushman et al., 1954; Todd and Low, 1960. Hauerina milletti Cushman: Cushman et al., 1954. + Hauerina serrata Cushman: Cushman et al., 1954. Massilina planata Cushman: Cushman et al., 1954; Todd and Low, 1960. Miliolinella australis (Parr): Cushman et al., 1954; Todd and Low, 1960. Parrina bradyi (Millett): Cushman et al., 1954; Todd and Low, 1960. *Pseudomassalina australis var. reticulata (Heron-Allen and Earland, 1915). Pyrgo denticulata (Brady): Cushman et al., 1954; Todd and Low, 1960. Pyrgo denticulata var. striolata (Brady): Cushman et al., 1954. “Pyrgo fornasini Chapman and Parr, 1935. +Pyrgo lucernula (Schwager): Cushman et al., 1954. Pyrgo millettii (Cushman): Cushman et al., 1954. Quinqueloculina anguina var. arenata Said: Cushman et al., 1954. Quinqueloculina bidentata d’Orbigny: Cushman et al., 1954; Todd and Low, 1960. *Quinqueloculina bosciana d’Orbigny, 1839. Quinqueloculina sulcata d’Orbigny: Cushman et al., 1954; Todd and Low, 1960. Schlumbergerina alveoliniformis (Brady): Cushman et al., 1954; Todd and Low, 1960. Triloculina affinis d’Orbigny, 1839. Triloculina trigonula (Lamarck): Cushman et al., 1954; Todd and Low, 1960. Triloculina cf. bassensis Parr: Cushman et al., 1954. Triloculina bikiniensis Todd: Cushman et al., 1954. Triloculina cf. bicarinata d’Orbigny: Cushman et al., 1954. Triloculina earlandi Cushman: Cushman et al., 1954; Todd and Low, 1960. Triloculina involuta Todd: Cushman et al., 1954. Triloculina irregularis (d’Orbigny): Cushman et al., 1954; Todd and Low, 1960. Triloculina kerimbatica (Heron-Allen and Earland): Cushman et al., 1954; Todd and Low, 1960. *Triloculina linneiana d’Orbigny, 1839. Triloculina marshallana Todd: Cushman et al., 1954. Triloculina oblonga (Montagu): Todd and Low, 1960. Triloculina cf. oblonga (Montagu): Cushman et al., 1954. Triloculina sp. A: Cushman et al., 1954. Triloculina spinata Cushman: Cushman et al., 1954. Triloculina subplanciana Cushman: Todd and Low, 1960. Triloculina terquemiana (Brady): Cushman et al., 1954. Triloculina tricarinata d’Orbigny: Cushman et al., 1954. Triloculinella labiosa (d’Orbigny): Cushman et al., 1954. Miliolinella laboisa (d’Orbigny): Todd and Low, 1960. Family SORITIDAE Amphisorus hemptrichii (Ehrenberg, 1840). Marginopora vertebralis Blainville (in part): Cushman et al., 1954. Marginopora vertebralis Blainville: Cushman et al., 1954; Todd and Low, 1960. Monalysidium politum Chapman: Cushman et al., 1954; Todd and Low, 1960. Sorites marginalis (Lamarck): Cushman et al., 1954; Todd and Low, 1960. Spirolina acicularis (Batsch): Cushman et al., 1954. Spirolina arietina (Batsch): Cushman et al., 1954; Todd and Low, 1960. Family ALVEOLINIDAE Borelis pulchra (d’Orbigny). Neoalveolina pulchra (d’Orbigny): Cushman et al., 1954. Borelis schlumbergeri (Reichel): Cole, 1957. “New Enewetak record. t+Not found in 1981 Enewetak collections. (This table continued on next page.) 39 CHAVE AND DEVANEY TABLE 1 (cont'd) Family NODOSARIIDAE Bolivinella folia (Parker and Jones): Todd and Low, 1960. Bolivinella folia var. ornata Cushman: Cushman et al., 1954. *Dentalina filiformis (d’Orbigny, 1826). } Frondicularia sp. A of Boomgaart: Cushman et al., 1954. Lagena globosa (Montagu): Cushman et al., 1954. t Lagena spiralis Brady: Cushman et al., 1954. Tt Lagena striato-punctata Parker and Jones: Cushman et al., 1954. T Lagena williamsoni (Alcock): Cushman et al., 1954. Nodosaria pauciloculata Cushman: Cushman et al., 1954. Family VAGINULINIDAE *Astacolus pacifica (Cushman and Hanazawa, 1936). *Astacolus planulata Galloway and Whissler, 1927. Robulus gibbus (d’Orbigny): Cushman et al., 1954. Robulus limbosus (Reuss): Cushman et al., 1954. *Robulus orbicularis (d’Orbigny, 1826). Robulus sp. A.: Cushman et al., 1954. Family POLYMORPHINIDAE *Ramulina globulifera Brady, 1879. Family GLANDULINIDAE Fissurina circularis Todd: Cushman et al., 1954; Todd and Low, 1960. TFissurina formosa (Schwager): Cushman et al., 1960; Todd and Low, 1960. } Fissurina lacunata (Burrows and Holland): Cushman et al., 1954; Todd and Low, 1960. Fissurina marginato-perforata (Seguenza): Todd and Low, 1960. Fissurina sp. B: Cushman et al., 1954. Fissurina milletti Todd: Cushman et al., 1954. Fissurina orbignyana flinti (Cushman): Todd and Low, 1960. Fissurina sp. C: Cushman et al., 1954. Fissurina sp. A: Cushman et al., 1954. Family SPIRILLINIDAE Alanwoodia campanaeformis (Brady, 1884). Conicospirillina semi-involuta Cushman: Todd and Low, 1960. Planispirillina denticulogranulata (Chapman). Spirillina denticulo-granulata Chapman: Todd and Low, 1960. Planispirillina tuberculatolimbata (Chapman). Spirillina tuberculato-limbata Chapman: Cushman et al., 1954; Todd and Low, 1960. Spirillina decorata Brady: Todd and Low, 1960. Spirillina decorata Brady var. of Sidebottom: Cushman et al., 1954. *Spirillina inaegualis Brady, 1879. T Spirillina spinigera Chapman: Todd and Low, 1960. Spirillina vivipara var. densepunctata Cushman: Cushman et al., 1954; Todd and Low, 1960. Spirillina vivipara var. revertens Rhumbler: Cushman et al., 1954. Family CERATOBULIMINIDAE *Ceratobulimina pacifica Cushman and Harris, 1927. Hoeglundina flinti (Galloway and Whissler, 1927). Hoeglundina elegans d’Orbigny: Cushman et al., 1954. *Lamarckina scabra (Brady, 1884). *Mississippina concentrica (Parker and Jones, 1864). Family ROBERTINIDAE *Alliatina excentrica (di Napoli Alliata, 1952). Family TURRILINIDAE + Buliminella elegantissima (d’Orbigny): Cushman et al., 1954. Buliminella milletti Cushman: Cushman et al., 1954. Buliminoides williamsonianus (Brady): Cushman et al., 1954. Family SPHAEROIDINIDAE t Sphaeroidina bullioides d’Orbigny: Cushman et al., 1954. Family BOLIVINITIDAE *Bolivina abbreviata Heron-Allen and Earland, 1924. Bolivina compacta Sidebottom: Cushman et al., 1954; Todd and Low, 1960. *New Enewetak record. +Not found in 1981 Enewetak collections. (This table continued on next page.) FORAMINIFERA AND NONPLANKTONIC PROTOZOANS 41 TABLE 1 (cont'd) Family BOLIVINITIDAE (cont'd) Bolivina pseudopygmaea Cushman: Cushman et al., 1954; Todd and Low, 1960. Bolivina rhomboidalis (Millett): Cushman et al., 1954; Todd and Low, 1960. Bolivina robusta Brady: Cushman et al., 1954. *Bolivina spinescens Cushman, 1911. Bolivina striatula Cushman: Cushman et al., 1954; Todd and Low, 1960. Bolivina subexcavata Cushman and Wickenden: Cushman et al., 1954. *Bolivina vadascens Cushman, 1933. Rectobolivina dimorpha var. pacifica (Cushman) Siphogenerina dimorpha var. pacifica Cushman: Cushman et al., 1954. Family ISLANDIELLIDAE *Cassidulinoides sp. Family BULIMINIDAE Bulimina fijiensis Cushman: Cushman et al., 1954. Fijiella simplex (Cushman, 1929). Reussella simplex (Cushman): Cushman et al., 1954, Todd and Low, 1960. Fijiella spinulosa (Reuss, 1850). Reussella sp. A: (Cushman): Cushman et al., 1954; Todd and Low, 1960. Mimosina pacifica Cushman: Cushman et al., 1954. Family UVIGERINIDAE Siphogenerina raphana (Parker and Jones): Cushman et al., 1954; Todd and Low, 1960. Trifarina albatrossi var. ornata (Cushman). Angulogerina albatrossi var. ornata Cushman: Cushman et al., 1954. Trifarina carinata (Cushman, 1927). Trifarina bradyi Cushman: as figured by Cushman et al., 1954. *Trifarina bradyi Cushman, 1942. *Trifarina scrobiculata (Cushman, 1921). Uvigerina porrecta Brady: Cushman et al., 1954. t+ Uvigerina proboscidea Schwager: Cushman et al., 1954. Uvigerina proboscidea var. vadescens Cushman: Cushman et al., 1954. Family DISCORBIDAE Discorbinella subbertheloti (Cushman). Discorbis subbertheloti Cushman: Cushman et al., 1954. Epistominella pulchra (Cushman): Cushman et al., 1954; Todd and Low, 1960. Epistominella sp.: Cushman et al., 1954. Neoconorbina crustata (Cushman). Discorbis crustata Cushman: Cushman et al., 1954. Neoconorbina tuberocapitata (Chapman). Discorbis tuberocapitata (Chapman): Cushman et al., 1954. * Patellinella carinata Collins, 1958. * Patellinella fijiana Cushman, 1933. T+ Patellinella inconspicua (Brady): Cushman et al., 1954. Rosalina concinna (Brady). Discorbis concinna (Brady): Cushman et al., 1954. Rosalina floridana (Cushman, 1922). Discorbis opima Cushman: Cushman et al., 1954. *Rosalina globularis d’Orbigny, 1826. Rosalina micens (Cushman). Discorbis micens Cushman: Cushman et al., 1954. *Rosalina villardeboana d’Orbigny, 1839. *Rotorbinella lobatulus Parr, 1950. Rugidia corticata (Heron-Allen and Earland): Cushman et al., 1954. Rugidia (?) spinosa Cushman: Cushman et al., 1954. Tretomphalus bulloides (d’Orbigny), 1839. Tretomphalus planus Cushman: Cushman et al., 1954. *Tretomphalus millettii Heron-Allen and Earland, 1915. Valvulineria rugosa (d’Orbigny). Discorbis rugosa (d’Orbigny): Cushman et al., 1954. Rosalina rugosa d’Orbigny: Todd and Low, 1960. “New Enewetak record. +Not found in 1981 Enewetak collections. (This table continued on next page.) 42 CHAVE AND DEVANEY TABLE 1 (cont'd) Family EPONIDAE *Eponides berthelotianus (d’Orbigny, 1839). Eponides lateralis (Terquem): Todd and Low, 1960. Eponides repandus (Fichtel and Moll, 1798). Poreponides cribrorepandus Asano and Uicho: Todd and Low, 1960. Family GLABRATELLIDAE Glabratella patelliformis (Brady). Discorbis patelliformis (Brady): Cushman et al., 1954. Neoconorbina tabernacularis (Brady): Todd and Low, 1960. Heronallenia sp.: Cushman et al., 1954. Family PEGIDIIDAE Pegidia dubia (d’Orbigny): Todd and Low, 1960. Family EPISTOMARIIDAE Epistomaroides polystomelloides (Parker and Jones): Cushman et al., 1954; Todd and Low, 1960. Family SIPHONINIDAE Siphonina tubulosa Cushman: Cushman et al., 1954. Siphoninoides echinata (Brady): Cushman et al., 1954; Todd and Low, 1960. Family AMPHISTEGINIDAE Amphistegina bicirculata Larsen, 1976. Amphistegina radiata var. venosa (Fichtel and Moll): Cushman et al., 1954. Amphistegina madagascariensis d’Orbigny: Cushman et al., 1954 (in part). Amphistegina lessonii (d’Orbigny): Halloch and Larsen, 1979. Amphistegina madagascariensis d’Orbigny: Cushman et al., 1954 (in part): Todd and Low, 1960 (in part?). Amphistegina lobifera Larsen: Halloch and Larsen, 1979. Ampbhistegina madagascariensis d’Orbigny: Cushman et al., 1954 (in part); Todd and Low, 1960 (in part?). Amphistegina papillosa Said. Ampbhistegina radiata var. papillosa Said: Cushman et al., 1954. Amphistegina sp. Amphistegina radiata (Fichtel and Moll): Cushman et al., 1954. Amphistegina radiata var.: Cushman et al., 1954. Family CIBICIDIDAE +Cibicides cicatricosus (Schwager): Cushman et al., 1954. Cibicides lobatulus (Walker and Jacob): Cushman et al., 1954; Todd and Low, 1960. Cibicides mayori (Cushman): Todd and Low, 1960. Cibicides refulgens Montfort: Todd and Low, 1960. Cibicidella variabilis (d’Orbigny): Cushman et al., 1954; Todd and Low, 1960. *Planulina ariminensis d’Orbigny, 1826. Planorbulinoides retinaculatus (Parker and Jones): Todd and Low, 1960. Family PLANORBULINIDAE Planorbulina acervalis Brady: Cushman et al., 1954; Todd and Low, 1960. Planorbulinella larvata (Parker and Jones): Todd and Low, 1960. Family ACERVULINIDAE Acervulina inhaerens Schultze: Cushman et al., 1954; Todd and Low, 1960. Gypsina globula (Reuss): Cushman et al., 1954; Todd and Low, 1960. Gypsina plana Carter: Todd and Low, 1960. Gypsina vesicularis (Parker and Jones) Cushman et al., 1954; Cole, 1957, Todd and Low, 1960. *Planogypsina squamiformis (Chapman, 1900). Family CYMBALOPORIDAE *Cymbaloporella tabellaeformis (Brady, 1884). Cymbaloporetta bradyi (Cushman): Cushman et al., 1954; Todd and Low, 1960. Cymbaloporetta squammosa (d’Orbigny): Cushman et al., 1954; Todd and Low, 1960. Pyropilus rotundatus Cushman: Cushman et al., 1954; Todd and Low, 1960. Family HOMOTREMIDAE Biarritzina monticularis (Carter). Carpenteria monticularis Carter: Todd and Low, 1960. Homotrema rubrum (Lamarck): Cushman et al., 1954; Todd and Low, 1960. Miniacina miniacea (Pallas): Cushman et al., 1954. +Family GLOBOROTALIIDAE Globorotalia crassaformis Galloway and Whissler, 1927. “New Enewetak record. +Not found in 1981 Enewetak collections. +Species names of family updated but not studied in 1981. (This table continued on next page.) FORAMINIFERA AND NONPLANKTONIC PROTOZOANS TABLE 1 (cont’d) Family GLOBOROTALIIDAE (cont’d) Globorotalia crassa (d’Orbigny): Cushman et al., 1954. Globorotalia cf. hirsuta (d’Orbigny): Cushman et al., 1954. Globorotalia menardii (d’Orbigny): Cushman et al., 1954; Todd and Low, 1960. Globorotalia truncatulinoides (d’Orbigny): Cushman et al., 1954. Globorotalia tumida (Brady): Cushman et al., 1954. +Family HANKENTINIDAE Hastigerina siphonifera (d’Orbigny, 1839). Globogerinella aequilateralis (Brady): Cushman et al., 1954. +Family GLOBIGERINIDAE Candeina nitida d’Orbigny: Cushman et al., 1954. Globigerina bulloides d’Orbigny: Cushman et al., 1954; Todd and Low, 1960. Globigerina eggeri Rhumbler: Todd and Low, 1960. Globogerina subcretacea Lomnicki: Cushman et al., 1954. Globigerina? sp.: Cushman et al., 1954. Globigerinoides conglobatus (Brady): Cushman et al., 1954; Todd and Low, 1960. Globigerinoides ruber (d’Orbigny): Cushman et al., 1954; Todd and Low, 1960. Globigerinoides sacculifer (Brady): Cushman et al., 1954; Todd and Low, 1960. Hastigerinella digitata (Rhumbler): Cushman et al., 1954. Orbulina universa d’Orbigny: Cushman et al., 1954; Todd and Low, 1960. Pulleniatina obliquiloculata (Parker and Jones): Cushman et al., 1954; Todd and Low, 1960. Sphaeroidinella dehiscens (Parker and Jones): Cushman et al., 1954. Family ROTALIIDAE Ammonia beccarii tepida (Cushman). Rotalia cf. beccarii var. tepida Cushman: Cushman et al., 1954. Streblus beccarii tepida (Cushman): Todd and Low, 1960. *Carpenteria balaniformis Gray, 1858. Carpenteria utricularis (Carter, 1876). Carpenteria proteiformis Goes: Todd and Low, 1960. Family CALCARINIDAE Calcarina hispida Brady: Cushman et al., 1954; Todd and Low, 1960; Todd, 1960. Calcarina spengleri (Gmelin): Cushman et al., 1954; Todd and Low, 1960; Todd, 1960. Family ELPHIDIDAE Elphidium advenum (Cushman): Cushman et al., 1954; Todd and Low, 1960. Elphidium advenum var. dispar Cushman: Cushman et al., 1954. Elphidium jenseni (Cushman): Cushman et al., 1954; Todd and Low, 1960. *Elphidium oceanicum Cushman, 1933. Elphidium simplex Cushman: Cushman et al., 1954; Todd and Low, 1960. Family NUMMULITIDAE *Cycloclypeus carpenteri Brady, 1856. Heterostegina suborbicularis d’Orbigny: Cushman et al., 1954; Cole, 1957; Todd and Low, 1960. Operculina ammonoidea (Gronovius): Cushman et al., 1954. Family CAUCASINIDAE Cassidella earlandi (Cushman). Virgulina earlandi Cushman: Cushman et al., 1954. Sigmavirgulina tortuosa (Brady). Bolivina tortuosa Brady: Cushman et al., 1954; Todd and Low, 1960. Family LOXOSTOMATIDAE Loxostomum convallarium (Millett): Cushman et al., 1954. Loxostomum karrerianum (Brady): Todd and Low, 1960. Loxostomum limbatum (Brady): Cushman et al., 1954; Todd and Low, 1960. Loxostomum mayori (Cushman): Cushman et al., 1954. Family CASSIDULINIDAE + Cassidulina angulosa Cushman: Cushman et al., 1954. Cassidulina costatula Cushman: Cushman et al., 1954. *Cassidulina crassa d’Orbigny, 1839. *Cassidulina gemma Todd: Todd, 1954. Cassidulina marshallana Todd: Cushman et al., 1954. *New Enewetak record. +Not found in 1981 Enewetak collections. Species names of family updated but not studied in 1981. (This table continued on next page.) 43 44 CHAVE AND DEVANEY TABLE 1 (cont’d) Family CASSIDULINIDAE (cont'd) Cassidulina minuta Cushman: Cushman et al., 1954. *Cassidulina murrhyna Schwager, 1866. Cassidulina oriangulata Belford, 1966. Cassidulina subglobosa Brady: Cushman et al., 1954 (in part). Cassidulina patula Cushman: Cushman et al., 1954. Cassidulina sulcata Belford, 1966. Cassidulina delicata Cushman: Cushman et al., 1954. + Cassidulina subglobosa Brady: Cushman et al., 1954. +Cassidulina subglobosa Brady var: Cushman et al., 1954. Ehrenbergina pacifica Cushman: Cushman et al., 1954. *Ehrenbergina reticulata Cushman, 1933.° Family NONIONIDAE * Astrononion tumidum Cushman and Edwards: Cushman et al., 1954. Nonion pacificum (Cushman): Todd and Low, 1960. t+ Nonionella sp. A: Cushman et al., 1954. Nonionella translucens Cushman: Cushman et al., 1954. + Pullenia salisburyi Stewart and Stewart: Cushman et al., 1954. Family ALABAMINIDAE Alabamina tubulifera (Heron-Allen and Earland). Epistominella tubulifera (Heron-Allen and Earland): Cushman et al., 1954; Todd and Low, 1960. + Gyroidina soldanii d’Orbigny: Cushman et al., 1954. + Oridorsalis tenera (Brady). Eponides tenera (Brady): Cushman et al., 1954. Family GAVELINELLIDAE Anomalina cf. glabrata Cushman: Cushman et al., 1954. Anomalinella rostrata (Brady): Cushman et al., 1954; Todd and Low, 1960. Cibicidoides pseudoungerianus (Cushman). Cibicides cf. pseudoungerianus (Cushman): Cushman et al., 1954. Family CARTERINIDAE Carterina spiculotesta (Carter): Cushman et al., 1954; Lipps and Enrico, 1973; Deutsch and Lipps, 1976; Conger et al., 1977. Phylum CILIOPHORA Class KINETOFRAGMINOPHORA Subclass GYMNOSTOMATA Order HAPTORIDA Lacrymaria sp.: Hirshfield et al., 1957. Order PLEUROSTOMATIDAE Loxophyllum(?) sp.: Hirshfield et al., 1957. Subclass VESTIBULIFERA Order COLPODIDA Colpoda sp.: Hirshfield et al., 1957. Class OLIGOHYMENOPHORA Subclass HYMENOSTOMATA Order SCUTICOCILIATA Cohnilembus verminus: Hirshfieid et al., 1957; Thompson, 1968. Cyclidium sp.: Hirshfield et al., 1957. Pseudochonilembus sp.: Hirshfield et al., 1957. Uronema sp.: Hirshfield et al., 1957. Subclass PERITRICHA Order PERITRICHIDA Syphidia sp.: Hirshfield et al., 1957. Vorticella sp.: Hirshfield et al., 1957. Class POLYHYMENOPHORA Subclass SPIROTRICHA Order HYPOTRICHIDA Euplotes sp.: Hirshfield et al., 1957. Stylonychia sp.: Hirshfield et al., 1957. *New Enewetak record. +Not found in 1981 Enewetak collections. +Species names of family updated but not studied in 1981. FORAMINIFERA AND NONPLANKTONIC PROTOZOANS TABLE 2 New Enewetak Records Species Location*® Depth Substrate Bolivina abbreviata L 50 m_s Fine sand Bolivina spinescens 1 52 m_ Fine sand Bolivina vadascens L 5 to 23 m__ Filamentous algae OS 100 to 140 m__Macro-algae Cassidulinoides sp. OS 400 m_ Coarse sediment Trifarina bradyi OS 400 m_ Coarse sediment Trifarina scrobiculata OS 400 m_ Coarse sediment Patellinella carinata OS 200 m_ Coarse sediment Patellinella fijiana L 55 m_ Filamentous algae Rosalina globularis CH 20 to 27 m___ Filamentous and macro-algae OS 80 to 100 m___ Filamentous and macro-algae Rosalina vilardeboana L 52 m_ Filamentous algae OS 80 to 140 m___ Filamentous and macro-algae Rotorbinella lobatulus EE 2to5 m_ Macro-algae OS 100 m_ Coarse sediment Tretomphalus millettii ie 16 m_ Algae OS 80 to 400 m_—__ Sand and algae on rocks Eponides berthelotianus OS 200 to 230 m___ Rock crevices Planulina ariminesis OS 80 to 140 m_ Rocks Planogypsina squamiformis OS 80 to 300 m___— Rocks and coarse sand Cymbaloporella tabellaeformis IE 13 m_ Rock with macro-algae Carpenteria balaniformis OSs 80 to 230 m___ Rocks Elphidium oceanicum L 60 m_ Fine sand Cycloclypeus carpenteri OS 140 to 200 m___ Rock crevices Cassidulina crassa OS 80 to 140 m_ Coarse sand, rock crevices Cassidulina gemma L 55 to60 m_s— Fine sand OS 80 to 140 m__ Rock crevices Cassidulina murrhyna OS 230 to 400 m_ Coarse sand Ehrenbergina reticulata OS 400 m_ Coarse sand Placopsilina bradyi OS 140 to 230 m__ Rocks Textularia sagitula var fistulosa OS 2m _ Macro-algae Trochamina nitida OS 300 m_ Coarse sand Nubeculina divaricata Os 300 m_ Coarse sand Ptychomiliola separans OS 12 m_ Fine sand Spiroloculina grateloupi L 55 m_ Fine sand Spiroloculina sp. B Os 100 to 200 m__ Rock crevices Articulina sagra OS 80 m_ Algae on rock Pseudomassalina australis var L 52to55 m_ Macro-algae reticulata CH 20 to 27 m__ Filamentous algae Pyrgo fornasini OS 400 m_ Coarse sand Quinqueloculina bosciana IL 52 to55 m__ Filamentous algae Quinqueloculina lamarckiana IL 15 to 16m _ Filamentous and macro-algae Quinqueloculina polygona Os 140 m___ Rock crevice Triloculina linneiana CH 2to 33 m_ Algae on rocks Dentalina filiformis OS 400 m_ Coarse sand Astacolus pacifica OS 300 to 400 m_ Coarse sand Astacolus planulata Os 400 m_ Coarse sand Robulus orbicularis OS 300 to 400 m_ Coarse sand Ramulina globulifera OS 400 m_ Coarse sand OS 200 to 300 m_ Coarse sand Spirillina inaequalis *L, lagoon; CH, channel; OS, outer slope. +Living form (followed by color of protoplasm): encrusting, test cemented to substrate; attached, holding on to substrate with pseudo- pods; free, not attached to substrate. +Total number of intact specimens in all 42 samples, each containing 1 cc of sand or scraped material. (This table continued on next page.) Formt Attached (brown) Attached (gray) Attached (brown) Attached, free (brown) Attached (brown) Attached (brown) Attached, free (brown) Attached, free (brown) Free (brown) Free Attached (light brown) Attached (reddish brown) Encrusting (brown) Encrusting (grayish brown) Attached (white) Attached (brown) Attached (grayish brown) Encrusting (tan) Attached (reddish brown) Attached (pink) Attached (brown) Free (red) Free (pink) 45 Number NNN AANNe Nore Lom) pe Onn Ww w _ NF PNHNM WWHWH DORR PR HWE DH NANWNH NNOWW NRKPORON DKW —" —y _ 46 CHAVE AND DEVANEY TABLE 2 (Cont’d) Species Location* Depth Substrate Formt Number Ceratobulimina pacifica OS 80 to 300 m__ Crevices in rocks Attached (brown) 4 Lamarckina scabra OS 200 m_ Rocks Encrusting (reddish brown) 6 Mississippina concentrica OS 300 m Coarse sand 1 Alliatina excentrica OS 400 m_ Coarse sand 2 *L, lagoon; CH, channel; OS, outer slope. +Living form (followed by color protoplasm): encrusting, test cemented to substrate; attached, holding on to substrate with pseudo- pods; free, not attached to substrate. +Total number of intact specimens in all 42 samples, each containing 1 cc of sand or scraped material. The taxonomy to the generic level is based on Loeblich and Tappan (1964, 1974, 1981). Species were identified by Chave using Ellis and Messina (1940 et seq.) and a large number of publications of which Graham and Mili- tante (1959), Barker (1960), Todd (1965), and Belford (1966) were especially useful. CILIOPHORA Several marine ciliates were collected at Enewetak from reef flat scum (Hirshfield et al., 1957). In addition, Scyphidium sp. was attached to unspecified invertebrates on the reef, and Cyclidium sp. was found with habitat unspecified (Hirshfield, op. cit.). Observations made by Patrick Colin from 1979 to 1980 off Runit Islet revealed discolored brownish areas on the lagoon bottom, which were found by Devaney to be formed by high concentra- tions of an undetermined peritrich ciliate that resembles Scyphidia. During 1966, 42 hymenostomme ciliates were col- lected from the ocean and lagoon sides of nine islets at Enewetak Atoll. Permanent silver-impregnated slides were made of these protozoans from cultures (Thompson and Sellers, 1967), but species were not listed. Subsequently, Thompson (1968) described Cohnilembus verminus from Enewetak. Berger (1964) noted that undetermined endocommen- sal ciliates were present in several unspecified regular sea urchins but were absent from at least two species (Euci- daris metularia and Parasalinia gratiosa) and from irregular urchins and other unspecified echinoderms examined. The classification for ciliophorans follows Corliss (1977). OTHER PROTOZOANS Fresh and brackish water protozoans at Enewetak revealed mainly amoeboid types from nearly 100 water and soil sites on nine islets. Additional fish, gecko, rat and bird blood, and other tissues were examined for protozoan parasites (Dillon and Kasweck, 1970). NEW ENEWETAK RECORDS In 1981, 42 samples were studied at Enewetak immediately after being collected from the lagoon at depths of from 1 to 61 m, in the channels from 1 to 33 m, and on the outer slopes from 1 to 400 m. Of the new Enewetak records, 23 species were represented by one to three specimens and 14 species were fairly com- mon but were obtained by methods not used during the 1946 collections (hard surfaces were scraped by hand to depths of 33 m, and rocks were brought up from deep areas by the submersible Makili’i). The final 10 species were common in sediment samples and were not reported by Cushman et al. (1954). Twenty-nine species listed by Cushman et al. (1954) were not found by the senior author. Twenty-seven of these species were listed as rare or from the 914 m sta- tion (op. cit.). Two species, Hauerina serrata and Astrononion tumidum, were listed as common in lagoon sediments (op. cit.). ACKNOWLEDGMENTS We acknowledge and are grateful to H. I. Hirshfield, New York University, for permission to include the records of ciliates given in his 1957 report to MPRL. J. Resig at the University of Hawaii made valuable suggestions and corrections in the Foraminifera part of this paper. The facilities at Enewetak and material obtained by the Hawaii Undersea Research Laboratory’s submersible Makali’i were made available to both authors through grants to Patrick Colin and John Harrison of MPRL from NOAA’s National Undersea Research Program, the Department of Energy, and the Defense Nuclear Agency. REFERENCES Barker, R. W., 1960, Taxonomic Notes on the Species Figured by H. B. Brady in his Report on the Foraminifera Dredged by the HMS Challenger During the Years 1873-1876, S.E.P.M. Spec. Pub. 9, Tulsa, Oklahoma, 238 pp. Belford, D. J., 1966, Miocene and Pliocene Smaller Forami- nifera from Papua and New Guinea, Austr. Bur. Min. Res. Geol. Geoph. Bull. 79, 306 pp. Berger, J., 1964, Mid-Pacific Research Laboratory Annual Rept., FY 1963-64, 3 pp. Cole, W. S., 1957, Larger Foraminifera from Eniwetok Atoll Drill Holes, U. S. Geol. Surv. Prof. Pap. 260-V, pp. 743-784. FORAMINIFERA AND NONPLANKTONIC PROTOZOANS 47 Conger, S. D., H. W. Green, and J. H. Lipps, 1977, Test Ultra- structure of Some Calcareous Foraminifera, J. Foramin. Res., 7: 278-196. Corliss, J. O., 1977, Annotated Assignment of Families and Gen- era to the Orders and Classes Currently Comprising the Corlissian Scheme of Higher Classification for the Phylum Ciliophora, Trans. Am. Micros. Soc., 96: 104-140. Cushman, J. A., R. Todd, and R. J. Post, 1954, Recent Forami- nifera of the Marshall Islands, U. S. Geol. Surv. Prof. Pap. 260-H, pp. 319-384. Dillon, R. D., and K. L. Kasweck, 1970, Mid-Pacific Research Laboratory Annual Rept., FY 1970-71, 3 pp. Deutsch, S., and J. H. Lipps, 1976, Test Structure of the Fora- minifera Carterina, J. Paleontol., 50: 312-317. Ellis, B. F., and A. R. Messina, 1940 et seq., Catalogue of Foraminifera, Spec. Publ. Am. Mus. Nat. Hist., New York. Graham, J. J., and P. J. Militante, 1959, Recent Foraminifera from the Puerto Galera Area Northern Mindoro, Philippines, Stanford University Public., 6: 1-171. Halloch P., and A. R. Larsen, 1979, Coiling Direction in Amphistegina, Marine Micropaleo., 4: 33-44. Hirshfield, H. I., L. Helson, and R. Charmatz, 1957, Eniwetok Protozoa, Report to MPRL, 100 pp. __, R. Charmatz, and L. Helson, 1968, Foraminifera in Samples Taken Mainly from Eniwetok Atoll in 1956, J. Protozool., 15: 497-502. Lipps, J. H., and E. E. Delaca, 1980, Shallow-Water Forami- niferal Ecology, Pacific Ocean in Quaternary Depositional Environments of the Pacific Coast, M. Field and others (Eds.), S.E.P.M. Pacific Coast Paleogeography Symposium, 4: 325-340. Lipps, J. H., and R. J. Enrico, 1973, Biology and Ecology of the Strange Spicular Foraminiferan Carterina, Geol. Soc. Amer. Abs., 5: 73. Loeblich, A. R., and H. Tappan, 1964, Protista 2, Treatise on Invertebrate Paleontology, R. Moore (Ed.), Part C., University of Kansas Press, Lawrence, Kansas, 900 pp. ___, 1974, Recent Advances in the Classification of the Forami- nifera, Foraminifera Vol. 1, R. H. Hedley and C. G. Adams (Eds.), Academic Press, London, 53 pp. , 1981, Suprageneric Revisions of Some Calcareous Forami- nifera, J. Foram. Res., 11: 159-164. Maguire, B., Jr., 1967, Small Freshwater Organisms on Eniwetok Atoll, Micronesica, 3: 151-157. Showers, W. I., and M. Atkinson, 1979, Functional Morphology, Biometry and Life History of the Foraminifera Rosalina globularis, MPRL Report for FY 1979, pp. 80-90. Thompson, J. C., Jr., 1968, A Description of Cohnilembus ver- minus from Eniwetok Atoll, J. Protozool., 15: 396-399. __, and C. M. Sellers, 1967, Mid-Pacific Research Laboratory Annual Rept., FY 1966-67, 13pp. Todd, R., 1960, Some Observations on the Distribution of Calcarina and Baculogypsina in the Pacific, Sci. Repts. Tohoku Univ. Ser. 2, Geol. Spec., 4: 100-107. , 1965, The Foraminifera of the Tropical Pacific Collections of the “Albatross” 1899-1900. Part 4, Rotaliform Families and Planktonic Families, U.S.N.M. Bull. 161, 127 pp. __, and D. Low, 1960, Smaller Foraminifera from Eniwetok Drill Holes, Geol. Surv. Prof. Pap. 260-X, pp. 799-861. Porifera of Enewetak Atoll Compiled by DENNIS M. DEVANEY (deceased) Bernice P. Bishop Museum Honolulu, Hawaii 96817 The first sponges reported from Enewetak were based on collections made between 1946 and 1948 from a number of tropical Pacific areas (de Laubenfels, 1954). The Pacific Science Board of the National Research Coun- cil sponsored the collections project. Six of the 13 species recorded from Enewetak were collected by dredging near the center of the lagoon (8 km north of the south anchorage; approximately 11° 29’ N, 165° 15’ E, de Lau- benfels, 1954) at a depth of 35 m. Another species came from the reef flat, and six were from unrecorded localities. Three new species were recorded from Enewetak, but only one (Lissodendoryx calypta) was unique to that atoll. Three sponges, each prey for different species of dorid nudibranchs, have also been recorded from Enewetak (Young, 1967). The color, form, and habitat of these sponges were given. Beginning in 1972, research was initiated on the biochemistry of Enewetak sponges (Middlebrook et al., 1972; Scheuer et al., 1974; Ravi and Scheuer, 1975; Scheuer et al., 1975; Chevolot et al., 1977; Yunker and Scheuer, 1978; Ravi et al., 1979). A sponge from the lagoon was used as a source of radionuclides concentrated in its siliceous spicules (Hurd and Lawson, 1975). The need for valid identification of sponges is clearly indicated in research where chemical studies have been made on species that remain undetermined (Chevolot et al., 1977; Yunker and Scheuer, 1978; Ravi et al., 1979). Attempts to get Enewetak sponge material identified have Chapter 5 resulted in 17 additional sponges receiving preliminary determinations by Klaus Ruetzler (personal communication, 1981). Examples of some or all of these species are de- posited at U. S. National Museum of Natural History (USNM), Mid-Pacific Research Laboratory (MPRL), and the B. P. Bishop Museum (BPBM). Brief descriptions (color, external morphology, colony form) and habitat information on these sponges were prepared by Janet Lamberson for MPRL. The presence of several boring clionid sponges in four selected Enewetak corals has been noted by Highsmith (1981). The boring organisms were the most common infaunal associates of the corals studied, with 86% of the corals showing sponge bioerosion effects. A calcarean “ear sponge” with fossil affinity collected on the ocean side of Biken (Leroy) is close to Murrayona phaneolepis but might be a new species (Reutzler, personal communication, 1977). It occurs on the roofs of small caves with sclerosponges, is flattened, and is green in color. Another calcarean sponge, Neocoelia crypta, and the sclerosponge, Astrosclera willeyana, have also been identi- fied from Enewetak (Basile et al., in press). Kinzie (1973) found sclerosponges in a large crevasse that is considered a very prominent feature on the leeward side of Enewetak Atoll. Two species of sclerosponges were first described from Enewetak, as well as other parts of the tropical Indo-Pacific, by Hartman and Goreau (1975, 1976). The Enewetak sponge fauna remains too poorly known at this time to reach zoogeographical conclusions and awaits the work of someone with extensive field and taxo- nomic research experience in the region to clarify the systematics of this fauna. The classification scheme herein is based on Berquist (1978). Table 1 is a checklist of Enewetak Porifera. TABLE 1 Checklist of Porifera at Enewetak Atoll Class CALCAREA Subclass CALCINEA Order LEUCELLIDA Family LEUCELLIDAE *Leucetta sp. Subclass PHARETRONIDA Order INOZOA Family MURRAYONIDAE *Murrayona cf. M. phanolepis Kirkpatrick, 1910. “New Enewetak record. (This table continued on next page.) DEVANEY TABLE 1 (cont'd) Order SPHINCTOZOA Family CRYPTOCOELIIDAE *Neocoelia crypta Vacelet, 1977. Class DEMOSPONGIAE Subclass TETRACTINOMORPHA Order CHORISTIDA Family STELLETIDAE Ancorina acervus (Bowerbank, 1862). Hezekia walkerri de Laubenfels, 1954. Family JASPIDAE Jaspis stellifera (Carter): de Laubenfels, 1954. Order SPIROPHORIDA Family TETILLIDAE *Cinachyra sp. Order HADROMERIDA Family SUBERITIDAE Terpios aploos de Laubenfels: de Laubenfels, 1954. Young, 1967. Family SPIRASTRELLIDAE Spirastrella decumbens Ridley: de Laubenfels, 1954. Spirastrella vagabunda Ridley, 1884. Anthosigmella vagabunda (Ridley): de Laubenfels, 1954. Family CLIONIDAE Aka sp. cf. A. diagonoxea Thomas: Highsmith, 1981. Aka spp. (3): Highsmith, 1981. Cliona mucronata Sollas: Highsmith, 1981. Cliona sp. cf. C. quadrata Hancock: Highsmith, 1981. Cliona sp. cf. C. viridis Schmidt: Highsmith, 1981. Cliothosa hancocki Topsent: Highsmith, 1981. Family TETHYIDAE Aaptos unispiculus (Carter): de Laubenfels, 1954. *Tethya sp. Family CHONDROSIIDAE Chondrilla australiensis Carter: de Laubenfels, 1954. * Chondrosia sp. Chondrosia (?) chucalla: Ravi et al., 1979. Order AXINELLIDA Family AGELASIDAE Agelas mauritiana (Carter): de Laubenfels, 1954. Subclass CERACTINOMORPHA Order HALICHONDRIDA Family HYMENIACIDONIDAE Prianos phlox de Laubenfels: Young, 1967. Order POECILOSCLERIDA Family CRELLIDAE *Grayella sp. *Yvesia spinulata (Hentschel, 1911). Family MYXILLIDAE Lissodendoryx calypta de Laubenfels: de Laubenefels, 1954. Family CLATHRIIDAE Clathria abietina (Lamarck): de Laubenfels, 1954. Order HAPLOSCLERIDA Family HALICLONIDAE *Haliclona sp. 1. *Haliclona sp. 2. *Toxadocia sp. Family NIPHATIDAE Cribrochalina olemda de Laubenfels, 1954. Family ADOCIIDAE * Adocia sp. Family CALLYSPONGIIDAE Callyspongia fistularia (Topsent): de Laubenfels, 1954. *Callyspongia (?) ridleyi Burton, 1934. “New Enewetak record. (This table continued on next page.) PORIFERA 51 TABLE 1 (cont'd) Order DICTYOCERATIDA Family DYSIDEIDAE *Dysidea (?) arenaria Bergquist, 1965. Family SPONGIIDAE Heteronema erecta Keller, 1889. Thorectopsamma mela de Laubenfels: de Laubenfels, 1954. *Heteronema sp. Ircinia halmiformis (Lendenfeld): de Laubenfels, 1954. Family DYSIDEIDAE *Megalopastas sp. Order DENDROCERATIDA Family APLYSILLIDAE Aplysilla gracialis de Laubenfels: Young, 1967. Order VERONGIDA Family VERONGIIDAE Psammaplysilla purpurea (Carter, 1880). Thorectopsamma xana de Laubenfels: de Laubenfels, 1954. Class SCLEROSPONGIAE Order CERATOPORELLIDA Family ASTROSCLERIDAE *Astrosclera willeyana Lister, 1900. Order TABULOSPONGIDA Family ACANTHOCHAETETIDAE Acanthochaetetes wellsi Hartman and Goreau: Hartman and Goreau, 1975. Stromatospongia micronesica Hartman and Goreau: Hartman and Goreau, 1976. *New Enewetak record. ACKNOWLEDGMENTS This review is more nearly complete due to Roger Cuf- fey (Pennsylvania State University) and especially Klaus Reutzler (USNM) who have permitted the inclusion of their species determinations which form the basis of new Enewetak records. Janet and Phil Lamberson, resident managers at MPRL in the late 1970s, were instrumental in collecting fresh material with notes that now permit field determination of at least some of the more common sponges at Enewetak. REFERENCES Basile, L. L., R. J. Cuffey, and D. F. Kosich, Sclerosponges, Pharetronids, and Sphinctozoans (Relic Cryptic Hard-Bodied Porifera) in the Modern Reefs of Enewetak Atoll, J. Paleont., in press. Bergquist, P. R., 1978, Sponges, University of California Press, Berkeley. Chevolot, L., S. Padua, B. N. Ravi, P. C. Blyth, and P. J. Scheuer, 1977, Isolation of 1-Methyl-4, 5-Dibromopyrrole- 2-Carboxylic Acid and Its 3’- (Hydantoyl) Propylamide (Midpa- camide) from a Marine Sponge, Heterocycles, 7: 891-894. De Laubenfels, M. W., 1954, The Sponges of the West-Central Pacific, Oregon State Monogr. Studies in Zool., No. 7, pp. 1-306. Hartman, W. D., and T. F. Goreau, 1975, A Pacific Tabulate Sponge, Living Representative of a New Order of Sclero- sponges, Postilla, 167: 1-13. —, 1976, A New Ceratoporellid Sponge (Porifera: Sclerospon- giae) from the Pacific, F. W. Harrison, and R. R. Cowdew (Eds.), pp. 329-347, Aspects of Sponge Biology, Academic Press, N. Y. Highsmith, R. C., 1981, Coral Bioerosion at Enewetak: Agents and Dynamics, Int. Revue Gesamen. Hydrobiol., 66: 335-375. Hurd, D. C., and D. S. Lawson, 1975, On the Possible Concen- tration of Radionuclides by the Siliceous Spicules from Enewetak Lagoon, MPML Ann. Rept. FY 1975. Kinzie, R. A. Ill, 1973, Sclerosponge Ecology, EMBL Ann. Rept. FY 1972-73. Middlebrook, R. E., L. W. Wittle, and E. D. Scura, 1972, Isola- tion and Purification of Naturally Occurring Compounds of Marine Organisms, EMBL Ann. Rept. FY 1971-72. Ravi, B. N., and P. J. Scheuer, 1975, Toxins and Biomedical Agents of Marine Invertebrates, MPML Ann. Rept. FY 1975. —_—, H. Perry Perzanowski, R. A. Ross, T. R. Erdman, and P. J. Scheuer, 1979, Recent Research in Marine Natural Products: the Puupehenones, Pure and Appl. Chem., 51: 1893-1900. Scheuer, P. J., B. J. Burreson, and M. G. Missakian, 1974, Antimicrobial and Antitumor Constituents of Sponges; etc., EMBL Ann. Rept. FY 1973-74. —_, R. F. Dietrich, and F. X. Woolard, 1975, Toxins and Biomedical Agents of Marine Invertebrates, MPML Ann. Rept. FY 1975. Young, D. K., 1967, New Records of Nudibranchia (Gastropoda: Opisthobranchia: Nudibranchia) from the Central and West- Central Pacific, Veliger, 10: 159-173. Yunker, M. B., and P. J. Scheuer, 1978, Mokupalides, Three Novel Hexaprenoids from a Marine Sponge, J. Amer. Chem. Soc., 100: 307-309. a ad Chapter 6 Sea Anemones of Enewetak Atoll CHARLES E. CUTRESS* and A. CHARLES ARNESONT “Department of Marine Sciences, University of Puerto Rico, Mayagiiez, Puerto Rico 00708 +Scripps Institution of Oceanography, La Jolla, California 92093 INTRODUCTION The sea anemones covered in this chapter were col- lected and/or photographed in part by Cutress during May and June 1955 and in part by Arneson from September through November 1980. Collection records are shown at the end of the chapter. Most identifications were made from preserved specimens, but a few were made from pho- tographs alone. Available photographs of Phymanthus strandesi and Physobrachia douglasi are inadequate. Heterodactyla hemprichi was not photographed. No published taxonomic work on Enewetak sea anemones exists. Dunn (1981), in a revision of the clown- fish anemones, refers by catalog numbers to preserved specimens from Enewetak which she examined. Allen (1972) refers to and illustrates seven species of host anemones that he observed at Enewetak from 1968 to 1971. Josephson (1966) made physiological observations on Calliactis polypus, and Johannes et al. (1972) refer to an unidentified digging anemone subsequently identified as Actinodendron plumosum. Table 1 provides a checklist of Enewetak sea anemones. TABLE 1 Checklist of Enewetak Sea Anemones Phylum CNIDARIA Class ANTHOZOA Order CORALLIMORPHARIA Family ACTINODISCIDAE Actinodiscus neglectus (Fowler, 1888) Rhodactis howesii Saville-Kent, 1893 Order ACTINIARIA Family EDWARDSIIDAE Edwardsia pudica Klunzinger, 1877 Edwardsia gilbertensis Carlgren, 1931 Family BOLOCEROIDIDAE Bunodeopsis medusoides (Fowler, 1888) Family ALICIIDAE Triactis producta Klunzinger, 1877 Family ACTINIIDAE Anthostella badia (Carlgren, 1900) Anthopleura nigrescens (Verrill, 1928) Physobrachia douglasi Saville-Kent, 1893: Allen, 1972, pp. 91, 166, 167, 169, 173, 177, 179 Fig. 95, 183, 227, 229, 252, 256, I: Entacmaea quadricolor (Rueppel and Leuckart) [pro parte], Dunn, 1981,p. 15, 27. Actiniogeton sesere (Haddon and Shackleton, 1893). Family ACTINODENDRONIDAE Actinodendron plumosum Haddon, 1898: Johannes et al., p. 542. Actinodendron glomeratum Haddon, 1898 Megalactis hemprichii Ehrenberg, 1834 53 (This table continued on next page.) 54 CUTRESS AND ARNESON TABLE 1 (cont'd) Family THALASSIANTHIDAE Heterodactyla hemprichii Ehrenberg, 1834 Cryptodendrum adhesivum Klunzinger, 1877: Dunn, 1981, p. 13. Family PHYMANTHIDAE Phymanthus strandesii Carlgren, 1900 Family STICHODACT YLIDAE Stichodactyla mertensii Brandt, 1835: Dunn (1981), p. 99: Stoichactis giganteum (Forsskal); Allen 1972, pp. 102, 105, 124, 166, 173, 178 Figs. 93-94, 184, 190, 193, 194 Fig. 102, 196, 200, 206 Fig 105, 237 Fig. 123, 252, 254, 255, 258, 259, Il Heteractis aurora (Quoy and Gaimard, 1833): Dunn, 1981, pp. 57, 66: Radianthus simplex (Haddon and Shackleton), Allen, 1972, pp. 124, 166, 177, 181, 190, 191 Fig. 100, 192 Fig 101, 193, 196, 197, 230, 250, 252, 258, 259, Il. Heteractis ritteri (Kwietniewski, 1897): Heteractis magnifica (Quoy and Gaimard) [pro parte], Dunn, 1981, p. 40: Radianthus ritteri (Kwietniewski): Allen, 1972, pp. 166, 169, 175 Figs. 90-91, 176, Fig 92, 200, 220 Fig. 113, 222 Fig. 114, 228 Fig. 118, 232 Fig. 120, 233, 236, 250, 257, Il. Heteractis macrodactylum (Haddon and Shackleton, 1893): Heteractis crispa (Ehrenberg) [pro parte], Dunn, 1981, p. 48: Radianthus malu (Haddon and Shackleton): Allen, 1972, pp. 152 Fig. 75, 166, 168, 169, 170 Figs. 85-86, 171 Fig. 87, 172 Fig. 89, 200, 209, 225 Fig. 115, 228, Fig. 117, 229 Fig. 119, 233 Fig. 121, 254, 255, Il. Heteractis gelam (Haddon and Shackleton, 1893): Macrodactyla doreensis (Quoy and Gaimard) [pro parte], Dunn (1981), p. 29: Radianthus gelam (Haddon and Shackleton): Allen, 1972, pp. 122 Fig. 56, 166, 168, 177, 186 Fig. 98, 187 Fig. 99, 200, 252, I. Family ISOPHELLIIDAE Telmatactis decora (Ehrenberg, 1834). Telmatactis vermiformis (Haddon, 1898). Family HORMATHIIDAE Calliactis polypus (Forsskal, 1795): Josephson, 1966, p. 305. Family SAGARTIIDAE Verrillactis paguri (Verrill, 1869). Family AIPTASIIDAE Aiptasia pulchella Carlgren, 1943. Order CERIANTHARIA Family ARACHNANTHIDAE Isarachnanthus bandanensis Carlgren, 1924. BIOGEOGRAPHY Discussion of the geographic ranges of tropical Pacific shallow-water sea anemones would, at this time, be mean- ingless. In the many papers on these anemones, species are often improperly or inadequately identified and improp- erly or inadequately synonymized. Two corallimorpharians, 24 actiniarians, and one ceri- antharian are presently known from Enewetak. Of these, nine have an Indo-Pacific range, and 18 are known only from the tropical Pacific. Cutress has also collected at Majuro, Marshall Islands; French Polynesia; and Hawaii. At Majuro (1 month), 19 actiniarians, one corallimorpharian, KEY TO SPECIES Corallimorpharia Polyps often gregarious, small to medium sized. Column short, sometimes rudimentary. and one ceriantharian were collected. All species were identical to those from Enewetak. In French Polynesia (3 months), 14 actiniarians, two corallimorpharians, and one ceriantharian were collected. All species were identical to those from Enewetak. In Hawaii (6 years), 25 actiniari- ans, one corallimorpharian, and one ceriantharian were col- lected. Of these, only six species were identical to those from Enewetak. With more collecting at Enewetak, additional species of sea anemones will certainly be found. This speculation is based on 10 species that are known from the Gilbert Islands or Kwajalein that have not yet been found at Enewetak. smooth. Tentacles very numerous, arranged in radial rows, 1. Discal tentacles dendritic, marginals simple. Crown diameter to 6 cm. General color ochre, brown, or greenish brown (Fig. 4c). Rhodactis howesii Saville-Kent SEA ANEMONES 55 All tentacles alike: simple, wart-like, or rudimentary. Crown diameter to 4 cm. General color greenish brown to brown, area about mouth iridescent blue, oral disc often with white radial streaks (Fig. 4a). Actinodiscus neglectus (Fowler) Actiniaria Polyps flaccid or firm, with or without adhesive pedal disc, small to very large, short and broad to long and vermiform. Column clean or covered with cuticle, smooth, with warts of one kind or another, or with inflatable outgrowths. Tentacles cyclically or radially arranged, very long to very short, all alike or discals and marginals dissimilar; simple with swellings or branched; pointed, blunt, or capitate. 10. LE . Tentacles branched, borne on long arm-like outgrowths of the oral disc. 2 Tentacles borne on the oral disc proper. 4 . Tentacles bilaterally arranged on sides of arms, tips conical. Crown diameter 30 cm. General color brown, oral disc with small white spots at base of arms (Fig. 2a). Megalactis hemprichii Ehrenberg Tentacles arranged more or less spirally all around arms. 3 . Tentacles about as broad as long, conical. Crown diameter and column length to 25 cm. General color ochre to brown, proximal column with lilac to brownish spots, oral disc with radial rows of white spots (Fig. 3d). Actinodendron plumosum Haddon Most of tentacles broader than long, blunt or capitate. Crown diameter and column length 20 cm. General color gray, no markings (Fig. 3f). Actinodendron glomeratum Haddon . Oral disc bearing branched tentacles and nematospheres. 5 Oral disc bearing unbranched tentacles only. 6 . Tentacles very numerous, short. Nematospheres simple, club-shaped, occurring as a distinct, broad, peripheral band. Crown diameter to 36 cm. Color of tentacles and nematospheres variable, char- treuse, pink, gray, or brown (all may be present in one individual) (Fig. 2c). Cryptodendrum adhesivum Klunzinger Tentacles numerous, short. Nematospheres globular, appearing like bunches of grapes among the peripheral tentacles. Crown diameter to 20 cm. Tentacles green to greenish brown. Nematospheres frequently violet (not illustrated). Heterodactyla hemprichii Ehrenberg . Tentacles radially arranged (more than one communicating with a single endocoel). 7 Tentacles cyclically arranged (never more than one tentacle communicating with a single endocoel). 12 . Tentacles of two distinct kinds, wart-like discals and normal-length marginals with prominent lateral projections (not illustrated). Phymanthus strandesi Carlgren Tentacles not of two distinct kinds. 8 . Tentacles very short, blunt, covering the oral disc from near mouth to margin. Crown diameter 50 cm (few 1 m). Oral disc and tentacles ochre to brown, sometimes tinged with pink (Fig. 2e). Stichodactyla mertensii Brandt Tentacles normal length to very long. 9 . Tentacles with adoro-lateral protuberances, usually only a few extra ones per endocoel. Crown diame- ter to 25 cm. Tentacles brownish violet, protuberances white. Column with patches of bright orange or yellow (Fig 2f). Heteractis aurora (Quoy and Gaimard) Tentacles smooth. 10 Column smooth or with few inconspicuous warts in upper part. 11 Column with numerous conspicuous warts. Tentacles, to 7.5 cm, pointed, flaccid, pale violet-gray, tipped with chartreuse (Fig. 2d). Heteractis macrodactylum (Haddon and Shackleton) Tentacles, to 7 cm, blunt, many extra ones over older endocoels, greenish gray to greenish brown, white tipped (irregular branching due to Chaetodon browsing). Crown diameter to 25 cm. Column medium brown (Fig. 2b). Heteractis ritteri (Kwietniewski) Tentacles, to 5 cm, blunt, sometimes with subterminal swellings, usually only a few extra ones over older endocoels, greenish brown, tips white or magenta. Crown diameter to 25 cm. Column length to 30 cm, thin-walled, orange becoming pale violet gray near margin (Fig. 3c). | Heteractis gelam (Haddon and Shackleton) 56 CUTRESS AND ARNESON 12. Column with inflatable, simple, or compound vesicles. 13 Column smooth or with solid warts. 14 13. Vesicles simple, of different sizes, scattered over proximal half of column. Tentacles deciduous. Column diameter and column length 1.5 cm. General color pale greenish brown, column often with alternate streaks of white, yellow, and violet (Fig. 4d). Bunodeopsis medusoides (Fowler) Vesicles branched and compound, forming a ring around mid-column. Tentacles not deciduous. Crown diameter and column length 1.5 cm. General color dark brown (chartreuse when carried by Lybia tesselata) (Fig. 4f). Triactis producta Klunzinger 14. Column smooth. Margin without vesicles. 15 Column with or without warts. Margin with vesicles. 16 15. Tentacles to 4 cm, usually with subterminal inflations. Crown diameter to 8 cm. Column off white to pale greenish brown to pink to violet. Tentacles greenish brown, tips often white (not illustrated). Physobrachia douglasi Saville-Kent Tentacles to 4 cm, evenly tapered to point. Mid-column with cinclides. Crown diameter and column length to 7 cm. General color pale to dark olive brown (Fig. le). Aiptasia pulchella Carlgren 16. Marginal vesicles prominent. Column without warts. Crown diameter and column length 1.5 cm. Gen- eral color reddish brown, larger vesicles white (Fig. 3b). Anthostella badia Carlgren Column with rows of prominent adhesive warts. 17 17. General color violet black, warts often white. Crown diameter to 2.5 cm (Fig. 3a). Anthopleura nigrescens (Verrill) Tentacles bright green to brownish green. Column pale pink. Crown diameter to 2.5 cm (Fig. 3e). Actinogeton sesere (Haddon and Shackleton) 18. Column elongate to vermiform, with prominent cuticle. 19 Column short, cuticle inconspicuous. 22 19. Tentacles number 48 or more. 20 Tentacles fewer than 48. PAA 20. Tentacles pointed. Column with thin sand-covered cuticle, length to 4 cm, diameter to 1.5 cm (Fig. 1c). Telmatactis vermiformis (Haddon) Tentacles blunt to capitate. Column with thick, shaggy cuticle, length and diameter to 8.0 and 2.5 cm (Fig. la). Telmatactis decora (Ehrenberg) 21. Column length and diameter 25 and 2.5 cm (Fig. 4e). Edwardsia pudica Klunzinger Column length and diameter to 4.0 and 0.4 cm (Fig. 4b). Edwardsia gilbertensis Carlgren 22. Column firm, lower part with one or more rings of prominent pores through which salmon-colored threads are frequently extruded. Color of column gray to rusty orange, often with streaks and blotches of dull plum red. Crown diameter to 5 cm. Lives with hermit crabs (Fig. 1d). Calliactis polypus (Forsskal) Column lacking pores, white to gray, often with fine streaks of rusty orange. often thicker than rest. Crown diameter to 2 cm. Lives with hermit crabs (Fig. 1b). Verrillactis paguri (Verrill) Ceriantharia Solitary, elongate, tube-dwelling polyps, with tentacles divisible into short labials and long marginals. Only species recorded from Enewetak: Tentacles numbering to 56, more commonly 48 or 52. Column length to 10 cm, crown diameter to 6 cm. General color reddish brown (copper) (Fig. 1f). Isarachnanthus bandanensis Carlgren COLLECTION RECORDS Actinodendron glomeratum Haddon Actinodiscus neglectus (Fowler) Enewetak: 60 feet, Sept. 1980. Lojwa: Sept. 1980. Actinodendron plumosum Haddon Actinogeton sesere (Haddon and Shackleton) Chop Top: Sept. 23, 1980. Ananij: May 8, 1955. SEA ANEMONES 57 Aiptasia pulchella Carlgren Heterodactyla hemprichii Ehrenberg Medren: May 15, 1955. No collection data now at hand but found ; y in 1955 and 1980. Anthopleura nigrescens (Verrill) Igurin: May 11, 1955. Isarachnanthus bandenensis Carlgren Medren: May 15, 1955; Anthostella badia Carlgren ; t t. 1980 AnanijaMay 11955: Bokoluo: 45 feet, Sept. ; Pain eticopuisnmediscidesi(Eowler) Megalactis hemprichii Ehrenberg a io: Sept. 1980. Ananij, Enewetak: June 21, 1955; Enewclals Masala. Sep S Chop Top: lagoon, 40 feet, Phymanthus strandesi Carlgren Sept. 22, 1980. Ananij: June 15, 1955. Calliactis polypus (Forsskal) Physobrachia douglasi Saville-Kent Medren: May 18, 1955. Ananij: June 16, 1955; Cryptodendrum adhesivum Klunzinger Benes Sap 8 UL Medren: June 25, 1955. Rhodactis howesii Saville-Kent Edwardsia gilbertensis Carlgren ESHER AEN) Bay VES Bokoluo: 45 feet, Sept. 1980. Stichodactyla mertensii Brandt Edwardsia pudica Klunzinger MENG BLAMAENS Sale Mele Medren, Runit: May 18, 1955; Telmatactis decora (Ehrenberg) Medren: 50 feet, Sept. 1980. Ananij: May 8, 1955; : : Enewetak: 60 feet, October 17, 1980. Heteractis aurora (Quoy and Gaimard) Bokoluo: Sept. 1980. Telmatactis vermiformis (Haddon) Heteractis gelam (Haddon and Shackleton) Meshers HEY Wey Ieee Medren: May 22, 1955. Triactis producta Klunzinger Heteractis macrodactylum (Haddon and Shackleton) a Japtan: Cement ship, 30 feet, Sept. 1980. Verrillactis paguri (Verrill) Heteractis ritteri (Kwietniewski) sehen MEY) TS, ee Medren: Sept. 1980. Plates 1-4 Fig. la, Telmatactis decora (Ehrenberg), live specimen from Hawaii; b, Verrillactis paguri (Verrill), live specimen from Hawaii; c, Telmatactis vermiformis (Haddon), preserved specimen from Majuro; d, Calliactis polypus (Forsskal), live specimen from Hawaii; e, Aiptasia pulchella Carlgren, live specimen from Hawaii; f, Isarachnanthus bandanensis Carlgren, live specimen from Hawaii. ed Page 58 Fig. 2a, Megalactis hemprichii Ehrenberg, live specimen in situ, Enewetak, 40 feet; b, Heteractis ritteri (Kwietniewski), live speci- men in situ, Medren, Enewetak, patch reef, 10 feet; c, Cryptodendrum adhesivum Klunzinger, preserved specimen from Enewetak; d, Heteractis macrodactylum (Haddon and Shackleton), live specimen in situ, Cement ship, Enewetak, 30 feet; e, Stichodactyla mertensii Brandt, live specimen in situ, Henry, outside reef Enewetak, 30 feet; f, Heteractis aurora (Quoy and Gaimard), live speci- men in situ, Enewetak. a Page 59 Fig. 3a, Anthopleura nigrescens (Verrill), preserved specimen from Hawaii; b, Anthostella badia Carlgren, preserved specimen from Majuro; c, Heteractis gelam (Haddon and Shackleton), live specimen in situ, with Amphriprion tinctus, Bokoluo, Enewetak patch reef, 40 feet; d, Actinodendron plumosa Haddon, live specimen in situ, Chop Top, Enewetak, 30 feet; e, Actinogeton sesere (Had- don and Shackleton), live specimen from Hawaii; f, Actinodendron glomeratum Haddon, live specimen in situ, off Enewetak, 60 feet. aa Page 60 Fig. 4a, Actinodiscus neglectus (Fowler), live specimen in situ, Lajwa, Enewetak, 20 feet; b, Edwardsia gilbertensis Carlgren, preserved specimen from Majuro; c, Rhodactis howesii Saville-Kent, preserved specimen from Enewetak; d, Bunodeopsis medusoides (Fowler), live specimen from Hawaii; e, Edwardsia pudica Klunzinger, live specimen in situ, Runit, Enewetak, 30 feet; _ f, Triactis producta Klunzinger, live specimen from Hawaii. ee Page 61 58 CUTRESS AND ARNESON 59 SEA ANEMONES Fig. 2. 60 CUTRESS AND ARNESON SEA ANEMONES 61 62 CUTRESS AND ARNESON REFERENCES Allen, Gerald R., 1972, The Anemonefishes: Their Classification and Biology, Neptune City, N. J., T. F. H. Publ. Inc. Ltd., 288 pp. Dunn, Daphne Fautin, 1981, The Clownfish Sea Anemones: Stichodactylidae (Coelenterata: Actiniaria) and Other Sea Anemones Symbiotic with Pomacentrid Fishes, Trans. Am. Philos. Soc., 71, Pt. 1: 1-115. Johannes, R. E., et al., 1972, The Metabolism of Some Coral Reef Communities: A Team Study of Nutrient and Energy Flux at Eniwetok, Bioscience, 22 (9): 541-543. Josephson, Robert K., 1966, Neuromuscular Transmission in a Sea Anemone, J. Exp. Biol., 45 (2): 305-319. Octocorallia of Enewetak Atoll [Editor’s Note: The section on the Octocorallia was compiled by Dennis M. Devaney (B. P. Bishop Museum, Honolulu, Hawaii), but the collection data listing was not completed due to his untimely death. This unfinished section, with complete references, is included in order to round out the categories of organisms recorded from Enewetak.—B. L. Burch] STOLONIFERA Two widely distributed Indo-West Pacific stoloniferans including the distinctive organ-pipe coral, Tubipora musica, are present at Enewetak. This species occurs most abun- dantly on windward reef flats or just behind the algal ridge and less commonly on lagoon reefs (Wells, 1954). For Pachyclavularia violacea, see below. TELESTACEA One undetermined species of Telesto is represented in the Mid-Pacific Research Laboratory (MPRL) reference col- lection. It was found attached on the reef near the Medren cargo pier. This appears to be the first record of Telesto from the Marshall Islands; members of this genus may occur on ship hulls and be distributed as fouling organisms. ALCYONACEA Alcyonaceans are the most numerous octocorals represented in Enewetak shallow waters with approxi- mately 20 species now known. The majority of these “soft corals” recorded herein are from the lagoon and sub- merged reef habitats. One alcyonacean was reported by Odum and Odum (1956) as one of the organisms encoun- tered during their earlier coral reef studies on the atoll. Enewetak soft corals (Sarcophyton and Heteroxenia) were utilized in work on the nerve net system of cnidarians (Bul- lock and Thorson, 1962; Josephson, 1965). Six shallow- water alcyonaceans and one stoloniferan were reported by Verseveldt (1972) based on collections made in the search for crustacean copepod symbionts in July 1969. The stoloniferan, Pachyclavularia violacea, and three of the alcyonaceans were hosts for several of these crustaceans (Humes, 1973; Devaney, Chapter 19, Table 2, this volume). 63 Chapter 7 Research at MPRL on marine natural products chemis- try (Ciereszko et al., 1968; Schmitz et al., 1974; Van- derah et al., 1978) has been based in part on soft corals. In response to this work, eight alcyonaceans were deter- mined from Enewetak by Verseveldt (1977) having been collected mainly through projects by the University of Hawaii’s Department of Chemistry. In this publication, one new species (Sinularia sandensis) was described together with other more widely distributed Indo-West Pacific species all representing new records for the Marshall Islands. This led to a revision of the genus Sinularia in Verseveldt (1980). Furthermore, species identified by Verseveldt for Ciereszko (University of Oklahoma) in 1975, include five that are new Enewetak records (Table 1). Examples were deposited in the MPRL refer- ence collection. Nephthea, Sarcophyton, and Lobophytum were genera reported in studies on the mucus chemistry and zooxanthellae of reef animals (Ciereszko and Ciereszko, 1978). COENOTHECALIA The blue-coral, Heliopora coerulea, distributed throughout much of the Indo-Pacific, was found at Enewetak and other Marshall Islands where colonies up to many meters in area were reported on the seaward reef flats (Wells, 1954). GORGONACEA The gorgonaceans (sea fans and sea whips) are represented by at least eight species in the suborders Scleraxonia and Holaxonia at depths from 5 to 45 m at Enewetak. These are known by collections made together with black corals to obtain information on their diversity and to measure the concentration and distribution of skele- tal radioiodine (Goldberg, 1975). He also included informa- tion on colony form, color, habitat, and collection localities. Determination of species was made by F. M. Bayer and updated in 1982. Representatives of all but one of these taxa are in the United States National Museum (Smith- sonian Institution). In another study of gorgonaceans from the Marshall Islands, Bayer (1949) recorded several species from Bikini Atoll. All but two of the Bikini specimens were 64 DEVANEY TABLE 1 Checklist of Octocorallia of Enewetak Atoll Phylum COELENTERATA | Family ALCYONIIDAE (Cont'd) Class ANTHOZOA Sinularia polydactyla (Ehrenberg, 1834): Subclass OCTOCORALLIA Verseveldt, 1972; Humes, 1973. Order STOLONIFERA Sinularia rigida (Dana, 1846): Verseveldt, 1977. Family CLAVULARIIDAE Sinularia sandensis (Verseveldt, 1977): Pachyclavularia violacea (Quoy and Gaimard, Verseveldt, 1977. 1833): Verseveldt, 1972: Humes, 1973. Family NEPHTHEIDAE Family TUBIPORIDAE Dendronephthya (Roxasia) mirabilis Henderson, Tubipora musica Linnaeus, 1758: Wells, 1954. 1909: Verseveldt, 1977. Order TELESTACEA Nephthea chabrolii Audouin, 1828: Verseveldt, Family TELESTIDAE 1972; Humes, 1973. * Telesto sp. * Nephthea pacifica Kukenthal, : Order ALCYONACEA * Nephthea albida (Holm, 1894). Family ALCYONIIDAE * Stereonephthya unicolor (Gray, 1862). * Cladiella pachyclados (Klunzinger, 1877). Family XENIIDAE Lobophytum borbonicum (Von Marenzeller, 1886): * Heteroxenia coheni Verseveldt, 1974. Verseveldt, 1977. Order COENOTHECALIA Lobophytum catalai Tixier-Durivault, 1957: Family HELIOPORIDAE Verseveldt, 1977. Heliopora coerulea (Pallas, 1766): Wells, 1954. Lobophytum denticulatum Tixier-Durivault, 1956: Order GORGONACEA Verseveldt, 1977. Suborder SCLERAXONIA Lobophytum pauciflorum (Ehrenberg, 1834): Family SUBERGORGIIDAE Odum and Odum, 1956; Verseveldt, 1972. * Subergorgia nuttingi Stiasny, 1937. Humes, 1973; Verseveldt, 1977. * Subergorgia reticulata (Ellis and Solander, 1786). Sarcophyton glaucum (Quoy and Gaimard, 1833): * Subergorgia suberosa (Pallas, 1766). Verseveldt, 1977. Suborder HOLAXONIA Sarcophyton trocheliophorum Von Marenzeller, Family ACANTHOGORGIIDAE 1886: Verseveldt, 1972; 1977. * Acanthogorgia sp. Sinularia densa (Whitelegge, 1897): Verseveldt, Family PARAMURICEIDAE 1980. Sinularia brongersmai Verseveldt, 1972: * Astrogorgia sp. Verseveldt, 1972. * Astrogorgia? sp. Sinularia leptoclados (Ehrenberg, 1834): Family PLEXAURIDAE Verseveldt, 1977. * Rumphella antipathes (Linnaeus, 1758). Sinularia marenzelleri (Wright and Studer, 1889): Family ELLISELLIDAE Verseveldt, 1972. * Ellisella sp. Le "New Enewetak record. dredged from depths of 100 m or deeper. One species, recorded as Subergorgia mollis, is very close taxonomi- cally to that determined as S. reticulata from Enewetak, but further work is necessary before this question is resolved (Bayer, personal communication). Otherwise, none of the species so far identified is common to the two atolls. At least one more shallow-water gorgonacean, as yet unrecorded from Enewetak, was described from Arno Atoll in the southern Marshall Islands (Bayer, 1955). The gorgonacean Rumphella antipathes has been utilized from specimens collected at Enewetak for biochemical studies (Ciereszko et al., 1968; Marsh and Ciereszko, 1973). During submersible dives off the seaward side of three islets (Biken, Mut, and Enewetak) during July 1981, the author observed nephtheids and gorgonaceans as common faunal elements at depths from approximately 80 to 180 m. Several of these, as yet undetermined, were col- lected and were deposited at the Bishop Museum. The classification of octocorals used herein follows that by Bayer (1956), although a revised classification has recently been proposed (Bayer, 1981). ACKNOWLEDGMENTS J. Verseveldt was most helpful in reviewing the discus- sion and checklist on the alcyonaceans and allowing the inclusion of the five new records that he determined. F. M. Bayer (USNM) provided updated names for the material collected by Goldberg and was most helpful in permitting their inclusion here. REFERENCES Bayer, F. M., 1949, The Alcyonaria of Bikini and Other Atolls in the Marshall Group, Part I: the Gorgonacea, Pac. Sci., 3: 195-214. __, 1955, Contributions to the Nomenclature, Systematics, and OCTOCORALLIA 65 Morphology of the Octocorallia, Proc. U. S. Nat. Mus., 105: 207-212. —, 1956, Octocorallia, Treatise on Invertebrate Paleontology, Part F Coelenterata, R. C. Moore (Ed.), University of Kansas Press, Lawrence, Kansas, p. F166-F190. __., 1981, Key to the Genera of Octocorallia Exclusive of Pen- natulacea (Coelenterata: Anthozoa) with Diagnoses of New - Taxa, Proc. Biol. Soc. Wash., 94: 902-947. Bullock, T. H., and J. Thorson, 1962, MPRL Ann. Rept. FY 1972, p. 2. Ciereszko, L. S., and E. M. Ciereszko, 1978, MPRL Ann. Rept. FY 1978, p. 15. __, M. A. Johnson, R. W. Schmidt, and C. B. Koons, 1968, Chemistry of Coelenterates, VI. Occurrence of Gorgosterol, AC30 Sterol, in Coelenterates and Their Zooxanthellae, Comp. Biochem. Physiol., 24: 899-904. Goldberg, W. M., 1975, MPRL Ann. Rept. FY 1975, p. 55. Humes, A. J., 1973, Cyclopoid Copepods (Lichomolgidae) from Octocorals at Eniwetok Atoll, Beaufortia, 21: 135-151. Josephson, R. K., 1965, MPRL Ann. Rept. FY 1964-65, p. 6. Marsh, M. E., and L. S. Ciereszko, 1973, Alpha-Glyceryl Ethers in Coelenterates, Proc. Oklahoma Acad. Sci., 53: 53-56. Odum, E. P., and H. T. Odum, 1956, Zonation of Corals on Jap- tan Reef, Eniwetok Atoll, Atoll Res. Bull. No. 52, pp. 1-3. Schmitz, F. J., D. J. Vanderah, and L. S. Ciereszko, 1974, Marine Natural Products: Nephenol and Epoxynephenol Ace- tate, Cembrene Derivatives from a Soft Coral, Chem. Com- mun., 1974: 407-408. Vanderah, D. J., N. Rutledge, F. J. Schmitz, and L. S. Ciereszko, 1978, Marine Natural Products: Cembrene-A and Cembrene-C from a Soft Coral, Nephthea sp., J. Org. Chem., 43: 1614-1616. Verseveldt, J., 1972, Report on a Few Octocorals from Eniwetok Atoll, Marshall Islands, Zool. Meded., 47: 457-465. —_, 1977, Octocorallia from Various Localities in the Pacific Ocean, Zool. Verh. (Leiden) No. 150, pp. 1-42. , 1980, A Revision of the Genus Sinularia May (Octocorallia, Alcyonacea), Zool. Verh. (Leiden) No. 179, pp. 1-128. Wells, J., 1954, Recent Corals of the Marshall Islands, U. S Geol. Survey Prof. Pap. 260-I, pp. 385-486. Chapter 8 Scleractinia (Stony Corals) of Enewetak Atoll DENNIS M. DEVANEY’ (deceased) and JUDITH C. LANG+ * B. P. Bishop Museum, Honolulu, Hawaii 96817 + University of Texas at Austin, Department of Zoology, Austin, Texas 78712, and Texas Memorial Museum, Austin, Texas 78705 The first record of stony corals from Enewetak Atoll, in which five species were reported, appears to have been that of Yabe et al. (1936). Wells (1954) later made a major systematic study of the corals of the Marshall Islands. Of 254 species and varieties in 57 genera recorded overall from this region, only 44 scleractinian species were listed as occurring at Enewetak. This disparity is understandable because much of the collecting effort was concentrated at Bikini Atoll. Most of these specimens are now deposited at the U. S. National Museum of Natural History (USNM). When the Mid-Pacific Research Laboratory (MPRL) was established in 1954, one of its first goals was to have a labeled set of stony corals available for use by investigators at the laboratory. Specimens collected by R. Iverson at Enewetak in 1955 were partitioned among 115 sclerac- tinian species by J. W. Wells and formed the nucleus of the MPRL reference collection. A list of the corals col- lected during the classical field study by Odum and Odum (1955) at Enewetak, and subsequently identified by Dr. Wells, was published by Odum and Odum in 1956. In the summer of 1970, R. H. Randall collected 357 scleractinian specimens for an investigation of growth form and variation (Randall, 1971). As a result of this effort, 21 specimens were added to the Enewetak reference collec- tion (the balance remains at the University of Guam Marine Laboratory). Weber and Haggerty (1976) collected stony corals of 48 genera and subgenera for stable isotopic stud- ies of skeletal calcification. By then the coral reference col- lection at Enewetak contained representatives of about 124 species and varieties. A more intensive effort was made to identify Enewetak scleractinians during the 1976 reef coral workshop. An international team of coral systematists from the United States, Australia, England, France, and the Netherlands gathered under the direction of J. C. Lang to (a) update 67 the reference collection with a set of labeled coral speci- mens representative of each major reef habitat at Enewetak and (b) prepare a reef coral field identification guide for general use by scientists working at Enewetak. It was also hoped that the specimens and accompanying field observations would be used in individual systematic publi- cations of the Enewetak scleractinians. To date, approximately 170 species in 54 genera have been identified among the 2500 stony coral specimens which were collected from 28 sites around Enewetak Atoll. These have been deposited in major museums (B. P. Bishop Museum, United States National Museum, British Museum [Natural History], Rijksmuseum van Natuurlijke Historie, Queensland Museum), as well as in the Enewetak reference collection. Although the field guide is still in preparation, a provi- sional checklist of scleractinian corals now believed to occur at Enewetak is presented in Table 1. In many fami- lies, the species-level classification generally follows that given in the recent publications about eastern Australian scleractinians by Veron and Pichon (1976, 1980, 1982), by Veron, Pichon, and Wijsman-Best (1977), and by Veron and Wallace (1984). Ecological and zoogeographic con- siderations will await the completed guide. Stony corals collected at Enewetak during the workshop have contributed to several revisionary sys- tematic studies. For example, range extensions were made by Wallace (1978) for species of Acropora; by Dinesen (1980) for a species of Leptoseris; and by Wijsman-Best (1980) for species in three faviid genera (Cyphastrea, Lep- tastrea, Echinopora). Lamberts (1980, 1982) has also used Enewetak specimens to revise the genus Astreopora. Six of the nine species of Astreopora (two of which are new species) recognized by Lamberts (1982) have been col- lected at Enewetak, while two others occur elsewhere in the Marshall Islands. In addition to these systematic studies, there are numerous publications on scleractinian biology or geology which have resulted from field work and collections made at Enewetak. A selection of these references, with the gen- era examined, is presented in Table 2. The authors of these nonsystematic papers are also cited in Table 1 under any species names given for these genera. 68 DEVANEY AND LANG TABLE 1 Checklist of Stony Corals of Enewetak Atoll Order SCLERACTINIA Suborder ASTROCOENIINA Family ASTROCOENIIDAE Stylocoeniella armata (Ehrenberg, 1834): Hildemann et al., 1975a. “+ Stylocoeniella guentheri (Bassett-Smith, 1890). Family THAMNASTERIIDAE “+ Psammocora contigua (Esper, 1795). Psammocora digitata Milne Edwards and Haime, 1851. Psammocora (Stephanaria) togianensis Umbgrove, 1940: Garth, 1964; Knutson et al., 1972; Buddemeier et al., 1974. Synonymy after Veron and Pichon (1976). *Psammocora explanulata van der Horst, 1922. *Psammocora nierstrazi van der Horst, 1921. *t Psammocora profundacella Gardiner, 1898. Family POCILLOPORIDAE Pocillopora damicornis (Linnaeus, 1758): Garth, 1964; Johannes et al., 1972a; Clausen and Roth, 1975; Coles and Jokiel, 1977; Stimson, 1978; Richmond, 1982; Roth et al., 1982; Richmond and Jokiel, 1984. Pocillopora setchelli Hoffmeister, 1929: Wells, 1954. Synonymy via J. Veron (personal communication). Pocillopora eydouxi Milne Edwards and Haime, 1860: Garth, 1964; Pomeroy and Kuenzler, 1969; Buddemeier et al., 1974; Hildemann et al., 1975a, 1975b. Pocillopora grandis Dana, 1846: Hildemann et al., 1975a, 1975b. Synonymy after Veron and Pichon (1976). Pocillopora meandrina Dana, 1846: Banner and Banner, 1968; Muscatine and D’Elia, 1978. Pocillopora elegans Dana, 1846: Wells, 1954; Odum and Odum, 1956; Garth, 1964; Hildemann et al., 1975a, 1975b; Coles et al., 1976; D’Elia and Webb, 1977; Stimson, 1978. Synonymy via J. Veron (personal communication). Pocillopora verrucosa (Ellis and Solander, 1786): Wells, 1954; Odum and Odum, 1956; Garth, 1964; Pilson, 1974; Hildemann et al., 1975a, 1975b; Stimson, 1978. Pocillopora danae Verrill, 1864: Odum and Odum, 1956; Knudsen, 1967; D’Elia, 1977. Synonymy via J. Veron (personal communication). Seriatopora hystrix Dana, 1846: Wells, 1954; Garth, 1964; Young, 1971; Hildemann et al., 1975a; Stimson, 1978. Seriatopora angulata Klunzinger, 1879: Wells, 1954. Synonymy after Veron and Pichon (1976). Stylophora pistillata (Esper, 1795). Stylophora mordax (Dana, 1846): Yabe et al., 1936; Wells, 1954; Odum and Odum, 1956. Synonymy after Veron and Pichon (1976). Family ACROPORIDAE *+ Acropora aculeus (Dana, 1846). Acropora acuminata (Verrill, 1864): Wells, 1954; Garth, 1964 (as “A. formosa or A. acuminata”); Hildemann et al., 1975a, 1975b. Acropora aspera (Dana, 1846): Stimson, 1978. Acropora hebes (Dana, 1846): Wells, 1954. Synonymy after Wallace (1978). + Acropora austera (Dana, 1846): Wallace, 1978. *+ Acropora cerealis (Dana, 1846). * Acropora cuneata (Dana, 1846). Acropora cythera (Dana, 1846): Wallace, 1978. Acropora reticulata (Brook, 1892): Vosburgh, 1977, 1978, 1982. Synonymy after Wallace (1978). Acropora danai (Milne Edwards and Haime, 1860). Acropora rotumana (Gardiner, 1898): Wells, 1954; Hildemann et al., 1975a, 1975b. Synonymy after Veron and Wallace (1984). Acropora digitifera (Dana, 1846): Stimson, 1978. Acropora echinata (Dana, 1846): Wells, 1954; Garth, 1964; Hildemann et al., 1975a. t Acropora florida (Dana, 1846): Humes and Stock, 1973; Hildemann et al., 1975a; Humes, 1981; Humes and Dojiri, 1982. Acropora formosa (Dana, 1846): Wells, 1954; Garth, 1964 (as “A. formosa or A. acuminata”); Cheney, 1975; Hildemann et al., 1975a, 1975b; Coles et al., 1976; Roth et al., 1982. Acropora gemmifera Brook, 1892: Odum and Odum, 1955; in part as Acropora humilis (Dana, 1846) in Wells, 1954; ? in part in: Odum and Odum, 1956; Garth, 1964; Hildemann et al., 1975a, 1975b; Stimson, 1978. Revised by Veron and Wallace (1984). Acropora granulosa (Milne Edwards and Haime, 1860). Acropora rayneri (Brook, 1892): Wells, 1954. Synonymy after Wallace (1978). *New Enewetak record. +Not recorded from Marshall Islands by this name in Wells (1954). (This table continued on next page.) SCLERACTINIA (STONY CORALS) 69 TABLE 1 (cont’d) Family ACROPORIDAE (cont’d) tAcropora horrida (Dana, 1846). Acropora angulata (Quelch, 1886): Hildemann et al., 1975a. Synonymy after Wallace (1978). Acropora humilis (Dana, 1846): in part in Wells, 1954; ? in part in: Odum and Odum, 1956; Garth, 1964; Hildemann et al., 1975a, 1975b; Stimson, 1978. Revised by Veron and Wallace (1984). Acropora hyacinthus (Dana, 1846): Wells, 1954; Garth, 1964; Pomeroy and Kuenzler, 1969; Bruce, 1969; Coles et al., 1976; Smith and Harrison, 1977; D’Elia, 1977; Stimson, 1978. Acropora corymbosa (Lamarck, 1816): in part in Wells, 1954; ? in part in: Odum and Odum, 1956; Garth, 1964; Stimson, 1978. Synonymy after Veron and Wallace (1984). Acropora recumbens (Brooks, 1892): Odum and Odum, 1956. Synonymy after Wallace (1978). Acropora? surculosa (Dana, 1846): Garth, 1964; Hildemann et al., 1975a. Synonymy after Veron and Wallace (1984). Acropora loripes (Brook, 1892): Young, 1971; Hildemann et al., 1975b. Acropora microphthalma (Verrill, 1869): Wallace, 1978. Acropora millepora (Ehrenberg, 1834). Acropora corymbosa (Lamarck, 1816): in part in Wells, 1954; ? in part in: Odum and Odum, 1956; Garth, 1964; Stimson, 1978. Synonymy via C. Wallace (personal communication). Acropora nasuta (Dana, 1846): Wells, 1954; Hildemann et al., 1975a, 1975b; Stimson, 1978. Acropora cymbicyathus (Brook, 1893): Wells, 1954; Odum and Odum, 1955; Garth, 1964. Synonymy after Wallace (1978). Acropora palifera (Lamarck, 1816): Wells, 1954. Acropora palmerae Wells, 1954: Odum and Odum, 1955; Bruce, 1969; Coles et al., 1976. Acropora paniculata Verrill, 1902: Garth, 1964. “t+ Acropora robusta (Dana, 1846). Acropora samoensis (Brook, 1891): in part as Acropora humulis (Dana, 1846) in Wells, 1954; ? in part in: Odum and Odum, 1956; Hildemann et al., 1975a, 1975b; Stimson, 1978. Revised by Veron and Wallace (1984). Acropora secale (Studer, 1878). Acropora diversa (Brook, 1891): Wells, 1954. Synonymy after Veron and Wallace (1984). “t+ Acropora “sp. called A. polymorpha (Brook, 1891)”. Acropora sp.? Acropora conferta (Quelch, 1886): Wells, 1954; Odum and Odum, 1956; Goreau, 1959; Hildemann et al., 1975a. Equals A. hyacinthus (Dana, 1846) or A. millepora (Ehrenberg, 1834) (C. Wallace, personal communication). Acropora delicatula (Brook, 1891): Coles et al., 1976. Acropora grandis (Brook, 1892): Hildemann et al., 1975a Probably not this species (C. Wallace, personal communication). Acropora syringoides (Brook, 1892): Stimson, 1978. Acropora striata (Verrill, 1866): Wells, 1954; Stimson, 1978. “Acropora tenella (Brook, 1892). * Acropora tenuis (Dana, 1846). Acropora teres (Verrill, 1866): Wells, 1954. Acropora tubicinaria (Dana, 1846): Odum and Odum, 1956. Acropora valida (Dana, 1846). ‘Acropora variabilis (Klunzinger, 1879): Garth, 1964; Hildemann et al., 1975a. Synonymy after Veron and Wallace (1984). *Acropora vaughani Wells, 1954. + Astreopora cucullata Lamberts, 1980: Lamberts, 1982. Astreopora gracilis Bernard, 1896: Lamberts, 1982. Astreopora listeri Bernard, 1896: Lamberts, 1982. Astreopora myriophthalma (Lamarck, 1816): Wells, 1954; Odum and Odum, 1956; Buddemeier et al., 1974; Highsmith, 1980b, 1981b; Lamberts, 1982. +Astreopora scabra Lamberts, 1982. Astreopora suggesta Wells, 1954: Lamberts, 1982. + Montipora aequituburculata, Bernard, 1897. Montipora composita Crossland, 1952: Odum and Odum, 1956. Monitpora?verrilli Vaughan, 1907: Odum and Odum, 1956; Highsmith, 1980b. Montipora?verrili [sic] Vaughan: Hildemann et al., 1975a. *+ Montipora australiensis Bernard, 1897. + Montipora berryi Bernard, 1897: Highsmith, 1980b, 1981a. * Montipora caliculata (Dana, 1846). “New Enewetak record. tNot recorded from Marshall Islands by this name in Wells (1954). (This table continued on next page.) 70 DEVANEY AND LANG TABLE 1 (cont’d) Family ACROPORIDAE (cont’d) * Montipora conicula Wells, 1954. * Montipora danae Milne Edwards and Haime, 1851. "+ Montipora digitata (Dana, 1846). *Montipora ehrenbergii Verrill, 1875. *Montipora floweri Wells, 1954. *+Montipora foliosa (Pallas, 1766). Montipora foveolata (Dana, 1846): Odum and Odum, 1956. Montipora socialis Bernard, 1897: Wells, 1954; Highsmith, 1980b. Synonymy after Veron and Wallace (1984). “+ Montipora hispida (Dana, 1846). Montipora hoffmeisteri Wells, 1954: Highsmith, 1980b. Montipora informis Bernard, 1897: Hildemann et al., 1975a. *+ Montipora lobulata Bernard, 1897. *Montipora marshallensis Wells, 1954. “+ Montipora sp. cf. M. monasteriata (Forskal, 1775). * Montipora myriophthalma Bernard, 1897. *+Montipora porosa Bernard, 1897. Montipora sp.? Montipora granulosa Bernard, 1897: Highsmith, 1980b. *Montipora tuberculosa (Lamarck, 1816). Montipora venosa (Ehrenberg, 1834): Wells, 1954. Montipora verrucosa (Lamarck, 1816): Hildemann et al., 1975a; Coles and Jokiel, 1977. Suborder FUNGHNA Family AGARICIIDAE * Gardineroseris planulata (Dana, 1846). As Pavona (Polyastra) planulata (Dana, 1846) from Arno in Wells (1954). Synonymy after Veron and Pichon (1980). Leptoseris hawaiiensis Vaughan, 1907: Dinesen, 1980. Leptoseris mycetoseroides Wells, 1954: Dinesen, 1980. *Pachyseris speciosa (Dana, 1846). Pavona cactus (Forskal, 1775). Pavona praetorta (Dana, 1846): Wethey and Porter, 1976a, 1976b. Synonymy after Veron and Pichon (1980). Pavona clavus (Dana, 1846): Highsmith, 1981a, 1981b. * Pavona maldivensis (Gardiner, 1905). As P. (Pseudocolumnastraea) pollicata Wells, 1954 from Bikini and Arno (Wells, 1954). Synonymy after Veron and Pichon (1980). *+Pavona prismatica Briiggemann, 1879. * Pavona varians Verrill, 1864. * Pavona venosa (Ehrenberg, 1834). Family SIDERASTREIDAE *Coscinaraea columna (Dana, 1846). Family FUNGIIDAE *+ Cycloseris erosa (Déderlein, 1901). Cycloseris hexagonalis Milne Edwards and Haime, 1849: Wells, 1954. Fungia (Ctenactis) echinata (Pallas, 1766). Fungia echinata (Pallas, 1766): Young, 1971; Hildemann et al., 1975a, 1975b. *+ Fungia (Danafungia) valida Verrill, 1864. Fungia (Fungia) fungites (Linneaus, 1758). Fungia fungites (Linnaeus, 1758): Pomeroy and Kuenzler, 1967; Buddemeier et al., 1974; Hildemann et al., 1975a, 1975b. *Fungia (Pleuractis) paumotensis Stuchbury, 1833. Fungia (Pleuractis) scutaria Lamarck, 1801. Fungia scutaria Lamarck, 1801: Wells, 1954; Coles et al., 1976; D’Elia, 1977. *Fungia (Verillofungia) concinna Verrill, 1864. *Fungia (Verrillofungia) repanda Dana, 1846. *Halomitra pileus (Linnaeus, 1758). As H. philippinensis Studer 1901 from Jaluit and Arno (Wells, 1954). Synonymy. after Veron and Pichon (1980). *New Enewetak record. +Not recorded from Marshall Islands by this name in Wells (1954). (This table continued on next page.) SCLERACTINIA (STONY CORALS) 71 TABLE 1 (cont'd) Family FUNGIIDAE (cont’d) *+ Herpetoglossa simplex (Gardiner, 1905). Herpolitha limax (Houttuyn, 1772). Herpolitha limax (Esper, 1795) [sic]: Young, 1971; Buddemeier et al., 1974. “t Lithophyllon sp. cf. L. edwardsi (Rousseau, 1854). “+ Podabacia crustacea (Pallas, 1776). Parahalomitra robusta (Quelch, 1886): Buddemeier et al., 1974. “t+ Zoopilus echinatus Dana, 1846. Family PORITIDAE “+ Alveopora verrilliana Dana, 1872. “+ Goniopora tenuidens (Quelch, 1886). Porites australiensis Vaughan, 1918: Wells, 1954. Porites cylindrica Dana, 1846. Porites andrewsi Vaughan, 1918: Garth, 1964; Hildemann et al., 1975a, 1975b. Synonymy after Veron and Pichon (1982). * Porites lichen Dana, 1846. Porites lobata Dana, 1846: Odum and Odum, 1955, 1956; Pomeroy and Kuenzler, 1969; Buddemeier et al., 1974. Porites lopata [sic]: Hildemann et al., 1975a, 1975b. Porites lutea Milne Edwards and Haime, 1851: Wells, 1954; Odum and Odum, 1956; Knutson et al., 1972; Knutson and Buddemeier, 1973; Buddemeier et al., 1974; Coles et al., 1976; Muscatine and D’Elia, 1978; Highsmith, 1979, 1980a, 1980b, 1981a, 1981b, 1981c. *Porites (Synaraea) monticulosa (Dana, 1846). As Porites (Synaraea) iwayamaensis Eguchi, 1938 from Jaluit, and as Porites (Synaraea) monticulosa (Dana, 1846) from Jaluit and Arno, in Wells (1954). Considered to be a junior synonym of Porites (Synaraea) rus [Forskal, 1775) by Veron and Pichon (1982). “+ Porites (Synaraea) rus (Forskal, 1775). Suborder FAVIINA Family FAVIIDAE “t+ Caulastrea furcata Dana, 1846. Cyphastrea chalcidicum (Forskal, 1775): Odum and Odum, 1956; Wijsman-Best, 1980. +Cyphastrea microphthalma (Lamarck, 1816): Bruce, 1979; Wijsman-Best, 1980. + Cyphastrea ocellina (Dana, 1846): Wijsman-Best, 1980. Cyphastrea serailia (Forskal, 1775): Wells, 1954; Odum and Odum, 1955; Wijsman-Best, 1980. Echinopora lamellosa (Esper, 1795): Odum and Odum, 1956; Wijsman-Best, 1980. *Favia amicorum complex. Veron and Pichon (1982) now consider most east Australian specimens of the Favia amicorum complex of Veron et al. (1977) to be Barabattoia amicorum (Milne Edwards & Haime, 1850). *Favia helianthoides Wells, 1954. “+ Favia matthaii Vaughan, 1918. Favia pallida (Dana, 1846): Wells, 1954; Odum and Odum, 1955; Odum and Odum, 1956; Highsmith, 1979, 1980b, 1981a, 1981b, 1981c; Haggerty et al., 1980. Favia speciosa (Dana, 1846): Wells, 1954; Knutson et al., 1972; Knutson and Buddemeier, 1973; Buddemeier et al., 1974. Favia stelligera (Dana, 1846): Wells, 1954; Odum and Odum, 1956; Buddemeier et al., 1974; Haggerty et al., 1980. Favites abdita (Ellis and Solander, 1786): Odum and Odum, 1956; Hildemann et al., 1975a. Favites flexuosa (Dana, 1846): Hildemann et al., 1975a. Favites halicora (Ehrenberg, 1834): Odum and Odum, 1956. “+ Favites rotundata Veron, Pichon and Wijsman-Best, 1977. *Favites russelli (Wells, 1954). As Plesiastrea russelli Wells, 1954 from Bikini (Wells, 1954). Synonymy after Veron et al. (1977). Favites sp.? Favites yamanarii Yabe and Sugiyama, 1935: Hildemann et al., 1975a. Veron et al. (1977) considered this species to be a junior synonym of Favites chinensis (Verrill, 1866). + Goniastrea edwardsi Chevalier, 1971. Goniastrea parvistella (Dana, 1846) sensu: Knutson et al., 1972; Knutson and Buddemeier, 1973; Buddemeier et al., 1974. Synonymy after Veron et al. (1977). *Goniastrea pectinata (Ehrenberg, 1834). Goniastrea retiformis (Lamarck, 1816): Yabe et al., 1936; Wells, 1954; Odum and Odum, 1956; Knutson et al., 1972; Buddemeier et al., 1974; Highsmith, 1979, 1980b, 1981a, 1981b, 1981c. “Hydnophora exesa (Pallas, 1776). Hydnophora microconos (Lamarck, 1816): Yabe et al., 1936; Wells, 1954; Buddemeier et al., 1974. “New Enewetak record. TNot recorded from Marshall Islands by this name in Wells (1954). (This table continued on next page.) 72 DEVANEY AND LANG TABLE 1 (cont'd) Family FAVIIDAE (cont'd) * Hydnophora rigida (Dana, 1846). + Leptastrea pruinosa Crossland, 1952: Wijsman-Best, 1980. Leptastrea purpurea (Dana, 1846): Pomeroy and Kuenzler, 1969; Wijsman-Best, 1980. Leptastrea transversa Klunzinger, 1879: Wijsman-Best, 1980. Leptoria phrygia (Ellis and Solander, 1786): Yabe et al., 1936; Wells, 1954. *+ Montastraea annuligera (Milne Edwards and Haime, 1849). *t+ Montastraea curta (Dana, 1846). Oulophyllia crispa (Lamarck, 1816): Highsmith, 1981a, 1981b. Oulophyllia aspera (Quelch, 1886): Buddemeier et al., 1974. Synonymy after Veron et al. (1977). Platygyra daedalea (Ellis and Solander, 1786). Platygyra rustica (Dana, 1846): Wells, 1954; Odum and Odum, 1956. Synonymy after Veron et al. (1977). t Platygyra lamellina (Ehrenberg, 1834): Buddemeier et al., 1974; Highsmith, 1980b, 1981b. “+ Platygyra pini Chevalier, 1975. *Platygyra sinensis (Milne Edwards and Haime, 1849). Plesiastrea versipora (Lamarck, 1816): Yabe et al., 1936; Wells, 1954; Odum and Odum, 1956. Family RHIZANGIIDAE *Culicia rubeola (Quoy and Gaimard, 1833). Family OCULINIDAE *Acrhelia horrescens (Dana, 1846). Family MERULINIDAE *Merulina ampliata (Ellis and Solander, 1786). * Scapophyllia cylindrica Milne Edwards and Haime, 1848. Family MUSSIDAE * Acanthastrea echinata (Dana, 1846). Lobophyllia corymbosa (Forskal, 1775): Odum and Odum, 1956; Young, 1971. * Lobophyllia hemprichii (Ehrenberg, 1834). *+ Scolymia sp. cf. S. vitiensis Briiggemann, 1877. *+ Symphyllia agaricia Milne Edwards and Haime, 1849. *+ Symphyllia radians Milne Edwards and Haime, 1849. Symphyllia sp.? Symphyllia nobilis (Dana, 1846): Wells, 1954. Veron and Pichon (1980) considered this species to be a junior synonym of Symphyllia cf. recta (Dana, 1846). Family PECTINIIDAE *Echinophyllia aspera (Ellis and Solander, 1786). *+ Pectinia lactuca (Pallas, 1766). Suborder CARYOPHYLLIINA Family CARYOPHYLLIIDAE *Euphyllia glabrescens (Chamisso and Eysenhardt, 1821). * Physogyra lichtensteini (Milne Edwards and Haime, 1851). “+ Plerogyra sinuosa (Dana, 1846). Suborder DENDROPHYLLIINA Family DENDROPHYLLIIDAE *Endopsammia sp. *+ Tubastraea coccinea Lesson, 1829. *Turbinaria crater (Pallas, 1766). B. R. Rosen and R. A. Kinzle (personal communication) working with Enewetak Turbinaria only recognize this species among a number of specimens examined. Turbinaria danae Bernard, 1896: Garth, 1964; Hildemann et al., 1975a, 1975b. Considered to be a junior synonym of Turbinaria frondens (Dana, 1846) by Veron and Pichon (1980). Turbinaria globularis Bernard, 1896: Odum and Odum, 1956. Considered to be a junior synonym of Turbinaria stellulata (Lamarck, 1816) by Veron and Pichon (1980). Turbinaria mesenterina (Lamarck, 1816): Odum and Odum, 1955. Considered to be a valid species by Veron and Pichon (1980). *New Enewetak record. +Not recorded from the Marshall Islands by this name in Wells (1954). SCLERACTINIA (STONY CORALS) TABLE 2 Selections of Nonsystematic Stony Coral Research at Enewetak Atoll* Type of study Coral genera‘ Reference Biochemistry Skeleta! organic Acropora, Fungia, Herpolitha, Lobophyllia, Young, 1971 composition Asexual dispersal Bioerosion Biomechanics Environmental health Invertebrate associates Productivity and trophic structure Radionuclides in skeletons Stable isotopic composition of skeletons Alkaline earth chemistry Seriatopora, Turbinaria Ecology Porites Astreopora, Favia, Goniastrea, Montipora, Oulophyllia, Pavona, Platygyra, Porites Acropora Acropora sp., Astreopora sp., Favia sp., Favites sp., Fungia sp., Montipora sp., Plesiastrea sp., Psammocora sp., Symphyllia sp., Turbinaria sp. Acropora sp., Pocillopora, Porites sp. Acropora Cyphastrea Acropora, Pocillopora, Porites, Psammocora, Seriatopora, Turbinaria Astreopora, Favia, Goniastrea, Montipora, Oulophyllia, Pavona, Platygyra, Porites Acropora Acropora sp., Pocillopora Acropora sp., Fungia sp., Pocillopora Acropora, Astreopora sp., Cyphastrea, Favia, Lepastrea,t Lobophyllia sp., Montipora sp., Pocillopora sp., Porites, Stylophora sp., Turbinaria Acropora, Astreopora, Cyphastrea, Echinopora, Favia, Favites, Goniastrea, Lobophyllia, Montipora, Platygyra, Plesiastrea, Pocillopora, Porites, Stylophora, Turbinaria Acropora Pavona Geochemistry Astreopora, Favia, Fungia, Goniastrea, Herpolitha, Hydnophora, Oulophyllia, Parahalomitra, Platygyra, Pocillopora, Porites, Psammocora Astreopora sp., Favia, Goniastrea, Porites Favia Acropora sp., Favia sp., Montipora sp., Platygyra sp., Pocillopora sp., Porites sp., Psammocora sp. Highsmith, 1980a Highsmith, 1980b, 1981a, 1981b, 1981c Vosburgh, 1977, 1978, 1982 Johannes et al., 1972a Banner and Banner, 1968 Bruce, 1969 Bruce, 1979 Garth, 1964 Highsmith, 1980b, 1980c, 1981b, 1981c Humes, 1981; Humes and Dojiri, 1982; Humes and Stock, 1973 Knudsen, 1967 Johannes et al., 1972b Odum and Odum, 1955 Odum and Odum, 1956 Smith and Harrison, 1977 Wethey and Porter, 1976a, 1976b Buddemeier et al., 1974 Knutson and Buddemeier, 1973 Haggerty et al., 1980. Buddemeier et al., 1982 “Any species names given for each of these genera can be found in the checklist of corals (Table 1). +Reclassified as a species of Plesiastrea in Odum and Odum (1956). (This table continued on next page.) 73 74 DEVANEY AND LANG TABLE 2 (cont’d) Type of study Coral genera*® Reference Immunology Immunorecognition Acropora, Cyphastrea sp., Favites, Hildemann et al., Fungia, Montipora, Pocillopora, 1975a, 1975b Porites, Psammocora sp., Seriatopora, Stylocoeniella, Turbinaria Physiology Abnormal growth Acropora Cheney, 1975 Calcification Acropora Goreau, 1959 Chemical uptake Acropora, Fungia, Pocillopora D’Elia, 1977 and release Pocillopora D’Elia and Webb, 1977 Acropora sp., Pocillopora, Porites Muscatine and D’Elia, 1978 Pocillopora Pilson, 1974 Acropora, Fungia, Leptastrea, Pomeroy and Pocillopora, Porites Kuenzler, 1969 Acropora sp. Pomeroy et al., 1974 Growth rate Astreopora, Favia, Fungia, Buddemeier et al., 1974 Goniastrea, Herpolitha, Hydnophora, Oulophyllia, Parahalomitra, Platygyra, Pocillopora, Porites, Psammocora Favia, Goniastrea, Porites Highsmith, 1979 Favia Haggerty et al., 1980 Favia, Goniastrea, Porites, Psammocora Knutson et al., 1972 Acropora, Pocillopora Roth et al., 1982 Mucus release Acropora sp., Porites sp. Coles and Strathmann, 1973 Temperature Pocillopora Clausen and Roth, 1975 adaptation and tolerance Montipora, Pocillopora Coles and Jokiel, 1977 Acropora, Fungia, Leptastrea, Coles et al., 1976 Porites, Pocillopora Sexual Reproduction Pocillopora Richmond, 1982 Pocillopora Richmond and Jokiel, 1984 Acropora, Pocillopora, Seriatopora Stimson, 1978 *Any species names given for each of these genera can be found in the checklist of corals (Table 1). | +Reclassified as a species of Plesiastrea in Odum and Odum (1956). ACKNOWLEDGMENTS Gérard Faure, Univ. de la Réunion (Thamnasteriidae, We are grateful to participants of the 1976 reef coral workshop for permitting inclusion of many of the new records of corals, which were based for the most part on workshop and MPRL reference collection material. We par- ticularly wish to thank the following for their reviews of this checklist: Maya B. Best, Rijksmuseum van Natuurlijke Historie (Favi- idae, Merulinidae, Mussidae, Oculinidae, Pectiniidae, Caryophylliidae) Astrocoeniidae) Paul L. Jokiel, Hawaii Institute of Marine Biology (Poriti- dae; nonsystematic scleractinian literature _—for Enewetak) Robert A. Kinzie Ill, University of Hawaii (Turbinaria) Austin E. Lamberts, Grand Rapids, Michigan (Astreopora) James E. Maragos, U. S. Army Corps of Engineers, Hawaii (Montipora) Michel Pichon, James Cook University of North Queens- land (Montipora, Fungiina, Turbinaria) SCLERACTINIA (STONY CORALS) 75 Richard H. Randall, University of Guam (Montipora) Brian R. Rosen, British Museum (Natural History) (Fungi- idae, Dendrophylliidae) John E. N. Veron, Australian Institute of Marine Sciences (Stylocoenielia, _Psammocora, Pocilloporidae, Acrhelia, Faviidae, Merulinidae, Mussidae, Oculinidae, Pectini- idae, Caryophylliidae) Carden C. Wallace, Dept. of Home Affairs and Environ- ment, Canberra (Acropora) John W. Wells, Cornell University (Astrocoeniidae, Dendro- phylliidae, | Fungiidae, Thamnasteriidae; historical records) Financial support for the workshop was provided by NSF grant No. OEC 76-22196 and by ERDA contract No. AT(26-1)-628. REFERENCES Banner, A. H., and D. M. Banner, 1968, Contributions to the Knowledge of the Alpheid Shrimp of the Pacific Ocean, Part XII. Collections from the Marshall and Caroline Islands, Micronesica, 4: 261-294. Bruce, A. J., 1969, Notes on Some Indo-Pacific Pontoniinae XIV. 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Wijsman-Best, M., 1980, Indo-Pacific Coral Species Belonging to the Subfamily Montastreinae Vaughan and Wells, 1943 (Scleractinia-Coelenterata), Part II: The Genera Cyphastrea, Leptastrea, Echinopora and _ Diploastrea, Zool. Meded. (Leiden), 55: 235-263. Yabe, H., T. Sugiyama, and M. Eguchi, 1936, Recent Reef- Building Corals from Japan and the South Sea Islands Under the Japanese Mandate, I. Sci. Rep. Tohoku Univ. Second Ser. (Geol.), Spec. Vol., 1: 1-66. Young, S. D., 1971, Organic Material from Scleractinian Coral Skeletons-1. Variation in Composition Between Several Species, Comp. Biochem. Physiol. B. Comp. Biochem., 40: 113-120. Brachiopods of Enewetak Atoll RICHARD E. GRANT Department of Paleobiology National Museum of Natural History Smithsonian Institution, Washington, D. C. 20560 Brachiopods have one of the longest continuous his- tories of any extant group of metazoans, originating in the early Cambrian period and still thriving in modern seas. Although brachiopods have long been considered rare and in evolutionary decline, modern studies show they are abundant, widespread, and highly diverse. Living bra- chiopods are classified into about 100 genera and occupy niches from tide pools to the abyss; their apparent rarity is partly the result of their patchy distribution. Those on Enewetak are small and inconspicuous, living in “cryptic” habitats under the shade of coral fronds, in recesses in the reef, or on lagoon pinnacles. Hitherto, the only published account of them at Enewetak is as fossils from drill holes (Cooper, 1964), although their presence alive on the reef is well-known (Grant, 1971; Zumwalt, 1978). Three species, Thecidellina congregata Cooper, Frenulina sanguinolenta Gmelin, and Argyrotheca arguta Grant, are known at Enewetak from shells. In addition, Lingula sp. is known by larvae taken in plankton tows in the lagoon (Gilmartin, 1958), and a species of Crania is expected based on specimens found at Bikini (Cooper, 1954). These taxa seem to constitute a suite of bra chiopods that characterizes the shallow waters of tropical islands, not only in the Pacific (Cooper, 1954) but also in the Caribbean and, with some different species, in the Red Sea (Jackson et al., 1971). (See Table 1 for checklist of Enewetak species and Fig. 1 for map of Enewetak.) METHODS Brachiopods were collected during September 1969 by the author using scuba equipment to depths of about 50 m. Further collections of Thecidellina congregata to a depth of about 65 m were made in 1972 by G. S. Zumwalt. His specimens are deposited in the United States National Museum of Natural History (USNM) and were used to sup- 77 Chapter 9 TABLE 1 Checklist of Enewetak Brachiopods Class INARTICULATA Order LINGULIDA Superfamily LINGULACEA Family LINGULIDAE Lingula sp., larvae: Gilmartin, 1958. Class ARTICULATA Order TEREBRATILIDA Superfamily TEREBRATELLACEA Family MEGATHYRIDIDAE Argyrotheca arguta Grant: Grant, 1983. (=Argyrotheca sp. Cooper, 1954.) Family DALLINIDAE *Frenulina sanguinolenta Gmelin, 1792. Order THECIDEIDA Superfamily THECIDEACEA Family THECIDELLINIDAE Thecidellina congregata Cooper, 1954. (= Thecidellina maxilla Hedley: Grant, 1972.) *New Enewetak record. plement material for this report. Each station where bra- chiopods were obtained has been assigned a number in the USNM register of localities, Department of Paleobiology. Station data are summarized in Table 2. The Enewetak brachiopods cannot be seen under water because they are small and live in dark habitats; therefore, collecting had to be somewhat random. Pieces of coral were broken out of likely looking habitats and brought to the surface for exam- ination, much in the manner described by Jackson et al. (1971) although not on so vast a scale. Most individuals were left attached to their coral slabs and were dried in the sun; those that became detached were preserved in alcohol for immediate study and were dried later. ECOLOGY Brachiopods were found on Enewetak at depths rang- ing from 5 m to 65 m, the limit of scuba equipment. At Bikini, where they were collected by dredging, they are reported as deep as 480 m (Cooper, 1954). The shal- lowest occurrence was a small Frenulina sanguinolenta on a 78 GRANT = 4 | | (@) 10 KM Seen eee a a = West : Spit ; P 12 \ VAN aT RIGILI 19 21,22 10 ZOre ®. CHINIMI “ I s. @M ; UTI Southwest . S On Passage 15 16-189 % e 2, ., PARRY of CoS Lz GRINEM Nia oe i 7@ ? "ORB \, f ; 14 of & IGURIN™ ENEWETAK South Channe/ Fig. 1 Map of Enewetak showing where brachiopods were collected. Numbers are last digits of USNM locality numbers 32007-22. coral frond at 5 m. The abundance and diversity of bra- chiopods increase with depth and with increased darkness in small caves and recesses in the reef. On the seaward side of Biken Island, one piece of coral-cum-sponge con- tained a density of Thecidellina congregata that by extrapo- lation amounts to 7000/m?, and another has 40 individu- als on a surface 7 XX 17 cm, amounting to about 3400/m? (Figs. 6 through 8). These densities compare to those found by Jackson et al. (1971) at Jamaica and Curagao where densities of 5000 to 10,000/m? were encountered. These figures may appear to exaggerate the density of brachiopods on these reefs, because the com- mon impression is that brachiopods are rare. Most divers do not encounter brachiopods, however, because they do not search for them deliberately, and brachiopods are essentially invisible under water. In addition, the distribu- BRACHIOPODS 79 TABLE 2 Station Data for Enewetak Brachiopods Collected in 1969 and 1972 Bottom Sample depth, depth, No. of Station® Locality m m Taxa specimens Grant’s 1969 Localities 32007 Enewetak Islet, ~1200 m off personnel pier 18 17 F. sanguinolenta 1 32008 ~1500 m off personnel pier: low bioherms 21 14 F. sanguinolenta 1 32009 Same bioherm as 32008 21 21 F. sanguinolenta 2 32010 Jinimi Islet ~400 m east 14 12 F. sanguinolenta 3 32010a dJinimi Islet, same area as 32010 21 21 F. sanguinolenta 3 32011 Pinnacle, 11 km offshore bearing 320° — 20 F. sanguinolenta 1 from personnel pier T. congregata 9 32012 Jinedrol Islet, ~2000 m offshore, bearing 18 17 F. sanguinolenta 29 285°, broken coral T. congregata 2 32013 Kidrenen Islet, ~1000 m offshore, bearing 26 24 F. sanguinolenta 11 25°, low bioherms and dead coral T. congregata 13 32013a Kidrenen Islet, ~400 m offshore, — 5 F. sanguinolenta 1 shallow slope 32014 Ikurin Islet, 1500 m, 10° off east end, 18 14 T. congregata 1 dying bioherms A. arguta 19 32015 Pinnacle Qk FIR (map 6033) 6 km off 35 30 F. sanguinolenta 3 personnel pier 32016 Medren Islet, ~ 1000 m offshore, bearing 21 21 T. congregata 2 270°, low bioherms 32017 Medren Islet, higher pinnacles 21 12 T. congregata 1 (same region as 32016) 32018 Medren Islet pinnacles (same region as 32016) 21 8 F. sanguinolenta 5 32019 Biken Islet, wall of channel just SW of islet 9 9 F. sanguinolenta 1 T. congregata 60 32020 Biken Islet, steep reef slope 120+ 14 T. congregata 1 32021 Biken Islet, steep reef slope 120+ 23 F. sanguinolenta 1 T. congregata 71 32022 Biken Islet, steep reef slope 120+ 35 T. congregata 210 A. arguta 13 Zumwalt’s 1972 Localities Medren Islet, ~1000 m NW of islet, 20-40 20-30 T. congregata deep channel reef Pole pinnacle, lagoonward extension of 37 37 T. congregata channel, 1 km offshore Biken Islet, steep seaward slope 120+ 37+ T. congregata *Stations 32007 to 32018 are lagoon stations, stations 32019 to 32022 are seaward stations. tion of brachiopods is patchy, so it can be said that under the best conditions of light and depth the local density of brachiopods, especially T. congregata, can be very great. Brachiopods tend to associate with bryozoans, a combi- nation that dates as far back as the Ordovician period. The undersides of many coral fronds also contain encrust- ing sponges along with the brachiopods, an association so consistent that Jackson et al. (1971) refer to a “brachiopod—coralline sponge community.” They note that true coral reefs of the modern type originated in the Juras- sic period and that the light dependency of scleractinians Necessarily produces a foliaceous framework with numerous cavities, because light-dependent corals cannot fill their own gaps. The association of Thecidellina and Argyrotheca (plus various terebratulinids) dates at least as far back as the Eocene in the Pacific area (Cooper, 1971), indicating that the cryptic environment produced by the foliaceous reef has been exploited for a long time. Surlyk (1972, Fig. 12) cites a Cretaceous association of a the- cidean and two species of Argyrotheca but shows them occupying exposed hardgrounds. Jackson et al. (1971) noted that the competition for space on the undersides of corals is keen, an observation that is confirmed at Enewetak. The shapes of the two 80 forms that are tightly attached, T. congregata and Argyrotheca arguta, are clearly adapted to avoid encrusta- tion by the ubiquitous sponges and algae; they grow away from the substrate and thus keep the commissure elevated from the encrusters. Frenulina sanguinolenta attaches by a pedicle that holds the shell well above the substratum (actually below it, on these undersides) and affords some rotational mobility that can help to ward off encrustation. Frenulina sanguinolenta is not as negative to light as is T. congregata. Several samples with F. sanguinolenta also contain abundant Halimeda algae, but other pieces in the collection have F. sanguinolenta and T. congregreta in association. The brachiopod found in the shallowest habitat was F. sanguinolenta in 5 m off Kidrenen (loc. 32013a) where a single tiny individual was taken from a shady place in a niche under an overhang. Apparently the some- what warmer water at this shallow depth was more a fac- tor than shelter from light in limiting the presence of brachiopods. Two representatives of the Class Inarticulata also prob- ably occur at Enewetak. The larva of Lingula sp. is reported in plankton hauls made in transects across the southern part of the lagoon (Gilmartin, 1958). Species of Lingula normally burrow in unconsolidated sediment in the intertidal’and shallow subtidal zones. This habitat was not explored thoroughly, but the presence of the larva indi- cates that such a search would produce specimens of a Lingula. Another inarticulate genus that can be expected to be found at Enewetak is Crania. An undetermined species in this genus was reported from Bikini where the fauna is essentially the same as at Enewetak (Cooper, 1954). Its small size and tightly cemented habit make it difficult to find (it looks like a small, very flat limpet); the largest specimen reported by Cooper is 3 mm wide, a pale salmon color that blends into the surroundings, and a shape that tends to conform to the contours of the substrate. ARTICULATA Argyrotheca arguta is characterized by its small size and white, clear, or translucent shell (Figs. 2 and 3) with weak to absent costae. It had been recognized from Enewetak as not belonging to any described species (Cooper, 1954) but was not described for fear that it was represented only by juveniles. The present Enewetak sam- ple is adequate, however, and contains small shells along with the majority that seem to cease growth between 2 and 3 mm in width (Table 3). The narrow outline, lack of costae (other species are typically strongly costate), and lack of pigmentation make it unique among species of Argyrotheca. It was described recently (Grant, 1983), is known only from Enewetak and Bikini so far, and may prove to be endemic to the Marshall Islands. The two Pacific species described by Blochmann (1910, 1914) from Australia and Tasmania, A. australis and A. mayi, are much longer than A. arguta and have the strong costae that characterize most species of the genus. GRANT Fig. 2. Argyrotheca arguta from underside of coral in lagoon off Ikurin Island at depth 15 m, locality 32014; juvenile in small depression (4X). Fig. 3 Argyrotheca arguta on same slab from 32014 as in Fig. 2; adult and juvenile shells in close proximity (4X). The colorless shell of A. arguta is another distinguishing feature. Argyrotheca arguta differs from the other pedi- culate species of Enewetak, Frenulina sanguinolenta, in its smallness, its flatter profile, its wide hinge, its tri- angular rather than round foramen, and, especially, its pel- lucid shell material. Argyrotheca arguta attaches to the substrate by means of a short pedicle. It attaches somewhat loosely, i.e., the pedicle is longer than is normal in other members of the genus. Argyrotheca arguta favors cryptic habitats under coral fronds and in recesses, as would be expected from its colorless and translucent shell. At Enewetak it was found BRACHIOPODS 81 TABLE 3 Measurements of A. arguta in mm from Ikurin (sta. 32014)° Width Length 0.8 - 1.5 1.4 1.6 1.2 1.8 1.4 Dall 2.0 2.5 2.6 2.8 3.2t *Most specimens are preserved still attached to their substrates, so the thickness measurement was impossible to obtain. +Holotype: USNM 265875 only at two localities, both in association with Thecidellina congregata. One locality is on the seaward side of the reef and the other on the lagoon side; both are on the southwest segment of the atoll [Biken and Ikurin where 13 and 19 specimens were found, respectively (Table 2)]. The shell of Frenulina sanguinolenta is small for bra- chiopods in general and even for a species of Frenulina, yet this is the largest brachiopod at Enewetak. Adults nor- mally are slightly wider than long, and the shape is that of a generalized terebratulide, with biconvex profile and api- cal circular foramen (Figs. 4 and 5) that earned the bra- Fig. 4 Frenulina sanguinolenta from pinnacle in lagoon, locality 32010; the largest specimen collected (w is 12.3 mm) and a smaller one (w is 6.9) with pedicle and loose pedicular fibers (2X). Fig. 5 Frenulina sanguinolenta from niches at 24 m depth in the lagoon, shown attached to substrate (4X). Shells are pink with white stripes or solid pink. chiopod the vernacular name “lamp shell” (the ancient oil lamp). The color is distinctive, normally white with pink stripes or pink mottling in irregular rows but occasionally a solid pink or red color without stripes. The dorsal valve bears a shallow median depression resulting in a sulcate anterior commissure. The dorsal loop consists of a large “hood” of thin shell material attached to a short slim median septum by means of two broad lateral bands. The ventral valve has deltidial plates outlining the anterior part of the foramen, which are conjunct in some specimens but disjunct in most. The species thus is variable, but unmistakable, in the Enewetak brachiopod fauna. Frenulina sanguinolenta is the most widespread and smallest living species of the genus (Hatai, 1940; Cooper, 1957). This species like other brachiopods at Enewetak prefers cryptic habitats. It is not as abundant locally as T. congregata but was found at more localities than any other brachiopod species, being present at 14 of the 18 localities sampled (Table 2). Most specimens remain attached to their substrates by the pedicle, so only the width measurement could be made without danger of dam- age to the shells. Table 4 compares the length, width, and thickness of 14 Enewetak specimens. However, the smal- lest specimen examined had a width of 0.8 mm. The beak curves toward the substrate, so the dorsal valve is held facing the coral frond or other “overhang” that the shells cling to. Relative to the sea floor, the dorsal is thus upper 82 TABLE 4 Shell Measurements of Frenulina sanguinolenta in mm Width Length Thickness Locality 2.9 2.9 1e2 32018 Sh7/ 3.8 1.6 32018 4.0 4.6 2.5 32018 4.1 4.0 7, 32018 4.8 4.4 2.0 32010 4.8 4.6 all 32008 6.3 6.3 3.4 32017 6.9 6.6 3.1 32010 6.9 6.4 3.0 32013 Us 6.9 3.5 32017 7.4 7.0 3.6 32017 8.6 8.8 = 32010 10.0 9.6 5.4 32017 11.4 12.3 5.7 32010 and the ventral lower as the shells hang from the ceilings of their cryptic recesses. The red color looks greenish under water, and the stripes render the shells nearly invisi- ble to the human eye in the dim and shimmering light. Thecidellina congregata was described originally from Bikini Atoll where it occurred as deep as 480 m (Cooper, 1954). It is the most abundant brachiopod at Enewetak, although not the most widespread. Thecidellina congregata can be identified by its cemented habit, with low cup- shaped ventral valve and lid-like dorsal valve (Figs. 6 through 9). Its color ranges from white (mostly dead shells) to bluish gray in the majority of living shells. It is not immediately distinguishable from T. maxilla, known from Bikini (Cooper, 1964), although close inspection reveals several differences. Thecidellina congregata is smaller (Table 5), has the blue-gray color, and the dorsal median septum is wider at the anterior than at the posterior, where that of T. maxilla maintains nearly a uniform width from back to front. Zumwalt (1978) conducted a detailed anatomical and functional study of this species, based upon material he collected at Enewetak. He found that both the diductor and the adductor muscles insert in the hemispondylium, which is a muscle platform that changes length and shape with growth and general configuration of the valve, to pro- duce the most advantageous angles for muscle action. Hence the hemispondylium varies from one individual to the next. As implied by the name, T. congregata can occur in abundant patches where extrapolated densities run to several thousand per square meter. Judging from the sam- ples taken at Enewetak and reports of other species of Thecidellina from elsewhere (e.g., Jackson et al., 1971), Thecidellina survives best in cryptic habitats. It feeds by opening the valves and exposing the lophophore to the seawater, which is then pumped across the filaments of the lophophore. The lophophore is considered a modified GRANT Fig. 6 Thecidellina congregata clustered on underside of coral frond from sea side of Biken Island at USNM locality 32022 (2X). These are the specimens whose measurements in the species description indicate a width range from 0.5 to 5.2 Fig. 7 Thecidellina congregata from shallow cave at depth about 33 m on sea side of Biken Island, USNM locality 32022; combination of living shells and dead empty ventral valves with narrow variation in size. trocholophe, with a deep median indentation along the sides of the dorsal median septum. The normal position for feeding seems to be with the dorsal valve open to a posi- tion nearly perpendicular to the margin of the ventral valve BRACHIOPODS 83 Fig. 8 Thecidellina congregata on coral frond from same place as Fig. 7. Specimen at broken edge has grooves on the dorsal valve reflecting grooved surface to which ventral valve is attached. Such dorsal reflection of ventral irregularities is common in brachiopods. Fig. 9 Thecidellina sp. in feeding position (specimen from Curagao, photo courtesy D. L. Meyer, shown in Grant, 1972). (Fig. 5; and Grant, 1972: pl. 5, Fig. 21). This position would be highly vulnerable to predators and damage by grazing fish, possibly accounting for the preference toward sheltered environments and development of a muscle sys- tem with a quick closing mechanism (Rudwick, 1961; Zumwalt, 1978). Thecidellina congregata occurs at 10 of the 18 locali- ties where brachiopods were recovered, fewer than the more widespread species F. sanguinolenta. It is, however, the most abundant species, being represented by at least 370 specimens (both dead and alive). Of this number, 340 were obtained from Biken Islet, on the seaward slope or the seaward end of the channel leading out to that slope. Its abundance increases with depth, to 65 m, but it is doubtful that the abundance continues to increase to depths as great as the 480 m reported by Cooper (1954). TABLE 5 Shell Measurements of Thecidellina congregata in mm* Length Width Length Width Length *Specimens cemented to substrate permit only length and width measurements, but these afford an idea of size and varia- tion. All measured specimens are from the underside of a thin coral frond from locality 32022 (Biken, 28 m). REFERENCES Blochmann, F., 1910, New Brachiopods from South Australia, in Verco, J. C., The Brachiopods of South Australia, Trans. R. Soc. S. Aust., 34: 88-89. __, 1914, Some Australian Brachiopods, R. Soc. Tasmania Pap. Proc., 1913: 112-115. Cooper, G. A., 1954, Recent Brachiopods. Bikini and Nearby Atolls, Part 2. Oceanography (biologic), U. S. Geological Sur- vey Prof. Paper, 260-G, pp. 315-318. ___, 1964, Brachiopods from Eniwetok and Bikini Drill Holes. Bikini and Nearby Atolls, Marshall Islands, U. S. Geological Survey Prof. Paper, 260-FF, pp. 1117-1120. ___, 1971, Eocene Brachiopods from Eua, Tonga. Late Eocene Fossils from Eua, Tonga, U.S. Geological Survey Prof. Paper, 640-F, pp. 1-9. __, 1973, Vema’s Brachiopods (Recent), Smithson. Contrib. Paleobio., 17: 1-51. , 1977, Brachiopods from the Caribbean Sea and Adjacent Waters, Stu. Trop. Oceanog. (Miami), 14: xii + 212. Dall, W. H., 1895, Report on Mollusca and Brachiopoda Dredged in Deep Water, Chiefly near the Hawaiian Islands, with Illus- trations of Hitherto Unfigured Species from Northwest Amer- ica, Proc. U. S. Nat. Mus., 17 (1032): 675-733. __, 1920, Annotated List of the Recent Brachiopoda in the Col- lection of the United States National Museum, with Descrip- tions of Thirty-Three New Forms, Proc. U. S. Nat. Mus., 57 (2314): 261-377. Gmelin, J. F., and C. Linné, 1792, Systema Naturae, (13th ed.), 1: 3021-3909. Gilmartin, M., 1958, Some Observations on the Lagoon Plankton of Eniwetok Atoll, Pac. Sci., 12: 313-316. Grant, 1971, Brachiopods in the Permian Reef Environment of West Texas, Proceedings of the North American Paleontology Convention, Part J, 1444-1481. ___, 1972, The Lophophore and Feeding Mechanism of the Pro- ductidina (Brachiopoda), J. Paleontol., 46: 213-248. ___, 1983, Argyrotheca arguta, a New Species of Brachiopod from the Marshall Islands, Western Pacific, Proc. Biol. Soc. Wash., 96: 178-180. 84 Hatai, K. M., 1940, Cenozoic Brachiopoda from Japan, Sci. Rep. Tohuku Univ., Second Ser. (Geol.), pp. 1-413. Jackson, J. B. C., T. F. Goreau, and W. D. Hartman, 1971, Recent Brachiopod—Coralline Sponge Communities and Their Paleontological Significance, Science, 173: 623-625. Rudwick, M. J. S., 1961, ‘Quick’ and ‘Catch’ Adductor Muscles in Brachiopods, Nature (Lond.), 191: 1021. GRANT Surlyk, F., 1972, Morphological Adaptations and Population Structures of the Danish Chalk Brachiopods, Konig. Danske Vid. Selek., Biol. Skrift., 19: 57. Zumwalt, G. S., 1978, The Functional Morphology of the Tropical Brachiopod Thecidellina congregata Cooper, 1954, unpub- lished M.S. thesis, University of California at Davis. Chapter 10 Reef-Dwelling Bryozoans of Enewetak Atoll ROGER J. CUFFEY* and ROBERT S. COXt "Department of Geosciences (Deike Bldg.), Pennsylvania State University, University Park, Pennsylvania 16802; +Museum of Paleontology, University of Michigan, Ann Arbor, Michigan 48109 INTRODUCTION Purpose The goals of this report are to list the bryozoans found on the modern reefs of Enewetak Atoll, illustrate the most conspicuous species, indicate their biogeographic implica- tions, and summarize their ecology (particularly ecozonal distributions and constructional roles). Significance The contribution made by the phylum Bryozoa to the Enewetak reefs, although known in general terms (Cuffey, 1970, 1973, 1978), has not until now been detailed at the species level for the benefit of those concerned with vari- ous aspects of the Enewetak ecosystem. Such information will also constitute a precise data base for comparative studies of fossil reefs containing these animals. Moreover, the Enewetak species are of special broader interest because they represent the first modern atoll-reef bryozoan fauna to be analyzed thoroughly. Their distributions thus will shed light on bryozoan involvement in reefs in general. Collections Over 40 localities on Enewetak Atoll were sampled by scuba diving and reef walking in late 1969. Sites were selected to provide balanced but comprehensive coverage of the major environments, particularly ecozones, developed on the atoll (Cuffey, 1973, pp. 26, 36-38; Cuf- fey, 1978, pp. 68, 72; Haggerty, Weber, Cuffey, and Deines, 1980, p. 97; Basile, Cuffey, and Kosich, 1984, Fig. 1). However, no samples were dredged from below 40 m (135 ft). 85 Literature Most useful in identifying the Enewetak bryozoans were the monographs by Canu and Bassler (1927, 1929), Harmer (1915, 1926, 1934, 1957), Osburn (1950, 1952, 1953), Levinsen (1909), and Soule and Soule (1973). Additional papers too numerous to list here include occa- sional relevant species descriptions as well. FAUNAL COMPOSITION Overall, 84 bryozoan species, 73 cheilostomes and 11 cyclostomes in 61 genera have been identified on Enewetak (Table 1). Of these species, six are new, being published elsewhere (Cox and Cuffey, submitted). In order to refer to these without creating taxonomic problems, they are denoted simply as “n. sp.” followed by the pro- posed trivial name in quotation marks and parentheses. Among the remaining species, several generic reassignments were required (incorporated in Table 1) due to changes in concepts since publication of the earlier monographs cited above. Certain species are particularly conspicuous among the Enewetak bryozoans and so comprise a distinctive assem- blage (Figs. 1 and 2) characteristic of this atoll and possi- bly others. Such species appear conspicuous for different reasons. Some are represented by several large colonies, others by many small colonies, still others by a large total substrate area encrusted, and others yet by a high propor- tion of the bryozoan fauna at individual localities. These species are also differently distributed within the atoll, as discussed below. The Enewetak bryozoan species are characterized by relatively few colony growth forms (Cuffey, 1973, pp. 29-31 and references therein). Most numerous by far (both colonies and species) are encrusting cheilostomes, thin sheets or crusts ranging from many square millimeters in area down to a few zooecia. Most of the Enewetak cheilo- stome families (Table 1) studied by Cox (1983; also Cox and Cuffey, submitted) grow as encrusting sheets. A few sheets consist of more than one layer of zooecia, but none encountered were thickened into multilaminar nodular masses (not even Celleporaria albirostris, which does form such masses in Bahamian bryozoan reefs; Cuffey, Gebelein, Fonda, Bliefnick, Kosich, and Soroka, 1977). More regular CUFFEY AND COX TABLE 1 Enewetak Atoll Bryozoans Systematics Biogeography’* Ecologyt Phylum BRYOZOA Class GYMNOLAEMATA Order CHEILOSTOMIDA Suborder ANASCINA Infraorder INOVICELLATA Family AETEIDAE Aetea truncata PAE Lg;d Infraorder MALACOSTEGA Family CALLOPORIDAE Alderina brevispina P Ol;s Family HINCKSINIDAE Antropora claustracrassa P Lg;d A. laguncula P Lg;d +A. ovata P Ol,Lg;d,s Cauloramphus cf. brunea P Lg;d +Cranosina coronata PI Ol,Ow,Lg;d,s,f Infraorder COELOSTEGA Family ONYCHOCELLIDAE +Smittipora americana Jee Ol,Ow,Lg;d,s Family MICROPORIDAE Caleschara levinseni P,I Ol;d Family THALAMOPORELLIDAE Thalamoporella distorta C Ol,Lg;d,s +T. expansa P Ol,Ow,Lg;d Infraorder PSEUDOSTEGA Family ASPIDOSTOMATIDAE Monoporella fimbriata P O1l,Ow,Lg;d,s Infraorder CELLULARINA Family FARCIMINARIIDAE Didymozoum marginatum EB Lg;d Nellia cf. tenuis |2e Lg;d Family BUGULIDAE Caulibugula ciliata P Lg;d Family BEANIIDAE Beania cupulariensis EG Lg;d Family SCRUPOCELLARIIDAE Caberea ellisi 2) Ol;s Scrupocellaria longispinosa P Ol,Lg;d,s Suborder CRIBRIMORPHINA Family CRIBRILINIDAE Cribrilaria calamorpha 2 Ol,Lg;d,s C. radiata PLC Ol,Lg;d,s C. simulator F Lg;d,s Figularia n. sp. (“cribricapitata” ) E Lg;d Suborder ASCOPHORINA Family ADEONIDAE Reptadeonella flagellifera I Ol,Lg;d,s £R. joloensis Pil Ol,Lg;d,s “Geography: P, Pacific Ocean; I, Indian Ocean; C, Caribbean and western Atlantic; E, Enewetak endemic; F, previously known only as fossils. tEcozone: Ol, Ow, oceanward reefs on leeward or windward sides of atoll, respectively; Lg, lagoonal reefs. Depth: d, deep reefs; s, shallow reefs; f, reef flats. +The several most conspicuous species on Enewetak (Figs. 1 and 2). (This table continued on next page.) REEF-DWELLING BRYOZOANS TABLE 1 (cont’d) Systematics Biogeography’* Ecologyt Family CELLEPORARIIDAE Celleporaria albirostris PLC Lg;d,s C. columnaris Pil Lg;s C. n. sp. (“eniwetokensis” ) E Lg;s C. granulosa Pil Lg;d,s C. labelligera PI Ol;d,s C. vagans PLE Lg;d,s Trematooecia turrita P,I,C Lg;s Family EXECHONELLIDAE Exechonella magna P Ol,Lg;d,s E. tuberculata PI Ol,Ow,Lg;d,s Family PETRALIELLIDAE Mucropetraliella robusta P Ol,Lg;d Robertsonidra argentea Pil Lg;s Family ARACHNOPUSIIDAE +Arachnopusia spathulata P Ol,Ow,Lg;d,s,f Family CHORIZOPORIDAE +Chorizopora ventricosa P Ol,Ow,Lg;d,s Family HIPPOTHOIDAE +Trypostega venusta P,I,C Ol,Ow,Lg;d,s Family HIPPOPORINIDAE Stephanosella bernardii PI Ol,Lg;d,s Family HIPPOPODINIDAE + Hippopodina feegeensis Pe Lg;s H. pulcherrima LC Ow,Lg;d,s Family CREPIDACANTHIDAE Crepidacantha grandis P Ol,Lg;d,s C. poissonii P,I,C Ol,Lg;d Family SMITTINIDAE Codonellina montferrandii P,LC Lg;d +Parasmittina alanbanneri P Ol,Lg;d,s P. crosslandi P,C Lg;d,s P. raigiformis P Lg;d,s P. spathulata P,C Lg;d,s P. cf. tropica PI Lg;s Porella n. sp. (“ikurenensis’ ) E Ol,Lg;d,s Rimulostoma signata P,I,C Lg;d,s Smittoidea pacifica P Ol,Ow,Lg;d,s Family TEUCHOPORIDAE Teuchopora verrucosa Pil Lg;d Family SCHIZOPORELLIDAE Arthropoma circinatum Pil Ol,Lg;d,s +Calyptotheca impar Pil Ol,Ow,Lg;d,s Escharina pesanseris PI,C Ol,Ow,Lg;d,s Stylopoma duboisii Pl Ol,Lg;d,s Family GIGANTOPORIDAE Cosciniopsis coelatus P Ol,Lg;d,s +C. n. sp. (“enewetakensis”) E Ol,Ow,Lg;d,s Gigantopora pupa RC Lg;d Thornelya n. sp. (“aniyaaniensis” ) E Lg;d,s T. n. sp. (“marshallensis” ) E Lg;d,s “Geography: P, Pacific Ocean; I, Indian Ocean; C, Caribbean and western Atlantic; E, Enewetak endemic; F, previously known only as fossils. tEcozone: Ol, Ow, oceanward reefs on leeward or windward sides of atoll, respectively; Lg, lagoonal reefs. Depth: d, deep reefs; s, shallow reefs; f, reef flats. +The several most conspicuous species on Enewetak (Figs. 1 and 2). (This table continued on next page.) CUFFEY AND COX TABLE 1 (cont'd) Systematics Family CLEIDOCHASMATIDAE +Cleidochasma porcellanum Hippoporella spinigera Family MICROPORELLIDAE Fenestrulina malusii Family RETEPORIDAE (encrusting species) Drepanophora incisor D. tuberculata +Rhynchozoon curtum R. solidum Family RETEPORIDAE (fenestrated species) Triphyllozoon cuspidatum Family SAVIGNYELLIDAE Savignyella lafontii Family TETRAPLARIIDAE Tetraplaria veleroae Family TUBUCELLARIIDAE Tubucellaria fusiformis Class STENOLAEMATA Order CYCLOSTOMIDA Suborder TUBULIPORINA Family DIASTOPORIDAE Stomatopora cf. grandipora Family TUBULIPORIDAE Idmidronea flexuosa Platonea philippsae Tubulipora pulcherrima Family ONCOUSOECIIDAE Oncousoecia cf. abrupta Proboscina cf. dichotoma Family DIAPEROECIIDAE Diaperoecia aff. rosea Suborder ARTICULINA Family CRISIIDAE Crisia elongata C. cf. kerguelensis Suborder RECTANGULINA Family LICHENOPORIDAE Disporella cf. buski Lichenopora novazelandiae Biogeography* Ecology} PI,C Ol,Ow,Lg;d,s Pil Lg;d Pe Ow,Lg;d,s I Lg;d P,C Lg;d,s F Lg,d,s c Lg;d,s P Ol,Lg;d PLE Lg;d P Ol,Lg;d,s Pil Ol;Lg;d,s P Ol;d Jee: Lg;d P Ol,Lg;d,s Pil Lg;d,s P Ol,Lg;d,s P Ol,Lg;d,s P Ol,Lg;d,s P,I,C Ol,Lg;d,s PI Ol,Lg;d,s RG Lg;d PI Lg;d "Geography: P, Pacific Ocean; I, Indian Ocean; C, Caribbean and western Atlantic; E, Enewetak endemic; F, previously known only as fossils. +Ecozone: Ol, Ow, oceanward reefs on leeward or windward sides of atoll, respectively; Lg, lagoonal reefs. Depth: d, deep reefs; s, shallow reefs; f, reef flats. +The several most conspicuous species on Enewetak (Figs. 1 and 2). in outline, but always small and much rarer, are thin encrusting disks, lichenoporid cyclostomes. Also along the substrate are encrusting threads (aeteid cheilostomes, dias- toporid cyclostomes); encrusting branches (some tubuli- porine cyclostomes); and recumbent, though slightly elevated, branching tubes (other tubuliporine cyclostomes; Cuffey, 1985). Erect rigid lattices are certain reteporid cheilostomes (Cuffey and McKinney, 1982). Erect flexible or jointed tufts include both extremely delicate crisiid cyclostomes and somewhat more robust cheilostomes (cel- lularine, savignyellid, cheiloporinid, and tubucellariid cheilostomes). None of the Enewetak reefal bryozoans were observed to grow as erect cylindrical or flattened branches. All the Enewetak bryozoans mentioned above are geo- logically modern. Deep drilling into the late Cenozoic REEF-DWELLING BRYOZOANS aay as ss ae Fig. 1 The most abundant Enewetak bryozoans (width of each field in mm; specimens numbered within “MAPENI” suite in Paleobryozoological Research Collection at Pennsylvania State University). (a) Cranosina coronata, field 2.43 mm wide, 27-A-1; (b) Cosciniopsis n. sp. (“enewetakensis”), field 1.54 mm wide, 16-G-12; (c) Chorizopora ventricosa, field 1.01 mm wide, 27-A-la; (d) Thalamoporella expansa, field 1.51 mm wide, 17-B-2; (e) Reptadeonella joloensis, field 1.02 mm wide, 16-G-8. 89 90 CUFFEY AND COX 2 yrat as oma) 1. a es Fig. 2. The other common Enewetak bryozoans (sizes and numbers as in Fig. 1). (a) Arachnopusia spathulata, field 0.84 mm wide, 14-A-37; (b) Hippopodina feegeensis, field 1.27 mm wide, 31-C-4; (c) Rhynchozoon curtum, field 0.78 mm wide, 3-B-12; (d) Smittipora americana, field 0.94 mm wide, 21-B-4; (e) Calyptotheca impar, field 0.89 mm wide, 21-A-3; (f) Trypostega venusta, field 0.86 mm wide, 15-A-2; (g) Cleidochasma porcellanum, field 0.78 mm wide, 21-A-8; (h) Antropora ovata, field 0.79 mm wide, 25-A-22; (i) Parasmittina alanbanneri, field 0.88 mm wide, 20-B-12. REEF-DWELLING BRYOZOANS 91 limestone cap atop Enewetak’s volcanic pedestal yielded fossil remnants of encrusting cheilostomes (Escharoides? sp. and possibly Cosciniopsis aff. coelatus), a tuft-like cheilostome (Nellia aff. oculata), an unidentified reteporid cheilostome, and a tubuliporine cyclostome (Diastopora? sp.) (Brown, 1964). BIOGEOGRAPHIC IMPLICATIONS These bryozoan records from Enewetak (Table 1) extend the published ranges of all the species onto that atoll and probably also into the Marshall Islands and Micronesia overall. [A small Enewetak collection years ago was not published (D. Soule and J. Soule, 1971, personal communication)]. Most of the Enewetak species have been recorded previously from elsewhere in the Indian Ocean or western Pacific; therefore, this atoll’s bryozoans do appear typically Indo-Pacific. Several of the Enewetak species range more widely than Indo-Pacific. A few are even circumtropical or nearly cosmopolitan, most noticeably Aetea truncata, Cleido- chasma_ porcellanum, Crepidacantha poissonii, Cribrilaria radiata, Crisia elongata, Escharina pesanseris, Fenestrulina malusii, Hippopodina feegeensis, Savignyella lafontii, and Trypostega venusta. Two Enewetak forms, Rhynchozoon solidum and Thalamoporella_ distorta, represent substantial range exten- sions from the Caribbean region. Moreover, three others, Cribrilaria calamorpha, C. simulator, and Rhynchozoon curtum, were previously known only as lower and mid- Tertiary fossils from lands near the Gulf of Mexico. It is interesting to compare the Enewetak fauna with other reefal bryozoans investigated by similar methods. Several species (25), more than a quarter of the Enewetak fauna, also occur on Caribbean or western Atlantic reefs. If genera are considered, even more (34), about half the fauna, are common to Enewetak and the Bermuda-— Bahamas reefs (Fonda and Cuffey, 1976; Cuffey and Fonda, 1977, 1979). Bryozoan endemism is difficult to establish with any confidence at present because so few reef sites have been studied comprehensively for these animals. However, it is noteworthy that among the 62 encrusting cheilostome species identified from Enewetak (Table 1; Cox, 1983), six are new to science. These six species may be narrowly endemic to Enewetak or instead may be more widely dis- tributed in the surrounding region (and were discovered here simply because this atoll was the first investigated bryozoologically). Comparable reefal-bryozoan surveys on several other atolls will be required before any degrees of endemism can be determined. ECOLOGIC DISTRIBUTIONS Like other Pacific atolls, Enewetak exhibits distinct eco- zones developed across its prevailing windward—leeward gradient (northeast-southwest, respectively; Cuffey, 1973, pp. 35-40). Recognition of similar zones in fossil reefs is essential for interpreting those deposits; hence, ecozonal distributions of species in modern reefs like Enewetak Atoll are of the greatest interest for comparative paleoecologic studies. Unlike certain Caribbean coral species, the Enewetak bryozoans do not fall into sharply defined, ecozonally re- stricted assemblages. Instead, the conspicuous species (Figs. 1 and 2) characterize somewhat different parts of the atoll in a more generalized fashion. Consequently, it is sufficient here to contrast oceanward reefs with lagoonal, and deep reefs (10 to 40 m or 30 to 135 ft), shallow reefs (0 to 10 m or O to 30 ft), and reef flats (intertidal). The oceanward reefs that were examined for bryozoans are nearly vertical coral-rock cliffs on the leeward side of the atoll and sloping coral pavements on its windward side. The lagoonal reefs, all surrounded by barren sand bottoms, range from low coral knolls to platform-like patch reefs to tall steep-sided pinnacles. The rarer Enewetak bryozoan species are sparsely scattered in a few ecozones but are without immediately obvious patterns (clearly a subject for further research). Overall, bryozoan colonies appear most abundant on the deeper Enewetak reefs (below 10 m or 30 ft) and on the leeward-oceanward, and lagoonal reefs. Cranosina coronata, recognized by its large open opesia and crenulated mural rims (Fig. la), forms by far the greatest number of colonies, particularly large ones, in our collections. It thus appears as the most conspicuous Enewetak bryozoan species, occurring almost ubiquitously on deep and shallow oceanward (leeward and windward) and lagoonal reefs, and locally also on the windward reef flat. Arachnopusia spathulata, much less numerous though still common, exhibits the same distribution; it has a cal- careous frontal perforated by large pores (Fig. 2a). Simi- larly distributed, except for not inhabiting reef flats, are the next most important Enewetak bryozoans in terms of colony numbers, Cosciniopsis n. sp. (“enewetakensis”) with large peristomes and granulate frontals (Fig. 1b) and Chorizopora ventricosa with transversely striate frontals (Fig. 1c). Two species each encrust substantially larger total sub- strate areas than any others and therefore appear espe- cially conspicuous in the Enewetak fauna. Thala- moporella expansa, distinguished by opesiules and opesium-flank swellings (Fig. 1d), is abundant on the deep oceanward and lagoonal reefs. Reptadeonella joloensis, with coarse areolar pores, conspicuous peristomes, and violet-stained frontals (Fig. le), is especially abundant on deep and shallow lagoonal reefs but is also found on lee- ward oceanward reefs. Two common species are restricted to lagoonal reefs, Hippopodina feegeensis (convex granulate frontals, nearly circular apertures, Fig. 2b) on shallow reefs, and Rhynchozoon curtum (large areolar pores, slit-like apertural sinus, Fig. 2c) on deep reefs. Four species—Smittipora americana (opesiular notches, Fig. 2d), Calyptotheca impar (raised interzooecial threads, Fig. 2e), Trypostega venusta (zooeciules and minute frontal pores, Fig. 2f), Cleidochasma porcellanum (keyhole-shaped 92 CUFFEY AND COX apertures, Fig. 2g)—are common on deep and shallow oceanward (leeward and windward) and lagoonal reefs. Comparably distributed, except not on windward ocean- ward reefs, are Antropora ovata (many tiny avicularia, Fig. 2h) and Parasmittina alanbanneri (long curved avicu- laria, Fig. 2i). Bryozoan roles in reef construction are as important:to the paleoecologist as ecozonal distributions (Cuffey, 1977). The Enewetak bryozoans are all cryptic or coelobitic, func- tioning mostly as hidden encrusters under coral fronds, coral-rock ledges, detached rubble, and occasional shells. A few are locally cavity dwellers, but none were observed to be cavity fillers, frame builders, or sediment producers. The Enewetak bryozoans thus are more restricted in con- structional roles than certain cavity-filling and rock-building Bahamian relatives (Storr, 1964; Cuffey, Gebelein, Fonda, Bliefnick, Kosich, and Soroka, 1977; Cuffey and Fonda, 1979). ACKNOWLEDGMENTS Assisting in collecting were M. Zeigler, R. Port, P. Kier, R. Grant, J. Pojeta, E. Kauffman, T. Waller, T. Phelan, A. Childs, R. Mehlen, and C. Dawson, with logistical support from Eniwetok Marine Biological Labora- tory, Pennsylvania State University, National Science Foun- dation, and Atomic Energy Commission. REFERENCES Basile, L. L., R. J. Cuffey, and D. F. Kosich, 1984, Sclero- sponges, Pharetronids, and Sphinctozoans (Relict Cryptic Hard-Bodied Porifera) in the Modern Reefs of Enewetak Atoll, J. Paleont., 58: 636-650. Brown, D. A., 1964, Fossil Bryozoa from Drill Holes on Eniwetok Atoll, U. S. Geol. Surv. Prof. Pap., 260-EE, pp. 1113-1116. Canu, F., and R. S. Bassler, 1927, Bryozoaires des Iles Hawaii, Bull. Soc. Sci. Seine-et-Oise, 7 (Suppl.): 1-66. ——, 1929, Bryozoa of the Philippine Region, U. S. Nat. Mus. Bull., 100(9): 1-685. Cox, R. S., 1983, Encrusting Cheilostome Bryozoans of Enewetak (Eniwetok) Atoll—lIdentification Manual and __ Distributional Data, unpublished report for Mid-Pacific Marine Laboratory and Bishop Museum (supplemented from M.S. thesis, Pennsyl- vania State University, University Park). ——, and R. J. Cuffey, Distribution, Ecology, and New Species of Encrusting Cheilostome Bryozoans on Enewetak Atoll, Micronesica, (submitted). Cuffey, R. J., 1970, Eniwetok Bryozoans, Eniwetok Marine Biol. Lab. Ann. Rept., 1969-1970, 3 pp. ——, 1973, Bryozoan Distribution in the Modern Reefs of Eniwetok Atoll and the Bermuda Platform, Pac. Geol., 6: 25-50. ——, 1977, Bryozoan Contributions to Reefs and Bioherms through Geologic Time, Amer. Assoc. Petrol. Geol. Stud. Geol., 4: 181-194. , 1978, A Note on Bryozoans in the Modern Reefs of Eniwetok Atoll and the Australian Great Barrier Reef, Pac. Geol., 13: 65-76. ——, 1985, Reef-Dwelling Cyclostome Bryozoans on Enewetak Atoll—Comparisons and Contrasts for Fossil Reefs, Geol. Soc. Amer. Abstr. Prog., 17: 283-284. __, and S. S. Fonda, 1977, Cryptic Bryozoan Species Assem- blages in Modern Coral Reefs off Andros and Eleuthera, Baha- mas, Proceedings of the Third International Coral Reef Sympo- sium, Miami, Florida, May 1977, 1: 81-86. ——, and S. S. Fonda, 1979, Bryozoan Species Assemblages in the Modern Bryozoan-Rich Reefs at Joulters Cays (Bahamas)—General Character and Paleoenvironmental Impli- cations, Geol. Soc. Amer. Abstr. Progr., 11: 407-408. —_, C. D. Gebelein, S. S. Fonda, D. M. Bliefnick, D. F. Kosich, and L. G. Soroka, 1977, Modern Tidal-Channel Bryozoan Reefs at Joulters Cays (Bahamas), Proceedings of the Third International Coral Reef Symposium, Miami, Florida, May 1977, 2: 339-345. —_, and F. K. McKinney, 1982, Reteporid Cheilostome Bryo- zoans from the Modern Reefs of Enewetak Atoll and Their Implications for Paleozoic Fenestrate Bryozoan Paleoecology, Pac. Geol., 16: 7-13. Fonda, S. S., and R. J. Cuffey, 1976, Bryozoan Assemblages in the Modern Bermuda Reefs—Comparative Data for Analysis of Cenozoic Bioherms, Geol. Soc. Amer. Abstr. Progr., 8: 175. Haggerty, J. A., J. N. Weber, R. J. Cuffey, and P. Deines, 1980, Environment-Related Morphologic and Geochemical Variability in the Modern Reef Corals Favia pallida and Favia stelligera on Enewetak Atoll, Pac. Geol., 14: 95-113. Harmer, S. F., 1915, The Polyzoa of the Siboga Expedition, Pt. 1: Entoprocta, Ctenostomata, Cyclostomata, Siboga-Expeditie, 28a: 1-180. __, 1926, The Polyzoa of the Siboga Expedition, Pt. 2: Cheilo- stomata Anasca, Siboga-Expeditie, 28b: 181-501. , 1934, The Polyzoa of the Siboga Expedition, Pt. 3: Cheilo- stomata Ascophora I, Family Reteporidae, Siboga-Expeditie, 28c: 502-640. __, 1957, The Polyzoa of the Siboga Expedition, Pt. 4: Cheilo- stomata Ascophora II, Siboga-Expeditie, 28d: 641-1147. Levinsen, G. M. R., 1909, Morphological and Systematic Stud- ies on the Cheilostomatous Bryozoa, Nat. Forfatteres Forlag, Copenhagen, pp. 1-431. Osburn, R. C., 1950, Bryozoa of the Pacific Coast of America, Pt. 1, Cheilostomata—Anasca, Allan Hancock Pacif. Exped., 14: 1-269. __, 1952, Bryozoa of the Pacific Coast of America, Pt. 2: Cheilostomata—Ascophora, Allan Hancock Pacif. Exped., 14: 271-611. __, 1953, Bryozoa of the Pacific Coast of America, Pt. 3: Cyclostomata, Ctenostomata, Entoprocta, and Addenda, Allan Hancock Pacif. Exped., 14: 613-841. Soule, D. F., and J. D. Soule, 1973, Morphology and Speciation of Hawaiian and Eastern Pacific Smittinidae (Bryozoa, Ectoprocta), Amer. Mus. Nat. Hist. Bull. 152, pp. 365-440. Storr, J. F., 1964, Ecology and Oceanography of the Coral-Reef Tract, Abaco Island, Bahamas, Geol. Soc. Amer. Spec. Pap. 79, pp. 1-78. Chapter 11 Sipunculans and Echiurans of Enewetak Atoll DENNIS M. DEVANEY (deceased) TABLE 1 Bemice PUBshop Museum Checklist of Enewetak Sipuncula and Echiura Honolulu, Hawaii 96817 Phylum SIPUNCULA Family SIPUNCULIDAE SIPUNCULA PRA eR D EE (Damosiphon) cumanense (Keferstein, 1867). In 1950, a new sipunculan species, Siphonosoma “Siphonosoma (Siphonosoma) rotumanum (Shipley, 1898) as eniwetoki, was described by Fisher from an_ intertidal *Siphonosoma (Siphonosoma) eniwetoki Fisher, 1950; lagoon area off Boken Islet. Studies on the bioerosion of Fisher, 1950; Stephen and Edmonds, 1972. coral and coral rubble by Highsmith (1981) revealed six Sipunculus indicus Peters, 1850. Family ASPIDOSIPHONIDAE Aspidosiphon muelleri Diesing, 1851: Highsmith, 1981. Cloeosiphon aspergillus (Quatrefages, 1865): Highsmith, identified aspidosiphonid sipunculans from Enewetak (reduced to five herein). These sipunculans were con- sidered to be “responsible for more excavations in rubble 1981. than in any of the coral species studied” (p. 360). The Lithacrosiphon cristatus (Sluiter, 1902). high proportion of both sipunculans and polychaetes in Lithacrosiphon gurjanovae Murina, 1967: Highsmith, rubble, compared to those living in coral, was attributed to 1981. their feeding on detritus and algae from the rubble sur- Lithacrosiphon odhneri Fischer, 1922: Highsmith, 1981. faces. Lithacrosiphon (?) uniscutatus (Ikeda, 1904): Highsmith, In another study, examination of deeper parts of the 1981. ; ; : : f lagoon in September 1980 by the author and Patrick Colin Paraspidosiphon gigas (Sluiter, 1884): Highsmith, 1981. revealed additional sipunculans which were determined by Fai orteeiee eels eae 1980. S. J. Edmonds (South Australian Museum). An airlift wes *Phascolosoma (Phascolosoma) nigrescens Keferstein used to collect the fauna from soft lagoonal sediment and 1865. : loose coral rock. Sipunculus indicus, an undetermined juvenile Sipunculus (not S. indicus), and Siphonosoma (Damosiphon) cumanense were recovered at depths be- Family BONELLIDAE tween 9 and 21 m. At 15 m under coral rock in the °A we eioboneliia aculetn Eicher 1952 lagoon, a specimen considered to be Paraspidosiphon “Family ECHIURIDAE x : johnstoni was also collected. In July 1981 a specimen of Phascolosoma nigrescens was recovered from hard substra- tum off the wall of a lagoonal crater (Mike) made by a “New Enewetak record. nuclear detonation. Presently, at least 10 identified species of sipunculans are considered to occur at Enewetak. A Islands without further locality. Geographic range of sipun- checklist is shown in Table 1. Additional species have been culans is shown in Table 2. found in a recent (1982) examination of bioturbation- causing organisms in the lagoon sediments (Suchanek and ECHIURA Phylum ECHIURA Order ECHIUROINEA Anelassorhynchus? sp. Colin, 1986). Stephen and Edmonds (1972) record at least five addi- Echiurans are previously unrecorded from Enewetak or tional sipunculans from the Marshall Islands: Siphonosoma the other Marshall Islands. However, the bifurcate-tipped vastum, Aspidosiphon spinalis, Paraspidosiphon steenstru- proboscis (sometimes up to a meter in length) of a bonellid pii, and Lithacrosiphon cristatus all from Jaluit Atoll. The echiuran has been observed in the shallower waters of last three species are reported in Ikeda (1924). Fischer Enewetak Lagoon extending from beneath attached coral (1928) described Siphonosoma parvum from the Marshall reef outcrops. Two specimens which appear to be of the 93 94 DEVANEY TABLE 2 Identified Enewetak Sipunculans Recorded and Their Geographic Range* Pan-tropical Marshall Islands (Enewetak) East African coast to Caroline Islands Atlantic boreal; W. Indian Ocean; Japan Indo-West Pacific Indonesia, tropical Pacific and Caribbean, Japan, Australia Pan-tropical Siphonosoma (D.) cumanense Siphonosoma (S.) rotumanum Sipunculus inducus Aspidosiphon muelleri Cloeosiphon aspergillus Lithacrosiphon cristatus Phascolosoma (P.) nigrescens *Based on Stephen and Edmonds, 1972; Edmonds, 1980. same form were collected at Kawajalein Atoll in 1982 by Scott Johnson. S. J. Edmonds has identified these as Achaetobonellia maculata Fisher, previously recorded from Onotaa in the Gilbert Islands. Also a complete specimen member of the family Echiuridae was collected in the inter- tidal area of a shallow channel beneath rocks. The latter is tentatively determined as a member of the genus Anelassorhynchus. Additional material is required before a final determination is possible (S. J. Edmonds, personal communication). REFERENCES Cutler, E. B., and N. J. Cutler, 1981, A Reconsideration of Sipuncula named by I. Ikeda and H. Sato, Publ. Seto. Mar. Bio. Lab., 26: 51-93. __, 1982, Revision of the genus Siphonosoma, Proc. Biol. Soc. Wash., 95: 748-762. Edmonds, S. J., 1980, A Revision of the Systematics of Aus- tralian Sipunculans (Sipuncula), Rec. S. Aust. Mus., 18: 1-74. Fischer, W., 1928, Uber . zwei neue Siphonosoma—Arten der Wurtemburgischen naturalien Sammlung zu Stuttgart, Zool. Anz., 76: 138-143. Fisher, W. K., 1950, The New Subgenera and a New Species of Siphonosoma (Sipunculoidea), Ann. Mag. Nat. Hist., Ser. 12, 3: 805-808. Highsmith, R. C., 1981, Coral Bioerosion at Enewetak: Agents and Dynamics, Int. Revue Gesamen. Hydrobiol., 66: 335-375. Ikeda, I., 1924, Further Notes of the Gephyrea of Japan with Descriptions of Some New Species from the Marshall, Caro- line, and Palau Islands, Jap. J. Zool., 1: 23-44. Stephen, A. C., and S. J. Edmonds, 1972, The Phyla Sipuncula and Echiura, Trustees Brit. Mus. (Nat. Hist.), London. Suchanek, T. H., and P. L. Colin, 1986, Rates and Effects of Bioturbation by Invertebrates and Fishes at Enewetak and Bikini Atolls, Bull. Mar. Sci., 33: 25-34. Chapter 12 Platyhelminthes, Nemertea, and Nematoda of Enewetak Atoll DENNIS M. DEVANEY (deceased) Bernice P. Bishop Museum Honolulu, Hawaii 96817 INTRODUCTION Except for some reports on parasitic nematodes and platyhelminthes in oysters, fish, and porpoises, little has been written about the platyhelminthes, nemertina, and nematoda of Enewetak Atoll. Little has been done on free- living nemertines or nematodes, although both groups must be abundant. The checklist in Table 1 gives the taxo- nomics of these species. TABLE 1 Checklist of Platyhelminthes, Nemertea, and Nematoda of Enewetak Atoll Phylum PLATYHELMINTHES Class CESTODA Subclass EUCESTODA Order TETRAPHYLLIDEA Family PHYLLOBOTHRIIDAE Anthobothrium sp.: Alexander, 1971. Crossobothrium spp. (8): Alexander, 1971. Family ONCHOBOTHRIIDAE Platybothrium spp. (3): Alexander, 1971. Phoreibothrium spp. (2): Alexander, 1971. Order TRYPANORHYNCHIDEA (8 undetermined spp.): Alexander, 1971. Order LECANICEPHALIDEA Family DISCOCEPHALIDAE Disculiceps spp. (2): Alexander, 1971. Family CEPHALOBOTHRIIDAE Tylocephalum sp.: Rifkin, 1970. Class TREMETODA Order DIGENEA Suborder PROSTOMATA Family FELLODISTOMIDAE Paradiscogaster eniwetokensis Martin 95 TABLE 1 (cont'd) Class TURBELLARIA Order POLYCLADIDA Suborder ACOTYLEA Family LEPTOPLANIDAE Stylochoplana (?) minuta Hyman: Hyman, 1959. Family PLANOCERIDAE Paraplanocera fritillata Hyman: Hyman, 1959. Suborder COTYLEA Family PROSTHIOSTOMIDAE Prosthiostomum exiguum Hyman: Hyman, 1959. Prosthiostomum griseum Hyman: Hyman, 1959. Order RHABDOCOELA Undetermined fresh water form: Maguire, 1967. Phylum NEMERTEA Class ANOPLA Order HETERONEMERTEA Baseodiscus delineatus (Delle Chiaje): Coe, 1947. Phylum NEMATODA Class SECERNENTEA (PHASMIDA) Order SPIRURIDA Family SPIRURIDAE Protospirura muricola: Jackson and Bastian, 1975. PLATYHELMINTHES (Tapeworms, Flukes, Flatworms) Bivalve mollusks were examined for the presence of lecanicephalid cestode larvae at Enewetak in June 1969. Of 12 bivalve species examined, only an oyster (Margari- tifera vulgaris) was found infected with an undetermined species of Tylocephalum together with another large larval cestode (Rifkin, 1970). Sharks are the definitive hosts of Tylocephalum, and at least four shark species at Enewetak were the reported hosts for over 20 other cestodes (Alex- ander, 1971). Specimens of another cestode were recovered from the feces of a large ray collected in the Enewetak lagoon during July 1981. A collection of worm parasites from fishes, fish-eating birds, and mammals was made in 1957 at Mid-Pacific ae DEVANEY Research Laboratory (MPRL) (Martin, 1957). As a result, one new species of trematode (fluke) was described from specimens of a butterfly fish (Martin and Hammerich, 1970). Four new species of turbellarian polyclad flatworms were reported by Hyman (1959) from Enewetak as a result of collections made by D. I. Reish in the summer of 1956 from reef flat rocks and algal thalli. An undetermined rhabdocoel was collected from a shell and coconut contain- ing fresh water found on the ground during the summer of 1963 (Maguire, 1967). NEMERTEA (Ribbon Worms) Three species of benthic nemertine worms found at Bikini and Enewetak in 1946 _ included only one—Baseodiscus delineatus, 30 cm long, 4 to 5 mm wide—from Enewetak (Coe, 1947). This specimen was collected in shallow water on the lagoon side of Bogon, (Boken) Islet. Subsequently, it was noted that nem- erteans made up 25% of the meiobenthos from interstitial sands examined at Enewetak. Two undetermined nemer- teans, collected on the reef flat beachrock near the quarry on Enewetak islet, have been deposited in the MPRL refer- ence collection. NEMATODA (Roundworms) The distribution and biological relations of a stomach nematode, Protospirura muricola, found in rats and house mice at Enewetak were reported by Jackson and Bastian (1975). Undetermined nematodes have been recovered from the gut of a porpoise and from the feces of a large ray collected at Enewetak. No study has yet been made on the free-living nematodes from the atoll’s marine or terres- trial environment. ACKNOWLEDGMENT Appreciation is extended to D. Hope for his sugges- tions. REFERENCES Alexander, C. G., 1971, Ann. Rept. EMBL 1970-71, pp. 53-55. Coe, W. R., 1947, Nemerteans of the Hawaiian and Marshall Islands, Occas. Pap. Bernice P. Bishop Museum, 19: 101-106. Hyman, L. H., 1959, A Further Study of Micronesian Polyclad Flatworms, Proc. U. S. Nat. Mus., 108: 543-597. Jackson, W. B., and R. K. Bastian, 1975, Ecological Investiga- tions of Rat Populations, Ann. Rept. EMBL 1975, pp. 57-62. Maguire, B., Jr., 1967, Small Freshwater Organisms on Eniwetok Atoll, Micronesica, 3: 151-157. Martin, W. E., 1957, Ann. Rept. EMBL 1956-57, p. 6. —_, and B. Hammerich, 1970, Paradiscogaster eniwetokenis n. sp. (Fellodistomidae: Trematoda) from Eniwetok, Marshall Islands, Pac. Sci., 24: 379-381. Rifkin, E., 1970, Ann. Rept. EMBL 1969-70, p. 5. Polychaetes of Enewetak Atoll DENNIS M. DEVANEY (deceased)* and JULIE H. BAILEY-BROCK+ *Bernice P. Bishop Museum, Honolulu, Hawaii 96817; Department of Zoology, University of Hawaii Honolulu, Hawaii 96822 POLYCHAETA The first record of polychaetes from Enewetak is based on collections made during Operation Crossroads between 1946 and 1952 (Hartman, 1954). Earlier reports on polychaetes from the Marshall Islands (Chamberlin, 1919; Gustafson, 1930) did not refer directly to Enewetak as a collection locality. Just over 70 polychaete species (20 of which could not be specifically determined) were identified by Hartman (1954). Most of the polychaetes were col- lected in shallow reef flat areas on lagoon and seaward sides of windward and leeward islands. Specimens from these collections have been deposited in the U. S. National Museum of Natural History (USNM) and Allan Hancock Foundation. Hartman suggested that many of the polychaetes had an erosive effect on the reefs because they are known to bore into coral rock. Most of the iden- tified species are represented in other parts of the Indo- West Pacific, and three new species, Dodecaceria laddi, Leaena minuta, and Sphaerodorum pacificum, were described from Enewetak (Hartman, 1954). Table 1 pro- vides a checklist of polychaetes of Enewetak Atoll. Due to the importance of polychaetes as _ bio eroders, food for fishes and invertebrates, and components of the coral reef biota, Mid-Pacific Research Laboratory contracted for further collections to be made between 1956 and 1957. As a result, a series of identified species was deposited in the reference collection at Enewetak and the USNM, and three publications added 10 new species to the atoll polychaete fauna (Reish, 1961, 1968; Woodwick, 1964). Of the 29 newly recorded species from Enewetak, 13 were cosmopolitan, eight were range exten- sions from the Indian Ocean and Red Sea, and five were from nontropical areas (Reish, 1968). This latter reference is valuable in enumerating and in giving keys to species within 29 families of polychaetes known from the Marshall Islands at that time. 97 Chapter 13 Two ectocommensal polychaetes have been recorded as symbionts from echinoderms at Enewetak. Hartman (1954) reported Gastrolepidia clavigera associated with the sea cucumbers Stichopus horrens at Bikini, Enewetak, and Rongelap and with Holothuria gyrifer at Enewetak. Two additional holothuroids (Actinopyga mauritiana and Holothuria atra) were recorded as hosts for this polynoid by Reish (1968). Devaney (1967) noted Hololepidella nigropunctata as a symbiont with the _ brittle-star Ophiocoma anaglyptica at Enewetak. Straughan (1969) made the first identifications of spirorbid tube worms from Enewetak. Two species were recognized. One, erroneously described as a new species, is the sinistrally coiled species Vinearia koehleri [as Spirorbis (Pileolaria) polyoperculatus Straughan, (1969)]; the other is a dextral species, Neodexiospira brasiliensis [as Spirorbis (Circeis) bellulus Bush]. Young (1969) recorded a nudibranch that preyed on the above two species and on a serpulid. Leviten (1976, 1978) recorded polychaetes from the families Eunicidae and Nereididae that served as prey for several conid gastropods on the seaward reef bench at Enewetak Islet and elsewhere in the Indo-West Pacific. Kohn (1981), in his continuing ecological studies on Conus, reported representatives of eight polychaete fami- lies that are prey for several of these gastropods. Some of the identified polychaetes are apparently the first records of these species from Enewetak where they occurred mainly in the sandy lagoon among algae (Halimeda spp.). An understanding of the habitat preferences and diver- sity of reef flat polychaetes along the seaward side of Enewetak Islet was greatly enhanced through a study con- ducted in 1976 and 1978 under laboratory support by Bailey-Brock et al. (1980). As a result of this work, it was determined that 44 polychaete taxa included 18 identified species that were new Enewetak records, with representa- tives of two additional families (Paraonidae and Scalibreg- midae). Over 20 of the 31 syllids had been previously unrecorded, but 12 have yet to be assigned specific names. Polychaetes representing 26 families were abundant infaunal components associated with selected Enewetak corals and coral rubble (Highsmith, 1981). The poly- chaetes were the most common invertebrates found in the coral heads, with over 80% of all individuals represented 98 DEVANEY AND BAILEY-BROCK TABLE 1 Checklist of Polychaetes of Enewetak Atoll* ANNELIDA Class POLYCHAETA Family POLYNOIDAE Gastrolepidia clavigera Schmarda, 1861: Hartman, 1954. Gastrolepidia claverigera [sic] Schmarda: Reish, 1968. Harmothoe (?) imbricata (Linnaeus, 1767): Hartman, 1954; Reish, 1968. Hololepidella nigropunctata (Horst, 1915): Devaney, 1967. Iphione muricata Savigny, 1818: Reish, 1968. Paralepidonotus ampulliferus (Grube, 1878). Paralepidonotus ampullifera Hartman, 1954; Reish, 1968. Family PALMYRIDAE Palmyra aurifera Savigny, 1818: Hartman, 1954; Reish, 1968. Family CHRYSOPETALIDAE Bhawania goodei Webster, 1884: Reish, 1968. Bhawania cryptocephala Gravier, 1901: Hartman, 1954. Chrysopetalum ehlersi Gravier, 1902: Hartman, 1954; Reish, 1968. Palaeonotus debilis (Grube, 1855). Palaeonotus debilis [sic]: Bailey-Brock et al., 1980. Family AMPHINOMIDAE Eurythoe complanata (Pallas, 1766): Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980. Eurythoe pacifica Kinberg, 1857: Pomeroy and Kuenzler, 1967. Pseudeurythoe oculifera (Augener, 1913): Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980. Family EODPHROSINIDAE Euphrosine myrtosa Savigny, 1818: Hartman, 1954; Reish, 1968. Family PHYLLODOCIDAE Anaitides madeirensis (Langerhans, 1880): Hartman, 1954; Reish, 1968. Eulalia viridis (Linnaeus,' 1767): Bailey-Brock et al., 1980. Eulalia tenax Grube, 1878: Hartman, 1954; Reish, 1968. Eumida sanguinea (Oersted, 1843): Reish, 1968. Genetyllis gracilis (Kinberg, 1866): Hartman, 1954; Reish, 1968. Phyllodoce marquesensis ? Monro, 1939: Hartman, 1954; Reish, 1968. Phyllodoce pruvoti Fauvel, 1930: Hartman, 1954; Reish, 1968. Family HESIONIDAE Hesione genetta Grube, 1866: Hartman, 1954; Reish, 1968. Leocrates chinensis Kinberg, 1866: Hartman, 1954; Reish, 1968. Podarke augustifrons (Grube, 1878): Reish, 1968. Irma? augustifrons Hartman, 1954. Family PILARGIDAE Synelmis albini (Langerhans, 1881) in Pettibone, 1966. Ancistrosyllis rigida Fauvel, 1919: Hartman, 1954; Reish, 1968. Family SYLLIDAE Subfamily SYLLINAE Langerhansia cornuta (Rathke, 1843). Ehlersia cornuta: Bailey-Brock et al., 1980. Langerhansia sp. A: Bailey-Brock et al., 1980. Langerhansia sp. B: Bailey-Brock et al., 1980. Haplosyllis aberrans (Fauvel, 1939): Hartman, 1954; Reish, 1968. Haplosyllis spongicola (Grube, 1855): Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980. Opisthosyllis australis Augener, 1913: Bailey-Brock et al., 1980. Opistosyllis [sic] australis Reish, 1968. Opisthosyllis brunnea Langerhans, 1879: Bailey-Brock et al., 1980. Opistosyllis [sic] brunnea Reish, 1958. Opisthosyllis corallicola (Hartmann-Schréder, 1965): Bailey-Brock et al., 1980. Opithosyllis longicirrata Monro, 1939. Opistosyllis [sic] longicirrata: Reish, 1968. Syllis gracilis Grube, 1840: Hartman, 1954; Reish, 1968. *The authors and dates for species have been checked using Hartman (1959) and Day (1967). References following the colon are Enewetak records. (This table continued on next page.) POLYCHAETES 99 TABLE 1 (cont'd) Family SYLLIDAE (cont’d) Trypanosyllis zebra (Grube, 1860): Hartman, 1954; Reish, 1968. Branchiosyllis exilis (Gravier, 1900) in Westheide, 1974. Typosyllis (?) exilis: Bailey-Brock et al., 1980. Typosyllis alternata (Moore, 1908): Bailey-Brock et al., 1980. Typosyllis armillaris (Miller, 1771): Reish, 1968; Bailey-Brock et al., 1980. Typosyllis bouvieri (Gravier, 1900): Bailey-Brock et al., 1980. Typosyllis brachychaeta (Schmarda, 1861): Hartman, 1954; Reish, 1968. Typosyllis cirropunctata (Michel, 1909): Reish, 1968. Typosyllis closterobranchia (Schmarda, 1861): MPRL Collection. Typosyllis hyalina (Grube, 1863): Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980. Typosyllis prolifera (Krohn, 1852): Reish, 1968; Bailey-Brock et al., 1980. Typosyllis taprobanensis (?) Willey, 1905: Bailey-Brock et al., 1980. Typosyllis variegata (Grube, 1860): Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980. Typosyllis sp. A: Bailey-Brock et al., 1980. Typosyllis sp. B: Bailey-Brock et al., 1980. Typosyllis sp. C: Bailey-Brock et al., 1980. Typosyllis sp. D: Bailey-Brock et al., 1980. Subfamily EUSYLLINAE Odontosyllis hyalina Grube, 1878: Reish, 1968. Subfamily EXOGONINAE Brania rhopalophora (Ehlers, 1897): Bailey-Brock et al., 1980. Brania sp. A: Bailey-Brock et al., 1980. Brania sp. B: Bailey-Brock et al., 1980. Exogone verugera Claparede, 1868: Reish, 1968; Bailey-Brock et al., 1980. Exogone sp. A: Bailey-Brock et al., 1980. Parasphaerosyllis indica Monro, 1937: Hartman, 1954; Reish, 1968. Sphaerosyllis ovigera Langerhans, 1879: Bailey-Brock et al., 1980. Sphaerosyllis pirifera Claparede, 1868: Reish, 1968. Sphaerosyllis sublaevis Ehlers, 1913: Bailey-Brock et al., 1980. Sphaerosyllis sp.: Hartman, 1954. Sphaerosyllis sp. A: Bailey-Brock et al., 1980. Subfamily AUTOLYTINAE Autolytus sp.: Bailey-Brock et al., 1980. Subfamily EURYSYLLINAE Eurysyllis pacificus (Hartman, 1954): Hartman, 1965. Sphaerodorum pacificum Hartman, 1954; Reish, 1965. Undetermined SYLLIDAE Undet. sp. A: Bailey-Brock et al., 1980. Undet. sp. Y: Bailey-Brock et al., 1980. Family NEREIDIDAE Websterinereis foli (Fauvel, 1930): Pettibone, 1971. Ceratocephala corallicola Reish, 1968: Reish, 1968. Ceratonereis tentaculata Kinberg, 1866; Perkins, 1980. Ceratonereis mirabilis Kinberg, 1866; Hartman, 1954; Reish, 1968; Leviten, 1976, 1978; Bailey-Brock et al., 1980; Kohn, 1981. Micronereis eniwetokensis Reish, 1961; Reish, 1961, 1968; Bailey-Brock et al., 1980; Paxton, 1983. Neanthes acuminata (Ehlers, 1868). Neanthes arenaceodentata (Moore, 1903): Reish, 1968. Neanthes dawydoui (Fauvel, 1937): Hartman, 1954; Reish, 1968. Nereis persica Fauvel, 1911. Nereis zonata persica Fauvel, 1911: Reish, 1968. Nereis zonata perisca [sic] Bailey-Brock et al., 1980. Nereis sp. A: Bailey-Brock et al., 1980. Nereis (?) sp.: Bailey-Brock et al., 1980. Perinereis cultrifera (Grube, 1840). Perinereis cultifera [sic] Grube: Reish, 1968. Perinereis helleri (Grube, 1878): Reish, 1968. Perinereis nigropunctata (Horst, 1889): Reish, 1968. (This table continued on next page.) 100 DEVANEY AND BAILEY-BROCK TABLE 1 (cont’d) Family NEREIDIDAE (cont’d) Platynereis dumerilii (Audouin and Milne Edwards, 1833): Kohn, 1981. Platynereis polyscalma Chamberlin, 1919: Reish, 1968. Platynereis pulchella Gravier, 1901: Reish, 1968; Bailey-Brock et al., 1980. Pseudonereis anomala Gravier, 1901: Hartman, 1954; Reish, 1968. Pseudonereis variegata (Grube, 1857): Bailey-Brock et al., 1980. Pseudonereis gallapagensis Kinberg, 1866: Hartman, 1954; Reish, 1968. Family NEPHTYIDAE Micronephtys sphaerocirrata (Wesenberg-Lund, 1949): Reish, 1968. Family SPHAERODORIDAE Sphaerodoridium capense (Day, 1963). Sphaerodoropsis capense Day: Bailey-Brock et al., 1980. Family GLYCERIDAE Glycera tesselata Grube, 1863: Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980; Kohn, 1981. Glycera sp.: Hartman, 1954. Family EUNICIDAE Eunice afra Peters, 1854: Hartman, 1954; Reish, 1968; Leviten, 1976, 1978; Bailey-Brock et al., 1980; Highsmith, 1981. Eunice antennata (Savigny, 1820): Hartman, 1954; Reish 1968; Highsmith, 1981. Eunice australis Quatrefages, 1865: Reish, 1968; Highsmith, 1981. Eunice (Nicidion) gracilis Crossland, 1904: Reish, 1968. Eunice vittata (Delle Chiaje, 1828): Kohn, 1981; Highsmith, 1981. Lysidice ninetta Audouin and Milne Edwards, 1833. Lysidice collaris Grube, 1868: Hartinan, 1954; Reish, 1968; Leviten, 1976, 1978; Bailey-Brock et al., 1980; Highsmith, 1981. Nematonereis unicornis (Grube, 1840): Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980; Highsmith, 1981. Palola siciliensis (Grube, 1840): Hartman, 1954; Reish, 1968; Leviten, 1976, 1978; Bailey-Brock et al., 1980; Kohn, 1981. Eunice siciliensis Grube: Highsmith, 1981. Paramarphysa orientalis Willey, 1905: Hartman, 1954; Reish, 1968. Family LUMBRINERIDAE Lumbrineris latreilli Audouin and Milne Edwards, 1834: Bailey-Brock et al., 1980. Lumbrineris sphaerocephala (Schmarda, 1861): Hartman, 1954; Reish, 1968. Family ARABELLIDAE Arabella iricolor (Montagu, 1804): Hartman, 1954; Reish, 1968. Arabella mutans (Chamberlin, 1919): Hartman, 1954; Reish, 1968; Kohn, 1981. Drilonereis sp. cf. D. major Crossland, 1924: MPRL Collection. Family LYSARETIDAE Oenone fulgida Savigny, 1818: Bailey-Brock et al., 1980; Highsmith, 1981. Aglaurides fulgida (Savigny): Hartman, 1954; Reish, 1968. Family DORVILLEIDAE Dorvillea pseudorubrovittata Berkeley, 1927: Highsmith, 1981. Dorvillea similis Crossland, 1924: Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980. Papilliodorvillea gardineri (Crossland, 1924): Reish, 1968. Dorvillea gardineri (Crossland): Hartman, 1954; Highsmith, 1981. Family ORBINIIDAE Naineris sp.: Hartman, 1954; Reish, 1968. Scoloplos sp.: Bailey-Brock et al., 1980. Family PARAONIDAE Levinsenia gracilis (Tauber, 1879); Hartley, 1981. Paraonis gracilis: Bailey-Brock et al., 1980. Family SPIONIDAE Laonice cirrata (?) (Sars, 1851): Hartman, 1954. Malacoceros (Malacoceros) indicus (Fauvel, 1928): Reish, 1968. tMicrospio microcera (Dorsey, 1977): Bailey-Brock et al., 1980. Polydora armata Langerhans, 1880: Woodwick, 1964; Reish, 1968; Bailey-Brock et al., 1980. Polydora cf. P. armata Langerhans: Hartman, 1954. Polydora tridenticulata Woodwick, 1964: Woodwick, 1964; Reish, 1968. ‘The authors and dates for species have been checked using Hartman (1959) and Day (1967). References following the colon are Enewetak records. tNeeds re-confirmation, possibly referrable to Microspio granulata Blake and Kudenov, 1978 (Linda Ward, personal communication). (This table continued on next page.) POLYCHAETES TABLE 1 (cont'd) Family SPIONIDAE (cont'd) Minuspio cirrifera (Wirén, 1883); Foster, 1971. Prionospio cirrifera Wirén, 1883: Reish, 1968. Pseudopolydora antennata (Claparede, 1870): Woodwick, 1964; Reish, 1968; Bailey-Brock et al., 1980. Pseudopolydora corallicola Woodwick, 1964: Woodwick, 1964; Reish, 1968. Pseudopolydora pigmentata Woodwick, 1964: Woodwick, 1964; Reish, 1968. Caraziella reishi (Woodwick, 1964): Blake, 1979. Pseudopolydora reishi: Woodwick, 1964; Reish, 1968; Bailey-Brock et al., 1980. Scolelepis (Scolelepis) bonnieri (Mesnil, 1896): Reish, 1968. Spio sp.: Hartman, 1954. Tripolydora spinosa Woodwick, 1964: Woodwick, 1964; Reish, 1968; Blake and Woodwick, 1981. Family CHAETOPTERIDAE Chaetopterus sp.: Reish, 1968. Mesochaetopterus sagittarius (Claparede, 1870): Bailey-Brock et al., 1980. Mesochaetopterus minutus Potts, 1914: Hartman, 1954; Reish, 1968; Kohn, 1981. Phyllochaetopterus ramosus Willey, 1905: Reish, 1968. Phyllochaetopterus racemosus: Bailey-Brock et al., 1980. Phyllochaetopterus socialis Claparede, 1870: Reish, 1968. Phyllochaetopterus pictus Crossland, 1903: Hartman, 1954. Phyllochaetopterus sp. cf. P. arabicus Grube, 1870: Kohn, 1981. Family SCALIBREGMIDAE Hyboscolex longiseta Schmarda, 1861: Bailey-Brock et al., 1980. Family MAGELONIDAE Magelona sp.: Reish, 1968. Family CIRRATULIDAE Cirratulus sp.: Hartman, 1954; Reish, 1968. Cirriformia punctata (Grube, 1859): Kohn, 1981. Cirriformia semicincta (Ehlers, 1905): Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980. Dodecaceria laddi Hartman, 1954: Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980. Family CTENODRILIDAE Ctenodrilus serratus (Schmidt, 1857): Reish, 1968. Family OPHELIIDAE Armandia intermedia Fauvel, 1901. Armandia lanceolata Willey, 1905: Reish, 1968. Polyophthalmus pictus (Dujardin, 1839): Hartman, 1954; Reish, 1968; Bailey-Brock et al., 1980. Family CAPITELLIDAE Capitella capitata (Fabricius, 1780): Reish, 1968; Bailey-Brock et al., 1980. Dasybranchus caducus (Grube, 1846): Hartman, 1954; Reish, 1968; Kohn, 1981. Dasybranchus lumbricoides Grube, 1878: Hartman, 1954; Reish, 1968. Dasybranchus sp.: Bailey-Brock et al., 1980. Decamastus sp.: Bailey-Brock et al., 1980. Heteromastides bifidus Augener, 1914: Reish, 1968; Bailey-Brock et al., 1980. Leiochrides sp.: Bailey-Brock et al., 1980. Notomastus latericeus Sars, 1851: Bailey-Brock et al., 1980. Pulliella sp.: Bailey-Brock et al., 1980. capitellid sp.: Bailey-Brock et al., 1980. Family ARENICOLIDAE Branchiomaldane vincentii Langerhans, 1881: Hartman, 1954; Reish, 1968. Family MALDANIDAE Axiothella australis Augener, 1914: Reish, 1968. Euclymene sp. maldanid sp.: Bailey-Brock et al., 1980. Family TEREBELLIDAE Eupolymnia trigonostoma (Schmarda, 1861). Eupolymnia trigostoma [sic] (Schmarda): Reish, 1968. Leaena minuta Hartman, 1954: Hartman, 1954: Hartman, 1954; Bailey-Brock et al., 1980. Terebella ehrenbergi Grube, 1870: Reish, 1968. Terebella ehrenbergi (?) Grube: Hartman, 1954. i01 (This table continued on next page.) 102 DEVANEY AND BAILEY-BROCK TABLE 1 (cont'd) Family TEREBELLIDAE (cont’d) Thelepus sp.: Reish, 1968. terebellid sp.: Bailey-Brock et al., 1980. Family SABELLIDAE Euchone eniwetokensis Reish, 1968: Reish, 1968. Notaulax phaeotaenia (Schmarda, 1861). Hypsicomus phaeotaenia [sic]: Hartman, 1954; Reish, 1968. Megalomma trioculatum Reish, 1968: Reish, 1968. Sabella notata Grube, 1878: Hartman, 1954; Reish, 1968. sabellid sp.: Bailey-Brock et al., 1980. Family SERPULIDAE Hydroides albiceps (Grube, 1870). Eupomatus albiceps Grube: Hartman, 1954; Reish, 1968. Pomatostegus stellatus (Abildgaard, 1789): Bailey-Brock unpubl. Filograna implexa Berkeley, 1835. Salmacina dysteri (Huxley, 1855): Young, 1969. Salmacina incrustans Clarapede, 1870: Reish, 1968. Salmacina sp.: Hartman, 1954. Serpula hartmanae Reish, 1968: Reish, 1968. Serpula sp.: Hartman, 1954. Spirobranchus giganteus (Pallas, 1766): Hartman, 1954; Reish, 1968. Vermiliopsis glandigerus Gravier, 1906: Reish, 1968; Bailey-Brock et al., 1980. Vermiliopsis sp.: Hartman, 1954. serpulid sp.: Bailey-Brock et al., 1980. Family SPIRORBIDAE Neodexiospira brasiliensis (Grube, 1872); Knight-Jones, 1984. Spirorbis bellulus Bush [sic]: Young, 1969. Vinearia koehleri (Caullery and Mesnil, 1897); Knight-Jones, 1984. Spirorbis (Pileolaria) n. sp.: Young, 1969. spirorbids, dextral: Hartman, Reish, 1968. spirorbids, sinistral: Hartman, 1954; Reish, 1968. spirorbid sp.: Bailey-Brock et al., 1980. by this group. Highsmith indicated that most of the polychaetes found in association with the corals were “nestlers” rather than bioeroders (borers). Eunicids and syl- lids were most numerous among the polychaete families, with the former having a far greater biomass. Polychaetes, together with sipunculans, were found to be the most com- mon invertebrates associated with rubble in Highsmith’s study. Six identified eunicids, two dorvilleids, including one new Enewetak record and one lysaretid, were among the species enumerated, together with undetermined members from other families. The family Aphroditidae was recorded for the first time from Enewetak in that paper, but no gen- era or species was listed. A great deal of work on the annelids of Enewetak remains to be done. This is espe- cially true of the soft bottom in the lagoon where this fauna has yet to be investigated to any extent. One hundred thirty-two polychaete species have been identified from Enewetak Atoll. Of them, 52 species (40%) are also found in the high Hawaiian Islands (Bailey-Brock and Hartman, 1984) and 43 (33%) are known from the Solomon Islands (Gibbs, 1971). Twenty-seven Enewetak species occur on the shallow coral reefs of Sumatra and Thailand, and 20 are from Easter Island, representing 21 and 15%, respectively (Kohn and Lloyd, 1973a, b). Col- lections from Heron Island reefs off Australia (Reichelt, 1979) comprised mostly errant polychaetes (Eunicidae and Syllidae) and shared only 10% of the species with Enewetak. All these collections were from _ similar habitats—hard substrata of shallow coral reefs—except for Reichelt’s, which was restricted to the infauna of sand and rubble from underneath small coral boulders. These numbers suggest faunal _ similarities of Enewetak polychaetes with both the Solomon and Hawaiian Islands. The high number for Hawaii probably reflects more inten- sive sampling and research there than elsewhere in the tropical Pacific. It is expected that the similarity between Enewetak and west Pacific locations will increase as the polychaetes from those areas become better known. Among the Enewetak polychaetes, two syllid species (Haplosyllis spongicola and Exogone verugera) and two eun- icids (Nematonereis unicornis and Palola siciliensis) have been found at the Indian and Pacific Ocean locations mentioned above. Other species with broad Pacific distributions are the amphinomid Eurythoe complanata, POLYCHAETES 103 which occurs also on Guam and Australian reefs (Bailey- Brock, personal observation), Typosyllis prolifera, Ceratonereis tentaculata (C. mirabilis), Lysidice ninetta (L. collaris), and Mesochaetopterus sagittarius (M. minutus). These syllids and nereidids are typically associated with algal turf and encrusting sponges. Lysidice ninetta and P. siciliensis are known to bore in coral, and the tube builder, M. sagittarius, forms dense aggregations in sand patches on intertidal reefs and benches. Many species, presently only identified to genus, may eventually further relate the Enewetak polychaete fauna to that occurring on other coral islands in the Pacific. Little is known about the interstitial annelids at Enewetak. However, nearly 50% of the meiobenthic fauna from sands around the atoll was found to be composed of polychaetes by MacCoy (1967). Further research is neces- sary to define this fauna systematically. MYZOSTOMATIDA At least two, as yet undetermined, myzostomatid species have been found on comatulid crinoids collected at Enewetak (Devaney, personal observation). REFERENCES Bailey-Brock, J. H., and O. Hartman, 1985, Class Polychaeta, Reef and Shore Fauna of Hawaii, D. M. Devaney and L. G. Eldredge (Eds.), Revised Edition, Bernice P. Bishop Mus. Spec. Publ., 64 (2-3) (in press). __, J. K. White, and L. A. Ward, 1980, Effects of Algal Turf and Depressions as Refuges on Polychaete Assemblages of a Windward Reef Bench at Enewetak Atoll, Micronesica, 16: 43-58. Blake, J. A., 1979, Four New Species of Caraziella (Polychaeta: Spionidae) from North and South America, with a Redescrip- tion of Two Previously Described Forms, Proc. Biol. Soc. Wash., 92: 466-481. , and K. H. Woodwick, 1981, The Morphology of Tripolydora spinosa Woodwick (Polychaeta: Spinoidae): An Application of the Scanning Electron Microscope to Polychaete Systematics, Proc. Biol. Soc. Wash., 94: 352-362. Chamberlin, R. V., 1919, The Annelida Polychaeta, Mem. Mus. Comp. Zool. Harvard, 48: 1-514. Day, J. H., 1967, A Monograph of the Polychaeta of Southern Africa: Part 1, Errantia, pp.1-458; Part 2. Sedentaria, pp. 459-878, Trustees British Museum (National History) London. Devaney, D. M., 1967, An Ectocommensal Polynoid Associated with Indo-Pacific Echinoderms, Primarily Ophiuroids, Occas. Pap. Bernice P. Bishop Mus., 23: 287-304. Foster, N. M., 1971, Spionidae of the Gulf of Mexico and Carib- "bean, Stud. Fauna Curacao Caribb. Islands, 36: 1-183. Gibbs, P. E., 1971, The Polychaete Fauna of the Solomon Islands, Bull. Br. Mus. (Nat. Hist.) Zool., 21: 101-211. Gustafson, G., 1930, Anatomische Studien uber die Polychaten- familien Amphinomidae und Euphrosynidae, Zool. Bidrag Uppsala, 12: 305-471. Hartley, J. P., 1981, The Family Paraonidae (Polychaeta) in Brit- ish Waters: A New Species and New Records with a Key to Species, J. Mar. Biol. Assn. U.K., 61: 133-149. Hartman, O., 1954, Marine Annelids from the Northern Marshall Islands, Geol. Sur. Prof. Pap., 260-Q, pp. 619-644. __, 1959, Catalogue of the Polychaetous Annelids of the World, Parts I and Il, Allan Hancock Found. Occ. Pap., No. 23, pp. 1-628. Highsmith, R. C., 1981, Coral Bioerosion at Enewetak: Agents and Dynamics, Int. Revue Gesamen. Hydrobiol., 66: 335-375. Knight-Jones, P., 1984, A New Species of Protolaeodora (Spiror- bidae: Polychaeta) from Eastern U.S.S.R., with a Brief Revi- sion of Related Genera, Zool. J. Linn. Soc., 80: 109-120. Kohn, A. J., 1981, Abundance, Diversity, and Resource Use in an Assemblage of Conus Species in Enewetak Lagoon, Pac. Sci., 34: 359-369. Kohn, A. J., and M. C. Lloyd, 1973a, Polychaetes of Truncated Reef Limestone Substrates on Eastern Indian Ocean Coral Reefs: Diversity, Abundance, and Taxonomy, Int. Revue Gesa- men. Hydrobiol., 58: 369-400. , 1973b, Marine Polychaete Annelids of Easter Island, Int. Revue Gesamen. Hydrobiol., 58: 691-712. Leviten, P. J., 1976, The Foraging Strategy of Vermivorous Conid Gastropods, Ecol. Monogr., 46: 157-178. ___, 1978, Resource Partitioning by Predatory Gastropods of the Genus Conus on Subtidal Indo-Pacific Coral Reefs: The Signif- icance of Prey Size, Ecology, 59: 614-631. MacCoy, C. V., 1967, Eniwetok Mar. Biol. Lab. Ann. Rept. FY 1967-68, pp. 11-13. Paxton, H., 1983, Revision of the Genus Micronereis (Polychaeta: Nereididae: Notophycinae), Rec. Aust. Mus., 35: 1-18. Perkins, T. H., 1980, Review of Species Previously Referred to Ceratonereis mirabilis, and Descriptions of New Species of Ceratonereis, Nephtys, and Goniada (Polychaeta), Proc. Biol. Soc. Wash., 93: 1-49. __, 1984, Revision of Demonax Kinberg, Hypsicomus Grube, and Notaulax Tauber, with a Review of Megalomma Johans- son from Florida (Polychaeta: Sabellidae), Proc. Biol. Soc. Wash., 97: 285-368. Pettibone, M. H., 1966, Revision of the Pilargidae (Annelida: Polychaeta), Including Descriptions of New Species and Redescription of the Pelagic Podarmus ploa Chamberlin (Polynoidae), Proc. U. S. Nat. Mus., 118: 155-208. __, 1971, Revision of Some Species Referred to Leptonereis, Nicon, and Laeonereis (Polychaeta: Nereididae), Smithson. Contr. Zool., 104: 1-53. Pomeroy, L. R., and E. J. Kuenzler, 1967, Phosphorus Turnover by Coral Reef Animals, Symposium on _ Radioecology, D. J. Nelson and F. C. Evans (Eds.), pp. 474-482, Univer- sity of Michigan Press, Ann Arbor, Mich. Reichelt, R., 1979, Infaunal Polychaetes of Reef Crest Habitats at Heron Island, Great Barrier Reef, Micronesica, 15: 297-307. Reish, D. J., 1961, A New Species of Micronereis (Annelida, Polychaeta) from the Marshall Islands, Pac. Sci., 15: 273-277. ___, 1968, The Polychaetous Annelids of the Marshall Islands, Pac. Sci., 22: 208-231. Straughan, D., 1969, Spirorbinae (Annelida: Polychaeta) from Eniwetok, Marshall Islands, Micronesica, 5: 151-153. Westheide, W., 1974, Interstitielle Fauna von Galapagos XI. Pisionidae, Hesionidae, Pilargidae, Syllidae (Polychaeta), Microfauna des Meeresbodens (Akad. Wiss. Lit. Mainz), 44: 1-146. Woodwick, K. H., 1964, Polydora and Related Genera (Annelida, Polychaeta) from Eniwetok, Majuro, and Bikini Atolls, Marshall Islands, Pac. Sci., 18: 146-159. Young, D. K., 1969, Okadaia elegans, a Tube-Boring Nudibranch Mollusc from the Central and West Pacific, Am. Zool., 9: 903-907. Mollusca of Enewetak Atoll E. ALISON KAY and SCOTT JOHNSON University of Hawaii at Manoa, Honolulu, Hawaii 96822 INTRODUCTION The mollusks of the Marshall Islands were first studied by Demond (1957)* for a summary of the ecology and zoogeography of central Pacific mollusks. Her summary was based on collections made between 1944 and 1957 in the Marshall, Caroline, Gilbert, and Mariana Islands. Although Demond’s paper does not specifically cite species from Enewetak, her work was derived in part from study of collections made in 1946 and 1947 at Enewetak by the late J. P. E. Morrison of the National Museum of Natural History during Operation Crossroads and from collections made in 1952 at Enewetak by H. S. Ladd, M. Russell, “Earlier records of the mollusks of the Marshall Islands include those of Escholtz (1826), Hoffman (1928), Hatai (1941), and Dietrich and Morris (1953). None mentions Enewetak. Chapter 14 and R. C. Townsend of the U. S. Geological Survey. These Enewetak collections are deposited in the National Museum of Natural History (USNM), Washington, D. C. Another reference collection of Enewetak mollusks was begun in 1955 when approximately 180 species collected by A. D. Hinckley were determined by Demond (Hiatt, 1956, 1957) and deposited at the Enewetak Marine Bio- logical Laboratory (EMBL). Subsequently, cephalopods col- lected and identified by S. J. Townsley (Hiatt, 1957) and other mollusks collected by EMBL visitors and by the authors were added to the EMBL collections. A checklist of the Mollusca of Enewetak Atoll is provided in Table 1. Records of the mollusks of Enewetak are found in a wide range of publications.t Monographic treatments of families and genera which record shells from Enewetak are those on Assiminea (Abbott, 1958); Strombidae (Abbott, 1960, 1961; Jung and Abbott, 1967); Vasidae (Abbott, +Abstracts from the annual EMBL (later known as MPML) reports are cited only if we could not trace subsequent publica- tion. TABLE 1 Annotated Checklist of Mollusks from Enewetak Atoll* Phylum MOLLUSCA Class GASTROPODA Subclass PROSOBRANCHIA Order ARCHAEOGASTROPODA Family HALIOTIDAE Haliotis clathrata Reeve, 1846; Talmadge, 1963: 132; as H. ruber clathrata Reeve. Haliotis gemma Reeve, 1846; Talmadge, 1963: 136, pl. 14, Fig. 8; as H. jacnensis Reeve. H. ovina Gmelin, 1791; [Cernohorsky, 1972: 33, pl. 2, Fig. 3). H. planata Sowerby, 1833; [Talmadge, 1963: 135, pl. 14, Fig. 2]. H. varia Linnaeus, 1758; [Cernohorsky, 1972: 34, pl. 2, Fig. 5]. “This checklist represents all of the mollusks presently known from Enewetak, both living and fossil. Specimens of species recorded as living at Enewetak are deposited in the National Museum of Natural History, Washington, D.C., and/or Bernice P. Bishop Museum, Hono- lulu, Hawaii. The following conventions are used in the checklist: (1) species found living at Enewetak are listed with a citation to a refer- ence figure in square brackets; where species have been previously recorded, the reference is also cited; (2) families are given in approxi- mate phylogenetic sequence, and within each family the genera and species are listed alphabetically. (This table continued on next page.) 106 KAY AND JOHNSON TABLE 1 (cont’d) Family SCISSURELLIDAE Scissurella concinna A. Adams, 1862; [Habe, 1951: 5, pl. 11, Figs. 10, 11]. S. coronata Watson, 1886; Ladd, 1966: 26-27, pl. 2, Figs. 7, 8; Holocene. S. declinans Watson, 1886; Ladd, 1966: 27, pl. 2, Fig. 6; Holocene. S. equatoria Hedley, 1899; Ladd, 1966: 27, pl. 2, Figs. 9, 10; Holocene. Family FISSURELLIDAE Diodora foveolata Garrett, 1872; Fig. 1A. D. granifera (Pease, 1861); Ladd, 1966: 31-32, pl. 3, Figs. 3, 4; Holocene; [Kay, 1979: Figs. 11E-F]. + D. marshallensis Ladd, 1966; Ladd, 1966: 31, pl. 3, Figs. 1, 2; late Miocene. D. ruppelli (Sowerby, 1834); [Kay, 1979: 42, Figs. 11C-D]. Emarginula bicancellata Montrouzier, 1860; Ladd, 1966: 28, pl. 2, Figs. 11, 12; Holocene. E. concinna A. Adams, 1852; Figs. 1B, 1C. TE. cf. E. clypeus A. Adams, 1851; Ladd, 1966: 28, pl. 2, Figs. 15, 16; Holocene. E. cf. E. dilecta A. Adams, 1852; Ladd, 1966: 28, pl. 2, Figs. 13, 14; as Emarginula sp. cf. peasei Thiele, 1915; Holocene. E. nigromaculata Thiele, 1915; Fig. 1D. E. souverbiana Pilsbry, 1890; Ladd, 1966: 28, pl. 2; Figs. 17, 18; Holocene. t Emarginula sp. B; Ladd, 1966: 29, pl. 2, Figs. 21, 22; Holocene. Puncturella sp. tT Hemitoma bikiniensis Ladd, 1966; Ladd, 1966: 29, pl. 2, Figs 24, 25; Holocene. TH. (Montfortia) sp. A; Ladd, 1966: 29, pl. 2, Figs. 26, 27; Holocene. Rimula mariei Crosse, 1866; Ladd, 1966: 30, pl. 2, Figs. 32, 33; as Rimula sp.; Miocene?; Holocene. + Scutus sp. A.; Ladd, 1966: 31, pl. 2, Figs. 34, 35; late Miocene. T Scutus sp. B.; Ladd, 1966: 31, pl. 2, Figs. 36, 37; late Miocene. Family PATELLIDAE Patella flexuosa (Quoy and Gaimard, 1834); Ladd, 1966: 32, pl. 3, Fig. 7; as P. stellaeformis Reeve, 1842; Holocene. Family ACMAEIDAE Patelloida sp. Family TROCHIDAE Angaria delphinus (Linnaeus, 1758); [Cernohorsky, 1972: 42, text Fig. 4]. Astele engebiensis Ladd, 1966; Ladd, 1966: 37, pl. 4, Figs. 11-13; early Miocene; Fig. 1F. Calliostoma sp; Zmarzly, 1984: 15; Fig. 1E. Clanculus atropurpureus (Gould, 1849); Zmarzly, 1984: 115; [Cernohorsky, 1978: 32, pl. 8, Fig. 3]. C. clanguloides (Wood, 1828); [Cernohorsky, 1972: 40, pl. 8, Fig. 8]. C. margaritarius Philippi, 1847; [Cernohorsky, 1972: 40, pl. 8, Fig. 9]. Euchelus angulatus Pease, 1867; Ladd, 1966: 34, pl. 3, Figs. 17-19; Holocene. E. atratus (Gmelin, 1791); [Cernohorsky, 1972: 40, pl. 8, Fig. 7]. E. foveolatus (A. Adams, 1852); [Marshall, 1979: 524, Figs. 2A—E]. E. instrictus (Gould, 1849); Ladd, 1966: 33, pl. 3, Figs. 11-13; Miocene?; Pleistocene. E. pauperculus (Lischke, 1872); [Cernohorsky, 1978: 33, pl. 8, Fig. 7]. TE. cf. E. quadricarinatus (Dillwyn, 1817); Ladd, 1966: 33, pl. 3, Figs. 9, 10; late? Miocene; Holocene. T Euchelus sp. A.; Ladd, 1966: 34, pl. 3, Figs 20-22; Holocene. +Gibbula engebiensis Ladd, 1966; Ladd, 1966: 36, pl. 4, Figs. 6, 7; early Miocene. G. marmorea (Pease, 1861); Ladd, 1966: 36, pl. 4, Figs. 8-10; as Fossarina hoffmeisteri Ladd, 1966; Miocene. Hybochelus cancellatus orientalis Pilsbry, 1904; Ladd, 1966: 34-35, pl. 3, Figs. 23, 24; Holocene. Isanda apicina (Gould, 1862); Ladd, 1966: 39-40, pl. 5, Figs. 1-4; Holocene. Monilea belcheri (Philippi, 1849); [Cernohorsky, 1972: 41, pl. 8, Fig. 11]. M. danieli Crosse 1862; [Cernohorsky, 1972: 41, pl. 8, Fig. 12; as M. philippiana Dunker, 1871). +M. marshallensis Ladd, 1966; Ladd, 1966: 40, pl. 5, Figs. 9-12; early Miocene. M. vernicosa Gould, 1861; Ladd, 1966: 40-41, pl. 5, Figs. 13, 14; as M. lifuana Fischer, 1878; Holocene; [Cernohorsky, 1978: pl. 9, Fig. 7]. Tectus fenestratus (Gmelin, 1791); [Cernohorsky, 1972: 39, pl. 6, Fig. 7]. T. hirasei Pilsbry, 1904; [Pilsbry, 1904: 32, pl. 5, Figs. 52, 52a). T. pyramis (Born, 1778); [Cernohorsky, 1972: 39, pl. 8, Fig. 4]. }Thalotia berauensis (Beets, 1941); Ladd, 1966: 34-35, pl. 3, Figs. 27, 28; late Miocene. T. gilberti (Fischer, 1878); [Cernohorsky, 1978: 34, pl. 8, Fig. 10; as Cantharidus]. Tosatrochus attenuatus (Jonas, 1844); [Ladd, 1966: 35, pl. 3, Fig. 29; as Thalotia aff. elongatus (Wood, 1828)]. Trochus histrio Reeve, 1848; Ladd, 1966: 37, pl. 4, Figs. 16-18; Holocene; [Cernohorsky, 1978: 32, pl. 8, Fig. 1). T. niloticus Linnaeus, 1767; introduced by the Japanese about 1935; [Cernohorsky, 1972: 38, pl. 8, Fig. 1]. +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) MOLLUSCA 107 TABLE 1 (cont’d) Family TROCHIDAE (cont'd) T. ochroleucus Gmelin, 1791; Ekdale, et al., 1979: Table 2; as Trochus maculatus; [Demond, 1957: 286, Fig. 2]. Turcica morrisoni Ladd, 1966; [Ladd, 1966: 36, pl. 3, Fig. 31; pl. 4, Figs. 1-5). Family STOMATELLIDAE Broderipia iridescens (Broderip, 1834); [Cernohorsky, 1978: 38, text Fig. 6]. Pseudostomatella decolorata (Gould, 1848); [Cernohorsky, 1972: 44, pl. 9, Fig. 4]. P. lyrata Pilsbry, 1890; [Hirase, 1951: pl. 64, Fig. 1). P. maculata (Quoy and Gaimard, 1834); [Cernohorsky, 1972: 43, pl. 9, Fig. 2]. Stomatella auricula (Lamarck, 1816); [Cernohorsky, 1972: 43, pl. 9, Fig. 4]. Stomatia heckliana (Crosse, 1871); [Cernohorsky, 1978: 36, pl. 9, Fig. 9]. S. cf. S. phymotis Helbling, 1779; [Cernohorsky, 1972: 43, pl. 8, Fig. 14). S. tuberculata (A. Adams, 1850); [Cernohorsky, 1978: 36, pl. 9, Fig. 8]. Synaptocochlea concinna (Gould, 1845); Ladd, 1966: 42, pl. 5, Figs. 20-23; Holocene. +S. marshallensis Ladd, 1966; Ladd, 1966: 42, pl. 5, Figs. 27, 28; late Miocene. TS. rosacea (Pease, 1867); Ladd, 1966: 42, pl. 5, Fig. 24; Holocene. Family SKENEIDAE Cyclostremiscus emery Ladd, 1966; Ladd, 1966: 80, pl. 16, Figs. 9-11; Miocene. TC. novemcarinatus Ladd, 1966; Ladd, 1966: 80, pl. 16, Figs. 12-14; early Miocene. C. striatus Kay, 1979; [Kay, 1979: 54, pl. 15A—E). t Haplocochlias sp. A.; Ladd, 1966: 76, pl. 14, Figs. 28, 29; late Miocene. Family TURBINIDAE + Astraea enewetokensis Ladd, 1966; Ladd, 1966: 43, pl. 6, Figs. 7—9; late Miocene. A. rhodostoma (Lamarck, 1822); [Cernohorsky, 1972: 47, pl. 10, Fig. 4]. + Astraea sp. A.; Ladd, 1966: 44, pl. 6, Figs. 13-15; late Miocene. T Astraea sp. C.; Ladd, 1966: 44, pl. 6, Figs. 19, 20; early Miocene; late Miocene; post-Miocene. tAstraea sp. D.; Ladd, 1966: 44, pl. 6, Figs. 21-23; Miocene. T Astraea sp. E.; Ladd, 1966: 44—45, pl. 6, Figs. 24-26; early Miocene. T Cynisca pacifica Ladd, 1966; Ladd, 1966: 49, pl. 6, Figs. 5-7; early Miocene. + Leptothyra candida (Pease, 1861); Ladd, 1966: 51, pl. 9, Figs. 7—9; Holocene. L. emenana Ladd, 1966; Ladd, 1966: 53, pl. 9, Figs. 24-26; Miocene; Pliocene. L. glareosa marshallensis Ladd, 1966; Ladd, 1966: 52, pl. 9, Figs. 16-20; Miocene; Holocene; [Cernohorsky, 1978: 39, pl. 10, Fig. 5; as Leptothyra nanina Souverbie]. TL. harlani Ladd, 1966; Ladd, 1966: 50-51, pl. 8, Figs. 14-20; Miocene. Wil. inepta (Gould, 1861); Ladd, 1966: 50, pl. 8, Figs. 14-22; Holocene. TL. cf. L. laeta (Montrouzier, 1863); Ladd, 1966: 51, pl. 9, Figs. 4-6; early Miocene. +L. maculosa (Pease, 1868); Ladd, 1966: 50, pl. 8, Figs. 8-13; post-Miocene. +L. picta (Pease, 1868); Ladd, 1966: 53, pl. 9, Figs. 21-23; Holocene. L. rubricincta (Mighels, 1845); [Kay, 1979: 58, Fig. 16B]. TL. verruca (Gould, 1845); Ladd, 1966: 51-52, pl. 9, Figs. 10-12; as L. balnearii Pilsbry, 1920; early Miocene; late Miocene; Holocene. TL. wellsi Ladd, 1966; Ladd, 1966: 52, pl. 9, Figs. 13-15; Miocene?. + Leptothyra sp. A.; Ladd, 1966: 53, pl. 9, Figs. 27-29; late Miocene. Turbo argyrostomus Linnaeus, 1758; Ladd, 1966: 48; late? Miocene; Holocene; [Cernohorsky, 1972: 45, pl. 9, Fig. 11]. T. petholatus Linnaeus, 1758; [Cernohorsky, 1972: 44, pl. 10, Fig. 1]. T. setosus Gmelin, 1791; [Cernohorsky, 1972: 45, pl. 10, Fig. 1]. Family PHASIANELLIDAE Gabrielona raunana raunana Ladd, 1966; Ladd, 1966: 54, pl. 10, Figs. 1-5; as Gabrielona raunana; Holocene; Robertson, 1973: 50-51, pl. 44, Figs. 4-6; pl. 45, Fig. 7; pl. 46, Figs. 1, 2. + Phasianella sp. (operculum only); Ladd, 1966: 53, pl. 10, Figs. 10, 11; late Miocene. Tricolia variabilis (Pease, 1861); Ladd, 1966: 54, pl. 10, Figs. 6, 7; late Miocene; Holocene. Family NERITIDAE Nerita albicilla Linnaeus, 1758; Stokes, 1966; Menge, 1973; [Cernohorsky, 1972: pl. 10, Fig. 8]. TN. insculpta Recluz, 1844; Ladd, 1966: 55, pl. 10, Figs. 15, 16; late Miocene. N. maxima Gmelin, 1791; [Maes, 1967: 107, pl. 4G]. N. plicata Linnaeus, 1758; Stokes, 1966; Menge, 1973; [Cernohorsky, 1972: pl. 11, Fig. 6]. N. polita Linnaeus, 1758; Ladd, 1966: 56, pl. 10, Figs. 17, 18; as N. aff. polita; Miocene; Stokes, 1966; [Kay, 1979: 64, Fig. 19C-E]. N. reticulata Karsten, 1789; [Cernohorsky, 1972: 52, pl. 11, Fig. 10). +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) 108 KAY AND JOHNSON TABLE 1 (cont'd) Family NERITIDAE (cont'd) + Pisulina adamsiana G. and H. Nevill, 1869; Ladd, 1977: 14, pl. 1, Figs. 1, 2; Holocene. Puperita cf. P. bensoni (Recluz, 1850); Fig. 1P, 1Q. T Smaraqdia colei Ladd, 1966; Ladd, 1966: 58-59, pl. 11, Figs. 5-7; Miocene. TS. joqjacartensis (Martin, 1916); Ladd, 1966: 58, pl. 10, Figs. 28-31; Miocene. + Smaraqdia sp. A.; Ladd, 1966: 59, pl. 11, Figs. 8, 9; late Miocene. [Nerita picea Recluz, 1841, reported by Stokes (1966) and Kay (1979) has not been confirmed]. Family NERITOPSIDAE Neritopsis radula (Linnaeus, 1758); Ladd, 1966: 55 pl. 10, Figs. 12-14; Miocene. Family PHENACOLEPADIDAE Phenacolepas granocostata (Pease, 1868); [Kay, 1979: 68, Fig. 21A]. P. scobinata (Gould, 1859); [Kay, 1979: 68, Fig. 21B]. Order MESOGASTROPODA Family LITTORINIDAE Littorina coccinea (Gmelin, 1791); Rosewater, 1970: 439, pl. 325, Figs. 11, 12. L. pintado (Wood, 1828); Rosewater, 1970: 447, pl. 325, Figs. 15, 16. L. scabra Linnaeus, 1758; Rosewater, 1970: 436, pl. 325, Figs. 8-10. L. undulata Gray, 1839; Rosewater, 1970: 298, pl. 325, Figs. 8-10. Nodilittorina millegrana (Philippi, 1848); Rosewater, 1970: 491, pl. 378, Figs. 13, 14. Peasiella tantilla (Gould, 1849); [Kay, 1979: 74, Fig. 24A]. + Tectarius rehderi Ladd, 1966; Ladd, 1966: 59, pl. 11, Figs. 11-13; Miocene. Family RISSOIDAE Alvania crystallina Garrett, 1873; Fig. 1L. A. isolata (Laseron, 1956); [Kay, 1979: 77, Fig. 27C]. Brookula sp., Fig. 1M. + Cinqula parryensis Ladd, 1966; Ladd, 1966: 61, pl. 11, Figs. 23, 24; Miocene. C. cf. C. roseocincta (Suter, 1908); Ladd, 1966: 61, pl. 11, Figs. 25, 26; Miocene; Holocene. t Merelina pisinna (Melville and Standen, 1890); Ladd, 1966: 63, pl. 12, Figs. 4, 5; Miocene. M. wanawana Kay, 1979; [Kay, 1979: 82, Figs. 28 A, B]. Parashiela beetsi Ladd, 1966; Ladd, 1966: 64, pl. 12, Figs. 8, 9; Holocene. T Pusillina jeffcoati (Ladd, 1966); Ladd, 1966: 62, pl. 11, Figs. 27-31; Miocene; Holocene; Ponder, 1985: 30. P. marmorata (Hedley, 1907); [Kay, 1979: 79, Figs. 27 E, F; as Vitricithna]. T Putilla morana Ladd, 1966; Ladd, 1966: 16, pl. 11, Figs. 16, 17; Miocene. + Rissoina abbotti Ladd, 1966; Ladd, 1966: 70, pl. 13, Figs. 19-21; Miocene. R. abrardi (Ladd, 1966); Ladd, 1966: 65, pl. 12, Figs. 15-18; as Zebina; early Miocene; Holocene; Ponder, 1985; 80. TR. ailinana Ladd, 1966; Ladd, 1966; 70-71, pl. 13, Fig. 26; Miocene. +R. alexisi Ladd, 1966; Ladd, 1966: 70, pl. 13, Figs. 13-16; Miocene. R. ambiqua (Gould, 1849); Ladd, 1966: 71, pl. 14, Figs. 23, 24; Holocene. +R. ambiqua parryensis Ladd, 1966; Ladd, 1966: 72, pl. 14, Figs. 25, 26; early Miocene. R. andamanica Weinkauff, 1885; Ladd, 1966: 67, pl. 12, Figs. 22-25; as R. mijana Ladd, 1966; early Miocene; Holocene; [Cernohorsky, 1978: 44, pl. 11, Fig. 7]. R. balteata Pease, 1869; Ladd, 1966: 69, pl. 13, Figs. 5-8; early Miocene; Holocene. +R. bikiniensis Ladd, 1966; Ladd, 1966: 69, pl. 13, Figs. 9, 10, 17, 18; Miocene; Ponder, 1985: 84. R. clathrata (A. Adams, 1853); [Cernohorsky, 1978: 47, pl. 12, Fig. 4]. +R. concinna A. Adams, 1851; Ladd, 1966: 72, pl. 13, Fig. 33; Miocene. R. ekkanana Ladd, 1966; Ladd, 1966: 71, pl. 13, Figs. 31, 32; Holocene. +R. emnanana Ladd, 1966; Ladd, 1966: 67, pl. 12, Figs. 31, 32; Miocene. R. fimbriata (Souverbie, 1872); [Cernohorsky, 1978: 45, pl. 11, Fig. 9]. R. cf. R. flexuosa Gould, 1861; Ladd, 1966: 66, pl. 12, Figs. 21, 22; Holocene. +R. herringi Ladd, 1966; Ladd, 1966: 74, Fig. 14, Figs. 13, 14; late Miocene. R. illustris Sowerby, 1894; Ladd 1966: 66, pl. 14, Figs. 5-8; as R. ephamilla Watson, 1886; early Miocene; Holocene. R. imbricata Gould, 1832; [Kay, 1979: 86, Fig. 29L; as Zebina]. TR. cf. R. indrai Beets, 1941; Ladd, 1966: 66-67, pl. 12, Figs. 23, 24; early Miocene. R. lamberti Souverbie, 1870; [Cernohorsky, 1978: 46, pl. 12: 2]. +R. lomaloana Ladd, 1966; Ladd, 1966: 72, pl. 13, Figs. 27, 28; Miocene. +R. marshallensis Ladd, 1966; Ladd, 1966: 73, pl. 13, Figs. 38-40; pl. 14, Figs. 1-4; Miocene. R. mejilana Ladd, 1966; Ladd, 1966: 67, pl. 12, Figs. 25, 26; early Miocene. tSpecies not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) MOLLUSCA TABLE 1 (cont’d) tSpecies not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). Family RISSOIDAE (cont'd) R. plicata A. Adams, 1851; Ladd, 1966: 74-75, pl. 14, Figs. 21, 22; early Miocene. R. reticulata (Sowerby, 1824); [Cernohorsky, 1978: 46, pl. 12: 1]. TR. cf. R. supracostata Garrett, 1873; Ladd, 1966: 68, pl. 12, Figs. 35, 36; early Miocene. R. tenuistriata Pease, 1868; Ladd, 1966: 68, pl. 12, Figs. 33, 34; early Miocene; Holocene. R. tornatilis Gould, 1861; Ladd, 1966: 69, pl. 13, Figs. 11, 12; as R. transenna Watson, 1886; early Miocene; Holocene; [Cernohorsky, 1978: 48, pl. 12: 8]. R. turricula Pease, 1861; Ladd, 1966: 72, pl. 13, Figs. 36, 37; Miocene; Holocene. Schwartziella triticea (Pease, 1861); [Kay, 1979: 86, Fig. 29]. + Stosicia gardnerae (Ladd, 1966); Ladd, 1966: 60, pl. 11, Figs. 14, 15; as Iravadia; Miocene; Ponder, 1985: 94. + Zebina cf. Z. cooperi (Oliver, 1915); Ladd, 1966: 65, pl. 12, Figs. 11, 12; as Zebina; Holocene. Z. killeblebana Ladd, 1966; Ladd, 1966: 65, pl. 12, Figs. 13, 14; early Miocene. Z. semiplicata (Pease, 1862); Ladd, 1966: 64, pl. 12, Fig. 10; as Z. metaltilana Ladd, 1966; early Miocene; Holocene; [Kay, 1979: 86, Fig. 29K]. Z. striatula (Gould, 1861); [Kay, 1979: 86, Fig. 29L]. Z. tridentata Michaud, 1830; [Kay, 1979: 86, Fig. 29J]. Family ASSIMINEIDAE t+ Assiminea nitida eniwetokensis Ladd, 1966; Ladd, 1966: 75, pl. 10, Fig. 8; Miocene. A. nitida marshallensis Abbott, 1958; Abbott, 1958: 256, pl. 15, Fig. 8; holotype from Japtan Island, Enewetak. Family BARLEEIDAE t Protobarleeia myersi (Ladd, 1966); Ladd, 1966: 62, pl. 11, Figs. 32, 33; as Amphithalamus (Ceratostraca?) myersi; late Miocene; Ponder, 1983: 243; [Ponder, 1983b: 243, Figs. 11E-I] Family CAECIDAE + Caecum berberense Ladd, 1972; Ladd, 1972: 22, pl. 5, Figs. 11, 12; early Miocene; Pliocene. C. cf. C. folini Kisch, 1959; Fig. 2B. C. cf. C. glabellum A. Adams, 1868; [Kay, 1979: 110, Fig. 42G]. TC. parryensis Ladd, 1972; Ladd, 1972: 23, pl. 5, Figs. 1-7; early Miocene. C. sepimentum de Folin, 1867; [Kay 1979: 109, Fig. 42B). +C. vertebrale Hedley, 1899; Ladd, 1972: 22, pl. 5, Figs. 8-10; Pleistocene; Holocene. } Elephantenellum sp. A.; Ladd, 1972: 23, pl. 5, Fig. 15; post-Miocene. t Fartulum cf. F. amputatum (Hedley, 1893); Ladd, 1972: 24, pl. 5, Figs. 16, 17; early Miocene. t Fartulum sp. A.; Ladd, 1972: 24, pl. 5, Fig. 18; post-Miocene. Micranellum schlangeri Ladd, 1972; Ladd, 1972: 23, pl. 5, Figs. 13, 14; Miocene; Holocene. Family VITRINELLIDAE Leucorhyncha caledonica Crosse, 1867; Ladd, 1966: 76, pl. 14, Figs. 30, 31; Holocene. L. crossei Tryon, 1888; Ladd, 1966: 78, pl. 14, Figs. 32, 33; Holocene. TL. stephensonii Ladd, 1966; Ladd, 1966: 76, pl. 14, Figs. 34, 35; early Miocene. Liotina cf. L. botanica (Hedley, 1914); Ladd, 1966: 46, pl. 7, Figs. 9-11; Holocene. L. loculosa (Gould, 1862); Ladd, 1966: 46, pl. 7, Figs. 12-14; Holocene. L. peronii (Kiener, 1839); [Cernohorsky, 1972: pl. 10, Fig. 7]. Liotina sp. A.; Ladd, 1966: 47, pl. 7, Figs. 15-17; early Miocene. Lophocochlias minutissimus (Pilsbry, 1921); Ladd, 1966: 77, pl. 15, Figs. 3-5; early Miocene. TL. paucicarinatus Ladd, 1966; Ladd, 1966: 77, pl. 15, Figs. 6-8; early Miocene. + Lydiphnis enewetokensis Ladd, 1966; Ladd, 1966: 79, pl. 16, Figs. 6-8; early Miocene; Pliocene. Marelepetopoma sp.; Fig. 1q. Microliotia dautzenbergi (Bavay, 1917); Fig. 1R. + Munditella parryensis Ladd, 1966; Ladd, 1966: 78, pl. 15, Figs. 12-14; late Miocene; Holocene. M. gualum (Hedley, 1899); Ladd, 1966: 78, pl. 15, Figs. 9-11; Pleistocene. Sansonia andamanica (Preston, 1908); [Iredale, 1917: 372, Fig. 8]; Fig. 1N. S. corayi (Ladd, 1966); Ladd, 1966: 63, pl. 12, Fig. 2; as Alvania (Taramellia); Holocene. TS. kenneyi (Ladd, 1966); Ladd, 1966: 63, pl. 12, Fig. 3; as Alvania (Taramellia); Miocene. Solariorbis tricarinata (Melville and Standen, 1896); Ladd, 1966: 79, pl. 16, Figs. 1-3; early Miocene. Teinostoma engebiense Ladd, 1966; Ladd, 1966: 78, pl. 15, Figs. 15-17; Holocene. T. marshallense Ladd, 1966; Ladd, 1966: 78, pl. 15, Figs. 18-20; early Miocene. T. parvulum Hedley, 1899; [Hedley, 1899: 553, Fig. 64]. Teinostoma sp.; Fig. 11, 1d. Family ORBITESTELLIDAE Orbitestella sp.; Fig. 1G, 1H. (This table continued on next page.) 110 KAY AND JOHNSON TABLE 1 (cont’d) Family OMALOGYRIDAE Omalogyra japonica (Habe, 1972); [Kay, 1979: 92, Figs. 32A—C]. Family CINGULOPSIDAE Rufodardanula sp.; Fig. 1K. Family TURRITELLIDAE t Turritella cf. T. cingulifera Sowerby, 1825; Ladd, 1972: 14, pl. 1, Fig. 1; early Eocene to early Miocene. Turritella sp. A.; Fig. 2C. Turritella sp. B.; Fig. 2D. Vermicularia sp. A.; Ladd 1972, pl. 2, Figs. 5-7; early Miocene; Fig. 2K. Family TRUNCATELLIDAE Truncatella guerinii A. and J. Villa, 1841; [Cernohorsky, 1972: 58, pl. 12, Fig. 16). Family VERMETIDAE Dendropoma gregaria Hadfield and Kay, 1972; [Kay, 1979: 103, Fig. 38). D. meroclista Hadfield and Kay, 1972; [Kay, 1979: 104, Fig. 39A]. D. platypus (Morch, 1861); [Kay, 1979: 106, Fig. 39B}. D. psarocephala Hadfield and Kay, 1972; [Kay, 1979: 106, Fig. 39D}. D. rhyssoconcha Hadfield and Kay, 1972; [Kay, 1979: 106, Fig. 39C]. Petaloconchus keenae Hadfield and Kay, 1972; [Kay. 1979: 108, Fig. 40A-B]. +P. lamellosus Ladd, 1972; Ladd, 1972: 20, pl. 3, Fig. 13; late Miocene. +P. merkana Ladd, 1972; 20, pl. 3, Figs. 4-12; early Miocene; Holocene. P. tokyoensis (Pilsbry, 1895); [Pilsbry, 1895: 61, pl. 1, Figs. 9-11]. +Serpulorbis cf. S. javanus Martin, 1879; Ladd, 1972: 20-21, pl. 3, Figs. 14, 51; late Miocene. S. variabilis Hadfield and Kay, 1972; [Kay, 1979: 108, Figs. 40 E, F]. Vermetus alii Hadfield and Kay, 1972; [Kay, 1979: 108, Figs. 40 C, D]. Family SILIQUARIDAE Stephopoma cf. S. roseum Quoy and Gaimard, 1833. Family PLANAXIDAE Planaxis sulcatus (Born, 1780); [Cernohorsky, 1972: 58, pl. 12, Fig. 17). P. zonatus A. Adams, 1851; Fig. 2A. Family CERITHIIDAE Bittium impendens (Hedley, 1899); Ladd, 1972: 30, pl. 7, Fig. 15; Holocene. B. sergentum (Jousseaume, 1930); Ladd, 1972: 30-31, pl. 7, Figs. 16-20; early Miocene; Holocene. B. zebrum (Kiener, 1841); Ladd, 1972: 31, pl. 7, Figs. 21, 22; as B. ianthinum; Holocene; Ladd, 1972: 31, pl. 4, Fig. 13; as B. eniwetokensis Ladd, 1972; early Miocene; Ladd, 1972: 32, pl. 4, Figs. 14-16; as B. toddae; late Miocene. Cerithium alveolus Hombron and Jaquinot, 1841; [Cernohorsky, 1972: 68, pl. 15, Fig. 8]. . atromarginatum Dautzenberg and Bouge, 1933; [Kay, 1979: Fig. 45 F]. . claviforme Schepman, 1901; Fig. 2L. . columna Sowerby, 1834; Ladd, 1972: 39, pl. 10, Fig. 2; Holocene. _ echinatum (Lamarck, 1822); Ladd, 1972: 37, pl. 9, Fig. 6; Holocene; as C. mutatum Sowerby, 1834; Houbrick, 1974: [Kay, 1979: 122, Fig. 45L; as C. mutatum Sowerby, 1834]. egenum Gould, 1849; Ladd, 1972: 39, pl. 9, Figs. 9, 10; Holocene. eniwetokensis (Ladd, 1972); Ladd, 1972: 37, pl. 9, Figs. 1, 2; Miocene; as Rhinoclavis. . floranensis (Ladd, 1972); Ladd, 1972: 37, pl. 9, Figs. 4, 5; Miocene; as Rhinoclavis. . cf. interstriatum Sowerby, 1855; [Kay, 1979: 122, Fig. 45R]. _ marshallensis (Ladd, 1972); Ladd, 1972: 34-35, pl. 8, Figs. 12, 13; Miocene; as Rhinoclavis. munitum Sowerby, 1855; Ladd, 1972: 39, pl. 10, Fig. 2; Holocene; as Cerithium aff. C. columna. nesioticum (Pilsbry and Vanatta, 1905); [Kay, 1979: 123, Fig. 45H]. nodulosum Bruguiere, 1792; Houbrick, 1971: 560-565; 1974: 20; [Cernohorsky, 1972, pl. 13, Fig. 8}. cf. C. planum Anton, 1839; [Cernohorsky, 1972: pl. 14, Fig. 7]. _ rostratum Sowerby, 1855; Ladd, 1972: 32, pl. 4, Fig. 17; as Colina rostrata; Pleistocene; Holocene. _ salebrosum Sowerby, 1855; Ladd, 1972: 37, pl. 9, Fig. 12; Pliocene; Holocene. . tenuifilosum Sowerby, 1866; Ladd, 1972: 38, pl. 9, Fig. 15; as C. tenellum Sowerby, 1855; Pleistocene; Holocene; [Cernohorsky, 1972: 70, pl. 14, Fig. 3]. C. trailii (Sowerby, 1855); [Cernohorsky, 1972: 71, pl. 16, Fig. 10). Clypeomorus batillariaeformis Habe and Kosuge, 1966; Ladd, 1972: 37, pl. 9, Figs. 7, 8 as Cerithium alveolus Hombron and Jaquinot, 1841; Holocene; Houbrick, 1974; as Cerithium sejunctum Iredale; [Houbrick, 1985: 51-52, Fig. 23). + ++ anaanaAnAAAAA AANA +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) MOLLUSCA 111 TABLE 1 (cont'd) Family CERITHIIDAE (cont'd) C. bifasciata bifasciata (Sowerby, 1855); Ladd, 1972: 40, pl. 9, Fig. 11; as Cerithium (Conocerithium) aff. egenum; late Miocene; [Houbrick, 1985: 24—27, Fig. 10]. C. brevis (Quoy and Gaimard, 1834); Houbrick, 1974: 21; as Cerithium morum; Houbrick, 1985: 50; [Houbrick, 1985: 44, pl. 19). C. nympha Houbrick, 1985; Houbrick, 1985: 119, Fig. 56e. +C. verbeeki (Woodward, 1879); Ladd, 1972: 40, pl. 10, Fig. 5; late Miocene. t+ Liocerithium kayae Ladd, 1972; Ladd, 1972: 40, pl. 4, Figs. 7-12; Miocene. + Plesiotrochus marshallensis Ladd, 1972; Ladd, 1972: 42, pl. 10, Figs. 13-15; Miocene. TP. pagodiformis Hedley, 1907; Ladd, 1972: 41, pl. 10, Fig. 10; post Miocene. TP. talinana Ladd, 1972; Ladd, 1972: 42, pl. 10, Figs. 11, 12; early Miocene. P. unicinctus A. Adams, 1853; Ladd, 1972: 41, pl. 10, Figs. 8, 9; as P. luteus (Gould, 1861); late Miocene; Holocene. +P. whitmorei Ladd, 1972; Ladd, 1972: 42, pl. 11, Figs. 2, 3; Miocene. + Plesiotrochus sp. A. Ladd, 1972; Ladd, 1972: 42, pl. 11, Fig. 1; Pliocene. Rhinoclavis articulata (Adams and Reeve, 1850); Houbrick, 1978: 302, pl. 1, Figs. 25-31. R. aspera (Linnaeus, 1758); Ladd, 1972: 34, pl. 8, Figs. 8, 9; late Miocene; Holocene; Houbrick, 1974: 21; 1978: 289-294, pl. 1, Figs. 9-14. R. diadema Houbrick, 1978; Houbrick, 1978: 318-320, pls. 33-35. R. fasciata (Bruguiere, 1792); Houbrick, 1974; as R. pharos; 1978: 295-302, pl. 2. +R. lowae Ladd, 1972; Ladd, 1972: 36, pl. 9, Fig. 3; Miocene. +R. powelli Ladd, 1972; Ladd, 1972: 35, pl. 8, Figs. 14, 15; late Miocene. R. sinensis (Gmelin, 1791); Ladd, 1972: 34, pl. 8, Fig. 11. R. aff. sinensis; late Miocene; Ladd, 1972: 39, pl. 10, Figs. 3, 4; as Cerithium tuberculatum (Linnaeus, 1791); Houbrick, 1974: 21; 1978: 53, pl. 1, Figs. 15-24. Royella sinon Bayle, 1880; [Houbrick, 1984: 12]. Family POTAMIDIDAE + Potamides wardi Ladd, 1972; Ladd, 1972: 25, pl. 6, Figs. 4, 5; Miocene. t Potamides sp. A. Ladd, 1972; Ladd, 1972: 25-26, pl. 6, Fig. 6; late Miocene. +Potamides sp. B. Ladd, 1972; Ladd, 1972: 26, pl. 6, Fig. 7; late Miocene. + Terebralia sulcata Born, 1778; Ladd, 1972: 27, pl. 6, Fig. 13; late Miocene. + Vicarya sp. Ladd, 1972; Ladd, 1972: 26, pl. 6, Fig. 11; early Miocene. Family MODULIDAE Modulus tectum (Gmelin, 1791); [Kay, 1979: 113, Fig. 43A). Family DIALIDAE Cerithidium asperulata (A. Adams, 1860); Ladd, 1972: 29, pl. 7, Fig. 6; as Obtortio dancei; early Miocene. Diala flammea (Pease, 1867); Ladd, 1972: pl. 7, Figs. 8-12; as Diala ludens Melvill and Standen, 1897; early Miocene; Holocene. TD. stricta Habe, 1960; Ladd, 1972: 30, pl. 7, Fig. 13; early Miocene. Finella failingi Ladd, 1972; Ladd, 1972: 28-29, Figs. 4, 5; as Obtortio; early Miocene. F. pupoides A. Adams, 1860; Ladd, 1972: 30, pl. 7, Fig. 14; as Obtortio pyrrhacme; Melvill and Standen, 1895; early Miocene. F. sulcifera (A. Adams, 1862); Ladd, 1972: 30, pl. 7, Fig. 14; as Diala; Holocene. + Finella sp. A. Ladd, 1972; Ladd, 1972: 29, pl. 7, Fig. 7; as Obtortio; early Miocene. Scalioa arenosa A. Adams, 1862; Fig. 2E. S. bella A. Adams, 1860; [Kay, 1979: 117, Fig. 441]. S. caledonica Crosse, 1870; Fig. 2F. S. glareosa A. Adams, 1862; Fig. 2G. Family LITIOPIDAE Styliferina goniochila (A. Adams, 1860); [Kay, 1979: 118, Fig. 44H]. [Placement of this species follows Ponder (1985)}. Family CERITHIOPSIDAE t Ataxocerithium eniwetokensis Ladd, 1972; Ladd, 1972: 32-33, pl. 4, Fig. 18; pl. 8, Fig. 1; Holocene. Cerithiopsis sp. cf. C. arga Kay, 1979; [Kay, 1979: 126, Fig. 46A]. Cerithiopsis sp. A.; Fig. 2Q. Cerithiopsis sp. B.; Fig. 2R. Joculator granata Kay, 1979; [Kay, 1979: 127, Fig. 46C). tJoculator sp. cf. J. ovata (Laseron, 1956); Ladd, 1972: 44, pl. 11, Fig. 12; early Miocene; Holocene. J. semipicta (Gould, 1861); Ladd, 1972: 43, pl. 11, Figs. 5, 6; late Miocene; Holocene. J. sumangi Ladd, 1972; Ladd, 1972: 44, pl. 11, Figs. 7-11; late Miocene. J. tribulationis (Hedley, 1909); Ladd, 1972: 43, pl. 11, Fig. 14; late Miocene; Holocene. J. turrigera (Watson, 1886); [Kay, 1979: 128, Figs. 46, D, E}. +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) KAY AND JOHNSON TABLE 1 (cont'd) Family CERITHIOPSIDAE (cont’d) J. uveanum (Melvill and Standen, 1896); [Kay, 1979: 128, Fig. 46H]. + Seila waluensis Ladd, 1972; Ladd, 1972: 44, pl. 11, Fig. 13; late Miocene. Family EULIMIDAE Balcis acicula (Gould, 1849); Lamberson, 1978; Fig. 3B. B. bryani (Pilsbry, 1917); [Kay, 1979: 161, Fig. 56l). B. solida (Sowerby, 1865); [Kay, 1979: 162, Fig. 56K]. B. thaanumi (Pilsbry, 1917); [Kay, 1979: 164, Fig. 56M]. } Balcis sp. A. Ladd, 1982; Ladd, 1982: 55, pl. 41, Figs. 14, 15; as Eulima; early Miocene. Hemileisotracum sp., Fig. 3C. Leiostraca metcalfei (A. Adams, 1853); [Kay, 1979: 165, Fig. 56Q]. Mucronalia nitidula Pease, 1861; [Kay, 1979: 165, Fig. 56T]. Pyramidelloides suta (Pilsbry, 1918); [Kay, 1979: 84, Fig. 29G). Stylapex sp., Fig. 3A. Stylifer linckiae Sarasin and Sarasin, 1887; [Kay, 1979: 167, Fig. 57E). Thyca crystallina (Gould, 1846); [Kay, 1979: 167, Fig. 56U). Family STROMBIDAE Lambis chiragra (Linnaeus, 1758); Abbott, 1961: 172, pl. 121, Figs. 10-12; Berg, 1974. L. crecata (Link, 1807); [Abbott, 1961: 157, pl. 121, Fig. 8]. L. lambis (Linnaeus, 1758); Abbott, 1961: 154, pl. 12, Fig. 4; Berg, 1974. L. scorpius (Linnaeus, 1758); Berg, 1974; [Abbott, 1961: 164, pl. 121, Fig. 5]. L. truncata (Humphrey, 1786); Abbott, 1961: 157, pl. 121: 9; Berg, 1974. Strombus dentatus Linnaeus, 1758; Abbott, 1960: 85, pl. 14: 23; Berg, 1974. S. erythrinus Dillwyn, 1817; Abbott, 1960: 81, pl. 20, Figs. 1-5. S. fragilis (Réding, 1798); [Abbott, 1960: 82, pl. 14, Fig. 30). S. gibberulus Linnaeus, 1758; Abbott, 1960: 143, pl. 14, Fig. 20; Berg, 1974; Ladd, 1972: 62, pl. 19, Fig. 9, pl. 20, Figs. 1-3; Holocene. . haemastoma Sowerby, 1842; Abbott, 1960: 117, pl. 17, Figs. 11, 12; Berg, 1974. . lentiginosus Linnaeus, 1758; Abbott, 1960, pl. 17, Figs. 11, 12. . luhuanus Linnaeus, 1758; Abbott, 1960: 136, pl. 14, Fig. 15; Berg, 1974; Gillary, 1974; Gillary and Gillary, 1979. . maculatus Sowerby, 1842; Abbott, 1960: 77, pl. 20, Figs. 24, 25. . micklei Ladd, 1972; Ladd, 1972: 60-61, pl. 18, Figs. 5-8; late Miocene. . microurceus (Kira, 1959); [Abbott, 1960: 71, pl. 20: 24, 25). . mutabilis Swainson, 1821; Abbott, 1960: 74, pl. 20: 15, 16; Berg, 1974; Ladd, 1972: 58-59, pl. 15, Figs. 10-15; Holocene. . sinuatus Lightfoot, 1786; [Abbott, 1960: 60, pl. 17, Fig. 8]. . taurus Reeve, 1857; Cernohorsky, 1972: 73, pl. 17, Fig. 5. . variabilis Swainson, 1820; [Abbott, 1960: 103, pl. 14, Figs. 21, 22]. . wilsoni Abbott, 1967; [Abbott, 1967: 455, pl. 328, Figs. 1-3]. TStrombus sp. C.; Ladd, 1972: 61, pl. 18, Figs. 10, 11; late Miocene. Terebellum terebellum (Linnaeus, 1758); Jung and Abbott, 1967: 453, pl. 321. Family VANIKORIDAE Vanikoro cancellata (Lamarck, 1822); Ladd, 1972: 52, pl. 13, Figs. 6-8; early Miocene, Holocene. V. queriniana (Recluz, 1843); Ladd, 1972: 52, pl. 13, Figs. 9, 10; early Miocene; Holocene. V. helicoidea (le Guillou, 1842); [Cernohorsky, 1972: 86, pl. 22, Fig. 1]. TV. cf. V. kanakarum Pilsbry, 1921; Ladd, 1972: 52, pl. 13, Fig. 11; Holocene. Family HIPPONICIDAE t Hipponix foliaceus Quoy and Gaimard, 1835; Ladd, 1972: 53, pl. 13, Figs. 15-21; Miocene. TH. revellei Ladd, 1972; Ladd, 1972: 54, pl. 14, Figs. 1-18; early Miocene; Pliocene. Sabia conica (Schumacher, 1817); Ladd, 1972: 54, pl. 13, Figs. 22-27: early Miocene; Holocene; [Kay, 1979: 179, Figs. 62F—H]. Family CAPULIDAE Cheilea equestris (Linnaeus, 1758); Ladd, 1972: 52, pl. 13, Fig. 12; Miocene. Crepidula aculeata (Gmelin, 1791); Found on pilings only; [Kay, 1979: 181, Figs. 63G, H]. Capulus danieli (Crosse, 1858); [Cernohorsky, 1972: 88, pl. 22, Fig. 5]. +Capulus sp. A.; Ladd, 1972: 55, pl. 14, Fig. 21; late Miocene. fCapulus sp. B.; Ladd, 1972: 55, pl. 14, Fig. 22: late Miocene. Family XENOPHORIDAE Xenophora cerea (Reeve, 1845); Ponder, 1983a: 23-25, Fig. 23. NHnNNHNHNN NHNN +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) MOLLUSCA 113 TABLE 1 (cont'd) Family LAMELLARIIDAE Lamellaria cf. L. berghi Deshayes, 1863; Fig. 3D. Family ERATOIDAE Proterato gemma (Bavay, 1917); (Cate, 1977: 353, Fig. 23). P. stalagmia Cate, 1975; [Cate, 1977: 347, Fig. 11). Family TRIVIIDAE Trivia hordacea Kiener, 1845; [Kay, 1979: 187, Fig. 65B). T. oryza (Lamarck, 1810); Ladd, 1977: 17, pl. 1, Figs. 10-12; Miocene and younger beds. Family CYPRAEIDAE Cypraea annulus Linnaeus, 1758; Ingram, 1947; [Burgess, 1970: 341, pl. 414]. arabica Linnaeus, 1758; Ladd, 1977: 19, pl. 2, Figs. 1, 2; Holocene; [Burgess, 1970: 215, pl. 18A]. _ argus Linnaeus, 1758; [Burgess, 1970: 253, pl. 28 B-C]. _ asellus Linnaeus, 1758; [Burgess, 1970: 118, pl. 8H]. . aurantium Gmelin, 1791; [Burgess, 1970: 118, pl. 8H). . beckii Gaskoin, 1836; [Burgess, 1970: 277, pl. 30L]. . bistrinotata (Schilder and Schilder, 1937); [Burgess, 1970: 271, pl. 30C). caputserpentis Linnaeus, 1758; Ingram, 1947; [Burgess, 1970: 177, pl. 12F]. carneola Linnaeus, 1758; Ingram, 1947; [Burgess, 1970: 201, pl. 32D}. catholicorum (Schilder and Schilder, 1938); [Burgess, 1970: pl. 13A]. caurica Linnaeus, 1758; [Burgess, 1970: 292, pl. 32D). childreni Gray, 1825; Ladd, 1977: 22, pl. 3, Figs. 11-13; Holocene; [Burgess, 1970: 262, pl. 29G]. chinensis Gmelin, 1791; [Burgess, 1970: 79, pl. 5H). . cicercula Linnaeus, 1758; [Burgess, 1970: 266, pl. 30A). clandestina Linnaeus, 1767; (Burgess, 1970: 93, pl. 7C]. . cribraria Linnaeus, 1758; [Burgess, 1970: 189, pl. 136K]. depressa Gray, 1824; Ingram, 1947; as Cypraea intermedia; [Burgess, 1970: 217, pl. 18E]. . dillwyni (Schilder, 1922); [Burgess, 1970: 273, pl. 30H]. . eglantina Duclos, 1833; [Burgess, 1970: 218, pl. 19A]. erosa Linnaeus, 1758; [Burgess, 1970: 175, pl. 12 A-D]. . fimbriata Gmelin, 1791; [Burgess, 1970: 139, pl. 9I]. globulus Linnaeus, 1758; [Burgess, 1970: 272, pl. 30D). goodalli Sowerby, 1832; [Burgess, 1970: 103, pl. 7aC]. helvola Linnaeus, 1758; Ladd, 1977: 23, pl. 5, Figs. 1-3; late Miocene; [Burgess, 1970: 154, pl. 10G]. . hirundo Linnaeus, 1758; [Burgess, 1970: 286, pl. 31F). humphreysii Gray, 1825; [Cernohorsky, 1967: 92, pl. 17: 92]. isabella Linnaeus, 1758; Ingram, 1947; [Burgess, 1970: 41, pl. 2B). cf. C. kamai (Beets, 1941); Ladd, 1977: 25, pl. 5, Figs. 13-15; early Miocene. . labrolineata Gaskoin, 1849; [Burgess, 1970: 143, pl. 9aF]. . leviathan Schilder and Schilder, 1938; [Burgess, 1970: 202, pl. 15, Fig. E, Ej). . limacina Lamarck, 1810; [Cernohorsky, 1967, pl. 13, Fig. 69]. lynx Linnaeus, 1758; [Burgess, 1970: 241, pl. 24C]. . maculifera (Schilder, 1932); [Burgess, 1970: 217, pl. 18D}. mappa Linnaeus, 1758; [Burgess, 1970: 248, pl. 26 A, C—D). . margarita (Schilder and Schilder, 1938); [Cernohorsky, 1967: 72, pl. 11: 52). . mariae (Schilder, 1927); [Burgess, 1970: 274, pl. 30I]. martini Schepman, 1907; [Burgess, 1970: 60, pl. 4D]. . mauritiana Linnaeus, 1758; [Burgess, 1970: 247, pl. 25B]. microdon Gray, 1828; [Burgess, 1970: 130, pl. 9D-E]. minoridens Melville, 1901; [Burgess, 1970: 132, pl. 9F}. moneta Linnaeus, 1758; Ingram, 1947; Menge, 1973; Renaud, 1977; [Burgess, 1970: 343, pl. 41B—H]. nucleus Linnaeus, 1758; [Burgess, 1970: 261, pl. 29H). onyx Linnaeus, 1758; [Burgess, 1970: pl. 6A). poraria Linnaeus 1758; Ingram, 1947; [Burgess, 1970: 152, pl. 10B]. . punctata Linnaeus, 1771; [Burgess, 1970: 104, pl. 7aE]. . schilderorum (Iredale, 1939); [Burgess, 1970: 190, pl. 14A]. . scurra Gmelin, 1791; [Burgess, 1970: 225, pl. 19D]. . staphylaea Linnaeus, 1758; [Burgess, 1970: 259, pl. 29E]. . Stolida Linnaeus, 1758; [Burgess, 1970: 290, pl. 31E]. QDNNAHANAHHAHAHAAADANNHNHHNAANDAANAANNNHHNHHAHHNHAAANHADA NHAHAHAHHHAHANHD +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) 114 KAY AND JOHNSON TABLE 1 (cont'd) Family CYPRAEIDAE (cont’d) . talpa Linnaeus, 1758; [Burgess, 1970: 199, pl. 14F]. . teres Gmelin, 1791; [Burgess, 1970: 109, pl. 8B). . testudinaria Linnaeus, 1758; [Burgess, 1970: 249, pl. 27A]. . tigris Linnaeus, 1758; [Burgess, 1970: 229, pl. 22A]. . ursellus Gmelin, 1791; [Burgess, 1970: 287, pl. 31). . ventriculus Lamarck, 1810; Ingram, 1947; [Burgess, 1970: 194, pl. 14D]. . vitellus Linnaeus, 1758; [Burgess, 1970: 206, pl. 17A]. Family OVULIDAE Calpurnus lacteus (Lamarck, 1810); [Cernohorsky, 1967: 107, pl. 21, Fig. 121]. C. verrucosus (Linnaeus, 1758); [Cernohorsky, 1967: 107, pl. 21, Fig. 122]. Pedicularia pacifica Pease, 1865; Ladd, 1977: 25, pl. 6, Figs. 5-7; Holocene. Phenacovolva brunneiterma Cate, 1969; [Cernohorsky, 1972: 91, pl. 23: 5]. Primovula margarita (Sowerby, 1828); [Cernohorsky, 1972: 90, pl. 23, Fig. 7]. Family ATLANTIDAE AO) Q)Ciey@)P) t Atlanta peronii Leseur, 1817; Ladd, 1977: 26; Holocene; [Ladd, 1977: 26, pl. 6, Figs. 14—20). Family NATICIDAE tAmpullina berauensis (Beets, 1941); Ladd, 1977: 27, pl. 7, Figs. 5-7; early Miocene. Eunaticina papilla (Gmelin, 1791); [Cernohorsky, 1972: 102, pl. 27, Fig. 5). Natica bougei Sowerby, 1908; [Cernohorsky, 1972: 98, pl. 25, Fig. 2]. N. qualteriana Recluz, 1844; Ladd, 1977: 30, pl. 9, Figs 3-10; as Naticarius marochiensis (Gmelin, 1791); Miocene; Holocene; [Cernohorsky, 1972: 95, pl. 24, Fig. 9]. N. onca (Roding, 1798); [Cernohorsky, 1972: 98, pl. 25, Fig. 6]. N. violacea Sowerby, 1825; [Cernohorsky, 1972: 96, pl. 25, Fig. 6]. Neverita cf. N. petiveriana Recluz, 1855; Fig. 3H, 3l. Neverita sp.; Fig. 3F, 3G. Polinices albumen (Linnaeus, 1758); [Cernohorsky, 1972: 99, pl. 26, Fig. 2]. P. aurantius (Roding, 1798); [Cernohorsky, 1972: 99, pl. 25, Fig. 12]. P. flemingiana (Recluz, 1844); [Cernohorsky, 1972: 98, pl. 25, Fig. 11). P. melanostomus (Gmelin, 1791); [Cernohorsky, 1972: 101, pl. 26, Fig. 8]. P. tumidus (Swainson, 1840); Ladd, 1977: 28, pl. 8, Fig. 3; as P. mamilla (Linnaeus, 1758); late Miocene. TSinum sp. A.; Ladd, 1977: 30, pl. 9, Figs. 1, 2; Holocene. Family CASSIDIDAE Casmaria erinaceus (Linnaeus, 1758); [Abbott, 1968: 190, pl. 14, Figs. 8, 10-12]. C. ponderosa (Gmelin, 1791); [Abbott, 1968: 195, pl. 14, Figs. 1, 2]. Cassis cornuta (Linnaeus, 1758); [Abbott, 1968: 47, pl. 3, Figs. 1—4]. Phalium bisulcatum (Schubert and Wagner, 1829); [Abbott, 1968: 126, pl. 8, Fig. 13). Family CYMATIIDAE Charonia tritonis (Linnaeus, 1767); [Kay, 1979: 215, Fig. 77F]. Cymatium aquatile (Reeve, 1844); [Kay, 1979: 220, Fig. 77A]. C. clandestinum (Lamarck, 1816); [Kay, 1979: 220, Fig. 79A]. . flaveola (Réding, 1798); [Hinton, 1978: 30, Fig. 7]. . gemmatum (Reeve, 1844); [Kay, 1979: 222, Fig. 79C]. . hepaticum (Linnaeus, 1758); [Cernohorsky, 1967: 48, pl. 4, Fig. 14). . intermedium (Pease, 1869); [Kay, 1979: 220, Fig. 77C]. . lotorium (Linnaeus, 1758); [Cernohorsky, 1967: 48, pl. 4, Fig. 14]. . muricinum (Réding, 1798); [Kay, 1979: 217, Fig. 77E]. . nicobaricum (Réding, 1798); Houbrick, 1974; [Kay, 1979: 216, Fig. 77D]. . pileare (Linnaeus, 1758); [Kay, 1979: 221, Fig. 77B). . pyrum (Linnaeus, 1758); [Kay, 1979: 218, Fig. 78B]. . rubeculum (Linnaeus, 1758); [Cernohorsky, 1967: 52, pl. 4, Fig. 13]. . vespaceum (Lamarck, 1822); [Kay, 1979: 223, Fig. 79E]. Distorsio anus (Linnaeus, 1758); [Kay, 1979: 223, Fig. 79K]. D. pusilla Pease, 1861; Ladd, 1977: 35, pl. 11, Figs. 12, 13; Holocene. Gyrineum roseum (Reeve, 1844); [Cernohorsky, 1972: 117, pl. 32, Fig. 3]. {The occurrence of Cymatium cf. C. caudatum (Gmelin, 1791) cited by Ekdale et al. (1979) has not been confirmed]. GIQIQ VI ViGle!eieioi® tSpecies not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) TS + > Gs MOLLUSCA 115 TABLE 1 (cont'd) Family BURSIDAE Bursa bufonia (Gmelin, 1791); Demond, 1957: 308; Kohn and Leviten, 1976; [Cernohorsky, 1967: 42, pl. 2, Fig. 5]. B. cruentata (Sowerby, 1841); [Kay, 1979: 227, Fig. 80B}. B. granularis (Réding, 1798); [Kay, 1979: 227, Fig. 80A]. B. rhodostoma (Sowerby, 1841); [Kay, 1979: 229, Fig. 80C—D}. B. rosa (Perry, 1811); [Kay, 1979: 229, Fig. 80E-F]. B. rubeta (Linnaeus, 1758); [Cernohorsky, 1972: 118, pl. 1, Fig. 5]. Tutufa bubo (Linnaeus, 1758); [Cernohorsky, 1972: 42, pl. 1, Fig. 1). Family TONNIDAE Malea pomum (Linnaeus, 1758); [Kay, 1979: 231, Fig. 814A). Tonna perdix (Linnaeus, 1758); [Kay, 1979: 233, Fig. 81D). +Order HETEROGASTROPODA Family ARCHITECTONICIDAE + Architectonica corwini Ladd, 1972; Ladd, 1972: 18, pl. 2, Figs. 14-18; Miocene. + Climacopoma? sp. A.; Ladd, 1972: 18, pl. 2, Figs. 8-10; early Miocene. Heliacus infundibuliformis (Gmelin, 1791); [Cernohorsky, 1978: 165, pl. 58, Fig. 7]. H. variegatus (Gmelin, 1791); Ladd, 1972: 19, pl. 3, Figs. 1-3; late Miocene. Philippia oxytropis A. Adams, 1855; [Kay, 1979: 100, Fig. 36A). P. radiata (Roding, 1798); Ladd, 1972: 19, pl. 2, Figs. 19-21; late Miocene. Family TRIPHORIDAE [The arrangement follows that of Marshall (1983)}. +Bouchetriphora otsuensis (Yokoyama, 1920); Ladd, 1972: 45, pi. 11, Fig. 15; as Triphora; early Miocene. B. pallida (Pease, 1871); Ladd, 1972: 45, pl. 11, Fig. 14; as Triphora; Holocene; (Kay, 1979: 148, Fig. 51L; as Triphora]. B. quadrimaculata (Hervier, 1897); Fig. 2d. Euthymella elegans (Hinds, 1843); Ladd, 1972: 50, pl. 13, Fig. 2; as Viriola; Holocene; [Cernohorsky, 1978: 173, pl. 61, Fig. 7]. E. pagoda (Hinds, 1843); Ladd, 1972: 50, pl. 12, Fig. 21; as Viriola; Holocene. E. cf. E. regalis (Jousseaume, 1884); Fig. 2M. Inella cf. I. aemulans (Hinds, 1843); [Kay, 1979: 133, Fig. 48B). I. japonica Kuroda and Kosuge, 1963; Fig. 2H. I. cf. I. lanceolata Kosuge, 1962; [Kosuge, 1962: 120, pl. 8, Fig. 16]. +I. maharatai Beets, 1941; Ladd, 1972: 46, pl. 12, Fig. 1; early Miocene; late Miocene. I. pyramidalis (Adams and Reeve, 1850); Ladd, 1972: 45, pl. 11, Figs. 16, 17; as Triphora (Inella); Miocene. Iniforis albogranosa Kosuge, 1961; Ladd, 1972: 47, pl. 12, Figs. 3—5; as Triphora (Iniforis); Holocene; [Kosuge, 1961: 313, pl. 19, Fig. 7]. I. formosulus (Hervier, 1897); [Cernhorsky, 1978: 170, pl. 60, Fig. 4]. I. ikukoae (Kosuge, 1963); [Kosuge, 1963; 258, pl. 18, Fig. 1). I. lifuana (Hervier, 1897); [Kosuge, 1961: pl. 19, Fig. 5]. I. ofuensis (Baker and Spicer, 1935); Ladd, 1972: 47, pl. 12, Figs. 8-12; Holocene. + Mastonia intermissa (Laseron, 1958); Ladd, 1972: 49, pl. 12, Figs. 17, 18; late Miocene. M. rubra (Hinds, 1843); [Cernohorsky, 1978: 170, pl. 60, Fig. 6]. +M. squamosa (Kosuge, 1962); Ladd, 1972: 48, pl. 12, Fig. 16; Holocene. M. ustulata (Hervier, 1897); Fig. 2N. Mastoniaeformis clavata (Pease, 1871); [Kay, 1979: 133, Fig. 48B; as Iniforis aemulans Hinds). M. concors (Hinds, 1843); [Kay, 1979: 133, Fig. 48D]. Mesophora cnodax (Jousseaume, 1884); Fig. 20. M. granosa (Pease, 1871); [Marshall, 1983: 45, Figs. 19E—G]. Metaxia cf. M. albicephala Kay, 1979; [Kay, 1979: 130, Fig. 48M]. Metaxia sp.; Fig. 2P. Nanophora cf. N. atratus (Kosuge, 1962); [Kosuge, 1962: 83, pl. 9, Fig. 5]. TN. cingulifera (Pease, 1861); Ladd, 1972: 48, pl. 12, Fig. 13; as Triphora (Mastonia); Holocene. +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). +The ordinal taxon “Heterogastropoda” is utilized here as a convenience only. Kosuge (1966) proposed the suborder Heterogas- tropoda to accommodate the Triphoridae, Mathildidae, Architectonicidae, and Epitoniidae, which he regarded as distinct from the Taenioglossa and Steonglossa. Haszpruner (1984, 1985) separates the Architectonicidae and Pyramidellidae as the Allogastropoda, dis- tinct from the Architaenioglossa (the Valvatacea, Cyclophoracea, and Viviparacea); the Neotaenioglossa (rissoids, cerithiids, hipponicids, strombids, cypraeids, naticids, and tonnids); the Heteroglossa (cerithiopsids, triphorids, epitoniids and eulimids); and the Stenoglossa (the Neogastropods, including muricids, cancellarids, and the Conacea). (This table continued on next page.) 116 KAY AND JOHNSON TABLE 1 (cont’d) Family TRIPHORIDAE (cont’d) N. cf. N. leucomys (Hervier, 1897); Fig. 2I. N. triticea (Pease, 1861); [Kay, 1979: 150, Fig. 51E). Nanophora sp.; Fig. 2H. Tetraphora hervieri (Kosuge, 1962); [Cernohorsky, 1978: 172, pl. 60, Fig. 14). Triphora granulata (Adams and Reeve, 1850); [Cernohorsky, 1978: 169, pl. 59, Fig. 10]. T. taeniolata (Hervier, 1897); [Cernohorsky, 1978: 172, pl. 61, Fig. 1; as Inella (Cautotriphora) monovitta Laseron, 1958]. Viriola abbotti (Baker and Spicer, 1935); [Kay, 1979: 139, Fig. 50E). V. bayani Jousseaume, 1884; [Kay, 1979: 139, Fig. 50J). V. cancellata (Hinds, 1843); Ladd, 1972: 50, pl. 13, Fig. 1; late Miocene; [Kay, 1979: 140, Fig. 50D). V. cf. V. corrugata (Hinds, 1843); [Cernohorsky, 1978: 173, pl. 61, Fig. 6; as Viriola interfilata (Gould)}. V. incisa (Pease, 1861); Ladd, 1972: 50, pl. 15, Fig. 22; Holocene. V. intergranosa (Hervier, 1897); [Cernohorsky, 1978: 123, pl. 61, Fig. 8]. Family EPITONIIDAE Epitonium alatum (Sowerby, 1844); [Kay, 1979: 153, Fig. 54C]. . auritum (Sowerby, 1844); Fig. 3E. . irregularis (Sowerby, 1844); [Cernohorsky, 1972: 197, pl. 56, Fig. 8]. . paumotensis (Pease, 1868); [Kay, 1979: 156, Fig. 54A). . perplexum (Deshayes, 1863); Ladd, 1972: 51, pl. 13, Fig. 3; Holocene; [Kay, 1979: 156, Fig. 54B). . symmetrica (Pease, 1868). . revolutum (Hedley, 1899); [Kay, 1979: 156, Fig. 54F). . cf. E. vestalis (Hinds, 1844); [Cernohorsky, 1978: 168, pl. 59, Fig. 7]. . umbilicatum (Pease, 1869); [Kay, 1979: 157, Fig. 54K). Opalia bicarinata (Sowerby, 1844); [Cernohorsky, 1978: 168, pl. 59, Fig. 9; as Nodiscala]. mommmmmmm Family JANTHINIDAE Janthina globosa Swainson, 1822; [Kay, 1979: 158, Fig. 55A, D]. J. janthina (Linnaeus, 1758); [Kay, 1979: 158, Fig. 55B). Order NEOGASTROPODA Family MURICIDAE Aspella producta (Pease, 1861); [Kay, 1979: 235, Figs. 83C, D]. Attiliosa caledonicus (Jousseaume, 1881); [Vokes and D’Attilio, 1982: 70, Fig. 5]. Chicoreus brunneus (Link, 1807); [Cernohorsky, 1967: 120, pl. 25, Fig. 148). C. microphyllus (Lamarck, 1816); [Radwin and D’Attilio, 1976: 39, pl. 4, Fig. 7]. C. ramosus (Linnaeus, 1758); [Cernohorsky, 1967: 122, pl. 25, Fig. 152]. Favartia guamensis Emerson and D’Attilio, 1979; [Emerson and D’Attilio, 1979, 4, Figs. 11, 12]. Homalocantha anatomica (Perry, 1811); [Radwin and D’Attilio, 1976, pl. 9, Fig. 5]. Marchia bipinnata (Reeve, 1845); [Radwin and D’Attilio, 1976: pl. 9, Fig. 1]. M. laqueata (Sowerby, 1841); [Radwin and D’Attilio, 1976: 58, pl. 9, Fig. 12]. M. martinetana (Roding, 1798); [Radwin and D’ Attilio, 1976: pl. 27, Fig. 3]. M. triptera (Born, 1778); [Radwin and D’Attilio, 1976: pl. 9, Fig. 12]. Naguetia trigonula (Lamarck, 1816); [Radwin and D’Attilio, 1976, pl. 15, Fig. 12]. N. triquetra (Born, 1778); [Radwin and D’Attilio, 1976, pl. 15, Fig. 11). Phyllocoma convoluta (Broderip, 1833); [Cernohorsky, 1967, pl. 24, Fig. 14]. Phyllonotus laciniatus (Sowerby, 1841); [Radwin and D’Attilio, 1976, pl. 6, Fig. 3]. tPterynotus sp.; Ladd, 1977: 39, pl. 14, Fig. 3; Holocene. Family THAIDIDAE Cronia fiscella (Gmelin, 1791); Ladd, 1977: 40, pl. 14, Fig. 8; late Miocene; [Cernohorsky, 1972: 128, pl. 36, Fig. 8; as Morula margariticola (Broderip, 1832)]. Drupa arachnoides (Lamarck, 1822); Bernstein, 1974; Leviten, 1974; Kohn and Leviten, 1976; Kohn, 1980a; {Emerson and Cernohorsky, 1973: pl. 2, Fig. 6; as Drupa ricinus ricinus). . Clathrata (Lamarck, 1816); Bernstein, 1974; Emerson and Cernohorsky, 1973: 33, pl. 2, Figs. 16-18. . elegans (Broderip and Sowerby, 1829); Bernstein, 1974; Emerson and Cernohorsky, 1973: 25, pl. 2, Fig. 12. . grossularia Réding, 1798; Emerson and Cernohorsky, 1973: 37, pl. 2, Figs. 23, 24; Kohn, 1980a. . morum Réding, 1798; Emerson and Cernohorsky, 1973: 17, pl. 2, Figs. 1-3; Bernstein, 1974; Kohn and Leviten, 1976; Kohn, 1980a; Leviten and Kohn, 1980. . ricina (Linnaeus, 1758); Emerson and Cernohorsky, 1973: 22, pl. 2, Figs. 7,8; Bernstein, 1974, Leviten, 1974; Kohn and Leviten, 1976; Kohn 1980a; Leviten and Kohn, 1980. io} Lofesits) (=) +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) MOLLUSCA 117 TABLE 1 (cont'd) Family THAIDIDAE (cont'd) D. rubusidaeus Réding, 1798; Emerson and Cernohorsky, 1973: 29, pl. 2, Figs. 13-15. Drupella elata (Blainville, 1832); Kohn, 1980a; as Drupella cornus (Roding, 1798); [Kay, 1979: 244, Fig. 87A). D. fenestrata (Blainville, 1832); [Cernohorsky, 1978: 68, pl. 19, Fig. 10). D. fragum (Blainville, 1832); [Moyer et al., 1982: Fig. 1]. D. rugosa (Born, 1778); Ladd, 1982: 44, pl. 11, Figs. 12, 13; as D. concatenata (Lamarck, 1822); Holocene; [Cernohorsky, 1972: 126, pl. 35, Fig. 5). Drupella sp., [Kay, 1979: 244, Fig. 87B; as Drupella ochrostoma (Blainville, 1832)). Maculotriton eximius (Reeve, 1844); [Cernohorsky, 1978: 71, pl. 20, Fig. 10). M. sculptile (Reeve, 1844); [Cernohorsky, 1972: 130, pl. 36, Fig. 12]. M. serriale (Deshayes, 1824); Leviten, 1974; as M. bracteatus (Hinds, 1844); Kohn and Leviten, 1976; Kohn, 1980a; [Cernohorsky, 1972: 129, pl. 36, Fig. 11). Morula anaxeres (Kiener, 1836); [Cernohorsky, 1972: 126, pl. 35, Fig. 10}. M. funiculata (Reeve, 1846); Ladd, 1977: 40; as Drupa (Drupella) cf. D. monilifera (Pease, 1860); Holocene; [Kay, 1979; 247, Fig. 83]. M. granulata (Duclos, 1832); Bernstein, 1974; Leviten, 1974; Kohn and Leviten, 1976; Ladd, 1977: 40, pl. 14, Fig. 7; Holocene; Kohn, 1980a; as Morula fiscella; Leviten and Kohn, 1980; [Cernohorsky, 1972: 127, pl. 36, Fig. 2]. M. nodicostata (Pease, 1868); [Cernohorsky, 1972: 127, pl. 36, Fig. 6]. M. spinosa (H. and A. Adams, 1853); [Cernohorsky, 1978: 128, pl. 36, Fig. 6]. M. squamosa (Pease, 1868); Figs. 3d, 3K. M. uva (Réding, 1798); Bernstein, 1974; Leviten, 1974; Kohn and Leviten, 1976; Kohn, 1980a; Leviten and Kohn, 1980; [Kay, 1979: 249, Fig. 87F]. Muricodrupa funiculus (Wood, 1828); Kohn and Leviten, 1976; [Kay, 1979: 238, Fig. 831]. Nassa serta (Bruguiere, 1789); [Cernohorsky, 1967: 135, pl. 29, Fig. 193]. Thais aculeata (Deshayes, 1844); [Cernohorsky, 1967: 130, pl. 28, Fig. 172; as Thais hippocastanum (Linnaeus, 1758)]. T. armigera (Link, 1807); Menge, 1973; Bernstein, 1974; Leviten and Kohn, 1980; [Kay, 1979: 251, Fig. 88G]. T. intermedia (Kiener, 1836); Bernstein, 1974; [Kay, 1979: 251, Fig. 88H]. T. tuberosa (Réding, 1798); Leviten, 1974; Kohn and Leviten, 1976; Leviten and Kohn, 1980; Kohn, 1980a; [Cernohorsky, 1967; 130, pl. 28, Fig. 174). Vexilla lineata A. Adams, 1853; [Kay, 1979: 252, Fig. 88E]. V. vexillum (Gmelin, 1791); [Kay, 1979: 252, Fig. 88D]. [The record of Purpura aperta (Blainville, 1832) cited by Kay (1979) has not been confirmed]. Family CORALLIOPHILIDAE Coralliophila costularis (Lamarck, 1816); [Cernohorsky, 1978: 73, pl. 21, Fig. 5]. C. d’orbignyana (Petit, 1851); [Kay, 1979: 255, Fig. 90A). C. erosa (Roding, 1798); [Cernohorsky, 1972: 131, pl. 37, Fig. 6]. +C. macneili Ladd, 1977; Ladd, 1977: 41-42, pl. 14, Figs. 12, 13; late Miocene. C. violacea (Kiener, 1836); [Kay, 1979: 256, Fig. 90C]. + Coralliophila (Coralliophila) sp.; Ladd, 1977: 41, pl. 14, Fig. 11; Holocene. + Coralliophila (Fusomurex) sp. A.; Ladd, 1977: 42, pl. 15, Fig. 3; late Miocene. + Coralliophila (Fusomurex) sp. B.; Ladd, 1977: 42, pl. 15, Fig. 4; early Miocene. Latiaxis inflata (Dunker, 1847); Ladd, 1977: 43, pl. 15, Fig. 5; late Miocene. Quoyula monodonta (Blainville, 1832); Ladd, 1977: 43-44, pl. 15, Fig. 7; Holocene; [Kay, 1979: 258, Fig. 90G; as Quoyula madreporarum Sowerby]. Rapa rapa (Linnaeus, 1758); [Cernohorsky, 1972: 130, pl. 37, Fig. 4]. Family BUCCINIDAE Caducifer truncata (Hinds, 1844); [Cernohorsky, 1972: 141, pl. 30, Fig. 4]. Cantharus eximius Reeve, 1846; [Cernohorsky, 1978: 76, pl. 22, Fig. 11]. C. farinosus (Gould, 1850); [Kay, 1979: 261, Fig. 92D]. C. pulcher (Reeve, 1846); [Cernohorsky, 1972: 142, pl. 38, Fig. 8). C. undosus (Linnaeus, 1758); Leviten, 1974; Kohn, 1980a; [Cernohorsky, 1972: 141, pl. 38, Fig. 5]. C. wagneri (Anton, 1839); [Cernohorsky, 1972: 142, pl. 39, Fig. 1]. Clivipollia fragaria (Wood, 1828); [Kay, 1979: 263, Fig. 92E]. Engina egregia (Reeve, 1844); [Cernohorsky, 1972: 143, pl. 39, Fig. 4]. E. lineata (Reeve, 1846); [Cernohorsky, 1972: 143, pl. 39, Fig. 4]. E. mendicaria (Linnaeus, 1758); [Cernohorsky, 1972: 144, pl. 38, Fig. 9]. +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) 118 KAY AND JOHNSON TABLE 1 (cont'd) Family BUCCINIDAE (cont’d) E. zonalis (Lamarck, 1822); [Salisbury, 1983a: 7]. Prodotia ignea (Gmelin, 1791); [Kay, 1979: 264, Fig. 92I]. P. iostoma (Gray, 1834); [Kay, 1979: 264, Fig. 92J]. Family COLUMBELLIDAE Aesopus spiculum (Duclos, 1846); [Cernohorsky, 1972: 138, pl. 41, Fig. 9]. Anachis miser (Sowerby, 1844); [Kay, 1979: 266, Fig. 94F]. Columbella cf. C. moleculina Duclos, 1835; Fig. 3N. + C. pardalina Lamarck, 1822; Ladd, 1977: 44, pl. 15, Fig. 11; late Miocene. Euplica amirantium (Smith, 1884); Fig. 3M. . deshayesii (Crosse, 1859); [Cernohorsky, 1972: 134, pl. 40, Fig. 6]. . ef. E. liocyma Pilsbry, 1904; Fig. 3Q. . cf. E. minuta Gould, 1860; Fig. 3R. . obesula Hervier, 1899; Fig. 3W. . turturina (Lamarck, 1822); Ladd, 1977: 44, pl. 15, Figs. 12, 13; Quaternary. . varians (Sowerby, 1832); Bernstein, 1974; Ladd, 1977: 45, pl. 15, Fig. 14; Holocene. tLavesopus eniwetokensis Ladd, 1977; Ladd, 1977: 45, pl. 16, Fig. 1; Holocene. Mitrella fusiformis (Pease, 1869); [Kay, 1979: 268, Fig. 94H-I]. M. cf. M. albina (Kiener, 1841); Fig. 3P. M. ligula (Duclos, 1840); [Cernohorsky, 1972: 135, pl. 41, Fig. 1). TM. sagitta (Gaskoin, 1851); Ladd, 1977: 47, pl. 16, Fig. 11; early Miocene to Holocene. Pyrene flava (Bruguiere, 1789); [Cernohorsky, 1972: 132, pl. 40, Fig. 2]. . cf. P. livescens (Reeve, 1859); Fig. 30. . obtusa (Sowerby, 1832); Ladd, 1977: 47, pl. 16, Fig. 10; Holocene; Fig. 3L. . ocellata (Link, 1807); [Cernohorsky, 1972: 132, pl. 40, Fig. 3]. . punctata (Bruguiere, 1789); Fig. 3V. . splendidula (Sowerby, 1874); [Habe, 1964: 89, pl. 28, Fig. 30). . testudinaria (Link, 1807); [Cernohorsky, 1972: 133, pl. 40, Fig. 4]. Seminella cf. S. dautzenbergi (Hervier, 1897); Fig. 3T. S. divaricata (Pilsbry, 1904); Fig. 3S. TS. smithi (Angas, 1877); Ladd, 1977: 47, pl. 16, Fig. 9 as Anachis; late Miocene. S. subphilodicia (Hervier, 1899); Fig. 3U. Zafra troglodytes (Souverbie, 1866); [Cernohorsky, 1972: 138, pl. 41, Fig. 8]. mommmmm de! ae} As} 38) as} 28) Zafrona pusilla (Pease, 1860); Ladd, 1977: 45, pl. 16, Fig. 2; as Zafrona lifuana (Hervier, 1899); Holocene. Family COLUBRARIIDAE Colubraria antiquata (Hinds, 1844); [Cernohorsky, 1972: 121, pl. 37, Fig. 2]. C. cf. C. cumingi (Dohrn, 1861); [Hirase, 1951: pl. 105, Fig. 1]. C. muricata (Lightfoot, 1786); [Kay, 1979: 271, Fig. 92L]. C. nitidula (Sowerby, 1833); [Cernohorsky, 1972: 121, pl. 37, Fig. 1]. C. tortuosa (Reeve, 1844); [Kay, 1979: 272, Fig. 92M]. Family MELONGENIDAE t Pugilina swartzi Ladd, 1977; Ladd, 1977: 51, pl. 17, Figs. 6, 8; early Miocene. Family NASSARIIDAE Nassarius comptus (A. Adams, 1852); [Cernohorsky, 1978: 87, pl. 27, Fig. 4]. N. concinnus (Powys, 1835); [Cernohorsky, 1978: 88, pl. 27, Fig. 9]. TN. eniwetokensis Ladd, 1977; Ladd, 1977: 52, pl. 17, Figs. 14, 15; late Miocene. N. gaudiosus (Hinds, 1844); Kohn, 1980a; [Cernohorsky, 1978: 86, pl. 27, Fig. 2]. N. glans (Linnaeus, 1758); [Cernohorsky, 1972: 151, pl. 45, Fig. 1]. N. graniferus (Kiener, 1834); [Cernohorsky, 1972: 146, pl. 42, Fig. 5]. TN. marshallensis Ladd, 1977; Ladd, 1977: 55, pl. 18, Figs. 6, 7; Miocene. N. papillosus (Linnaeus, 1758); [Kay, 1979: 275, Fig. 95A]. N. pauperus (Gould, 1850); [Cernohorsky, 1972: 152, pl. 44, Fig. 8]. TN. tiarula (Kiener, 1841); Ladd, 1977: 52, pl. 17, Fig. 9; pl. 21, Fig. 9; Holocene. Family FASCIOLARIIDAE Dolicholatirus lancea (Gmelin, 1791); [Cernohorsky, 1972: 161, pl. 46, Fig. 7]. Fasciolaria filamentosa (Roding, 1798); [Cernohorsky, 1972: 153, pl. 45, Fig. 3]. Fusinus undatus (Gmelin, 1791); [Cernohorsky, 1972: 162, pl. 48, Fig. 2]. Latirus craticulatus (Linnaeus, 1758); [Cernohorsky, 1972: 156, pl. 46, Fig. 5]. L. maculatus (Reeve, 1847); [Cernohorsky, 1978: 92, pl. 29, Fig. 1). +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) MOLLUSCA TABLE 1 (cont’d) 119 Family FASCIOLARIIDAE (cont’d) L. nodatus (Gmelin, 1791); [Kay, 1979: 278, Fig. 96H]. L. noumeensis (Crosse, 1870); [Cernohorsky, 1978: 92, pl. 29, Fig. 3]. L. polygonus (Gmelin, 1791); Leviten, 1974; as L. barclayi (Reeve, 1847); [Cernohorsky, 1972: 154, pl. 45, Fig. 7]. Peristernia chlorostoma (Sowerby, 1825); Ladd, 1977: 56, pl. 18, Fig. 12; Holocene. +P. eniwetokensis Ladd, 1977; Ladd, 1977: 57, pl. 18, Figs. 14, 15; Miocene. P. funiculata (Tapparone-Canefri, 1882); [Cernohorsky, 1978: 142, pl. 29, Fig. 4]. P. incarnata (Kiener, 1840); [Cernohorsky, 1972: 161, pl. 47, Fig. 5). P. nassatula (Lamarck, 1822); Ladd, 1977: 56, pl. 18, Fig. 10; early Miocene. Family VASIDAE Vasum ceramicum (Linnaeus, 1758); [Abbott, 1959: pl. 1, Fig. 1). V. turbinellum (Linnaeus 1758); Leviten, 1974; Kohn and Leviten, 1976; as V. cornigerum (Lamarck, 1822); Kohn, 1980a; [Abbott, 1959: pl. 1, Figs. 2, 3]. Family OLIVIDAE Belloliva simplex (Pease, 1867); Ladd, 1977: 58, pl. 18, Fig. 19; early Miocene; Holocene. Oliva annulata (Gmelin, 1791); [Cernohorsky, 1967: pl. 46, Fig. 334]. O. carneola (Gmelin, 1791); [Cernohorsky, 1967: pl. 46, Fig. 337]. O. miniacea (Réding, 1798); [Cernohorsky, 1967: pl. 47, Fig. 340]. O. panniculata Montfort, 1810; [Zeigler and Porreca, 1969: 76, pl. 10, Fig. 19]. O. paxillus Reeve, 1850; [Cernohorsky, 1967: 192, pl. 48, Fig. 345]. Family HARPIDAE + Eocithara? sp. B.; Rehder, 1973: 231; Ladd, 1977: 67; late Miocene. Harpa amouretta Roding, 1798; Rehder, 1973: 240, pl. 189, Figs. 6-11. H. gracilis Broderip and Sowerby, 1829; [Rehder, 1973, pl. 189, Figs. 3-5]. H. harpa (Linnaeus, 1758); Rehder, 1973: 237, pl. 187: Figs. 7-10. Family MARGINELLIDAE Granulina cf. G. vitrea Laseron, 1957; Fig. 4A. Haloginella micros (Bavay, 1922); Fig. 4E. H. cf. H. keppelense Laseron, 1957; Fig. 4F. Kogomea hervieri (Bavay, 1922); Fig. 4G. K. ovata Habe, 1951; Fig. 4B. Kogomea sp.; Fig. 4C. t Marginella ringicula Sowerby, 1900; Ladd, 1982: 59, pl. 37, Figs. 11, 12; Miocene. Persicula cf. P. pacifica (Pease, 1868); [Cernohosky, 1972: pl. 48, Fig. 7]. Tringinella sp.; Fig. 4D. Volvarina cf. V. avena (Kiener, 1834); Ladd, 1982: 59, pl. 37, Figs. 11, 12); Miocene. V. fusiformis (Hinds, 1844); [Kay, 1979: Fig. 99B]. V. cf. V. volunta (Laseron, 1957); [Laseron, 1957: 305, Fig. 83). Family MITRIDAE Cancilla carnicolor (Reeve, 1844); [Cernohorsky, 1978: 104, pl. 33, Fig. 7]. C. filaris (Linnaeus, 1758); [Cernohorsky, 1967, pl. 36, Fig. 241). C. fulgetrum (Reeve, 1844); [Cernohorsky, 1967: pl. 30, Fig. 188; as M. boissaci Montrouzier, 1858]. C. praestantissima (Roding, 1798); [Cernohorsky, 1967: pl. 37, Fig. 250]. Imbricaria conovula (Quoy and Gaimard, 1833); [Cernohorsky, 1978: 105, pl. 34, Fig. 5]. I. conularis (Lamarck, 1811); [Cernohorsky, 1967: 160, pl. 37, Fig. 254]. I. olivaeformis (Swainson, 1821); [Cernohorsky, 1967: 160, pl. 37, Fig. 255]. I. punctata (Swainson, 1821); Kohn and Leviten, 1976; [Cernohorsky, 1967: 160, pl. 37, Fig. 256]. I. vanikorensis (Quoy and Gaimard, 1833); [Cernohorsky, 1976: 160, pl. 37, Fig. 257]. Mitra acuminata Swainson, 1824; Cernohorsky, 1976: 494, [pl. 258, Figs. 26, 27]. . aurora Dohrn, 1861; [Cernohorsky, 1976: 441, pl. 257, Figs. 21, 22]. . avenacea Reeve, 1845; [Cernohorsky, 1976: 411, pl. 351). . cardinalis (Gmelin, 1791); Cernohorsky, 1976: 312, pl. 253, Figs. 5, 6. . chrysalis Reeve, 1844; Cernohorsky, 1976: 402, pl. 256, Figs. 16, 17. . chrysostoma Broderip, 1836; [Cernohorsky, 1976: pl. 256, Figs. 5, 6]. coarctata Reeve, 1844; [Cernohorsky, 1976: pl. 256, Fig. 8}. . coffea Schubert and Wagner, 1829; [Cernohorsky, 1976: 320, pl. 255, Figs. 3, 4]. . columbelliformis Kiener, 1838; [Cernohorsky, 1976: 491, pl. 258, Figs. 23-25). . contracta Swainson, 1820; [Cernohorsky, 1976: 393, pl. 256, Fig. 7]. . coronata Lamarck, 1811; [Cernohorsky, 1976: 437, pl. 257, Figs. 16-18]. SSSSSSESESEE +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) 120 KAY AND JOHNSON TABLE 1 (cont'd) Family MITRIDAE (cont’d) M. cucumerina Lamarck, 1811; Bernstein, 1974; Kohn and Leviten, 1976; Kohn, 1980a; [Cernohorsky, 1976: 400, pl. 256, Figs. 13-15). M. edentula Swainson, 1823; [Cernohorsky, 1976: pl. 257, Figs. 35, 36]. M. eremitarum (Réding, 1798; [Cernohorsky, 1976: 324, pl. 255, Figs. 7, 8]. M. fastigium Reeve, 1845; Bernstein, 1974; as Mitra brunnea Pease, 1868; [Cernohorsky, 1976: 494, pl. 258, Figs. 28, 29]. M. ferruginea Lamarck, 1811; [Cernohorsky, 1976: 397, pl. 256, Figs. 9, 10]. M. fraga Quoy and Gaimard, 1833; Ladd, 1977: 60, pl. 19, Fig. 9; Holocene; [Cernohorsky, 1976: 404, pl. 256, Figs. 18, 19]. M. fulvescens Broderip, 1836; [Cernohorsky, 1976: 431, pl. 257, Figs. 10, 11). M. imperialis Réding, 1798; Cernohorsky, 1976: 321, pl. 255, Figs. 15, 16. M. incompta (Solander in Lightfoot, 1786); [Cernohorsky, 1976: 322, pl. 253, Fig. 11). M. litterata Lamarck, 1811; Bernstein, 1974; Leviten, 1974; Kohn and Leviten, 1976; as Strigatella,; Kohn, 1980a; as Strigatella; Leviten and Kohn, 1980; [Cernohorsky, 1976: 482, pl. 258, Figs. 9-11). . luctuosa A. Adams, 1853; [Cernohorsky, 1976: 443, pl. 257, Figs. 23, 24). . mitra (Linnaeus, 1758); [Cernohorsky, 1976: 307, pl. 253, Fig. 1]. . nivea (Broderip, 1836); [Cernohorsky, 1976: 335, pl. 291). papalis (Linnaeus, 1758); [Cernohosky, 1976: 308, pl. 253, Fig. 3]. paupercula (Linnaeus, 1758); Leviten and Kohn, 1980; [Cernohorsky, 1976: 477, pl. 258, Figs. 1-3]. peculiaris Reeve, 1845; [Cernohorsky, 1976: 424, pl. 452). petrosa Sowerby, 1874; [Cernohorsky, 1976: pl. 292, Fig. 13; “albino form of M. ustulata”). pyramis (Wood, 1828); [Cernohorsky, 1976: 424, pl. 371). rubritincta Reeve, 1844; [Cernohorsky, 1976: 399, pl. 256, Figs. 11, 12]. scutulata (Gmelin, 1791); [Cernohorsky, 1976: 488, pl. 258, Figs. 17-20]. stictica (Link, 1807); [Cernohorsky, 1976: 309, pl. 253, Figs. 3, 4]. tabanula Lamarck, 1811; [Cernohorsky, 1976: 408, pl. 256, Fig. 22). . ticaonica Reeve, 1844; [Cernohorsky, 1976: 429, pl. 257, Figs. 4, 5]. turgida Reeve, 1845; Ladd, 1977: 60, pl. 19, Figs. 10, 11; late Miocene. typha Reeve, 1845; [Cernohorsky, 1976: 499, pl. 450). ustulata Reeve, 1844; [Cernohorsky, 1976: 337, pl. 292]. . vultuosa Reeve, 1845; [Cernohorsky, 1976: 442, pl. 391, Fig. 5). Neocancilla clathrus (Gmelin, 1791); [Cernohorsky, 1967: 154, pl. 35, Fig. 238). N. papilio (Link, 1807); [Cernohorsky, 1967: 154, pl. 32, Fig. 210). Pterygia crenulata (Gmelin, 1791); [Cernohorsky, 1967: 161, pl. 38, Fig. 259]. P. dactylus (Linnaeus, 1767); [Cernohorsky, 1967: 162, pl. 38, Fig. 260). P. fenestrata (Lamarck, 1811); [Cernohorsky, 1967: 162, pl. 38, Fig. 261]. P. nucea (Gmelin, 1791); [Cernohorsky, 1967: 162, pl. 38, Fig. 262]. Scabricola casta (Gmelin, 1791); [Cernohorsky, 1967: 160, pl. 37, Fig. 258]. S. caerulea (Reeve, 1844); [Cernohorsky, 1972: 173, pl. 51, Fig. 6]. S. desetangsii (Kiener, 1838); [Cernohorsky, 1967: 161, pl. 37, Fig. 251; as Swainsonia suffecta (Dautzenberg and Bouge, 1923)]. S. variegata (Gmelin, 1791); [Cernohorsky, 1978: 101, pl. 32, Fig. 4]. Subcancilla interlirata (Reeve, 1844); [Cernohorsky, 1978: 105, pl. 34, Fig. 3). S. verrucosa (Reeve, 1845); [Cernohorsky, 1967: 158, pl. 36, Fig. 249; as Cancilla]. ZESSSSS5SS55S5N5NN88 Family COSTELLARIIDAE Vexillum acupictum (Reeve, 1844); [Cernohorsky, 1967: 164, pl. 38, Fig. 256). V. amabilis (Reeve, 1845); [Cernohorsky, 1967: 90, pl. 44, Fig. 320]. TV. approximatum (Pease, 1860); Ladd, 1977: 66, pl. 20, Fig. 19; early Miocene. V. bernhardina (Réding, 1798); [Cernohorsky, 1967: 182, pl. 44, Fig. 322]. V. cadaverosum (Reeve, 1844); [Cernohorsky, 1967: 166, pl. 39, Fig. 269]. V. cancellarioides (Anton, 1839); Bernstein, 1974; Leviten, 1974; Kohn and Leviten, 1976; Leviten and Kohn, 1980; as Pusia nodosa (Swainson, 1840); Kohn, 1980a; as Pusia cancellarioides (Anton, 1839); (Kay, 1979: 325, Fig. 109B]. . catenatum (Broderip, 1836); [Cernohorsky, 1978: 116, pl. 40, Fig. 4]. . cernohorskyi Ladd, 1977; Ladd, 1977: 63, pl. 20, Figs. 6, 7; late Miocene. . corbiculum (Sowerby, 1870); [Cernchorsky, 1978: 112, pl. 37, Fig. 8]. . coronatum (Helbling, 1779); [Cernohorsky, 1967: 166, pl. 39, Fig. 27]. . costatum (Gmelin, 1791); [Cernohorsky, 1967: 167, pl. 39, Fig. 273]. SES BS SS SS +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) MOLLUSCA 121 TABLE 1 (cont'd) Family COSTELLARIIDAE (cont'd) . crocatum (Lamarck, 1811); [Cernohorsky, 1967: 167, pl. 39, Fig. 273}. V. deshayesi (Reeve, 1844); Ladd, 1977: 63, pl. 20, Fig. 8; late Miocene. . echinatum (A. Adams, 1853); [Cernohorsky, 1967: 168, pl. 40, Fig. 280]. . emiliae (Garrett, 1880); [Kay, 1979: 327, pl. 109F}. . exasperatum (Gmelin, 1791); [Cernohorsky, 1967: 168, pl. 40, Fig. 281). . exquisitum (Garrett, 1872); Ladd, 1977: 66, pl. 21, Fig. 1; Holocene. festum (Reeve, 1845); Ladd, 1977: 64, pl. 20, Fig. 10; Miocene. . filistriatum (Sowerby, 1874); [Cernohorsky, 1978: 110, pl. 36, Fig. 7]. fortiplicatum (Pease, 1868); [Cernohorsky, 1972: 176, pl. 52, Fig. 6). cf. V. gembacanum (Martin, 1884); Ladd, 1977: 65, pl. 20, Fig. 14; Miocene. goubini (Hervier, 1898); [Cernohorsky, 1981: 101, pl. 3, Fig. 9]. granosum (Gmelin, 1791); [Cernohorsky, 1967: 170, pl. 40, Fig. 286]. gruneri (Reeve, 1844); [Cernohorsky 1967: 171, pl. 41, Fig. 287]. humilis (Hervier, 1897); [Cernohorsky, 1981: 96, pl. 1, Fig. 4]. infaustum (Reeve, 1845); [Cernohorsky, 1967: 183, pl. 45, Fig. 326]. interruptum (Anton, 1839); [Cernohorsky, 1978: 112, pl. 37, Fig. 10). . interstriatum (Sowerby, 1870); [Kay, 1979: 320, Fig. 108P]. lautum (Reeve, 1845); [Cernohorsky, 1972: 177, pl. 52, Fig. 10). . leucozonias (Deshayes, 1834); Ladd, 1977: 64, pl. 29, Fig. 11; Holocene. . luculentum (Reeve, 1845); [Cernohorsky, 1967: 172, pl. 41, Fig. 29]. . mica (Reeve, 1845); [Cernohorsky, 1967: 172, pl. 41, Fig. 292). . michaui (Crosse and Fischer, 1864); [Cernohorsky, 1967: 171, pl. 41, Fig. 288; as V. intertaeniatum (Sowerby, 1874)]. . microzonias (Lamarck, 1811); [Cernohorsky, 1967: 183, pl. 45, Fig. 328]. . millecostatum (Broderip, 1836); [Cernohorsky, 1967: pl. 41, Fig. 321; as Pusia adamsoni (Reeve, 1845)]. modestum (Reeve, 1845); [Kay, 1979: 322, Fig. 108H]. moelleri (Kuster, 1840); [Cernohorsky, 1972: 178, pl. 51, Fig. 9]. . obeliscus (Reeve, 1844); [Cernohorsky, 1967: 174, pl. 41, Fig. 296]. obtusispinosum (Sowerby, 1874); [Cernohorsky, 1967: 174, pl. 42, Fig. 297]. pacificum (Reeve, 1845); Ekdale et al. 1979: as V. speciosum (Reeve, 1844); [Cernohorsky, 1967: 174, pl. 42, Fig. 298). . pagodula (Hervier, 1897); [Cernohorsky, 1981: 95, pl. 1, Fig. 3]. . patriarchalis (Gmelin, 1791); [Cernohorsky, 1967: 184, pl. 45, Fig. 329]. . polygonum (Gmelin, 1791); [Cernohorsky, 1978: 108, pl. 36, Fig. 3]. . roseum (Broderip, 1836); Ladd, 1977: 65, pl. 20, Fig. 15; Pleistocene. . rubrum (Broderip, 1836); [Kay, 1979: 331, Fig. 110E). . sanguisugum (Linnaeus, 1758); [Cernohorsky, 1967: 176, pl. 43, Fig. 206]. speciosum (Reeve, 1844); Ekdale et al., 1979; [Cernohorsky, 1972: 177, pl. 52, Fig. 9]. suavis (Souverbie, 1875); [Cernohorsky, 1978: 116, pl. 40, Fig. 3). tuberosum (Reeve, 1845); [Cernohorsky, 1967: 186, pl. 45, Fig. 333]. turben (Reeve, 1844); [Cernohorsky, 1978: 117, pl. 39, Fig. 8]. . turrigerum (Reeve, 1845); [Cernohorsky, 1972: 175, pl. 52, Fig. 4). tusum (Reeve, 1845); [Cernohorsky, 1972: 178, pl. 52, Fig. 11). . unifascialis (Lamarck, 1811); [Cernohorsky, 1967: 180, pl. 44, Fig. 316]. . vulpecula (Linnaeus, 1758); [Cernohorsky, 1967: 180, pl. 44, Fig. 317]. . zelotypum (Reeve, 1845); [Cernohorsky, 1972: 177, pl. 52, Fig. 8). Family TURRIDAE Anacithara nanisca (Hervier, 1897); Fig. 41. A. notabilis (E. A. Smith, 1888); Fig. 4J. A. stricta Hedley, 1922; [Fig. 4H]. Carinapex minutissima (Garrett, 1873); [Kay, 1979: 344, Fig. 1150]. Clavus bilineatus (Reeve, 1845); Fig. 4L. . canicularis (Roding, 1798); [Cernohorsky, 1972: 185, pl. 53, Fig. 13]. . laetus (Hinds, 1843); [Cernohorsky, 1978: 153, pl. 55, Fig. 1]. . lamberti (Montrouzier, 1860); [Cernohorsky, 1978: 153, pl. 55, Fig. 2]. . nodulosa (Pease, 1860); [Kay, 1965: 80, pl. 14, Figs. 9, 10]. . opalus (Reeve, 1845); [Cernohorsky, 1978: 152, pl. 54, Fig. 11]. . pica (Reeve, 1843); [Cernohorsky, 1978: 152, pl. 54, Fig. 9]. < SSSSSSSS SSS SSS SS SSS SSS SS ws SS Ss PSAP S13 JIGS OPO) O) ©) +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) 122 KAY AND JOHNSON TABLE 1 (cont’d) Family TURRIDAE (cont’d) C. unizonalis (Lamarck, 1822); [Cernohorsky, 1972: 186, pl. 53, Fig. 14). C. viduus (Reeve, 1845); [Cernohorsky, 1978: 159, pl. 54, Fig. 7]. Clavus sp. A.; Fig. 4M. Clavus sp. B.; Fig. 4K. Daphnella aureola (Reeve, 1843); [Cernohorsky, 1978: 159, pl. 57, Fig. 1). D. flammea (Hinds, 1843); [Cernohorsky, 1978: 158, pl. 56, Fig. 12]. D. cf. D. granata Hedley, 1922; [Hedley, 1922: 328, pl. 58, Fig. 162). D. olyra (Reeve, 1845); [Cernohorsky, 1978: 159, pl. 57, Fig. 3]. D. rissoides (Reeve, 1843); [Powell, 1966: 124, pl. 19, Fig. 18). Etrema alphonsianum (Hervier, 1896); [Hedley, 1922: pl. 47, Fig. 71]. E. crassilabrum (Reeve, 1843); [Cernohorsky, 1978: 156, pl. 56, Fig. 5}. E. cf. E. polydesma Hedley, 1922; [Hedley, 1922: 282, pl. 47, Fig. 83]. E. scalarina (Deshayes, 1863); [Deshayes, 1863: 109, pl. 12, Figs. 12-14] E. trigonostomum (Hervier, 1896); Fig. 4R. Eucithara angiostoma (Pease, 1868); [Kay, 1979: 352, Fig. 116B]. . celebensis (Hinds, 1843); [Cernohorsky, 1972: 189, pl. 54, Fig. 7]. . cithara (Gould, 1849); Fig. 4N. . debilis (Pease, 1868); Fig. 4P. . marshallensis (Ladd, 1982); Ladd, 1982: 68, pl. 22, Figs. 15-19; Miocene. . reticulata (Reeve, 1846); [Cernohorsky, 1972: 186, pl. 54, Fig. 3). . cf. E. stricta Hedley, 1922; [Hedley, 1922: 304, pl. 51, Fig. 123). . stromboides (Reeve, 1846); [Cernohorsky, 1972: 186, pl. 54, Fig. 1]. . vexillum (Reeve, 1846); [Hirase, 1951: pl. 115, Fig. 18]. Eucithara sp.; Fig. 4P. Eucyclotoma tricarinata (Kiener, 1840); [Cernohorsky, 1978: 160, pl. 57, Fig. 8]. Iredalea pygmaea (Dunker, 1860); [Cernohorsky, 1972: 186, pl. 55, Fig. 4]. Kermia cf. K. bifasciata (Pease, 1860); [Kay, 1979: 360, Fig. 118F]. moammmmmm K. canistra (Hedley, 1922); [Cernohorsky, 1978: 162, pl. 57, Fig. 10). K. cf. K. concinna (Hedley, 1922); Fig. 4S. K. daedalea (Garrett, 1873); [Kay, 1979: 361, Fig. 118]]. K. edychroa (Hervier, 1896); Fig. 4U. K. cf. K. granicostata (Reeve, 1846); Fig. 4Q. K. maculosa (Pease, 1862); [Cernohorsky, 1978: 161, pl. 57, Fig. 11; as Kermia barnardi Brazier, 1876]. Lienardia apiculata (Montrouzier, 1864); [Kay, 1979: 353, Fig. 116L]. L. crassicostata (Pease, 1860); [Kay, 1979: 354, Fig. 116F]. L. cf. L. falsaria Hedley, 1922; [Cernohorsky, 1978: 156, pl. 56, Fig. 3]. L. goubini (Hervier, 1896); Fig. 4V. L. idiomorpha (Hervier, 1897); Fig. 4W. L. roseotincta (Montrouzier, 1872); [Cernohorsky, 1978: 156, pl. 56, Fig. 4). L. rubida (Hinds, 1844); [Cernohorsky, 1972: 189, pl. 54, Fig. 9]. Lophiotoma acuta (Perry, 1811); [Powell, 1964: 303, pl. 180, Figs. 1, 2]. L. albina (Lamarck, 1822); [Powell, 1964: 306, pl. 180, Figs. 11, 12]. TL. eniwetokensis Ladd, 1982; Ladd, 1982: 62, pl. 19, Figs. 2, 3; early Miocene. Lovellona atramentosa (Reeve, 1849); [Kay, 1979: 348, Fig. 115A). Macteola interrupta (Reeve, 1845); [Cernohorsky, 1978: 157, pl. 56, Fig. 9]. M. segesta (Chenu, 1850); [Kay, 1979: 355, Fig. 116M]. Macteola sp.; not figured. Microdaphne morrisoni Rehder, 1980; [Rehder, 1980: 88, pl. 11, Figs. 5, 6). Mitrolumna stepheni (Melville and Standen, 1897); [Cernohorsky, 1978: 163, pl. 58, Fig. 5]. M. metula (Hinds, 1843); [Cernohorsky, 1978: 162, pl. 58, Fig. 2; as Mitromorpha lachryma (Reeve, 1845)]. Nodotoma harucoa (Bartsch, 1941); [Johnson, 1964: pl. 5, Fig. 4; as Clathurella lacunosa Gould, 1860]. Philbertia nexa (Reeve, 1845); [Cernohorsky, 1978: 162, pl. 58, Fig. 1]. P. cf. P. crassilabrum (Reeve, 1843); Fig. 4T. P. rufolirata (Hervier, 1896); [Hervier, 1896: 99, pl. 2, Fig. 5]. P. tincta (Reeve, 1846); [Cernhorsky, 1978: 162, pl. 57, Fig. 15). Tritonoturris cumingii (Powys, 1835); [Kay, 1979: 363, Fig. 118Q]. T. robillardi (H. Adams, 1869); [Powell, 1966: pl. 20, Fig. 23). Turridrupa albofasciata (Smith, 1877); [Powell, 1967: 416, pl. 303, Figs. 3, 4]. +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) MOLLUSCA 123 TABLE 1 (cont'd) Family TURRIDAE (cont'd) T. astricta (Reeve, 1843); [Powell, 1967: 417, pl. 305, Fig. 4]. Turridrupa bijubata (Reeve, 1843); [Powell, 1967: 418, pl. 303, Figs. 1, 2]. T. cerithina (Anton, 1839); [Powell, 1967: 420, pl. 305, Fig. 1]. T. diffusa (Powell, 1967); Powell, 1967: 422, pl. 305, Fig. 5. T. jubata (Hinds, 1843); [Powell, 1967: 423, pl. 301, Fig. 3). T. weaveri Powell, 1967; Powell, 1967: 423, pl. 303, Fig. 5 Turris acuta (Perry, 1811); [Powell, 1964: 303, pl. 180). T. albina (Lamarck, 1822); [Powell, 1964: 306, pl. 180, Figs. 11, 12]. T. cryptorraphe (Sowerby, 1825); [Powell, 1964: 335, pl. 181, Figs. 14, 15]. T. spectabilis (Reeve, 1843); Powell, 1964: 336, pl. 181, Figs. 16, 17. Xenuroturris cingulifera (Lamarck, 1822); [Powell, 1964: 322, pl. 175, Fig. 12]. X. kingae Powell, 1964; [Powell, 1964: 325, pl. 252, Fig. 6]. Family CONIDAE Conus ammiralis Linnaeus, 1758; [Cernohorsky, 1978: 124, pl. 3, Fig. 2]. . arenatus Hwass in Bruguiere, 1792; [Cernohorsky, 1978: 131, pl. 43, Fig. 4]. . aulicus Linnaeus, 1758; [Cernohorsky, 1978: 122, pl. 1, Fig. 5). . aureus Hwass in Bruguiere, 1792; [Cernohorsky, 1978: 121, pl. 41, Fig. 1]. . auricomus Hwass in Bruguiere, 1792; [Cernohorsky, 1978: 123, pl. 1, Fig. 9]. . aurisiacus Linnaeus, 1758; [Cernohorsky, 1967: 217, pl. 55, Fig. 412]. boeticus Reeve, 1849; [Cernohorsky, 1967: pl. 56, Fig. 415]. . bullatus Linnaeus, 1758; [Cernohorsky, 1972: pl. 55, Fig. 8). capitaneus Linnaeus, 1758; [Cernohorsky, 1978: 127, pl. 5, Fig. 1). catus Hwass in Bruguiere, 1792; Bernstein, 1974; Leviten, 1974; Kohn and Leviten, 1976; Kohn, 1980a; Leviten and Kohn, 1980; [Kay, 1979: 370, Fig. 121H)]. . chaldaeus (Roding, 1798); Bernstein, 1974; Leviten, 1974; Kohn and Leviten, 1976; Kohn, 1980a; Leviten and Kohn, 1980; [Kay, 1979: 370, Fig. 1211]. . circumactus Iredale, 1929; [Kay, 1979: 370, Fig. 1214]. . circumcisus Born, 1778; [Cernohorsky, 1978: 141, pl. 49, Fig. 1]. . coelinae Crosse, 1858; [Cernohorsky, 1978: 130, pl. 43, Fig. 8]. . consors Sowerby, 1833; Kohn, 1980a; [Walls, n.d.: 241]. . coronatus Gmelin, 1791; Bernstein, 1974; Leviten, 1974; Kohn and Leviten, 1976; Kohn, 1980a; Leviten and Kohn, 1980; [Cernohorsky, 1978: 133, pl. 44, Fig. 6). . crocatus Lamarck, 1810; [Cernohorsky, 1978: 122, pl. 1, Fig. 7]. . cylindraceus Broderip and Sowerby, 1830; [Cernohorsky, 1978: 143, pl. 50, Fig. 2]. . distans Hwass in Bruguiere, 1792; [Cernohorsky, 1978: 126, pl. 3, Fig. 7]. . ebraeus Linnaeus, 1758; Menge, 1973; Bernstein, 1974; Leviten, 1974; Kohn and Leviten, 1976; Kohn, 1980a; Leviten and Kohn, 1980; [Kay, 1979: 371, Fig. 121K]. . eburneus Hwass in Bruguiere, 1792; Demond, 1957: 327; Ladd, 1982: 73, pl. 26, Fig. 5; early Miocene; Kohn, 1980a; Kohn, 1980b. . episcopus Hwass in Bruguiere, 1792; [Hinton, 1978: pl. 69, Fig. 5]. . flavidus Lamarck, 1810; Bernstein, 1974; Kohn and Leviten, 1976; Levitten and Kohn, 1980; [Kay, 1979: 373, Fig. 122E]. . floccatus Sowerby, 1839; [Cernohorsky, 1972: 192, pl. 55, Fig. 6]. frigidus Reeve, 1848; Leviten, 1974; Kohn and Leviten, 1976; Kohn, 1980a; Leviten and Kohn, 1980; [Cernohorsky, 1967: 220, pl. 56, Fig. 419). generalis Linnaeus, 1767; [Cernohorsky, 1978: 124, pl. 3, Fig. 3]. geographus Linnaeus, 1758; [Cernohorsky, 1978: 137, pl. 7, Fig. 4]. glans Hwass in Bruguiere, 1792; [Cernohorsky, 1967: 222, pl. 56, Fig. 421]. imperialis Linnaeus, 1758; [Kay, 1979: 373, Fig. 122J]. legatus Lamarck, 1810; [Hinton, 1978: pl. 69, Fig. 10]. leopardus (Roding, 1798); Kohn, 1980a; Kohn, 1980b; [Kay, 1979: 374, Fig. 122I]. litoglyphus Hwass in Bruguiere, 1792; [Kay, 1979: 374, Fig. 122B}. litteratus Linnaeus, 1758; Kohn, 1980a; Kohn, 1980b; [Cernohorsky, 1978: 129, pl. 5, Fig. 9]. lividus Hwass in Bruguiere, 1792; Kohn, 1980a; Kohn, 1980b; [Kay, 1979: 374, Fig. 122B}. luteus Sowerby, 1833; [Hinton, 1972: pl. 40, Fig. 11]. magnificus Reeve, 1843; [Cernohorsky, 1978: 123, pl. 41, Fig. 2]. magus Linnaeus, 1758; [Cernohorsky, 1978: 138, pl. 47, Fig. 2]. marmoreus Linnaeus, 1758; Kohn, 1980a; Kohn, 1980b; [Cernohorsky, 1978: 125, pl. 42, Fig. 1]. . miles Linnaeus, 1758; [Kay, 1979: 375, Fig. 122H]. ADOIMOVOOONOEMOOOOOG OF OD O HOOEVQ YOOHOOOND OQ POOOOOVMOO® (This table continued on next page.) 124 KAY AND JOHNSON TABLE 1 (cont’d) Family CONIDAE (cont’d) C. miliaris Hwass in Bruguiere, 1792; Leviten, 1974; Kohn and Leviten, 1976; Kohn, 1980a; Leviten and Kohn, 1980; [Cernohorsky, 1978: 132, pl. 44, Fig. 4]. . moreleti Crosse, 1858; [Kay, 1979: 375, Fig. 122F]. . musicus Hwass in Bruguiere, 1792; [Cernohorsky, 1967: 224, pl. 57, Fig. 424]. . mustelinus Hwass in Bruguiere, 1792; [Cernohorsky, 1978: 127, pl. 5, Fig. 2]. . nussatella Linnaeus, 1758; [Kay, 1979: 376, Fig. 122D}. obscurus Sowerby, 1833; [Kay, 1979: 376, Fig. 122C]. omaria Hwass in Bruguiere, 1792; [Cernohorsky, 1978: 123, pl. 1, Fig. 8]. pertusus Hwass in Bruguiere, 1792; [Kay, 1979: 378, Fig. 123C]. . pulicarius Hwass in Bruguiere, 1792; Kohn, 1980a; Kohn, 1980b; [Kay, 1979: 378, Fig. 123C]. . quercinus Solander in Lightfoot, 1786; [Kay, 1979: 379, Fig. 123J]. rattus Hwass in Bruguiere, 1792; Bernstein, 1974; Kohn, 1980a; Leviten and Kohn, 1980; [Kay, 1979: 379, Fig. 123E). . retifer Menke, 1829; Leviten and Kohn, 1980; [Cernohorsky, 1978: 122, pl. 1, Fig. 4]. . sanguinolentus Quoy and Gaimard, 1834; [Cernohorsky, 1967: 226, pl. 58, Fig. 434). . scabriusculus Dillwyn, 1817; [Cernohorsky, 1967: 226, pl. 58, Fig. 435]. sponsalis Hwass in Bruguiere, 1792; Bernstein, 1974; Leviten, 1974; Kohn and Leviten, 1976; Kohn, 1980a; Leviten and Kohn, 1980; [Kay, 1979: 380, Fig. 123G]. striatus Linnaeus, 1758; [Kay, 1979: 380, Fig. 123]]. sugillatus Reeve, 1844; Kohn, 1980a; Kohn, 1980b; [Cernohorsky, 1967: 228, pl. 59, Fig. 437]. tenuistriatus Sowerby, 1858; [Cernohorsky, 1967: 230, pl. 59, Fig. 438]. terebra Born, 1778; [Cernohorsky, 1978: 140, pl. 48, Fig. 7]. tessulatus Born, 1778; [Cernohorsky, 1978: 125, pl. 3, Fig. 5). textile Linnaeus, 1758; [Cernohorsky, 1978: 121, pl. 1, Fig. 2]. . tulipa Linnaeus, 1758; [Cernohorsky, 1978: 137, pl. 7, Fig. 5]. varius Linnaeus, 1758; [Cernohorsky, 1978: 135, pl. 7, Fig. 3]. vexillum Gmelin, 1791; [Kay, 1979: 381, Fig. 12a]. . virgo Linnaeus, 1758; [Cernohorsky, 1978: 131, pl. 43, Fig. 6]. . vitulinus Hwass in Bruguiere, 1792; [Kay, 1979: 382, Fig. 123K]. Family TEREBRIDAE Hastula lanceata (Linnaeus, 1758); [Cernohorsky, 1967: 210, pl. 54, Fig. 403]. H. penicillata (Hinds, 1844); [Cernohorsky, 1967: 212, pl. 54, Fig. 406]. H. solida (Deshayes, 1857); Miller, 1966; as Terebra; [Cernohorsky, 1967: 212, pl. 54, Fig. 408}. H. strigilata (Linnaeus, 1758); [Cernohorsky, 1967: 212, pl. 54, Fig. 409]. Terebra affinis Gray, 1834; Miller, 1966; Ladd, 1982: 82, pl. 30, Figs. 4, 5; Quaternary. . amoena Deshayes, 1859; [Cernohorsky, 1967: 196, pl. 49, Fig. 350). . areolata (Link, 1807); [Cernohosky,1967: 196, pl. 49, Fig. 354]. argus Hinds, 1844; Miller, 1966; [Cernohorsky, 1967: 196, pl. 49, Fig. 354). babylonia Lamarck, 1822; Miller, 1966; [Cernohorsky, 1967: 198, pl. 49, Fig. 355). cerithina Lamarck, 1822; [Cernohorsky, 1967: 198, pl. 49, Fig. 356]. chlorata Lamarck, 1822; [Cernohorsky, 1967: 198, pl. 49, Fig. 357]. cingulifera Lamarck, 1822; Miller, 1966; [Cernohorsky, 1967: 198, pl. 49, Fig. 358). columellaris Hinds, 1844; [Cernohorsky, 1967: 198, pl. 49, Fig. 359]. conspersa Hinds, 1844; [Cernohorsky, 1967: 198, pl. 49, Fig. 360). . crenulata (Linnaeus, 1758); Miller, 1966; [Cernohorsky, 1967: 199, pl. 50, Fig. 361]. . dimidiata (Linnaeus, 1758); Miller, 1966; [Cernohorsky, 1967: 199, pl. 50, Fig. 362]. . felina (Dillwyn, 1817); Miller, 1966; as Terebra tigrina Gmelin, 1791; [Cernohorsky, 1967: 199, pl. 50, Fig. 364]. . fenestrata Hinds, 1844; [Cernohorsky, 1967: 200, pl. 50, Fig. 365). . funiculata Hinds, 1844; [Cernohorsky, 1967: 200, pl. 50, Fig. 369]. guttata (Roding, 1798); Miller, 1966; [Cernohorsky, 1967: 202, pl. 50, Fig. 372]. kilburni Burch, 1965; [Cernohorsky, 1967: 202, pl. 50, Fig. 372]. maculata (Linnaeus, 1758); Miller, 1966; [Cernohorsky, 1967: 204, pl. 51, Fig. 376]. nebulosa Sowerby, 1825; Miller, 1966; [Cernohorsky, 1967: 204, pl. 51, Fig. 380). . paucistriata (E. A. Smith, 1873); [Cernohorsky, 1967: 205, pl. 51, Fig. 383]. pertusa (Born, 1798); [Cernohorsky, 1967: 205, pl. 52, Fig. 384]. quoygaimardi Cernohorsky and Bratcher, 1976; [Cernohorsky, 1978: 146, pl. 52, Fig. 3]. subulata (Linnaeus, 1767); Miller, 1966; [Cernohorsky, 1967: 205, pl. 52, Fig. 385]. . textilis Hinds, 1844; [Cernohorsky, 1967: 206, pl. 52, Fig. 391]. . undulata Gray, 1834; [Cernohorsky, 1967: 208, pl. 52, Fig. 391). DAIOAVAIOGOEO OOGOGQ VBDOOOQVOQOO®& NASA AAAS AAAS (This table continued on next page.) MOLLUSCA 125 TABLE 1 (cont'd) Family TEREBRIDAE (cont’d) tTerebra (Strioterebra) sp. A. Ladd, 1982; Ladd, 1982: 82, pl. 30, Fig. 3; late Miocene. tTerebra (Oxymeris) sp. B. Ladd, 1982; Ladd, 1982: 84, pl. 30, Fig. 14; late Miocene. Terenolla pygmaea (Hinds, 1844); [Cernohorsky, 1978: 150, pl. 53, Fig. 5]. [The records of T. cancellata Quoy and Gaimard, 1832, T. laevigata Gray, 1834, and T. picta Hinds, 1844, cited by Miller (1966) have not been confirmed.] Subclass OPISTHOBRANCHIA Order ENTOMOTAENIATA Family PYRAMIDELLIDAE Chemnitzia cf. C. gabrieli (Hedley, 1909); Fig. 5A. Chrysalida alveata (A. Adams, 1861); Fig. 5E. Contraxiala obliqua Laseron, 1956; [Ponder, 1985: 108, Fig. 75A—B). Costabieta horrida (Garrett, 1873); [Cernohorsky, 1978: 48, Figs. 13, 14; as Rissoina]. Evalea debilis (Pease, 1868); [Pease, 1868: 292, pl. 24, Fig. 2]. E. densestriata (Garrett, 1873); Fig. 5B. E. peasei (Dautzenberg and Bouge, 1933); [Kay, 1979: 407, Fig. 132A]. Herviera gliriella (Melvill and Standen, 1896); [Kay, 1979: 407, Fig. 132C). Miralda gemmifera (Dautzenberg and Fischer, 1906); [Dautzenberg and Fischer, 1906: 195, pl. 6, Fig. 2]. Odostomia exilis (Garrett, 1873); Fig. 5C. O. gulicki Pilsbry, 1918; [Kay, 1979: 411, Fig. 132J]. O. oxia Watson, 1886; [Kay, 1979: 41, Fig. 132I]. Otopleura mitralis (A. Adams, 1855); [Cernohorsky, 1972: 201, pl. 57, Fig. 6]. O. nodicincta (A. Adams, 1855); [Cernohorsky, 1972: 201, pl. 57, Fig. 5]. Pyramidella pusilla A. Adams, 1854; [Dall and Bartsch, 1906: 324-325, pl. 24, Fig. 6]. P. sulcata (A. Adams, 1855); [Cernohorsky, 1972: 200, pl. 57, Fig. 2]. Pyrgulina cf. P. densecostata (Garrett, 1873); Fig. 5D. P. oodes (Watson, 1886); [Kay, 1979: 414, Fig. 132N]. Syrnola lutea (Garrett, 1873). S. polita (Pease, 1868). Turbonilla cf. T. infantula Dall and Bartsch, 1906; [Dall and Bartsch, 1906: 338, pl. 20, Fig. 2]. T. lirata (A. Adams, 1855); [Kay, 1979: 414, Fig. 133F]. T. orthoplicatulata Nomura, 1936; [Nomura, 1936: 74, pl. 12, Figs. 101a, 101bj. T. pseudomala Nomura, 1938; [Nomura, 1938: 29, pl. 4, Figs. 33a, 33b). T. cf. T. tenuissima Hedley, 1909; [Hedley, 1909: 450, pl. 42, Fig. 78). Order SOLEOLIFERA Family ONCHIDIIDAE Onchidiella evelinae Marcus and Burch, 1965; Marcus and Burch, 1965: 253, Figs. 36-39. Peronia personii (Cuvier, 1804); Marcus, 1965: 264. Order CEPHALASPIDEA Family ACTAEONIDAE Pupa alveola (Souverbie, 1863); [Cernohorsky, 1972: 204, pl. 58, Fig. 3]. P. sulcata (Gmelin, 1791); [Cernohorsky, 1972: 203, pl. 58, Fig. 2). Pupa sp.; Fig. 5H. [The record of Acteon variegatus (Bruguiere, 1789) cited by Ekdale et al. (1979) has not been confirmed.] Family RINGICULIDAE Ringicula cf. R. dolearis Gould, 1860; Fig. 5F. Family AMPLUSTRIDAE Hydatina amplustre (Linnaeus, 1758); Johnson and Boucher, 1983: 254; [Bertsch and Johnson, 1981: 16]. H. physis (Linnaeus, 1758); Johnson and Boucher, 1983: 254; [Bertsch and Johnson, 1981: 16]. Micromelo guamensis (Quoy and Gaimard, 1825); Johnson and Boucher, 1983: 255; [Bertsch and Johnson, 1981: 17]. Family SMARAGDINELLIDAE Phanerophthalmus smaragdinus (Ruppell and Leuckart, 1828); Marcus and Burch, 1965: 238; as Lathophthalmus smaragdinus (Ruppell and Leuckart, 1828); [Kay, 1979: 422, Fig. 136B]. Smaragdinella calyculata (Broderip and Sowerby, 1829); Marcus and Burch, 1965: 236-238, Fig. 1; [Kay, 1979: 423, Fig. 136D]. Family BULLIDAE Bulla vernicosa Gould, 1859; Johnson and Boucher, 1983: 255; [Kay, 1979: 423, Fig. 137E]. Cylichnatys angusta (Gould, 1859); Fig. 5M. tSpecies not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) 126 KAY AND JOHNSON TABLE 1 (cont'd) Family ATYIDAE Atys cylindricus (Helbling, 1779); [Cernohorsky, 1972: 208, pl. 59, Fig. 3]. A. debilis Pease, 1860; [Kay, 1979: 424, Fig. 137C]. A. cf. A. kuhnsi Pilsbry, 1917; [Kay, 1979: 425, Fig. 137B). A. naucum (Linnaeus, 1758); [Cernohorsky, 1972: 208, pl. 59, Fig. 2]. Haminoea crocata Pease, 1860; [Kay, 1979: 426, Fig. 137F]. H. curta (A. Adams, 1850); [Kay, 1979: 427, Fig. 137]]. H. cymbalum (Quoy and Gaimard, 1835); Marcus and Burch, 1965: 243; as Lamprohaminoea cymbalum (Quoy and Gaimard, 1835); [Bertsch and Johnson, 1981: 18]. H. galba Pease, 1861; [Kay, 1979: 427, Fig. 137G]. H. cf. H. japonica Pilsbry, 1895; Fig. 5G. H. linda Marcus and Burch, 1965; Marcus and Burch, 1965: 241, Figs. 12-16; [Johnson, 1982: 89]. H. musetta Marcus and Burch, 1965; Marcus and Burch, 1965: 239, Fig. 6. Family RETUSIDAE Coleophysis minima (Yamakawa, 1911); Fig. 5L. Family AGLAJIDAE Chelidonura hirundinina (Quoy and Gaimard, 1833); [Kay, 1979: 429, Fig. 138E]. C. inornata Baba, 1949; Johnson and Boucher, 1983: 255; [Johnson, 1982: 88). Odontoglaja guamensis Rudman, 1978a; [Rudman, 1978a: 90, Fig. 1]. Philinopsis gardineri Eliot, 1903; Johnson and Boucher, 1983: 255; [Bergh, 1905: pl. 3, Fig. 6]. Family GASTROPTERIDAE Gastropteron brunneomarginatum Carlson and Hoff, 1974; [Carlson and Hoff, 1974: 347, pl. X, Fig. 1]. Family SCAPHANDRIDAE Acteocina cf. A. hawaiensis Pilsbry, 1921; [Kay, 1979: 431, Fig. 137M]. A. sandwicensis Pease, 1860; [Kay, 1979: 431, Fig. 137N]. Adamnestia japonica (A. Adams, 1862); Fig. 51. Nakamigawaia spiralis Kuroda and Habe, 1961; Fig. 5d. Order ANASPIDEA Family APLYSIIDAE Aplysia dactylomela Rang, 1828; [Kay, 1979: 437, Fig. 131A]. A. parvula Guilding in Morch, 1863; Johnson and Boucher, 1983: 255; [Kay, 1979: 439, Fig. 131C]. A. pulmonica Gould, 1852; Marcus, 1965: 266. Dolabella auricularia (Lightfoot, 1786); Johnson and Boucher, 1983: 256; [Kay, 1979: 440, Fig. 140D). Dolabrifera dolabrifera (Rang, 1828); Marcus and Burch, 1965: 244; [Bertsch and Johnson, 1981: 24). Stylocheilus longicaudus (Quoy and Gaimard, 1824); Marcus and Burch, 1965: 244; Johnson and Boucher, 1983: 256; [Kay, 1979: 440, Fig. 141B]. Order SACOGLOSSA Family PLAKOBRANCHIDAE Elysia bayeri Marcus, 1965; Johnson and Boucher, 1983: 257; [Carlson and Hoff, 1978: 91, Fig. 4]. . marginata (Pease, 1871); Johnson and Boucher, 1983: 257; [Carlson and Hoff, 1978: 102, Fig. 14]. . halimedae MacNae, 1954; Johnson and Boucher, 1983: 257; [Carlson and Hoff, 1978: 99, Fig. 12]. . livida Baba, 1955; Johnson and Boucher, 1983: 257; [Carlson and Hoff, 1978: 100, Fig. 13]. . mercieri (Pruvot-Fol, 1930); [Carlson and Hoff, 1978: Fig. 15]. . obtusa Baba, 1938; Johnson and Boucher, 1983: 257; [Baba, 1949: 131, pl. 9, Figs. 28, 29]. . ratna Marcus, 1965; Johnson and Boucher, 1983: 258; [Carlson and Hoff, 1978: 107, Fig. 18]. . vatae Risbec, 1928; Johnson and Boucher, 1983: 258; [Carlson and Hoff, 1978: 108, Fig. 19). Plakobranchus ocellatus van Hasselt, 1824; Johnson and Boucher, 1983: 256; [Bertsch and Johnson, 1981: 20-21]. Family HERMAEIDAE Stiliger smaragdinus Baba, 1949; Johnson and Boucher, 1983: 256; [Baba, 1949: 129, pl. 7, Fig. 22]. Family CALYPHYLLIDAE Cyerce elegans Bergh, 1888; [Kay, 1979: 455). Order NOTASPIDEA Family PLEUROBRANCHIDAE Berthellina citrina (Ruppell and Leuckart, 1828); Willan, 1984: 37—40, Figs. 33-36; [Bertsch and Johnson, 1981: 26-27]. Berthella pellucida (Pease, 1860); Willan, 1984: 40-43, Figs. 6, 7, 17, 18, 29, 37-39, 44. B. martensi (Pilsbry, 1896); Willan, 1984: 43-45, 51, 52, Figs. 8-12, 21-28, 30-32, 40-42, 44. Pleurohedra haraldi Marcus and Marcus, 1970; Willan, 1984: 38-40, Figs. 1-5, 13-26, 19, 20. mmmmmmm (This table continued on next page.) MOLLUSCA 127 TABLE 1 (cont’d) Order NUDIBRANCHIA Family DORIDIDAE Doriopsis pecten (Collingwood, 1881); Young, 1967: 161; [Bertsch and Johnson, 1981: 34-35]. D. viridis Pease, 1861; Young, 1967: 160, Fig. 1. Family KENTRODORIDIDAE Jorunna alisonae Marcus, 1976; Young, 1967: 168; as Jorunna tomentosa (Cuvier, 1804). J. funebris (Kelaart, 1858); Johnson and Boucher, 1983: 260; [Marcus, 1976: 26, Figs. 20a, 20b]. Family DISCODORIDIDAE Carminodoris nodulosa (Angas, 1864); [Bertsch and Johnson, 1981: 39]. Discodoris fragilis (Alder and Hancock, 1864); Johnson and Boucher, 1983: 261; [Bertsch and Johnson, 1981: 40]. Family ROSTANGIDAE Rostanga lutescens (Bergh, 1905); Johnson and Bertsch, 1985: 405-410, Fig. 1. Family TRIPPIDAE Trippa echinata (Pease, 1860); [Kay and Young, 1969: 189-190}. T. intecta (Kelaart, 1858); [Willan and Coleman, 1984: 34, Fig. 98]. Family HALGERDIDAE Halgerda elegans Bergh, 1905; [Bergh, 1905: pl. 2, Fig. 4a]. H. wasinensis Eliot, 1904; Johnson and Boucher, 1983: 262; [Rudman, 1978b: 60, Fig. 1). Sclerodoris paliensis Bertsch and Johnson, 1982; Johnson and Boucher, 1983: 262; [Bertsch and Johnson, 1981: 45]. S. tuberculata Eliot, 1904; [Rudman, 1978b: 72, pl. 1]. Family PLATYDORIDIDAE Platydoris cruenta (Quoy and Gaimard, 1832); Johnson and Boucher, 1983: 259; [Risbec, 1928: 75, pl. 2, Fig. 7]. P. scabra (Cuvier, 1904); Johnson and Boucher, 1983: 260; [Edmunds, 1971: 354, Fig. 7]. Family CADLINIDAE Cadlinella ornatissima (Risbec, 1928); Johnson and Boucher, 1983: 276; [Baba, 1949: 145, pl. 19, Fig. 71]. Family CHROMODORIDIDAE Chromodoris cf. C. albonotata Bergh, 1874; [Bertsch and Johnson, 1981: 52]. C. albopunctatus (Garrett, 1879); Johnson and Boucher, 1983: 263; [Bertsch and Johnson, 1981: 58; as C. cf. C. imperialis (Pease, 1860)]. C. albopustulosa (Pease, 1860); Young, 1967: 168; [Bertsch and Johnson, 1981: 53]. C. aspera (Gould, 1852); Young, 1967: 168; as Chromodoris lilacina (Gould, 1852); Johnson and Boucher, 1983: 266; as C. inornata Pease, 1871; [Bertsch and Johnson, 1981: 54; as C. lilacina]. briqua Marcus and Burch, 1965; Marcus and Burch, 1965: 145-250, Figs. 22-24. colemani Rudman, 1982; [Rudman, 1982: 215, Fig. 176]. decora (Pease, 1860); Johnson and Boucher, 1983: 264; [Bertsch and Johnson, 1981: 55]. elisabethina Bergh, 1877; Johnson and Boucher, 1983: 265; [Johnson, 1982: 92]. fidelis (Kelaart, 1858); Marcus and Burch, 1965: 245; Johnson and Boucher, 1983: 265; Rudman, 1985: 276; [Rudman, 1985: 276, Fig. 12E]. galactos Rudman and Johnson, 1985; Rudman, 1985: 286-289, Figs. 12G, 26B, 27A, 28. geometrica Risbec, 1928; Young, 1967: 166; Johnson and Boucher, 1983: 265; [Risbec, 1928: 148, pl. 6, Fig. 10). C. marginata (Pease, 1860); Johnson and Boucher, 1983: 267; [Kay, 1979: 467, Fig. 150D}. C. verrieri (Crosse, 1875); [Rudman, 1985: 262, Fig. 12, 13A, 14, 15A]. C. vibrata (Pease, 1860); [Bertsch and Johnson, 1981: 56-57]. Durvilledoris lemniscata (Quoy and Gaimard, 1832); Johnson and Boucher, 1983: 273; as Thorunna clitonata (Bergh, 1905); [Rudman, 1984: 231]. Glossodoris atromarginata (Cuvier, 1804); Johnson and Boucher, 1983: 263; as Casella atromarginata; [Baba, 1949: 145, pl. 19, Fig. 71]. G. youngbleuthi (Kay and Young, 1969); [Bertsch and Johnson, 1981: 68-71; as Chromolaichma youngbleuthi]. Hypselodoris bullockii (Collingwood, 1881); [Willan and Coleman, 1984: 26, Fig. 67]. H. decorata (Risbec, 1928); Johnson and Boucher, 1983: 270; [Risbec, 1928: 151, pl. 7, Fig. 4]. H. infucata (Ruppell and Leuckart, 1828); Johnson and Boucher, 1983: 271; [Bertsch and Johnson, 1981: 62-63]. H. kayae Young, 1967; Young, 1967: 164, Fig. 8. H. maculosa (Pease, 1871); [Pease, 1871: 16]. H. mouaci (Risbec, 1930); Marcus and Burch, 1965; as H. hilaris; Johnson and Boucher, 1983: 272; [Risbec, 1930: 278, pl. 1, Fig. 21. Miamira sinuata (van Hasselt, 1824); [Bertsch and Johnson, 1981: 67]. Noumea decussata Risbec, 1928; Johnson and Boucher, 1983: 273; as Thorunna decussata; [Risbec, 1928: 167, pl. 8, Fig. 6]. N. flava (Eliot, 1904); [Eliot, 1904: 399, pl. 24, Figs. 8, 9]. (This table continued on next page.) 99Q 99990 128 KAY AND JOHNSON TABLE 1 (cont’d) Family CHROMODORIDIDAE (cont'd) N. norba Marcus and Marcus, 1970; Johnson and Boucher, 1983: 275; as Thorunna norba; [Marcus and Marcus, 1970: 161, Fig. 19]. N. pusilla (Bergh, 1874); [Bergh, 1874: pl. 1, Fig. 18]. Risbecia imperialis (Pease, 1860); [Bertsch and Johnson, 1981: 50-51; as Chromodoris godeffroyana (Bergh, 1879). R. tryoni (Garrett, 1873); Johnson and Boucher, 1983: 268; (Garrett, 1873: 232, pl. 4]. Thorunna australis (Risbec, 1928); Johnson and Boucher, 1983: 270; as Hypeselodoris australis; [Johnson, 1982: 92; as Hypselodoris australis). T. daniellae (Kay and Young, 1969); [Kay and Young, 1969: 207, Fig. 48]. T. furtiva Bergh, 1874; [Rudman, 1984: 277]. T. purpuropedis Rudman and Johnson, 1985; Rudman, 1985: 289-291, Figs. 12H, 27, 29, 30. Family HEXABRANCHIDAE Hexabranchus sanguineus (Ruppell and Leuckart, 1828); Marcus, 1965: 271; as H. marginatus (Quoy and Gaimard, 1832); Johnson and Boucher, 1983: 259; [Bertsch and Johnson, 1981: cover]. Family DENDRODORIDIDAE Dendrodoris coronata Kay and Young, 1969; Johnson and Boucher, 1983: 276; [Kay and Young, 1969: 218, Fig. 70). D. elongata Baba, 1936; Johnson and Boucher, 1983: 276; [Baba, 1949: 156, pl. 28, Fig. 104). D. erubescens (Bergh, 1905); Marcus and Burch, 1965: 250. D. nigra (Stimpson, 1855); Marcus and Burch, 1965: 250; Young, 1967: 168; Johnson and Boucher, 1983: 277; [Bertsch and Johnson, 1981: 72]. D. rubra (Kelaart, 1858); [Orr, 1981: 39]. D. tuberculosa (Quoy and Gaimard, 1932); [Kay and Young, 1969: 219, Fig. 72]. Family PHYLLIDIIDAE Phyllidia pustulosa Cuvier, 1804; Johnson and Boucher, 1983: 282; [Bertsch and Johnson, 1981: 74-75]. P. varicosa Lamarck, 1801; Johnson and Boucher, 1983: 282; [Bertsch and Johnson, 1981: 74-75]. Family GYMNODORIDIDAE Gymnodoris ceylonica (Kelaart, 1858); Johnson and Boucher, 1983: 277; [Risbec, 1928: 182, pl. 7, Fig. 11]. G. citrina (Bergh, 1877); Marcus and Burch, 1965: 245; as Gymnodoris bicolor (Alder and Hancock, 1864); Young, 1967: 169; Johnson and Boucher, 1983: 278; [Baba, 1949: 40, pl. 11, Figs. 37, 38]. G. okinawae Baba, 1936; Johnson and Boucher, 1983: 278; [Bertsch and Johnson, 1981: 79]. G. plebeia (Bergh, 1877); Johnson and Bouclier, 1983: 279; [Kay and Young, 1969: 224, Fig. 76]. G. striata (Eliot, 1908); Johnson and Boucher, 1983: 279; [Johnson, 1982: 93; as Analogium striatum]. Nembrotha cristata Bergh, 1877; [Willan and Coleman, 1984: 12, Fig. 9]. Roboastra gracilis (Bergh, 1877); [Baba, 1949: 136, pl. 13, Fig. 45]. Family TRIOPHIDAE Plocamopherus ceylonicus (Kelaart, 1858); Johnson and Boucher, 1983: 281; [Thompson, 1975: 509, pl. la]. Family NOTODORIDIDAE Aegires citrinus Privot-Fol, 1930; [Risbec, 1953: 60). A. punctilucens (d’Orbigny, 1837); [Baba, 1974: Fig. 1]. A. villosus Farran, 1905; Johnson and Boucher, 1983: 281; [Edmunds, 1971: 379, Fig. 18]. Family GONIODORIDIDAE Goniodoridiella savignyi Pruvot-Fol, 1933; Johnson and Boucher, 1983: 282; [Baba, 1960: 301, pl. 8}. Goniodoris joubini Risbec, 1928; Johnson and Boucher, 1983: 282; [Risbec, 1928: 175, pl. 5, Fig. 2]. Family DORIDOMORPHIDAE Doridomorpha gardineri Eliot, 1906; [Rudman, 1982: Fig. 28a]. Family VAYSSIEREIDAE Okadaia elegans Baba, 1930; Young, 1967: 171; 1969. Family EMBLETONIIDAE Embletonia gracilis Risbec, 1928; [Gosliner and Griffiths, 1981: 142-148). Family ARMINIDAE Dermatobranchus fortunata (Bergh, 1888); Johnson and Boucher, 1983: 282; [Bergh, 1888: 353, pl. 10). Family BORNELLIDAE Bornella anguilla Johnson, 1985; Johnson, 1985: 17—26; [Willan and Coleman, 1984: 50, Fig. 162). B. stellifer (Adams and Reeve in Adams, 1848); Johnson and Boucher, 1983: 282; as Bornella adamsii Gray, 1850; (Willan and Coleman, 1984: 50, Fig. 161; as Bornella adamsii Gray, 1850.] Family MARIANINIDAE Marianina rosea (Pruvot-Fol, 1930); Johnson and Boucher, 1983: 283; [Carlson and Hoff, 1973: 172; as Aranucus bifidus Odhner, 1936)). (This table continued on next page.) MOLLUSCA TABLE 1 (cont'd) Family COTHONIDAE Phestilla lugubris (Bergh, 1870); Johnson and Boucher, 1983: 284; [Bertsch and Johnson, 1981: 90-91; as Phestilla sibogae Bergh, 1905}. Family CORYPHELLIDAE Flabellina alisonae Gosliner, 1980; Gosliner, 1980: 40; Johnson and Boucher, 1983: 283; [Bertsch and Johnson, 1981: 88]. Family GLAUCIDAE Glaucus atlanticus Forster, 1777; Johnson and Boucher, 1983: 285; [Bertsch and Johnson, 1981: 100). Family AEOLIDIIDAE Baeolidia nodosa (Haefelfinger and Stamm, 1959); [Gosliner, 1980: 66-69]. Cerberilla affinis Bergh, 1888; [McDonald and Nybakken, 1975: 380-381]. Favorinus japonicus Baba, 1949; Johnson and Boucher, 1983: 283; [Bertsch and Johnson, 1981: 96-97]. F. mirabilis Baba, 1955; [Baba, 1955: 53, pl. 17, Fig. 46]. Herviella claror Burn, 1963; Marcus and Burch, 1965: 251; Figs. 28-30; Young, 1967: 172. H. mietta Marcus and Burch, 1965; Marcus and Burch, 1965: 251; Young, 1967: 171. Phidiana bourailli (Risbec, 1928); Johnson and Boucher, 1983: 285; [Rudman, 1980: 146, Fig. 1B}. P. indica (Bergh, 1896); Johnson and Boucher, 1983: 285; [Bertsch and Johnson, 1981: 95; as Caloria militaris (Alder and Hancock, 1864)). Phyllodesmium hyalinum Ehrenberg, 1831; Johnson and Boucher, 1983: 284; [Rudman, 1981: 22, Fig. 8]. Pteraeolidia ianthina (Angas, 1864); Johnson and Boucher, 1983: 284-285; [Bertsch and Johnson, 1981: 98-99]. Spurilla major (Eliot, 1903); Johnson and Boucher, 1983: 285; [Bertsch and Johnson, 1981: 89]. Subclass PULMONATA Order BASOMMATOPHORA Family MELAMPIDAE Melampus flavus (Gmelin, 1791); [Cernohorsky, 1972: 213, pl. 61, Fig. 1). Family SIPHONARIIDAE Siphonaria guamensis (Quoy and Gaimard, 1833); Marcus and Burch, 1965: 256; [Hubendick, 1946: 41, pl. 6, Figs. 30-32]. S. normalis (Gould, 1846); Menge, 1973; Leviten, 1974; [Cernohorsky, 1972: 210, pl. 60, Fig. 2]. Order ORTHURETHRA Family ACHATINELLIDAE Lamellidea pusilla (Gould, 1847). Family PUPILLIDAE Gastrocopta pediculus (Shuttleworth, 1852); Reigle, 1964: 128. G. servilis (Gould, 1843). Order SIGMURETHRA Family SUBULINIDAE Subulina octona (Bruguiere, 1792). Lamellaxis gracilis (Hutton, 1834); Reigle, 1964: 128. Suborder AULACOPODA Family CHAROPIDAE +Vatusila eniwetokensis (Ladd, 1958); Ladd, 1958: 190, pl. 30, Figs. 9-12; Miocene g; as Ptychodon; Solem, 1982: 195, Figs. 85e-f. Class BIVALVIA Family ARCIDAE Anadara antiquata (Linnaeus, 1758); [Cernohorsky, 1972: 215, pl. 61, Fig. 3). Arca avellana Lamarck, 1819; [Kilburn, 1983: 514, Figs. 2, 3]. A. ventricosa Lamarck, 1819; [Kay, 1979: 500, Figs. 161C—D]. Barbatia amygdalumtostum (Réding, 1798); [Kira, 1962: 122, pl. 43, Fig. 14; as Barbatia bicolorata (Dillwyn)}. . decussata (Sowerby, 1833); [Kay, 1979: 500, Figs. 162G—H]. . divaricata (Sowerby, 1833); [Kay, 1979: 501, Figs. 161E—G). . nuttingi Dall, Bartsch, and Rehder, 1938; [Kay, 1979: 503, Figs. 162I-—J}. . tenella (Reeve, 1844); [Kay, 1979: 504, Figs. 162C—D). . cf. B. wendti (Lamy, 1907); Fig. 6A. Family GLYCYMERIDIDAE Glycymeris reevei (Mayer, 1868); Fig. 6B. Family MYTILIDAE Botula silicula (Lamarck, 1819); [Kay, 1979: 509; Figs. 164E-—F]. Lithophaga zittelliana Dunker, 1860; Fig. 6E. Donnnwn 129 +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) 130 KAY AND JOHNSON TABLE 1 (cont’d) Family MYTILIDAE (cont’d) Mytilus edulis Linnaeus, 1758; on floating plastic fishing floats; [Abbott, 1954: pl. 35, Fig. Mj. Modiolus auriculatus Krauss, 1848; [Salvat and Rives, 1975: 367, Fig. 404). M. philippinarum (Hanley, 1843); [Kira, 1962: 126, pl. 46, Fig. 18}. M. vagina (Lamarck, 1819); Fig. 6C. Septifer bryanae (Pilsbry, 1929); [Kay, 1979: 512, Fig. 164P]. Family PINNIDAE Atrina vexillum (Born, 1778); [Rosewater, 1961: pls. 156, 157]. Pinna muricata Linnaeus, 1758; Rosewater, 1961:191, pl. 141). Streptopinna saccata (Linnaeus, 1758); Rosewater, 1961: 22, pl. 169. Family PTERIIDAE Electroma electrina (Reeve, 1857); Fig. 6D. Pinctada margaritfera (Linnaeus, 1758); [Kay, 1979: 516, Figs. 166D-—E]. Pteria penguin (Roding, 1798); [Cernohorsky, 1978: 177, pl. 62, Fig. 1]. Family ISOGNOMONIDAE Isognomon dentifera Krauss, 1848; [Salvat and Rives, 1975: 370, Fig. 415; as I. pectinata (Reeve, 1858)]. I. incisum (Conrad, 1837); [Kay, 1979: 520, Figs. 167C-—D (not E-F)]. I. legumen (Gmelin, 1791); [Kay, 1979: 520, Figs. 167G—H]. I. perna (Linnaeus, 1758); [Kay, 1979: 521, Figs. 167C—D]. Melina ephippium (Linnaeus, 1758); [Cernohorsky, 1978: 178, pl. 63, Fig. 2]. Family MALLEIDAE Malleus regula (Forsskal, 1775); [Kay, 1979: 521, Figs. 167I-K]. Family PECTINIDAE Chlamys coruscans (Hinds, 1845); Waller, 1972: 231, pl. 1, Figs. 1-7, 12]. C. wilhelmini Bavay, 1904; Waller, 1972: 236, pl. 2, Figs. 30-33, 36-37; as C. marshallensis Waller, 1972. Comptopallium vexillum (Reeve, 1853); Waller, 1972: 243, pl. 4, Figs. 64-75. tExcellichlamys spectabilis (Reeve, 1853); Waller, 1972: 249; Holocene; [Waller, 1972: 246, pl. 5, Figs. 85-92]. Gloripallium pallium (Linnaeus, 1758); Waller, 1972: 239, pl. 3, Figs. 45-56. Juxtamusium coudeini (Bavay, 1902); Waller, 1972: 250, Figs. 111-127; as Juxtamusium maldviensis (E. A. Smith, 1903). Mirapecten mirificus (Reeve, 1853); [Kay, 1979: 526, Figs. 169A-B]. Pedum spondyloideum (Gmelin, 1791); Waller, 1972: 254, pl. 8, Figs. 136-143. Family SPONDYLIDAE Spondylus hystrix Bolten in Roding, 1798; [Kay, 1979: 530, Figs. 176G—H; and see S. nicobaricus Schreiberg, 1795 in Cernohorsky, 1978: 180, pl. 63, Fig. 4]. S. linguaefelis Sowerby, 1874; [Kay, 1979: 531, Figs. 171D-E]. S. squamosus Schreiber, 1793; [Hirase, 1951: pl. 15, Fig. 2]. S. tenebrosus Reeve, 1856; [Kay, 1979: 532, Figs. 171A-C]. S. cf. S. varians Sowerby, 1829; [Cernohorsky, 1978: 180, pl. 64, Fig. 3]. Family PLICATULIDAE Plicatula simplex Gould, 1861; [Johnson, 1964: 149, pl. 25, Fig. 4]. Family LIMIDAE Lima annulata Lamarck, 1819; [Cernohorsky, 1972: 220, pl. 62, Fig. 2]. L. concentrica Sowerby, 1888; [Cernohorsky, 1978: 181, pl. 65, Fig. 1]. L. fragilis (Gmelin, 1791); [Kay, 1979: 534, Figs. 172A-B). L. lima vulgaris (Link, 1807); [Cernohorsky, 1972: 219, pl. 61, Fig. 5]. Family OSTREIDAE Ostrea cf. O. hanleyana Sowerby, 1871; [Kay, 1979: 537, Figs. 173D-E]. O. cf. O. sandvicensis Sowerby, 1871; [Kay, 1979: 539, Figs. 173A-B]. Family CHAMIDAE Chama broderipii Reeve, 1846; [Reeve, 1846: Chama No. 2]. C. iostoma Conrad, 1837; [Cernohorsky, 1978: 183, pl. 65, Fig. 5). C. lazarus Linnaeus, 1758; [Cernohorsky, 1978: 183, pl. 65, Fig. 3]. C. pacifica Broderip, 1835; Fig. 6M. Family LUCINIDAE Codakia punctata (Linnaeus, 1758); [Cernohorsky, 1972: 221, pl. 62, Fig. 6]. Ctena bella (Conrad, 1837); [Kay, 1979: 543, Figs. 176A-B). Lucina edentula (Linnaeus, 1758); [Kay, 1979: 543, Figs. 177A-B]. L. cf. L. erythraea Issel, 1869; [Bouchet and Danrigal, 1982: Fig. 20]. +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) MOLLUSCA 131 TABLE 1 (cont'd) Family LUCINIDAE (cont’d) L. cf. L. lorisi Viader, 1937; Fig. 6K. Family FIMBRIIDAE Fimbria fimbriata (Linnaeus, 1758); Nicol, 1950: 83, Figs. 1, 2, 4, 6, 7. Family LASAEIDAE Nesobornia bartschi Chavan, 1969; [Kay, 1979: 549, Fig. 179G}. Family GALEOMMATIDAE Scintilla hiloa Dall, Bartsch and Rehder, 1938; [Kay, 1979: 550, Fig. 1791]. Family SPORTELLIDAE Hitia cf. H. ovalis Dall, Bartsch, and Rehder, 1938; (Kay, 1979: 552, Figs. 179E, F). Family CARDIIDAE Acrosterigma orbita (Broderip and Sowerby, 1838); [Kay, 1979: 556, Figs. 180F—G; as Trachycardium]. Corculum cardissa (Linnaeus, 1758); [Cernohorsky, 1972: 225, pl. 65, Fig. 2]. Discors lyratum (Sowerby, 1841); [Cernohorsky, 1972: 225, pl. 63, Fig. 2]. Fragum fragum (Linnaeus, 1758); [Cernohorsky, 1972: 223, pl. 63, Fig. 5). F. loochooanum Kira, 1959; [Kira, 1962: 154, pl. 55, Fig. 13). Laevicardium biradiatum (Bruguiere, 1789); [Cernohorsky, 1972: 224, pl. 63, Fig. 6]. L. pulcherrima Sakurai and Habe, 1966; Fig. 6H. Microfragum festivum (Deshayes, 1855); [Kira, 1962: 154, pl. 55, Fig. 10). Family MACTRIDAE Mactra sulcataria Reeve 1854; [Hirase, 1951: pl. 50, Fig. 7]. Family MESODESMATIDAE Atactodea glabrata (Gmelin, 1791); [Kira, 1962: 166, pl. 58, Fig. 33; as Actactodea striata (Gmelin). Ervilia (Spondervillia) bisculpta (Gould, 1861); [Kay, 1979: 558, Figs. 181E-F (not Figs. 181C-—D)}. Sponderuillia cf. S. simplex Laseron, 1953; Fig. 6d. Family TELLINIDAE Macoma dispar (Conrad, 1837); [Kay, 1979: 559, Figs. 182I-J]. M. obliquilineata (Conrad, 1837); [Kay, 1979: 559, Figs. 182 G—H]. M. obliquaria (Deshayes, 1854); Fig. 6L. Pinquitellina robusta (Hanley, 1844); [Kay, 1979: 561, Figs. 182C-—D). Tellina crucigera Lamarck, 1818; [Kay, 1979: 564, Figs. 183A-B]. . gargadia Linnaeus, 1758; [Afshar, 1969: 40, pl. 11, Figs. 6-10). . cf. T. inflata Gmelin, 1791; Fig. 61. cf. T. ovalis Sowerby, 1825; Fig. 6G. . linguaefelis Linnaeus, 1758; [Cernohorsky, 1972: 229, pl. 65, Fig. 4]. . palatam Iredale, 1929; [Kay, 1979: 563, Figs. 182E-F]. perna (Spengler, 1798); [Kay, 1979: 563, Figs. 183E-F]. . pudica Hanley, 1844; Fig. 6F. . scobinata Linnaeus, 1758; [Cernohorsky, 1972: 229, pl. 65, Fig. 5). . staurella (Lamarck, 1818); [Cernohorsky, 1972: 228, pl. 7, Fig. 4]. semen Hanley, 1844; [Boss, 1969: 136, pl. 15, Figs. 1—4]. . substruncata Hanley, 1845; [Habe, 1964: 201, pl. 62, Fig. 8]. . virgata Linnaeus, 1758; [Cernohorsky, 1972: 228, pl. 7, Fig. 5]. Family PSAMMOBIIDAE Asaphis violascens (Forsskal, 1775); [Cernohorsky, 1972: 231, pl. 66, Fig. 7]. Gari maculosa (Lamarck, 1818); [Cernohorsky, 1972: 231, pl. 66, Fig. 3]. G. squamosa (Lamarck, 1818); [Cernohorsky, 1972: 231, pl. 66, Fig. 4]. Family SEMELIDAE Semele australis (Sowerby, 1833); [Kay, 1979: 566, Figs. 183M—N]. Semelangulus crebrimaculatus Sowerby, 1867; [Kay, 1979: 565, Figs. 183J—K]. S. cf. S. tokubei Habe, 1961; [Habe, 1964: 119, pl. 61, Fig. 19]. Family TRAPEZIIDAE Trapezium bicarinatum (Schumacher, 1817); [Cernohorsky, 1972: 232, pl. 68, Fig. 4]. T. oblongum (Linnaeus, 1758); [Kay, 1979: 566, Figs. 184E-F]. Family TRIDACNIDAE Hippopus hippopus (Linnaeus, 1758); Rosewater, 1965: 361, pl. 267, Figs. 1, 3; Fankboner and Renaud, 1971. + Tridacna crocea Lamarck, 1819; Rosewater, 1965: 393; Pleistocene?; [Rosewater, 1965: 392, pl. 292). T. gigas (Linnaeus, 1758); Rosewater, 1965: 369, pl. 267, Fig. 2. T. maxima (Roding, 1798); Rosewater, 1965: 384, pl. 267, Figs. 9-12. T. squamosa Lamarck, 1819; Rosewater 1965: 380, pl. 267, Figs. 4-7. SANNA +Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). (This table continued on next page.) 132 KAY AND JOHNSON TABLE 1 (cont'd) Family GLOSSIDAE Meiocardia tetragona (Adams and Reeve, 1848); [Kira, 1962: 148, pl. 53, Fig. 24]. Family VENERIDAE Dosinia cf. D. miticula Viader, 1937; Fig. 6N. Gafrarium pectinatum (Linnaeus, 1758); [Cernohorsky, 1972: 234, pl. 67, Fig. 1]. G. tumidum (Roding, 1798); [Cernohorsky, 1972: 234, pl. 67, Fig. 2]. Glycodonta marica (Linnaeus, 1758); [Cernohorsky, 1972: 237, pl. 67, Fig. 5]. Lioconcha castrensis (Linnaeus, 1758); [Cernohorsky, 1972: 235, pl. 5, Fig. 7]. L. ornata (Dillwyn, 1817); [Cernohorsky, 1972: 236, pl. 67, Fig. 6]. Periglypta puerpera (Linnaeus, 1771); [Cernohorsky, 1972: 233, pl. 66, Fig. 5). P. reticulata (Linnaeus, 1758); [Cernohorsky, 1972: 233, pl. 66, Fig. 6). Pitar pellucidus (Lamarck, 1818); [Cernohorsky, 1972: 236, pl. 68, Fig. 2]. Venus toreuma Gould, 1850; [Kay, 1979: 570, Fig. 184L]. Family GASTROCHAENIDAE Gastrochaena cuneiformis Spengler, 1783; [Kay, 1979: 570, Fig. 185A). Family PHOLADIDAE Martesia striata (Linnaeus, 1758); [Kay, 1979: 573, Fig. 185C]. Class POLYPLACOPHORA Order CHITONID Family ACANTHOCHITONIDAE Cryptoplax elioti Pilsbry, 1901. Family CHITONIDAE tLuculina russelli Ladd, 1966; Ladd, 1966: 23, pl. 1, Figs. 13-15; probably Pliocene. Family ISCHNOCHITONIDAE Ischnochiton sp. Family SCHIZOCHITONIDAE {Schizochiton marshallensis Ladd, 1966; Ladd, 1966: 22, pl. 1, Figs. 4-9; early Miocene. Family LEPIDOPLEURIDAE Leptochiton acuminatus (Thiele, 1909). Class CEPHALOPODA Order OCTOPODA Family OCTOPODIDAE Octopus cyanea Gray, 1849; [Kay, 1979: 589, Figs. 194B-E]. Order TEUTHOIDEA Family OMMASTREPHIDAE Hyaloteuthis pelagica (Bosc, 1802); Voss, 1954: 365. Subclass NAUTILOIDEA Family NAUTILIDAE Nautilus macromphalus Sowerby, 1849; Shells occasionally washed up on windward beaches; [Cernohorsky, 1972: 240, pl. 7, Fig. 8]. N. pompilius Linnaeus, 1758; Shells occasionally washed up on windward beaches; [Cernohorsky, 1972: 240, pl. 7 big? 7]! {Species not now living at Enewetak; horizons recorded by Ladd (1966, 1972, 1977, 1982). 1959); Cassididae (Abbott, 1968); Mitridae (Cernohorsky, 1976); Drupa (Emerson and Cernohorsky, 1973); Rhinoclavis_ (Houbrick, 1978); Clypeomorus (Houbrick, 1985); Harpidae (Rehder, 1973); Gabrielona (Robertson, 1973); Pinnidae (Rosewater, 1961); Tridacnidae (Rosewa- ter, 1965); and Littorinidae (Rosewater, 1970). Collections of opisthobranchs made by J. B. Burch and W. Heard were described by Marcus and Burch (1965) and Marcus (1965). Young (1967, 1969) reported on his own collec- tion of opisthobranchs, and Johnson and Boucher (1983) recorded 76 species from collections they made at Enewetak. Enewetak pectinids were discussed by Waller (1972). A paper by Ingram (1947) and a manuscript by Ingram and Morrison (MS, USNM) discuss the cowries of the Marshall Islands and include records from Enewetak. Most studies of Enewetak mollusks have been ecologi- cally oriented. As early as 1956, Kohn made collections and observations on the gastropod genus Conus, and his work on Conus (and that of his students) continues (Kohn, 1967, 1971, 1980a, 1980b; Kohn and Leviten, 1976; Leviten and Kohn, 1980; Leviten, 1974). Ecological stud- ies on other groups include those on the Tridacnidae (Stasek, 1963a, 1963b; Fankboner, 1971a, 1971b, 1972; Leviten and Osenberg, 1979); Terebridae (Miller, 1966); Nerita (Stokes, 1966); Muricidae (Menge, 1974; Bernstein, 1974); Cypraea moneta (Renaud, 1977); Strombidae MOLLUSCA (Berg, 1974); Vermetidae (Hopper, 1975, 1978); Morula (Sinclair, 1975); Stylocheilus (Sarver, 1978); Phestilla (Switzer, 1971); Siphonaria (Menge, 1973); and soft bot- tom communities (Ekdale et al., 1979). Enewetak has also been the site of research on the genetic variability and the physiology of the Tridacnidae (Valentine and Ayala, 1974; Ayala et al., 1973; Valentine et al., 1973; Fankboner, 1971a, 1971b, 1974), of physio- logical studies of the Strombidae (Gillary, 1974; Géillary and Gillary, 1979), and of studies of growth and mortality rates in marine snails (Turk, 1979a, 1979b). MODERN MOLLUSCAN FAUNA Nonmarine Mollusks The nonmarine molluscan fauna, comprised of eight species, conforms to the description of C. M. Cooke, Jr. (in Harry, 1966) of a typical atoll land snail fauna: “ very restricted. . . made up mostly of species that are accidentally distributed by man.” Thus, of the eight species we record, five (both Gastrocopta, both Subulinidae, and Lamellidea pusilla) are probably adventive and widely dis- tributed among the islands of the Pacific as a result of the activities of man (see Christensen and Kirch, 1981; Harry, 1966; and Solem, 1964 for discussion of Pacific basin land snails and their dispersal by humans). The remaining three nonmarine molluscan species are strandline species: Melampus flavus and Truncatella guernii are widely distrib- uted on Pacific islands; Assiminea nitida marshallensis is endemic to the Marshall Islands, according to Abbott (1958). The list of Enewetak nonmarine mollusks is similar 133 to that compiled for neighboring Rongelap in the Marshall Islands (Reigle, 1964). Of the 11 species recorded from Rongelap, three of the five adventive species also occur at Enewetak, and the two atolls also share two of the strand- line species. Marine Mollusks At least three factors affect the records of the presence or absence of marine mollusks at Enewetak and, indeed, on any atoll. The three factors are the collecting effort, kinds of habitat available, and the chance occurrence or recruitment of young to the island. Given 40 years of collection activity and the numerous collectors who have visited Enewetak, the approximately 1000 species of marine mollusks that are recorded as occurring at Enewetak today are probably fairly representa- tive of the molluscan fauna that occurs between the inter- tidal zone and depths of about 30 m. In terms of numbers of species, the Enewetak molluscan fauna is about ¥% that of the island of Okinawa, 2500 miles northwest; just slightly larger than that recorded for the Hawaiian Islands, 2000 miles northeast; and two and a half to three times larger than those reported for two Pacific atolls, Fanning and Funafuti (Table 2). The fauna is dominated by gastropods, and the gastropod:bivalve ratio of 90:10 is the highest of all Pacific faunas for which there are records (Table 2). The gastro- pod record includes 947 species in 109 families. Neogas- tropods comprise 44% of the gastropod species, mesogas- tropods (including Heterogastropoda), 32%; euthyneurans, 17%; and archaeogastropods, about 8% of the mollusks. TABLE 2 Characteristics of Pacific Island Marine Molluscan Faunas Taxonomic Enewetak Okinawa* Hawaiit Funafuti} Fanning! Group No. % No. % No. % No. % No. % Archaeogastropods 76 8 7s) 1183 50 6 SOL 16 5 Mesogastropods S23moe 472 34 292) ODL Sa a Sia Oe Si Neogastropods 421 44 611 44 344 40 106 33 O8mrsS Euthyneura ees a 149 8 W7il 7X0) 34 8 33 Total 994 1411 857 329 295 Gastropods Bivalves 115 426 141 80 52 Cephalopods 4 n.a 5 3 Polyplacophora 3 16 4 1 1 Total mollusk species 1116 1853 1007 413 348 Gastropod:Bivalve Ratio 90:10 77:23 86:14 80:20 85:15 “Kuroda (1960). +Kay (1979). +Hedley (1899). Kay and Switzer (1974). 134 KAY AND JOHNSON The families with the greatest number of species are the Turridae (78), Conidae (65), Mitridae (58), Cypraeidae (55), and the Costellariidae (51). The bivalve record includes 30 families and 115 species. Among the bivalves, 51% of the species are infau- nal, 41% are epifaunal byssate forms, and 6% are free-living species. The family Tellinidae has the largest number of species (17), followed by the Veneridae (10), Arcidae (9), and Cardiidae (8). Composition There are some noticeable anomalies in the composi- tion of the Enewetak fauna in that several well-known Indo-West Pacific mollusks are conspicuous by their absence. We do not record the turbinid Leptothyra picta, the neritid Smaragdia, the rissoid Merelina pisinna, the ver- metid Dendropoma maxima, the mesogastropod limpet Hipponix, the thaid Vexilla, the buccinid Cantharus fumosus, or the heterogastropod Architectonica. Several species such as Gyrineum gyrineum and Morum denisoni which are recorded from Kwajalein are absent at Enewetak. Some of the absences are explicable. If a habitat is absent, then the species occurring in that habitat are also absent. Thus Smaragdia associated with sea grass beds and gastropods associated with volcanic substrates such as Hipponix and the muricid Vexilla (Vermeij et al., 1984) are not expected to occur at Enewetak. Other more sub- tle ecological restrictions have been suggested by Waller (1972) who has shown that several species of Enewetak Pectinidae are extremely restricted in their habitats. These species include Comptopallium vexillum which lives in asso- ciation with terrigenous sediments and marine grasses around elevated islands and is found only as juveniles in the deeper portions of the lagoon of the atoll; and Juxtamusium maldivense, which also lives in the deeper portions of the lagoon, in association with abundant Halimeda on which it may depend for attachment and con- cealment. The absence of gastropods such as the reef-associated Leptothyra picta, Merelina pisinna, Dendropoma maxima, and Architectonica is not easily explained and, for the present, is attributed to chance as has been suggested for certain aspects of the marine fauna of the Line Islands (Kay, 1971), the Hawaiian Islands (Kay, 1979, 1980), the northern Marianas (Vermeij et al., 1984), and among the islands of the Indian Ocean (Taylor, 1971). Biogeography With one exception, the subspecies Assiminea nitida marshallensis Abbott, 1958, all the Recent mollusks identi- fied to species have been recorded elsewhere in the Indo- West Pacific. As Waller (1972) has noted, the significance of the latter observation lies in the demonstration that, for Pacific island mollusks, miles of open water without strong current systems have not resulted in much differentiation. The marine mollusks of Enewetak are clearly of Indo- West Pacific derivation, and many species have ranges which span the Indo-West Pacific from the Hawaiian Islands on the east to the coast of east Africa on the west. A component of this fauna is restricted to the Pacific Plate (sensu Springer, 1982): 14 of the 56 cowries and five of the 19 strombids are Pacific Plate endemics. Four species require special mention. Shells of two species of Nautilus and specimens of Mytilus edulis are reported from windward beaches, but there is no evidence that either Nautilus or Mytilus lives at Enewetak. One species, Trochus niloticus, was introduced by the Japanese about 1935, and these mollusks are very common. FOSSIL MOLLUSKS Fossil Record The Enewetak molluscan fossil record was documented by Ladd (1966, 1972, 1977, 1982) who described and/or recorded 274 species primarily from two drill cores on the atoll. Ladd dealt only with chitons and gastropods and, unfortunately, was unable to complete his study of the Enewetak fossil material. Of the gastropods recorded by Ladd (1966 et seq.), archaeogastropods comprise 25%; mesogastropods (including Heterogastropoda), 56%; and neogastropods, 17% of the fossil record. Although relatively numerous in terms of species, the Enewetak fossil record is far from complete. Apart from the euthyneurans, bivalves, cephalopods, and scaphopods which have not been considered, the neogastropods are underrepresented, and there are biases in other respects. There are proportionately more lagoonal forms such as ris- soids and cerithiids than there are seaward reef flat species such as cones and muricids. High intertidal littorinids and neritids are not well represented. Four of the six fossil ne- ritids are subtidal forms and only one littorinid (Tectarius) is recorded. Still another bias is in size—82% of the mea- surements for shell sizes are less than 10 mm in greatest dimension and the largest fossil shell is only 34 mm in length. Despite these deficiencies, the fossil record, together with the current checklist, allows at least three conclusions: 1. Species composition at Enewetak has changed with time. 2. Many species were widely distributed in the Pacific in the Miocene. 3. Previously wide species ranges have become re- stricted with time. Geological Record The Enewetak geological record comprises a Tertiary sequence extending back into the Eocene and a Quater- nary surface limestone section. Ladd (1966) recognized six subdivisions of the Tertiary, the late Eocene (Tertiary b) to the Pliocene (Tertiary h), based on the system developed for the East Indies by van der Vlerk and Umbgrove (1927) MOLLUSCA 135 Fig. 1 a, Diodora foveolata, length 10 mm; b-c, Emarginula concinna, length 6.75 mm; d, E. nigromaculata, length 10 mm; e, Calliostoma sp., height 8 mm; f-g, Orbitestella sp., diameter 1.5 mm; h-i, Teinostoma sp., diameter 1.5 mm; j, Astele engebienesis, height 3 mm; k, Rufodardanula sp., height 1.5 mm; 1, Alvania crystallina, height 3 mm; m, Brookula sp., height 1.25 mm; n-o, Puperita bensoni, height 6 mm; p, Sansonia andamanica, height 2 mm; q, Marelepetopoma sp., height 1.5 mm; r, Microliotia dautzenbergi, diameter 1.5 mm. 136 KAY AND JOHNSON Fig. 2. a, Planaxis zonatus, height 7 mm; b, Caecum cf. C. folini, length 2.5 mm; c, Turritella sp. A., height 18 mm; d, Turritella sp. B., height 17 mm; e, Scaliola arenosa, height 3 mm; f, S. caledonica, height 2 mm; g, S. glareosa, height 3 mm; h, Inella japonica, height 3 mm; i, Nanophora cf. N. leucomys, height 2.5 mm; j, Bouchetriphora quadrimaculata, height 3 mm; k, Vermicularia sp., height 10 mm; |, Cerithium claviforme, height 25 mm; m, Metaxia sp., height 3 mm; n, Cerithiopsis sp. A., height 4 mm; o, Cerithiopsis sp. B., height 4 mm; p, Euthymella regalis, height 8 mm; q, Mastonia ustulata, height 3 mm; r, Mesophora cnodax, height 5 mm; s, Nanophora sp., height 3 mm. MOLLUSCA Fig. 3 a, Stylapex sp., height 3.5 mm; b, Balcis acicula, height 8 mm; c, Hemileiostracum sp., height 2 mm; d, Lamellaria sp., length 8 mm; e, Epitonium auritum, height 19 mm; f-g, Neverita sp., height 5 mm; h-i, Morula squamosa, height 22 mm; j-k, Neverita cf. N. petiveriana, height 4.5 mm; 1, Pyrene obtusa, height 12 mm; m, Euplica amirantium, height 4 mm; n, Columbella cf. C. moleculina, height 13 mm; o, Pyrene cf. P. livescens, height 10 mm; p, P. albina, height 12 mm; q, Euplica cf. E. liocyma, height 6.5 mm; r, E. cf. E. minuta, height 6 mm; s, Seminella divaricata, height 4 mm; t, S. dautzenbergi, height 4 mm; u, S. subphilo- dicia, height 4 mm; v, Pyrene punctata, height 13 mm; w, P. obesula, height 4 mm. 137 138 KAY AND JOHNSON Fig. 4 a, Granulina vitrea, height 3 mm; b, Kogomea ovata, height 4 mm; c, Kogomea sp. A., height 1.5 mm; d, Tringinella sp., height 1.5 mm; e, Haloginella micros, height 3 mm; f, H. keppelensis, height 7 mm; g, Kogomea hervieri, height 1.5 mm; h, Anacithara stricta, height 3 mm; i, A. nanisca, height 3 mm; j, A. nota- bilis, height 3 mm; k, Clavus sp. B., height 7 mm; 1, C. bilineata, height 10 mm; m, Clavus sp. A. n. Eucithara cithara, height 12 mm; 0, E. daedalea, height 6 mm; p, E. sp., height 15 mm; q, Kermia cf. K. granicostata, height 6 mm; r, Etrema trigonostoma, height 2 mm; s, Kermia concinna, height 10 mm; t, Philbertia crassilabrum, height 12 mm; u, Kermia edychroa, height 4.5 mm; v, Lienardia goubini, height 5 mm; w, Kermia idiomorpha, height 3 mm. MOLLUSCA 139 Fig. 5 a, Chemnitzia gabrieli, height 5.25 mm; b, Chrysalida alveata, height 4 mm; c, Evalea densestriata, height 12 mm; d, Pyrgulina densecostata, height 2.5 mm; e, Chrysalida alveata, 4 mm; f, Ringicula doleari, height 1.75 mm; g, Haloa japonica, height 5 mm; h, Pupa sp. 8 mm; i, Nakagamigawaia spiralis, height 4 mm; j, Adamnestia japonica, height 8 mm; k, Eocylichna soyae, height 10 mm; 1, Coleophysis minima, height 3 mm; m, Cylichnatys angustata, height 5 mm. and Leupold and van der Vlerk (1931). Mollusks are rare and poorly represented in limestones of late Eocene age (Tertiary b) that form the near-basement pavement and include only poorly preserved valves of Pecten and Arca, minute turbinids, molds of small gastropods, and one iden- tifiable mollusk, a species of Turritella related to a living species also reported from the upper Tertiary of Indonesia (Ladd, 1966). In what is questionably considered early Oli- gocene, three shells of Rissoina ailinana Ladd, 1966 and nine specimens of the naticid Ampullina berauensis described from the Miocene of East Borneo (Ladd, 1966) are represented. Above the early limestones are 610 m (2000 ft) of fossil reef of Miocene age which contain the bulk of the Enewetak fossil record, about 170 species. The post-Miocene sections consist of more than 183 m (600 ft) of consolidated limestone and about 9 m (30 ft) of uncon- solidated sediments (Ladd, 1966) from which 105 species are reported. Paleoecology Reef-associated limestones that underlie Enewetak do not record slow, continuous subsidence but rather periods 140 KAY AND JOHNSON Fig. 6 a, Barbatia wendti, length 6 mm; b, Glycymeris reevei, length 15 mm; c, Modiolus vagina, length 45 mm; d, Electroma electrina, length 10 mm; e, Lithophaga zitelleliana, length 40 mm; f, Tellina pudica, length 6 mm; g, T. ovata, length 37 mm; h, Laevicardium pulcherrimum, length 18 mm; i, Tellina inflata, length 17 mm; j, Spondervilia simplex, length 2.75 mm; k, Tellina sp., length 5 mm; 1, Macoma obliquaria, length 6 mm; m, Chama sp., length 20 mm; n, Dosinia miticula, length, 10 mm. MOLLUSCA 141 when the atoll stood above the sea as a high island. Leopold’s (1969) report of spores and pollen indicates a richer and more varied flora on Enewetak in the Miocene than now occurs. The record of a high island fossil land shell, Vatusila (Ladd, 1958 as Pytochodon; Solem, 1982), is evidence of conditions quite different from those today. Leopold (1969) also demonstrated conclusively from analysis of pollen grains that mangroves occurred at Enewetak through the Miocene. The limestone sections “"_ . are all reef associated. Many of the fossiliferous beds, especially those rich in mollusks, appear to have been deposited in lagoons, but other environments—reef wall, open shoal, and off-reef deposits—are also found” (Ladd, 1966). Miocene Record The Miocene record, the richest and most diverse of the fossil records, includes 170 species, 26% archaeogas- tropods, 58% mesogastropods (including heterogastro- pods), and 17% neogastropods, representing 90 genera and 42 families of mollusks. The Miocene fauna is very different from that found at Enewetak today. The archaeogastropods are represented by eight families; today there are 13 families. Turbinids, trochids, and neritids comprise 71% of the Miocene species record; trochids, fissurellids, stomatellids, and tur- binids represent 68% of the present archaeogastropod fauna. Among the mesogastropods, the rissoids (29 species) are the most numerous family comprising 30% of the record; the cerithiids (15 species), 19%; and vitrinellids and dialids (12 species) about 13% of the record. Today the cypraeids, cerithiids, rissoids, stombids, vitrinellids, and cymatids dominate the mesogastropods. Some of the differences between the Miocene and modern faunas undoubtedly reflect biases in the fossil record as such. Other differences indicate the presence of habitats such as sea grass beds, mangrove swamps, brack- ish water lagoons, and high island shorelines which are not now present on the atoll. Thus four genera present in the Miocene record are extinct on Enewetak today but are found elsewhere in the Indo-West Pacific in environments which do not presently occur at Enewetak. The neritid Smaragdia (three species in the Miocene record) is associ- ated with sea grass beds at Kwajalein, Marshall Islands, in the Caroline Islands, and as far east as the Hawaiian Islands (Kay, 1979). The sessile limpet-like mesogastropod Hipponix (two species in the Miocene record) is found on the volcanic shores of the Hawaiian Islands, on Guam, and in the northern Mariana Islands (Vermeij et al., 1984). The cerithiids Terebralia and Potamides (five species in the Miocene record) occur in mangrove associations in the Philippines and elsewhere in the Indo-Malayan archipelago. In all, 12 species or 7% of the Miocene fauna recorded by Ladd (1966, et seq.) are associated with habitats not now represented at Enewetak. Other extinctions may represent range restrictions for reasons other than habitat change. Nerita insculpta, Tectarius, Protobarleeia, Pugilina, and Eocithara are not recorded after the Miocene. Nerita insculpta is now found in Indonesia and the Northern Marianas but apparently not on islands to the east. Only one species of Tectarius, T. grandinatus, now occurs on the Pacific Plate, and it is re- stricted to southeast Polynesia (Rosewater, 1970). Protobarleeia myersi now survives in Fiji and Australia (Ponder, 1983b). Although the Melongenidae do occur in the Indo-West Pacific, no species of Pugilina is known from the Pacific Plate. Still other extinctions are even more puzzling. A number of species recorded from the Miocene of Enewetak survive on other Pacific islands, including Kwajalein. Among the species are Leptothyra laeta, L. candida, L. verruca, Merelina pisinna, Rissoina concinna, R. sup- racostata, Mitra turgida, and Vexillum approximatum. Post-Miocene Record The post-Miocene record consists of 105 species of mollusks: 23% archaeogastropods, 61% mesogastropods, and 16% neogastropods. Most of these mollusks are extant. Twenty-three or 22% are extinct at Enewetak today, although some survive elsewhere in the Indo-West Pacific, among them Synaptocochlea rosacea, Leptothyra picta, and Nannophora cingulifera. Faunal Relations The closest tie in the fossil record between Enewetak and another locale is with Bikini. About 34% of the species found in the drill holes at Enewetak were also found in drill holes at Bikini. Apart from the Bikini rela- tionship, the strongest tie for which there is a published record is with Fiji with 13% of the Enewetak Miocene species shared. The record is less strong with Guam and Saipan (4% of the Miocene record) and Java (3%). Several possibly far-flung relationships were cited by Ladd (1966). Among them he notes that Haplocochlias (one species in the Miocene) is recognized as a western American genus, and the turbinid Cynisca (one species in the Miocene) is known only from South Africa. Neither of these relationships has been subsequently confirmed. How- ever, the shells of Sansonia kenneyi, described from the Miocene of Enewetak, are indistinguishable from Miocene shells from Italy and Pliocene shells from France (Le Renard, personal communication). Species Survival Ladd (1966) pointed out early that many living mol- lusks in the Marshall Islands have been there since late Tertiary time. More recently Stanley (1979) and Stanley et al. (1980) commented on the “great stability of the Indo-West Pacific fauna” and suggest that species have tended to survive for unusually long periods in the region. The Enewetak fossil record together with the modern record provides strong evidence for the generalization: 62 species from the Miocene record are identified as living today at Enewetak. They represent 35% of that record. 142 KAY AND JOHNSON From the Holocene record, 65% survive today on Enewetak. Of the 140 extinct species listed by Ladd (1966, et seq.), 84% became extinct in the Miocene, 14.8% in the Holocene. The Enewetak record may be contrasted with that reported by Newman (1986) who calculated from Ladd (1982: Table 4) that of 126 extinctions since the Eocene, 56 or 44.4% became extinct in the Pleistocene and only five (3.5%) of the extant species survived from the Miocene. This last figure is too low for Enewetak and may be peculiar to the fossil record of Fiji and the New Hebrides (Vanuatu). There is no single pattern of extinction and survival such as Wells (1982) found for the Midway corals, where at the end of the Miocene there was complete replacement of the Miocene genera. At Enewetak, the Turbinidae and Rissoidae show a pattern in which equal numbers of species have been extinct since the Miocene, occur both in the Miocene and today, and appear only in the modern record. The trochid, neritid, and littorinid records, in con- trast, are almost entirely modern. ACKNOWLEDGMENTS The checklist of Enewetak mollusks (Table 1) is based on the references cited in the Introduction, the Enewetak collections in the USNM, the EMBL reference collection (now housed at the B. P. Bishop Museum, Honolulu, Hawaii), material collected for an earlier checklist by A. S. Bernstein and A. Miller which has been incorporated into the Enewetak collections at B. P. Bishop Museum, and observations by the authors. These observations are based in part on examination of private shell collections of mili- tary personnel stationed at Enewetak between 1978 and 1983. We are especially grateful to Carol Hopper, Richard Houbrick, and the late Joseph Rosewater for their contri- butions to the checklist. Numerous people have contributed to the listing through collection and/or identification of mollusks from Enewetak and by way of their assistance with the literature. Among them are A. Adams, H. Bertsch, W. O. Cernohorsky, C. Christensen, P. Colin, the late D. Devaney, A. Ferreira, R. Houbrick, J. Lamberson, J. Le Renard, V. O. Maes, W. B. Rudman, E. Vokes, and T. Waller. Lisa Boucher and Lori Bell have been of particular help in locating unrecorded species. Special thanks go to Rich Nakagawa for the use of his unpublished listing of Kwajalein shells and to Jeannete Hammon, Karen deGroff, Dave Johnson, D. J. MacDonald, Lee Rousseau, and Jim and Judy Wedge for additional information and assistance in finding Kwajalein material. The senior author is grateful to Kenneth Boss and Ruth Turner of the Museum of Comparative Zoology, Harvard University; Philippe Bouchet, Laboratoire des Invertébrés marins et de Malacologie, Paris; George Davis, Academy of Natural Science, Philadelphia; Oliver Paget and Erhard Wawra, Naturalhistoriske Museum, Vienna; the late Joseph Rosewater, Department of Invertebrates, U. S. National Museum, Smithsonian Institution; and J. D. 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Ayala, 1973, Mass Extinctions and Genetic Polymorphisms in the “Killer Clam,” Tridacna, Geol. Soc. Am. Bull., 84: 3411-3414. van der Vlerk, I. M., and J. H. F. Umbgrove, 1927, Tertiaire gidsforaminiferan van Nederlandsch Oost-Indie, Wetensch. Medeceel, 6: 1-31. Vermeij, G., E. A. Kay, and L. G. Eldredge, 1984, Molluscs of the Northern Mariana Islands with Special Reference to the Selectivity of Oceanic Dispersal Barriers, Micronesica, 19: 27-55. Vokes, E. H., and A. D’Attilio, 1982, Review of the Muricid Genus Attiliosa (Mollusca: Gastropoda), Veliger, 25: 67-71. Voss, G. L., 1954, Decapodous Cephalopod Mollusks from the Marshall Islands, Pac. Sci., 8: 363-366. Waller, T. R., 1972, The Pectinidae (Mollusca: Bivalvia) of Eniwetok Atoll, Marshall Islands, Veliger, 14: 221-264. Walls, J. G., n.d., Cone Shells, T. F. H. Publications, Inc., Nep- tune City, New Jersey. Wells, J. W., 1982, Fossil Corals frerm Midway Atoll, Geol. Surv. Prof. Paper 680G, 7 pp. Willan, R. C., 1984, The Pleurobranchidae (Opisthobranchia: Notaspidea) of the Marshall Islands, Central-West Pacific Ocean, Veliger, 27: 37-53. __, and N. Coleman, 1984, Nudibranchs of Australasia, Austral- asian Marine Photo Index, Sydney. Young, D. K., 1967, New Records of Nudibranchia (Gastropoda: Opisthobranchia: Nudibranchia) from the Central and West Central Pacific with a Description of a New Species, Veliger, 10: 159-173. ——, 1969, Okadaia elegans, a Tube-Boring Nudibranch Mollusc from the Central and West Central Pacific, Amer. Zool., 9: 903-907. Zeigler, R. F., and H. C. Porreca, 1969, Olive Shells of the World, Zeigler and Porreca, W. Henrietta, New York. Zmarzly, D. L., 1984, Distribution and Ecology of Shallow-Water Crinoids at Enewetak Atoll, Marshall Islands, with an Anno- tated Checklist of Their Symbionts, Pac. Sci., 38: 105-122. Chapter 15 Insects and Allies (Arthropoda) of Eneweiak Atoll G. ALLAN SAMUELSON and GORDON M. NISHIDA Bernice P. Bishop Museum Honolulu, Hawaii 96817 INTRODUCTION Insects and related terrestrial arthropods of Enewetak Atoll remain inadequately investigated. Woodbury (1962) included an inventory of arthropods known from the atoll in which he indicated that ectoparasites were taken from 10 species of birds; altogether he reported 45 species or subspecies authentically recorded from the atoll. The present list increases the number of arthropods to 191 species (or subspecies) and must still be considered prelim- inary, as records are lacking for many expected groups. Before this list can be completed, further collections and their study are required, especially of the soil fauna, ecto- parasites and nidicoles of reptiles, birds, and rodents, and the forms associated with humans and human habitations. A checklist of Enewetak insects and related arthropods is provided in Table 1. Investigations Enewetak Atoll appears to have received little or no attention from investigators of insects and other terrestrial arthropods until the beginning of American occupation in 1944. Published records and dated specimens from earlier times are either given without exact localities or refer to other parts of the Marshall Islands. The relative isolation of the atoll probably accounted for the inattention. Even dur- ing the period of the Japanese Mandate, when insects were collected elsewhere in the Marshalls and generally throughout Micronesia, nothing was specifically included from Enewetak. The entomological activities of the Japanese during that period are best summarized in a long series of papers reporting on results of Teiso Esaki’s Micronesian Expeditions of 1936-1940 (cf. Esaki et al., 1955). Entomological field work on Enewetak by Ameri- cans began in 1944 and continued through the postwar 147 years to the present. Surveys conducted by or with the support of the military, the U. S. Commercial Company, and the Pacific Science Board (PSB) gave considerable impetus to establishing the series Insects of Micronesia in 1954. This series has become the most important source for reporting on the Micronesian fauna. The earliest account treating Enewetak insects is apparently Townes (1946), a mimeographed report for the U. S. Commercial Company, which contains records of about 20 species. The first such account published in a serial is apparently Van Zwaluwenburg (1948) in which an elaterid beetle is reported. Titles containing records of Enewetak insects and allies are cited in tne bibliography. Visits to Enewetak Atoll by entomologists and others providing records of insects and related groups from the atoll are outlined in Table 2. So far as is known, all of these visits follow the period of Japanese occupation. Spellings of the islets or motus are those currently used by the Marshallese, and they may vary from names previously used in gazetteers or in Bryan (1971). For a time, an insect collection assembled by L. D. Tuthill was kept at the Mid-Pacific Research Laboratory (MPRL) at Enewetak. Those specimens eventually went to Bishop Museum for inclusion in studies reported in Insects of Micronesia. This collection, along with those conducted under the auspices of the PSB, are to be ultimately depos- ited in collections of participating PSB institutions: Califor- nia Academy of Sciences in San Francisco, Field Museum of Natural History in Chicago, Museum of Comparative Zoology at Harvard University in Cambridge, and National Museum of Natural History in Washington, as weli as Bishop Museum in Honolulu. Environment Enewetak Atoll is an _ isolated outlier on the northwestern extremity of the Ralik Chain in the drier, northern portion of the Marshall Islands. Vegetation is accordingly poorer there than on the wetter, southern atolls of both the Ralik and Ratak Chains (cf. Hathaway, 1953; St. John, 1960). A drier climate and a less diverse flora would offer fewer niches for insects; thus one would 148 SAMUELSON AND NISHIDA TABLE 7 Checklist of Enewetak Insects and Related Arthropods Class ARACHNIDA Order SCORPIONIDA Family ISCHNURIDAE Hormurus australasiae Fabricius, 1775: Townes, 1946. Order PSEUDOSCORPIONIDA Family ATEMNIDAE Oratemnus samoanus whartoni Chamberlin, 1947: Beier, 1957. Family CHERNETIDAE Haplochernes insulanus Beier, 1957: Beier, 1957. Family CHTHONIIDAE Lechytia sakagamii Morikawa, 1952: Beier, 1957. Order ARANEAE Family ARANEIDAE *Araneus theis (Walckenaer, 1841). Family CLUBIONIDAE *Chiracanthium diversum Koch, 1873. Family DIPLURIDAE Masteria hirsuta Koch, 1873: Roewer, 1963. Family EUSPARASSIDAE *Heteropoda venatoria (Linnaeus, 1767). Family LINYPHIIDAE *Erigone sp. Family OONOPIDAE Opopaea foveolata Roewer, 1963: Roewer, 1963. *Opopaea lena Suman, 1965. Family PHOLCIDAE *Smeringopus elongatus (Vinson, 1863). Family SALTICIDAE *Flacillula minuta (Berland, 1929). *Vitia albipalpis Marples, 1957. Family SCYTODIDAE Scytodes striatipes (Koch, 1873): Roewer, 1963. Family THERIDIIDAE Latrodectus geometricus (Koch, 1841): Townes, 1946. Order ACARI Suborder GAMASIDA Family DERMANYSSIDAE *Ornithonyssus sp. Suborder IXODIDA Family ARGASIDAE Ornithodoros capensis Neumann, 1901: Bushman, Parker and Johnson, 1963. Suborder ACTINEDIDA Family ERYTHRAEIDAE Balaustium sp. Belaustuim sp.: Townes, 1946. Class DIPLOPODA (MYRIAPODA in part) Order PROTEROSPERMOPHORA Family STRONGYLOSOMIDAE *Oxidus gracilis (Koch, 1847). Class INSECTA Order THYSANURA Family LEPISMATIDAE Not identified: Townes, 1946. Order COLLEMBOLA Family ENTOMOBRYIDAE *Seira bipunctata (Packard, 1873). Family ISOTOMIDAE *Isotomurus tricuspis Borner, 1906? “New Enewetak record. (This table continued on next page.) INSECTS AND ALLIES (ARTHROPODA) TABLE 1 (cont’d) Order ODONATA Family LIBELLULIDAE Pantala flavescens (Fabricius, 1798): Jackson, 1968. *Tramea lacerata Hagen, 1861. Order BLATTODEA Family BLATTELLIDAE *Blattella lituricollis (Walker, 1868). *Lupparia notulata (Stal, 1860). Family BLATTIDAE *Melanozosteria soror (Brunner, 1865). * Neostylopyga rhombifolia (Stoll, 1813). *Periplaneta americana (Linnaeus, 1758). *Periplaneta australasiae (Fabricius, 1775). Family PYCNOSCELIDAE *Pycnoscelis surinamensis (Linnaeus, 1767). Order ORTHOPTERA Family ACRIDIDAE Aiolopus thalassinus tamulus (Fabricius, 1798). Aiolopus tamulus (Fabricius, 1798): Townes, 1946. Family GRYLLIDAE * Ornebius bimaculatus (Shiraki, 1930). Gryllodes sigillatus (Walker, 1869): Townes, 1946. “Speonemobius sp. prob. tigrinus (Saussure, 1877). Family TETTIGONIIDAE Phisis pallida (Walker, 1869): Townes, 1946. Order DERMAPTERA Family CARCINOPHORIDAE Euborellia annulipes (Lucas, 1847): Brindle, 1972. Family CHELISOCHIDAE Chelisoches morio (Fabricius, 1775): Townes, 1946. Family LABIDURIDAE Labidura riparia (Pallas, 1773): Townes, 1946; Brindle, 1972. Order PSOCOPTERA Family CAECILIIDAE Caecilius casarum Badonnel, 1931: Thornton et al., 1972. Family ECTOPSOCIDAE Ectopsocus maindroni Badonnel, 1935: Thornton et al., 1972. Family LEPIDOPSOCIDAE Cyptophania marginata Thornton, Lee and Chui, 1972: Thornton et al., 1972. Family PSOCIDAE Ptycta angulata Thornton, Lee and Chui, 1972: Thornton et al., 1972. Order MALLOPHAGA Family MENOPONIDAE * Actornithophilus bicolor (Piaget, 1880). Actornithophilus ceruleus (Timmermann, 1954): Woodbury, 1962. * Actornithophilus incisus (Piaget, 1880). * Actornithophilus ochraceus (Nitzsch, 1818). * Austromenopon atrofuluum (Piaget, 1880). * Colpocephalum angulaticeps Piaget, 1880. Family PHILOPTERIDAE * Halipeurus mirabilis Thompson, 1940. * Pectinopygus gracilicornis (Piaget, 1880). * Quadraceps birostris (Giebel, 1874). * Quadraceps hopkinsi Timmermann, 1952. * Quadraceps separatus (Kellogg and Kuwana, 1902). * Saemundssonia albemarlensis (Kellogg and Kuwana, 1902). * Saemundssonia puellula Timmermann, 1965. * Saemundssonia remota Timmermann, 1951. * Saemundssonia snyderi (Kellogg and Paine, 1910). * Trabeculus hexakon (Waterston, 1914). “New Enewetak record. (This table continued on next page.) 149 150 SAMUELSON AND NISHIDA TABLE 1 (cont’d) Order THYSANOPTERA Family PHLAEOTHRIPIDAE * Haplothrips gowdeyi (Franklin, 1908). Order HETEROPTERA Family ANTHOCORIDAE Physopleurella mundula (White, 1877): Herring, 1967. Family COREIDAE Liorhyssus hyalinus (Fabricius, 1794): Gross, 1963. Family CYDNIDAE Geotomus pygmaeus (Dallas, 1851): Ruckes, 1963. Family GERRIDAE Halobates micans (Eschscholtz, 1822): Cheng, 1981. Hermatobates sp.: Cheng, 1977. Family LYGAEIDAE *Germalus sp. Nysius picipes Usinger, 1937: Usinger, 1949, 1951; Barber, 1958. Nysius pulchellus Stal, 1859: Usinger, 1949, 1951; Barber, 1958. Pachybrachius nigriceps (Dallas, 1852): Barber, 1958. Family MIRIDAE Campylomma eniwetok Schuh, 1984: Schuh, 1984. Campylomma marshallensis Usinger, 1952: Schuh, 1984. Trigonotylus dohertyi (Distant, 1904): Carvalho, 1956. Family NABIDAE Nabis capsiformis Germar, 1839: Gross, 1963. Family PENTATOMIDAE Oechalia schellenbergii (Guerin-Méneville, 1831): Ruckes, 1963. Oechalia consocialis (Boisduval): Usinger, 1952. Platynopus melacanthus (Boisduval, 1835): Ruckes, 1963. Order HOMOPTERA Family APHIDIDAE Aphis gossypii Glover, 1855: Essig, 1956. Aphis craccivora Koch, 1854. Aphis medicaginis Koch, 1854: Essig, 1956. Family CICADELLIDAE Balclutha incisa (Matsumura, 1902): Linnavouri, 1975. Balclutha saltuella (Kirschbaum, 1868): Linnavousi, 1975. Exitianus fusconervosus (Motschulsky, 1863). Exitianus capicola Stal, 1855: Linnavouri, 1960 (in part). Exitianus plebeius (Kirkaldy, 1906): Linnavouri, 1960. Exitianus capicola Stal, 1855: Linnavouri, 1960 (in part), Orosius argentatus (Evans, 1940): Linnavouri, 1975. Nesaloha cantonis Oman, 1943: Oman, 1943. Orosius cantonis (Oman, 1943): Ghauri, 1966. Recilia affinis (Osborn, 1934): Linnavouri, 1975. Recilia hopponis (Matsumura, 1914): Linnavouri, 1975. Superfamily COCCOIDEA Ceroplastes cirripediformis Comstock, 1881: Beardsley, 1966. Coccus hesperidum Linnaeus, 1758: Beardsley, 1966. Icerya purchasi Maskell, 1878: Beardsley, 1966. Lepidosaphes esakii Takahashi, 1939: Beardsley, 1966. Phenacoccus solani Ferris, 1918: Beardsley, 1966. Pseudococcus microadonidum Beardsley, 1966: Beardsley, 1966. Saissetia coffeae (Walker, 1852): Beardsley, 1966. Family DELPHACIDAE Sogatella kolophon (Kirkaldy, 1907): Fennah, 1971. Family DERBIDAE Lamenia caliginea charon Fennah, 1956: Fennah, 1956, 1971. Order COLEOPTERA Family ANTHICIDAE Anthicus vexator Werner, 1965: Werner, 1965. *New Enewetak record. (This table continued on next page.) INSECTS AND ALLIES (ARTHROPODA) 151 TABLE 1 (cont'd) Family CARABIDAE Egadroma smaragdula (Fabricius, 1798): Darlington, 1970. Family CERAMBYCIDAE Prosoplus atlanticus atlanticus Breuning, 1938: Gressitt, 1956. Family CHRYSOMELIDAE *Aphthona bicolorata Jacoby, 1904. Family COCCINELLIDAE Coelophora inaequalis (Fabricius, 1775): Chapin, 1965 [including variety novemmaculata (Fabricius, 1781): Chapin, 1965]. Harmonia arcuata (Fabricius, 1787): Chapin, 1965. Nephus roepkei (Fluiter, 1938): Chapin, 1965. *Scymnus nigrosuturalis Kamiya, 1961. Family DERMESTIDAE Dermestes ater De Geer, 1774: Beal, 1961. Family ELATERIDAE Conoderus pallipes Eschscholtz, 1829: Van Zwaluwenburg, 1948, 1957. Simodactylus fasciolatus Fairmaire, 1863: Van Zwaluwenburg, 1957. Family MYCETOPHAGIDAE Typhaea stercorea Linnaeus, 1758: Michitaka Chijo, 1970. Family NITIDULIDAE Carpophilus davidsoni Dobson, 1952: Gillogly, 1962. Carpophilus hemipterus (Linnaeus, 1758): Gillogly, 1962. Carpophilus marginellus Motschulsky, 1858: Gillogly, 1962. Carpophilus mutilatus Erichson; 1843: Gillogly, 1962. Carpophilus pilosellus Motschulsky, 1858: Gillogly, 1962. Urophorus humeralis (Fabricius, 1798): Gillogly, 1962. Family OEDEMERIDAE * Eobia kanack (Fairmaire, 1849). * Eobia matsumurai Kono, 1937. * Eobia trukana Kono, 1937. * Eobia new species, Macnamara and Arnett, in preparation. * Pselaphanca apicata (Fairmaire, 1881). “New genus and species, Macnamara and Arnett, in preparation. Family PROPALTICIDAE Propalticus insularis John, 1960, 1971. Family SCOLYTIDAE Cryphalomorphus nubilus Wood, 1960: Wood, 1960. Hypothenemus eruditus Westwood, 1836: Wood, 1960. Family TENEBRIONIDAE Gebieniella carinata (Eschscholtz, 1831): Kulzer, 1957. *Gonocephalum adpressiforme Kaszab, 1951. Order NEUROPTERA Family CHRYSOPIDAE Chrysopa ramburi Schneider, 1851: Adams, 1959. Order LEPIDOPTERA Family ARCTIIDAE Utetheisa pulchelloides Hampson, 1907: Townes, 1946. Family HESPERIIDAE Badamia exclamationis (Fabricius, 1775): Townes, 1946. Family NOCTUIDAE Spodoptera exempta (Walker, 1856). Laphygma exempta (Walker, 1856): Townes, 1946. Spodoptera litura (Fabricius, 1775). Prodenia litura (Fabricius, 1775): Townes, 1946. Family NYMPHALIDAE Hypolimnas bolina (Linnaeus, 1764): Townes, 1946. Precis villida (Fabricius, 1787): Townes, 1946. Precis villida bismarckiana (Hagen): Clark, 1951. *New Enewetak record. (This table continued on next page.) SAMUELSON AND NISHIDA TABLE 1 (cont'd) Family SPHINGIDAE *“Agrius convolvuli (Linnaeus, 1758): Riotte, in preparation. Cephonodes armatus Rothschild and Jordan, 1903: Townes, 1946. *Cephonodes picus (Cramer, 1777): Riotte, in preparation. Gnathothlibus erotus (Cramer, 1777). Chromis erotus (Cramer, 1777): Townes, 1946. Family TORTRICIDAE Adoxophyes fasciculana (Walker, 1866): Clarke, 1976. Order DIPTERA Family AGROMYZIDAE Ophiomyia cornuta (de Meijere, 1910): Spencer, 1963. Pseudonapomyza spicata (Malloch, 1914): Spencer, 1963. Family CALLIPHORIDAE Chrysomya megacephala (Fabricius, 1784): James, 1962. Phaenicia cuprina (Wiedemann, 1930): James, 1962. Family CANACEIDAE Nocticanace marshallensis Wirth, 1951: Wirth, 1951. Family CERATOPOGONIDAE Dasyhelea esakii Tokunaga, 1940: Tokunaga and Murachi, 1959. Dasyhelea flavescens Tokunaga and Murachi, 1959: Tokunaga and Murachi, 1959. Dasyhelea flavibasalis Tokunaga, 1940: Tokunaga and Murachi, 1959. Dasyhelea peliliouensis Tokunaga, 1940: Tokunaga and Murachi, 1959. Forcipomyia tuthilli Tokunaga, 1959: Tokunaga and Murachi, 1959. Family CHIRONOMIDAE Clunio tuthilli: Tokunaga, 1964. Pontomyia natans Edwards, 1926: Cheng and Hashimoto, 1978. *Tanytarsus halophilae Edwards, 1926. *Telmatogeton pusillum Edwards, 1935. Thalassomya maritima Wirth, 1947: Tokunaga, 1964. Family CHLOROPIDAE *Cadrema pallida (Loew, 1866). *“Gaurax” bicoloripes (Malloch, 1932). Family EPHYDRIDAE *Allotrichoma alium Cresson, 1929. *Discocerina mera Cresson, 1939. *Hecamede persimilis Hendel, 1913. *Hostis guamensis Cresson, 1945. *Placopsidella cynocephala Kertész, 1901. Family HIPPOBOSCIDAE * Olfersia spinifera (Leach, 1817). Family MILICHIIDAE * Desmometopa varipalpis Malloch, 1927. * Milichiella lacteipennis (Loew, 1866). Family MUSCIDAE Atherigona flavipalpis Malloch, 1928: Snyder, 1965. Atherigona (Acritochaeta) orientalis Schiner, 1868. Atherigona excisa (Thompson, 1868): Snyder, 1965. Musca (Musca) domestica Linnaeus, 1758: Snyder, 1965. Family PHORIDAE Diplonevra (Dohrniphora) cornuta (Bigot, 1857): Beyer, 1967. Megaselia (Meguselia) scalaris (Loew, 1866): Beyer, 1967. Family SARCOPHAGIDAE Boettcherisca karnyi (Hardy, 1927): Souza Lopes, 1958. Parasarcophaga (Liosarcophaga) misera (Walker, 1849): Souza Lopes, 1958. Phytosarcophaga gressitti (Hall and Bohart, 1948): Souza Lopes, 1958. Family SCATOPSIDAE *Holoplagia guamensis (Johannsen, 1946) or near? Family SCIARIDAE Bradysia tritici (Coquillett, 1895): Steffan, 1969. “New Enewetak record. (This table continued on next page.) INSECTS AND ALLIES (ARTHROPODA) 153 TABLE 1 (cont'd) Family SCIARIDAE (cont’d) Corynoptera latistylata (Hardy, 1956): Steffan, 1969. Plastosciara latipons Hardy, 1956: Steffan, 1969. Family STRATIOMYIDAE Brachycara ventralis Thomson, 1869: James, 1962. Family SYRPHIDAE Ishiodon scutellaris (Fabricius, 1805): Shiraki, 1963. Family TIPULIDAE Lirnonia (Dicranomyia) pectinunguis Tokunaga, 1940: Alexander, 1972. Order HYMENOPTERA Family ANTHOPHORIDAE * Xylocopa sonorina Smith, 1879. Family BRACONIDAE Zele sp.: Townes, 1946. Family EDLOPHIDAE Hemiptarsenus semialbiclavus (Girault, 1916): Yoshimoto and Ishii, 1965. Family EVANIIDAE *Szepligetella sericea (Cameron, 1883). Family FORMICIDAE Monomorium pharaonis (Linnaeus, 1758): Townes, 1946. Family ICHNEUMONIDAE Echthromorpha agrestoria insidiator (Smith, 1863): Townes, 1958. Family MEGACHILIDAE Megachile diligens hedleyi Rainbow, 1897: Krombein, 1950. Megachile (Eutricharaea) fullawayi Cockerell, 1914: Krombein, 1950. Family SPHECIDAE *Chalybion bengalense (Dahlbom, 1845). Pison punctifrons Shuckard, 1838: Krombein, 1949. Solierella peckhami (Ashmead, 1897). Solierella rohweri (Bridwell, 1920): Krombein, 1949. *Tachysphex bengalensis Cameron, 1889. Family VESPIDAE *Odynerus sp. Pachodynerus nasidens (Latreille, 1812): Townes, 1946. *Polistes fuscatus aurifer (Saussure, 1853). *Ropalidia marginata (Lepeletier, 1836). “New Enewetak record. expect fewer indigenous insect species on Enewetak than on, for instance, Arno Atoll. However, the introduction of exotic plant and insect species on Enewetak complicates the picture. The most notable aspect concerning the environment at Enewetak is obviously the man-made alterations and distur- bances over many parts of the atoll. These had extreme impacts on the indigenous biota and tended to become more severe as the sequence of human occupation pro- gressed on the atoll. The advent of Micronesians and the development of a native culture were followed by coloniza- tion and by trade with non-Micronesian elements. Thus, at least certain islets were affected by the changes associated with the earlier phases of human occupation, including the establishment of copra plantations. The later periods included construction of military bases and air fields, war- fare, cleanup operations following warfare, and testing of nuclear devices and associated activities. The devastation by warfare that occurred on certain islets during World War II was subsequently magnified by applications of insec- ticides (Townes, 1946) and cleanup operations in which entire islets were cleared by bulldozing. One of the early programs of the U. S. military was to establish vegetative growth on the bare sand to hold it down (Bryan, 1944; personal communication). The postwar testing of nuclear devices accounted for further and sometimes complete devastation of certain islets. Some sites were also bulldozed following tests. All these postwar activities and the traffic of personnel and materials, mainly through Hawaii, facilitated the importa- tion of exotic plants and insects. Many species of plants were intentionally introduced or reintroduced to Enewetak from nursery stock via Kwajalein (Fosberg, 1959). Over 50 species of exotic plants were reported for Enewetak during that period (St. John, 1960) marking a dramatic increase of exotics over previous periods when, for example, only 24 exotics were estimated for a lusher part of the Marshall Islands (Arno) in 1850 (Hathaway, 1953). The plants in some measure may serve as an indication of the numbers of insects and related groups that were also introduced to 154 SAMUELSON AND NISHIDA TABLE 2 Collectors of Insects and Related Groups on Enewetak Atoll Name Month/Year Islets Allen, R. Aug. 1945 Enewetak, Runit Bohart, R. Jan. 1945 Enewetak? Bryan, E. H., dr. Aug. 1944 Enewetak, Enjebi, Japtan, Medren Feb. 1975 Alembel, Ananij, Bijile, Enewetak, Enjebi, Ikuren, Japtan, Lojwa, Medren, Runit Bushman, J. B. Mar.—May 1962 Aej, Enewetak, Enjebi, Inedral, daptan, Jinimi, Medren, Mijikadrek, Ribewon Cheng, L. Sept. 1975 Enewetak, Ikuren, Japtan, Medren Dybas, H. S. Nov.—Oct. 1944 Enewetak, Japtan Edgar, S. A. Nov.—Oct. 1944 Enewetak, Japtan Fosberg, F. R. May 1946 Japtan May 1951 Enjebi Hagen, E. Nov. 1944 Enewetak, Japtan Jackson, W. B. 1964-67 (summers) Enjebi, Runit Knudsen, J. W. Apr. 1965 Enewetak Aug. 1967 ? Lamberson, P. ? Enjebi Morrison, J. Apr.—June 1946 Biken, Ikuren, Kidrinen, Louj, Lujor, Mut Oakley, R. G. May 1946 Aomon, Enjebi, Japtan (and others ‘ with Townes?) Oshiro, Y. Dec. 1950—Jan. 1951 Bokombako, Elugelab, Enjebi Parker, D. Mar.—Apr. 1962 Aej Reish, D. J. July 1956 Enewetak Townes, H. K. May 1946 Aomon, Enewetak, Enjebi, Ikuren, Japtan Tuthill, L. D. Aug.—Sept. 1956 Ananij, Ikuren, Japtan, Medren Jan.—Feb. 1957 Japtan, Medren July 1957 “Nan” Sept. 1957 Medren Aug. 1958 Japtan Wheeler, M. R. Aug.—Sept. 1955 Enjebi, Ikuren, Japtan, Medren Young, F. N. May 1945 Medren Enewetak during the same periods. Hence, the indigenous (including endemic) biota of Enewetak—the species that represent the true unadulterated character of the atoll—probably suffered profoundly through both physical impacts on the environment and _ biological impacts, including competition for space and resources from adven- tive forms. In view of the disturbances to Enewetak Atoll, it seems relevant to note the times and places where species were collected or observed, thereby providing a summary of the first four decades of entomological records for the atoll. This information is provided in the section on Collection Records. Fauna Townes (1946) estimated that a large dry atoll like Enewetak might have 500 species of insects. This estimate could be somewhat high. The indigenous insect fauna of Enewetak wouid be expected to be less than that of Arno Atoll. The fauna of Arno was also estimated at around 500 species (Usinger and LaRivers, 1953), and at least 296 species have already been recorded for that atoll (Gressitt, 1954). Cole (1949, 1951) listed 56 species of insects collected from Bikini Atoll in 1947. Bikini lies in the same zone as Enewetak and would be expected to sup- port a nearly identical fauna. Cole’s survey was the only one conducted for Bikini and, unfortunately, it occurred after applications of DDT on the islets and after the first atomic bomb tests there. These perturbations must have had severe effects on the fauna and on Cole’s results. Cole’s lists were admittedly incomplete; however, 35 species listed by him, including 12 species of ants, have not yet been reported for Enewetak. Most of these 35 species probably also occur or have occurred on Enewetak. INSECTS AND ALLIES (ARTHROPODA) We expect that, when fully studied, the arthropod fauna of Enewetak Atoll will comprise 350 to 400 species. This estimate takes into consideration that the 191 taxa reported herein do not include many groups that are expected to occur in cryptic habitats, such as soil and decaying plant materials and those associated with vertebrates. In comparing the fauna of Enewetak Atoll with the total number of species reported for the Marshall Islands, we find about 32% (191 of 592) represented on Enewetak (Table 3). Many of the exotic species carried to Kwajalein and Enewetak in recent years are, of course, reflected in this percentage, which would be lower if only the truly indigenous and endemic elements were considered. We estimate that around 43% (82 of 191) of the species presently listed for Enewetak could be adventive elements. The extent of transfer into neighboring atolls by these adventives is not known. The percentage of species endemic to the Marshalls is certainly not high at 4% (25 of 592); we expect this figure 166 to increase somewhat after future collections from cryptic habitats are studied. These endemics are listed in Table 4. The proportion of species endemic to Enewetak Atoll is very low at 1%% (3 of 191); this percentage, too, is expected to increase following future studies. Of the 25 species endemic to the Marshalls, 52% (13) are restricted to the western or Ralik Chain, 24% (6) are restricted to the eastern or Radak Chain, and 24% (6) are found on both chains (Table 5). Present records indicate that 64% (16 of 25) of these endemics are restricted to single atolls; they represent only 242% (16 of 592) of the total Marshall Islands’ fauna. A rather low proportion, 912% (57 of 592), of the Marshall Islands fauna is restricted to the central and east- ern Carolines—Marshalls—Gilberts sector of the Pacific (Table 6). This includes the most characteristic species to be expected on atolls; they are listed in Table 7. About half (31 of 57) are strictly limited to atolls and low islands, and these include the taxa endemic to the Marshalls. The others (26 of 57) also occur on high islands proximal to TABLE 3 Numbers of Insects and Allies Reported for the Marshall Islands and Numbers for Enewetak Atoll Only, Including Counts of Endemic Taxa Marshal! Enewetak Endemic to Endemic to Group Islands Atoll Marshalls Enewetak Arachnida 42 19 2 Myriapoda 4 1 Insecta: Thysanura 1 1 Collembola’® 2 2 Odonata 7 2 Blattodea* 9 7 Mantodea 1 Phasmatodea 1 Orthoptera 10 5 Isoptera 6 Dermaptera 7 3 Embioptera 1 Psocoptera 16 4 Mallophaga 22 16 Anoplura 3 Thysanoptera 2 1 Heteroptera 30 15 2 1 Homoptera 65 18 1 Coleoptera 116 29 9 Neuroptera 2 1 Lepidoptera 40 11 Diptera 142 40 10 2 Siphonaptera 1 Hymenoptera 62 16 1 Totals 592 191 25 3 “Group is poorly surveyed, with Enewetak records accounting for most or all Marshall Island records. 156 SAMUELSON AND NISHIDA TABLE 4 Taxa Endemic to the Marshall Islands Arachnida Pseudoscorpionida Garypus ornatus Beier—Garypidae (Bikini) Xenolpium oceanicum latum Beier—Olpiidae (Lae, Ujae, Utirik) Insecta Heteroptera Riptortus saileri Usinger—Coreidae (Kwajalein) Campylomma eniwetok Schuh—Miridae (Enewetak) Homoptera Ugyops kinbergi magas Fennah—Delphacidae (Ailinglapalap, Jaluit) Coleoptera Ceresium robustum Gressitt—Cerambycidae (Arno) Ceresium unicolor marshallum Gressitt—Cerambycidae (Majuro) Oopsis marshallensis Gressitt—Cerambycidae (various atolls) Prosoplus hibisci Gressitt—Cerambycidae (Majuro, Utirik) Prosoplus major Gressitt—Cerambycidae (Arno) Melanoxanthus lariversi Van Zwaluwenburg—Elateridae (Arno) Simodactylus marshallensis Ohira—Elateridae (Kwajalein) Chariotheca costata Kulzer—Tenebrionidae (Arno, Ailinglapalap) Tagalus angustus Kulzer—Tenebrionidae (Arno) Diptera Nocticanace marshallensis Wirth—Canaceidae Dasyhelea nigristigmata Tokunaga and Murachi— Ceratopogonidae (Ailinglapalap) Dasyhelea sp. No. 1 Tokunaga and Murachi—Ceratopogonidae (Arno) Forcipomyia tuthilli Tokunaga—Ceratopogonidae (Enewetak) Clunio tuthilli Tokunaga—Chironomidae (Enewetak) Cricotopus sp. No. 1 Tokunaga—Chironomidae (Kwajalein) Plastosciara jaluitensis Steffan—Sciaridae (Jaluit) Scythropochroa trispinosa Steffan—Sciaridae (Kwajalein) Limonia beardsleyi Alexander—Tipulidae (Namu, Namorik) Limonia sentifera Alexander—Tipulidae (Namu, Arno) Hymenoptera Cirrospiloideus fullawayi Yoshimoto and Ishii—Eulophidae (Ebon) atolls where they are indigenous in the central and eastern Carolines (Yap, Truk, Ponape, Kosrae). None of the Marshalls or Gilberts are correspondingly close to a high island. The high islands of the Carolines are centers of endemism with relatively rich faunas and may have been sources for many of the forms now strictly limited to atolls. Thus, the presence of isolated high islands in the vicinity of atolls may have considerable influence on the nature of an atoll fauna. Factors influencing an atoll fauna would include the above, as well as the degree of isolation of the atoll, land area, climate, frequency of typhoons, effects of wind and sea currents, and presence of migratory birds. Finally, human activities have considerable bearing on the nature TABLE 5 Distribution of Insects and Allies Endemic to the Marshall Islands Ralik Ratak Chain Chain Both Single Group (Western) (Eastern) chains atoll Arachnida 1 1 1 Heteroptera Z 2 Homoptera 1 Coleoptera 1 5 3 5 Diptera 8 1 1 7 Hymenoptera 1 1 Totals 13 6 6 16 of an atoll fauna; these affect faunal composition through environmental alteration and allow increased opportunities for establishment of adventive species. Direct air and ship- ping routes between the Marshalls and Hawaii have obvi- ously permitted movements of species between the two island groups within the past 40 years (Table 8). Of the adventive elements on Enewetak that also occur in Hawaii, we note that they represent around 28% (54 of 191) of the atoll’s fauna. Furthermore, when considering the Enewetak adventives alone, we see that a higher propor- tion, 66%, also occurs in Hawaii (54 of 82). Hawaii, as an example of a center of commerce in the Pacific, has wit- nessed a marked increase of established adventives over recent decades (cf. Beardsley, 1962, 1979), and that is due largely to increased air traffic. Gressitt (1954) stated that atolls and low islands in the Carolines, Marshalls, and Gilberts have relatively poor but uniform faunas. This appears so within the Marshalls, at least at the generic level and frequently at the species level, when the adventive forms established on Kwajalein and Enewetak Atolls are discounted. Zoogeography The geographical affinities of major groups represented in the Marshall Islands, or otherwise in Micronesia, are only partly ascertained at this time. Many groups are not yet studied to the point where they can be analyzed defini- tively. This is also true for the faunas of adjacent regions; all have influences on the Micronesian fauna through past or present movements of species. In the following discussion, the general geographical distributions are given for all species reported for the Marshall Islands to classify the potential inventory and sources of the species for Enewetak Atoll. The numbers of species counted for the Marshalls and Enewetak are pro- vided at the beginning of most of the following sections. Also, most major groups occurring in Micronesia are sum- marized to give some perspective to the Marshall Islands fauna; this includes, when known, the number of species or subspecies reported for Micronesia and the percentage INSECTS AND ALLIES (ARTHROPODA) 157 TABLE 6 Distribution of Marshalls’ Fauna Restricted to the Carolines (Central and Eastern*)—Marshalls—Gilberts Sector of the Pacific Ocean TABLE 7 Taxa Reported for the Marshalls and Restricted to the Central and Eastern Carolines (Excluding Palau)—Marshalls-Gilberts Sector of the Pacific Ocean Atolls: Of Atolls island and high Atolls: groups islands Marshalls within within Groupt only sector sector Arachnida Pseudoscorpionida 2 1 1 Opiliones 1 Insecta Psocoptera 2, Heteroptera Coreidae 1 Lygaeidae 1 Miridae 1 1 Homoptera Cixiidae 2 Pseudococcidae 2 Delphacidae 1 1 Derbidae 1 Coleoptera Cerambycidae 5 3} Chrysomelidae 1 Coccinellidae il Elateridae 2 Oedemeridae 1 Scolytidae 1 Tenebrionidae 2 1 Lepidoptera Noctuidae 1 Diptera Canaceidae 1 Ceratopogonidae 3 Zz Chironomidae 2 1 Culicidae 1 Drosophilidae 1 Platystomatidae 1 Sciaridae 2 Stratiomyidae 1 Tipulidae 2 Hymenoptera Cynipoidea 1 Eucharidae 1 Eulophidae 1 1 Totals 25 6 26 “Central and Eastern Carolines = all atolls and high islands E of Palau; this division is the Federated States of Micronesia. +Table 4 lists taxa in column 1, Table 7 lists taxa in columns 2 and 3. Arachnida Pseudoscorpionida Lechytia sakagamii Morikawa (CA, M + Marcus) Nesidiochernes maculatus Beier (K, M) Opiliones Zalmoxis solitaria (Roewer) (P,M) Insecta Psocoptera Ectopsocus villosus Thornton and Wong (T, P, K, M) Heterocaecilius campanula Lee and Thornton (P, CA, M, G) Hemiptera Nysius picipes Usinger—Lygaeidae (M, G + Wake) Campylomma marshallensis Usinger—Miridae (M, G + Kanton) Homoptera Myndus apicalis (Metcalf)—Cixiidae (CA, T, P, K, M) Myndus praecanus Fennah—Cixiidae (P, M) Pseudococcus marshallensis Beardsley—Pseudococcidae (K, M) Pseudococcus microadonidum Beardsley—Pseudococcidae (T, P, M, G) Ugyops superciliata Fennah—Delphacidae (CA, P, K, M) Flaccia dione Fennah—Derbidae (CA, K, M) Coleoptera Paremeopedus minimus (Blair)—Cerambycidae (CA, T, P, M) Prosoplus a. atlanticus Breuning—Cerambycidae (P, M) Prosoplus xyalopus (Karsch)—Cerambycidae (Y, CA, M, G) Brontispa chalybeipennis (Zacher)—Chrysomelidae (P, K, M) Scymnus uncinus Chapin—Coccinellidae (M, G) Eobia new species Macnamara and Arnett—Oedemeridae (M, G + Ocean) Cryphalomorphus nubilus Wood—Scolytidae (CA, K, M) Uloma lariversi Kulzer—Tenebrionidae (K, M) Lepidoptera Nagia hieratica Hampson—Noctuidae (Y, P, M, G) Diptera Dasyhelea flavibasalis Tokunaga—Ceratopogonidae (P?, M) Dasyhelea pallivittae Tokunaga—Ceratopogonidae (K, M) Smittia yapensis Tokunaga—Chironomidae (Y, T, CA, M) Aedes marshallensis Stone and Bohart—Culicidae (CA, K, M, G) Microdrosophila errator Wheeler and Takada—Drosophilidae (P, K, M) Scholastes carolinensis Enderlein—Platystomatidae (P, M) Cephalochrysa rugulosa James—Stratiomyidae (P, K, M) Hymenoptera Pseudeucoila gressitti Yoshimoto—Cynipoidea (T, M) Parachalcura maculata Watanabe—Eucharidae (CA, M) Hemiptarsenus carolinensis Yoshimoto and Ishii—Eulophidae (Y, CA, M, G) “CA, Caroline Atolls; G, Gilberts; K, Kosrae; M, Marshalls; P, Ponape; T, Truk; Y, Yap. 158 SAMUELSON AND NISHIDA TABLE 8 Proportions of Indigenous and Adventive Species for Groups Represented on Enewetak Atoll Indig- Also Total enous, found species including Adven- in Group reported endemic tive Hawaii Arachnida 19 10 9 6 Myriapoda 1 1 1 Insecta: Thysanura 1 1 Collembola 2 2 Odonata 2 1 1 Z Blattodea 7 7 7 Orthoptera 5 5 2 Dermaptera 3 3 3 Psocoptera 4 2 2 2 Mallophaga 16 16 13} Thysanoptera 1 il 1 Heteroptera 15 11 4 6 Homoptera 18 7 11 8 Coleoptera 29 18 11 10 Neuroptera 1 1 Lepidoptera 11 8 3 2 Diptera 40 30 10 23 Hymenoptera 16 5 11 9 Totals 191 109 82 95° *Forty-one are indigenous to Enewetak; 54 are adventive. of those endemic to Micronesia. These figures will certainly change as studies continue on this and adjacent faunas. ARACHNIDA The Micronesian arachnid fauna is partially studied, though the spiders and mites remain incompletely sur- veyed. The Amblypygi just enter the Micronesian area at Palau, with one Papuan—Malayan species reported, Charon grayi Gervais (Takashima, 1950), Scorpionida Marshalls 2, Enewetak 1—The Hormurus we list for Enewetak is widely distributed in the Pacific and also occurs from China to Australia. The tropicopolitan Isometrus maculatus (DeGeer) is reported for the Marshalls, Gilberts, Palau; it is commonly found in _ dwell- ings but is not yet reported for Enewetak. Micronesian fauna, studied by Chapin (1957), comprises three species. A second Hormurus of mainly Australian—Melanesian distri- bution reaches Palau. Pseudoscorpionida Marshalls 10, Enewetak 3—Seven of the 10 pseudo- scorpions reported for the Marshalls are restricted to Micronesia. Two of these are endemic to the Marshalls but are not reported for Enewetak; they are Garypus ornatus Beier (Bikini) and Xenolpium oceanicum latum (Beier) (Lae, Ujae, Utirik). The Lechytia we list for Enewetak is re- stricted to atolls or low islands, including the Carolines (Ulithi) and Marcus Island. Nesidiochernes maculatus Beier is restricted to the eastern Carolines (Kosrae) and Marshalls and is not yet reported for Enewetak. Three species are either widespread in Micronesia or at least recorded from distant localities. One of these is the Haplochernes we list for Enewetak. The three extra Micronesian taxa comprise Oratemnus samoanus whartoni Chamberlin reported for Enewetak, western Carolines, and Okinawa; the nominate O. s. samoanus Beier of mainly Polynesian distribution; and a species of Geogarypus also occurring on Taiwan. Micronesian fauna, studied by Beier (1957), comprises 45 species or subspecies of which all are confined to the Pacific basin. Opiliones Marshalls 1, Enewetak 0—Only one species, Zalmoxis solitaria (Roewer), is recorded for the Marshalls; its distri- bution is restricted to the eastern Carolines (Ponape) and Marshalls (Jaluit). Micronesian fauna, studied by Goodnight and Goodnight (1957), comprises six endemic species in five genera. One genus, Parasamoa, is endemic to Ponape and is related to a Samoan genus and also to another in the Seychelles; the remaining four genera have representa- tives in the Malayan subregion. Araneae Marshalls 19, Enewetak 12—Spiders reported for the Marshalls are widespread (9 species), distributed in the Malayan subregion and Pacific (3), confined to Oceania (5), or undetermined (2). Of the species we list for Enewetak, the Araneus, Chiracanthium, Flacillula, | Heteropoda, Latrodectus, Smeringopus, and Vitia are widespread; the Masteria, two Opopaea, and Scytodes are confined to Oceania; and the Erigone is of unknown distribution, as it is unidentified. At least five of the 12 spiders reported for Enewetak also occur in Hawaii, a probable avenue for the introduction of some of the adventive species. Roewer (1963) treated seven families of Micronesian spiders total- ing 13 species, 10 of which are restricted to Micronesia; eight of the species we treat for Enewetak are not included in Roewer’s treatment. Acari Many additional records are expected for the Marshalls; suborders are treated ‘below. Gamasida (Mesostigmata): Marshalls 4, Enewetak 1—Laelaps echidnina Berlese and L. nuttalli Hirst are reported for the Marshalls but not Enewetak; both are associated with Rattus and are cosmopolitan or nearly so. The Ornithonyssus we list for Enewetak is not identified, but the only species thus far reported for Micronesia is the INSECTS AND ALLIES (ARTHROPODA) 159 cosmopolitan O. bacoti (Hirst), a species associated with rodents, particularly rats. These mites are among the seven parasitic Mesostigmata reported for Micronesia by Wilson (1967). Glyptholaspis asperrima (Berlese) is one of five macrochelids reported for Micronesia by Krantz (1967); it is described from Java and reported for Palau and the Marshalls (Arno); some of the specimens are from a palm log (Palau). Ixodida (Metastigmata): Marshalls 2, Enewetak 1—The ticks, reviewed by Kohls (1957), now comprise at least six species in Micronesia. Kohls reported Rhipicephalus sanguineus (Latreille) for the Marianas and Gilberts; earlier, Schnee (1904) reported it for the Marshalls (Jaluit). This cosmopolitan tick is associated mainly with the dog. The Ornithodoros we list for Enewetak is from collections from two species of noddies and their nests and possibly from three additional species of sea birds from Enewetak (Bush- man, Parker, and Johnson, 1963). This tick is widespread on oceanic islands of both hemispheres, including Hawaii. Actinedida (Prostigmata): Marshalls 4, Enewetak 1—Balaustium and Calorema are not identified further; the former is reported for Enewetak and the latter for Kwajalein. Two species of Tetranychus are reported for Kwajalein by Sugerman (1972a, 1979): T. cinnabarinus (Boisduval) is known from North America and India, and T. tumidus Banks is reported for the United States, Puerto Rico, and Guam. Oribatida: Records are lacking for the Marshalls, though the group is certainly expected there. Micronesian oribatids are beginning to be reported upon by Sengbusch (1982a, b), who described two lohmnanniids endemic to the Carolines (Yap, Ponape) belonging to an ancient group of Gondwanan origin. MYRIAPODA The Micronesian myriapod fauna is poorly sampled, with only chilopods and diplopods reported for the Marshalls. Pauropods are reported only from Palau and Guam (Remy, 1957). Symphylans apparently remain unreported for Micronesia, though they may be expected on remote islands. Hanseniella orientalis (Hansen), for example, is an Indo-Australian species that also reaches Samoa and the Marquesas (Adamson, 1932; Silvestri, 1939). Four other species are reported for Hawaii (Scheller, 1961). Chilopoda Marshalls 1, Enewetak 0—The only centipede defi- nitely reported for the Marshalls is Scolopendra morsitans Linnaeus, a widespread species reported for Jaluit (Schnee, 1904). Townes (1946) stated that Scolopendra occurs on practically every island and is moderately com- mon, but Enewetak was not specifically indicated. Diplopoda Marshalls 3, Enewetak 1—Two unidentified millipedes representing Polyxenus and Trigonoiulus are reported for the Marshalls. Oxidus gracilis (Koch), recently collected from Enewetak, occurs throughout the Pacific; it is com- mon in Hawaii and is easily transported in soil through commerce (Williams, 1931). INSECTA Apterygota Only two orders (Thysanura and Collembola) of these primitive, wingless insects have been reported for Enewetak to date. Although only a few species have been reported, further records can be expected, possibly includ- ing proturans and diplurans. Thysanura Marshalls 1, Enewetak 1—At least one species of Lep- ismatidae is reported for the Marshalls, questionably identi- fied as Lepisma saccharina Linnaeus (Schnee, 1904). Townes (1946) observed a lepismatid on Enewetak, but it was not identified. Collembola Marshalls 2, Enewetak 2—The Isotomurus and Seira we list for Enewetak were recently collected and appear to be the first records of the order for the Marshalls. Uchida (1944) studied earlier collections from Micronesia (Mari- anas and Carolines), with more than half of the 13 species treated appearing to be endemic. Mari Mutt (1979) added another endemic species to the Carolines fauna (Ponape). Pterygota These are insects that have attained flight, including those that have secondarily lost the capability. The bulk of the insects belong to this group, and they are discussed below either at the ordinal or familial level. Odonata Marshalls 7, Enewetak 2—Only one damselfly (Zygop- tera), Ischnura aurora (Brauer), reaches the Marshalls but not Enewetak; it has Oriental affinities. Six dragonflies (Anisoptera) reach the Marshalls, and two are recorded for Enewetak. Of the latter, the Pantala we list for Enewetak is tropicopolitan and widespread in the Pacific, including Hawaii. The Tramea we list is a sight record by E. H. Bryan, Jr.; it is a North American species that reached Hawaii late last century and was likely transported from Hawaii to Enewetak. The four other anisopterans reaching the Marshalls are widespread in Micronesia and beyond, with three found in Australia or on continental islands bordering the western Pacific, and one ranging from Africa through Asia and Australia. The Micronesian fauna, stud- ied by Asahina (1940) and reviewed by Lieftinck (1962), comprises 48 species or subspecies, with endemism about 52%. The monotypic genus Pacificothemis is endemic to the eastern Carolines (Ponape). 160 SAMUELSON AND NISHIDA Blattodea Marshalls 9, Enewetak 7—All the cockroaches reported for the Marshalls are widespread in the Pacific and either include at least the Oriental region (4 species) or occur around the world (5). The Micronesian fauna has not been surveyed, but seven species from Enewetak are present in Bishop Museum. Of these, the Blattella, Lupparia, and Melanozosteria we list are Oriental; the Neostylopyga and Pycnoscelis are Oriental now spread through the tropics; and both Periplaneta are African, with one species now tropicopolitan (australasiae) and one now cosmopolitan (americana). All seven reported for Enewetak include Hawaii in their distributions. Two additional species reported for the Marshalls are the cosmopolitan Blattella| germanica (Linnaeus) and the Oriental Melanozosteria nitida (Brunner). Mantodea Marshalls 1, Enewetak O—Only one Melanesian mantid, Orthodera burmeisteri Wood-Mason, reaches the Marshalls but it is not reported for Enewetak. Beier (1972) reviewed the Micronesian fauna, which comprises five spe- cies. The only mantid endemic to Micronesia is from Palau, and it has Philippine affinities. Phasmatodea Marshalls 1, Enewetak O—Only one species, Megacrania batesi Kirby, is reported for the Marshalls but not Enewetak; it is distributed through Melanesia, Indonesia, and the Philippines. The Pacific walkingsticks were generally surveyed by Nakata (1961). At least nine species are reported for Micronesia, including four endemic to the Carolines and one to the Marianas, but the group there remains poorly collected. Orthoptera Marshalls 10, Enewetak 5—The species reaching the Marshalls are circumtropical (1), widespread in the Old World (1), Oriental (3), Indo-Australian (2), or restricted to Oceania (3). Although the Micronesian fauna has not been surveyed, five species are known from Enewetak, either reported earlier (Townes, 1946; Sugerman, 1972a) or present in Bishop Museum. Of the five we list for Enewetak, the Aiolopus is an acridid of Indo-Australian dis- tribution; the Phisis is a tettigoniid also found in Polynesia (Samoa, Tahiti); and there are three gryllids: the Gryllodes is circumtropical, the Ornebius is Oriental, and the Speonemobius is Oceanian. The Gryllodes and Ornebius reported for Enewetak are also found in Hawaii. The fol- lowing are reported for the Marshalls but are not definitely recorded for Enewetak—Acrididae: Locusta danica (Linnaeus), L. migratoria manilensis (Mayne), Oxya hyla intricata (Stal); Gryllacrididae: Gryllacris sp. near aurantiaca Brunner von Wattenwyl; and Tettigoniidae: Phaneroptera furcifera Stal Isoptera Marshalls 6, Enewetak O—Three of the termites known from the Marshalls are identified: Kalotermes repandus Hill is restricted to Oceania, Coptotermes formosanus Shiraki is an Oriental species spread into the Pacific and elsewhere, and Cryptotermes brevis (Walker) is tropicopolitan (Cole, 1951; Sugerman, 1972a, 1972b). The last two are also reported for Hawaii. The three unidentified termites appear to represent Glyptotermes, Nasutitermes, and Prorhinotermes. The Micronesian fauna has not been monographed. Dermaptera Marshalls 7, Enewetak 3—All of the earwigs reported for the Marshalls are cosmopolitan species. All but one of the seven are reported for Kwajalein. Three of the seven comprise the total earwig fauna for the Caroline atolls, and the greater number in the Marshalls, particularly Kwajalein, is likely a result of heavier international traffic. At least four of the seven also occur in Hawaii, including the three that we list for Enewetak. Micronesian fauna, treated by Brindle (1972), comprises 24 species, with endemism about 46%. Embioptera Marshalls 1, Enewetak O0—Oligotoma (Aposthonia) micronesiae Ross, described from the Marianas, is also reported for the Marshalls (Jaluit) and Gilberts (Butaritari). It and a related species found in the Carolines and Polynesia, O. (A.) oceanica Ross, along with an undescribed member of the subgenus from Ponape, all pos- sibly have Philippines affinities. Micronesian fauna, reviewed by Ross (1955), comprises five species of which two appear to be endemic to Micronesia. Two Oriental species that have spread to other tropical areas also reach Micronesia. Psocoptera Marshalls 16, Enewetak 4—All of these species range beyond the Marshalls, with seven of the 16 restricted to Micronesia, two extending only to Hawaii, three restricted to Oceania, one also in the Oriental region, and three widespread. Of the four psocids reported for Enewetak, one is confined to Micronesia, one to Oceania, and two are widespread. Only the Caecilius and Ectopsocus of the four we list are also reported for Hawaii. Micronesian fauna, reviewed by Thornton et al. (1972), comprises 90 species. Endemism in the fauna is about 44%, with 19 endemic species in the Marianas, 19 more in the Carolines, and two in the Bonins. The number of endemics is diminished in the eastern part of the Carolines, with three on Ponape and one on Kosrae. Psocids are fairly well represented on Caroline atolls, with one of the nine species there endemic; five of the nine also occur in the Marshalls. INSECTS AND ALLIES (ARTHROPODA) 161 Mallophaga Marshalls 22, Enewetak 16—Collections from 11 avian species were made on Enewetak (Elbel, personal communi- cation), and at least 16 species of chewing lice are now known from the atoll. These are mainly from collections made by J. B. Bushman and coworkers, subsequently stud- ied by R. E. Elbel et al. Amerson and Emerson (1971) reported most of the same species for the central South Pacific. Earlier, Woodbury (1962) reported Actorni- thophilus ceruleus (Timmerman) for Enewetak. Six addi- tional chewing lice recorded for the Marshalls, but not Enewetak, are included in the National Museum of Natural History collection: Actornithophilus limosae Kellogg, A. umbrinus Burmeister, Eidmanniella albescens Piaget, Menopon gallinae Linnaeus, Trinoton querquedulae (Lin- naeus), and Pectinopygus sulae (Rudow). Mallophaga are coincident with their avian hosts, and all of the host spe- cies sampled on Enewetak range beyond the Marshalls, with most being widespread over parts of the Pacific. Thir- teen of the 16 species listed for Enewetak are also reported for Hawaii. Anoplura Marshalls 3, Enewetak 0—Two lice reported for the Marshalls are cosmopolitan and associated with man: Pediculus humanis (Linnaeus) and Pthirus pubis (Linnaeus). Another species, Hoplopleura pacifica Ewing, is associated with Rattus and is widespread in the Pacific; it is also reported for the Malayan subregion, Madagascar, Carib- bean, and southeastern United States. Micronesian fauna comprises six species which are mostly widespread and coincident with their hosts; they are mostly associated with man or domestic animals (Ferris, 1959; Wilson, 1972). Thysanoptera Marshalls 2, Enewetak 1—These species are either cosmopolitan, the Haplothrips we list for Enewetak, or Oriental, Taeniothrips vitticornis (Karny); the latter is reported for Kwajalein by Sugerman (1972a). Both are also reported for Hawaii. Micronesian fauna remains incompletely surveyed. Kurosawa (1940) treated 14 species based largely on T. Esaki’s collections; at that time no thrips were reported for the Marshalls. Heteroptera Many of the larger families represented in Micronesia have been studied. Among the groups remaining to be reported upon are the aquatic and semiaquatic families occurring in fresh water; however, they are probably largely absent on isolated atolls. Families treated for Micronesia but lacking records for the Marshalls include Acanthosomidae (Ruckes, 1963), Enicocephalidae (Usinger and Wygodzinsky, 1960), Neididae (Gross, 1963), Plataspi- dae (Ruckes, 1963), Saldidae (Drake, 1961), and Tingidae (Drake, 1956). As with this and most of the following insect orders, each family represented in the Marshalls is reviewed, and when there is a relevant treatise for the group, the author is cited. Anthocoridae (Herring, 1967): Marshalls 3, Enewetak 1—These are species of Pacific or Ethiopian-New World distribution, with one restricted to Micronesia. All three are widespread in Micronesia. The Physopleurella we list for Enewetak also occurs in Hawaii. Micronesian fauna comprises 20 species, with about 70% endemism. Aradidae (Matsuda and Usinger, 1957): Marshalls 1, Enewetak O—Chiastoplonia pygmaea China, described from Samoa, is also reported for Marshalls and Carolines. Three species including the one reported for the Marshalls also occur on Caroline atolls; all three are widespread in Micronesia. Micronesian fauna comprises 40 species, with endemism about 93%. Cimicidae (Usinger and Ferris, 1960): Marshalls 1, Enewetak 0—The bed bug, Cimex hemipterus (Fabricius), is a tropicopolitan species generally distributed in Micronesia. It is reported for the Marshalls but not for Enewetak. Coreidae (Gross and Schaffner, 1963): Marshalls 3, Enewetak 1—These are species either endemic or of Melanesian or cosmopolitan distribution. The endemic species, Riptortus saileri Usinger, is known only from Kwajalein. The Melanesian species, Leptocoris isolata (Dis- tant), occurs throughout the Marshalls but is not reported elsewhere in Micronesia. Micronesian fauna comprises 15 species, with about 13% endemism. Cydnidae (Ruckes, 1963): Marshalls 1, Enewetak 1—Geotomus pygmaeus (Dallas) is a species of Malayan—Oceanian distribution, including Hawaii. Only one other species, from Guam, is reported for Micronesia. Gerridae (Cheng, 1977): Marshalls 2, Enewetak 2—Two marine species were recently collected from Enewetak. Halobates micans (Eschscholtz) is commonly found on the open ocean; it is circumtropical in distribution and is the only member of the genus also occurring in the Atlantic. The Hermatobates remains unidentified; it is apparently undescribed and of limited distribution in the Pacific. Lygaeidae (Barber, 1958): Marshalls 7, Enewetak 4—tThese species are restricted to Micronesia (3), or are of Malayan—Melanesian (2), Indo-Australian—Pacific (1), or Oriental—Pacific (1) distribution. Nysius picipes Usinger, one of the species reported for Enewetak, is restricted to atolls and low islands. The Pachybrachius is the only species of four we list for Enewetak that is also reported for Hawaii; it is mainly of Indo-Australian—Pacific distribu- tion. Micronesian fauna comprises 43 species, with ende- mism about 40%. Miridae (Carvalho, 1956; Schuh, 1984): Marshalls 7, Enewetak 3—These species are tropicopolitan (1), mainly of Old World distribution (1), Oriental—Pacific (2), Australia—Pacific (1), or limited to Oceania (2). Of the Enewetak species, Campylomma eniwetok Schuh is notable because it is one of only three insects thus far reported as endemic to the atoll (the other Enewetak endemics are midges: Ceratopogonidae and Chironomidae). Campy- lomma marshallensis Usinger is restricted to the Marshalls and Gilberts, plus one record for Kanton, Phoenix Islands; the Trigonostylus we list is tropicopolitan. Micronesian 162 SAMUELSON AND NISHIDA fauna comprises 89 species, with endemism about 81%. Four species are restricted to atolls; all are in Campylomma: the two mentioned above, plus C. ulithiensis Carvalho (Ulithi) and C. wakeana Schuh (Wake). Nabidae (Gross, 1963): Marshalls 1, Enewetak 1—Nabis capsiformis Germar is a cosmopolitan species. Micronesian fauna comprises four species of which one is endemic to the Carolines (Palau, Ponape). Pentatomidae (Ruckes, 1963): Marshalls 2, Enewetak 2—tThe Platynopus we list for Enewetak is restricted to Melanesia and Micronesia, and the Oechalia we list is of Australian—Pacific distribution. Micronesian fauna com- prises 33 species, with endemism about 55%. All of these species include high islands in their distributions. Reduviidae (Wygodzinsky and Usinger, 1960): Marshalls 1, Enewetak 0—The only species recorded from isolated atolls is the one reaching the Marshalls, Ploiaria insolida (B. White), which is Philippines—Pacific in distribution. Micronesian fauna comprises 30 species, with endemism about 67%. Veliidae: Marshalls 1, Enewetak 0—Halovelia maritima Bergroth is marine and mainly of Australian—Indonesian distribution; it is also reported for Bikini (Cole, 1951). Homoptera This is the only large insect order reasonably stucied in its entirety for Micronesia. Some of the families are well represented on Micronesian islands and have a high pro- portion of endemic taxa. The following families were treated for Micronesia but representatives are not reported for the Marshalls: Aleyrodidae (Takahashi, 1956), Cercopi- dae (Synave, 1957), Cicadidae (Esaki and Miyamoto, 1975), and Membracidae (Kato, 1960). Aphididae (Essig, 1956): Marshalls 4, Enewetak 2—All are widespread or cosmopolitan species also reported for Hawaii. An additional aphid reported for the Marshalls is Hysteroneura setariae (Thomas) (Sugerman, 1972a). Micronesian fauna comprises 16. species, mostly widespread; endemics include one species and one sub- species of Malayan or Oriental affinities. Cicadellidae (Linnavuori, 1960, 1975): Marshalls 10, Enewetak 7—The species we list for Enewetak are widespread in both hemispheres (1), of New World—Pacific distribution (1), Oriental—Pacific (3), Australia—Pacific (1), or limited to Oceania (1). At least one is reported for Hawaii: Recilia affinis (Osborn). Balclutha lucida (Butler) is reported for Arno and Ebon but not Enewetak; it is of New World-Pacific distribution. Two undetermined cicadel- lids are reported for Kwajalein by Sugerman (1972a, 1979): Empoasca, a genus well represented in the Pacific and Carneocephala, a New World genus not included in Linnavuori’s treatment. Micronesian fauna comprises 87 species, with endemism about 66%. Cixiidae (Fennah, 1956, 1971): Marshalls 2, Enewetak O—Both Myndus apicalis (Metcalf) and M. praecanus Fen- nah are endemic to the central eastern Carolines and Marshalls. Micronesian fauna comprises 29 species, all endemic. Coccoidea (Beardsley, 1966, 1975): Marshalls 36, Enewetak 7—These are mostly species of widespread dis- tribution of which 25 also occur in Hawaii. Seven species are restricted to Micronesia; of these, five are widespread in Micronesia and two are endemic to the eastern Caro- lines and Marshalls. Many of these scale insects are com- mon on cultivated plants and have become widely dis- tributed. All of the species reported for Enewetak also occur on Kwajalein, and five of the seven also occur in Hawaii. Micronesian fauna comprises 150 species, with endemism about 23%. Delphacidae (Fennah, 1956, 1971): Marshalls 9, Enewetak 1—These species are restricted to Micronesia (2), Micronesia plus Hawaii (1), or mainly to Oceania plus continents and continental islands bordering the western Pacific (6). The Sogatella we list for Enewetak is reported for Australia, Philippines, and Micronesia. Of the restricted taxa, Ugyops kinbergi mayas Fennah is endemic to the Marshalls and occurs on both island chains and U. superciliatus Fennah is limited to the eastern Carolines and Marshalls. Micronesian fauna comprises 63 species with endemism about 44%. Three species are endemic to atolls elsewhere: the Gilberts, Wake, and Ocean Island, a raised atoll. Derbidae (Fennah, 1956, 1971): Marshalls 3, Enewetak 1—Species or subspecies restricted to Micronesia of which the Lamenia we list for Enewetak is endemic to the Marshalls; Flaccia dione Fennah is limited to Caroline atolls, Kosrae, and Marshalls; and Swezeyia zephrus Fen- nah occurs through much of Micronesia. Lamenia and Swezeyia possibly have Philippine affinities, while Flaccia has ties to Melanesia (Fiji). Micronesian fauna comprises 56 species, with endemism about 95%. Derbids are well represented on Caroline atolls, with 20 species of which four are endemic to atolls; another species is endemic to the Gilberts. Psyllidae (Tuthill, 1964): Marshalls 1, Enewetak 0—Mesohomotoma hibisci (Froggatt) ranges from Australia and continental islands bordering the western Pacific to Oceania where it is widespread; it is reported for Kwajalein (Sugerman, 1972a). Micronesian fauna comprises 22 species, with endemism about 68%. Two additional psyllids besides M. hibisci occur on Caroline atolls but are not reported for the Marshalls. Coleoptera Most of the large families of beetles have been sur- veyed. The outstanding exception is the Curculionidae or weevils, a family well represented on Pacific islands, with only the Guam components of the Micronesian fauna ade- quately studied. A few species are reported for the Marshalls but not Enewetak. Families monographed for Micronesia, but not yet reported for the Marshalls, include the Cantharidae (Wittmer, 1958), Cryptophagidae (Michi- taka Chijo, 1970), Cybocephalidae (Endrédy-Younga, 1971), Endomychidae (Strohecker, 1958), Lampyridae (Wittmer, 1958), Malachiidae (Wittmer, 1958, 1970), Pla- INSECTS AND ALLIES (ARTHROPODA) 163 typodidae (Wood, 1960), Prionoceridae (Wittmer, 1958), Rhysodidae (Bell and Bell, 1981), and Scaphidiidae (Lobl, 1981). Anobiidae (Ford, 1958): Marshalls 1, Enewetak 0—Lasioderma serricorne (Fabricius), the cigarette beetle, is reported for Kwajalein; it is a cosmopolitan species. Micronesian fauna comprises nine species, with endemism about 67%. Anthicidae (Werner, 1965): Marshalls 3, Enewetak 1—All are species of Anthicus and are confined to the Pacific Basin. Anthicus vexator Werner, reported for Enewetak, occurs on Caroline atolls and the Marshalls and Gilberts. It is also reported for the Phoenix Islands and Hawaii. Anthicus insularis Werner is_ restricted to Micronesia and A. oceanicus LaFerté is widely distributed in Micronesia, extending to Polynesia, Ryukyus, and possi- bly into the Indian Ocean. Micronesian fauna comprises nine species, with endemism about 44%. Anthribidae: Marshalls 1, Enewetak O—Araecerus sp. is reported for Kwajalein (Sugerman, 1972a) and is prob- ably one of several species widespread in the Pacific. The genus is mainly Oriental—Pacific in distribution. Bostrychidae (Michio Chtjo, 1958): Marshalls 1, Enewetak O—Dinoderus bifoveolatus (Wollaston), widely distributed in warmer regions, is reported for Kwajalein. Micronesian fauna comprises 10 species, with none endemic; most are widespread. Carabidae and _ Cicindelidae (Darlington, 1970): Marshalls 5, Enewetak 1—These species are of Oriental—Pacific distribution (1) or Indo-Australian—Pacific (1), of Neotropical affinities (2), or possibly restricted to Oceania (1). The Egadroma we list for Enewetak belongs to the second category. Two species were not treated by Darlington; they are Callida insularis Boheman and Selenophorus sp.; both were reported for Jaluit by Schnee (1904). Micronesian fauna comprises 49 species, with endemism about 27%, representing 11 different stocks, most resulting in only a single endemic species. Curiously, one cicindelid, Therates labiatus (Fabricius), ranging from Melanesia to the Philippines, also occurs on Nukoro, one of the Caroline atolls, but nowhere else in Micronesia. Cerambycidae (Gressitt, 1956): Marshalls 12, Enewetak 1—These species are mostly restricted to Micronesia and have Oceanian—Malayan affinities; close relatives may extend to the Indian Ocean, but they are generally absent on continental Asia. Of these, five are endemic to the Marshalls: two Ceresium are restricted to single atolls, C. unicolor marshallum Gressitt (Majuro) and C. robustum Gressitt (Arno); Prosoplus major Gressitt and P. _hibisci Gressitt are from various atolls, also in the eastern chain (Ratak); and Oopsis marshallensis Gressitt is the most widespread of the endemics, occurring on both island chains. The Prosoplus we list for Enewetak occurs in the eastern Carolines (Ponape) and Marshalls. Only Sybra alternans (Wiedemann) ranges beyond Micronesia; it is also reported for Indonesia, Philippines, and Hawaii. Microne- sian fauna comprises 105 species, with about 87% re- stricted to Micronesia, including 7% endemic to atolls or low islands. Correspondence of species occurring on the different Micronesian atoll groups is fairly low, with three of 12 species from Caroline atolls also present in the Marshalls and one of the four species from the Gilberts likewise found in the Marshalls. Most of the atoll species in the Carolines also occur on adjacent. high islands; however, Sciadella atollorum Gressitt is restricted to atolls. One of the four Gilberts species, Sybra catalana Gressitt, is endemic there. Chrysomelidae (Gressitt, 1955): Marshalls 3, Enewetak 1—These species are either confined to Oceania and their affinities are Melanesian—Malayan (2) or they are Oriental—Pacific in distribution (1). Aphthona_bicolorata Jacoby is associated with euphorbiaceous plants and is fairly common in parts of Oceania, including Enewetak. Brontispa chalybeipennis (Zacher) is associated with the coconut palm and had a rather restricted range, within the eastern Carolines (Ponape, Kosrae) and the Marshalls before it reached Hawaii. Pagria signata (Motschulsky) belongs to the second category. Micronesian fauna comprises about 38 species, with endemism about 66%. None of the species is restricted to atolls. Cleridae: Marshalls 1, Enewetak 0—One cosmopolitan species, Necrobia rufipes DeGeer, ihe red-legged ham bee- tle, is reported for Jaluit (Schnee, 1904). Coccinellidae (Chapin, 1965): Marshalls 8, Enewetak 4—tThese species either are found on continents, Asia and/or Australia (4), or are confined to the Pacific Basin, with two restricted to Micronesia and two also found on subcontinental islands of the western Pacific. One of the restricted species, Scymnus uncinus Chapin, is limited to the Marshalls and Gilberts. One additional species, from India, was released in the Marshalls (Majuro, Uliga) and Guam but was not recovered. Micronesian fauna comprises 48 species, with endemism about 46%. Cucujidae: Marshalls 1, Enewetak 0—The cosmopoli- tan Oryzaephilus surinamensis (Linnaeus), the saw-tootned grain beetle, is reported for Kwajalein (Sugerman, 197 2a). Curculionidae (Zimmerman, 1942, 1948, 1964): Marshalls 6, Enewetak O—These species are possibly cosmopolitan (Sitophilus sp.), of Indonesian—Pacific affini- ties (Oxydema sp.), restricted to Oceania (3), or undeter- mined (1). Of the restricted species, Sphenophorus sulcipes Karsch is described from the Marshalls; Platysimus insularis (Boheman) is reported for Polynesia, Fiji, the Marshalls and Gilberts; and Trigonops hirsuta Zimmerman, described from Guam, is reported for Kwajalein. Weevils of Guam and the Marianas have received attention by Zimmerman (1942, 1948). Dermestidae (Beal, 1961): Marshalls 3, Enewetak 1—Alll of the species reported for the Marshalls are also found in Hawaii, and none is restricted to the Pacific. The Dermestes we list for Enewetak is cosmopolitan and widespread in Micronesia. Orphinus terminale (Sharp) is of Malayan-Pacific distribution and Trogoderma anthrenoides (Sharp) is a Neotropical species that has been established in Hawaii and has also reached Saipan and Kwajalein. Micronesian fauna comprises 12 species, mostly wide- 164 SAMUELSON AND NISHIDA spread; only one species, Orphinus nesioticus Beal, from Guam, is endemic. Elateridae (Van Zwaluwenburg, 1948, 1957; Ohira, 1971): Marshalls 11, Enewetak 2—All identified species are of Oceanian distribution. The two species we list for Enewetak are either reported for New Britain, eastern Carolines, and Marshalls (Simodactylus) or range through Micronesia and Polynesia (Conoderus). Two other species are endemic to the Marshalls and are restricted to single atolls: Melanoxanthus lariversi Van Zwaluwenburg from Arno and _ Simodactylus marshallensis Ohira from Kwajalein. Two unidentified elaterids, Ampedus sp. and Anchastus sp., are reported for Kwajalein (Sugerman, 1972a, 1979). Micronesian fauna comprises 70 species, with endemism about 71%. Histeridae: Marshalls 1, Enewetak 0—Platysoma sp. is reported for Jaluit (Schnee, 1904). The genus is well represented in the Indo-Malayan area, but some species occur in the New World. Hydrophilidae: Marshalls 1, Enewetak 0—At least one species, Dactylosternum abdominale (Fabricius), is reported for the Marshalls (Sugerman, 1972a); it is possibly the same as one or both of the Cyclonotum spp. reported by Schnee (1904) for Jaluit. The species is widely distributed in the New World and is established in Hawaii. Mycetophagidae (Michitaka Chijo, 1970): Marshalls 2, Enewetak 1—The Typhaea we list for Enewetak is a cosmopolitan species. Litargus vestitus Sharp, reported for Arno and Jaluit, is an Oceanian species found throughout Micronesia and parts of Polynesia. Both species are reported for Hawaii. Micronesian fauna comprises three species, with none endemic. Nitidulidae (Gillogly, 1962): Marshalls 14, Enewetak 6—These are mostly widespread species, but one is re- stricted to Micronesia and Polynesia; six are cosmopolitan; three are found on various continents; one is also from Australia; two are also from New Guinea; and one occurs in Polynesia, Philippines, and Sri Lanka. Micronesian fauna comprises 39 species, with endemism about 33%. None is restricted to atolls; correspondence of species among dif- ferent atoll groups is fairly high with six of eight of the Caroline atolls species and seven of the nine Gilberts spe- cies also found in the Marshalls. Oedemeridae (Macnamara and Arnett, in preparation): Marshalls 8, Enewetak 6—These species are more or less of Malayan—Oriental distribution. Of these, only one is rather restricted, occurring only in the Marshalls, Gilberts, and Ocean Island. The remaining seven tend to be widespread in Micronesia and include some that also occur on Melanesian and Polynesian islands. Micronesian fauna comprises 19 species, with most restricted to Micronesia. Propalticidae (John, 1960, 1971): Marshalls 1, Enewetak 1—Propalticus insularis John is restricted to Micronesia. Micronesian fauna comprises eight species, with endemism 63%. Scarabaeidae (Cartwright and Gordon, 1971): Marshalls 6, Enewetak 0—These species are of the subfamily Apho- diinae and are either restricted to Micronesia (2); nearly restricted, occurring also in Samoa (1); found in the Malayan and/or Melanesian subregions (2); or have a gen- eral Old World distribution (1). Micronesian fauna comprises 27 species (10 are aphodiines), with endemism about 37%. Of the endemic species, four are aphodiines, four are melolonthines, one is a ruteline, and one is a dynastine. None is restricted to atolls. Both species occur- ring on Caroline atolls (aphodiines) also occur in the Marshalls. One of the two Gilberts species, also an apho- diine, occurs in the Marshalls. The other species, a dynas- tine, Papuana huebneri (Fairmaire), is the only Micronesian record for that Melanesian group. Scolytidae (Wood, 1960): Marshalls 9, Enewetak 2—Of these species, only one is endemic to a more or less restricted area, Kosrae and the Marshalls including Enewetak; one species reaches Hawaii; one is Malayan; two are cosmopolitan; and the rest tend to be widespread in Micronesia and also occur in warmer regions bordering the western Pacific. Micronesian fauna comprises 53 species, with endemism about 28%. Staphylinidae: Marshalls 2, Enewetak O0—Two rove beetles are reported for Kwajalein by Sugerman (1972a): Lispinus impressicollis Motschulsky, ranging throughout the Old World tropics and extending into the Indian and Pacific Oceans, and a species questionably placed in Medon, a widespread genus with many species. Tenebrionidae (Kulzer, 1957): Marshalls 16, Enewetak 2—Of these species, two are endemic to the Marshalls; one also occurs on Kosrae, eastern Carolines; three are more widespread in Micronesia; three are Oceanian; two are Malayan—Pacific and comprise the Gebieniella and Gonocephalum that we list for Enewetak; one is Oriental; and four are cosmopolitan. Of the species endemic to the Marshalls, Tagalus angustus Kulzer is restricted to Arno and Chariotheca costata Kulzer is restricted to Arno and Ailinglapalap. Micronesian fauna comprises 92 species, with about 62% restricted to Micronesia, including 4% endemic to atolls and low islands. Correspondence of species between atoll groups is fairly low, with four of 13 Caroline atolls species and one of the three Gilberts species also occurring in the Marshalls. One of the Caro- line atolls species, Bradymerus faraulepensis Kulzer, is endemic to a single atoll (Faraulep). Strepsiptera Micronesian fauna unstudied. Neuroptera Only the Chrysopidae has been reported for the Marshalls. Two other families treated for Micronesia are still unreported for the Marshalls: Hemerobiidae (Car- penter, 1961) and Myrmeleontidae (Adams, 1959). Chrysopidae (Adams, 1959): Marshalls 2, Enewetak 1—Both species are widespread in Micronesia and are also of Australian—Malayan distribution. Chrysopa ramburi Schneider, reported for Enewetak, is the senior synonym INSECTS AND ALLIES (ARTHROPODA) 165 of C. jaluitana Kempny (1904). Chrysopa basalis Walker is also reported for the Marshalls (Ailinglapalap). Trichoptera The Micronesian fauna is largely unsurveyed, although a few species are known to occur in the Marianas and Carolines, including remote Kosrae (Gressitt, 1954). Lepidoptera The Micronesian fauna is poorly studied, except the Sphingidae and certain microlepidopteran families. Three families that have been treated for Micronesia, but have not been found as yet in the Marshalls, are Agonoxenidae (Clarke, 1984), Chlidanotidae (Clarke, 1976), and Oeco- phoridae (Clarke, 1984). Arctiidae: Marshalls 2, Enewetak 1—The Utetheisa we list for Enewetak ranges through much of Micronesia and the South Pacific (Schnee, 1904). An undetermined lithos- sine is reported for Kwajalein (Sugerman, 197 2a). Cosmopterygidae: Marshalls 1, Enewetak O—An undetermined Trissodorus is reported for the Marshalls but without mention of Enewetak (Townes, 1946). This species might be T. honorariella (Walshingham), a Pandanus leaf perforator recorded from Sri Lanka and the Pacific, including Hawaii. Danaidae: Marshalls 1, Enewetak 0—Danaus plexippus (Linnaeus) is reported for the Marshalls (Jaluit) by Seitz (1904); it is also reported by Townes (1946) but without specific locality. This is a widespread North American species, also occurring in Hawaii and elsewhere in the Pacific, including Australia. Gelechiidae: Marshalls 1, Enewetak O—Stoeberhinus testaceus Butler, of Indonesian—Pacific distribution, including Hawaii, is reported for Kwajalein (Sugerman, 1972a). Geometridae: Marshalls 1, Enewetak O—An undeter- mined Thalassodes is reported for Jaluit by Seitz (1904). A number of species occur in the South Pacific. Hesperiidae: Marshalls 1, Enewetak 1—The Badamia we list for Enewetak (Townes, 1946) is widely distributed in the Pacific, including Hawaii (as a recent introduction); it also occurs in the Oriental and Australian regions. Lycaenidae: Marshalls 1, Enewetak 0—The Lampides reported by Sugerman (1979) is probably L. boeticus (Lin- naeus), a blue butterfly distributed throughout the Old World and Pacific, including Hawaii. Noctuidae (Fukushima, 1947, in part): Marshalls 13, Enewetak 2—These species are mostly widespread: Anticarsia irrorata (Fabricius), Mocis frugalis (Fabricius), Nagia linteola (Guenée), Plusia chalcites (Esper), Spodoptera exempta (Walker), S. litura (Fabricius), and S. mauritia (Boisduval) occur through warmer regions of the Old World and extend into the Pacific; Achaea janata Linnaeus, Anua coronata (Fabricius), Bocana manifestalis Walker, Calogramma festiva (Donovan), and Platysenta_ illecta (Walker) are Oriental or Indo-Australian—Pacific; and Nagia hieratica Hampson is restricted to Micronesia. The last is described from material taken in the Marshalls and Gilberts and is also reported for the Carolines (Yap, Ponape). Of the two Spodoptera we list for Enewetak, one (exempta) is common in Hawaii. we Nymphalidae: Marshalls 2, Enewetak 2—These butterflies are Indo-Australian—Pacific (Hypolimnas) or Australian—Pacific (Precis) in distribution. The former, H. bolina, exhibits much variation, and specimens from the Marshalls have received the subspecific or infrasubspecific names: inconstans, jaluita, or pallescens. Clark (1951) treated the butterflies of the Marshalls. Olethreutidae (Clarke, 1976): Marshalls 3, Enewetak O0—tThese species are either Oriental extending into the Pacific and Indian Oceans (2) or are restricted to Micronesia (1). Heleana p. physalodes (Meyrick) and Stratherotis leucaspis (Meyrick) belong to the first category. The restricted species, Icelita monela Clarke, is reported for the southern Marianas, Marshalls, and Gilberts. Micronesian fauna comprises 58 species, with endemism about 55%. Pyralidae: Marshalls 7, Enewetak 0—These are mostly widespread species: Marasmia trapezalis Guenée occurs throughout the world; Hymenia recurvalis (Fabricius), Nacoleia diemenalis Guenée, and Syngamia floridalis (Zeller) are of Old World—Pacific distribution; Herpetogramma licarsisalis (Walker) is Oriental; Piletocera signiferalis (Wal- lengren) ranges from Australia to Polynesia (Marquesas); and one is undetermined (Diaphania). The last is a large world-distributed genus of warmer regions. All of these species are reported for Kwajalein by Sugerman (1972a, 1979). The Hymenia appears to be the only one of the above that is also reported for Hawaii. Sphingidae (Riotte, in preparation): Marshalls 4, Enewetak 4—We list all of these species for Enewetak; they are either restricted to Oceania (1) or also occur on continental islands or continents bordering the western Pacific (3). The restricted one is Cephonodes armatus R. and J. Micronesian fauna comprises 18 species, with none endemic. All seem to have Old World affinities and many tend to be widespread in the Eastern Hemisphere. Although these insects are large-bodied, they are highly vagile and are able to reach oceanic islands far from con- tinents; this may help to explain the apparent absence of endemic forms in Micronesia. Tineidae: Marshalls 2, Enewetak 0—Decadarchis simulans (Butler) and Opogona omoscopa (Meyrick) range from Africa to Australia and into the Pacific, including Hawaii. Both are based on single records for the Marshalls and are tentatively identified (Sugerman, 1972a, 1979). Tortricidae (Clarke, 1976): Marshalls 1, Enewetak 1—Adoxophyes faciculana (Walker) mainly ranges through the Malayan and Melanesian subregions, with the only records for Micronesia being from Kosrae and Enewetak. Micronesian fauna comprises seven species, with endemism about 86%. 166 SAMUELSON AND NISHIDA Diptera While many of the families represented in Micronesia have been surveyed, a few remain incompletely studied at this time; these include the Asilidae, Cecidomyiidae, Doli- chopodidae, Tachinidae, and various acalyptrate families. Families monographed for Micronesia, but not represented in the Marshalls, include the Asteiidae (Sabrosky, 1957), Bibionidae (Hardy, 1956), Clusiidae (Steyskal and Sasakawa, 1966), Coelopidae (Hardy, 1957), Empididae (Quate, 1960), Mycetophilidae (Colless, 1966), Micropezi- dae (Aczel, 1959), Nycteribiidae (Theodor, 1966), Pipun- culidae (Hardy, 1956), Scenopinidae (Hardy, 1958), Simuliidae (Stone, 1964), Streblidae (Maa, 1966), and Tabanidae (Stone, 1960). Agromyzidae (Spencer, 1963): Marshalls 2, Enewetak 2—Species either ranging from Melanesia through Indonesia to islands in the Indian Ocean (Ophiomyia) or restricted to the Pacific, Australia, and Taiwan (Pseudonapomyza). Both species are widely distributed in Micronesia. Micronesian fauna comprises 19 species, with endemism about 32%. Both species reported for the Marshalls also occur in the Gilberts and Caroline atolls; however, the Caroline atolls have three additional species not recorded for the Marshalls; no additional species are recorded for the Gilberts. Asilidae: Marshalls 2, Enewetak 0—Two unidentified species representing two genera (Clinopogon and Stenopogon) are reported for the Marshalls (Cole, 1951; Sugerman, 1972a, 1972b). Both genera are represented in the Oriental region. Calliphoridae (James, 1962): Marshalls 6, Enewetak 2—These are mostly species of broad distribution, being circumtropical (1), Holarctic and reaching many other areas (1), Indo-Australian (2), ranging from Africa to the Pacific (1), or Malayan—Pacific (1). The two species reported for Enewetak are circumtropical [Phaenicia cuprina (Wiedemann)] or Indo-Australian [Chrysomya megacephala (Fabricius)]. All of the above are widespread in Micronesia except the Holarctic Phaenicia sericata (Meigen). Micronesian fauna comprises 11 species, with endemism low at around 9% (one endemic species occurs in the Bonins). The nonendemic fauna is at least partly coincident with man in the Pacific. Over half of the Micronesian fauna is represented in the Marshalls. All three species reported on Caroline atolls and three of the four species reported for the Gilberts also occur in the Marshalls. Canaceidae (Wirth, 1951): Marshalls 2, Enewetak 1—These are either endemic to the Marshalls (Nocticanace marshallensis Wirth) or restricted to the Marshalls and eastern Carolines (Kusaie) (Procanace townesi Wirth). The latter is not yet reported for Enewetak. Micronesian fauna comprises two species, with endemism 100%. Ceratopogonidae (Tokunaga and Murachi, 1959): Marshalls 23, Enewetak 5—These species are mostly con- fined to Micronesia (19), including three endemic to the Marshalls; four also occur on continental islands bordering Asia. Of the three endemics, Forcipomyia tuthilli Tokunaga is one of three of the only known insects restricted to Enewetak Atoll at this time; Dasyhelea nigristigmata Toku- naga and Murachi is restricted to Ailinglapalap and Dasyhelea sp. No. 1 T. and M. is restricted to Arno. Two species are limited to the eastern Carolines (Ponape or Kosrae) and the Marshalls: Dasyhelea flavibasalis Tokunaga and D. pallivittae Tokunaga. Micronesian fauna comprises 147 species, with endemism about 90%. Ten of the 16 species reported from Caroline atolls also occur in the Marshalls, as do five of the nine Gilberts species, showing that there is a fair correspondence of these midges on Micronesian atolls. Chironomidae (Tokunaga, 1964): Marshalls 16, Enewetak 5—These species are mostly confined to Micronesia, including two endemic to the Marshalls. Eight occur beyond Micronesia and mostly range no farther than the Malayan subregion, Japan, or Polynesia. Of the Marshall Island endemics, Clunio tuthilli Tokunaga is re- stricted to Enewetak, and Cricotopus sp. No. 1 Tokunaga is restricted to Kwajalein. Clunio tuthilli is notable because, thus far, it is one of only three known insects restricted to Enewetak. Micronesian fauna comprises 100 species, with endemism about 79%. Some species are marine. Chloropidae: Marshalls 3, Enewetak 2—Two species from Enewetak are among the Micronesian specimens under study by C. W. Sabrosky; both are presumably widespread, including Hawaii. Another chloropid, Eutropha noctilux (Walker), is reported for Kwajalein (Sugerman, 1972a). Culicidae (Bohart, 1956): Marshalls 4, Enewetak O—tThese are mostly widespread through the tropics or subtropics (2), ef Australian—Malayan plus Melanesian dis- tribution (1), or restricted to the eastern Carolines (Kos- rae), Marshalls, and Gilberts (1). The last mentioned is Aedes marshallensis Stone and Bohart. Micronesian fauna comprises 47 species, with endemism about 64%. The fauna of the Caroline atolls possesses 10 species, of which only three are reported for the Marshalls; the two species reported for the Gilberts also occur in the Marshalls. Dolichopodidae: Marshalls 5, Enewetak O—Chryso- soma complicatum Becker and C. fraternum Van Duzee are apparently restricted to Pacific islands, with the latter described from Hawaii, while C. leucopogon (Wiedemann) ranges from Africa through Asia into the Pacific. Campsicnemus and Cymatopus are each represented by an unidentified species from the Marshalls. The former genus contains numerous endemics in Hawaii. These species are variously listed by Cole (1951) and Sugerman (1972a, 1972b, 1979). Drosophilidae (Wheeler and Takada, 1964): Marshalls 5, Enewetak 0—These species are cosmopolitan or tropi- copolitan (2), Old World tropics (1), Australian— Malayan-Pacific (1), or restricted to Micronesia (1). The restricted species is endemic to the eastern Carolines (Ponape, Kosrae) and the Marshalls. Micronesian fauna comprises 70 species, with endemism about 59%. About 43% of all Micronesian species are restricted to single INSECTS AND ALLIES (ARTHROPODA) 167 island groups. Only one species is endemic to atolls (Kapingamarangi, eastern Carolines). Of the species recorded from Caroline atolls, only two of the eight also occur in the Marshalls, but both of the two Gilberts species occur in the Marshalls. Ephydridae: Marshalls 5, Enewetak 5—Five species from Enewetak are among the Micronesian specimens under study by Wirth. These species are fairly widespread in the Pacific, and all occur at least on continental islands of the western Pacific, including two also found in Aus- tralia and one ranging west to India and the Seychelles. Hippoboscidae (Maa, 1966): Marshalls 3, Enewetak 1—All are ectoparasites of birds; two are pantropical and one, Ornithoica pusilla (Schiner), is restricted to the Marshalls (Arno) and Polynesia (Tokelaus, Tuamotus). Olfersia spinifera (Leach) has been recorded from Enewetak and also leeward Hawaiian islands. Micronesian fauna, six species, comprises two species parasitic on marine birds, two generally parasitic on migratory land birds, and two generally parasitic on breeding land birds. The dispersal of hippoboscids is apparently keyed to the migration routes of hosts. Maa mentioned two routes, through Siberia to New Zealand via the Marianas and through the Philippines into the Carolines via Palau. The second is the more important for hippoboscids. The fauna has Oriental affinities. Lauxaniidae: Marshalls 2, Enewetak 0—Two unidenti- fied species are reported for the Marshalls representing Homoneura and possibly Prosopomyia (Cole, 1951; Suger- man, 1972a, 1972b, 1979). The former has many species in the Oriental region; the latter is Palearctic. Lonchaeidae: Marshalls 3, Enewetak O—Lamprolon- chaea aurea (Macquart) is reported for Bikini (Cole, 1951) and L. metatarsata (Kertész) is reported for Kwajalein (Sugerman, 1979). The former is of Mediterranean origin, and the latter is of Malayan—Pacific distribution. An undetermined Lonchaea is also reported for Kwajalein. Milichiidae: Marshalls 3, Enewetak 2—The two species we list for Enewetak are virtually cosmopolitan. A third species, Desmometopa singaporensis Kertész, reported for Kwajalein (Sugerman, 1979), is Oriental—Pacific in distribution. Muscidae (Snyder, 1965): Marshalls 8, Enewetak 3—These are either widespread or cosmopolitan species (4), those more or less restricted to the Pacific and con- tinents and islands bordering the western Pacific (3), or undetermined (1). The Enewetak muscids all belong to the first category. An undetermined Lispe from Kwajalein, listed by Sugerman (1979), is tentatively assigned to the last category. Micronesian fauna comprises 84 species, with endemism about 63%. Only one species, Atherigona tobi Snyder, is restricted to atolls or low islands. Seven Marshalls muscids occur on Caroline atolls, which have 16 species reported. All three of the Gilberts species also occur in the Marshalls. Neriidae (Aczel, 1959): Marshalls 1, Enewetak O—Telostylinus longicoxa (Thomson) is widespread in Micronesia. Micronesian fauna comprises four species, with endemism 100%; three are of limited distribution in Micronesia, being endemic to Palau, Yap, or Palau and western Caroline atolls. Otitidae: Marshalls 1, Enewetak O—Pseudeuxesta prima (Osten Sacken), reported for Kwajalein (Sugerman, 1972a), is Indonesian—Pacific in distribution, including Hawaii. Phoridae (Beyer, 1967): Marshalls 4, Enewetak 2—tThese species are either restricted to Micronesia (2) or nearly cosmopolitan (both Enewetak records). One of the restricted species occurs only in the eastern Carolines (Ponape, Kosrae) and the Marshalls. Micronesian fauna comprises 25 species, with endemism about 64%. Most of the extra-Micronesian species are either widespread or Malayan-Pacific in distribution, but two species range no farther than Hawaii. Platystomatidae: Marshalls 2, Enewetak O—Scholastes carolinensis Enderlein, reported for Kwajalein and Lib, is possibly restricted to the Carolines and Marshalls, while Plagiostenopterina aenea (Wiedemann), reported for Kwajalein, is Indo-Australian (Sugerman, 1972a, 1972b). Psychodidae (Quate, 1959): Marshalls 2, Enewetak O—These are species also reaching Melanesia or Polynesia (Hawaii). Micronesian fauna comprises 34 species, with about 71% endemism. Sarcophagidae (de Souza Lopes, 1958, 1963): Marshalls 5, Enewetak 3—These are species widespread in Micronesia and that also occur in Hawaii (1), Malayan subregion (2), or continents and islands bordering the western Pacific basin (1) or are species of nearly worldwide distribution (1). Micronesian fauna comprises 14 species, with about 36% endemism. Correspondence of species on atolls is fairly uniform, with four of five species reported for the Gilberts also occurring in the Marshalls. Scatopsidae (Hardy, 1956): Marshalls 2, Enewetak 1—The Enewetak record is new, a specimen questionably determined as Holoplagia guamensis (Johannsen) by W. W. Wirth. Holoplagia guamensis is otherwise distributed in the Marianas (Guam) and Carolines, including atolls, and also Hawaii, the only non-Micronesian locality. The other species reaching the Marshalls, but not Enewetak, is Swammerdamella albimana Edwards; it also occurs in Fiji and Samoa. Micronesian fauna comprises six species of which one is endemic to the Carolines, three are Micronesian—Hawaiian, one also occurs in Melanesia— Polynesia, and one is cosmopolitan. Sciaridae (Steffan, 1969): Marshalls 7, Enewetak 3—Two of these species are endemic to the Marshalls, two are not widely distributed in Micronesia but also occur in Hawaii, two also occur in the Oriental region, and one is nearly cosmopolitan. Of the endemics, Scythropochroa trispinosa Steffan is restricted to Kwajalein and Plastosciara jaluitensis Steffan is restricted to Jaluit. Micronesian fauna comprises 22 species, with endemism about 77%, but some species may have broader distribution than presently indicated. The fauna apparently has Oriental and possibly Papuan affinities. 168 SAMUELSON AND NISHIDA Sphaeroceridae (Richards, 1963): Marshalls 4, Enewetak O—These are species restricted to Micronesia (2), or otherwise occurring in the Malayan subregion (1), or cosmopolitan (1). Micronesian fauna comprises 21 species, with endemism about 38%. The single species reported for Caroline atolls, as well as the two species reported for the Gilberts, also occurs in the Marshalls. Stratiomyidae (James, 1962): Marshalls 4, Enewetak 1—These are either island species restricted to Micronesia (1), those occurring also in Melanesia and islands of the Indian Ocean (2), or widespread species of the tropics and subtropics (1). The restricted species, Cephalochrysa rugulosa James, occurs in the eastern Carolines (Ponape, Kosrae) and the Marshalls. The widespread species, Hermetia illucens (Linnaeus), occurring in all warm regions, is of American origin. Micronesian fauna comprises 24 species, with endemism about 63%. The three species occurring on Caroline atolls and the two occurring in the Gilberts are also found in the Marshalls, making the atoll stratiomyid fauna fairly uniform. Syrphidae (Shiraki, 1963): Marshalls 3, Enewetak 1—The Ischiodon we list for Enewetak is a widespread Indo-Australian species occurring all over Micronesia. Also reported for the Marshalls are Syritta orientalis Macquart, ranging through southern Asia into the Pacific, and Eristalis calliphoroides Shiraki, restricted to the Carolines and Marshalls. Micronesian fauna comprises 27 species, with endemism about 70%. Tachinidae: Marshalls 2, Enewetak O—The Nearctic Lespesia archippivora (Riley) probably reached Kwajalein via Hawaii. An unidentified Exorista, also reported for Kwajalein, belongs to a genus well represented in the Old World. Both species are reported by Sugerman (1972a). Tephritidae (Hardy and Adachi, 1956): Marshalls 3, Enewetak O—These species are circumtropical [Dioxyna sororcula (Wiedemann)], ranging from Africa through Asia into Australia and the Pacific (Spathulina acroleuca Schiner), or Malayan—Pacific in distribution (Dacus frauenfeldi Schiner). Of these, only D. sororcula is reported for Hawaii. Tethinidae: Marshalls 1, Enewetak 0—Dasyrhicnoessa insularis (Aldrich), described from Hawaii, is reported for Lib Island (Sugerman, 1972b). Tipulidae (Alexander, 1972): Marshalls 9, Enewetak 1—Excepting the circumtropical Limonia umbrata (de Meijere), all of these species are confined to the Pacific, with most restricted to Micronesia; two are endemic to the Marshalls. The most widely distributed of the preceding, Styringomyia didyma Grimshaw, ranging from New Guinea to the Tuamotus and northward in the Pacific, was described from Hawaii. The two endemic species are Limonia_ beardsleyi Alexander from Namu, Kili, and Namorik and L. sentifera Alexander from Namu and Arno. Micronesian fauna comprises 71 species, with endemism about 72%. One of the species from the Marshalls also occurs on Caroline atolls for which five species are reported, including two that are more or less endemic (one may also occur on Ponape). The three species reported for the Gilberts also occur in the Marshalls. Siphonaptera (Hopkins, 1961): Marshalls 1, Enewetak O—Only Ctenocephalides f. felis (Bouche), the cat flea, is reported thus far for the Marshalls but not Enewetak. Micronesian fauna comprises six species that are mostly cosmopolitan species associated with man, domestic mammals (particu- larly Carnivora), and rats. One species, a bat flea, is found mainly in the Oriental region and reaches Japan and apparently Guam (a single record). Hymenoptera The Micronesian fauna is partly studied but remains poorly surveyed in general. Groups that are especially in need of study include the Braconidae, Formicidae, and many families of Chalcidoidea and Proctotrupoidea. Fami- lies essentially monographed for Micronesia but not reported for the Marshalls include Colletidae (Krombein, 1950), Encyrtidae (Yoshimoto and Ishii, 1965), Halictidae (Krombein, 1950), Mymaridae (Doutt, 1955), Scoliidae (Krombein, 1949, 1950), Stephanidae (Townes, 1958), and Trichogrammatidae (Doutt, 1955). Anthophoridae (Krombein, 1950, in part): Marshalls 1, Enewetak 1— Xylocopa sonorina Smith is a carpenter bee of Oriental affinities introduced to Hawaii and elsewhere in the Pacific. Micronesian fauna comprises six species, including two of Ceratina endemic to the Marianas or Carolines. Bethylidae: Marshalls 1, Enewetak 0O—An unidentified Scleroderma is reported for Kwajalein (Sugerman, 1972a). The genus is widespread and occurs in Hawaii. Braconidae: Marshalls 4, Enewetak 1—Four genera are known for the Marshalls, and all are single records with none of the species identified (Cole, 1951; Sugerman, 1972a, 1979; Townes, 1946). They are Chelonus, Macrocentrus, Microplitis, and Zele. Only the last is reported for Enewetak. All are widespread genera whose species parasitize Lepidoptera. Chelonus and Macrocentrus are known from Hawaii. Watanabe (1945) studied the Esaki collections from the Carolines and Marianas, includ- ing three endemic species of Chelonus and two of Macrocentrus. Chalcidae: Marshalls 1, Enewetak 0—An unidentified species, possibly Chalcis, is reported for Jaluit (Schnee, 1904). Cynipidae: Eucoilinae (Yoshimoto, 1962): Marshalls 3, Enewetak 0—These species, all in Pseudeucoila, are either restricted to Micronesia (2) or also occur in Samoa and Hawaii (1). Micronesian fauna comprises 13 species, with endemism about 77%. Two other species of Pseudeucoila reported for Caroline atolls are known from high islands in the Carolines and elsewhere but are not among those reported for the Marshalls. Eucharidae (Watanabe, 1958): Marshalls 1, Enewetak O—A single species, Parachalcura maculata Watanabe, is restricted to the Marshalls and Caroline atolls. Micronesian fauna comprises nine species, with all species endemic. Eulophidae (Yoshimoto and Ishii, 1965): Marshalls 5, Enewetak 1—These are species either restricted to INSECTS AND ALLIES (ARTHROPODA) 169 Micronesia (4) or also occurring in Australia, Philippines, and Hawaii (1). The last is also reported for Enewetak. One of the restricted species, Cirrospiloideus fullawayi Y. and I., is reported only from Ebon. Micronesian fauna comprises 35 species, with endemism about 60%. Only three of the 11 species occurring on Caroline atolls and one of the two species occurring in the Gilberts are also present in the Marshalls, making the atoll eulophid fauna irregular. Evaniidae (Townes, 1958): Marshalls 2, Enewetak 1—These species are either circumtropical [Evania appendigaster (Linnaeus)] or Malayan—Melanesian and extending farther into the Pacific, including Enewetak and Hawaii (Szeptigetella). Micronesian fauna comprises three species, with none endemic. Formicidae: Marshalls 22, Enewetak 1—Ants from Enewetak are poorly sampled, and the 13 species listed by Cole (1949) for Bikini should also occur there. The sole species we list for Enewetak, Monomorium pharaonis (Lin- naeus), is cosmopolitan. Species reported for the Marshalls are widespread (11), Oriental (8), more or less restricted to the Pacific (2), or Australian (1). At least 13 of these ants also occur in Hawaii, being mainly “tramp species“ that are widely distributed by commerce. Wilson and Taylor (1967) treated the Polynesian fauna, which includes many species of possible occurrence in Micronesia. Ichneumonidae (Townes, 1958): Marshalls 2, Enewetak 1—The polytypic Echthromorpha agrestoria Swederus ranges from Africa through Asia into the Pacific and includes three subspecies endemic to Micronesia, plus a further subspecies (insidiator) of Indonesian—Melanesian dis- tribution that is reported for Enewetak. The other species, Trathala flavoorbitalis (Cameron), is Oriental. Both species occur in Hawaii. Micronesian fauna comprises 33 species, with endemism probably less than 60%. Only 12% are reported for isolated atolls, but none occurs exclusively on atolls. Megachilidae (Krombein, 1950): Marshalls 3, Enewetak 2—Two species of Megachile reported for Enewetak are either of Oceanian distribution (hedleyi) or possibly of Philippine origin and are now found throughout Micronesia and Hawaii (fullawayi). A third species, M. umbripennis Smith, is Oriental and was not seen in Micronesia until after World War Il; it is now reported for the Marianas and Marshalls and also occurs in Hawaii. Micronesian fauna comprises eight species, with endemism about 38%. Scelionidae: Marshalls 2, Enewetak 0—Two unidenti- fied species are reported for Kwajalein (Sugerman, 1979); they represent Telenomus and possibly Caloteleia. Sphecidae (Krombein, 1949, 1950): Marshalls 11, Enewetak 4—The species from Enewetak are from con- tinental areas, probably reaching Pacific islands, including Hawaii, through human activities. Species reported for the Marshalls are widespread in the Old World (2), Oriental region (3), New World (3), or limited to Pacific islands (3). Species in the last category all belong to Pison. Nine of 11 species reaching the Marshalls also occur in Hawaii, includ- ing the four reported for Enewetak. It is probable that at least some of the sphecids gained entry to the Marshalls via Hawaii. Micronesian fauna comprises 30 species, with endemism about 17%. Vespidae (Krombein, 1949): Marshalls 4, Enewetak 4—The species of Pachodynerus and Polistes we list for Enewetak are from North America, introduced to Hawaii and Micronesia; the Ropalidia is Malayan. The distribution of Odynerus sp. is undetermined, but there are three endemic congeners described from the Marianas. Microne- sian fauna comprises 12 species, with endemism about 58%. COLLECTION RECORDS Insects and Allies of Enewetak Atoll Taxa are listed in the same order as given in the checklist (Table 1). Collection records reported are abbre- viated and include, when known, the islet, month, year, and surname of collector. The specific islet was unclear in some cases, and these are listed as “?Islet” or by a descriptive phrase (see first entry). The name of the deter- miner is included when the particular specimens cited were previously unreported for Enewetak. Class ARACHNIDA Order SCORPIONIDA Family ISCHNURIDAE Hormurus australasiae Fabricius “On every island visited”: May 1946, Townes. Order PSEUDOSCORPIONIDA Family ATEMNIDAE Oratemnus samoanus whartoni Chamberlin Enewetak Islet: Nov. 1944, Dybas and Edgar. Japtan Islet: Oct.—Nov. 1944, Dybas. Family CHERNETIDAE Haplochernes insulanus Beier Japtan Islet: Nov. 1944, Dybas. Family CHTHONIIDAE Lechytia sakagamii Morikawa Enewetak Islet: Nov. 1944, Dybas. Japtan Islet: Nov. 1944, Dybas. Order ARANEAE Family ARANEIDAE Araneus theis (Walckenaer) Ikuren Islet: Sept. 1975, Cheng. Medren Islet: Sept. 1975, Cheng—det. by L. J. Pinter. Family CLUBIONIDAE Chiracanthium diversum Koch Japtan Islet: Sept. 1975, Cheng. Medren Islet: Sept. 1975, Cheng—det. by L. J. Pinter. Family DIPLURIDAE Masteria hirsuta Koch Enewetak ?Islet: June 1946, Morrison. Family EODSPARASSIDAE Heteropoda venatoria (Linnaeus) Enewetak Islet: Sept. 1975, Cheng—det. by L. J. Pinter (determined from cast exoskeleton). 170 SAMUELSON AND NISHIDA Family MICRYPHANTIDAE Erigone sp. Enewetak Islet: Sept. 1975, Cheng—det. by L. J. Pinter. Family OONOPIDAE Opopaea foveolata Roewer Enewetak ?Islet: Nov. 1944, Dybas. Opopaea lena Suman Enewetak Islet: Sept. 1975, Cheng. Medren Isiet: Sept. 1975, Cheng—det. by L. J. Pinter. Family PHOLCIDAE Smeringopus elongatus (Vinson) Enewetak Islet: Sept. 1975, Cheng—det. by L. J. Pinter. Family SALTICIDAE Flacillula minuta (Berland) Japtan Islet: Sept. 1975, Cheng—det. by L. J. Pinter. Vitia albipalpis Marples Enewetak Islet: Sept. 1975, Cheng—det. by L. J. Pinter. Family SICARIIDAE Scytodes striatipes (Koch) Japtan Islet: Nov. 1944, Dybas; July 1946, Townes. Family THERIDIIDAE Latrodectus geometricus (Koch) Enjebi Islet: Date?, Lamberson—det. by L. J. Pinter. ?lslet: May 1946, Townes. Order ACARI Family ARGASIDAE Ornithodoros capensis Neumann Aej, Ananij, Boken, Jinimi, Ribewon, and other Islets: Spring 1962, Bushman or coworkers—det. by D. E. Johnson. Family DERMANYSSIDAE Ornithonyssus sp. ?Islet: 1962, Bushman or coworkers—det. by N. Wilson. Family ERYTHRAEIDAE Balaustium sp. ?Islet: May 1946, Townes. Class DIPLOPODA Order PROTEROSPERMOPHORA Family STRONGYLOSOMIDAE Oxidus gracilis (Koch) Enewetak Islet: Sept. 1975, Cheng—det. by F. G. Howarth. Class INSECTA Order THYSANURA Family LEPISMATIDAE Not identified ?Islet: May 1946, Townes. Order COLLEMBOLA Family ENTOMOBRYIDAE Seira bipunctata (Packard) Enewetak Islet: Sept. 1975, Cheng—det. by P. Bellinger. Family ISOTOMIDAE Isotomurus tricuspis Borner ? ?Islet: Sept. 1975, Cheng—det. by P. Bellinger. Order ODONATA Family LIBELLULIDAE Pantala flavescens (Fabricius) Enjebi Islet: Date?, Jackson. Tramea lacerata Hagen Medren Islet: Feb. 1975, Bryan—sight record by E. H. Bryan, dr. Order BLATTODEA Family BLATTELLIDAE Blattella lituricollis (Walker) 2 Enjebi Islet: Jan. 1951, Oshiro. Japtan Islet: Aug. 1956, Tuthill—det. by S. Asahina. Lupparia notulata (Stal) i Bokombako Islet: Dec. 1950, Oshiro. Japtan Islet: Jan. 1957, Tuthill; Feb. 1957, Tuthill—det. by S. Asahina. Family BLATTIDAE Melanozosteria soror (Brunner) Ananij Islet: Aug. 1956, Tuthill. Bokombako Islet: Dec. 1950, Oshiro—det. by S. Asahina. Enewetak Islet: Sept. 1975, Cheng—det. by G. M. Nishida. Neostylopyga rhombifolia (Stoll) Enewetak Islet: Sept. 1975, Cheng—det. by G. M. Nishida. Periplaneta americana (Linnaeus) Enewetak Islet: Sept. 1975, Cheng—det. by F. G. Howarth. Enjebi Islet: Jan. 1951, Oshiro—det. by S. Asahina. Periplaneta australasiae (Fabricius) Enjebi Islet: May 1946, Oakley and Townes. Japtan Islet: Aug. 1956, Tuthill; Sept. 1956, Tuthill. Medren Islet: Aug. 1956, Tuthill. Family PYCNOSCELIDAE Pycnoscelis surinamensis (Linnaeus) Ananij Islet: Aug. 1956, Tuthill. Elugelab Islet: Jan. 1951, Oshiro Enewetak Islet: Nov. 1944, Dybas. Enjebi Islet: May 1946, Oakley. Japtan Islet: Aug. 1956, Tuthill; Jan. 1957, Tuthill—det. by S. Asahina. Medren Islet: Sept. 1975, Cheng—det. by F. G. Howarth. Order ORTHOPTERA Family ACRIDIDAE Aiolopus thalassinus tamulus (Fabricius) Aomon Islet: May 1946, Townes Enewetak Islet: May 1946, Townes; Sept. 1975, Cheng. Japtan Islet: May 1946, Townes; Sept. 1975, Cheng—det. by G. M. Nishida. Family GRYLLIDAE Ornebius bimaculatus (Shiraki) Enewetak Islet: Sept. 1975, Cheng—det. by F. G. Howarth. Gryllodes sigillatus (Walker) Enewetak Islet: Sept. 1975, Cheng—det. by F. G. Howarth. Speonemobius sp. prob. tigrinus (Saussure) Japtan Islet: Sept. 1975, Cheng—det. by A. B. Gurney. Family TETTIGONIIDAE Phisis pallida (Walker) “All islands or atolls visited”: May 1946, Townes. Order DERMAPTERA Family CARCINOPHORIDAE Euborellia annulipes (Lucas) Enewetak Islet: Sept. 1975, Cheng—det. by G. M. Nishida. Enjebi Islet: May 1946, Townes and Oakley [published as 1949]: Nov. 1947, Dybas; Aug. 1956, Tuthill. Family CHELISOCHIDAE Chelisoches morio (Fabricius): “on every island visited”: May 1946, Townes. Family LABIDURIDAE Labidura riparia (Pallas). Enewetak Islet: Sept. 1975, Cheng—det. by G. M. Nishida. Enjebi Islet: May 1946, Townes; Jan. 1951, Oshiro. ? Islet: May 1946, Townes; Jan. 1951, Oshiro. Order PSOCOPTERA Family CAECILIIDAE Caecilius casarum Badonnel INSECTS AND ALLIES (ARTHROPODA) 71 Japtan Islet: Feb. 1957, Tuthill. Family ECTOPSOCIDAE Ectopsocus maindroni Badonnel Enewetak Islet: Sept. 1975, Cheng—det. by I. W. B. Thorn- ton. Japtan Islet: May 1946, Townes. Family LEPIDOPSOCIDAE Cyptophania marginata Thornton, Lee and Chui Enewetak Islet: Nov. 1944, Dybas. ?Islet: Nov. 1944, Edgar. Family PSOCIDAE Ptycta angulata Thornton, Lee and Chui Ikuren Islet: Aug. 1956, collector? Medren Islet: Sept. 1975, Cheng—det. by I. W. B. Thornton. Order MALLOPHAGA Family MENOPONIDAE Actornithophilus bicolor (Piaget) Medren Islet: May 1962, Bushman—det. by K. C. Emerson. Actornithophilus ceruleus (Timmermann) Aej Islet: Mar.—Apr. 1962, Parker and Bushman—det. by R. D. Price. ?Islet: 1962, Woodbury. Actornithophilus incisus (Piaget) Aej Islet: Mar.—Apr. 1962, Parker and Bushman—det. by K. C. Emerson. ?lslet: 1962, Parker and Bushman. Actornithophilus ochraceus (Nitzsch) Medren Islet: Apr. 1962, Parker and Bushman—det. by R. D. Price. Austromenopon atrofuluum (Piaget) Aej Islet: Mar.-Apr. 1962, Parker and Bushman—det. by R. D. Price. Enewetak Islet: Feb. 1962, Parker and Bushman; Mar. 1962, Parker and Bushman. Islet: 1962, Parker and Bushman—det. by R. D. Price. Colpocephalum angulaticeps Piaget Ribewon Islet: May 1962, Bushman—det. by R. D. Price. Medren Islet: May 1962, Bush—det. by R. D. Price. Family PHILOPTERIDAE Halipeurus mirabilis Thompson dinimi Islet: May 1962, Bushman—det. by T. Clay. Pectinopygus gracilicornis (Piaget) Ribewon Islet: May 1962, Bushman—det. by K. C. Emerson. Medren Islet: May 1962, Bushman—det. by K. C. Emerson. Quadraceps birostris (Giebel). Aej Islet: Mar. 1962, Parker and Bushman—det. by K. C. Emerson. Enewetak Islet: Feb.—Mar. 1962, Parker and Bushman—det. by R. E. Elbel. Quadraceps hopkinsi Timmermann Aej Islet: Mar.—Apr. 1962, Parker and Bushman—det. by R. E. Elbel. Quadraceps separatus (Kellogg and Kuwana) Aej Islet: Mar.—-Apr. 1962, Parker and Bushman—det. by R. D. Price. Quadraceps sp. [Not counted in totals.] ?Islet: 1962, Woodbury. Saemundssonia albemarlensis (Kellogg and Kuwana) Aej Islet: Mar. 1962, Parker and Bushman—det. byKenG: Emerson. Saemundssonia puellula Timmermann dinimi Islet: May 1962, Bushman—det by R. E. Elbel. Saemundssonia remota Timmermann Aej Islet: Mar.—Apr. 1962, Parker and Bushman—det. by R. E. Elbel. Saemundssonia snyderi (Kellogg and Paine) Enewetak Islet: Mar. 1962, Parker and Bushman—det. by R. E. Elbel. Trabeculus hexakon (Waterston) dinimi Islet: May 1962, Bushman—det. by T. Clay. Order THYSANOPTERA Family PHLAEOTHRIPIDAE Haplothrips gowdeyi (Franklin) Enewetak Islet: Sept. 1975, Cheng—det. by F. A. Bianchi. Order HETEROPTERA Family ANTHOCORIDAE Physopleurella mundula (White) Japtan Islet: Aug. 1956, Tuthill. Family COREIDAE Liorhyssus hyalinus (Fabricius) Elugelab Islet: Jan. 1951, Oshiro. Family CYDNIDAE Geotomus pygmaeus (Dallas) ?Islet: Aug. 1956, Tuthill. Family GERRIDAE Halobates micans (Eschscholtz) Enewetak Islet: Sept. 1975, Cheng. Hermatobates sp. ?Islet: 1977, Cheng. Family LYGAEIDAE Germalus sp. Enewetak Islet: Sept. 1975, Cheng—det. by W.C. Gagne. Nysius picipes Usinger Aomon Islet: May 1946, Oakley and Townes. Enjebi Islet: May 1946, Oakley and Townes. Japtan Islet: May 1946, Oakley and Townes; date?, Townes. Nysius pulchellus Stal Ananij Islet: Aug. 1956, Tuthill. Enewetak Islet: Sept. 1975, Cheng—det. by W. C. Gagné. Enjebi Islet: Jan. 1951, Oshiro. Japtan Islet: Aug. 1956, Tuthill; Jan. 1957, Tuthill; Sept. 1975, Cheng. Medren Islet: Aug. 1956, Tuthill; Sept. 1975, Cheng. ?Islet: May 1946, Oakley; Dec. 1950, Oshiro. Pachybrachius nigriceps (Dallas) Enewetak Islet: Sept. 1975, Cheng—det. by W. C. Gagné. Japtan Islet: Aug. 1956, Tuthill; Sept. 1956, Tuthill; Jan. 1957, Tuthill; Sept. 1975, Cheng—det. by W. C. Gagné. Family MIRIDAE Campylomma eniwetok Schuh Japtan Islet: Feb. 1957, Tuthill. Campylomma marshallensis Usinger Enewetak Islet: Sept. 1975, Cheng. Japtan Islet: Sept. 1975, Cheng—det. by W.C. Gagné. Trigonotylus dohertyi (Distant) ?Islet: May 1946, Townes. Family NABIDAE Nabis capsiformis Germar 7 Elugelab Islet: Jan. 1951, Oshiro. Japtan Islet: May 1946, Townes; Aug. 1956, Tuthill; Jan.Feb. 1957, Tuthill; Sept. 1975, Cheng—det. by W. C. Gagné. Family PENTATOMIDAE Oechalia schellenbergii (Guerin-Meéneville) Enjebi Islet: Dec. 1950, Oshiro. ?Islet: Aug. 1945, Allen; May 1946, Townes. Platynopus melacanthus (Boisduval) Bokombako Islet: Dec. 1950, Oshiro. Order HOMOPTERA Family APHIDIDAE Aphis gossypii Glover Enjebi Islet: May 1946, Oakley. Japtan Islet: May 1946, 172 SAMUELSON AND NISHIDA Townes. ?Islet: Jan. 1945, R. Bohart; May 1946, Oakley. Aphis craccivora Koch Enjebi Islet: May 1951, Fosberg. Japtan Islet: May 1946, Oakley. Family CICADELLIDAE Balclutha incisa (Matsumura) ?Islet: Dec. 1950, Oshiro. Balclutha saltuella (Kirschbaum) Japtan Islet: Aug. 1958, Tuthill. Exitianus fusconervosus (Motschulsky) Aomon Islet: May 1946, Townes. Elugelab Islet: Jan. 1951, Oshiro. Ikuren Islet: May 1946, Townes. Exitianus plebeius (Kirkaldy) Ananij Islet: Aug. 1956, Tuthill. Aomon Islet: May 1946, Townes. Elugelab Islet: Jan. 1951, Oshiro. Enewetak Islet: Sept. 1975, Cheng. Ikuren Islet; May 1946, Townes; Aug. 1956, Tuthill. Japtan Islet: Sept. 1975, Cheng. Medren Islet: Aug. 1956, Tuthill; Sept. 1975, Cheng—det. by W. C. Gagne. Orosius argentatus (Evans) Aomon Islet: May 1946, Townes. Enewetak Islet: May 1946, Townes. Japtan Islet: May 1946, Oakley; Aug. 1958, Tuthill. Recilia affinis (Osborn) Ikuren Islet: Aug. 1956, Tuthill. Recilia hopponis (Matsumura) Japtan Islet: Aug. 1958, Tuthill. Superfamily COCCOIDEA Ceroplastes cirripediformis Comstock Medren Islet: Sept. 1957, Tuthill. Coccus hesperidum Linnaeus Enjebi Islet: May 1946, Oakley. Japtan Islet: May 1946, Townes. Icerya purchasi Maskell ?“Nan” Islet: July 1957, Tuthill. Lepidosaphes esakii Takahashi daptan Islet: May 1946, Townes. Phenacoccus solani Ferris Enjebi Islet: May 1946, Townes. Pseudococcus microadonidum Beardsley Enjebi Islet: May 1946, Townes. Saissetia coffeae (Walker) Medren Islet: Sept. 1957, Tuthill. Family DELPHACIDAE Sogatella kolophon (Kirkaldy) Japtan Islet: Aug. 1956, Tuthill. Family DERBIDAE Lamenia caliginea charon Fennah Ikuren Islet: Aug. 1956, Tuthill. Japtan Islet: May 1946, Townes [published as Mar.]; Aug. 1956, Tuthill. ?Islet: Nov. 1944, Edgar. Order COLEOPTERA Family ANTHICIDAE Anthicus vexator Werner Enewetak Islet: Nov. 1944, Dybas. Japtan Islet: Nov. 1944, Hagen; Sept. 1975, Cheng—det. by G. A. Samuelson. Family CARABIDAE Egadroma smaragdula (Fabricius) Enewetak Islet: Sept. 1975, Cheng—det. by G. A. Samuelson. Enjebi Islet: Jan. 1951, Oshiro. Family CERAMBYCIDAE Prosoplus atlanticus atlanticus Breuning Elugelab Islet: Jan. 1951, Oshiro. Family CHRYSOMELIDAE Aphthona bicolorata Jacoby. ?Islet: Sept. 1975, Cheng—det. by G. A. Samuelson. Family COCCINELLIDAE Coelophora inaequalis Enewetak Islet: Sept. 1975, Cheng—det. by G. A. Samuelson. Japtan Islet: August 1956, Tuthill. Medren Islet: Aug. 1956, Tuthill. Harmonia arcuata (Fabricius) Enewetak Islet: Aug. 1945, Allen. Japtan Islet: May 1946, Townes. Nephus roepkei (Fluiter) Enewetak Islet: Sept. 1975, Cheng—det. by G. A. Samuelson. Enjebi Islet: May 1946, Townes. Medren Islet: Aug. 1956, Tuthill. Scymnus nigrosuturalis Kamiya Japtan Islet: Sept. 1975, Cheng—det. by G. A. Samuelson. Family DERMESTIDAE Dermestes ater De Geer Japtan Islet: May 1946, Townes. Family ELATERIDAE Conoderus pallipes Eschscholtz Enewetak Islet: May 1946, Oakley and Townes. Enjebi Islet: Dec. 1950, Oshiro. Japtan Islet: Sept. 1975, Cheng—det. by G. A. Samuelson. ?Islet: May 1946, Oakley and Townes. Simodactylus fasciolatus Enjebi Islet: May, 1946, Oakley. Japtan Islet: May 1946, Townes; Sept. 1975, Cheng—det. by G. A. Samuelson. Runit Islet: Aug. 1945, Allen. Family MYCETOPHAGIDAE Typhaea stercorea Linnaeus Enewetak Islet: Nov. 1944, Edgar. Family NITIDULIDAE Carpophilus davidsoni Dobson Japtan Islet: May 1946, Fosberg; Sept. 1956, Tuthill. Carpophilus hemipterus (Linnaeus) Enewetak Islet: Nov. 1944, Dybas. Carpophilus marginellus Motschulsky Japtan Islet: May 1946, Fosberg. Carpophilus mutilatus Erichson Japtan Islet: Nov. 1944, Dybas. Carpophilus pilosellus Motschulsky Enewetak Islet: Nov. 1944, Dybas and Edgar. Urophorus humeralis (Fabricius) Elugelab Islet: Jan. 1951, Oshiro. Enewetak Islet: Nov. 1944, Dybas. Japtan Islet: Nov. 1944, Dybas. May 1946, Oakley; May 1946, Fosberg; Sept. 1956, Tuthill. Family OEDEMERIDAE Eobia kanack (Fairmaire) Japtan Islet: Sept. 1975, Cheng—det. by G. A. Samuelson. Eobia matsumurai Kono Japtan Islet: Sept. 1975, Cheng—det. by G. A. Samuelson. Eobia trukana Kono Japtan Islet: Aug.—Sept. 1956, Jan.—Feb. 1957, Tuthill. Enewetak Islet: May 1946, Townes. Eobia new species Japtan Islet: August 1956, Jan. 1957, Tuthill. ?Islet: Sept. .1975, Cheng—det. by G. A. Samuelson. Pselaphanca apicata (Fairmaire) Japtan Islet: Aug.—Sept. 1956, Jan. 1957, Tuthill. Medren Islet: Sept. 1975, Cheng—det. by G. A. Samuelson. New genus and new species: Japtan Islet: Aug.—Sept. 1956, Jan.—Feb. 1957, Tuthill. Family PROPALTICIDAE Propalticus insularis John Japtan Islet: Nov. 1944, Dybas. INSECTS AND ALLIES (ARTHROPODA) 173 Family SCOLYTIDAE Cryphalomorphus nubilus Wood Ikuren Islet: May 1946, Townes. Hypothenemus eruditus Westwood daptan Islet: Nov. 1944, Dybas. Family TENEBRIONIDAE Gebieniella carinata (Eschscholtz) ?Islet: Nov. 1944, Edgar. Gonocephalum adpressiforme Kaszab Enewetak Islet: Sept. 1975, Cheng—det. by G. A. Samuelson. Order NEUROPTERA Family CHRYSOPIDAE Chrysopa ramburi Schneider Aomon_ Islet: May 1946, Townes. Bokombako Islet: Dec. 1950, Oshiro. Japtan Islet: May 1946, Townes. Order LEPIDOPTERA Family ARCTIIDAE Utetheisa pulchelloides Hampson “Occurs on every island visited”: May 1946, Townes. Family HESPERIIDAE Badamia exclamationis (Fabricius) ?Islet: May 1946, Townes. Family NOCTUIDAE Spodoptera exempta (Walker) ?Islet: May 1946, Townes. Spodoptera litura (Fabricius) “Occurs on every island visited”: May 1946, Townes. Family NYMPHALIDAE Hypolimnas bolina (Linnaeus) “On every island visited”: May 1946, Townes. Precis villida (Fabricius) Aomon Islet: May 1946, Townes. Enjebi Islet: May 1946, Townes. Medren Islet: May 1945, Young. Family SPHINGIDAE Agrius convolvuli (Linnaeus) _ Enjebi Islet: Jan. 1951, Oshiro. Japtan Islet: Aug.—Sept. 1956, Tuthill. Medren Islet: Aug. 1956, Tuthill. ?Islet: Aug. 1967, Knudsen. Cephonodes armatus Rothschild and Jordan Islet: May 1946, Townes. Cephonodes picus (Cramer) Enewetak Islet: May 1946, Townes and Oakley. Enjebi Islet: May 1946, Townes; May 1946, Townes and Oakley; Jan. 1951, Oshiro. Gnathothlibus erotus (Cramer): ?Islet: May 1946, Townes. Family TORTRICIDAE Adoxophyes fasciculana (Walker) Elugelab Islet: Jan. 1951, Oshiro. Japtan Islet: Aug. 1956, Tuthill; Jan. 1957, Tuthill. Order DIPTERA Family AGROMYZIDAE Ophiomyia cornuta (de Meijere) Enewetak Islet: Sept. 1975, Cheng. Japtan Islet: May 1954, Townes [published as 1956]; Aug. 1956, Tuthill?; Jan-Feb. 1957, Tuthill; Sept. 1975, Cheng—det. by J. A. Tenorio. Medren Islet: Aug. 1955, Wheeler. Pseudonapomyza spicata (Malloch) ?Islet: May 1946, Townes. Family CALLIPHORIDAE Chrysomya megacephala (Fabricius) Bokombako Islet: Dec. 1950, Oshiro. Enjebi Islet: Dec. 1950, Oshiro. Phaenicia cuprina (Wiedemann) 7 Enewetak Islet: May 1946, Townes; Jan. 1951, Oshiro. Enjebi Islet: May 1946, Oakley. Family CANACEIDAE Nocticanace marshallensis Wirth Aomon Islet: May 1946, Townes. Enewetak Islet: Sept. 1975, Cheng. Japtan Islet: May 1946, Townes; Sept. 1975, Cheng—det. by W. W. Wirth. Family CERATOPOGONIDAE Dasyhelea esakii Tokunaga Japtan Islet: Aug—Sept. 1956, Tuthill; Jan.—Feb. 1957, Tuthill; Sept. 1975, Cheng—det. by W. W. Wirth. Dasyhelea flavescens Tokunaga and Murachi Enewetak ?Islet: Aug. 1956, Tuthill. Dasyhelea flavibasalis Tokunaga Japtan Islet: Aug. 1956, Tuthill. Dasyhelea peliliouensis Tokunaga Japtan Islet: Aug. 1956, Tuthill. Forcipomyia (Forcipomyia) tuthilli Tokunaga Japtan Islet: Aug. 1956, Tuthill. Family CHIRONOMIDAE Clunio tuthilli Tokunaga Enewetak Islet: Sept. 1975, Cheng. Japtan Islet: Sept. 1956, Tuthill; Sept. 1975, Cheng—det. by W. W. Wirth. Medren Islet: Sept. 1956, Tuthill. Pontomyia natans Edwards 11 30N 162 15E (lagoon): date?, Cheng? Tanytarsus halophilae Edwards Japtan Islet: Sept. 1975, Cheng—det. by W. W. Wirth. Telmatogeton pusillum Edwards daptan Islet: Sept. 1975, Cheng—det. by W. W. Wirth. Thalassomya maritima Wirth Enewetak Islet: Sept. 1975, Cheng. Enjebi Islet: Dec. 1950, Oshiro. Ikuren Islet: Sept. 1975, Cheng. Japtan Islet: Aug. 1956, Tuthill; Sept. 1975, Cheng—det. by W. W. Wirth. Family CHLOROPIDAE Cadrema pallida (Loew) % Enewetak ?Islet: Jan. 1951, Oshiro. Enjebi Islet: Aug.—Sept., Wheeler. Ikuren Islet: Aug.—Sept., Wheeler. Japtan Islet: May 1946, Townes; Aug.—Sept. 1956, Tuthill; Jan-Feb. 1957, Tuthill; Aug.—Sept., Wheeler—det. by C. Sabrosky. “Gaurax” bicoloripes (Malloch) Japtan Islet: Aug.—Sept. 1955, Wheeler; Aug. 1956, Tuthill; Jan. 1957, Tuthill—det. by C. Sabrosky (generic position uncertain, not Oscinosoma). Family EPHYDRIDAE Allotrichoma alium Cresson Japtan Islet: Aug.—Sept. 1955, Wheeler—det. by W. W. Wirth. Discocerina mera Cresson daptan Islet: Aug. 1956, Tuthill—det. by W. W. Wirth. Hecamede persimilis Hendel Japtan Islet: Aug.—Sept. 1955, Wheeler. Medren Islet: Aug.—Sept. 1955, Wheeler. ?Islet: Dec. 1950, Oshiro—det. by W. W. Wirth. Hostis guamensis Cresson Japtan Islet: Aug. 1956, Tuthill; Sept. 1975, Cheng—det. by W. W. Wirth. Placopsidella cynocephala Kertész Enjebi Islet: Jan. 1951, Oshiro—det. by W. W. Wirth. Ikuren Islet: Sept. 1975, Cheng. Japtan Islet: Sept. 1975, Cheng—det. by J. A. Tenorio. Medren Islet: Aug.—Sept. 1955, Wheeler—det. by W. W. Wirth; Sept. 1975, Cheng—det. by J. A. Tenorio. 174 SAMUELSON AND NISHIDA Family HIPPOBOSCIDAE Olfersia spinifera (Leach) Ribewon Islet: May 1962, Bushman—det. by T. C. Maa. Family MILICHIIDAE Desmometopa varipalpis Malloch Medren Islet: Aug.—Sept. 1955, Wheeler. ?Islet: Dec. 1944, Edgar; May 1946, Townes—det. by C. Sabrosky. Milichiella lacteipennis (Loew) es Enewetak ?lIslet: Dec. 1950, Oshiro. Japtan Islet: Aug.—Sept. 1955, Wheeler—det. by C. Sabrosky. Family MUSCIDAE Atherigona flavipalpis Malloch _ Elugelab Islet: Jan. 1951, Oshiro. Enjebi Islet: Jan. 1951, Oshiro. Japtan Islet: Aug. 1956, Tuthill. Atherigona (Acritochaeta) orientalis Schiner Japtan Islet: Aug. 1956, Tuthill. Musca (Musca) domestica Linnaeus Ananij Islet: Aug. 1956, Tuthill. Ikuren Islet: Aug. 1956, Tuthill. Japtan Islet: Aug. 1956, Tuthill. Medren Islet: Aug. 1956, Tuthill. ?Islet: Dec. 1950, Oshiro. Family PHORIDAE Diplonevra (Dohrniphora) cornuta (Bigot) Japtan Islet: Aug. 1956, Tuthill. Megaselia (Megaselia) scalaris (Loew) Japtan Islet: Aug. 1956, Tuthill. Family SARCOPHAGIDAE Boettcherisca karnyi (Hardy) daptan Islet: Nov. 1944, Edgar. Parasarcophaga (Liosarcophaga) misera (Walker) Enewetak Islet: Sept. 1975, Cheng—det. by J. A. Tenorio. Enjebi Islet: Jan. 1951, Oshiro. Japtan Islet: Aug. 1956, Tuthill. Phytosarcophaga gressitti (Hall and Bohart) ?Islet: Jan. 1951, Oshiro. Family SCATOPSIDAE Holoplagia guamensis (Johannsen) or near ? Ikuren Islet: Sept. 1975, Cheng—det. by J. A. Tenorio. Family SCIARIDAE Bradysia tritici (Coquillett) Japtan Islet: Aug. 1956, Tuthill. Corynoptera latistylata (Hardy) Japtan Islet: Aug. 1956, Tuthill. Plastosciara latipons Hardy Japtan Islet: Aug. 1956, Tuthill; Sept. 1956, Tuthill. Family STRATIOMYIDAE Brachycara ventralis Thomson Japtan Islet: May 1946, Townes. Family SYRPHIDAE Ischiodon scutellaris (Fabricius) Enewetak Islet: Nov. 1944, Dybas; May 1946, Oakley and Townes. Japtan Islet: Jan. 1957, Tuthill. ?Islet: Jan. 1951, Oshiro; date?, Fosberg. Family TIPULIDAE Limonia (Dicranomyia) pectinunguis Tokunaga Ananij Islet: Aug. 1956, Tuthill. Elugelab Islet: Jan. 1951, Oshiro. Japtan Islet: Aug. 1956, Tuthill; Sept. 1975, Cheng—det. by F. G. Howarth. Order HYMENOPTERA Family ANTHOPHORIDAE Xylocopa sonorina Smith Enewetak Islet: Aug. 1944?, Bryan; Feb. 1975?, Bryan— sight records by E. H. Bryan, Jr. Family BRACONIDAE Zele sp.: ?Islet: May 1946, Townes. Family EULOPHIDAE Hemiptarsenus semialbiclavus (Girault) ?Islet: May 1946, Townes. Family EVANIIDAE Szeptigetella sericea (Cameron) Medren Islet: Sept. 1975, Cheng—det. by G. Nakahashi. Family FORMICIDAE Monomorium pharaonis (Linnaeus) Enjebi Islet: May 1946, Townes. Family ICHNEUMONIDAE Echthromorpha agrestoria insidiator (Smith) Bokombako Islet: Dec. 1950, Oshiro. Elugelab Islet: Jan. 1951, Oshiro. Family MEGACHILIDAE Megachile diligens hedleyi Rainbow Enjebi Islet: May 1946, Oakley; May 1946, Oakley and Townes. Japtan Islet: May 1946, Townes. Megachile (Eutricharaea) fullawayi Cockerell Enewetak Islet: Sept. 1975, Cheng—determined by G. Nak- ahashi. Enjebi Islet: May 1946, Oakley. Family SPHECIDAE Chalybion bengalense (Dahlbom) Enewetak Islet: Sept. 1975, Cheng—det. by G. Nakahashi. Pison punctifrons Shuckard Japtan Islet: May 1946, Townes. Solierella peckhami (Ashmead) Enjebi Islet: May 1946, Townes and Oakley. Tachysphex bengalensis Cameron Japtan Islet: Sept. 1975, Cheng—det. by G. Nakahashi. Family VESPIDAE Odynerus sp. Enewetak Islet: Sept. 1975, Cheng—det. by G. Nakahashi. Pachodynerus nasidens (Latreille) ?Islet: May 1946, Townes. Polistes fuscatus aurifer (Saussure) Japtan Islet: Sept. 1975, Cheng—det. by G. Nakahashi. Ropalidia marginata (Lepeletier) Enewetak Islet: Sept. 1975, Cheng. Japtan Islet: Sept. 1975, Cheng—det. by G. Nakahashi. ACKNOWLEDGMENTS We are indebted to the late Dennis M. Devaney, Bishop Museum, Honolulu, for encouraging our participa- tion in this volurne. The late Edwin H. Bryan, Jr., Bishop Museum, and Leonard D. Tuthill (retired), University of Hawaii, Honolulu, related first-hand information about the environment and human activities on Enewetak Atoll. Robert E. Elbel, University of Utah, Salt Lake City, fur- nished records from earlier investigations by John B. Bush- man and others of the University on vertebrates and ecto- parasites from the atoll. Lanna Cheng, Scripps Institution of Oceanography, La Jolla, California; Robert E. Elbel; and Curtis W. Sabrosky and Willis W. Wirth, National Museum of Natural History, Washington, D. C., provided informa- tion on problematical species or determined relatively recently collected specimens from the atoll. The following also assisted us in providing identifications or related infor- mation: Syoziro Asahina, Tokyo; Peter F. Bellinger, Cali- INSECTS AND ALLIES (ARTHROPODA) 175 fornia State University, Northridge; F. A. Bianchi, Bishop Museum; K. C. Emerson, Sanibel, Florida; Neal Evenhuis, Bishop Museum; Wayne C. Gagné, Bishop Museum; Ash- ley B. Gurney, National Museum of Natural History; Francis G. Howarth, Bishop Museum; C. H. C. Lyal, Brit- ish Museum (Natural History), London; Grace Nakahashi, Bishop Museum; Lawrence J. Pinter, Naval Facilities Engineering Command, San Francisco; Roger D. Price, University of Minnesota, St. Paul; J. C. E. Riotte, Bishop Museum; George A. Schultz, Hampton, New Jersey; Joaquin A. Tenorio, Plant Industry and Extension Service, Saipan; and Ian W. B. Thornton, LaTrobe University, Bun- doora. Ann Marsteller, Hawaiian Sugar Planters’ Associa- tion library, Aiea, located needed references. JoAnn M. Tenorio, Bishop Museum, read the manuscript and pro- vided useful criticism. Frank J. Radovsky, Bishop Museum, helped to initiate our participation in this volume. 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Johnson, 1963, A Report on the Team Effort at Eniwetok Atoll, Pacific Ocean, University of Utah for U. S. Army Chemical Corps, Dugway Proving Ground (typed report). Carpenter, F. M., 1961, Neuroptera: Hemerobiidae, Insects Micronesia, 8: 35-43. Cartwright, O. L., and R. D. Gordon, 1971, Coleoptera: Scara- baeidae, Insects Micronesia, 17: 257-296. Carvalho, J. C. M., 1956, Heteroptera: Miridae, Insects Micronesia, 7: 1-100. Chapin, E. A., 1957, Scorpionida, Insects Micronesia, 3: 65-70. __, 1965, Coleoptera: Coccinellidae, Insects Micronesia, 16: 189-269. Cheng, L., 1977, The Elusive Sea Bug Hermatobates, Pan-Pac. Entomol., 53: 87-97. __, and J. D. Collins, 1980, Observations on Behavior, Emer- gence and Reproduction of the Marine Midges Pontomyia (Diptera: Chironomidae), Mar. Biol., 58: 1-5. , and H. Hashimoto, 1978, The Marine Midge Pontomyia (Chironomidae) with a Description of Females of P. oceana Tokunaga, Syst. Entomol., 3: 189-196. 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J., 1956, Hemiptera: Tingidae, Insects Micronesia, 7: 101-116. ___, 1961, Hemiptera: Saldidae, Insects Micronesia, 7: 287-305. Enderlein, G., 1924, Beitrage zur Kenntnis der Platystominen, Mitt. Zool. Mus. Berl., 11: 97-153. Endrédy-Younga, S., 1971, Coleoptera: Cybocephalidae, Insects Micronesia, 16: 281-285. Esaki, T., E. H. Bryan, Jr., and J. L. Gressitt, 1955, Bibliogra- phy, Insects Micronesia, 2: 1-68. __, and S. Miyamoto, 1975, Homoptera: Cicadidae, Insects Micronesia, 6: 633-656. Essig, E. O., 1956, Homoptera: Aphididae, Insects Micronesia, 6: 15-37. Fennah, R. G., 1956, Homoptera: Fulgoroidea, Insects Micronesia, 6: 39-211. __, 1971, Homoptera: Fulgoroidea, Supplement, Insects Micronesia, 6: 563-699. Ferris, G. F., 1959, Anoplura, Insects Micronesia, 8: 9-12. Ford, E. J., Jr., 1958, Coleoptera: Anobiidae, Insects Micronesia, 16: 75-83. . Fosberg, F. R., 1959, Additional Records of Phanerogams from the Northern Marshall Islands, Atoll Res. Bull., 68: 1-9. 176 SAMUELSON AND NISHIDA Fukushima, I., 1947, Noctuidae of Micronesia, Mushi, 18: 1-22. Goodnight, C. J., and M. L. Goodnight, 1957, Opiliones, Insects Micronesia, 3: 71-83. Ghauri, M. S. K., 1966, Revision of the Genus Orosius Distant (Homoptera: Cicadelloidea), Bull. Br. Mus. (Nat. Hist.) Ento- mol., 18: 231-252. Gillogly, L. R., 1962, Coleoptera: Nitidulidae, Insects Micronesia, 16: 133-188. Gressitt, J. L., 1954, Introduction, Insects Micronesia, 1: 1-257. , 1955, Coleoptera: Chrysomelidae, Insects Micronesia, 17: 1-60. , 1956, Coleoptera: Cerambycidae, Insects Micronesia, 17: 61-183. Gross, G. F., and J. C. Schaffner, 1963, Heteroptera: Coreidae (Alydini by J. C. Shaffner), Neididae, and Nabidae, Insects Micronesia, 7: 357-390. Hardy, D. E., 1956, Diptera: Bibionidae and Scatopsidae, Insects Micronesia, 12: 87-102. ——, 1956, Diptera: Dorilaidae (Pipunculidae), Insects Micronesia, 13: 1-9. _—, 1957, Diptera: Coelopidae, Insects Micronesia, 14: 41-46. __, 1958, Omphralidae (Scenopinidae), Insects Micronesia, 13: 11-13. —, and M. Adachi, 1956, Diptera: Tephritidae, Insects Micronesia, 14: 1-28. Hathaway, W. H., 1953, The Land Vegetation of Arno Atoll, Marshall Islands, Atoll Res. Bull., 16: 1-68. Herring, J. L., 1967, WHeteroptera: Anthocoridae, Insects Micronesia, 7: 391-414. Hopkins, G. H. E., 1961, Siphonaptera, Insects Micronesia, 14: 91-107. Jackson, W. B., 1968, Dragonfly Populations at Eniwetok Atoll, Bioscience, 18: 1123. James, M. T., 1962, Diptera: Insects Micronesia, 13: 75-127. John, H., 1960, Eine Ubersicht iiber die Familie Propalticidae (Coleoptera), Pac. Insects, 2: 149-170. —_, 1971, Coleoptera: Propalticidae, Insects Micronesia, 16: 287-294. Kato, M., 1960, Homoptera: Membracidae, Insects Micronesia, 6: 345-351. Kempny, P., 1904, Beitrag zur Neuropternfauna der Marshall- Inseln, nebst Beschreibung zweier neuer Chrysopa-Arten, Verh. Zool.-Bot. Ges. Wien., 54: 352-355. Kohls, G. M., 1957, Ixodoidea, Insects Micronesia, 3: 85-104. Krantz, G. W., 1967, Acarina: Mesostigmata. Macrochelidae, Insects Micronesia, 3: 149-154. Krombein, K. V., 1949, The Aculeate Hymenoptera of Micronesia I. Scoliidae, Mutillidae, Pompilidae and Sphecidae, Proc. Hawaii. Entomol. Soc., 13: 367-410. __, 1950, The Aculeate Hymenoptera of Micronesia II. Colleti- dae, Halictidae, Megachilidae, and Apidae, Proc. Hawaii. Entomol. Soc., 14: 101-142. Kulzer, H., 1957, Coleoptera: Tenebrionidae, Insects Micronesia, 17: 185-256. Kurosawa, M., 1940, Thysanoptera of Micronesia, Tenthredo, 3: 45-57. Lieftinck, M. A., 1962, Odonata, Insects Micronesia, 5: 1-95. Stratiomyidae; Calliphoridae, Linnavuori, R., 1960, Homoptera: Cicadellidae, Insects Micronesia, 6: 231-344. —, 1975, Homoptera: Cicadellidae, Supplement, Insects Micronesia, 6: 611-632. Lobl, I., 1981, Coleoptera: Scaphidiidae, Insects Micronesia, 15: 69-80. Maa, T. C., 1966, Diptera: Hippoboscidae; Streblidae, Insects Micronesia, 14: 251-274. Macnamara, J. P., and R. H. Arnett, Jr., Coleoptera: Oedemeri- dae, Insects Micronesia (in preparation). Maguire, B., Jr., 1967, Small Freshwater Organisms on Eniwetok Atoll, Micronesica, 3: 151-157. Mari Mutt, J. A., 1979, A Review of the Genus Uchidanura with Description of a New Species (Collembola: Neanuridae), Pac. Insects, 20: 53-58. Matsuda, R., and R. L. Usinger, 1957, Heteroptera: Aradidae, Insects Micronesia, 7: 117-172. Nakata, S., 1961, Some Notes on the Occurrence of Phasmato- dea in Oceania, Pac. Insects Monogr., 2: 107-121. Ohira, H., 1971, Coleoptera: Elateridae, Supplement, Insects Micronesia, 16: 295-300. Quate, L. 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A., 1963, Diptera: Agromyzidae, Insects Micronesia, 14: 135-162. Steffan W. A., 1969, Diptera: Sciaridae, Insects Micronesia, 12: 669-732. Supplement, Insects INSECTS AND ALLIES (ARTHROPODA) 177 Steyskal, G., and M. Sasakawa, 1966, Diptera: Clusiidae, Insects Micronesia, 14: 243-249. Stone, A., 1960, Diptera: Tabanidae, Insects Micronesia, 13: BESS: ___, 1964, Diptera: Simuliidae, Insects Micronesia, 12: 629-635. Strohecker, H. F., 1958, Coleoptera: Endomychidae, Insects Micronesia, 16: 105-108. Sugerman, B. B., 1972a, Insects and Other Arthropods from Kwajalein Atoll (Marshall Islands), Proc. Hawaii. Entomol. Soc., 21: 271-286. ___, 1972b, Insects of Lib Island (Marshall Islands), Proc. Hawaii. Entomol. Soc., 21: 287-289. ___, 1979, Additions to the List of Insects and Other Arthropods from Kwajalein Atoll (Marshall Islands), Proc. Hawaii. Entomol. Soc., 23: 147-151. Synave, H., 1957, Homoptera: Cercopidae, Insects Micronesia, 6: 213-230. Takahashi, R. 1956, Micronesia, 6: 1-13. 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Watanabe, C., 1945, Notes on Some Micronesian Braconidae, Mushi, 16: 47-58. 1958, Hymenoptera: Eucharidae, Insects Micronesia, 19: 19-34. Werner, F. G., 1965, Coleoptera: Anthicidae, Insects Micronesia, 16: 255-269. Wheeler, M. R., and H. Takada, 1964, Diptera: Drosophilidae, Insects Micronesia, 14: 163-242. Williams, F. X., 1931, Handbook of the Insects and Other Inver- tebrates of Hawaiian Sugar Cane Fields, Honolulu: Hawaiian Sugar Planters’ Association, 400 pp. Wilson, E. O., and R. W. Taylor, 1967, The Ants of Polynesia, Pac. Insects Monogr., 14: 1-109. Wilson, N., 1967, Acarina: Mesostigmata. Dermanyssidae, Laela- pidae, Spinturnicidae Parasitic on Vertebrates, Insects Micronesia, 3: 133-148. 1972, Anoplura, Supplement, Insects Micronesia, 8: 145-148. Wirth, W. W., 1951, A Revision of the Dipterous Family Cana- ceidae, Occas. Pap. Bernice P. Bishop Mus., 20: 245-275. Wittmer, W., 1958, Coleoptera: Lampyridae, Cantharidae, Malachiidae, and Prionoceridae, Insects Micronesia, 16: 67-74. ——_, 1970, Coleoptera: Malachiidae, Supplement, Insects Micronesia, 16: 279-280. Wood, S. L., 1960, Coleoptera: Platypodidae and Scolytidae, Insects Micronesia, 18: 1-73. Woodbury, A. M., 1962, A Review of the Ecology of Eniwetok Atoll, Pacific Ocean, University of Utah, Institute of Environ- mental Biological Research, pp. 1-123. Wygodzinsky, P., and R. L. Usinger, 1960, Heteroptera: Reduvi- idae, Insects Micronesia, 7: 231-283. Yoshimoto, C. M., 1962, Hymenoptera: Eucoilinae (Cynipoidea), Insects Micronesia, 19: 89-107. ——, and T. Ishii, 1965, Hymenoptera: Chalcidoidea: Eulophidae, Encyrtidae (part), Pteromalidae, Insects Micronesia, 19: 109-178. Zimmerman, E. C., 1942, Curculionidae of Guam, Bull. Bernice P. Bishop Mus., 172: 73-146. ___, 1948, Notes on Marianas Islands Curculionidae (Coleoptera), Proc. Hawaii Entomol. Soc., 13: 305-315. __, 1964, Platysimus in Southeastern Polynesia (Coleoptera: Curculionidae: Otiorhynchinae: Celeuthetini), Pac. Insects, 6: 747-752. | Chapter 16 Pycnogonida of Enewetak Atoll C. ALLAN CHILD Department of Invertebrate Zoology (Crustacea) National Museum of Natural History Smithsonian Institution, Washington, D. C. 20560 INTRODUCTION The Pycnogonida (sea spider) fauna of the western and mid-Pacific island chains is almost unknown. Although generally benthic organisms, a few are considered plank- tonic and are associated with hosts in parasitic or commen- sal relationships. Most are known to feed on a wide variety of soft-bodied marine life with coelenterates reported as food more often in the literature than any other group. Pycnogonids should therefore be, and prob- ably are, quite at home on the coral reefs of atolls and vol- canic islands of these Pacific chains. The fact that there is only one paper (Child, 1982) on pycnogonids from the Marshall Islands, and particularly Enewetak Atoll, attests not only to the scarcity of pycnogonids but also to a lack of collections containing pycnogonids from the small and usually remote Pacific islands. Pycnogonids are collected fortuitously during the course of benthic sampling for other marine organisms. They are almost impossible to see in their regular habitats because of their microscopic size and their cryptic colora- tion, which renders them almost invisible. Pycnogonids have been found in the laboratory during the microscopic examination of benthic samples, sometimes long after the collection was made, and it is usually impossible to return to the same collection locality for more samples. That large numbers are found in benthic samples demonstrates the careful collecting and sorting procedure during the field efforts. The pycnogonids of the Marshall Islands and Enewetak Atoll predictably will not be as diverse or varied as the fauna of large islands or continental shores. The diversity of benthic habitats on coral atolls is reduced and almost totally excludes the muddy estuaries and weed-encrusted rocky shores of continental masses and large islands. Many groups of organisms are faunistically impoverished on coral atolls. Barnard (1965), in discussing predominantly benthic amphipods, states that “Micronesia primarily offers an epi- 179 faunal environment in the shallow sea, with scarce rem- nants of muddy coastal shelves fringing larger islands and continents. Hence the lack of [diverse] shallow sea bot- toms, the diminution of environmental variability and the decrease of food from runoff should be limiting factors.” This impoverishment would extend to those groups of pyc- nogonids preying on algal-attached sessile organisms. The lack of large expanses of foliose algae and sea grass beds on atolls, including Enewetak, limits the availability of many kinds of pycnogonid foods that use these plants as substrates. The embryology of most pycnogonids, at least for those groups about which we know anything of their life habits, is not conducive to wide island arc dispersal. The eggs and young of many pycnogonids are carried by the male on their ovigers—appendages specifically modified for this purpose—and have no known planktonic dispersal. Their metamorphosis is directly from larva to adult, but in some recorded instances, the eggs are deposited in a host organism, such as a bivalve mollusk, where the young lead a parasitoid existence until they reach a free-living adult stage. Dispersal from island group to island group is prob- lematical, and many pycnogonid species are thought to be endemics, some in quite restricted areas. Many of these endemics may be due to a lack of collecting in adjacent areas. I believe though, that when more is known about pycnogonid distribution, many of these species will remain recorded as endemics. Only five pycnogonid species in four genera are known from Enewetak Atoll (Table 1). A sixth, Nymphon micronesicum (family Nymphonidae), is also known from Bikini (Child, 1982). This is a very small faunule when con- sidering the great diversity of this class of arthropods in the oceans of the world. There are approximately 900 recognized species of pycnogonids in 76 genera and nine families (a very small faunule when the great diversity of this class of arthropods is considered), and we probably know about one-half or less than half of the world’s species. The littoral shores of the world have not been sampled to any great extent, and with a few exceptions, it is in this environment that the greatest diversity, if not the greatest numbers, of pycnogonids live. Further collecting in littoral and sublittoral habitats on Enewetak Atoll and other 180 TABLE 1 Checklist of Pycnogonida of Enewetak Atoll Phylum ARTHROPODA Class PYCNOGONIDA Order PANTOPODA Family AMMOTHEIDAE * Ammothella stauromata Child, 1982. Family CALLIPALLENIDAE * Callipallene sp. cf. novaezelandiae (Thomson, 1884). Family ENDEIDAE * Endeis nodosa Hilton, 1942. Family PHOXICHILIDIIDAE * Anoplodactylus glandulifer Stock, 1954. * Anoplodactylus marshallensis Child, 1982. *New Enewetak and Micronesian record. islands and atolls of Micronesia will undoubtedly add many species to the faunal checklist shown in Table 1. Collec- tions of coral rubble from lagoon and outer reef habitats will probably yield additional new distributional records and new species. Scraping for fouling organisms on pier pilings, navigation buoys, and ship’s hulls is another method for sampling common pycnogonid habitats and should increase our knowledge of that fauna. Pycnogonid habitats have hardly been touched on most of the oceanic islands of the Pacific. COLLECTION DATA Ammothella stauromata, Anoplodactylus glandulifer, and Anoplodactylus marshallensis were all collected by the author in 1969 from pier pilings on the north end of Enewetak Island (site name Fred). Callipallene sp. cf. novaezelandiae and Endeis nodosa were collected by the author in 1969 and by others in 1974 and 1975 from lagoon rubble and coral rocks in fairly shallow water. The Callipallene specimens were found in rubble on a lagoon coral knoll near Jinedrol Island (site name Alvin) and in coral rubble from Enewetak Island. Endeis nodosa came from coral rocks in the lagoon off Enewetak Island and from pier pilings on the lagoon side of Runit Island (site name Yvonne). No specimens have been reported from the rich outer reef habitats of Enewetak Atoll. Ammothella stauromata and Anoplodactylus marshall- ensis are recently described species (Child, 1982) and are known only from Enewetak Atoll. Anoplodactylus glandulifer Stock, 1954, has the widest known distribution of the five species. It is known in widely scattered localities from the Red Sea and the east coast of Africa to Burma and Singapore and for the first time in the central Pacific islands at Enewetak. Endeis nodosa Hilton, 1942, has been known only from Hawaii, on Oahu Island, and the Enewetak specimens extend its known range considerably to the west. The Callipallene specimens show slight differ- ences from Thomson’s (1884) species novaezelandiae, but if they are in fact this species, the previously known south- ern hemisphere distribution of C. novaezelandiae (east coast of Africa to New Zealand) would be extended to the CHILD northern hemisphere at Enewetak. The record depths for all five species are shallow to littoral. In general, the four genera have a worldwide distribu- tion, predominantly in the tropics, but there are species of Endeis, Anoplodactylus, and Callipallene in deep sea habi- tats and even in cold Antarctic waters. Anoplodactylus is the largest genus represented in terms of species known with about 75 now recognized. It has several known representatives in most papers on the perimeter Pacific island groups, including Australia and New Zealand, and appears in the few reports that exist on western and mid- Pacific islands. There are close relations of the Enewetak species of Anoplodactylus in Japan and the Indian Ocean. The specific food preferences of most pycnogonid species are unknown; so there can be no comparisons of habitat-food preference among the various species. The genus Ammothella, with 28 known species, is tropical-temperate in distribution and has island represen- tatives in scattered localities around some parts of the Pacific perimeter. Callipallene, with 28 known species, is one of many genera in a large family which seems to have an area of proliferation, at the generic level, in the Austral-Indonesian area (referring to the island arc from Australia and New Zealand through the Philippines). Of the 24 genera in the family Callipallenidae, 13 genera have representatives in this area. One genus (Parapallene) has the majority of its known species found in this area, and three genera (Pycnopallene, Spasmopallene, and Stylopallene) are endemic. Therefore, it would not be surprising at all to find a number of coral reef species of callipallenids at Enewetak and the Marshall Islands. The genus Endeis, with 17 reported species, has seven from the Austral-Indonesian region. ACKNOWLEDGMENTS The author is grateful to the Atomic Energy Commis- sion which operated the Enewetak Marine Biological Laboratory under the management of the University of Hawaii, to the base support of Kentron Inc., and to the U. S. Air Force for field support while specimens for the Smithsonian Institution were collected. I also wish to thank my fellow divers from the Smithsonian and the U. S. Geological Survey for their support and specimen contributions. REFERENCES Barnard, J. L., 1965, Marine Amphipoda of Atolls in Micronesia, Proc. U. S. Nat. Mus., 177: 459-552. Child, C. A., 1982, Pycnogonida from the Marshall Islands, Proc. Bio. Soc. Wash., 95: 270-281. Hilton, W. A., 1942, Pycnogonids from Hawaii, Occas. Pap. Bernice P. Bishop Mus., 17: 43-55. Stock, J. H., 1954, Pycnogonida from Indo-West Pacific, Aus- tralian, and New Zealand Waters, Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16, Vidensk. Medd. Dansk Naturh. Foren., 116: 1-168. Thomson, G. M., 1884, On the New Zealand Pycnogonida, with Descriptions of New Species, Trans. and Proc. New Zealand Institute, 16: 242-248. Chapter 17 Stomatopod Crustacea of Enewetak Atoll MARJORIE L. REAKA®* and RAYMOND B. MANNING+ “Department of Zoology, University of Maryland, College Park, Maryland 20742; +Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560 INTRODUCTION This study provides a checklist and discusses the biogeographic relationships of the 12 species of Stoma- topoda (mantis shrimps) now known to occur on Enewetak Atoll. Five species are widely distributed throughout the Indo-West Pacific region, and three species have somewhat more restricted ranges in the Indo-West Pacific and Indian Ocean. Four species are endemic to the Central Pacific (two to Enewetak), and three taxa need further taxonomic investigation, which may demonstrate further endemism. Stomatopods from Enewetak are dwarfed in body size compared to their mainlaind relatives. Small size has strong consequences for life history and evolutionary pat- terns in stomatopods, and in particular is likely to generate endemism. We provide information on the color patterns of the stomatopods from Enewetak, showing which traits are the most reliable indicators of species identity for taxo- nomic and field research and which traits are most likely to be influenced by body size, sex, or habitat. Several anomalies in usually invariant color traits are found in stomatopods from Aomon Island, which was closer to sources of radiation from atomic testing than more south- ern islands in the atoll. We also summarize what is known about the habitat and fighting behavior of the coral- dwelling mantis shrimps from Enewetak. For each of the above topics, we identify or discuss all previous literature on the stomatopods of Enewetak. CHECKLIST The stomatopod Crustacea from Enewetak Atoll were first recorded by Manning (1971) and more recently by Manning and Reaka (1982): these studies included the description of three new species. Here we identify four species (Gonodactylus ternatensis, Pseudosquilla ciliata, 181 P. ornata, and Lysiosquilla maculata) that have not been recorded previously from Enewetak, bringing to 12 the total number of species known from the atoll (Table 1). We also identify here all pertinent literature in which par- ticular species of stomatopods from Enewetak have been discussed. BIOGEOGRAPHIC COMPARISONS Pseudosquilla ciliata is the only species of those found on Enewetak that also occurs in the Atlantic. Five of the species known from Enewetak (G. chiragra, G. platysoma, P. ciliata, P. ornata, and L. maculata) are widely distrib- uted throughout the Indo-West Pacific region. All of these are relatively large species of stomatopods, and Pseudosquilla has a particularly long larval dispersal stage (Reaka, 1979a, 1980a; Reaka and Manning, 1987a). Three species have somewhat more restricted distribu- tions in the Western Pacific, Indo-Malayan, and Indian Ocean regions than the previously mentioned group. Haptosquilla glyptocercus occurs from Enewetak to Japan, northern Australia, and as far west as the Andaman Islands. Gonodactylus smithii is known from Oceania and the South China Sea and from Australia to the western Indian Ocean; however, populations from the western Indian Ocean and Red Sea may be referable to another species, G. acutirostris De Man. Gonodactylus ternatensis is known with certainty only from localities between Samoa and Indonesia. Of the remaining species, G. espinosus and G. in- cipiens are known only from island localities in the Central Pacific, while G. insularis and G. micronesica apparently are endemic to Enewetak Atoll. These four species, as well as H. glyptocercus (which also has a relatively restricted distribution), are the smallest species that occur on Enewetak. Figure 1 shows that the typical body sizes of popula- tions of all of the coral-dwelling stomatopods from Enewetak are diminutive compared to those from many other geographic localities, particularly the Indo-West Pacific mainland areas. This fact is emphasized by compar- ing lineages of the same or closely related species in dif- ferent regions. For example, individuals of G. smithii attain considerably smaller sizes in Enewetak than in Thailand or 182 REAKA AND MANNING TABLE 1 Checklist of Stomatopoda from Enewetak Atoll Phylum CRUSTACEA Class MALACOSTRACA Order STOMATOPODA Family GONODACTYLIDAE Gonodactylus chiragra (Fabricius): Manning, 1971. Gonodactylus espinosus Borradaile: Manning, 1971. Gonodactylus incipiens Lanchester: Reaka, 1973; Dingle et al., 1973; Caldwell and Brunenmeister, 1975; Reaka, 1975a; Caldwell and Dingle, 1977; Reaka, 1979a. Gonodactylus viridis Serene: Caldwell and Dingle, 1972. Gonodactylus childi Manning: Manning, 1972. Gonodactylus insularis Manning and Reaka: Manning and Reaka, 1982 Gonodactylus falcatus (Forskal): Manning, 1971; Caldwell and Dingle, 1972; Dingle et al., 1973; Reaka, 1973; Reaka, 1975a; Reaka, 1975b; Reaka, 1976; Reaka, 1979a; Manning and Reaka, 1982. Gonodactylus micronesica Manning: Manning, 1971; Dingle et al., 1973; Reaka, 1973. Gonodactylus platysoma Wood-Mason: Manning, 1971; Caldwell and Dingle, 1972; Reaka, 1973; Caldwell and Brunenmeister, 1975; Reaka, 1975a; Reaka, 1975b; Reaka, 1979a. Gonodactylus smithii Pocock: Manning, 1971; Caldwell and Dingle, 1972; Reaka, 1973; Dingle et al., 1973; Caldwell and Brunenmeister, 1975; Reaka, 1975a; Reaka, 1975b; Reaka, 1979a; Reaka, 1979b. “Gonodactylus ternatensis De Man, 1902 (new record based on specimen in USNM collections). Family PROTOSQUILLIDAE Haptosquilla glyptocercus (Wood-Mason): Manning, 1971; Caldwell and Dingle, 1972; Dingle et al., 1973; Caldwell and Brunenmeister, 1975; Reaka, 1975a; Reaka, 1975b; Reaka, 1976; Caldwell and Dingle, 1977; Reaka, 1979a; Reaka, 1979b. Family PSEUDOSQUILLIDAE *Pseudosquilla ciliata (Fabricius, 1787) (new record based on specimen in USNM collection). “Pseudosquilla ornata Miers, 1880 (new record based on specimen in Rijksmuseum van Natuurlijke Historie, Leiden). Family LYSIOSQUILLIDAE “Lysiosquilla maculata (Fabricius, 1793) (new record based on specimen determined by R. Kinzie for Mid-Pacific Marine Laboratory reference collection). “New records for species at Enewetak. Australia (and we suggest that this may be an endemic population). Individuals of G. incipiens from Enewetak are smaller than their cognates from Thailand (G. viridis) or Australia (G. affinis). Gonodactylus insularis in Enewetak is a dwarf species compared to its close relatives in Thailand (G. mutatus), Australia (G. falcatus), and Hawaii (G. aloha). These data cannot be accounted for by collecting tech- niques, sample sizes, or latitude and temperature. (See Reaka and Manning, 1987a, for further discussion). One lineage, that of H. glyptocercus, does not fit this pattern since it is not smaller in Enewetak than in Thailand. We have discussed elsewhere reproductive and life his- tory traits and their relationship to body size, dispersal ability, and evolutionary rates in stomatopods, including those from Enewetak (Reaka, 1975a, 1976, 1978, 1979a, 1979b, 1980a; Reaka and Manning, 1981, 1987a). Life history traits are scaled to body size within related lineages of stomatopods, and the life history patterns of species from Enewetak conform to these overall patterns. In partic- ular, however, small body size and restricted dispersal abil- ity generate rapid evolutionary changes and are likely to be associated with endemism. The tendency for dwarfism and endemism in the stomatopods from Enewetak provides another example of this trend. COLOR PATTERNS Despite their cryptic habits, stomatopods often are flamboyantly colored (Reaka, 1975b, 1980b, 1981; Reaka and Manning, 1981). Some of these color patterns vary dramatically among individuals of the population, while other traits are invariable in color. In another study on the coral-dwelling stomatopods of Enewetak (Reaka and Man- ning, 1987b), we provide the first quantitative analysis in any crustacean of the degree of color polymorphism in all morphological traits, and we test a series of hypotheses about the function of color patterns in stomatopods. Invari- ant and species-specific color traits can facilitate accurate identification of morphologically similar species in the field and also can be used to resolve taxonomic differences among sibling species (Manning, 1964, 1971; Manning and Reaka, 1979, 1981la, 1981b, 1982). Therefore, we pro- vide here a summary of the traits that will be either the most useful or the most likely to lead one astray in field identifications of the common coral-dwelling stomatopods in Enewetak. The meral spot (an often brightly colored oval indenta- tion on the dorsomedial surface of each of the raptorial second maxillipeds) is one of the most reliable characteris- tics for identifying species (Table 2). The meral spot is STOMATOPOD CRUSTACEA 183 (T) 70 Gap) Gt = 4Pc¢ Pcs CA) = oGs SOC) i = : / (Ge) Ca / ePc ae 5 P oGg = 3c é i; O one GE : Ei NCEE) 5; : =p of ; °Gz 40 \ ’ rs A@s > ’ 8s a a) Oo (aa) Ze = je LW = 10 15 20 ATM UDE (|) MS) 30 Fig. 1 Median body lengths of populations of stomatopods from different localities as a function of latitude. Body lengths are listed in descending order. Lines (discussed in text) connect con- specific or cognate populations that are especially pertinent to this discussion. [From Reaka and Manning, 1987b.] ge Phuket, Thailand (T) Gt is Gonodactylus ternatensis, Pc is Pseudosquilla ciliata, Gc is G. chiragra, Gs is G. smithii, Gv is G. viridis, Gm is G. mutatus, Hg is Haptosquilla glyptocercus O Townsville, Australia (A) Gs is G. smithii, Gg is G. graphurus, Gf is G. falcatus, Ga is G. affinis, Gh is G. hendersoni © Pacific Costa Rica (CR) Gf is G. festae, Gc is G. costaricensis, Gz is G. zacae, Gb is G. bahiahondensis, Gs is G. stanschi A Hawaii (H) Pc is P. ciliata, Ga is G. aloha, Gh is G. hendersoni A Enewetak (E) Gp is G. platysoma, Gs is G. smithii, Hg is H. glyptocercus, Gic is G. incipiens, Gis is G. insularis, Gm is G. micronesica, @ Gulf of California (GC) Gz is G. zacae @ Florida (F) Pc is P. ciliata REAKA AND MANNING 184 [@ Yysuees5 umoiq “8 YsImol[a A] INGEDS (9) Nagao yuid yuid sBe] Buryjem [sjjoyeois1w pue ajqqni pajuawies abie] ul pue sjeyiqey 1edeap ut ystuaaib ‘ajqqni asOo] |[ewWs ul pue syeyiqey moyjeys ul YysimoyaA [2 ystueasb ‘9 ysiueas5 spadyixeu umolq *S YysImola A] uaai6 Nagao moljaA *s ysimoyja A] JaUIO pediyixew +Nqaqu9-3N 14 pug jelsojder HSIN1d LHOME +q34y Jo [Ajoeq [sjjoyeosstu pue ajqqni +SNYJW NO ANITLNO pajuewias ab1e] GNNOYOOVE MOTTISA ul pue ‘ujdap pedijrxew LHOWE HLIM 31dund yyim aseaiuj] = pug jeojdes gaN1d LNAOsadlul +qg4y 4sjods yieq jo snpodoig pedyyixew pug [euojdes dau aay jo sndied +SONIY OLY LNAONOO UOWT]esS 0} ALIHM GNV YOV1d 4+NOISNANI uaeiB aniq padiixew Ad G3GNNOYYNS dy YOIMSALNV SuOT}eLIeA puz [eojdes +wvayo ‘G34 AWINYVO ALIHM ALIHM ‘MOTTSA + yuid ysitym uo 7jods era [synpe [synpe ystuees5 +Naqu9o TV.LSId [anf ur ystuaaiB * anf YVvd10 “anf ysimola A] ‘JUIHM IWSVa YvaO juasqe ysuaai9] ysImoyjad 1ea[p] safeos euuajuy [sjjoyeos91w pue ajqqni Pajuewias a61e| ul abuelo pue pai ‘ajqqni asooy [jews [synpe par UI Iea]d ‘p aBueso 4+3.LIHM (aehs| “anf 1ea[D] Yuval pai ‘1ea]9 JO pal ‘8 1ea[D] aeuuajuy [s}eyiqey 1edaap ul ysimoyjaA [synpe paz ‘syeyiqey moyjeys +NOOUVN aay “anf 1ea[9] aay 1eaj9 ul YsIppayy] sajnuuajuy sn3.1290)d 415 “H HYWWS pwosijpjd *} DO}SIUOAIIULSE) SHDINsUl “} Suatdjou! “D wel 2SaldedG uly s}Tery 1005 @ ATaVL 185 STOMATOPOD CRUSTACEA *saloeds 1ayjo ul s[enpiaipul Aue jo asou} dejiano jou op yey} siojoo anbiuqt WWUNO 0} GQ syeygey 1edeap ‘Ww Gg Q> ale s}e}Iqey moTeYs ‘s[enpiAIpu! [jews 10 saluaan{ ‘an ‘s}leI} a[qelsen JO SIOJOD UOWIWOD }sOW a}edIPU! ($}9}492I1Q OU) S1a}ja| eased Jamo] ‘sajoeds ujyyim Aem (GQ;Q > d) a[qejoipaid Ajjeoisijeys e ul Aen jeu} s}ie1} G1 S}aydeIQ Ul S1d}}2] aSeD amo] ‘s}!1e1} a[qeiseAul AjiuBis siajjaj ased 1add¢, uawopqe pue xeIOY} UO Ula}jed a1ydAjbosa1y y1ep [2 ystuees5 ‘8 ysjumoig] umMmo1Q ‘1ea]9 umoiq ‘1e2[9 umoiq +INISav ONIGNVG [sjjoyeos91w pue ajqqni pejuewes abe ul pue ‘saioeds antssaibbe Yum szeyiqey Ul ‘syeyiqey 1edeep ul ‘s[enplalpul 1a61e| ul saseeiou! Bulj49eds] [2 ystuees5 ‘§ ystumoig] (sy]npe UO SIOJOD 12y4}0 “anf ysimola A] [s}]npe UO SIOJ[OD 12430 “anf ysimola A] dy uaei5 aniq +:LNAWOS TVSVd NO LOdS Gay ANINYVO TIVYSAO ONITLLOW AAVSAH AYSA ‘XVYOHL SSOYOV GNV4d SLIHM umolqg aal[o NEEL (9) NEE?t9) dil N4dYo +:LNAWOdS TvSvVd NO SLOdS 4NTd WuyvVd CNV LHS! HUM JONVYO +SONIY OIYLNAONOSD F.LIHM CNV ‘NMOud day ‘3N1d wuvd Ad GJGNNOYENS HOLVd JONVYO MOTISA “WOddv HL9 NO «LOdS Ad» TWHALVT +SONIY OIU.LNAONOO ALIHM GNV “ANTE A1Vd ‘day “ANd Wuvd Ad GaaNNOWYENS HOLVd AONVHO dae *«LOdS “JAd,, TWYALV1 NAWOddv YOIMSL -SOd GNV XVHOHL HLOd SSOYUOV GNVd SLIHM INGER (9) NEERt9) NEED (9) NEE? t9) 4NOSTIL GNV ‘WOddY HL9 NO AVNIYVO Gay [syeqiqey Jadaap ul saseaidul Surpueq ‘sjen -PIAIpul 1aB1e] ul aseaioul BuljjjOowW pue 5ulj49eds] uawopge pue xe10U} uo sBuryiew o1ydA[bora1y4 yep jelpaw ysiuaes5 {ue} uaeiB moyjad uaaib moyjaA [sa1oads aAIssaib5e yno -4yIM syeyiqey Ul Pai0joo ‘saloads aatsseibbe yum szeyqey ul 1ea[D] +qgu {s}]npe pas “anf 1e2[D] [satoeds antssaibbe ynoyjiM asou} ul pue syeyiqey 1edaap ul ysimo]aA ‘saisads aalssaibbe yim syeyiqey ul pue Moyfeys ul ead ‘s}jnpe uaeaib uaaiB aniq “anf 1e2[D] +NOSTSL YOIMALNV NO @ CNV ‘WOddV HL9 NO @ ‘SLOdS MOV Td LNANINO'd 0 Sulpueq asiansuel} “‘Bulyyour “‘Bulyyoeds 1002 Apog podoxa jepodoiy) podopua [epodoiy, podoin jo diy paxi0j pue juawiBas jeseg sjuawbas uosja} pue Jeurwopge 49 juawbas 21928104} 4g 186 REAKA AND MANNING whitish pink in G. incipiens, yellow with an anterior red infusion in G. insularis, white in G. micronesica and G. platysoma, carmine red surrounded by a_black-and- white ring in G. smithii, and cream colored in H. glypto- cercus. Our analysis, however, shows that the color of the meral spot ranges from blue greenish to salmon among individuals classified as G. incipiens; morphological differ- ences, especially in the structure of the telson, among members of this population also suggest that this taxon needs further investigation. Additionally, individuals in the Enewetak population of G. smithii have a carmine meral spot, whereas members of other populations of G. smithii bear a dark purplish red meral spot. This fact, in combina- tion with the observations that G. smithii from Enewetak are diminutive in body size, produce smaller eggs relative to their body size, and invest a smaller proportion of their body volume in a brood of eggs than do other populations of G. smithii (Reaka, 1975a, 1979a; Reaka and Manning, 1987b), suggest that this population also is in need of tax- onomic investigation and may represent an endemic species. Several other invariant traits also are useful for species identifications, particularly when used in combination with the meral spots. Haptosquilla glyptocercus sports uniquely maroon antennules and white antennae among the species at Enewetak. The antennal scales of G. platysoma are white proximally but green distally. The propodus of the raptorial appendage in G. smithii is iridescent blue purple, set against a bright yellow background outline where it folds against the merus. Lateral dark spots characterize the propodus of the raptorial appendage in G. incipiens (although their number increases with depth and type of habitat). The propodus and dactyl of the raptorial maxil- liped are uniquely red in G. micronesica. The dactyl of the raptorial appendage in G. platysoma is bright bluish blue-green. Gonodactylus platysoma also is remarkable for the colorful concentric “eye spots” on each side of the eighth thoracic and sixth abdominal segments. On the thorax this spot is composed of an outer white band, fol- lowed by a pale blue, red, dark blue, and finally an inner- most red orange circlet of color. On the sixth abdominal segment, the “eye spot” is formed of an outer band of white, followed by red brown and dark blue rings and an innermost patch of yellow orange. The basal segment of each uropod in G. platysoma is orange with a light and dark blue spot. The basal segment of each uropod in G. smithii bears a brilliant red spot which is conspicuously exposed against dark green body coloration when the uro- pods are flared (particularly when the telson is coiled defensively in front of the body during a fight); these spots vanish when the uropods are tucked under the telson in a submissive posture. In addition, G. insularis is character- ized by a lineage-specific set of four black spots—two on the sixth abdominal segment and two on the anterior tel- son. The uropodal exopods (and usually the endopods) are uniquely red in G. insularis, as are the dorsal carinae on the sixth abdominal segment and telson in G. micronesica (Table 2). In contrast to the invariable and often unique traits dis- cussed above, however, the color of most traits varies among individuals or with sex, developmental stage, or habitat in most species. When the range of variability among individuals is considered for all species (Reaka and Manning, 1987b), all of the major morphological traits can be ranked for relative variability of color, providing an indi- cation of which traits are likely to be the most—or the least—reliable indicators of species identity in general. Except for the specific cases identified in Table 2 as being invariable, the uropodal exopods, antennules, antennae, uropodal endopods, and antennal scales, respectively, form a set of traits that are moderately variable in color. Simi- larly, when all species are considered (with invariable exceptions being noted in Table 2), the color of the maxil- lipeds, walking legs, and uropodal forked tips varies still more among individuals; body color is in general the most polymorphic of all traits. Descriptions of species and field identifications based upon the latter characters are most likely to be biased by small sample sizes. Color patterns vary predictably with sex in some cases (Table 2). The color of the body, walking legs, and maxil- lipeds are more likely to be greenish in males but may shift toward other hues (especially brownish, yellowish) in females of some species (G. smithii, G. incipiens, H. glypto- cercus). In G. incipiens, the antennae are more likely to be red or orange in males but clear in females. The color pattern of certain traits also varies predict- ably with size or age in some species. The incidence of blue green or green coloration on the antennal scales or on the endopods, exopods, or forked tips of the uropods increases in larger individuals of several species (G. incipiens, G. insularis, G. smithii). The color of the uropodal endopod shifts toward red (matching the red of the invariant exopod) as individuals of G. insularis become larger. Also, the antennae and antennules of G. platysoma change from clear to red as individuals mature. White speckling on the body increases with size in G. incipiens and G. smithii. Several color characteristics shift with habitat within species, where these shifts cannot be explained by the dis- tributions of individuals of different sexes and sizes in dif- ferent habitats. In G. incipiens, the frequency of white transverse banding on the body, dark spots on the propodus of the raptorial appendage, and yellow coloration on the antennules and distal fork of the uropods increases from shallow (0.5 m) to deeper (0.5 to 2.0 m) habitats. The degree of color polymorphism among individuals also increases with depth (antennal scales, maxillipeds, distal forks of the uropods) in G. incipiens. Additionally, the uropodal endopods in G. incipiens shift toward clear coloration in habitats that contain assemblages of aggres- sive species of stomatopods. Color patterns also correlate with habitat and suscepti- bility to predation among species. Because they are highly vagile or occupy deep, reefward, or open exposed environ- ments, G. incipiens, G. platysoma, and G. micronesica are likely to encounter considerable fish predation. On the STOMATOPOD CRUSTACEA 187 other hand, G. insularis, G. smithii, and H. glyptocercus are less vagile, often occur in more protected and shore- ward habitats, and undoubtedly suffer less predation than G. incipiens, G. platysoma, and G. micronesica. Gonodac- tylus platysoma and G. incipiens are the most heavily speckled and mottled species, and transverse banding is most developed in these two species and G. micronesica. Green is the predominant color in 17%, 47%, and 50%, respectively, of the traits of G. incipiens, G. platysoma, and G. micronesica. Green coloration is found on only 8%, 8%, and 0%, respectively, of the traits in G. insularis, G. smithii, and H. glyptocercus (see Reaka and Manning, 1987b, for details). Thus, we conclude that fish predation is likely to be a selective factor promoting drab greenish coloration and speckling and banding patterns that camou- flage the individual against algae and the grainy calcareous substrate. On the other hand, species with elaborate aggressive displays (see section on behavior) bear the most brightly colored structures; 62% and 58%, respectively, of the traits in G. smithii and G. insularis, but only 33%, 29%, and 20%, respectively, of those in nonagressive species (G. incipiens, G. micronesica, and G. platysoma, respectively) include predominantly red and yellow hues. (See Reaka and Manning, 1987b, for analysis.) Haptosquilla glypto- cercus is one of the most uncolorful species, with only 9% of its traits (the antennules) bearing bright coloration; indi- viduals of this species show extremely aggressive behavior but lack complex displays. Similarly, bright colors are concentrated on anterior structures associated with threatening and attacking behav- ior (see Table 2 and quantified data in Reaka and Man- ning, 1987b). Red, maroon, or orangish antennules are found in G. smithii, G. micronesica, G. platysoma, H. glyp- tocercus, and G. incipiens, respectively. Red antennae are frequent in G. insularis, G. smithii, and G. platysoma. Gonodactylus insularis and G. smithii usually bear red or yellowish antennal scales. Maxillipeds often are yellowish in G. incipiens, G. insularis, and G. smithii. Gonodactylus smithii bears a red meral spot, and the carpus and prop- odus in G. micronesica and G. platysoma are red. Bright colors also characterize posterior structures used in defen- sive combat (red uropodal spots in G. smithii; red or yel- low uropodal endopods and exopods in G. insularis and G. smithii, respectively; red telson carinae in G. micronesica). Color polymorphism of the body is greatest in G. smithii and G. insularis; G. insularis and G. incipiens have the most color polymorphic appendages. Therefore, color polymorphism among individuals in the population is greatest either in species with elaborate aggressive com- munication (e.g., graded displays in G. insularis, G. smithii) or in species that occur in association with these species (G. incipiens). These observations suggest that color polymorphism may function for individual recog- nition of aggressive communication. Several deviant color patterns, particularly in G. platysoma, were found on the northern island of Aomon, which was closer to sources of radiation from atomic testing than many of the southern islands in the atoll. Deviations in the red coloration of the anterior rap- torial appendage, in the blue spots on the orange back- ground of the basal segment of the uropod, and in the sequence of concentric colors on the lateral “eye spots” were observed in the population of G. platysoma collected from Aomon. Comparisons with other populations of this species collected from farther south on the atoll revealed that these deviations were not explained by body size, sex, or habitat. BEHAVIOR Stomatopods are highly visual, active predators that have some of the most complex behavior known in inver- tebrates (Caldwell and Dingle, 1975; Reaka, 1980b; Reaka and Manning, 1981). The gonodactylids and protosquillids on Enewetak (Table 1) inhabit preformed burrows in coral or calcareous substrate that cannot be enlarged; these species show the most intense and complex aggressive behavior. Members of the genus Pseudosquilla also show relatively elaborate behavior and strong aggression and occupy either holes in coral or excavations under coral. Lysiosquillids burrow in sand, can enlarge their burrows, have less elaborate behavior, and can grow to very large sizes (Caldwell and Dingle, 1975; Reaka, 1981; Reaka and Manning, 1981, 1987a). The second maxilliped of stomatopods is enlarged into a raptorial appendage that is used to smash and stab com- petitors for burrows, predators, or hard-bodied prey. Intense, potentially lethal fighting behavior is observed among members of the same and different species in both the field and laboratory. Aggressive behaviors include strikes (a smashing blow with the hammer-like raptorial appendage or a stab with the open, needle-sharp dacty]); a variety of aggressive lunges and chases; and graded threat displays in which the raptorial appendages are lowered and spread, exposing the conspicuous meral spots. Defensive behaviors include coiling the armored telson in front of the body with the uropods spread wide. In coral- dwelling species this stance may effectively block the entrance of a burrow against an intruder. In addition, these mantis shrimps exhibit several retreat behaviors, including submissive postures in which the head and telson are lowered and their associated structures are tucked along- side or under the body. The agonistic behavior of different species of coral- dwelling stomatopods in Enewetak has been addressed in several studies (Caldwell and Dingle, 1972, 1977; Reaka, 1973; Dingle, Highsmith, Evans, and Caldwell, 1973; Caldwell and Brunenmeister, 1975). We provide new data and synthesize all previous research on the aggressive behavior of these mantis shrimps in a comprehensive study elsewhere (Reaka and Manning, 1987b), but we provide a brief summary here. Haptosquilla glyptocercus delivers the most strikes, but this relatively uncolorful species is exceptionally prone to retreat from opponents and is less likely to use threatening displays than G. smithii or G. insularis. Sporting a yellow 188 REAKA AND MANNING meral spot and other bright coloration, G. insularis uses the highest number of threat displays in its interactions, strikes relatively often, and rarely retreats in a contest; consequently, this species can be classified as a very aggressive species. Gonodactylus smithii also is highly aggressive, striking but also retreating quite frequently. Members of this species perform some of the most expan- sive threat displays, thus exposing one of the most dramatic meral spots (carmine surrounded by black and white concentric circles) seen in any stomatopod. Inter- specific tests indicate that H. glyptocercus dominates G. insularis, and that G. insularis wins contests with G. smithii. Gonodactylus incipiens and G. platysoma are characterized by low levels of fighting activity and subordi- nate social relationships to the other species in Enewetak. HABITAT Field work in 1972 allowed us to quantitatively analyze the distribution of shallow water stomatopods from several major types of habitat in Enewetak (Reaka, 1973; Reaka and Manning, 1987b). These habitats include: 1. Dead coral rubble 2. Soft rubble and vermetid rocks 3. The calcareous bases of “microatolls” (colonies of Porites in which the center has died and often was riddled with boring organisms) 4. Open reef benches of consolidated calcareous accre- tion (Table 3) Dingle et al. (1973), Caldwell and Dingle (1975), and Caldwell and Brunenmeister (1975) provide additional information about the occurrence of stomatopods in partic- ular habitats on Enewetak. Coral rubble habitat is found on the reef platform as isolated heads, in small patch reefs, or along the reefward algal ridge on the east side of Enewetak. These habitats usually are <0.5 m deep and contain G. incipiens, G. micronesica, and H. glyptocercus. Stomatopods also are collected from coral rubble in somewhat deeper habi- tats (<1 m deep) in the channel between Enewetak and the sandy islets on the northeast side of Enewetak, between these sandy islets and Japtan, and on the northeast side of Japtan. These rubble habitats contain G. incipiens, G. micronesica, G. platysoma (small), G. smithii (small), and H. glyptocercus. In addition to the above species, a population now classified as G. insularis (Manning and Reaka, 1982) was found in the rubble on the northeast flank of the sandy islets (Dingle et al., 1973; Reaka, 1980a). By the summer of 1972, however, the rubble had been partly buried by sand, and no G. insularis could be found. Our later analysis suggests some morpho- logical differences between the population found in 1971 and G. insularis (Manning and Reaka, 1982), and the pos- sibility remains that the former population represents an additional (possibly now extinct?) species in the closely related G. falcatus lineage. Of the species found in dead coral rubble, G. incipiens is the most common (45% of the 40 individuals collected), followed by H. glyptocercus (25%), small G. smithii (18%), G. micronesica (10%), and small G. platysoma (39%). Stomatopods inhabit soft rubble and vermetid rocks on the shoreward and reefward sides (<1.5 m deep) of the moat on the north side of Aomon. In 1972 G. insularis inhabited only the shoreward side of this moat (<1 m deep). Gonodactylus incipiens and small G. platysoma and G. smithii also are found here; the distribution of small G. smithii extends into the somewhat deeper reefward areas of the moat. Of the stomatopods collected from the Aomon moat (N = 33), G. insularis is the most common (45%), followed by small G. smithii (27%), G. incipiens (15%), and small G. platysoma (12%). Stomatopods also can be collected from the bases of the microatolls at depths of 0.5 to 2.0 m on the reef flat northeast of Enewetak. Gonodactylus smithii is abundant (72% of the 43 individuals quantified) and reaches large body sizes in this habitat. Although their numbers were not thoroughly quantified, our field notes indicate that G. platysoma also attains large sizes and is “abundant.” The smaller species occurring in this habitat, G. incipiens and H. glyptocercus, also reach relatively large sizes but are less common (9% each of the quantified individuals; however, quantification was less systematic in this than in other habitats, and further sampling is needed). Caldwell and Dingle (1975) suggest that large individuals of G. platysoma occupy relatively open caverns under microatolls, while individuals of G. smithii and the smaller species inhabit borings further inside the center of the Porites colony. Despite their abundance, individuals of G. smithii are rarely observed away from their protective coral colony (Reaka, personal observation). Individuals of two species, G. platysoma and G. in- cipiens, are commonly observed moving rapidly about on the open reef adjacent to the microatolls, particularly on low or incoming tides. As in the microatolls where they find refuge, the G. incipiens and G. platysoma observed on the open reef flat are relatively large in size. Our field notes indicate that individuals of G. platysoma are particu- larly abundant in this habitat. The shallow reef benches (<0.5 m deep) on the east side of Enewetak and the east side of Japtan also yield stomatopods. These reef benches are scoured by wave action and frequently are exposed at low tides. Stomatopods (G. incipiens, H. glyptocercus) are collected from tubular holes in the solid reef substrate or as they dart about in the tide pools at low tide. Gonodactylus incipiens was the most common stomatopod in this habitat (7/9 or 78% of the quantified individuals), although further sampling is needed to assess the relative abundance of H. glyptocercus. In 1972 a heavy rainfall during a mid-day low tide caused a massive reef kill in this shallow exposed habitat (Reaka, 1980a; Leviten and Kohn, 1980). Despite their high activity levels and probable high metabolic rates, stomatopods are remarkably tolerant of harsh physical con- ditions (low Os, high temperatures, etc.; Reaka, personal observation). During this reef kill, mortality among the Distribution and Major Types of Habitat of Stomatopods TABLE 3 (E is east; N is north; NE is northeast) G. incipiens G. insularis G. micronesica G. platysoma G. smithii H. glyptocercus = Total Locally Relative abundance of No. females Body size, No. females Body size, No. females Body size, No. females Body size, No. females Body size, No. females _—_ Body size, No. co-occurring co-occurring species Habitats and sites and males* mmt and males* mmt and males” mmt and males" mmf and males” mmt and males* mmt individuals species in major habitats (1—4) rubble : Pk reefs and isolated heads 22,16 [3] _—-15 to 19 (18) 19 [1] 17 (17) [4] G. incipiens on a reef platform E side of H. glyptocercus Enewetak (<0.5 m deep), often near shore Along algal ridge of reef crest, E side 39, 26 [5] 11 to 17 (16) Present’ [5+] G. incipiens 45% G. incipiens of Enewetak (<0.5 m deep) G. micronesica 25% H. glyptocercus In channel (~1 m deep) and on 49, 16 [5] 17 to 24 (22) 22 [3] 16 to 20 (18) 19, “relatively 16, “small” 29, 16 [5] 19 to 20 (19) [144+] G. incipiens 18% G. smithii (small) outer reef flat between Enewetak common” “small” G. micronesica 10% G. micronesica and sandy islets (“Sand Island”) G. platysoma (small) 3+% G. platysoma (small) NE side of Enewetak G. smithii (small) On NE flank of Sand Island 29 [2] 28 to 34 (31) Present in 19717 19 [1] 22 (22) 29 [2] “small” [5+] G. incipiens_ (between Enewetak and Japtan) G. micronesica (<1 m deep) G. smithii (small) G. insularis** On N side of Japtan (<1 m deep) 12, 26 [3] 15 to 18 (17) 52, 46 [9] 15 to 27 (24) [12] G. incipiens H. glyptocercus 2. Soft rubble and vermetid rocks Shoreward side of moat on N 32 [5] 29 to 32 (30) 122, 36 [15] 8 to 33 (20) 19, 36 [4] 15 to 41 (28) [7] 20, “small” [31] G. incipiens 45% G. insularis side of Aomon (<1 m deep) G. insularis 27% G. smithii (small) G. platysoma (small) 15% G. incipiens G. smithii (small) 12% G_ platysoma (small) Reefward side of moat on N side Present! 19 [2] 13, “small” [2+] G. smithii (small) of Aomon (<1.5 m deep) G. incipiens 3. Bases of microatolls Reef flat (0.5 to 2 m deep) 19, 36 [4] 23 to 36 (33.5) 19, 36 50 to 60 (57) 139, 186[31] 18to62(31) 19,16[4] 20 to 34 (30) [43+] G. incipiens (large) 72% G. smithii (mod. large) NE side of Enewetak “abundant” G. platysoma (large) 9% G. platysoma (large)t+ G. smithii (mod. large) 9% G. incipiens (large) 4. Open reef flat H. glyptocercus 9% H. glyptocercus Reef flat (0.5 to 2 m deep) 32, 56 [9] 21 to 42 (33) “abundant” “large” [9+] G. incipiens (large) (Both common) near microatolls, large G. platysoma (large) coral and other rubble, often considerable algae, NE side of Enewetak Holes in solid reef platform, 42, 36 [7] 18 to 31 (23) 22 [2] 24 to 26 (25) [9] G. incipiens 78% G. incipiens often scoured, with little H. glyptocercus 22% H. glyptocercus or no algae or in open tidepools in the same area (<0.5 m deep), E side of Enewetak and E side of Japtan Total No. Individuals [43+] [15+] [4+] (9+) [47] (16) (134+) “Number in brackets is total number collected. Totals sometimes exceed the sum of the sexes because sex was not recorded for some individuals. +Number in parentheses is median body length. tAt low tide. §12 with eggs, 24 mm. ‘1 Field notes indicate that the species was observed, although individuals were not collected. “Had been present in 1971, but not present in 1972. ++G. platysoma undoubtedly is far more abundant than indicated by this datum. 190 REAKA AND MANNING stomatopods was low compared to that in other crusta- ceans and gastropods (Reaka and Manning, 1987b). Our field studies show that considerable microhabitat overlap, particularly for individuals of similar body sizes, occurs among the species of coral-dwelling stomatopods of Enewetak (Table 3). Gonodactylus incipiens is the most ubiquitous species, and H. glyptocercus, G. smithii, G. platysoma, G. micronesica, and G. insularis, respec- tively, occupy increasingly restricted habitats (number of sites and habitat type). Of the 10 sites sampled comprehensively in 1972, the most common co-occurring species assemblage includes G. incipiens, G. smithii, G. platysoma. Although not very common, G. micronesica occurs in this assemblage as well. When G. insularis is present, it also is a member of the G. incipiens, G. smithii, G. platysoma assemblage. Although H. glyptocercus frequents a wide range of habitat types, this species co- occurs most often with G. incipiens alone, thus forming a second major species association. ACKNOWLEDGMENTS Field work was supported by grants from the Pauley Foundation and the Enewetak Marine Biological Labora- tory to M. L. Reaka. We thank L. B. Holthuis for identify- ing and D. M. Devaney for informing us of the new record of Pseudosquilla ornata. We are grateful to A. J. Kohn for helpful discussion, and M. L. Reaka thanks E. O. Wilson for providing space and facilities at the Museum of Com- parative Zoology, Harvard University, during manuscript preparation. 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Rept. 1972-1973, pp. 51-54. , 1975a, The Ecology and Evolution of Molting, Life History Patterns, and Agonistic Behavior in Stomatopod Crustacea, Ph.D. thesis, Department of Zoology, University of California, Berkeley. __, 1975b, Molting in Stomatopod Crustacea: Stages of the Molt Cycle, Setagenesis, and Morphology, J. Morphol., 146: 55-80. ——, 1976, Lunar and Tidal Periodicity of Molting and Reproduc- tion in Stomatopod Crustacea; a Selfish Herd Hypothesis, Biol. Bull., 150: 468-490. , 1978, The Effects of an Ectoparasitic Gastropod, Caledoniella montrouzieri, upon Molting and Reproduction of a Stomatopod Crustacean, Gonodactylus viridis, Veliger, 21: 251-254. ——, 1979a, The Evolutionary Ecology of Life History Patterns in Stomatopod Crustacea, S. E. Stancyk (Ed.), Reproductive Ecology of Marine Invertebrates, University of South Carolina Press, Columbia. __, 1979b, Patterns of Molting Frequencies in Coral-Dwelling Stomatopod Crustacea, Biol. Bull., 156: 328-342. __, 1980a, Geographic Range, Life History Patterns, and Body Size in a Guild of Coral-Dwelling Mantis Shrimps, Evolution, 34: 1010-1030. ——, 1980b, On Learning and Living in Holes by Mantis Shrimp, An. Behav., 28: 111-115. __, 1981, The Hole Shrimp Story, Nat. Hist., 90: 36-43. —_, and R. B. Manning, 1981, The Behavior of Stomatopod Crustacea, and Its Relationship to Rates of Evolution, J. Crust. Biol., 1: 309-327. ——_, and R. B. Manning, 1987a, The Significance of Body Size, Larval Dispersal, and Habitat for Rates of Morphological Evo- lution in Stomatopod Crustacea, Smith. Contr. Zool., 448: 1-46. —_, and R. B. Manning, 1987b, The Distributional and Behavioral Ecology, and the Ecology of Color Patterns in the Stomatopod Crustacea from Enewetak Atoll, Smith. Contr. Mar. Sci., (in press). Cirripedia of Enewetak Atoll RICHARD H. TITGEN Bernice P. Bishop Museum Honolulu, Hawaii 96817 INTRODUCTION Barnacle research at Enewetak Atoll, Marshall Islands, has been incidental to other marine studies. There have been just two published reports. Zullo et al. (1972) reported specimens of Balanus amphitrite ampbhitrite Darwin, 1854, and Balanus eburneus Gould, 1841, in the collections of the California Academy of Sciences. Tomlin- son (1973) reported the acrothoracican barnacles Lithoglyptes bicornis Aurivillius, 1892 (as Lithoglyptes spi- natus Tomlinson and Newman, 1960), Lithoglyptes mitis Tomlinson, 1969, and Cryptophialus heterodontus Tomlin- son, 1969, from shells in the U. S. National Museum of Natural History’s Division of Malacology. Both of the Balanus spp. are cosmopolitan in warm and temperate seas; they are commonly transported as fouling organisms and were introduced to Enewetak (W. A. Newman, per- sonal communication). The acrothoracican barnacles reported by Tomlinson (1973) also have relatively wide distributional ranges, but these were established by natural means. However, because acrothoracicans lack plankto- Chapter 18 trophic larvae, their long-range dispersal mechanisms are not understood. Nine species of barnacles are known from Enewetak Atoll, four of which are new records supplied by W. A. Newman. In comparing barnacle records from Enewetak Atoll and the other Marshall Islands (Henry, 1957; Tomlin- son, 1969, 1973; Zullo et al., 1972; Newman and Tomlin- son, 1974; and Grygier, 1981a) to species reported from the Caroline Islands (Hiro, 1937, 1938; Boschma, 1953, 1955; Henry, 1957; Newman, 1960a, 1972; Tomlinson, 1969, 1973; Ross and Newman, 1973; and Newman and Ross, 1976), there is a strong indication that more barna- cles should occur at Enewetak Atoll than have been reported. The proximity of the Caroline Islands to the Marshall Islands suggests that many of the Caroline Island species, or closely related ones, may occur in the Marshall Islands and at Enewetak Atoll. However, it will take a directed effort to collect and identify the barnacle species, rather than the incidental collecting that has occurred pre- viously, because tropical barnacles tend to be cryptic and sparsely distributed as compared to temperate forms (New- man, 1960b). Because so few of the species that may occur at Enewetak Atoll have been reported, the included checklist (Table 1) also lists species that probably occur at Enewetak Atoll. TABLE 1 Checklist of Shallow-Water Cirripedia Found, or Likely to Be Found, in the Waters Surrounding Enewetak Atoll* Subphylum CRUSTACEA Class MAXILLOPODA Subclass CIRRIPEDIA Order ASCOTHORACICA Suborder SYNAGOGOIDIDA Family SYNAGOGIDAE + Gorgonolaureus bikiniensis Utinomi, 1962. Order THORACICA Suborder LEPADOMORPHA Family HETERALEPADIDAE +Heteralepas hataii Hiro, 1937. +Heteralepas spp. +Paralepas palinuri (Barnard, 1924). “Systematic hierarchy taken from Bowman and Abele (1982). +Species likely to be found at Enewetak Atoll. 191 (This table continued on next page.) 192 TITGEN TABLE 1 (cont'd) Family IBLIDAE T Ibla cumingi sibogae Hoek, 1907. Family LEPADIDAE + Conchoderma auritum (Linnaeus, 1767). t Conchoderma virgatum (Spengler, 1790). +Lepas anatifera Linnaeus, 1758. TLepas anatifera striata de Graaf, 1952. t Lepas anserifera anserifera Linnaeus, 1767. t Lepas hilli (Leach, 1818). Family MICROLEPADIDAE Tt Microlepas spp. Family POECILASMATIDAE t Octolasmis lowei (Darwin, 1851). t Trilasmis fissum (Darwin, 1851). Family SCALPELLIDAE t Lithotrya nicobarica Reinhardt, 1850. Suborder BALANOMORPHA Superfamily CHTHAMALOIDEA Family CHTHAMALIDAE Subfamily EURAPHIINAE t Euraphia hembeli Conrad, 1837. + Euraphia intertexta (Darwin, 1854). Superfamily CORONULOIDEA Family CORONULIDAE Subfamily CHELONIBIINAE t Chelonibia caretta (Spengler, 1790). t Chelonibia testudinaria (Linnaeus, 1758). Subfamily CORONULINAE t Coronula diadema (Linnaeus, 1767). Subfamily PLATYLEPADINAE Tt Platylepas decorata Darwin, 1854. + Platylepas hexastylos (Fabricius, 1798). Family TETRACLITIDAE Subfamily TETRACLITELLINAE + Tetraclitella divisa (Nilsson-Cantell, 1921). Subfamily TETRACLITINAE + Tesseropora pacifica (Pilsbry, 1928). Superfamily BALANOIDEA Family ARCHAEOBALANIDAE Subfamily ARCHAEOBALANINAE + Acasta spp. Family BALANIDAE Balanus amphitrite amphitrite Darwin, 1854: Zullo et al., 1972. Balanus eburneus Gould, 1841: Zullo et al., 1972. + Balanus trigonus Darwin, 1854. ; +Megabalanus tintinnabulum (Linnaeus, 1758). Family PYRGOMATIDAE Subfamily PYRGOMATINAE + Cantellius spp. + Hiroa stubbingsi Ross and Newman, 1973. t+ Nobia spp. + Savignium crenatum (Sowerby, 1823). + Savignium dentatum (Darwin, 1854). + Savignium elongatum (Hiro, 1931). + Savignium milleporum (Darwin, 1854). Order ACROTHORACICA Suborder PYGOPHORA Family CRYPTOPHIALIDAE +Cryptophialus cordylacis Tomlinson, 1969. Crytophialus heterodontus Tomlinson, 1969; Tomlinson, 1973. +Cryptophialus rossi Tomlinson, 1973. +Species likely to be found at Enewetak Atoll. +New Enewetak Atoll records. . (This table continued on next page.) CIRRIPEDIA 193 TABLE 1 (cont’d) Family LITHOGLYPTIDAE + Balanodytes taiwanus Utinomi, 1950. + Kochlorine ulula Tomlinson, 1973. t Lithoglyptes bicornis Aurivillius, 1892. Lithoglyptes spinatus Tomlinson and Newman, 1960; Tomlinson, 1973. Lithoglyptes mitis Tomlinson, 1969: Tomlinson, 1973. + Lithoglyptes wilson! Tomlinson, 1969. + Weltneria reticulata Tomlinson, 1969. Order RHIZOCEPHALA Suborder KENTROGONIDA Family SACCULINIDAE + Sacculina actaeae Guerin-Ganivet, 1911. t Sacculina bipunctata Kossmann, 1872. + Sacculina carpiliae Guerin-Ganivet, 1911. + Sacculina inconstans Boschma, 1952. +Sacculina punctata Boschma, 1934. +Sacculinid sp. “Systematic hierarchy taken from Bowman and Abele (1982). +Species likely to be found at Enewetak Atoll. +New Enewetak Atoll records. ASCOTHORACICA Ascothoracicans generally have been considered the most primitive barnacles, but there is argument to separate them as a subclass coordinate with the Cirripedia (Grygier, 1983). The bases for the argument are that the Ascothora- cica have nauplii that lack “frontolateral horns; a relatively anamorphic rather than highly metamorphic development; an ascothoracid larva capable of feeding at all stages rather than a single cyprid stage lacking a complete gut and functional mouthparts; wholly prehensile first antennae rather than ones also provided with cement glands; and natatory thoracic limbs that show no indication of ever having been used as cirri. Furthermore, it has recently been shown that the ascothoracican sperm, the most gen- eralized known for the Crustacea, is distinct from that of the remainder of the cirrlpeds” (Newman, 1982). Though the evidence is strong for the separation of the Ascothora- cica from the Cirripedia, no consensus exists. It is included herein for this reason and because only one species has been reported from the Marshall Islands (Utinomi, 1962). Ascothoracicans are ecto- and endoparasites of various anthozoans and echinoderms. Often highly modified for a Parasitic existence, they show little resemblance to the “typical” thoracic barnacle. They lack calcareous plates, do not cement themselves to the substratum, and retain more characteristics of the free-living ancestors of the cirripeds than any other group of crustaceans. Gorgonolaureus bikiniensis Utinomi, 1962, is the single ascothoracican reported from the Marshall Islands. Described as a new genus and species, it is known only from Bikini Atoll and is reported to be a parasite on the gorgonacean Paracis squamata. This shallow-water gor- gonian has not been reported from Enewetak Atoll (Lang and Devaney, this volume). The only other described congeneric species is Gorgonolaureus muzikae Grygier, 1981, reported from deep water in Hawaii (Grygier, 1981b). THORACICA Thoracicans include the typical barnacles usually recog- nized by their form and the presence of calcareous plates. There are both free-living and symbiotic taxa, and they can be divided morphologically into pedunculate and sessile forms. The pedunculate barnacles (Lepadomorpha) are composed of a peduncle that attaches to the substratum and a capitulum that is usually surrounded by calcareous plates. The sessile barnacles (Verrucomorpha and Balanomorpha) lack the peduncle, and the aperture is guarded by an operculum formed by one or two pairs of calcareous plates. Many thoracic barnacles have wide distributional ranges. This is due in part to the majority having plank- tonic larval stages (Newman and Tomlinson, 1974) and some, such as several oceanic lepadomorph species, natur- ally attaching to floating debris. In addition, some of the natural distributions have been greatly extended because certain species tend to “foul” ship bottoms and floating objects and can be carried great distances. Six of the nine barnacle species reported from Enewetak Atoll are thoracicans. This high percentage is not unusual because thoracicans are generally larger than species in the other barnacle orders and have a more obvi- ous “barnacle” appearance, thus making them more recog- nizable and liable to be collected. The other three species, belonging to the Acrothoracica, are small, burrow in cal- cium carbonate, and do not have the typical barnacle 194 TITGEN appearance. They were found because Tomlinson (1973) intentionally looked for their burrows in mollusk shells. ACROTHORACICA Acrothoracicans are small, nonparasitic naked barna- cles that burrow primarily in calcareous substrata. These substrata include live coral, coral rock, and mollusk shells. Sexes are separate, and the burrows are created by the females. The dwarf males are short-lived, non-feeding organisms (Tomlinson, 1969) that are usually found near the attachment disc of the female. Newman and Tomlinson (1974) presented an interest- ing question on the dispersal mechanisms of acrothoracican barnacles. The adults are poor dispersers because they are obligate inhabitants of calcium carbonate. Therefore, it is difficult to explain the distribution of species such as Lithoglyptes bicornis Aurivillius, 1892, which is found in the Caribbean Sea, the Red Sea, Australia, Japan, the Marshall Islands, and the Line Islands. The difficulty arises because the naupliar dispersal stages are usually passed in the egg, and the cyprids are non-feeding and weak swim- mers (Newman and Tomlinson, 1974). A single paper has been published reporting on acrothoracicans from Enewetak Atoll (Tomlinson, 1973). Not collected as barnacle material, the specimens were discovered in shells of the turbinid snail Turbo argyro- stomus Linnaeus, 1758, housed at the U. S. National Museum of Natural History’s Division of Malacology. The three species of acrothoracican barnacles reported from Enewetak Atoll have relatively wide distributions con- sidering their presumed limited powers of dispersal. Cryptophialus heterodontus Tomlinson, 1969, has a western Pacific distribution and is known from Australia, the Marshall Islands (Tomlinson, 1969), and Okinawa (Tomlinson, 1973). Lithoglyptes mitis Tomlinson, 1969, appears to have an insular distribution, having been col- lected at Fiji (Tomlinson, 1969), the Loyalty Islands, the Marshall Islands, New Caledonia, Ninafou Island (2Niuafou), and Samoa (Tomlinson, 1973). As mentioned, Lithoglyptes bicornis Aurivillius, 1892, has an unusually wide distribution for an acrothoracican barnacle. Over part of its range it is known by its synonyms Lithoglyptes spi- natus Tomlinson and Newman, 1960, and Lithoglyptes ampulla Aurivillius, 1892 (Newman and Tomlinson, 1974). Only a few species of acrothoracicans have been reported from the Marshall, Caroline, and Gilbert Islands (Tomlinson, 1969, 1973) that have not also been found at Enewetak Atoll. Due to the cryptic habitat of burrowing barnacles and the fact that they are seldom looked for, it is probable that these and/or other acrothoracicans occur at Enewetak Atoll. RHIZOCEPHALA Rhizocephalans are endoparasites of crustaceans, pri- marily decapods. The adults. are highly modified and bear no resemblance to other cirripeds. However, they are clas- sified with barnacles because their naupliar larvae have the characteristic frontolateral horns, and there is a typical cyprid larval stage. There are no published accounts of rhizocephalans from Enewetak Atoll or the Marshall Islands. The only record is of a sacculinid found on a male specimen of Galathea affinis Ortmann, 1892, collected at Sta. No. JWK-412, Engebi Island, Enewetak Atoll, found on an Acropora sp. (Haig, personal communication). Apparently, no other rhizocephalans were found on the decapods col- lected at Enewetak Atoll (J. S. Garth and A. J. Bruce, per- sonal communication). However, Sacculina punctata Boschma, 1934, and Sacculina bipunctata Kossmann, 1872, have been reported from the Caroline Islands. Sacculina carpiliae Guerin-Ganivet, 1911, Sacculina incon- stans Boschma, 1952, and possibly Sacculina actaeae Guerin-Ganivet, 1911, have been reported from the Gilbert Islands (Boschma, 1953, 1955); and an _ undescribed species of Lernaeodiscus has been reported from Hawaii (Edmondson, 1946; Boschma, 1953). Although present records indicate that infection by rhizocephalans at Enewetak Atoll is of low incidence, some undoubtedly occurs. In general, little work has been done on rhizo- cephalans in recent years, probably because of the histo- logical work necessary for taxonomic identification. DISCUSSION The most noticeable aspect of barnacle research at Enewetak Atoll, and the Marshall Islands in general, is the paucity of work that has been done. The more obvious noncommensal forms have been collected and identified, along with a few of the more cryptic species. However, in comparison to the Caroline Islands, the smaller commensal, burrowing, and parasitic forms have been largely missed. This is characteristic of studies in which barnacle research is incidental to other work being carried out. Because the cirriped fauna of Enewetak Atoll is so poorly known, little can be said of its biogeography. Its fauna, however, is Indo-West Pacific in origin, with some elements of the circumtropical-temperate fauna. Enewetak Atoll is composed of low islands and is located on the Pacific Plate. For these reasons the fauna can be expected to be less diverse than at high islands, such as the Caro- lines, or islands such as Palau and the Philippines that are not on the Pacific Plate (Springer, 1982; W. A. Newman, personal communication). While some patchiness is to be expected, the low islands of the Marshalls would be expected to have few, if any, of Springer’s (1982) Type 2 or 3 endemics. However, some care must be taken in interpreting barnacle endemicity and distributional ranges. Several of the taxa are so poorly known that distributional records often indicate where a certain amount of work has been done, not where the natural limits to distribution occur. Many of the species reported from the Caroline Islands could occur at Enewetak Atoll. Most have reasonably wide western Pacific distributional ranges, and the proximity of CIRRIPEDIA 195 the island groups makes this feasible. In addition, Springer (1982) suggests that the Caroline Islands may act as a con- duit onto the Pacific Plate from the western Pacific. The species most likely to occur at Enewetak Atoll are the bar- nacle commensals of wide-ranging nectonic organisms such as marine mammals and reptiles. These commensals include Conchoderma auritum (Linnaeus, 1767), Concho- derma virgatum (Spengler, 1790), Lepas hilli (Leach, 1818), Chelonibia caretta (Spengler, 1790), Coronula diadema (Linnaeus, 1767), and Platylepas hexastylos (Fabricius, 1798). Another group, which is undoubtedly better represented at Enewetak Atoll than records indicate, includes the coral-inhabiting species in the balanid subfam- ily Pyrgomatinae. Our knowledge of barnacles from Enewetak Atoll can be summarized as incomplete taxonomically. However, zoogeographically Enewetak Atoll is representative of the Indo-West Pacific faunal region with a close relationship to the western Pacific. A small but obvious component of the fauna is the cosmopolitan fouling community, which includes many of the familiar thoracic barnacles that are often so abundant in shallow water. The majority of species, however, display a more restricted distribution. ACKNOWLEDGMENTS I would like to thank William E. Newman for critically reading the manuscript and for his many valuable com- ments and suggestions. He was especially helpful in sug- gesting species that are likely to occur at Enewetak Atoll. | would also like to thank John S. Garth, A. J. Bruce, and Janet Haig for providing information on the possible pres- ence or absence of rhizocephalans on Enewetak decapods. REFERENCES Boschma, H., 1953, The Rhizocephala of the Pacific, Zool. Meded. (Leiden), 32: 185-201. —_, 1955, The Described Species of the Family Sacculinidae, Zool. Verh. (Leiden), 27: 1-76. Bowman, T. E., and L. G. Abele, 1982, Classification of the Recent Crustacea, The Biology of Crustacea, Vol. 1, Systemat- ics, the Fossil Record, and Biogeography, L. B. Abele (Ed.), Academic Press, New York, pp. 1-27. Edmondson, C. H., 1946, Reef and Shore Fauna of Hawaii, Bernice P. Bishop Mus. Spec. Publ. 22, 381 pp. Grygier, M. J., 198la, Redescription of Gorgonolaureus bikiniensis (Crustacea: Ascothoracica), with a Reevaluation of Its Familial Affinities, Micronesica 17: 67-76. —, 1981b, Gorgonolaureus muzikae sp. nov. (Crustacea: Ascothoracida) Parasitic on a Hawaiian Gorgonian, with Spe- cial Reference to Its Protandric Hermaphroditism, J. Nat. Hist., 15: 1019-1045. —_, 1983, Ascothoracida and the Unity of the Maxillopoda, Crustacean Issues 1, Crustacean Phylogeny, Frederick R. Schram (Ed.), A. A. Balkema, Rotterdam, pp. 73-104. Henry, D. P., 1957, Some Littoral Barnacles from the Tuamotu, Marshall, and Caroline Islands, Proc. U. S. Natl. Mus., 107(3381): 25-38. Hiro, F., 1937, Cirripeds of the Palao Islands, Palao Tropical Bio logical Station Studies, 1: 37-72. __.,, 1938, Studies on the Animals Inhabiting Reef Corals, II, Cir- ripeds of the Genera Creusia and Pyrgoma, Palao Tropical Biological Station Studies, 3: 391-416. Newman, W. A., 1960a, Five Pedunculete Cirripeds from the Western Pacific, Including Two New Forms, Crustaceana, 1: 100-116. ——, 1960b, On the Paucity of Intertidal Barnacles in the Tropical Pacific, Veliger, 2: 89-94. __, 1972, Lepadids from the Caroline Islands (Cirripedia Thora- cica), Crustaceana, 22: 31-38. ——, 1982, Evolution Within the Crustacea, Part 3: Cirripedia, The Biology of Crustacea, Vol. 1, Systematics, the Fossil Record, and Biogeography, L. B. Abele (Ed.), Academic Press, New York, pp. 197-221. __., and A. Ross, 1976, Revision of the Balanomorph Barnacles; Including a Catalog of the Species, San Diego Soc. Nat. Hist. Mem., 9: 1-108. __, and J. T. Tomlinson, 1974, Ontogenetic Dimorphism in Lithoglyptes (Cirripedia, Acrothoracica), Crustaceana, 27: 204-208. Ross, A., 1961, A New Cirriped from the Hawaiian Islands, Crustaceana, 2: 209-212. Ross, A., and W. A. Newman, 1973, Revision of the Coral- Inhabiting Barnacles (Cirripedia: Balanidae), Trans. San Diego Soc. Nat. Hist., 17: 137-173. Springer, V. G., 1982, Pacific Plate Biogeography, with Special Reference to Shorefishes, Smithson. Contrib. Zool., 367: 1-182. Tomlinson, J. T., 1969, The Burrowing Barnacles (Cirripedia: Order Acrothoracica), U. S. Natl. Mus. Bull., 296: 1-162. ——, 1973, Distribution and Structure of Some Burrowing Barna- cles, with Four New Species (Cirripedia: Acrothoracica), Wasmann J. Biol., 31: 263-288. Utinomi, H., 1962, Gorgonolaureus, a New Genus of Ascothora- cid Barnacle Endoparasitic in Octocorallia, Proc. U. S. Natl. Mus., 113: 457-464. Zullo, V. A., 1979, Marine Flora and Fauna of the Northeastern United States, Arthropoda: Cirripedia,) NOAA _ Technical Report NMFS Circular 425. —_, D. B. Beach, and J.T. Carlton, 1972, New Barnacle Records (Cirripedia, Thoracica), Proc. Calif. Acad. Sci., 39: 65-74. Chapter 19 Nonplanktonic Copepoda of Enewetak Atoll Compiled by DENNIS M. DEVANEY (deceased) Bernice P. Bishop Museum Honolulu, Hawaii 96817 The copepod fauna of Enewetak is based on studies carried out on symbiotic (commensal and parasitic) species as well as those found as part of the plankton community (Chapter 20, this volume). A checklist of copepod fauna is found in Table 1. Copepods associated with marine invertebrates at Enewetak have been recorded since 1970 mainly through the efforts of A. J. Humes. Twenty-five of these copepods, their hosts, and references are given in Table 2. These include two harpactacoid, 19 poecilostomatoid (as cyclo- poid), and four siphonostomatoid (as cyclopoid) forms. As a result of these publications, one new genus and 14 new species were reported from Enewetak; additional species remain to be described (Humes, personal communication). Parasitic copepods from fishes include two poecilo- stomatoid (as cyclopoid) and 20 siphonostomatoid (as caligoid) species, three of which were new to science (Lewis, 1964; 1968). These copepods and their fish hosts TABLE 1 Checklist of Nonplanktonic Copepoda of Enewetak Atoll “Subclass COPEPODA Order HARPACTICOIDA Family CANUELLIDAE Sunaristes dardani Humes and Ho, 1969: Humes, 1971b. Family TEGASTIDAE Tesgastes acroporanus Humes, 1981: Humes, 1981. Order POECILOSTOMATOIDA Family BOMOLOCHIDAE Nothobomolochus gibber (Shiino, 1957): Lewis, 1968. Pseudotaeniacanthus sp.: Lewis, 1968. Family LICHOMOLGIDAE Acanthomolgus fissisetiger (Humes and Ho, 1968): Humes 1973b. Anisomolgus insolens (Humes and Ho, 1968): Humes, 1973b. Lichomolgus tridacnae Humes, 1972: Humes, 1972b. Metaxymolgus aculeatus (Humes and Ho, 1968): Humes, 1973b. Octopicola regalis Humes, 1974: Humes, 1974. Paradoridicola adelphus (Humes and Ho, 1968): Humes 1973b. Paramolgus eniwetokensis Humes, 1973: Humes, 1973b. Paramolgus ostentus Humes, 1973: Humes, 1973b. Schedomolgus lobophorus Humes and Ho, 1968: Humes and Stock, 1973. Synstellicola acanthasteris (Humes, 1970). Stellicola acanthasteris Humes, 1970: Humes, 1970a. Synonomy based on Humes (1976). Family MYICOLIDAE Anthessius alatus Humes and Stock, 1965: Humes, 1972b. Anthessius amicalis Humes and Stock, 1965: Humes, 1972b. Anthessius solidus Humes and Stock, 1965: Humes, 1972b. Family PSEUDANTHESSIDAE Pseudanthessius comanthi Humes, 1972: Humes, 1972a. Senariellus tensus Humes, 1977: Humes, 1977. Family SABELLIPHILIDAE Scambicornus idoneus (Humes and Cressey, 1961): Humes, 1980. Family TAENIACANTHIDAE Clavisodalis heterocentroti Humes, 1970: Humes, 1970b. Family XARIFIDAE Xarifia breviramea Humes and Dojiri, 1982: Humes and Dojiri, 1982. Xarifia sabiuraensis Misaki, 1978: Humes and Dojiri, 1982. Order SIPHONOSTOMATOIDA Family CALIGIDAE Anuretes serratus Shiino, 1954: Lewis, 1968. Caligus alaihi Lewis, 1968: Lewis, 1968. Caligus asymmetricus Kabata, 1965: Lewis, 1968. Caligus bonito Wilson, 1905: Lewis, 1968. Caligus(?) confusus Pillai, 1961: Lewis, 1968. ‘The higher classification used herein (to Family level) follows Bowman and Abele (1982). 197 (This table continued on next page.) 198 DEVANEY TABLE 1 (cont’d) Family CALIGIDAE (cont’d) Caligus coryphaenae Steenstrup and Lutken, 1861: Lewis, 1968. Caligus kapuhili Lewis, 1967: Lewis, 1968. Caligus laticaudus Shiino, 1960: Lewis, 1968. Caligus ligatus Lewis, 1964: Lewis, 1968. Caligus productus Dana, 1853: Lewis, 1968. Caligus pseudokalumai Lewis, 1968: Lewis, 1968. Dentigryps litus Lewis, 1964: Lewis, 1964; 1968. Lepeophtheirus dissimulatus Wilson, 1905; Lewis, 1968. Lepeophtheirus(?) plectropomi Nunes-Ruivo and Fourmanoir, 1956: Lewis, 1968. Pseudanuretes pomacanthi Lewis, 1968: Lewis, 1968. Pseudocaligus similis Lewis, 1968: Lewis, 1968. Family DISSONIDAE Dissonus heronensis Kabata, 1966: Lewis, 1968. Dissonus similis Kabata, 1966: Lewis, 1968. Family EDRYPHORIDAE Alebion gracilis Wilson, 1905: Lewis, 1968. Family NANASPIDIDAE Nanaspis manca Humes, 1973: Humes, 1973a; 1980. Nanaspis pusilla Humes, 1973: Humes, 1973a; 1980. Nanaspis spinifera Humes, 1973: Humes, 1973a; 1980. Family PANDARIDAE Pandarus cranchii Leach, 1819: Lewis, 1968. Family STELLICOMITIDAE Astroxynus culcitae Humes, 1971: Humes, 1971a. TABLE 2 Invertebrate Host and Symbiotic Copepoda Recorded from Enewetak Atoll Hosts Copepod symbionts References CNIDARIA Octocorallia (Soft corals) Alcyonacea Lobophytum pauciflorum Lobophytum pauciflorum Lobophytum pauciflorum Lobophytum pauciflorum Nephthea chabrolii Humes (1973b) Humes (1973b) Humes (1973b) Humes (1973b) Humes (1973b) Anisomolgus insolens Paradoridicola adelphus Paramolgus eniwetokensis Paramolgus ostentus Metaxymolgus aculeatus Stolonifera Pachyclavularia violacea Acanthomolgus fissisetiger | Humes (1973b) Zoantharia Scleractinia (Stony corals) Acropora florida Schedomolgus lobophorus Humes and Stock (1973) Acropora florida Tegastes acroporanus Humes (1981) Xarifia breviramea Xarifia sabiuraensis ARTHROPODA Humes and Dojiri (1982) Humes and Dojiri (1982) Acropora florida Acropora florida Crustacea Coenobitoidea (Hermit crabs) Dardanus guttatus Dardanus lagopodes Dardanus megistos Dardanus scutellatus Sunaristes dardani Sunaristes dardani Sunaristes dardani Sunaristes dardani MOLLUSCA Humes (1971b) Humes (1971b) Humes (1971b) Humes (1971b) Bivalvia (Tridacnid clams) Hippopus hippopus Hippopus hippopus Anthessius alatus Anthessius amicalis Humes (1972b) Humes (1972b) Tridacna gigas Tridacna gigas Tridacna maxima Tridacna squamosa Tridacna squamosa Tridacna squamosa Tridacna squamosa Cephalopoda Octopus cyaneus Anthessius alatus Lichomolgus tridacnae Anthessius alatus Anthessius alatus Anthessius amicalis Anthessius solidus Lichomolgus tridacnae Octopicola regalis (This table continued on next page.) Humes (1972b) Humes (1972b) Humes (1972b) Humes (1972b) Humes (1972b) Humes (1972b) Humes (1972b) Humes (1974) are given in Table 3. Cressey and Cressey (1979) also reported three new species of parasitic copepods on syn- odontid lizard fishes from Bikini Atoll, northwest of Enewetak. NONPLANKTONIC COPEPODA TABLE 2 (cont’d) Hosts Copepod symbionts References Asteroidea (Sea stars) Acanthaster planci Culcita novaeguineae Crinoidea (Feather stars) Comanthus bennetti Echinoidea (Sea urchins) Heterocentrotus trigonarius Echinothrix calamaris Holothuroidea (Sea cucumbers) Thelenota ananas Thelenota ananas Thelenota ananas Holothuria atra ECHINODERMATA Synstellicola acanthasteris Astroxynus culcitae Pseuanthessius comanthi Clavisodalis heterocentroti Senariellus tensus Nanaspis manca Nanaspis pusilla Nanaspis spinifera Scambicornus idoneus Humes (1970a) Humes (197 1a) Humes (1972a) Humes 1970b) Humes (1977) Humes (1973a; 1980) Humes (1973a; 1980) Humes (1973a; 1980) Humes (1980) 199 While no free-living nonplanktonic copepods have been reported from Enewetak, nearly 60 species were recorded from Ifaluk Atoll in the Caroline Islands (Vervoort, 1964). Many of these and others are likely to occur at Enewetak. TABLE 3 Parasitic Copepods on Fish at Enewetak Atoll Hosts “Pisces Elasmobranchii (Sharks) Carcharhinidae Parasitic copepod References CHORDATA Carcharhinus amblyrhynchos _ Alebion gracilis (C. menisorrah) Galeocerdo cuvier (C. cuvieri) Osteichthyes (Bony fishes) Belonidae Platybelone argalus platyura (Belone platyura) Holocentridae Neoniphon sammara (Holocentrus sammara) Sargocentron spiniferum (Holocentrus spinifer) Aulostomidae Aulostomus chinensis Fistulariidae Fistularia commersonii (F. petimba) Serranidae Epinephelus fuscoguttatus Epinephelus hoedtii (E. kohleri) Plectropomus sp. (P. leopardus) Pandarus cranchii Nothobomolochus gibber Caligus alaihi Caligus ligatus Dentigryps litus Pseudocaligus similis Lepeophtheirus plectropomi? Lepeophtheirus plectropomi? Dentigryps litus “Systematic arrangement after Randall and Randall, this volume. (This table continued on next page.) Lewis (1968) Lewis (1968) Lewis (1968) Lewis (1968) Lewis (1968) Lewis (1968) Lewis (1968) Lewis (1968) Lewis (1968) Lewis (1968) 200 DEVANEY TABLE 3 (cont'd) Hosts Parasitic copepod References Carangidae Caranyx melampygus Caligus confusus? Lewis (1968) Caranyx melampygus? Caligus coryphaenae Lewis (1968) Mullidae Parupeneus cyclostomus? Lepeophtheirus dissimulatus Lewis (1968) Chaetodontidae Chaetodon auriga Caligus kapuhili Lewis (1968) Chaetodon lunula Caligus kapuhili Lewis (1968) Pomacanthidae Pomacanthus imperator Pseudanuretes pomacanthi Lewis (1968) Acanthuridae Acanthurus nigricauda Pseudotaeniacanthus sp. Lewis (1968) (A. gahhm) Acanthurus olivaceus Caligus laticaudus Lewis (1968) Naso vlamingii Anuretes serratus Lewis (1968) Scombridae Euthynnus affinis Caligus coryphaenae Lewis (1968) (E. yaito) Grammatorcynus bilineatus Caligus asymmetricus Lewis (1968) Caligus coryphaenae Lewis (1968) Gymnosarda unicolor (G. nuda) Katsuwonus pelamis +Sarda orientalis +Sarda orientalis +Sarda orientalis +Sarda orientalis Caligus pseudokalumai Caligus coryphaenae Caligus asymmetricus Caligus bonito Caligus croyphaenae Caligus productus Cressey and Cressey (1980); Lewis (1968) Lewis (1968) Lewis (1968) Lewis (1968) Lewis (1968) Lewis (1968) Balistidae Balistoides viridescens Balistoides viridescens Tetraodontidae Arothron meleagris Dentigryps litus Dissonus heronensis Dissonus similis Lewis (1964; 1968) Lewis (1968) Lewis (1968) +Considered misidentification of host (Bruce B. Collette, personal communication). REFERENCES Bowman, T. A., and L. G. Abele, 1982, Classification of the Recent Crustacea, L. G. Abele (Ed.), The Biology of Crustacea Volume 1 Systematics, the Fossil Record, and Biogeography, Academic Press, New York, pp. 1-27. Cressey, R., and H. B. Cressey, 1979, The Parasitic Copepods of Indo-West Pacific Lizardfishes (Synodontidae), Smithson. Con- trib. Zool., 296: 1-71. __, 1980, Parasitic Copepods of Mackerel- and Tuna-Like Fishes (Scombridae) of the World, Smithson. Contrib. Zool., 311: 1-186. Humes, A. G., 1970a, Stellicola acanthasteris n. sp. (Copepoda, Cyclopoida) Associated with the Starfish Acanthaster planci (L.) at Eniwetok Atoll, Publ. Seto Mar. Biol. Lab., 17: 329-338. ——, 1970b, Clavisodalis heterocentroti gen. n. et sp. n., a Cyclo- poid Copepod Parasitic on an Echinoid at Eniwetok Atoll, J. Parasitol., 56: 575-583. ——, 1971a, Cyclopoid Copepods (Stellicomitidae) Parasitic on Sea Stars from Madagascar and Eniwetok Atoll, J. Parasitol., 57: 1330-1343. ——, 1971b, Sunaristes (Copepoda, Harpacticoida) Associated with Hermit Crabs at Eniwetok Atoll, Pac. Sci., 25: 529-532. —_, 1972a, Pseudanthessius comanthi n. sp. (Copepoda, Cyclopoida) Associated with a Crinoid at Eniwetok Atoll, Pac. Sci., 26: 373-380. __, 1972b, Cyclopoid Copepods Associated with Tridacnidae (Mollusca, Bivalvia) at Eniwetok Atoll, Proc. Biol. Soc. Wash., 84: 345-358. __, 1973a, Nanaspis (Copepoda, Cyclopoida) Parasitic on the Holothurian Thelenota ananas (Jaeger) at Eniwetok Atoll, J. Parasitol., 59: 384-395. __, 1973b, Cyclopoid Copepods (Lichomolgidae) from Octo- corals at Eniwetok Atoll, Beaufortia, 21: 135-151. —_, 1974, Octopicola regalis n. sp. (Copepoda, Cyclopoida, Lichomolgidae) Associated with Octopus cyaneus from New Caledonia and Eniwetok Atoll, Bull. Mar. Sci., 24: 76-85. ___, 1976, Cyclopoid Copepods Associated with Asteroid Echino- derms in New Caledonia, Smithson. Contrib. Zool., 217: 1-19. 1977, Pseudanthesiid Copepods (Cyclopoida) Associated with Crinoids and Echinoids (Echinodermata) in the Tropical Western Pacific Ocean, Smithson. Contrib. Zool., 243: 1-43. ___, 1980, A Review of the Copepods Associated with Holothuri- ans Including New Species from the Indo-Pacific, Beaufortia, 30: 31-123. NONPLANKTONIC COPEPODA —_, 1981, A New Species of Tegastes (Copepoda, Harpac- ticoida) Associated with a Scleractinian Coral at Enewetak Atoll, Proc. Biol. Soc. Wash., 94: 254-263. __., and M. Dojiri, 1982, Xarifiidae (Copepoda) Parasitic in Indo- Pacific Scleractinian Corals, Beaufortia, 32: 139-228. __., and J. H. Stock, 1973, A Revision of the Family Lichomolgi- dae Kossmann, 1877, Cyclopoid Copepods Mainly Associated 201 with Marine Invertebrates, Smithson. Contrib. Zool., 127: 1-368. Lewis, A. G., 1964, The Caligid Copepod Genus Dentigryps (Crustacea: Calligoida), Proc. U. S. Nat. Mus., 115: 347-380. __, 1968, Copepod Crustaceans Parasitic on Fishes of Eniwetok Atoll, Proc. U. S. Nat. Mus., 125: 1-78. Vervoort, W., 1964, Free-Living Copepods from Ifaluk Atoll in the Caroline Islands., U. S. Nat. Mus. Bull., 236: 1-431. a Chapter 20 Lagoon Plankton of Enewetak Atoll RAY P. GERBER Biology Department, St. Joseph’s College North Windham, Maine 04062 INTRODUCTION The zooplankton community of atoll lagoons is uniquely different from the zooplankton community in the ocean waters outside the atoll for at least two reasons. First, there is a greater abundance of zooplankton in the lagoon, no doubt reflecting the better food conditions pro- vided by exported reef detritus and phytoplankton produc- tion (Gerber and Marshall, 1982). Second, the species composition of the lagoon zooplankton largely consists of species adapted to the restricted shallower conditions found therein. Earlier studies on the zooplankton of atoll lagoons were mostly concerned with determining abun- dance, either to evaluate their food potential to corals (e.g., Kramer, 1897; Russell, 1934; Motoda, 1938; Odum and Odum, 1955; Johannes et al., 1970; Johannes and Gerber, 1974) or in understanding the water exchange pat- terns between lagoons and the surrounding sea (e.g., John- son, 1949, 1954; Gilmartin, 1958; Michel, 1969; Michel et al., 1971; Tranter and George, 1972). In these studies little attempt was made to identify the organisms to species. However, these works are important because they provide a basic understanding of the ecology of zooplank- ton in lagoon environments. For a more detailed discussion on the ecological role of zooplankton in atoll lagoons see Chapter 10 by Gerber and Marshall, Volume 1. Studies on the species composition of lagoon zooplank- ton at Enewetak Lagoon and nearby atolls have for the most part focused on the holoplanktonic organisms, forms that are permanently planktonic all of their lives; these forms are the main attention of this chapter. Because infor- mation is lacking on the life history of many of the species mentioned, some meroplanktonic forms—organisms that are temporarily planktonic—are no doubt included in this checklist (Table 1). In the works discussed below, a variety of sampling methods have been used to collect the zoo- plankton, and each method has its inherent advantages and disadvantages. In a sense it is perhaps fortunate that no one method is best to sample the various types of zoo- 203 plankton. One important aspect lacking in the data is ade- quate seasonal samplings to provide information on how species composition and abundance change over time. Whereas most of the lagoon plankton studies were based on samples collected over a few days in one season, the present author has sampled more extensively, during two winter periods and one summer period. DINOPHYTA Dinophyceae (Dinoflagellata) Nineteen species of dinoflagellates were identified from plankton tows taken off the wide passage and/or deep entrance at Enewetak during the summer of 1956 (Hirsh- field et al., 1957). Their species list and nomenclature were originally based on Lebour (1925) and Schiller (1937). They nave been revised here using Parke and Dixon (1976). More recently, three additional planktonic species were recorded from Enewetak at a mid-lagoon and shallow station during two winters and one summer period (Gerber, 1981). SARCODINA Radiolaria Fifty species (including 18 undetermined) of radiolari- ans were collected from plankton hauls made during the summer of 1956 (Hirshfield et al., 1957) (Table 1). Gilmar- tin (1958) and Gerber (1981) have recorded radiolarians in plankton samples from Enewetak Lagoon but have not attempted identification. Radiolarian fragments were recorded in the gut contents of calanoid copepods from Enewetak Lagoon (Gerber and Marshall, 1974). For radiolarian classification see Campbell and Moore (1954). CILIOPHORA Tintinnina Nine species of tintinnids were recorded from plankton hauls taken in the wide passage and/or deep entrance at Enewetak during the summer of 1956 (Hirshfield et al., 1957). Four of the species from Enewetak were con- sidered to be new and are not known to have been described. In February 1976, plankton tows in Enewetak 204 GERBER TABLE 1 Checklist of Plankton of Enewetak Atoll Phylum DINOPHYTA (DINOFLAGELLATES) Class DINOPHYCEAE Order DINOPHYSIALES Family AMPHISOLENIACEAE Amphisolenia spp.: Hirshfield et al. 1957. Family DINOPHYSIACEAE Dinophysis rontundatum (Clap et Lach): Hirshfield et al. 1957. Dinophysis schuettii (Murray et Whitting): Hirshfield et al. 1957. Phalacroma (?) perradictyum Stein: Hirshfield et al. 1957. Family ORNITHOCERCIACEAE Ornithocercus splendidus Murray and Whitting: Hirshfield et al. 1957. Ornithocercus sp. Family CERATOCORY ACEAE Ceratocorys horrida Stein: Hirshfield et al. 1957. Order PYROCYSTALES Family PYROCYSTACEAE Pyrocystis fusiformis Thompsen ex Murray: Gerber, 1981. Pyrocystis pseudonoctiluca Thompsen ex Murray: Gerber, 1981. Dissodinium lunula (Schutt) Taylor: Gerber, 1981. Order PERIDINIALES Family PERIDINIACEAE Protoperidinium divergens (Ehrenberg) Balach. Peridinium divergens (Ehrenberg): Hirshfield et al. 1957. Protoperidinium depressum (Bailey) Balach. Peridinium depressum (Bailey): Hirshfield et al. 1957. Protoperidinium globulum (Stein) Balach. Peridinium globulum (Stein): Hirshfield et al. 1957. Peridinium oceanicum (Vanhoffen) Balach. Peridinium oceanicum (Vanhoffen): Hirshfield et al. 1957. Family GONYAULACACEAE Gonyaulax birostris (Stein): Hirshfield et al. 1957. Family CERATIACEAE Ceratium candelabrum (Ehrenberg) Stern: Hirshfield et al. 1957. Ceratium furca (Ehrenberg) Clap. et Lachm.: Hirshfield et al. 1957. Ceratium gibberum Gourret: Hirshfield et al. 1957. Ceratium lineatum (Ehrenberg) Cleve: Hirshfield et al. 1957. Ceratium tripos (O. F. Muller) Nitzsch: Hirshfield et al. 1957. Ceratium tripos O. F. Muller var. B: Hirshfield et al. 1957. Phylum SARCODINA Class ACTINOPODEA Order PORULOSIDA Suborder ACANTHARINA Acanthometron sp.: Hirshfield et al. (1957). Acantholonche sp. A.: Hirshfield et al. (1957). Acantholonche sp. B: Hirshfield et al. (1957). Acanthochiasma fusiformis Haeckel: Hirshfield et al. (1957). Acanthostaurus purpurascens Haeckel: Hirshfield et al. (1957). Aspidomma sp.: Hirshfield et al. (1957). Dodecapsis tricinata Haeckel: Hirshfield et al. (1957). Dorataspis sp.: Hirshfield et al. (1957). Hexalaspis heliodiscus Haeckel: Hirshfield et al. (1957). Hexacolpus infundibulum Haeckel: Hirshfield et al. (1957). Hexonaspis hastataz Haeckel: Hirshfield et al. (1957). Hystrichaspis dorsata Haeckel: Hirshfield et al. (1957). Phractaspis sp.: Hirshfield et al. (1957). Quadrilonche mesostauras Haeckel: Hirshfield et al. (1957). Tessarapelma concretuira Haeckel: Hirshfield et al. (1957). Tessarapelma concretuira Haeckel: Hirshfield et al. (1957). Zygostaurus sagitalis Haeckel: Hirshfield et al. (1957). Zygacantha sp.: Hirshfield et al. (1957). (This table continued on next page.) LAGOON PLANKTON TABLE 1 (cont’d) Suborder SPUMELLINA Diplosphaera hexagonalis Haeckel: Hirshfield et al. (1957). Euchitonia lanceolata Haeckel: Hirshfield et al. (1957). E. muelleri Haeckel: Hirshfield et al. (1957). Flustrella semispiralis Haeckel: Hirshfield et al. (1957). Heliodiscus asteriscus Haeckel: Hirshfield et al. (1957). Hexalonche philosophica Haeckel: Hirshfield et al. (1957). Hexastylus phaenaxonia Haeckel: Hirshfield et al. (1957). Hymeniastrum euclidis Haeckel: Hirshfield et al. (1957). Monozonium alatum Haeckel: Hirshfield et al. (1957). Ommatospyris virginea Haeckel: Hirshfield et al. (1957). O. hexagonium Haeckel: Hirshfield et al. (1957). Panartus pluteus Haeckel: Hirshfield et al. (1957). Sphaerozoum sp.: Hirshfield et al. (1957). Spongaster tetras Ehrenberg: Hirshfield et al. (1957). Spongodiscus velella Haeckel: Hirshfield et al. (1957). Stylospongia sp.: Hirshfield et al. (1957). Tetrapyle sp.: Hirshfield et al. (1957). Order OSCULOSIDA Suborder NASSELLINA Adelocyrtis sp.: Hirshfield et al. (1957). Archiscenium sp.: Hirshfield et al. (1957). Callimitra agnesae Haeckel: Hirshfield et al. (1957). Corocalyptra elisabethae Haeckel: Hirshfield et al. (1957). Dictyocephalus sp.: Hirshfield et al. (1957). Dictyoceras sp.: Hirshfield et al. (1957). Eucyrtidium hexagonatum Haeckel: Hirshfield et al. (1957). Eucyrtidium sp.: Hirshfield et al. (1957). Lychnocanium pyriforme Haeckel: Hirshfield et al. (1957). Monostephus sp.: Hirshfield et al. (1957). Pterocanium tricolpum Haeckel: Hirshfield et al. (1957). Pterocorys campanula Haeckel: Hirshfield et al. (1957). Semantis sp.: Hirshfield et al. (1957). Theocorys sp.: Hirshfield et al. (1957). Tricylidium dictyospyris Haeckel: Hirshfield et al. (1957). Zygostephanium sp.: Hirshfield et al. (1957). Phylum CILIOPHORA Class POLYHYMENOPHORA Order OLIGOTRICHIDA Suborder TINTINNINA 205 Family CODONELLIDAE Tintinnopsis sp.: Hirshfield et al. 1957. Codonella cuspidata Kofoid and Campbell: Gold and Morales (1977). Codonaria oceanica (Brandt) Kofoid and Campbell: Gold and Morales (1977). Family CODONELLOPSIDAE Codonellopsis americana Kofoid and Campbell: Gold and Morales (1977). Family CYTTAROCYCLIDAE Farella azorica (Cleve) Jorgensen: Hirshfield et al. 1957. Family PTYCHOCYLIDAE Epiplocyclis sp.: Hirshfield et al. 1957. Poroecus apicatus Kofoid and Campbell: Gold and Morales (1977). Poroecus apiculatus (Cleve) Cleve: Gold and Morales (1977). Poroecus curtus Kofoid and Campbell: Gold and Morales (1977). Family EPIPLOCYLIDAE Epiplocylis labiosa Kofoid and Campbell: Gold and Morales (1977). Epiplocylis pacifica Kofoid and Campbell: Gold and Morales (1977). Family PETALOTRICHIDAE Craterella torulata Jorgensen: Gold and Morales (1977). Petalotricha major Jorgensen: Gold and Morales (1977). Family RHABDONELLIDAE Protorhabdonella simplex (Cleve) Jorgensen: Gold and Morales (1977). Protorhabdonella striatura Kofoid and Campbell: Gold and Morales (1977). (This table continued on next page.) 206 GERBER TABLE 1 (cont'd) Family RHABDONELLIDAE (cont’d) Rhabdonella amor (Cleve) Brandt: Gold and Morales (1977). Rhabdonella elegens (Jorgensen) Kofoid and Campbell: Gold and Morales (1977). Rhabdonella inflata Kofoid and Campbell: Hirshfield et al. (1957). Rhabdonella poculum (Ostenfeld and Schmidt): Hirshfield et al. (1957). Rhabdonellopsis apophysata Cleve: Gold and Morales (1977). Family XYSTONELLIDAE Xystonella sp.: Hirshfield et al. (1957). Family UNDELLIDAE Undella hemispherica Laachmann: Gold and Morales (1977). Proplectella biorbiculata Hada: Gold and Morales (1977). Proplectella perpusilla Kofoid and Campbell: Gold and Morales (1977). Family DICTYOCYSTIDAE Dictyocysta occidentalis Kofoid and Campbell: Gold and Morales (1977). Family TINTINNIDAE Amphorella brandti Jorgensen: Gold and Morales (1977). Canthariella brevis Kofoid and Campbell: Hirshfield et al. (1957). Dadayiella (?) ganymedes Entz, Sr.: Hirshfield et al. (1957). Steenstrupiella gracilis (Jorgensen) Kofoid and Campbell: Gold and Morales (1977). Steenstrupiella steenstrupii (Claparede and Lachmann) Kofoid and Campbell: Gold and Morales (1977). Tintinnus fraknoii Daday: Gold and Morales (1977). Tintinnus lususundae var tenuis (Kofoid and Campbell) Hada: Gold and Morales (1977). Tintinnus lususundae var turgescens: Gold and Morales (1977). Tintinnus sp.: Hirshfield et al. (1957). Phylum CNIDARIA Class HYDROZOA Order SIPHONOPHORA Suborder CALYCOPHORA Family DIPHYIDAE *Abylopsis eschscholtzii Huxley: Sears, 1950. Abylopsis tetragona Otto: Sears, 1950; Gerber, 1981. *Agalma okeni Eschscholtz: Sears, 1950. Bassia bassensis Quoy and Gaimard: Sears, 1950; Gerber, 1981. Chelophyces contorta Lens and Van Riemsdijk: Sears, 1950; Gerber, 1981; Alvarino (personal communication). Diphyes chamissonis Huxley: Sears, 1950; Gerber, 1981. Diphyes dispar Chamiso and Eysenhardt: Sears, 1950; Gerber, 1981. *Lensia conoidea Keferstein and Ehlers: Sears, 1950. Lensia subtillis Chum: Gerber, 1981. Lensia subtiloides Lens and Van Riemsdijk: Gerber, 1981. Suborder PHYSOPHORIDA Family PHYSALIIDAE +Physalia utriculus (Gmelin, 1790) Order HYDROIDA Suborder LEPTOMEDUSAE (CALYPTOBLATINA) Family AEQUOREIDAE +Aequorea sp. Suborder CHONDROPHORA Family VELELLIDAE +Velella sp. Family PORPITIDAE +Porpita pacifica Lesson, 1826. Class SCYPHOZOA Order SEMAEOSTOMAE Family ULMARIDAE +Aurelia sp. of A. labiata Chamisso and Eysenhardi, 1820. Family PELAGIDAE +Pelagia noctiluca (Forsskal, 1775) * Only recorded outside lagoon but in the immediate vicinity of Enewetak Atoll. + These were identified in the ocean waters neighboring Bikini Atoll (Chiba et al., 1955). (This table continued on next page.) LAGOON PLANKTON 207 TABLE 1 (cont’d) Order RHIZOSTOMAE Suborder KOLPOPHORAE Family MASTIGIIDAE +Mastigias ocellatus (Modeer, 1791). Family THYSANOSTOMATIDAE Thysanostoma flagellatum (Haeckel): Arneson, 1980. Phylum CTENOPHORA Class TENTACULATA Order LOBATA Family MNEMIIDAE Mnemiopais sp.: Gilmartin, 1958; Gerber, 1981. Phylum MOLLUSCA Class GASTROPODA Order THECOSOMATA (PTEROPODA) Family CAVOLINIDAE Creseis acicula Rang: Gerber, 1981. Creseis virgula Rang: Gilmartin, 1958. Creseis sp.: Johnson, 1954. Family LIMACINIDAE Limacina sp.: Gerber, 1981. Order MESOGASTROPODA (HETEROPODA) Family ATLANTIDAE Atlanta sp.: Gilmartin, 1958. Phylum ARTHROPODA Class CRUSTACEA Subclass COPEPODA Order CALANOIDA Family CALANIDAE Mesocalanus tenuicornis Dana: Mahnken, 1966; Gerber, 1981. Calanus minor (Claus): Mahnken, 1966; Barnett, 1967; Gerber, 1981. Canthocalanus pauper (Giesbrecht): Johnson, 1954; Barnett, 1967; Gerber, 1981. Neocalanus gracilis (Dana): Gilmartin, 1958; Barnett, 1967. Undinula vulgaris (Dana): Johnson, 1954; Gilmartin, 1958; Mahnken; 1966, Barnett, 1967; Hobson and Chess, 1978; Gerber, 1981. Cosmocalanus darwini (Lubbock): Johnson, 1954; Mahnken, 1966; Barnett, 1967; Gerber, 1981. Family EDCALANIDAE Eucalanus attenuatus (Dana): Johnson, 1954; Gerber, 1981. Eucalanus monochus Giesbrecht: Gerber, 1981. Mecynocera clausi Thomson: Barnett, 1967; Gerber, 1981. Rhincalanus sp. Dana: Mahnken, 1966. Family PARACALANIDAE Paracalanus parvus (Claus): Barnett, 1967. Acrocalanus gibber Giesbrecht: Gilmartin, 1958; Barnett, 1967; Gerber, 1981. Acrocalanus gracilis Giesbrecht: Mahnken, 1966; Barnett, 1967; Gerber, 1981. Acrocalanus longicornis Giesbrecht: Barnett, 1967; Gerber, 1981. Acrocalanus monachus Giesbrecht: Mahnken, 1966; Barnett, 1967; Gerber, 1981. Calocalanus pavo (Dana): Johnson, 1954; Mahnken, 1966; Barnett, 1967; Gerber, 1981. Calocalanus pavoninus Farran: Gerber, 1981. Calocalanus plumulosus (Claus): Barnett, 1967; Gerber, 1981. Calocalanus styliremus Giesbrecht: Barnett, 1967; Gerber, 1981. Family CLAUSOCALANIDAE Clausocalanus arcuicornis (Dana): Mahnken, 1966; Barnett, 1967; Gerber, 1981. Clausocalanus farrani Sewell: Gerber, 1981. Clausocalanus furcatus (Brady): Mahnken, 1966; Gerber, 1981. Clausocalanus pergens Farran: Johnson, 1954; Barnett, 1967; Gerber, 1981. Family EDCHAETIDAE Euchaeta rimana (Bradford): Gerber, 1981. Euchaeta marina (Prestandrea): Gilmartin, 1958; Mahnken, 1966; Hobson and Chess, 1978. + These were identified in the ocean waters neighboring Bikini Atoll (Chiba et al., 1955). (This table continued on next page.) 208 GERBER TABLE 1 (cont’d) Family SCOLECITRICHIDAE Scolecithricella dentate (Giesbrecht): Barnett, 1967; Gerber, 1981. Scolecithricella marginata (Giesbrecht): Mahnken, 1966; Barnett, 1967. Scolecithricella tenuisserata (Giesbrecht): Barnett, 1967. Scolecithrix danae (Lubbock): Johnson, 1954; Gilmartin, 1958; Mahnken, 1966; Gerber, 1981. Family TEMORIDAE Temora discaudata Giesbrecht: Mahnken, 1966; Gerber, 1981. Family PSEUDODIAPTOMIDAE Pseudodiaptomus cornutus Nicholl: Barnett, 1967. Family METRIDIIDAE Metridia sp. Boeck: Hobson and Chess, 1978. Pleuromamma abdominalis (Lubbock): Barnett, 1967. Pleuromamma gracilis (Claus): Barnett, 1967; Gerber, 1981. Pleuromamma xiphias (Giesbrecht): Hobson and Chess, 1978. Family CENTROPAGIDAE Centropages calaninus (Dana): Gilmartin, 1958; Mahnken, 1966; Barnett, 1967; Gerber, 1981. Centropages elongatus Giesbrecht: Johnson, 1954; Mahnken, 1966; Barnett, 1967; Gerber, 1981. Centropages gracilis (Dana): Barnett, 1967; Gerber, 1981. Centropages orsinii Giesbrecht: Mahnken, 1966; Barnett, 1967; Gerber, 1981. Family LUCICUTIIDAE Lucicutia flavicornis (Claus): Mahnken, 1966; Barnett, 1967; Gerber, 1981. Lucicutia ovalis Wolfenden: Gerber, 1981. Family HETERORHABDIDAE Heterorhabdus papilliger (Claus): Barnett, 1967. Family ARIETELLIDAE Paramisophria ? sp. Scott: Barnett, 1967. Family PSEUDOCYCLOPIDAE Pseudocyclops sp. Brady: Barnett, 1967. Family CANDACIIDAE Candacia aethiopica (Dana): Gilmartin, 1958; Mahnken, 1966; Barnett, 1967; Gerber, 1981. Candacia bispinosa (Claus): Barnett, 1967; Gerber, 1981. Candacia catula (Giesbrecht): Barnett, 1967; Gerber, 1981. Candacia discaudata A. Scott: Barnett, 1967. Paracandacia simplex (Giesbrecht): Barnett, 1967. Paracandacia truncata (Dana): Mahnken, 1966; Barnett, 1967; Gerber, 1981. Family PONTELLIDAE Calanopia minor A. Scott: Johnson, 1954; Barnett, 1967; Gerber, 1981. Labidocera laevidentata (Brady): Gilmartin, 1958; Barnett, 1967; Gerber, 1981. Pontellina plumata (Dana): Gilmartin, 1958; Gerber, 1981. Pontellopsis sp. G. Brady: Gilmartin, 1958. Family ACARTIIDAE Acartia fossae Gurney: Gerber, 1981. Acartia hamata Mori: Johnson, 1954; Mahnken, 1966; Barnett, 1967. Acartia negligens Dana: Mahnken, 1966; Barnett 1967; Gerber, 1981. Acartia spinicaudata Giesbrecht: Barnett, 1967. Family TORTANIDAE Tortanus gracilis (Brady): Johnson, 1954; Gilmartin, 1958; Mahnken, 1966; Barnett, 1967; Gerber, 1981. Tortanus murrayi A. Scott: Johnson, 1954. Order CYCLOPODA Family OITHONIDAE Oithona nana (Claus): Barnett, 1967; Gerber, 1981. Oithona plumifera Baird: Johnson, 1954; Mahnken, 1966; Barnett, 1967; Gerber, 1981. Oithona pseudofrigida (Giesbrecht): Barnett, 1967; Gerber, 1981. Oithona rigida Giesbrecht: Barnett, 1967; Gerber, 1981. Oithona tenuis Rosendorn: Barnett, 1967; Gerber, 1981. Family CLAUSIDIIDAE Saphirella tropica Wolfenden: Gerber, 1981. Family ONCAEIDAE Oncaea media Giesbrecht: Barnett, 1967; Gerber, 1981. Oncaea venusta Philippi: Johnson, 1954; Barnett, 1967; Gerber, 1981. Lubbockia squillimana Claus: Johnson, 1954; Gerber, 1981. (This table continued on next page.) LAGOON PLANKTON 209 TABLE 1 (cont’d) Family SAPPHIRINIDAE Sapphirina stellata Giesbrecht: Johnson, 1954; Gerber, 1981. Sapphirina tropica Wolfenden: Gerber, 1981. Copilia mirabilis Dana: Johnson, 1954; Gilmartin, 1958; Mahnken, 1966; Barnett, 1967; Gerber, 1981. Copilia quadrata Dana: Barnett, 1967. Family CORYCAEIDAE Corycaeus agilis Dana: Mahnken, 1966; Barnett, 1967; Gerber, 1981. Corycaeus andrewsi Farran: Mahnken, 1966. Corycaeus asiaticus F. Dahl: Barnett, 1967; Gerber, 1981. Corycaeus catus F. Dahl: Barnett, 1967; Gerber, 1981. Corycaeus crassiusculus Dana: Mahnken, 1966; Barnett, 1967; Gerber, 1981. Corycaeus flaccus Giesbrecht: Gerber, 1981. Corycaeus gracilis Dana: Mahnken, 1966. Corycaeus latus Dana: Gerber, 1981. Corycaeus lautus Dana: Gerber, 1981. Corycaeus limbatus Brady: Barnett, 1967; Gerber, 1981. Corycaeus longistylis Dana: Barnett, 1967; Gerber, 1981. Corycaeus medius Gurney: Barnett, 1967; Gerber, 1981. Corycaeus pumilus M. Dahl: Mahnken, 1966. Corycaeus robustus Giesbrecht: Mahnken, 1966; Barnett, 1967. Corycaeus speciosus Dana: Mahnken, 1966; Barnett, 1967; Gerber, 1981. Corycaeus tenuis Giesbrecht: Gerber, 1981. Corycaeus typicus (Kroyer): Barnett, 1967; Gerber, 1981. Corycaeus vitreus Dana: Mahnken, 1966; Barnett, 1967; Gerber, 1981. Farranula carinata (Giesbrecht): Gerber, 1981. Corycaeus carinatus Giesbrecht: Barnett, 1967. Farranula concinna (Dana): Gerber, 1981. Corycaeus concinnus Dana: Mahnken, 1966; Barnett, 1967. Farranula gibbula (Giesbrecht): Gerber, 1981. Corycaeus gibbulus Giesbrecht: Mahnken, 1966; Barnett, 1967. Order HARPACTICOIDA Suborder OLIGOARTHRA Family ECTINOSOMATIDAE Microsetella norvegica (Boeck): Barnett, 1967. Microsetella rosea (Dana): Barnett, 1967; Gerber, 1981. Family CLYTEMNESTRIDAE Clytemnestra rostrata Brady: Gerber, 1981. Clytemnestra scutellata Dana: Gerber, 1981. Family MACROSETELLIDAE Macrosetella gracilis (Dana): Gilmartin, 1958; Barnett, 1967; Gerber, 1981. Family LONGIPEDIIDAE Longipedia coronata Claus: Gerber, 1981. Longipedia weberi A. Scott: Gerber, 1981. Family HARPACTICIDAE Harpacticus spp. M. Edwards: Gerber, 1981. Family PELTIDIIDAE Peltidium spp. Philippi: Gerber, 1981. Family TEGASTIDAE Tegestes sp. Norman: Gerber, 1981. Family THALESTRIDAE Eudactylopus andrewi Sewell: Gerber, 1981. Eudactylopus anomala Sewell: Gerber, 1981. Eudactylopus fasciatus Sewell: Gerber, 1981. Dactylopoda sp. G. O. Sars: Gerber, 1981. Family DIOSACCIDAE Amphiascus coralicola Sewell: Gerber, 1981. Amphiascopsis cinctus (Claus): Gerber, 1981. Metamphiascopsis hirsutus (Thompson and A. Scott): Gerber, 1981. Family METIDAE Metis sp. Philippi: Gerber, 1981. (This table continued on next page.) 210 GERBER TABLE 1 (cont'd) Family LAOPHONTIDAE Laophonte cornuta Philippi: Gerber, 1981. Order MONSTRILLOIDA Family MONSTRILLIDAE Monstrilla sp.: Barnett, 1967; Gerber, 1981. Subclass MALACOSTRACA Order MYSIDACEA Family MYSIDAE Subfamily SIRIELLINAE Siriella affinis Hanson: Murano, 1983. Siriella aequiremis Hanson: Murano, 1983. Siriella sp.: Hobsen and Chess, 1978. Subfamily MYSINAE Anisomysis chessi Murano: Murano, 1983. Anisomysis constricta Murano: Murano, 1983. Anisomysis enewetakensis Murano: Murano, 1983. Mysinae sp.: Hobsen and Chess, 1978. Subfamily GASTROSACCINAE Anchialina grossa Hansen: Gerber, 1981. Anchialina typica (Kreyer): Gerber, 1981. Gastrosaccus bengalensis Hansen: Murano, 1983. Gastrosaccus indicus Hansen: Gerber, 1981. Gastrosaccus pacificus Hansen: Gerber, 1981. Gastrosaccus parvus Hansen: Gerber, 1981. Gastrosaccus sp.: Gerber, 1981. Metamblyopsis sp.: Gerber, 1981. Pseudanchialina inermis Illig: Gerber, 1981; Murano, 1983. Order AMPHIPODA Suborder HYPERIIDEA Family HYPERIIDAE Lestrigonus bengalensis Giles: Gerber, 1981 as Hyperia dysschistus Stebbing. Lestrigonus latissima (Borallius): Gerber, 1981 as Hyperia hydrocephalia Vosseler. Family SYNOPIIDAE Synopia ultramarina Dana: Gerber, 1981. Synopia variabilis Spandl: Bernard, 1965. Order EUPHAUSIACEA Family EUPHAUSIIDAE Pseudeuphausia latifrons (G. O. Sars): Gerber, 1981. Order DECAPODA Family LUCIFERIDAE Lucifer chacei Bowman, 1967. Phylum CHAETOGNATHA Class SAGITTOIDEA Family SAGITTIDAE Pterosagitta draco (Krohn): Alvarino, personal communication: Gerber, 1981. Sagitta bedfordii: Alvarino, personal communication. Sagitta bipunctata Quoy and Gaimard: Alvarino, personal communication; Gerber 1981. Sagitta enflata Grassi: Johnson, 1954; Gilmartin, 1958; Alvarino, personal communication; Gerber, 1981. Sagitta ferox Doncaster: Alvarino, personal communication. Sagitta neglecta Aida: Gilmartin, 1958; Alvarino, personal communication; Gerber, 1981. Sagitta oceania: Alvarino, personal communication. *Sagitta pacifica Tokioka: Alvarino, personal communication. Sagitta regularis Aida: Gilmartin, 1958; Alvarino, personal communication; Gerber, 1981. Sagitta robusta Doncaster: Johnson, 1954; Alvarino, personal communication. Sagitta serratodentata Krohn: Johnson, 1954; Gerber, 1981. Phylum CHORDATA Subphylum UROCHORDATA Class THALIACEA Order DOLIOLIDA (CYCLOMY ARIA) Family DOLIOLIDAE Doliolum sp.: Kinzie, 1976. * Only recorded outside lagoon but in the immediate vicinity of Enewetak Atoll. (This table continued on next page.) LAGOON PLANKTON 211 TABLE 1 (cont'd) Order SALPIDA (DESMOMY ARIA) Family SALPIDAE Salpa sp.: Gilmartin, 1958. Thalia democratica (Forsskal): Gilmartin, 1958. Class APPENDICULARIA (COPELATA: LARVACEA) Family FRITILLARIDAE +Fritillaria megachile Fol: Gerber, 1981. Fritillaria sp.: Gerber, 1981. Family OIKOPLEURIDAE Oikopleura intermedia Lohmann: Gerber, 1981. +Oikopleura fusiformis Fol: Gerber, 1981. +Oikopleura longicaudata (Vogt): Gerber, 1981. +Oikopleura parva Lohmann: Gerber, 1981. +Oikopleura rufescens Fol: Gerber, 1981. + These were identified in the ocean waters neighboring Bikini Atoll (Chiba et al., 1955). Lagoon and the channel adjacent to Japtan Islet resulted in an additional 26 species (Gold and Morales, 1977), thus a total of 34 tintinnids is known from this area. Identifica- tion and nomenclature for the tintinnids are based on Campbell and Moore (1954), Corliss (1977), and Kofoid and Campbell (1939). For discussion of other Ciliophora and Foraminifera, see Chapter 4, this volume. CNIDARIA Hydrozoa Siphonophora Calycophorid siphonophores collected in net samples form a small fraction of the zooplankton in this area and have been most extensively sampled by Sears (1950) dur- ing the spring and summer of 1946. Two species were found within Enewetak Lagoon and eight outside, with only one common to both areas. At nearby Bikini, Rongelap, and Rongerik Atolls, 27 additional species were recorded with about one-third of those recorded in the lagoons. These appear to be largely oceanic species but include Abyla leukartii, A. trigona, Chelophyes appendicu- lata, Diphyes arctica, D. bojani, Enneagonum_hyalinuar, Eudoxides spiralis, Galetta australis, G. chuni, G. bigelowi, Lensia campanella, L. cossack, L. fowleri, L. grimaldii, L. hotspur, L. multicristata, Rosacea plicata, Sulculeolaria monoica, and S. quadridentata. Additional records of siphonophores identified from Enewetak Lagoon were made by Alvarino (personal communication) from samples collected in 1946 and by Gerber (1981) from 35 samples collected during the winters of 1972, 1974, and the sum- mer of 1974. In this last study, eight species were recorded from the Enewetak Lagoon (which includes the three species previously recorded); most species increased in abundance during the summer. Only one physophorid siphonophore, the Portuguese man-of-war, Physalia, has been collected and identified from Enewetak Lagoon where occasionally it appears commonly, sometimes washing up on lagoon islet beaches. Siphonophores were also recorded at Enewetak Lagoon by Johannes and Gerber (1974) during the sum- mer of 1972 and at Majuro Atoll Lagoon by Hobson and Chess (1973), but no species were identified. Taxonomy and identification of siphonophores can be found in Sears (1950) and Totton and Bargmann (1965). Hydroida Various Anthomedusae have been reported from net tows made in the lagoon by Gilmartin (1958) and Gerber (1981) but not identified. At least one species of Anthomedusae has been identified to genus (C. Arneson) but is unavailable. Representatives of this group commonly occur in the lagoon plankton and are usually damaged beyond recognition in net collections. Two species of chon- drophorans have been identified from the lagoon and are in the Mid-Pacific Marine Laboratory (MPML) reference collection. SCYPHOZOA Semaeostomae Two species have been collected from Enewetak Lagoon and have been identified and placed in the MPML reference collection. Rhizostomae One species of this group, preserved in the MPML reference collection, has been identified from the lagoon. It was collected by Trench and Colley in August 1978 off Bokoluo Island (Alice) and in the lagoon in October 1980 by C. Arneson. Two key works on the identification and taxonomy of the medusae (Hydroida and Scyphozoa) are Kramp (1961) and Mayer (1910). 212 GERBER CTENOPHORA Tentaculata Lobata One specimen preserved in the MPML reference collec- tion has been identified to genus and was collected in the lagoon off Enewetak Island December 28, 1975. Cteno- phores were collected from the lagoon in April and December 1955 (Gilmartin, 1958) and in the winters of 1972, 1974, and summer 1974 by Gerber (1981) but not identified to species. MOLLUSCA Gastropoda Two species of the holoplanktonic Thecosomata, collec- tively known as pteropods, have been reported from Enewetak Lagoon. Johnson (1949) and Gilmartin (1958) each reported one species from Enewetak and nearby Bikini, Rongelap, and Rongerik Lagoons. Pteropods were collected from shallow stations at Enewetak and Majuro Lagoons by Hobson and Chess (1973, 1978) but were not identified further. Gerber (1981) identified two species which commonly occurred in lagoon samples from shallow and deep stations; they were especially common in the summer months. Only one unidentified species of the heteropod genus Atlanta was recorded by Gilmartin (1958) from Enewetak Lagoon where it is not very abundant. Members of this genus have been reported in low abundance in nearby oceanic waters (Chiba and Hirakawa, 1972). The classic taxonomic works on this and the above group have been done by Tesch (1946, 1949, 1950). ARTHROPODA Crustacea Ostracoda Planktonic ostracods have been recorded from Enewetak Lagoon (Gerber and Marshall, 1974; Hobson and Chess, 1978; Gerber, 1981) and Majuro Lagoon (Hob- son and Chess, 1973). Those recorded by Gerber (1981) and most of those reported by Hobson and Chess (1973) belong to the order Myodocopa, but further identification was not made. Though they occur fairly regularly in net collections, ostracods are represented by only a few indi- viduals per sample. Copepoda Approximately 58 species of calanoid copepods have been identified from Enewetak Lagoon. Johnson (1954) identified the 12 most common species from Enewetak, Rongelap, Rongerik, and Bikini Lagoons but unfortunately did not indicate which species came from each lagoon. His collections at Enewetak, Rongerik, and Rongelap Lagoons were made in June and at Bikini Lagoon from March to May in 1946. Twelve species of calanoids were identified by Gilmartin (1958) at Enewetak Lagoon from samples col- lected in April, November, and December 1955; eight of these species were not recorded by Johnson (1954). Two plankton samples were collected at Enewetak Lagoon in September 1959 and 1961 by Mahnken (1966) in conjunc- tion with a detailed zooplankton study of Rongelap Lagoon. He identified 25 species from Enewetak Lagoon and 45 species from Rongelap Lagoon which included most of the species previously reported from Enewetak. In January and February 1966, Barnett (1967) studied the distribution of copepods at several locations in Enewetak Lagoon. He recorded about 46 species of calanoid copepods, most of which also occurred in Rongelap Lagoon. Gerber (1981) sampled Enewetak Lagoon during January and February 1972 and 1974, respectively, and in June through August 1974. He identified 45 species from a mid-lagoon station; most of those species had been previ- ously recorded from this lagoon. Even after these studies at Enewetak, there are species of calanoid copepods which have been recorded from Rongelap that have not been identified from Enewetak Lagoon. Those species are Euchirella sp., Labidocera bataviae Scott, Pontella fera, Pontella tenuiremis, Pontellina globosa, Pontellopsis kram- eri, Pontellopsis macronyx, and Undeuchaeta plumosa. At Majuro Atoll Lagoon, two species of calanoid copepods (Candacia discaudata and Undinula vulgaris) which also occur at Enewetak Lagoon were recorded from the stomach contents of fish; a third species collected there, Labidocera acuta, has not been recorded from the other lagoons (Hobson and Chess, 1973). In a study of the feed- ing habits of various planktivorous fishes at Enewetak, Hobson and Chess (1978) identified three of the large size copepods to species and four others to the generic level; these had been reported previously. Calanoid copepods in reduced abundance were recorded by Johnson (1954) from outside Bikini Lagoon, but these species were not reported. Chiba et al. (1955) collected zooplankton in the open ocean near Bikini Atoll and throughout the Marshall Islands from May to June 1954. They presented 80 species of calanoid copepods, about half of which have been recorded from Enewetak Lagoon; the remaining species consisted of oceanic and deep water forms. There are many important works dealing with the taxonomy and identification of calanoid copepods which occur at Enewetak Lagoon. The most important works are Bradford (1974), Dakin and Colefax (1940), Delsman (1939), Frost and Fleminger (1968), Giesbrecht (1889), Grice (1961), Mori (1937), Scott (1909), Sewell (1929), Vervoort (1964), and Wilson (1942). Cyclopoid copepods were first recorded from Enewetak and nearby atolls by Johnson (1954) during the sampling periods mentioned above. He recorded four species plus two genera consisting of several unidentified species. Gilmartin (1958) identified four species at Enewetak Lagoon, and Mahnken (1966) identified 12 species from Enewetak and five at Bikini and Rongelap Lagoons. These LAGOON PLANKTON 213 researchers used rather coarse mesh nets which did not adequately sample this group consisting mostly of smaller forms; this perhaps accounts for the few species recorded. About 27 species were recorded at Enewetak Lagoon by Barnett (1967) and included those species identified there previously; Gerber (1981) identified 28 species which included most of Barnett’s species and several additional species. Only one cyclopoid copepod, Corycaeus curtus, recorded at Rongelap has not been found at Enewetak Lagoon. In the surrounding waters of Bikini Atoll, 29 species were found by Chiba et al. (1955), but only 13 of these occur within the lagoon at Enewetak. Four cyclopoid genera were recorded at Majuro Atoll Lagoon from the stomach contents of plankton feeding fishes (Hobson and Chess, 1973), all of which have been recorded from Enewetak and include Oncaea sp., Corycaeus sp., Sapphirina sp., and Saphirella sp. For identification and taxonomy of cyclopoid copepods at Enewetak, the follow- ing works should be consulted: Dahl (1912), Dakin and Colefax (1940), Farran (1911), Giesbrecht (1889), Mori (1937), Motoda (1963), Scott (1909), and Wilson (1942). Few harpacticoid copepods are strictly holoplanktonic, and of these, five species have been found in the Enewetak Lagoon. They include members of the first three families presented in the checklist under this order. Gilmar- tin (1958) identified one of these species, Barnett (1967) identified three species, and Gerber (1981) found four of these species. At a shallow station (2 m depth) behind a windward reef area of the lagoon, Gerber (1981) collected in net tows 16 species of harpacticoids that are believed to be epibenthic types. These were restricted to the shallow station and may have been inadvertently washed from adjacent coral reefs. Harpacticoid copepods were also in the lagoon plankton and in the gut contents of various fishes (e.g., Gerber and Marshall, 1974; Hobson and Chess, 1978), but these were not identified. A thorough study of the epibenthic harpacticoid copepods has not been made at any location in the Marshall Islands. The five pelagic species noted above were not identified from net tow samples in the neighboring waters of Bikini Atoll (Chiba et al., 1955). This suggests that they are probably neritic types, surviving best in shallower protected areas such as atoll lagoons. Both Barnett (1967) and Gerber (1981) identified members of a single genus belonging to the other Monstril- loida. Whether the same species were represented in both studies is not known, because no identifications were made to the species level. Monstrilloid copepods occurred only rarely in net collections in both studies, and no reference indicates that they occur outside of atoll lagoons in the Marshall Islands. Information on the identification and tax- onomy of harpacticoid and monstrilloid copepods from Enewetak are scarce, but the following works are helpful: Dakin and Colefax (1940), Lang (1948), Scott (1909), and Vervoort (1964). Mysidacea A detailed investigation of the mysids at Enewetak Atoll has not been carried out. A planktonic mysid was collected in net tows by Gilmartin (1958) at Enewetak Lagoon, but the specimen was not identified. Swarms of mysids belonging to two genera (Siriella and Mysinae) were observed and collected over lagoon patch reefs at Enewetak by Hobson and Chess (1978); Gerber (1981) identified eight species of mysids from net tows at a nearby shallow station; and Murano (1983) identified seven species from net tows and emergence traps at depths between 2 and 10 m. Further samplings, especially at night, would no doubt reveal additional specimens. Uniden- tified mysids were recorded in the plankton and in the gut contents of plankton feeding fishes at nearby Majuro Atoll Lagoon by Hobson and Chess (1973). Plankton tows in the oceanic waters near Bikini Atoll by Chiba et al. (1955) revealed two species of mysids, Siriella thompsoni and Siriella sp., and Tattersall (1951) described two additional species of Siriella from the Marshall Islands (S. vulgaris ros- trata, S. anomala) and Anchialina typica and A. penicillata. As noted earlier, the genus Siriella was first recorded inside the lagoon at Enewetak by Hobson and Chess (1978), and later two other lagoon Siriella were identified to species by Murano (1983). Several excellent works by Hansen (1910) and Tattersall (1957) have proved useful in identifying the mysids from Enewetak. Isopoda The isopods which have been collected in plankton net hauls from Enewetak Lagoon have not been identified to genera (e.g., Gilmartin, 1958; Hobson and Chess, 1978; and Gerber, 1981). This group is generally scarce in such samplings indicating that they are most likely of epibenthic origin rather than planktonic. Planktonic isopods from other atoll lagoons or in the oceanic waters in the Marshall Islands have not been collected. Amphipoda At least three and possibly four species of holoplank- tonic amphipods that belong to the suborder Hyperiidea have been reported from the Enewetak Lagoon (Gerber, 1981). Earlier plankton collections by Gilmartin (1958) revealed one unidentified species, and Hobson and Chess (1978) recorded but did not identify several species of hyperiids. Many more nonplanktonic species occur within the lagoon (Barnard, 1965), and these are mostly gam- marids which occur in the epibenthic shallow water habitat discussed elsewhere in this book. Barnard (1965) reported one gammarid species, Synopia variabilis, which was col- lected from Enewetak and Ifaluk Lagoons by both plankton haul and dip-netting with a night light; it is included in the checklist. Hyperiid amphipods have been recorded at Majuro Atoll (Hobson and Chess, 1973) where one species, Lestrigonus bengalensis, was identified from net tows and made up an important dietary component of antherinid fish. In the oceanic waters around the Marshall Islands, Chiba et al. (1955) collected nine species of planktonic amphipods, none of which appear to occur in Enewetak Lagoon. These are Oxycephalus porcellus, Phronima_ pacifica, Phronima sp., Leptocotis tenuirostris, 214 GERBER Parascelus edwardsi, Phronimopsis spinifera, Hyperioides sibaginis, Hyperia sp., and Anchylomera blosseuillii. For the taxonomy and identification of the amphipods from Enewetak, see Barnard (1965), Bowman and Gruner (1973), and Dakin and Colefax (1940). Euphausiacea A single species of euphausiid has been reported from Enewetak Lagoon where it occurs regularly at both shallow and deep stations (Gerber, 1981). This species was also the dominant euphausiid recorded inside the Great Barrier Reef Lagoon (Russell, 1934). Gilmartin (1958) reported euphausiids in his plankton hauls at Enewetak Lagoon but did not identify them. In the surrounding oceanic waters of Enewetak and Bikini Atolls, seven species of typically oceanic euphausiids were reported by Chiba et al. (1955), and none of these occurs in the atoll lagoons. They are Stylocheiron carinatum, S. submii, Euphausia gracilis, E. gibba, . krohnii, Thysanopoda obtusifrons, and T. tricuspidata. Useful works concerning the identification and taxonomy of euphausiids include Dakin and Colefax (1940), Delsman (1939), Hansen (1910), Mauchline and Fisher (1969), and Mauchline (1980). CHAETOGNATHA At least nine species of chaetognaths have been reported and identified from Enewetak Lagoon, in addition to seven species from outside the lagoon or in the nearby ocean waters. They are an important planktonic group and appear in most net samplings. Johnson (1949, 1954) reported three species from net tows taken during 1946 at Enewetak Lagoon (June), and the lagoons at Rongelap (June), Rongerik (June), and Bikini (March to May) and from outside Bikini Atoll (March to May). Distribution of chaetognaths within Bikini Atoll was also studied. A translagoon study of plankton at Enewetak from April through December 1955 by Gilmartin (1958) revealed four species, most of which were abundant in the net samples. Gerber (1981) reported at least six species from a mid- lagoon station in the winter and summer (1972, 1974) with increased abundances in summer. Collections made in 1946 and later revealed six species from the lagoon and several more from outside, east of the lagoon (Alvarifo, personal communication). In the neighboring waters around Bikini Atoll, extensive plankton collections were made in May and June 1954 by Chiba et al. (1955) and later by Tsuruta (1963). These researchers identified at least 16 species; the eight not found within the lagoons include Sagitta decipiens, S. hexaptera, S. lyra, S. minima, S. pul- chre, S. pseudoserratodentata, Krohnitta pacifica, and K. subtilis. Three species of chaetognaths were reported from the gut contents of lagoon fish at Majuro Atoll (Hob- son and Chess, 1973) and at Enewetak Atoll (Gerber and Marshall, 1974). Important sources for the identification of chaetognaths include Alvarifo, (1967), Sund (1959), and Thomson (1947). CHORDATA Thaliacea This group is much less abundant than the Larvacea in the Enewetak Lagoon and from nearby atolls. They were first reported from Enewetak by Gilmartin (1958) who identified two species from April through December in his translagoon plankton study of 1955. An additional species has been collected from the lagoon in 12 m of water and identified by R. A. Kinzie (MPML reference collection). Appendicularia (Larvacea) This is an abundant zooplankton group in the atoll lagoons and surrounding waters of the Marshall Islands. At least seven species have been reported from Enewetak Lagoon, five of which have been collected from outside the atoll. Johnson (1954) reported densities of appendiculari- ans which were considerably higher for Enewetak and nearby atoll lagoons than in the nearby open ocean; he did not identify genera or species. At Enewetak Lagoon, Gil- martin (1958) also reported high densities of this group especially during April and in early November and identi- fied them as Oikopleura sp. Gerber similarly (1981) found high densities of larvaceans in the lagoon especially in the summer; at least six species are recorded belonging to two families. In the waters near Bikini Atoll, Chiba et al. (1955) identified 16 species, five of which are common to Enewetak Lagoon. These additional species include Fritillaria borealis forma sargassi, F. formica, F. hap- lostoma, F. pellucida, F. venusta, Oikopleura cophocerca, Oikopleura sp., Kowaleuskaia tenuis, Megalocercus huxleyi, Stegosoma magnum, and Pelagopleura sp. Appendicularia have been reported at Majuro Atoll Lagoon from the gut contents of plankton feeding fishes (Hobson and Chess, 1973); the species were not identified. The identification and taxonomy of the Thaliacea and Larvacea are included in the works of Bjornberg and Forneris (1955), Delsman (1939), and Fraser (1947). CONCLUSIONS There appears to be little difference of the holoplank- tonic zooplankton species composition between Enewetak and nearby atoll lagoons such as Rongelap. This observa- tion is largely based on the copepod data which have been most extensively studied at these two lagoons. Clearly, this faunal similarity reflects the rather similar habitats of the two lagoons and their similar locations with respect to the prevailing ocean currents. The information available comparing the copepod species assemblege inside the lagoon with that of the sur- rounding ocean outside the lagoon demonstrates that they are different. The lagoon zooplankton populations are par- tially composed of oceanic species known to occur in the neighboring waters outside the atoll, in addition to neritic species which occur only in shallow protected environ- ments. It has also been found that the density of zooplank- LAGOON PLANKTON 215 ton in the lagoons is considerably higher than in the sur- rounding ocean waters (Chap. 10, Volume 1). Thus, the zooplankton of lagoons in the Marshall Islands is composed of similar species, and we would expect only slight varia- tions in community structure from lagoon to lagoon reflect- ing small habitat and geographic differences. Included in this lagoon zooplankton, and yet ignored thus far, is the meroplanktonic component comprised mostly of the larvae of the many reef invertebrates and fishes as well as some adventurous benthic forms which occasionally move into the water column. Various researchers (e.g., Gerber and Marshall, 1974; Hobson and Chess, 1978) have shown that the meroplankton some- times dominates the zooplankton and becomes an impor- tant food of reef and lagoon fishes. ACKNOWLEDGMENTS I thank Dennis M. Devaney and Ernst S. Reese for inviting me to participate in this book. Thanks is also due to the Mid-Pacific Marine Laboratory whose facilities I used during my zooplankton studies. For use of some chaeto- gnath and siphonophore data, I thank Angeles Alvarino; and I thank H. Hirchfield and L. Helson, for use of their protozoan data. I also thank Thomas Bowman for informa- tion he provided and Bowman and Beatrice Burch for their helpful editorial comments. Anne Barrington, Pamela Labbe, and Gloria Chamard have done a fine job typing the drafts and final copies of this chapter—many thanks. REFERENCES Alvarifio, A., 1967, The Chaetognatha of the NAGA Expedition (1959-1961) in the South China Sea and Gulf of Thailand, Part I-Systematics, University of California, Scripps Inst. Oceanogr., La Jolla, California, NAGA Report, 4: 1-186. Barnard, J. L., 1965, Marine Amphipoda of Atolls in Micronesia, Proc. U. S. Natl. Mus., 117 (3516): 459-552. Barnett, A. M., 1967, Distributions of Copepoda in Enewetak Lagoon, Marshall Islands, M. S. thesis, University of Washing- ton. Bjornberg, T. K. S., and L. Forneris, 1955, Resulatados Cientifi- cos do Cruzeiro do “Baependi” e do “Vega” a’llha de Trin- dade, Copelata I, Contrib. Avulsas Inst. Oceanogr. Sao Paulo, 1: 1-68. Bowman, T. E., 1967, The Planktonic Shrimp Lucifer chacei, sp. nov., Sergestidae: Luciferinae; the Pacific Twin of the Atlan- tic, Lucifer faxoni, Pac. Sci., 21: 266-271. —, 1973, Pelagic Amphipods of the Genus Hyperia and Closely Related Genera (Hyperidea: Hyperiidae), Smithsonian Contrib. Zool., 136: 1-76. —_, and H. E. Gruner, 1973, The Families and Genera of Hyperiidea (Crustacea: Amphipoda), Smithson. Contrib. Zool., 146: 1-64. Bradford, J. M., 1974, Euchaeta marina (Prestandrea) (Copepoda, Calanoida) and Two Closely Related New Species from the Pacific, Pac. Sci., 28: 159-169. Campbell, A. S., and R. C. Moore, 1954, Protista 3 (Chiefly Radiolarians and Tintinnines), Treatise on Invertebrate Paleon- tology, Part D, University of Kansas Press, Lawrence. Chiba, T., and K. Hirakawa, 1972, Classification and Communities of the Zooplankton in the South-Western Pacific Ocean, with Special Reference to Copepods, J. Shimonoseki Coll. Fish., 21: 67-80. —_, A. Tsuruta, and H. Maeda, 1955, Report on Zooplankton Samples Hauled by Larva-Net During the Cruise of Bikini- Expedition, with Special Reference to Copepods, J. Shi- monoseki Coll. Fish., 5: 189-213. Corliss, J. O., 1977, Annotated Assignment of Families and Gen- era to the Orders and Classes Currently Comprising the Corlissian Scheme of Higher Classification for the Phylum Ciliophora, Trans. Am. Microsc. Soc., 96: 104-140. Dahl, M., 1912, Die Copepoden der Plankton-Expedition, 1, Corycaeinen mit Berusksichtigung aller Bekannten Arten, Ergebn. d. Plankton-Exped. d. Humboldt-Stift, Bd. 2, G. F., 1-G, 1-134, Taf. 1-15. Dakin, W. J., and A. N. Colefax, 1940, The Plankton of the Aus- tralian Coastal Waters off New South Wales, Part. I, Publ. Univ. Sydney, Dept. Zool. Monograph, 1: 1-215. Delsman, H. C., 1939, Preliminary Plankton Investigations in the Java Sea, Trebia, 17: 139-181. Farran, G. P., 1911, Plankton from Christmas Island, Indian Ocean, I, On Copepoda of the Family Corycaedae, Zool. Soc. London, 1913: 285-296. Fraser, J. H., 1947, Thaliacea I, Il, Family Salpidae and Family Doliolidae, Fiches d’identification du Zooplancton 9: 1-4, 10: 1-4, Conseil International pour I'Exploration de la Mer. Frost, B., and A. Fleminger, 1968, Revision of the Genus Clausocalanus, Bull. Scripps Inst. Oceanogr., Vol. 12. Gerber, R. P., 1981, Species Composition and Abundance of Lagoon Zooplankton at Eniwetak Atoll, Marshall Islands, Atoll Res. Bull., 247: 1-22. ___, and N. Marshall, 1974, Ingestion of Detritus by the Lagoon Pelagic Community at Enewetak Atoll, Limnol. Oceanogr., 19: 815-824. 7 __, and N. Marshall, 1982, Characterization of the Natural Suspended Particulate Organic Matter and Feeding of Lagoon Zooplankton at Enewetak Atoll, Marshall Islands, Bull. Mar. Sci. 32: 290-300. Giesbrecht, W., 1889, Elenco dei Copepodi Pelagici Raccolti dal Tenente di Vascello Gaetano Chierchia Durante il Viaggio della R. Corvetta Vettor Pisani Negli Anni 1882-1885, e dal Tenenete di Vascello Francesco Orsini nel Mar Rosso, nel 1884, Atti della Reale Accademia Nazionale dei Lincei, Rendi- conti Series 42): 284-287, 330-338; 5{1): 811-815; 5(2) 24-29. Gilmartin, M., 1958, Some Observations on the Lagoon Plankton of Enewetak Atoll, Pac. Sci., 12: 313-316. Gold, K., and E. A. Morales, 1977, Studies on the Tintinnida of Enewetak Atoll, J. Protozool., 24: 580-587. Grice, G. D., 1961, Calanoid Copepods from the Equatorial Waters of the Pacific Ocean, United States Department of the Interior, Fish and Wildlife Service, Fish. Bull., 61: 171-246. Hansen, H. J., 1910, The Schizopoda of the Siboga Expedition, Siboga-Expeditie Vol. 37. Hirshfield, H. I., L. Helson, and R. Charmatz, 1957, Enewetak - Protozoa., Rept. to EMBL. Hobson, E. S., and J. R. Chess, 1973, Feeding Oriented Move- ments of the Atherinid Fish Pranesus pinguis at Majuro Atoll, Marshall Islands, Fish. Bull., 71: 777-786. Hobson, E. S., and J. R. Chess, 1978, Trophic Relationships Among Fishes and Plankton in the Lagoon at Enewetak Atoll, Marshall Islands, Fish. Bull., 76: 133-153. 216 GERBER Johannes, R. E., and R. P. Gerber, 1974, Import and Export of Net Plankton by an Enewetak Coral Reef Community, in Proceedings of the Second International Coral Reef Sympo- sium, Australia, 1973: pp. 97-104. —__, S. L. Coles, and N. T. Kuenzel, 1970, The Role of Zoo- plankton in the Nutrition of Some Scleractinian Corals, Limnol. Oceanogr., 15: 579-586. Johnson, M. W., 1949, Zooplankton as an Index of Water Exchange Between Bikini Lagoon and the Open Sea, Trans. Am. Geophys. Union, 30: 238-244. ——, 1954, Plankton of the Northern Marshall Islands, Bikini and Nearby Atolls, U. S. Geol. Surv. Professional Paper 260-F, 301-314. Kramer, A., 1897, Ueber den Bau der Korallenriffe und die Planktonvertheibeng an den Samoanischen Kusten nebst ver- gleichenden Bemerkungen, Kiel and Leipzig. Kramp, P. L., 1961, Synopsis of the Medusae of the World, J. Mar. Biol. Assoc. U. K., 40: 7-469. Kofoid, C. A., and A. S. Campbell, 1939, Reports on the Scien- tific Results of the Expedition to the Eastern Tropical Pacific, in Charge of A. Agassiz, by the U. S. Fish. Comm. Steamer Albatross from Oct. 1904-Mar. 1905, The Ciliata: Tintinnoinea, Bull. Mus. Comp. Zool., Harvard University, 84: 1-473. Lang, K., 1948, Monographie der Harpacticiden, H. Ohlssons, Lund, Sweden. Lebour, M., 1925, Dinoflagellates of the Northern Seas, Mar. Biol. Assoc. U. K., Plymouth. Mahnken, C., 1966, Distributions of Copepoda at Rongelap Atoll, Marshall Islands, M. S. thesis, University of Washington. Mauchline, J., 1980, The Biology of Mysids, Euphausiids, Adv. Mar. Biol., 18: 1-681. —, and L. R. Fisher, 1969, The Biology of Euphausiids, Adv. Mar. Biol., 7: 1-454. Mayer, A. G., 1910, Medusae of the World, Hydromedusae and Scyphomedusae, Carnegie Inst. Wash., Publ. 109, 3 vols. Michel, A., 1969, Plancton des Lagoon et des Abords Exterieurs de L’Atoll de Mururoa, Cah. Pac., 13: 81-132. __, C. Colin, R. Desrosieres, and C. Oudot, 1971, Observations sur L’Hydrologie et le Plancton des Abords et de la Zone des Passes de L’Atoll de Rangiroa (Archipel des Tuamotu, Ocean Pacifique Central), Cah. ORSTOM, Ser. Oceanogr., 9: 375-402. Mori, T., 1937, The Pelagic Copepoda from the Neighboring Waters of Japan, Soyo Co. Inc., Tokyo, 1964. Motoda, S., 1938, Quantitative Studies on the Macroplankton off Coral Reefs of Palao Port, Trans. Sapporo Nat. Soc., 15: 242-246. ——, 1963, Corycaeus and Farranula (Copepoda, Cyclopoida) in Hawaiian Waters, Seto Mar. Biol. Lab., 11: 209-262. Murano, M., 1983, Mysidacea Fauna from Enewetak Lagoon, Micronesia, Bull. Plankton Soc. Jpn., 30: 81-90. Odum, H. T., and E. P. Odum, 1955, Trophic Structure and Pro- ductivity of a Windward Coral Reef Community on Enewetak Atoll, Ecol. Monogr., 25: 291-320. Parke, M., and P. S. Dixon, 1976, Checklist of British Marine Algae—Third Revision, J. Mar. Biol. Assoc. U. K., 56: 527-594. Russell, F. S., 1934, The Zooplankton, III, A Comparison of the Abundance of Zooplankton in the Great Barrier Reef Lagoon with that of Some Regions in Northern European Waters, Great Barrier Reef Expedition, 1928-1929, Sci. Rept., 2: 159-201. Schiller, J., 1937, Dr. L. Rabenhorst’s Kryptogamen-Flora von Deutschland, Osterreich und der Schweiz, Flagellate, Dinoflagettae, Zehnter Band, Dritte Abteilung. Scott, A., 1909, The Copepoda of the Siboga Expedition, Part I., Freeswimming, Littoral, and Semiparasitic Copepoda, Siboga Exp. Monogr., 29: 5-320. Sears, M., 1950, Notes on Siphonophores, I, Siphonophores from the Marshall Islands, J. Mar. Res., 9: 1-16. Sewell, R. B., 1929, The Copepoda of Indian Seas, Calanoidae, Mem. Indian Mus., 10: 1-221. Sund, P. N., 1959, A Key to the Chaetognatha of the Tropical Eastern Pacific Ocean, Pac. Sci., 13: 269-285. Tattersall, O. S., 1957, Mysidacea, Discovery Rep., 28: 3-189. Tattersall, W. M., 1951, A Review of the Mysidacea of the United States National Museum, Bull. U. S. Natl. Mus., 201: 1-291. Tesch, J. J., 1946, The Thecosomatous Pterpods, I, The Atlan- tic, Dana Rep., 28: 1-82. __, 1949, Heteropoda, Dana Rep., 34: 1-54. , 1950, The Gymnosomata, II, Dana Rep., 36: 1-55. Thomson, J. M., 1947, The Chaetognatha of South-Eastern Aus- tralia, Commonwealth of Australia, Council for Scientific and Industrial Research (Div. of Fisheries-Rep. No. 14), Bull. No. 222. Totton, A. K., and H. E. Bargmann, 1965, A Synopsis of the Siphonophora, Br. Mus. (Nat. Hist.), London. Tranter, D. J., and J. George, 1972, Zooplankton Abundance at Kavaratti and Kalpeni Atolls in the Laccadives, Symp. Corals and Coral Reefs, Proc. 1969 (Mar. Biol. Assoc. India): pp. 239-256. Tsuruta, A., 1963, Distribution of Plankton and Its Characteris- tics in the Oceanic Fishing Ground, with Special Reference to Their Relation to Fishing, J. Shimonoseki Coll. Fish., 12: 13-214. Vervoort, W., 1964, Free-Living Copepoda from Ifaluk Atoll in the Caroline Islands. U. S. Nat. Mus. Bull., 236: 1-431. Wilson, C. B., 1942, The Copepods of the Plankton Gathered During the Last Cruise of the Carnegie, Biology-l, Carnegie Inst. Wash. Pub., 536: 1-237. Chapter 21 Ostracoda (Myodocopina) of Enewetak Atoll LOUIS S. KORNICKER Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution Washington, D. C. 20560 INTRODUCTION Ostracods in the superorder Myodocopa, excluding the planktonic suborder Halocypridina, which is outside the scope of the present report, have not previously been reported from Micronesia. A comprehensive checklist of Ostracoda from southeast Asia (Hanai et al., 1980) is con- cerned with the area between 25°N and 11°S latitude and 140°E and 85°E longitude. The area includes the western edge of Micronesia, but a brief perusal of the checklist revealed no listing of specimens from Micronesia. A small collection of Enewetak ostracods in the suborder Myodocopina at the National Museum of Natural History, Smithsonian Institution, contains members of all five families that comprise the suborder (Table 1). Although the species in the collection are capable of swim- ming, and do so occasionally, they spend most of their existence crawling on, or burrowing in, the substrate. An exception to this is members of the mainly pelagic genus Cypridina, which is represented in the collection by two species. Some of the Enewetak specimens are illustrated in Fig. 1. The feeding habits of the families differ: the Cylindro- leberididae is composed of filter feeders, the Sarsiellidae and Rutidermatidae of carnivores, the Philomedidae of detritus feeders, and the Cypridinidae of scavengers, detritus feeders, and also carnivores (Kornicker, 1975). BIOGEOGRAPHIC COMPARISONS AT GENERIC LEVEL The genera Skogsbergia, Harbansus, Rutiderma, Sarsiella, Parasterope, and Cylindroleberis are worldwide in distribution; only Parasterope is present in Antarctic waters (Kornicker, 1975); and none has been reported from the Arctic Ocean. 217 TABLE 1 Checklist of Enewetak Atoll Ostracoda (Myodocopina) Phylum ARTHROPODA Subphylum CRUSTACEA Class OSTRACODA Superorder MYODOCOPA Order MYODOCOPIDA Suborder MYODOCOPINA Superfamily CYPRIDINOIDEA Family CYPRIDINIDAE *+Cypridina spp. *+Paravargula sp. *+Skogsbergia sp. Family PHILOMEDIDAE *+Harbansus sp. Family RUTIDERMATIDAE *+Rutiderma sp. Family SARSIELLIDAE *+Ancohenia sp. *+Anscottiella sp. *+Sarsiella sp. Family CYLINDROLEBERIDIDAE *+Parasterope sp. *+Cylindroleberis sp. * New Micronesian record. + New Enewetak record. The pelagic genus Cypridina is primarily Indo-Pacific. Ancohenia is previously known from Maunalua Bay, Oahu Island, Hawaii (Kornicker, 1976:7) and Vuna Point, Taveuni, Fiji (Kornicker, 1981:12). Anscottiella has previously been collected off Ceylon and Thailand (Poulsen, 1965:65; Kornicker, 1975:607; Kornicker and McKenzie, 1976:348). Paravargula has previously been collected off the tip of South Africa and from the East-Indian region (Poulsen, 1962:204; Kornicker, 1975:3). 218 KORNICKER Fig. 1 a, Harbansus species, female, length 0.87 mm, USNM 158308; b, Rutiderma species, female, length 0.93 mm, USNM 158309; c, Anscottiella species, male, length 0.70 mm, USNM 158313; d, Anscottiella species, female, length 0.80 mm, USNM 158317; e, Sarsiella species, female, length 0.78 mm, USNM 158310; f, Ancohenia species, male, length 1.01 mm, USNM 158312. OSTRACODA (MYDOCOPINA) 219 Station Data Specimens of Cypridina and Anscottiella were collected by C. Allan Child, National Museum of Natural History, Smithsonian Institution, in October 1969, by scraping the piling of a marine pier at the north end of the lagoon side of Enewetak Island in 4% to 2 m of water at low tide. The remaining specimens were collected by Edmund S. Hobson and James R. Chess, Southwest Fisheries Center, Tiburon Laboratory, National Marine Fisheries Service, in May 1979, from day and night airlift and emergence traps placed in the lagoon in the lee of Bokandrock Island in about 5 m of water and in the lee of a high section of interisland reef midway between Enewetak and Medren Islands in about 8 m of water. These samples were obtained from substrates of sand, coral, and coral rubble. Both species collected by C. Allan Child from the marine pier were also present in the samples of E. S. Hobson and J. R. Chess. ACKNOWLEDGMENTS I thank C. Allan Child, Edmund S. Hobson, and James R. Chess for collecting the specimens upon which this report is based. I also thank Raymond B. Manning and Anne C. Cohen for reviewing the manuscript and Carolyn Gast for rendering the shaded drawings of the carapaces. REFERENCES Hanai, T., N. Ikeya, and M. Yajima, 1980, Checklist of Ostracoda from Southeast Asia, Univ. Mus., Univ. Tokyo, Bull. 17. Kornicker, L. S., 1970, Myodocopid Ostracoda (Cypridinacea) from the Philippine Islands, Smithsonian Contrib. Zool., 39: 1-32. ——, 1975, Antarctic Ostracoda (Myodocopina), Smithsonian Contrib. Zool., 163: 1-720. ——, 1976, Benthic Marine Cypridinacea from Hawaii (Ostracoda), Smithsonian Contrib. Zool., 231: 1-24. —_, 1981, A Restudy of the Ostracode Genus Pleoschisma Brady, 1890 (Myodocopina), Smithsonian Contrib. Zool., 332: 1-16. , and K. G. McKenzie, 1976, Redescription of Eurypylus petrosus Brady 1869, and a Key to the Genera of Sarsiellidae (Myodocopina: Ostracoda), Proc. Biol. Soc. Washington, 89: 347-352. Poulsen, E. M., 1962, Ostracoda-Myodocopa, 1: Cypridinidae, Dana Report 57, 414 pp., Copenhagen: Carlsberg Founda- tion. —, 1965, Ostracoda-Myodocopa, 2: Cypridiniformes—Ruti- dermatidae, Sarsiellidae and Asteropidae, Dana Report 65, 484 pp., Copenhagen: Carlsberg Foundation. Chapter 22 Crustacea Decapoda (Penaeidea, Stenopodidea, Caridea, and Palinura) of Enewetak Atoll DENNIS M. DEVANEY’ (deceased) and A. J. BRUCET “Bernice P. Bishop Museum, Honolulu, Hawaii 96817; tNorthern Territory Museum of Arts and Sciences, GPO Box 4646, Darwin, N. T. 5794, Australia INTRODUCTION Biological surveys made during 1946 and 1947 in the northern Marshall Islands included faunal collections that resulted in a publication by Chace (1955) on several groups of shrimps. In this work, six species of sergestid, pasiphaeid, processid, and alpheid shrimps were recorded from Enewetak. Since then the number of taxa from the atoll has increased until now nearly 150 decapod shrimp and lobster species are recognized. Pontoniines and alpheids account for 73% (106 species) of the total number. There are several new records not only for Enewetak but also for the Marshalls, and in some cases, for the Pacific. A taxonomic checklist (Table 1) and a sec- tion on collection data for new records are provided. PENAEIDEA Two penaeid species in the genus Metapenaeopsis, one a widely distributed Indo-West Pacific form and the other as yet unidentified, are now recorded for the first time from Enewetak. Among the sicyonid shrimps, the widely distributed Indo-West Pacific species, Sicyonella mal- divensis, has been collected at Enewetak on the surface using a light at night. One planktonic sergestid, an undeter- mined species of Lucifer, was reported by Chace (1955) and was collected at Enewetak in the same manner. In 1967, Bowman recorded L. chacei as a new species from a number of Indo-West Pacific locations including Enewetak and Rongelap Atolls in the Marshalls. TABLE 1 Checklist of Enewetak Penaeidea, Stenopodidea, Caridea, and Palinura*® Subphylum CRUSTACEA Class MALACOSTRACA Order DECAPODA Suborder DENDROBRANCHIATA Infraorder PENAEIDEA Superfamily PENAEOIDEA Family PENAEIDAE T Metapenaeopsis borradailei (De Man, 1910). + Metapenaeopsis sp. Family SICYONIIDAE + Sicyonella maldivensis Borradaile, 1910. Superfamily SERGESTOIDEA Family SERGESTIDAE Lucifer chacei Bowman, 1967: Bowman, 1967. Lucifer sp.: Chace, 1955. “The higher classification used here follows that of Bowman and Abele (1982). +New records for Enewetak. (This table continued on next page.) 222 DEVANEY AND BRUCE TABLE 1 (cont’d) a ee ee eee Suborder PLEOCYEMATA Infraorder STENOPODIDEA Family STENOPODIDAE tT Microprosthema plumicorne (Richters, 1880). + Microprosthema scabricaudatum (Richters, 1880). T Odontozona sculpticaudata Holthuis, 1947. + Odontozona sp. nov. + Stenopus hispidus (Olivier, 1811). + Stenopus tenuirostris De Man, 1888. + Stenopus zanzibaricus Bruce, 1976. Infraorder CARIDEA Superfamily PASIPHAEOIDEA Family PASIPHAEIDAE Leptochela irrobusta Chace, 1976: Chace, 1976. Leptochela robusta Stimpson, 1860: Chace, 1955. Superfamily RHYNCHOCINETOIDEA Family RHYNCHOCINETIDAE Rhynchocinetes hiatti Holthuis and Hayashi, 1967: Holthuis and Hayashi, 1967. Rhynchocinetes hendersoni Kemp, 1925: Holthuis and Hayashi, 1967. Rhynchocinetes marshallensis Edmondson, 1952: Edmondson, 1952. T Rhynchocinetes aff. R. rigens Gordon, 1936. + Rhynchocinetes sp. Superfamily PALAEMONOIDEA Family PALAEMONIDAE Subfamily PALAEMONINAE + Brachycarpus biunguiculatus (Lucas, 1846). } Leandrites cyrtorhynchus Fujino and Miyake, 1969. Subfamily PONTONIINAE t Anchistus australis Bruce, 1977. Anchistus miersi (De Man, 1888): Fankboner, 1972. Tt Araiopontonia odontorhyncha Fujino and Miyake, 1970. + Conchodytes biunguiculatus (Paulson, 1875). t Conchodytes meleagrinae Peters, 1852. + Conchodytes tridacnae Peters, 1852. Tt Coralliocaris brevirostris Borradaile, 1898. t Coralliocaris graminea (Dana, 1852). + Coralliocaris nudirostris (Heller, 1861). T Coralliocaris superba (Dana, 1852). Tt Coralliocaris venusta Kemp, 1922. Ctenopontonia cyphastreophila Bruce, 1979: Bruce, 1979b. + Fennera chacei Holthuis, 1951. Tt Harpiliopsis beaupresii (Audouin, 1826). Tt Harpiliopsis depressa (Stimpson, 1860). Tt Harpiliopsis spinigera (Ortmann, 1890). t Jocaste japonica (Ortmann, 1890). Tt Jocaste lucina (Nobili, 1901). t Onycocaris quadratophthalma (Balss, 1921). Palaemonella pottsi (Borradaile, 1915): Bruce, 1970. Palaemonella rotumana (Borradaile, 1898): Bruce, 1970. + Palaemonella tenuipes Dana, 1852. t Palaemonella sp. Paranchistus armatus (H. Milne Edwards, 1837): Bruce, 1975; 1979a. Paranchistus biunguiculatus (Borradaile, 1898): Rosewater, 1965. Paratypton siebenrocki Balss, 1914: Bruce, 1969. t Periclimenaeus sp. t Periclimenes agag Kemp, 1922. t+#Periclimenes (?)amboinensis (De Man, 1888). t Periclimenes bayeri Holthuis, 1981. Periclimenes brevicarpalis (Schenkel, 1902): Bruce, 1979a. +New records for Enewetak. +The status of P. amboinensis (De Man) is not clear. It may be synonymous with P. cornutus Borradaile. (This table continued on next page.) CRUSTACEA DECAPODA 223 TABLE 1 (cont'd) Subfamily PONTONIINAE (cont'd) T Periclimenes commensalis Borradaile, 1915. t+ Periclimenes cristimanus Bruce, 1965. + Periclimenes denticulatus Nobili, 1906. + Periclimenes elegans (Paulson, 1875). t+ Periclimenes ensifrons (Dana, 1852). } Periclimenes grandis (Stimpson, 1860). +Periclimenes holthuisi Bruce, 1969. Periclimenes longirostris (Borradaile, 1915): Bruce, 1979a. Periclimenes ornatellus Bruce, 1979: Bruce, 1979a. Periclimenes ornatus Bruce, 1969: Bruce, 1979a. Periclimenes pilipes Bruce and Zmarzly, 1983: Bruce and Zmarzly, 1983. + Periclimenes seychellensis Borradaile, 1915. Periclimenes soror Nobili, 1904: Bruce, 1978; 1979a. T Periclimenes spiniferus (De Man, 1902). Periclimenes tenuipes Borradaile, 1898: Bruce, 1979a. + Periclimenes tenuis Bruce, 1969. +Philarius gerlachei (Nobili, 1905). +Philarius imperialis (Kubo, 1940). + Pontoniopsis comanthi Borradaile, 1915. + Stegapontonia commensalis Nobili, 1906. Thaumastocaris streptopus Kemp, 1922: Bruce, 1979a. Family GNATHOPHYLLIDAE +Gnathophyllum americanum Guérin-Meneville, 1855. t+ Hymenocera picta Dana, 1852. Superfamily ALPHEOIDEA Family ALPHEIDAE Alpheopsis diabolis Banner, 1956: Banner and Banner, 1968. Alpheopsis equalis Coutiere, 1896: Banner and Banner, 1968. Alpheus acutofemoratus Dana, 1852: Banner and Banner, 1968; Highsmith, 1981. Alpheus alcyone De Man, 1902: Banner and Banner, 1968. Alpheus alpheopsides Coutiere, 1905: Banner and Banner, 1968. Alpheus amirantei sizou Banner and Banner, 1967: Banner and Banner, 1968; Highsmith, 1981. Alpheus bidens (Olivier, 1811): Banner and Banner, 1968. Alpheus bradypus Coutiere, 1905: Banner and Banner, 1968. Alpheus brevipes Stimpson, 1860: Banner and Banner, 1968. Alpheus bucephalus Coutiere, 1905: Banner and Banner, 1968; Highsmith, 1981. Alpheus clypeatus Coutiere, 1905: Banner and Banner, 1968. Alpheus collumianus Stimpson, 1861: Highsmith, 1981. Alpheus collumianus medius Banner, 1956: Banner and Banner, 1968. Alpheus diadema Dana, 1852: Banner and Banner, 1968. Alpheus dolerus Banner, 1956: Banner and Banner, 1968. Alpheus ehlerii De Man, 1909: Banner and Banner, 1968. Alpheus frontalis H. Milne-Edwards, 1837: Banner and Banner, 1968. Alpheus gracilis Heller, 1861. Alpheus gracilis gracilis Heller, 1861: Banner and Banner, 1968. Alpheus idiocheles Coutiere, 1905: Banner and Banner, 1968. + Alpheus leviusculus Dana, 1852. Alpheus lobidens polynesica Banner and Banner, 1974. Alpheus crassimanus Heller, 1865: Banner and Banner, 1968. Alpheus lottini Guerin, 1829: Banner and Banner, 1968. Alpheus obesomanus Dana, 1852: Banner and Banner, 1968. Alpheus ovaliceps Coutiere, 1905: Banner and Banner, 1968. Alpheus pachychirus Stimpson, 1860: Banner and Banner, 1968. Alpheus pacificus Dana, 1852: Banner and Banner, 1968. Alpheus paracrinitus Miers, 1881: Banner and Banner, 1968. Alpheus paralcyone Coutiere, 1905: Banner and Banner, 1968. Alpheus parvirostris Dana, 1852: Banner and Banner, 1968; Highsmith, 1981. Alpheus samoa Banner and Banner, 1966: Banner and Banner, 1968. +New records for Enewetak. (This table continued on next page.) 224 DEVANEY AND BRUCE TABLE 1 (cont'd) a a EE EEE eee Family ALPHEIDAE (cont'd) Alpheus strenuus strenuus Dana, 1852. Alpheus strenuus Dana, 1852: Banner and Banner, 1968. Alpheus sulcatus Kingsley, 1878. Alpheus macrochirus Richters, 1880: Banner and Banner, 1968. Athanas areteformis Coutiere, 1903: Banner and Banner, 1968. Athanas djiboutensis Coutiere, 1897: Chace, 1955; Banner and Banner, 1960, 1968. Athanas dorsalis (Stimpson, 1861): Banner and Banner, 1960, 1968. Athanas esakii Kubo, 1940: Banner and Banner, 1968. Athanas indicus (Coutiere, 1903): Banner and Banner, 1960, 1968. Athanas marshallensis Chace, 1955: Chace, 1955; Banner and Banner, 1960, 1968. Athanas rhothionastes Banner and Banner, 1960: Banner and Banner, 1960, 1968. Athanas verrucosus Banner and Banner, 1960: Banner and Banner, 1960, 1968. Automate dolichognatha De Man, 1888. Automate gardineri Coutiére, 1902: Banner and Banner, 1968. Automate johnsoni Chace, 1955: Chace, 1955. Metalpheus paragracilis (Coutiere, 1897). Alpheus paragracilis Coutiére, 1897: Banner and Banner, 1968. Metalpheus rostratipes (Pocock, 1890). Alpheus rostratipes Pocock, 1890: Banner and Banner, 1968. Salmoneus serratidigitus (Coutiere, 1896). Salmoneus sibogae (De Man, 1910): Banner and Banner, 1968. Salmoneus tricristatus Banner, 1959: Banner and Banner, 1968. Synalpheus carinatus (De Man, 1888): Banner and Banner, 1968. Synalpheus charon (Heller, 1861): Banner and Banner, 1968. Synalpheus near coutierei Banner, 1953: Highsmith, 1981. Synalpheus demani Borradaile, 1900: Banner and Banner, 1968. Synalpheus hastilicrassus Coutiere. 1905: Banner and Banner, 1968. Synalpheus heroni Coutiere, 1909.+ Synalpheus paraneomeris Coutiere, 1905: Banner and Banner, 1968. Synalpheus stimpsonii (De Man, 1888): Banner and Banner, 1968. Synalpheus tumidomanus (Paulson, 1875): Banner and Banner, 1968; Highsmith, 1981. Family HIPPOLYTIDAE + Hippolyte (?)ventricosa H. Milne-Edwards, 1837. + Hippolyte sp. + Ligur uveae (Borradaile, 1899). + Lysmata vittata (Stimpson, 1860). + Lysmata sp. + Saron marmoratus (Olivier, 1811). + Saron neglectus De Man, 1902. + Thor amboinensis De Man, 1888. + Thor maldivensis Borradaile, 1915. T Thor paschalis (Heller, 1862). Family PROCESSIDAE Nikoides steinii (Edmondson, 1935). Nikoides nanus Chace, 1955: Chace, 1955. } Nikoides multispinatus Hayashi, 1981. + Processa (?)japonica (De Haan, 1844). + Processa molaris Chace, 1955. + Processa neglecta Hayashi, 1975. Superfamily PANDALOIDEA Family PANDALIDAE Micropandalus hardingi Bruce, 1983: Bruce, 1983. Family THALASSOCARIDIDAE +Thalassocaris crinata (Dana, 1852). Superfamily CRANGONOIDEA Family CRANGONIDAE tPontophilus sp. aff. P. sabsechota Kemp, 1911 + Vercoia gibbosa Baker, 1904. +New records for Enewetak. (This table continued on next page.) CRUSTACEA DECAPODA 225 TABLE 1 (cont'd) Infraorder PALINURA Superfamily PALINUROIDEA Family SYNAXIDAE t Palinurellus wieneckii (De Man, 1881). Family PALINURIDAE t Panulirus longipes femoristriga (von Martens, 1872). Panulirus pencillatus (Olivier, 1791): Ford et al., 1979; Cooke and MacDonald, 1981; Ebert and Ford, 1986. Family SCYLLARIDAE + Parribacus antarcticus (Lund, 1793). +New records for Enewetak. STENOPODIDEA Identified stenopodid shrimps determined from Enewetak include the pantropical banded coral shrimp, Stenopus hispidus; S. tenuirostris, from the western Indian Ocean, Philippines, and the western Pacific; S. zanzibar- icus, from the western Indian Ocean and central Pacific (Canton Island); Odontozona sculpticaudata, known from Malayan and northeastern Australian waters; and two species of Microprosthema. One of these, M. scabricauda- tum, has been recorded from the western Indian Ocean, New Guinea, and the Ryukyu’s, whereas M. plumicorne has previously been recorded only from Mauritius in the western Indian Ocean. All of these stenopodid shrimps are apparently new records from the Marshall Islands. J. W. Goy (personal communication) has also identified S. hispidus from Jaluit Atoll (USNM specimens). Recently, an undescribed species of Odontozona was found and awaits description. CARIDEA One widely distributed Indo-West Pacific pasiphaeid shrimp, a species of Leptochela, is known from Enewetak and several other Marshall Islands (Chace, 1955; 1976). It was said to have been the most common decapod found during the 1946 and 1947 survey and was collected at night on the surface using a light. Four species of rhynchocinetid shrimps are recorded: Rhynchocinetes hiatti occurs at Enewetak as well as several other Pacific island areas; R. hendersoni, reported as a new species (R. marshallensis) from Enewetak by Edmond- son (1952), is a wide-spread Indo-West Pacific form (Hol- thuis and Hayashi, 1967); a species approaching R. rigens is newly recorded from the atoll; a fourth species is as yet undetermined and awaits description (Holthuis, personal communication). Palaemonid shrimps known from Enewetak are represented by 43 identified and 10 unidentified species in 18 genera. All but two species are members of the sub- family Pontoniinae. Rosewater (1965) reported a species of Paranchistus and Fankboner (1972) a species of Anchistus as symbionts from tridacnid clams. Anchistus australis and three species of Conchodytes can now be added to the list of symbionts associated with tridacnid and ostreid bivalves at Enewetak. Paratypton siebenrocki, a species forming galls in Acropora coral, was recorded from Enewetak by Bruce (1969). Two widely distributed Indo-West Pacific species of Palaemonella (P. pottsi and P. rotumana) were reported from Enewetak by Bruce (1970), the former a commensal on the crinoid Comanthus bennetti but more recently found on two additional crinoid species (Table 2). Periclimenes soror, a symbiont on many tropical asteroids, was first recorded from Enewetak by Bruce (1978). Laboratory sponsored projects in 1974 led to the collec- tion and description of eight pontoniine shrimps, one being new and five being widely distributed tropical species pre- viously unrecorded from the Marshall Islands (Bruce, 1979a). Four of these species were associated with sea anemones; one each was found on a sea star, sponge, and tridacnid clam; and one is apparently a free-living form. A reef coral workshop held at the Mid-Pacific Research Laboratory (MPRL) in 1976 led to the collection and description of a new genus and species of pontoniine shrimp associated with a stony coral (Bruce, 1979b). Stud- ies on the commensals and parasites associated with coma- tulid crinoids (feather-stars) by Ann Fielding and Dennis Devaney in 1976 resulted in finding seven pontoniine and three alpheid shrimps as well as a galatheid and a brachy- uran crab (Table 2). One of the pontoniines, Periclimenes pilipes, is a species described as new from Enewetak (Bruce and Zmarzly, 1983). A study detailing the biological relationships between Enewetak crinoids, their symbiotic crustaceans, and other invertebrates has recently been pub- lished (Zmarzly, 1984). Two species of Harpiliopsis and one species of Fennera are symbionts on_pocilliporid corals. Similarly, the five species of Coralliocaris and two species of Philarius, herein recorded from Enewetak for the first time, are also symbionts of Acropora corals in other parts of the Indo-West Pacific (Bruce, 1974). Thirty of the 48 identified pontoniine shrimps recorded from Enewetak are also known from Heron Island, Australia (Bruce, 1981). The remainder are, with few exceptions, wide- spread tropical Indo-West Pacific or pantropical species. All but three (Anchistus demani, Onycocaris stenolepis, and Pontonia hurii) of the 20 pontoniines recorded by Holthuis 226 DEVANEY AND BRUCE TABLE 2 Symbiotic Decapod Crustacea Associated with Crinoids at Enewetak Atoll No. symbionts/ Symbiont* Host crinoidt No. hosts Palaemonidae Araiopontonia odontorhyncha —- Comanthus bennetti 1/7 Palaemonella pottsi Comanthus parvicirrus 1/2 Comaster gracilis 3/6 Periclimenes? amboinensis Comanthus bennetti 4/7 Comanthus parvicirrus 1/2 Periclimenes commensalis Comanthus bennetti 5/7 Periclimenes tenuis Comanthus bennetti 1/7 Periclimenes pilipes Comanthus parvicirrus 1/2 Comanthina schegeli 1/2 Pontoniopsis comanthi Comanthus bennetti 1/7 Comanthus parvicirrus 1/2 Alpheidae Athanas sp. Comaster gracilis 2/6 Synalpheus carinatus Comanthina schegeli 2/2 Comanthus parvicirrus 1/2 Synalpheus stimpsoni Comanthus bennetti 1/2 Comanthus parvicirrus 1/7 Comaster gracilis 5/6 Galatheidae Allogalathea elegans Comanthina schegeli 2/2 Comanthus bennetti 3/7 Parthenopidae Harrovia elegans Comaster gracilis 2/6 "Determinations or verifications of palaemonids made by AJB; alpheids by A. H. Banner; galatheid by K. Baba; parthenopid by Ann Fielding. +Determinations of Comanthina schegeli and Comanthus _parvicirrus resulted from work by Debbie Zmarzly, Scripps Institution of Oceanography, during her more extensive research on crinoid ecology and symbionts in 1980. (1953) from other atolls in the Marshall Islands have been found at Enewetak. The gnathophyllid shrimp, Hymenocera picta, has been observed at Kwajalein and Enewetak atolls. At Enewetak, a pair was seen at night well back in a cave (S. Johnson, personal communication) and a specimen photographed in the Enewetak quarry. An ovigerous specimen of Gnathophyllum americanum was collected in the Enewetak lagoon. Both gnathophyllids are widely distributed throughout the Indo-West Pacific region; the latter species is also found in the tropical eastern Pacific and Caribbean. Three species of alpheid shrimps were recorded at Enewetak by Chace (1955) based on collections from the northern Marshall Islands. More intensive study on Enewetak alpheids was made by A. H. Banner in Febru- ary and March 1957. The results of this latter work (Banner and Banner, 1960; 1968) increased the known number of alpheids from Enewetak to 49 species including two new species of Athanas (three species of Athanas listed only from “Marshall Islands” in these two papers include specimens from Enewetak). The 1968 paper is especially important in providing station data with collec- tion locality, depth, and habitat.* Beginning in 1976, exam- ination of symbionts associated with comatulid crinoids at Enewetak revealed two species of Synalpheus (S. carinatus and S. stimpsoni) and an undetermined species of Athanas (Table 2). One or more species of alpheids have also been observed together with gobiid fishes in burrows made in Enewetak Lagoon sediment. Seven alpheid species were found by Highsmith (1981) as associates of six different corals. Only two, Alpheus acutofemoratus and A. par- virostris, were found in more than two coral heads of the same species, and the former alpheid was found in all but one of the corals studied. Altogether, 52 identified alpheid species in seven genera are now known from Enewetak of ‘In 1961 a fire destroyed many of the specimens upon which these two papers were based (Banner and Banner, 1961). For- tunately, representatives of most of the species were deposited in the MPRL reference collection and are still available. CRUSTACEA DECAPODA 227 which two are new records. Nearly 80% (42 species) are broad ranging Indo-West Pacific forms, many recorded as far as the Red Sea as well as Indian and Pacific Oceans. An additional 10% (5 species) are pantropical with the bal- ance showing a more restricted distribution. An additional 22 alpheid taxa were recorded from atolls other than Enewetak in the Marshall Islands (Banner, 1957; Banner and Banner, 1968). Several of these are now considered synonyms of species recorded herein from Enewetak, while the others may be expected to occur there (Table 3). Eight identified hippolytid shrimps—members of the genera Hippolyte, Ligur, Lysmata, Saron, and Thor, each with a broad Indo-West Pacific distribution—are newly recorded from Enewetak. Two of these, S. marmoratus and S. neglectus, were also recorded from the southern Marshall Islands (Holthuis, 1953). Also, a new species of Hippolyte, to be reported at a later date (Holthuis, per- sonal communication) and at least one undescribed Lysmata, have been found at Enewetak. Among processid shrimps now known from Enewetak, one species of Nikoides was previously recorded from this atoll as well as Bikini, whereas a second species, N. sibo- gae, was collected only at Bikini (Chace, 1955). A third species, N. multispinatus, is newly recorded and was origi- nally reported from the Great Barrier Reef and Ishigakijima Island, Japan. Three species of Processa_ including P. molaris—a new species recorded by Chace from Rongelap and Bikini—are now recorded from Enewetak. A new genus and species of symbiotic pandalid shrimp, Miropandalus hardingi, has been found on branching black coral in waters off the seaward side of Enewetak Atoll (Bruce, 1983). Thalassocaris crinita, is newly recorded from Enewetak. This species is known from Indonesia and other Indo- TABLE 3 Non-Enewetak Marshall Island Alpheid Shrimp Records Species + Alpheus amirantei Coutiere + A. arnoa Banner A. collumianus + probabilis Banner 4A. crockeri Armstrong + A. deuteropus Hilgendorf T# A. gracilipes Stimpson ¢ A. gracilis var. simplex Banner + A. lutini Coutiere +A. malleodigitus (Spence-Bate) +A. miersi Coutiere + A. nanus (Banner) A. paracrinitus var. Tbengalensis Coutiere +A. percyi Coutiere +A. styliceps Coutiere +A. superciliaris Coutiere + A. ventrosus H. Milne Edwards + Alpheus sp. No. IV + Aretopsis amabilis de Man + Metabetaeus minutus (Whitelegge) + Synalpheus coutierei Banner +S. laticeps Coutiere + S. lophodactylus Coutiere Present status*® Atoll Valid Arno Valid Arno A. collumianus Stimpson Arno (Banner and Banner, 1981) Valid Bikini Valid Arno Valid Arno, Bikini, Rongerik A. gracilis Heller Arno (Banner and Banner, 1982) A. obesomanus Dana Arno (Banner and Banner, 1966) Valid Bikini, Rongelap Valid Bikini Metalpheus Arno rostratipes (Pocock) (Banner and Banner, 1964) A. paracrinitus Miers Arno (Banner and Banner, 1982) Valid Arno, Rongerik Valid Bikini, Rongelap Valid Rongerik A. lottini Guérin Arno (Banner and Banner, 1981) Incomplete specimen, Rongelap still undescribed Valid Rongelap Valid Arno Valid Arno Valid Rongelap Valid Aro *D. M. Banner (personal communication). +Recorded in Banner, 1957. +Recorded in Banner and Banner, 1968. 228 DEVANEY AND BRUCE Pacific waters including the Marshall Islands (Chace, 1955). Two crangonid shrimps were taken in the Enewetak Lagoon. One is possibly a new species of Pontophilus, whereas the other, Vercoia gibbosa, was previously known only from south Australia. PALINURA At least four palinuran lobsters are represented at Enewetak. Panulirus penicillatus is a common inhabitant of the windward reef flat and was recorded from Arno Atoll in the southern Marshall Islands (Holthuis, 1953). This spiny lobster was the focus of population and other life his- tory studies at Enewetak during 1978 and 1979 (Ford et al., 1979; Ebert and Ford, 1986). Juvenile (puerulus and post-puerulus) stages of this wide-spread Indo-West Pacific rock lobster have been recorded by Cooke and McDonald (1981) from Enewetak. Three specimens of P. longipes femoristriga were identified from a collection made in a surge channel near the outer reef. Although not previously reported from the Marshall Islands, this subspecies is reported widely from Japan, the Moluccas, New Guinea, eastern Australia and Polynesia (George and Holthuis, 1965). Panulirus versicolor, recorded by Holthuis (1953) from Arno, has not been reported at Enewetak. Parribacus antarcticus and Palinurellus wieneckii are now known from Enewetak. Although each of the last two species has a broad Indo-West Pacific distribution, only P. antarcticus had previously been recorded from the Marshall Islands (Arno) by Holthuis (op. cit.). The latter has also been recognized in waters off Kwajelein Atoll (S. Johnson, per- sonal communication). COLLECTION RECORDS Taxa are listed in the same order as in the checklist under their respective families. Higher categories may be found in the checklist. These records are not known to have been previously published. When known, the islet, month, year, habitat, depth, collector, identifier, and number of specimens are given. Many of the specimens collected between 1956 and 1971 and determined by L. B. Holthuis (LBH) are deposited at the Allan Hancock Foundation. Other specimens have been deposited in several institutions, mainly MPRL, BPBM (Honolulu), RMNH (Leiden), and USNM. Identifiers AJB and DMD refer to the authors; other identifiers are named in full. Family PENAEIDAE Metapenaeopsis borradailei (De Man) Enewetak Islet: Sept. 1980, lagoon, north end of islet, in sandy substrate using airlift, 12 m, DMD and P. Co lin, id. LBH; 1 spec. (juv. F). Medren Islet: Aug. 1956, lagoon side, light at night, id. LBH; 1 spec. Metapenaeopsis sp. Aomon, Bijile, Lojwa Islets: July 1959, lagoon side, sand bottom, 1.8 to 7.3 m, F. C. Ziesenhenne, id. LBH; 1 spec. Enewetak Islet: Feb. 1982, on sand at night (“burying shrimps”), 5 m, S. Johnson, id. LBH; 1 spec. (M). Medren Islet: July 1959, lagoon side, sand bottom, 2.4 to 4.7 m, J. S. Garth, id. LBH; 1 spec. Family SICYONIIDAE Sicyonella maldivensis Borradaile Medren Islet: Aug. 1956, lagoon:side, light at night, id. LBH; 9 spec. (3 lots). Family STENOPODIDAE Microprosthema plumicorne (Richters) Enewetak Islet: Dec. 1981, lagoon, under dead Acropora, 4 m, S. Johnson, id. P. Galloway and DMD; 1 spec., ovig.; BPBM. Microprosthema scabricaudatum (Richters) Alembel Islet: July 1959, lagoon side, Acropora coral, J. S. Garth, id. LBH; 1 spec.; BPBM. Odontozona sculpticaudata Holthuis Medren Islet: Feb. 1982, lagoon side, under dead coral, 5 m, S. Johnson, id. DMD; 1 spec., ovig.; BPBM. Odontozona sp. Lagoon: Sept. 1982, in submerged part of cement ship at night, 5 m, S. Johnson; 1 spec.; April 1983, exposed on rather silty bottom next to bulkhead of submerged part of cement ship, at night (ca. 2030 h), 5 m, S. John- son; 2 spec. Stenopus hispidus (Olivier) Biken Islet: July 1975, oceanside, rotenone station, 9 m, P. B. Lamberson, id. DMD; 1 spec.; BPBM. Reefer 8 Pinnacle: June 1982, in cave, 8 m, S. Johnson, id. DMD; 2 spec. (1 ovig., 1 male); BPBM. Stenopus tenuirostris De Man Enewetak Atoll: May 1983, lagoon, under sheet of aluminum debris, 3 m, S. Johnson; id. DMD; 1 spec.; BPBM. Stenopus zanzibaricus Bruce Enewetak Islet: May 1983, lagoon, under sheet of alumi- num debris, 3 m, S. Johnson, id. DMD; 2 spec.; BPBM. Family RHYNCHOCINETIDAE Rhynchocinetes hiatti Holthuis and Hayashi Enewetak Atoll: May 1978, algal ridge, recovered from fish poison (rotenone) station, id. AJB; 5 spec. (juv.). Rhynchocinetes rigens Gordon Japtan Islet: Feb. 1982, lagoon, in submerged metal locker, 12 m, S. Johnson, id. LBH; 2 spec. Rhynchocinetes sp. aff. R. rigens Japtan Islet: May 1978, lagoon side, 100 yds. off pier at base of coral head, id. LBH; 1 spec. Lagoon Mooring Buoy Pinnacle: Feb. 1982, on ledge, night, 10 m, S. Johnson, id. LBH; 1 spec; BPBM. Family PALAEMONIDAE Anchistus australis Bruce Ananij Islet: Aug. 1968, in Tridacna squamosa (Lam.), J. W. Knudsen, id. AJB; 1 spec. (ovig.). Jinderol Islet: March 1961, probably in Tridacna noae (Ré6ding) [Tridacna maxima (Réding)], R. Palumbo, id. LBH; 2 spec., ovig. Medren Islet: July 1959, reef between south end of islet and wreck on reef, under large pancake coral cemented to reef, F. Ziesenhenne, id. LBH; 1 spec., ovig. CRUSTACEA DECAPODA 229 Anchistus miersi (De Man) Alembel Islet: July 1959, north side, Acropora coral, J. S. Garth, id. LBH; 3 spec. Bokoluo Islet: Feb. 1968, in Tridacna sp., J. W. Knudsen, id. AJB; 2 spec. (ovig.). dinedrol Islet: March 1961, lagoon side, probably in Hippopus hippopus, shallow water, R. Palumbo, id. LBH; ? spec. Ariopontonia odontorhyncha Fujino and Miyake Medren Islet: July 1975, ocean side, on crinoid Comanthus bennetti, 12 m, Lamberson, id. AJB; 1 spec. Medren Islet: Feb. 1976, lagoon, ca. % mile off islet, on crinoid C. bennetti, 6 to 9 m, D. M. Devaney and A. Fielding, id. AJB; 2 spec. (1 male, 1 ovig. female); BPBM. Brachycarpus biunguiculatus (Lucas) Biken Islet: May 1946, 2 miles south of islet, leeward side of reef, light at night, L. P. Schultz, id. LBH; 19 spec. Conchodytes biunguiculatus (Paulson) Pole Pinnacle: Nov. 1977, lagoon, 30.4 m, coll. and id. Ann Fielding; ? spec. Conchodytes meleagrinae Peters Enewetak Islet: April 1961, lagoon side, in Pinctada? galtsoffi, 6 m, E. S. Reese, id. J. Yaldwyn; 2 spec. (1 male, 1 female). Conchodytes tridacnae Peters Lojwa Islet: Feb. 1971, inside Tridacna gigas, J. Shoup, id. LBH; 4 spec. (2 ovig.). July 1976, inside Tridacna maxima, id. Ann Fielding; ? spec. Coralliocaris brevirostris Borradaile Enewetak Islet: July 1967, in Acropora corymbosa, just behind algal ridge, J. W. Knudsen, id. AJB; 1 spec., ovig. Coralliocaris graminea (Dana) Alembel Islet: July 1959, lagoon, north side, Acropora, J. S. Garth, id. LBH; 2 spec. Ananij Islet: Feb. 1957, lagoon side, dead Acropora, 1.8 m, A. H. Banner, id. LBH; 3 spec. Ananij Islet: Jan. 1960, lagoon side, north end, Acropora, E. S. Reese, id. LBH; 2 spec. Ananij Islet: Jan. 1960, lagoon side, from Seriatopora, E. S. Reese, id. LBH; 2 spec. Biken Islet: Feb. 1957, ocean side, inner edge reef flat, on Acropora and Pocillopora, 0.5 m, A. H. Banner, id. LBH; 2 spec., (1 male, 1 ovig.). Ikuren Islet: July 1959, ocean side, Acropora, J. S. Garth, id. LBH; 2 spec. Medren Islet: July 1959, lagoon side, living Acropora, J. Coatsworth, id. LBH; 1 spec. Coralliocaris nudirostris (Heller) Boken Islet: July 1957, southeast side, living coral, J. S. Garth, id. LBH; 1 spec. (female). Enewetak Islet: April 1961, in Acropora hyacinthus, E. S. Reese, id. LBH; 2 spec. Medren Islet: July 1959, lagoon side, coral, 3.6 to 4.7 m, J. Coatsworth, id. LBH; 2 spec. Coralliocaris superba (Dana) Biken Islet: Aug. 1956, ocean side in coral head on reef flat, common, found among coral branches, id. LBH; 1 spec. Boken Islet: July 1957, southeast side, living coral, J. S. Garth, id. LBH; 1 spec. Bokoluo Islet: March 1968, in Acropora digitifera, on ocean reef flat, J. W. Knudsen, id. AJB; 1 spec., (male). Enewetak Islet: April 1961, in Turbinaria? danae, E. S. Reese, id. LBH; 1 damaged spec., identity not fully certain. Enjebi Islet: July 1959, shore, F. C. Ziesenhenne, id. LBH; 2 spec. Japtan Islet: Jan. 1960, lagoon side, north end, Acropora, E. S. Reese, id. LBH; 2 spec. Jinimi Islet: July and Aug. 1961, lagoon side, 4.7 m, J. Shoup, id. LBH; 2 spec. Coralliocaris venusta Kemp Enewetak Islet: Lagoon coral mound, % mile off islet, from live Acropora, 10 m, coll. S. Swerdloff, id. AJB; 1 spec. (ovig.). Enewetak Islet: Aug. 1969, lagoon, in Acropora sp., 10 m, J. W. Knudsen, id. AJB; 1 spec. (ovig.). Fennera chacei Holthuis Enewetak Islet: Feb. 1976, ocean side, coll. with crinoid Comanthus schelegi, 23 m, P. Lamberson, id. AJB; 1 spec. Harpiliopsis beaupresii (Andouin) Biken Islet: July and Aug. 1961, lagoon side, 2.4 m, J. Shoup, id. LBH; 5 spec. Boken Islet: July 1957, southeast side, Pocillopora coral, J. S. Garth, id. LBH; 5 spec. Enjebi Islet: July 1959, shore, F. C. Ziesen- henne, id. LBH; 2 spec. Japtan Islet: Jan. 1960, lagoon side, Pocillopora coral, 3 m, E. S. Reese, id. LBH; 2 spec. Medren Islet: July 1957, lagoon side, coral heads, 3 to 4.7 m, J. S. Garth, id. LBH; 1 spec. Medren Islet: July 1959, lagoon side, from Pocillopora damicornis, 3.6 to 4.7 m, J. Coatsworth, id. LBH; 2 spec. Harpiliopsis depressa (Stimpson) Ananij Islet: Feb. 1957, lagoon side, Pocillopora coral, 1.8 m, A. H. Banner, id. LBH; 2 spec. Biken Islet: July to Aug. 1961, lagoon side, 2.4 m, J. Shoup, id. LBH; 10 spec. Enewetak Islet: April 1961, in Seriatopora? hystrix, E. S. Reese, id. LBH; 2 spec. Enewetak Islet: July and Aug. 1961, lagoon side, 7.7 and 12 m, J. Shoup, id. LBH; 8 spec. (2 lots). Ikuren Islet: July 1959, coral, J. S. Garth, id. LBH; 3 spec. Harpiliopsis spinigera (Ortmann) Lojwa Islet: July 1967, behind reef on south side of islet, dead and overgrown coral with small amount of live coral, in Stylophora pistillata (mordax), 2 m, J. W. Knudsen, id. AJB; 1 spec. Jocaste japonica (Ortmann) Biken Islet: July and Aug. 1961, lagoon side, 2.4 m, J. Shoup, id. LBH; several spec. Jocaste lucina (Nobili) Alembel Islet: July 1959, north side, living Acropora coral, J. S. Garth, id. LBH; 1 spec. Ananij Islet: Feb. 1957, lagoon side, dead Acropora coral, 1.8 m, A. H. Banner, id. LBH; 2 spec. Ananij Islet: Jan. 1960, north end, lagoon side, Acropora coral, E. S. Reese, id. LBH; 2 spec. (2 lots). Biken Islet: Aug. 1956, ocean side, for- malin washings from coral, D. Reish, id. LBH; 4 spec. Biken Islet: Feb. 1957, ocean side, inner edge reef flat, from coral, A. H. Banner, id. LBH; 1 spec. Biken Islet: July and Aug. 1961, lagoon side, 2.4 m, J. Shoup, id. 230 DEVANEY AND BRUCE LBH; 3 spec. Boken Islet: July 1957, southeast side, liv- ing coral, J. S. Garth, id. LBH; 5 spec. Enjebi Islet: July 1959, shore, F. C. Ziesenhenne, id. LBH; 2 spec. Ikuren Islet: July 1959, ocean side, living Acropora coral, J. S. Garth, id. LBH; 1 spec. Japtan Islet: Jan. 1960, lagoon side, north end, Acropora coral, E. S. Reese, id. LBH; 2 spec. (2 lots). Jinimi Islet: July and Aug. 1961, lagoon side, 4.7 m, J. Shoup, id. LBH; 10 spec. Medren Islet: July 1959, lagoon side, living Acropora, J. Coatsworth, id. LBH; 3 spec. Leandrites cyrtorhynchus Fujino and Miyake Japtan Islet: Feb. 1982, lagoon off Japtan pier, in metal shrimp locker, 12 m, S. Johnson, id. LBH; 6 spec. (1 ovig.). Onycocaris quadratophthalma (Balss) Ikuren Islet: Aug. 1956, ocean side, sponges from under surfaces of rocks, D. J. Reish, id. LBH; 5 spec. Palaemonella pottsi (Borradaile) Enewetak Islet: Feb. 1976, ocean side, on crinoid Comanthus parvicirrus, 30 m, P. Lamberson, id. AJB; 1 spec. Enewetak Islet: Feb. 1976, lagoon (Mini Power Plant Pinnacle), on 3 crinoids Comaster gracilis, 6 to 12 m, D. M. Devaney, id. AJB: 6 spec. (3 lots: 1 ovig. and 1 juv.; 1 male; 3). Palaemonella rotumana (Borradaile) Biken Islet: Feb. 1957, ocean side, inner edge of reef flat, from coral, 0.5 m, A. H. Banner, id. LBH; 1 spec. Bokonbako Islet: July 1957, ocean side, old coral heads, 0.3 m, D. J. Reish, id. LBH; 1 spec. Palaemonella tenuipes Dana Enewetak Islet: July 1957, ocean side, reef flat, J. S. Garth, id. LBH; 4 spec. Medren Islet: Feb. 1957, lagoon side, mixed coral, 2.4 m, A. H. Banner, id. LBH; 1 spec. Medren Islet: July 1957, lagoon side, coral, 3 to 4.7 m, J. S. Garth, id. LBH; 2 spec. Palaemonella sp. Ananij Islet: Feb. 1957, lagoon side, Acropora coral, 1.8 m, A. H. Banner, id. LBH; 1 spec. Paranchistus armatus (H. Milne Edwards) Alembel Islet: Sept. 1956, lagoon side, from Tridacna gigas, id. LBH; 1 spec. Jinedrol Islet: March 1961, lagoon side, probably in Tridacna gigas, shallow water, R. Palumbo, id. LBH; 1 spec. ovig. Periclimenaeus sp. Medren Islet: Feb. 1957, lagoon, south end, patch reef about 600 ft offshore, surrounding bottom sand, three dead, overgrown coral heads (Pocillopora, Acropora, Porites) broken up, 2.4 m, A. H. Banner, id. LBH; 1 spec. Periclimenes agag Kemp Enewetak Lagoon: Jan. 1982, under dead coral, 5 m, S. Johnson, AJB; 1 spec. Periclimenes ?amboinensis (De Man) Enewetak Islet: Feb. 1976, ocean side, on crinoid Comanthus parvicirrus, 31 m, P. Lamberson, id. AJB; 1 spec. (juv.). Medren Islet: Feb. 1976, lagoon, ca. % mile off islet, on 3 crinoids Comanthus bennetti, 6 to 9 m, DMD and A. Fielding, id. AJB; 6 spec. (3 lots, each with male and ovig. female). Periclimenes bayeri Holthuis Enewetak Islet: April 1961, from Pocillopora verrucosa or P. elegans coral, E. S. Reese, id. LBH; 3 spec. Enewetak Islet: July and Aug. 1961, ocean side, Pocillopora? elegans coral, 1.5 m, J. Shoup, id. LBH; 2 spec. Enewetak Islet: July and Aug. 1961, lagoon side, 7.7 m, J. Shoup, id. LBH; 2 spec. Periclimenes commensalis Borradaile Enewetak Islet: Feb. 1976, lagoon pinnacle ca. ¥2 to 1 mile off islet, on crinoid Comanthus bennetti, 5 m, DMD and A. Fielding, id. AJB; ? spec. Medren Islet: Feb. 1976, lagoon, ca. % mile off islet, on 4 crinoids Comanthus bennetti, 6 to 9 m, DMD and A. Fielding, id. AJB; 54 spec. (4 lots: 16; 4 plus post-larva: 22 includ- ing 1 ovig.; 9 including 2 ovig.). Periclimenes cristimanus Bruce Enewetak Islet: Aug. 1968, rock quarry, on sea urchin Echinothrix calamaris, J. W. Knudsen, id. AJB; 6 spec. (2 lots, one with 5 including ovig., another with 1 ovig.). Periclimenes denticulatus Nobili Billae and Runit Islet: July 1967, acroporid coral and algae in lagoon dredge hauls between the two islets, 29 to 32 m, J. W. Knudsen (Stn. 353), id. AJB; 2 spec. Periclimenes elegans (Paulson) Alembel Islet: Feb. 1957, reef flat, coral, 0.6 m, A. H. Banner, id. LBH; 1 spec. (discarded—moldy). Ananij Islet: July 1957, ocean side, reef flat, J. S. Garth, id. LBH; 1 spec. Biken Islet: July 1959, southeast side between ocean and lagoon, rock with sand substrate, dead coral, J. S. Garth, id. LBH; many spec. Bokoluo Islet: Aug. 1968, ocean reef, from Porites sp., J. W. Knudsen, id. AJB; 1 spec., ovig. Bokonbako Islet: July 1957, lagoon side, old coral heads, D. Reish, id. LBH; 1 spec. Enewetak Islet: July 1957, ocean side, reef flat, J. S. Garth, id. LBH; 5 spec. Enewetak Islet: Aug. 1968, J. W. Knudsen, id. AJB; 4 spec. (1 ovig.). Enjebi Islet: July 1957, ocean side, reef flat, J. S. Garth, id. LBH; 1 spec. Japtan Islet: Aug. 1968, inner reef flat pool, J. W. Knudsen, id. AJB; 7 spec. (3 ovig.). Japtan Islet: Aug. 1968, lagoon tide pools, J. W. Knudsen, id. AJB; 13 spec. (5 male, 8 female with 7 of these ovig.). Japtan Islet: Aug. 1968, reef flat pool, poison station, J. W. Knudsen, id. AJB; 7 spec., (2 ovig.). Lojwa Islet: July 1967, in Stylophora pistillata (mordax) 1.5 m, J. W. Knudsen, id. AJB; 1 spec. Lojwa Islet: Ocean reef pool, J. W. Knudsen, id. AJB; 1 spec. (male). Periclimenes ensifrons (Dana) Lujor Islet: June 1946, outside lagoon, intertidal, M. W. Johnson, id. LBH; 1 spec. (ovig.). Periclimenes grandis (Stimpson) Kedrol Islet: May 1946, intertidal, in pot holes, M. W. Johnson, id. LBH; 1 spec. Periclimenes holthuisi Bruce Enewetak Islet: March 1982, lagoon side, found in exposed tentacles of poritid coral (Alveopora or CRUSTACEA DECAPODA 231 Goniopora), 25 m, S. Johnson, id. DMD; 2 spec., (1 ovig., 1 juv.). Periclimenes seychellensis Borradaile Medren Islet: July 1957, lagoon side, algae on coral, D. J. Reish, id. LBH; 1 spec. Periclimenes spiniferus (De Man) Jinimi Islet: July and Aug. 1961, lagoon side, 4.7 m, J. Shoup, id. LBH; 4 spec. Periclimenes tenuis Bruce Enewetak Islet: Feb. 1976, lagoon pinnacle ca. % to 1 mile off islet; on crinoid Comanthus bennetti, 5 m, DMD and A. Fielding, id. AJB; 7 spec. Philarius gerlachei (Nobili) Alembel Islet: July 1959, near shore. F. C. Ziesen- henne, id. LBH; 1 spec. Boken Islet: July 1957, southeast side, living coral, J. S. Garth, id. LBH; 1 spec. Enewetak Islet: April 1961, in Acropora hya- cinthus coral, E. S. Reese, id. LBH; 2 spec. (1 male, 1 female). Jinimi Islet: July and Aug. 1961, lagoon side, 4.7 m, J. Shoup, id. LBH; 2 spec. Medren Islet: July 1957, lagoon, coral, 3 to 4.7 m, J. S. Garth, id. LBH; 1 spec. Medren Islet: July 1959, lagoon side, coral, 3.6 to 4.7 m, J. Coatsworth, id. LBH; 1 spec. Philarius imperialis (Kubo) Ananij Islet: Jan. 1960, lagoon side, north end, Acropora, E. S. Reese, id. LBH; ? spec. Japtan Islet: Jan. 1960, lagoon side, north end, Acropora, E. S. Reese, id. LBH; 4 spec. Philarius ?imperialis (Kubo) Enewetak Islet: April 1961, in Acropora hyacinthus and Pocillopora damicornis, E. S. Reese, id. LBH; 9 spec. (2 lots). Enewetak Islet: July to Aug. 1961, lagoon side, 7.7 and 12 m, J. Shoup, id. LBH; 3 spec. (2 lots). Pontoniopsis comaanthi Borradaile Enewetak Islet: Feb. 1976, ocean side, on crinoid Comanthus parvicirrus, 30 m, P. Lamberson, id. AJB; 2 spec. (1 male and 1 ovig. female). Medren Islet: Feb. 1976, lagoon, ca. ¥2 mile off islet, on crinoid Comanthus bennetti, 6 to 9 m, DMD and A. Fielding, id. AJB; 2 spec. (1 male and 1 ovig. female). Stegopontonia commensalis Nobili Enewetak Islet: Aug. 1968, on sea urchin Diadema setosum, J. W. Knudsen, id. AJB; 1 spec. Family GNATHOPHYLLIDAE Gnathophyllum americanum Guerin-Meneville Enewetak Islet: July 1982, lagoon, under dead coral, 5 m, S. Johnson, id. AJB; 1 spec. (ovig.). Hymenocera picta Dana Enewetak Islet: 1982, quarry, 1 to 2 m, via color photo by S. Johnson. Family ALPHEIDAE Alpheus leviusculus Dana Japtan Islet: July 1957, seaward reef flat, rock with loose dead coral and coral sand, 0.5 m tide, J. S. Garth, id. A. H. Banner Synalpheus heroni Coutiere. Enewetak Atoll: March 1957, data destroyed in fire, id. A. H. Banner; 1 spec. Synalpheus heroni Coutiere Family HIPPOLYTIDAE Hippolyte ?ventricosa H. Milne Edwards Medren Islet: July 1957, lagoon side, algae attached to old coral head, D. J. Reish, id. LBH; 2 spec. Hippolyte sp. Medren Islet: Sept. 1956, sediment bucket suspended at deep water pier, 22 day exposure, D. J. Reish, id. LBH; 2 spec. Ligur uveae (Borradaile) Cement Ship: Sept. 1982, inside portion of submerged part of ship, at night, 2 to 5 m, S. Johnson, id. AJB; 2 spec. (ovig.). Lysmata vittata (Stimpson) Enewetak Islet: May 1946, surface, L. P. Schultz, id. LBH; 2 spec. (ovig.). Lysmata sp. Enewetak Islet: Oct. 1982, algal ridge, from rotenone station, J. Randall, id. AJB; 1 spec. (ovig.). Medren Islet: July 1957, lagoon, old coral heads from 1.5 m of water, D. J. Reish, id. LBH; 1 spec., (incomplete). Saron marmoratus (Olivier) Biken Islet: Aug. 1956, ocean side, far end of island, formalin wash from brown staghorn coral, D. J. Reish, id. LBH; 2 spec. Biken Islet: July 1959, southeast side, between ocean and lagoon, rock with sand substrate, dead coral, J. S. Garth, id. LBH; 6 spec. Enewetak Islet: July and Aug. 1957, lagoon side, 5 m, J. Shoup, id. LBH; 2 spec. Jinimi Islet: July and Aug. 1957, lagoon side, 5 m, J. Shoup, id. LBH; 2 spec. Medren Islet: Feb. 1957, lagoon side, bottom sand, from two overgrown heads of Pocillopora and one _ branching Acropora plus bases, 2.4 m, A. H. Banner, id. LBH; 2 spec. Medren Islet: July 1957, lagoon side, coral, 3 to 4.7 m, J. S. Garth, id. LBH; 2 spec. Saron neglectus De Man Ananij Islet: Jan. 1960, lagoon side, off north end, from Seriatopora hystrix, E. S. Reese, id. LBH; 3 spec. Medren Islet: July 1957, lagoon side, coral heads and dead coral, 3 to 4.7 m, J. S. Garth, id. LBH; 2 spec. Thor amboinensis (De Man) Billae Islet: Aug. 1968, on anemone Phymanthus pau- motensis, 1 to 2 m, J. W. Knudsen, id. AJB; 1 spec. Medren Islet: Feb. 1957, lagoon side, mixed coral, 2.4 m, A. H. Banner, id. LBH; 2 spec. (1 ovig.). Medren Islet: Feb. 1982, lagoon side, under dead coral, 5 m, S. Johnson, id. A. Fielding; 1 spec. Thor maldivensis De Man Medren Islet: July 1957, lagoon side, coral heads and dead coral, 3 to 4.7 m, T. Goreau and R. Neshida, id. LBH; 1 spec. (ovig.). Thor paschalis (Heller) Ananij Islet: Feb. 1957, lagoon side, Acropora, 1.8 m, A. H. Banner, id. LBH; 1 spec. Bokonbako Islet: July 1957, ocean side, old coral heads, 0.3 m, D. J. Reish, id. LBH; 2 spec. Medren Islet: Feb. 1957, lagoon side, south end, patch reef about 600 ft offshore, surrounding 232 DEVANEY AND BRUCE bottom sand, 3 dead, overgrown coral heads, Pocillopora, Acropora, Porites, broken up, 2.4 m, A. H. Banner, id. LBH; 2 spec. Medren Islet: July 1957, ocean side, old coral heads, 0.3 m, D. J. Reish, id. LBH; 2 spec. Medren Islet: July 1957, coral heads and dead coral, 3 to 5 m, T. Goreau and R. Neshida, id. LBH; 1 spec. Family PROCESSIDAE Nikoides steinii (Edmondson) Boken Islet: Sept. 1956, ocean side, opposite channel entrance, cemented coarse coral sand rock, D. J. Reish, id. LBH; 1 spec. Nikoides multispinatus Hayashi Enewetak Islet: Jan. 1982, lagoon at night, 5 m, under dead Acropora coral, S. Johnson, id. K. I. Hayashi, 2 spec. (ovig.). Processa ?japonica (De Man) Medren Islet: Lagoon side, dredging opposite middle of island, sand bottom, 2 to 3 fms. F. C. Ziesenhenne, id. LBH; 1 spec. (ovig.). Processa molaris Chace Biken Islet: Feb. 1957, ocean side, inner edge of outer reef flat, from coral, 0.5 m, A. H. Banner, id. LBH; 1 spec. (ovig.). Enjebi Islet: July 1959, shore, 0.3 m tide, F. C. Ziesenhenne, id. LBH; 1 spec. Medren Islet: July 1957, lagoon side, coral, 3 to 4.5 m, J. S. Garth, id. LBH; 1 spec. Processa neglecta Hayashi Medren Islet: June 1957, lagoon side, sand and worm tubes, 3 m, D. J. Reish, id. LBH; 2 spec. Family PANDALIDAE Miropandalus hardingi Bruce Ribewon Islet: June 1982, on branch of antipatharian (black coral) in small cave, 23 m, S. Johnson, id. AJB; 1 spec (M). Family THALASSOCARIDIDAE Thalassocaris crinata (Dana) Lagoon (11°26’N, 162°18’E): Aug. 1968, dredge, 60 m, J. Knudsen, id. AJB; 2 spec. Family CRANGONIDAE Pontophilus sp. aff. P. sabsechota Kemp Medren Islet: July 1959, lagoon side, dredging opposite middle of island, sand bottom, 3.5 to 6 m, F. C. Ziesenhenne, id. LBH; 1 spec. Vercoia gibbosa Baker Medren Islet: 1982, small lagoon pinnacle at night, L. Boucher, id. AJB; 1 spec. Family SYNAXIDAE Palinurella wieneckii (De Man) Enewetak Pinnacle: June 1981, in small cave, 10 m, coll. and id. S. Johnson; 1 spec. (carapace of molt). Family PALINURIDAE Panulirus longipes femoristriga (von Martens) Enewetak Islet: Oct. 1969, ocean side surge channel, poison station, 0 to 8 m in channel, C. A. Child, id. LBH; 3 spec. USNM. Panulirus penicillatus (Olivier) Runit Islet: Sept. 1969, outer reef flats Lithothamnion ridge, rocks, and tidepools, low to flood tide, night, A. C. Child, id. LBH; 1 spec. USNM. Family SCYLLARIDAE Parribacus antarcticus (Lund) Medren Islet: March 1976, ocean reef flat, in hollow on reef, low tide at night, Medrano, id. J. Lamberson; 1 spec. MPRL. Runit Islet: Sept. 1969, outer reef flats, Lithothamnion ridge, rocks, and tidepools, low to flood tide, at night, A. C. Child, id. LBH; 1 juv. spec. USNM. ACKNOWLEDGMENTS Many new Enewetak records herein are based on the identification made by L. B. Holthuis and duly shown in the section on collection records. Collections from Enewetak based on the Pacific Science Board investiga- tions during 1946 were made available to Holthuis by Fenner A. Chace III, (National Museum of Natural History). Additional specimens collected since the MPRL began operations in 1956 are also recorded. Collectors responsi- ble for this material include A. H. Banner, A. C. Child, J. Coatsworth, A. Fielding, J. S. Garth, T. Goreau, M. W. Johnson, S. Johnson, J. W. Knudsen, P. Lamberson, R. Neshida, R. Palumbo, E. S. Reese, D. J. Reish, L. P. Schultz, J. Shoup, S. Swerdloff, F. C. Ziesenhenne. REFERENCES Banner, A. H., 1957, Contributions to the Knowledge of the Alpheid Shrimp of the Pacific Ocean Part II, Collection from Arno Atoll, Marshall Islands, Pac. Sci., 11: 190-206. ___, and D. M. Banner, 1960, Contributions to the Knowledge of the Alpheid Shrimp of the Pacific Ocean Part V, The Indo- Pacific Members of the Genus Athanas, Pac. Sci., 14: 129-155. —_, 1961, Contributions to the Knowledge of the Alpheid Shrimp of the Pacific Ocean, Part VIII, Losses of Specimens in the Fire of the Hawaii Marine Laboratory, Pac. Sci., 16: — 235-240. ——_, 1964, Contributions to the Knowledge of the Alpheid Shrimp of the Pacific Ocean, Part IX, Collections from the Phoenix and Line Islands, Pac. Sci., 18: 83-100. —_, 1966, Contributions to the Knowledge of the Alpheid Shrimp of the Pacific Ocean, Part X, Collections from Fiji, — Tonga, and Samoa, Pac. Sci., 20: 145-188. —_, 1968, Contributions to the Knowledge of the Alpheid Shrimp of the Pacific Ocean, Part XII, Collections from the Marshall and Caroline Islands, Micronesica, 4: 261-294. Banner, D. M. and A. H. Banner, 1981, Annotated Checklist of the Alpheid Shrimp of the Red Sea and Gulf of Aden, Zool. Verhandel., No. 190; pp. 1-99. —. 1982, The Alpheid Shrimp of Australia, Part Ill, The — Remaining Alpheids, Principally the Genus Alpheus and the © Family Ogyrididae, Rec. Aust. Mus., 34: 1-358. Bowman, T. E., 1967, The Planktonic Shrimp, Lucifer chacei sp. _ nov. (Sergestidae, Luciferinae), the Pacific Twin of the Atlan- — tic Lucifer faxoni, Pac. Sci., 21: 266-271. | CRUSTACEA DECAPODA 233 __., and L. A. Abele, 1982, The Classification of the Recent | Crustacea, D. E. Bliss (Ed.), The Biology of Crustacea, Academic Press, Inc., New York and London. Bruce, A. J., 1969, Notes on Some Indo-Pacific Pontoniinae, Part XIV, Observations on Paratypton siebenrocki Balss, Crustaceana (Leiden), 17: 171-186. __, 1970, Observations on the Indo-West Pacific Species of the Genus Palaemonella Dana, 1852 (Decapoda, Pontoniinae), Crustaceana (Leiden), 19: 273-287. —_, 1974, A Synopsis of the Pontoniinid Shrimp Fauna of Cen- tral East Africa, J. Mar. Biol. Assoc. India, 16: 462-490. __., 1975, Pontonia armata H. Milne Edwards (Decapoda, Natan- tia, Pontoniinae) — a Correction, Crustaceana (Leiden), 29: 49-54. __, 1978, Periclimenes soror Nobili, a Pontoniin Shrimp New to the American Fauna, with Observations on Its Indo-West Pacific Distribution, Tethys, 8: 299-306. , 1979a, A Report on a Small Collection of Pontoniine Shrimps from Eniwetok Atoll, Crustaceana (Leiden), 5: 209-230. ——, 1979b, Ctenopontonia cyphastreophila, a New Genus and Species of Coral Associated Pontoniine Shrimp from Eniwetok Atoll, Bull. Mar. Sci., 29: 423-435. —_., 1981, Pontoniine Shrimps of Heron Island, Atoll Res. Bull. No. 245, pp. 1-33. ——, 1983, Miropandalus hardingi, New Genus, New Species, a Bizarre Commensal Pandalid Shrimp from the Marshall Islands, J. Crust. Biol. 3: 482-490. —— and D. L. Zmarzly, 1983, Periclimenes pilipes, New Species, a Crinoid Associated from Eniwetak Atoll, Marshall Islands (Crustacea: Decapoda: Pontoniinae), J. Crust. Biol., 3: 644-654. Chace, F. A., Jr., 1955, Notes on Shrimps from the Marshall Islands, Proc. USNM, 105(3349): 1-22. ___, 1976, Shrimps of the Pasiphaeid Genus Leptochela with Descriptions of Three New Species (Crustacea: Decapoda: Caridea), Smithson. Contrib. Zool., 222: 1-51. Cooke, W. J., and C. D. MacDonald, 1981, The Puerulus and Post-Puerulus of the Hawaiian Spiny Lobster Panulirus mar- ginatus, Proc. Biol. Soc. Wash., 94: 1226-1232. Ebert, T. A., and R. F. Ford, 1986, Population Ecology and Fishery Potential of the Spiny Lobster Panulirus pencillatus at Enewetak Atoll, Marshall Islands, Bull. Mar. Sci., 38: 56-67. Edmondson, C. H., 1952, Additional Central Pacific Crustaceans, Occas. Pap. Bernice P. Bishop Mus., 21: 67-86. Fankboner, P. V., 1972, On the Association Between the Pon- toniid Shrimp Achistus miersi De Man (Decapod, Palaemoni- dae) and Giant Clams (Lamellibranchia, Tridacnidae), Wasman J. Biol., 30: 35-42. Ford, R. E., T. A. Ebert, M. McCollum and D. Bartley, 1979, Population Characteristics and Fishery Potential of Spiny Lob- sters at Enewetak Atoll, MPRL Ann. Rept. FY 1978-79. George, R. W. and L. B. Holthuis, 1965, A Revision of the Indo- West Pacific Spiny Lobsters of the Panulirus japonicus Group, Zool. Verhand, No. 72, pp. 1-36. Highsmith, R. C., 1981, Coral Bioerosion at Enewetak: Agents and Dynamics, Int. Revue Gesamen. Hydrobiol. 66: 335-375. Holthuis, L. B., 1953, Enumeration of the Decapod and Stoma- topod Crustacean from Pacific Coral Islands, Atoll Res. Bull. No. 24, pp. 1-66. , and K. I. Hayashi, 1967, A New Species of Shrimp Rhynchocinetes hiatti (Crustacea, Decapoda), Annot. Zoolog. Japon., 40: 161-170. Rosewater, J., 1965, The Family Tridacnidae in the Indo-Pacific, Indo-Pacific Moll., 1: 347-393. Zmarzly, D. L., 1984, Distribution and Ecology of Shallow-Water Crinoids at Enewetak Atoll, Marshall Islands, with an Anno- tated Checklist of Their Symbionts, Pac. Sci., 38: 105-122. Chapter 23 Crustacea Decapoda (Brachyura and Anomura) of Enewetak Atoll JOHN S. GARTH,* JANET HAIG,” and JENS W. KNUDSEN+ *Allan Hancock Foundation, University of Southern California, Los Angeles, California 90089; +Pacific Lutheran University, Department of Biology Takoma, Washington 98447 INTRODUCTION A preliminary report on the Crustacea Decapoda (Bra- chyura and Anomura) of Enewetak Atoll, Marshall Islands, was published by the first author in 1964. It reported the collecting of 19 families, 81 genera, and 147 species of brachyuran and anomuran crabs, mostly at Enewetak Atoll. The report was based on collections made by Donald J. Reish in 1956 and 1957; A. H. Banner in 1957; John S. Garth and Fred C. Ziesenhenne in 1957 and 1959, with the assistance of J. Coatsworth, L. Donaldson, T. Goreau, E. Held, R. Neshida, R. Palumbo, J. Roberts, E. Ryan, and A. Smith; and R. A. Boolootian, E. S. Reese, B. Sather, J. Shoup, and R. A. Stevenson in 1960 and 1961. Only those species were listed, however, that occurred in the two specialized habitats discussed: those found in association with branch- ing corals and those obtained by dredging in the lagoon. (Reporting of Anomura was of a preliminary nature, giving number of families, genera, and species but listing only four species, two of them to genus only, from the lagoon- bottom habitat.) The overwhelming presence of the family Xanthidae was noted. These comprised 45% of the genera and 56% of the species of Brachyura encountered, a cir- cumstance attributed to the abundance and variety of corals in which many of the xanthid species reside. The present report, although still of a preliminary nature, is based additionally upon the more extensive col- lections made by Jens W. Knudsen, who visited Enewetak annually from 1965 through 1969 and again in 1971 and 1972; by Alan Havens, who visited the atoll in 1968, 1969, and 1970; and by C. Allan Child, who collected at Enewetak for the Smithsonian Institution in 1969. Smaller 235 collections used in preparing the report were those of W. A. Bartos in 1944, F. C. Ziesenhenne in 1946, G. J. Bakus and B. H. Bussing in 1965, C. V. MacCoy in 1967, S. L. Brunenmeister and E. Chave in 1974 and 1975, A. Fielding in 1976 and 1978, and P. Colin and D. M. Devaney in 1980. Of the decapod crustaceans obtained in the Marshall Islands during Operation Crossroads in 1946 and 1947, the Portunidae (Stephenson and Rees, 1967), the Xanthidae (under study by John S. Garth), and the Porcellanidae (under study by Janet Haig) from Enewetak could be included. Although the earlier report (Garth, 1964) could compare what was then known of Enewetak Brachyura and Anomura with the limited work done in the Marshall Islands by the Pacific Science Board’s Coral Atoll Survey (Holthuis, 1953), a much bet- ter comparison of the Enewetak fauna with those of other atolls of the northern Marshall Islands can now be made with reference to the earlier Operation Crossroads collec- tions. The Porcellanidae from Marshall Islands other than Enewetak are also being studied by Haig. Records of previous anomuran and brachyuran collect- ing in the Marshall Islands at atolls other than Enewetak are those of Balss (1938) from Ailinglablab, Ebon, Jaluit, Kwadelin (Kwajalein), Namu, and Namorik of the Ralik group; Likieb and Majeru (Majuro) of the Ratak group; of Miyake (1939) from the above plus Arno of the Ratak group; of Miyake (1943) (Porcellanidae only) from Jaluit; of Holthuis (1953) from Ailuk, Arno, Bikar, Jemo, Kwajalein, Lae, Likiep, Pokak, Taka, Ujae, Ujelang, Utrik, and Wotho; and of Stephenson and Rees (1967) (Portunidae only) from Bikini, Rongelap, and Rongerik. LAND CRABS The first decapod crustaceans encountered at Enewetak, as at any mid-Pacific atoll, and the only ones likely to be seen by many visitors, are the semiterrestrial and terrestrial species. These are the species that spend most of their lives at high-tide mark or above it, returning to the sea only to deposit their eggs, which require sea- water for hatching and for nurturing the larval stages. Anomuran crabs having this habit are hermit crabs of the 236 GARTH, HAIG, AND KNUDSEN family Coenobitidae, genus Coenobita, and the coconut crab, Birgus latro. Brachyuran crabs found in the terrestrial environment are grapsoid crabs of the families Gecarcini- dae, Ocypodidae, and a few Grapsidae. The spray zone on rocky shore is inhabited by the Sally Lightfoot crab, here Grapsus tenuicrustatus rather than Grapsus grapsus Linnaeus, 1758. The highest eleva- tions of sandy beaches are inhabited by the grapsid crabs Cyclograpsus and Pseudograpsus; the intermediate eleva- tions (with burrows extending below water level) are in- habited by the ghost crab, Ocypode ceratophthalma. The mole crab, Hippa pacifica, an anomuran, lives in the surf zone, burying itself quickly and emerging suddenly to grasp its food with its first pair of legs. Analogous situations on muddy beaches (rare at Enewetak) are inhabited by the ocypodid crabs Macrophthalmus and Uca. Where soil accu- mulates, as at the bases of coconut palms, the land crab Gecarcinus, family Gecarcinidae, burrows. The collector who turns inland will encounter the grapsids Geograpsus crinipes and G. grayi among leaf litter and the Metasesarma and Sesarma among roots and low branches. It was no accident that the collector responsible for most of the records of land crabs in the Pacific Science Board’s Coral Atoll Survey (Holthuis, 1953) was F. R. Fos- berg, a terrestrial botanist, or that the first crabs to reach the senior author soon after arriving at Enewetak were brought to him by Edward Held and Ralph Palumbo, members of the University of Washington group studying the effects of radiation on terrestrial organisms. Thus every habitat available to them, both terrestrial and marine, has been colonized by these ubiquitous arthropods, the decapod crustaceans. CRABS AS SYMBIONTS The relationships between crabs and the corals in which they may be found vary widely, from parasitism and commensalism (or mutualism) to facultative symbiosis. In parasitism, the crab burrows into the coral or the coral grows around the crab, in both cases enclosing it. In com- mensalism, the crab, although apparently free-living, invariably selects a living coral of a particular kind as its host. Facultative symbiosis is a relationship in which the crab, while often found in living coral, is also found in dead coral, in coral rubble, or even on a noncoral, rubbly substrate. The true parasites, long thought to include only the coral gall crabs and coral-burrowing crabs of the family Hapalocarcinidae, are now known to include some of the apparently free-living crabs of the family Xanthidae as well—in particular, the genera Trapezia and Tetralia, the former found on pocilloporid, the latter on acroporid corals. These were shown by Knudsen (1967) to feed on coral polyps, which they macerate with their specially adapted dactyls before ingesting. The genera Domecia and Cymo, different species of which occur on pocilloporid and acroporid corals, apparently have similar feeding habits, although only those of Domecia have been investigated (Patton, 1967). Other crabs found in coral apparently use it only for shelter and protection from predators. These include not only xanthid crabs like Liomera (=Carpilodes) and Pseudoliomera—some species of which rarely, if ever, occur elsewhere—but also a host of genera such as Chlorodiella, Phymodius, and Pilodius, which are found abundantly in dead coral and coral rubble. These are joined by crab genera of other families: the smaller swim- ming crabs of the genus Thalamita, family Portunidae; a number of spider crabs, including Perinea and Tylocarcinus, family Majidae; hermit crabs of the genus Calcinus, family Diogenidae; Coralliogalathea and Galathea, family Galatheidae; and porcelain crabs of the genera Pachycheles and Petrolisthes, family Porcellanidae. Small xanthid crabs of the genus Actumnus have been observed to carry a small piece of live coral as they move about the ocean bottom, much as a hermit crab carries a gastropod shell. Actumnus antelmi Ward, the species origi- nally described as having this habit, occurs at Enewetak, but its coral-carrying propensity was not observed (Lam- berts and Garth, 1977). Crabs that use not a coral polyp but an actinian coelenterate, which they carry in each claw to fend off attackers, are members of the genera Lybia and Polydectus. Several species of diogenid hermit crabs of the genus Dardanus carry anemones on their shells. In both cases, the crab benefits from the stinging nematocysts of the coelenterate, and the anemone achieves mobility and probably food as well. The porcelain crab Neopetrolisthes, family Porcellanidae, inhabits several species of large anemones. Crabs associated with living mollusks include the pin- notherid Xanthasia murigera, found in the mantle cavity of the giant clam, Tridacna gigas. Crabs associated with living echinoderms include the parthenopid Harrovia elegans, found on the crinoid Comanthus, and the galatheids Allogalathea elegans and Galathea amboinensis, also found on crinoids. The portunids Lissocarcinus orbicularis and L. holothuricola found in the respiratory tree or cloaca of the sea cucumbers Holothuria atra Jaeger and Actinopyga mauritiana (Quoy and Gaimard) proved impossible to parti- tion between their holothurian hosts. CRABS AS SUBJECTS FOR RESEARCH The Anomura and Brachyura of Enewetak Atoll have proven valuable as subjects for research. The terrestrial hermit crabs, Coenobita, were used by University of Wash- ington School of Fisheries personnel in determining re- sidual activity from tests conducted by the Atomic Energy Commission in the late 1940s and early 1950s (Held, 1960). Reese (1968a) used the coconut crab, Birgus latro, in life history studies showing use of a shell by the glau- cothoe larva. Hermit crabs, Coenobita, and the ghost crab, Ocypode ceratophthalma, were used by Reese in motion picture studies of locomotion. Experiments conducted by CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 237 Knudsen (1967) showed that the coral-inhabiting xanthid crabs, Trapezia and Tetralia, ate coral polyps after first macerating them and, hence, were true parasites rather than commensals. The coral burrowing crabs, family Hapalocarcinidae (Fize and Serene, 1957), were used by Knudsen in unpublished studies. The portunid crab, Thalamita integra, was used by Pomeroy and Kuenzler (1967) in studies of phosphorus turnover by coral reef animals. Highsmith (1981) involved the xanthid crabs, Tetralia and Maldivia, in studies of coral erosion by inver- tebrates and fishes. Intertidal crabs of the family Xanthidae were utilized by Havens (1974) in studies of competitive exclusion (the partitioning of food resources). Wenner (1977) and Wenner and Fusaro (1979) conducted studies of population structure and dynamics using the Pacific mole crab, Hippa pacifica. The xanthid crabs—Dacryopilumnus eremita, Eriphia sebana, Phymo- dius ungulatus, Pilumnus longicornis, and Trapezia speciosa—were shown by Danforth (1967, 1970) to host epicarid isopod parasites of hitherto undescribed species. The hermit crabs Dardanus were shown by Humes (1971) to be hosts of harpacticoid copepods. The hermit crabs Calcinus and Diogenes were used by Orians and King (1964) in studies on shell selection and invasion rates. Systematic studies have been published on the genus Petrolisthes, family Porcellanidae, by Haig (1981); on coral-inhabiting crabs by Garth (1964); and on swimming crabs, family Portunidae, by Stephenson and Rees (1967), with additional studies by Garth, Haig, and Knudsen in progress. POISONOUS CRABS During the 1970s Garth and Alcala (1977) showed beyond a reasonable doubt that numerous reef-inhabiting crabs of the Indo-West Pacific are harmful when eaten because they are poisonous. Included among these are several common Enewetak species: Daldorfia horrida (Lin- naeus), Atergatis floridus (Linnaeus), Eriphia sebana (Shaw and Nodder), and Zozymus aeneus (Linnaeus). The first is an elbow crab, family Parthenopidae; the remaining three are members of the family Xanthidae, as are most of the crabs found to be toxic to man and to domestic animals. The poison, a saxitoxin, is chemically indistinguishable from that produced by certain mollusks. It causes vomiting, followed by locomotory and neurological symptoms which, if not treated, result in paralysis and death. Crab-caused fatalities have been documented, and the crabs were identified by competent taxonomists in the Ryukyu Islands, the Philippines, and Palau. Native popula- tions of many South Sea island groups have traditions of killer crabs and vernacular names for the poisonous species. Although no poisonings from crabs are known to have occurred at Enewetak Atoll, crabs known to be poi- sonous elsewhere are common on the reef at Enewetak. Caution is urged in the handling of these crabs (a person who has handled such a crab might experience numbness after touching his tongue to his hand). Abstinence from their culinary use is also advised. FOSSIL CRUSTACEANS Fossil anomuran and brachyuran decapod species obtained by U. S. Geological Survey drillings at Enewetak and reported by Roberts (1964) include Callichirus armatus (A. Milne Edwards), Callichirus articulatus (Rathbun), Actaeodes hirsutissimus (Riippell), and Etisus laevimanus (Randall). Although known from elsewhere in the Indo-West Pacific, these four species have not been found living at Enewetak. This could mean either that subtle changes have occurred in the reef environment that render Enewetak no longer a suitable habitat or, as seems more likely, that the suite of species inhabiting Enewetak is changing constantly as new species are introduced, become established, are eliminated by competitors, and become locally extinct until reintroduced in another cycle. It is also possible that further and more diligent searching may yet uncover these four species at Enewetak in the living state. COLLECTING DECAPOD CRUSTACEANS’ Enewetak Atoll ascends abruptly to the surface where the North Equatorial Current, the prevailing trade winds, and the oceanic waves strike the atoll. Waves refract all the way around the atoll reef and penetrate the lagoon through channels or over the algal ridge. Therefore, we felt that every compass point, from windward to leeward, wouid have unique physical factors that influenced distribu- tion of both coral species (plus morphotypes of coral species) and decapod crustaceans. This proved useful. Since several other experts were to receive crustacean specimens beyond our interest, much time was spent hunt- ing new corals, crinoids, algae, and sediments that har- bored decapods. The intertidal zone is shallow (based on Kwajalein information), yet while zones are compressed, a centimeter elevation on the reef flat would usually yield some different brachyuran. We also believed zones directly below and above the tidal range were relatively shallow but became thicker (or deeper) down into the lagoon depth, or to the height of land and its vegetation on islands. This, in theory, kept us close to the intertidal zone where we thought the greatest species density of decapods was to be, and apparently is, found. We collected on all but three of the 34 named islands. The northwest chain from Bogallua Island to Bogon Island was considered too dangerous for camping trips as opposed to 1-day M-boat runs. Our radio carried less than 5 miles and the Equa- torial Current flowed westward. Later, the Garth—-Knudsen teams found the restricted access limited the work on the northwest chain as compared to that done on the more accessible islands. “This section was provided by Jens Knudsen. 238 GARTH, HAIG, AND KNUDSEN The windward algal ridge receives the largest waves which supersaturate incoming water with oxygen. Algae grow here in a profusion of species and mass. Initially we used pry bars to loosen slabs from the algal ridge. These were placed in plastic bags or buckets while another team member attempted to secure all free decapods. Slabs were carried to the nearest island for cracking and collecting of decapods. Subsequently, a rigid, heavy-welded pipe crack- ing table with a car-decking top was designed and built for the purpose of cracking coral or slabs at any site including the algal ridge (Fig. 1). Buckets, hammers, and other equipment were secured to the table. A marker allowed note taking (even under water), so station numbers were issued and recorded as needed. Naturally someone held the table when very large waves were running. The yield of species was dramatically increased by the use of the cracking table. Illuminated by a Coleman lantern, the reef-flat and algal ridge were also collected at night with excellent success. The algal ridge off Enewetak Island was also collected dur- ing waveless doldrums—until two large tsunamic waves came shoreward, throwing Knudsen 30 feet back onto the reef flat. Small amounts of formaldehyde were applied full strength to the reef flat at low—low tide. The preservative diluted in patches of water and entered worm burrows, evicting numerous decapods which otherwise would have been impossible to collect. Behind the reef flat, formal- dehyde was injected 6 inches deep in coral rubble. Soon afterwards, decapods that resembled chips of coral worked up to the surface and were captured with a guppy net. On our first trip, we snorkeled every day and averaged about 5 to 10 miles of swimming per day in local areas. Always new corals or new formations, new wave patterns, reef drainage currents, etc., provided new records. Plastic bags were placed over corals or crinoids, and the coral and/or crinoid was removed with decapod species intact. The author was towed by our slow moving outboard skiff to survey miles of lagoon margin. A hand signal meant new coral formation—or possible shark attack. We snorkeled down to about 60 feet in quest of some crinoids and corals. Collecting was successfully attempted with a dredge built and outfitted at Pacific Lutheran University (PLU). We used the dredge, powered by a skiff, in shallow water (to 30 feet). In deeper lagoon water, we used an M-boat in reverse, with the dredge rope and buoy ready to go over- board when fouled. Markers placed and recorded to allow work to continue the next day were never found again. However, many rare and some new records of crabs were worth the effort. A cable and winch, as opposed to pulling the cable by hand, would have greatly facilitated lagoon studies. Islands possessed many species of decapods on land and even in trees. Since islands are scattered around much of the reef, and refracting waves strike islands differently, Fig. 1 Cracking table on the algal ridge at Enewetak, about 1000 ft from land. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 239 the intertidal and island sediments had to be sampled all the way around and at all elevations (the highest being 13 feet above sea level). Several islands were studied in great detail with some new records derived. Can-traps, baited ‘traps, lantern light, etc., helped secure the nocturnal indi- viduals. Crepuscular decapods were the least collected. Two locations were found where real mud occurred, and there Uca, the fiddler crab, was collected for the first time. Some of my studies, such as the ecology and distribu- tion of brachyurans at the north end of Enewetak Island, became so large and required so much space that they were self-defeating. Because of the high class loads (18 to 22 contact hours at PLU until about 1970), research data had to be set aside so long it lost its relevancy. Despite such failings, evidence has been produced of a much richer and grander decapod fauna than any had guessed existed. Yet I am convinced another 50 species must lurk there waiting. I would like to capture them . . . personally! Students serving well at Enewetak include Jack Shan- non, M. D.; David Pearson, Ph.D., Penn State; Douglas Lambrecht, M. D.; Douglas Holt; Richard Myking, teacher; John Rankin, teacher; David Soderlund, Ph.D., Cornell University; and Erik Severeid, business overseas. Checklist of Anomura and Brachyura from Enewetak Atoll Order DECAPODA Suborder PLEOCYEMATA Infraorder ANOMURA Superfamily THALASSINOIDEA Family CALLIANASSIDAE “tCallianassa sp. *tCalliax sp., aff. novaebritanniae (Borradaile, 1900). Callichirus armatus (A. Milne Edwards, 1870). +Callianassa armata: Roberts, 1964. Callichirus articulatus (Rathbun, 1906). +Callianassa articulata: Roberts, 1964. *tCallichirus vigilax (De Man, 1916). “+ Thomassinia sp. Family CALLIANIDEIDAE “+ Callianidea [undescribed sp.]. *“tNew genus [undescribed sp. 1]. *“+New genus [undescribed sp. 2]. Family AXIIDAE “t+ Enoplometopus (Enoplometopus) sp. Enoplometopus (Holometopus) holthuisi Gordon, 1968: Holthuis, 1983. *“tNew genus [? undescribed sp.]. Superfamily HIPPOIDEA Family HIPPIDAE “tHippa adactyla Fabricius, 1787. Hippa pacifica (Dana, 1852): Wenner, 1977; Fusaro, 1978; Wenner and Fusaro, 1979; Wenner and Haley, 1981. Family ALBUNEIDAE *+Paralbunea dayriti (Serene and Umali, 1965). {Albunea ?elioti Benedict, 1904. {/Albunea sp. Superfamily PAGUROIDEA Family COENOBITIDAE Birgus latro (Linnaeus, 1767): Gross, 1964; Reese, 1968a; Reese and Kinzie, 1968; Page and Willason, 1982. Coenobita brevimanus Dana, 1852: Gross, 1964; Lawrence, 1976; Page and Willason, 1982. “ft Coenobita cavipes Stimpson, 1858. Coenobita perlatus H. Milne Edwards, 1837: Held, 1960; Gross, 1964; Reese, 1969; Lawrence, 1976; Page and Willason, 1982. Coenobita rugosus H. Milne Edwards, 1837: Lawrence, 1976; Page and Willason, 1982. Family DIOGENIDAE Aniculus aniculus (Fabricius, 1787): Reese, 1969; Forest, 1984. “ft Aniculus sp. “New Enewetak record. +New Marshall Islands record. +Fossil record. J. S. Garth manuscript lists. 240 GARTH, HAIG, AND KNUDSEN Calcinus elegans (H. Milne Edwards, 1836): Reese, 1969. Calcinus gaimardii (H. Milne Edwards, 1848): Reese, 1969. “tCalcinus guamensis Wooster, 1984. {Calcinus sp. indet. #2. “t+ Calcinus imperialis Whitelegge, 1901. Calcinus laevimanus (Randall, 1839): Reese, 1962; Reese, 1968b; Reese, 1969. Calcinus latens (Randall, 1839): Provenzano, 1963; Orians and King, 1964; Reese, 1969; Humes, 1971. Calcinus seurati Forest, 1951: Reese, 1969. “tCalcinus sp., aff. spicatus Forest, 1951. “} Clibanarius sp., aff. boschmai Buitendijk, 1937. Clibanarius corallinus (H. Milne Edwards, 1848): Reese, 1969. *Clibanarius eurysternus (Hilgendorf, 1879). “+ Clibanarius zebra rhabdodactylus Forest, 1953. 1Clibanarius zebra var. rhabdodactylus. “+ Clibanarius sp. *+ Dardanus crassimanus (H. Milne Edwards, 1836). *Dardanus deformis (H. Milne Edwards, 1836). “+ Dardanus gemmatus (H. Milne Edwards, 1848). Dardanus guttatus (Olivier, 1812): Humes, 1971. Dardanus lagopodes (Forsskal, 1775): Humes, 1971. Dardanus sanguinolentus (Quoy and Gaimard, 1824): Provenzano, 1963. {Dardanus sanguinolentus. Dardanus megistos (Herbst, 1804): Humes, 1971. Dardanus scutellatus (H. Milne Edwards, 1848): Provenzano, 1963; Orians and King, 1964; Garth, 1964; Humes, 1971. Dardanus woodmasoni (Alcock, 1905): Garth, 1964. Diogenes gardineri Alcock, 1905: Provenzano, 1963; Orians and King, 1964. I Diogenes sp. “t+ Diogenes pallescens Whitelegge, 1897. *+Paguristes sp. “} Trizopagurus strigatus (Herbst, 1804). Family PAGURIDAE “tCatapagurus sp. *“+Pagurixus anceps (Forest, 1954): McLaughlin and Haig, 1984. {Pagurus sp. (in part). {Pagurus (Pagurixus) sp. 1. Pagurixus boninensis (Melin, 1939): McLaughlin and Haig, 1984. Pagurixus maorus (Nobili, 1906): McLaughlin and Haig, 1984. 1 Pagurus sp. (in part). {Pagurus (Pagurixus) sp. 2. “+New genus, sp. Superfamily GALATHEOIDEA Family GALATHEIDAE Allogalathea elegans (Adams and White, 1848): Baba, 1977; Baba, 1979; Bruce and Zmarzly, 1983. {1Galathea elegans. “t+ Coralliogalathea humilis (Nobili, 1905). {1Galathea tridentirostris Miyake, 1953. *+Galathea aegyptiaca Paulson, 1875. “+ Galathea affinis Ortmann, 1892. 1Galathea spinosorostris Dana, 1852 (in part). {Galathea sp., aff. australiensis Stimpson, 1858. “+ Galathea amboinensis De Man, 1888. “+ Galathea sp., aff. tanegashimae Baba, 1969. {Galathea spinosorostris Dana, 1852 (in part). “+ Phylladiorhynchus serrirostris (Melin, 1939). {Galathea serrirostris. Family PORCELLANIDAE *Neopetrolisthes maculatus (H. Milne Edwards, 1837). *Pachycheles johnsoni Haig, 1965. | Pachycheles sculptus (H. Milne Edwards, 1837). “New Enewetak record. +New Marshall Islands record. J. S. Garth manuscript lists. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 241 “t+ Pachycheles pisoides (Heller, 1865). *+ Pachycheles spinipées (A. Milne Edwards, 1873). *+ Petrolisthes asiaticus (Leach, 1820). *+ Petrolisthes bispinosus Borradaile, 1900. Petrolisthes borradailei Kropp, 1984: Kropp, 1984. 1 Petrolisthes lamarckii (Leach, 1820) (in part). “+ Petrolisthes coccineus (Owen, 1839). “+ Petrolisthes decacanthus Ortmann, 1897. Petrolisthes elegans Haig, 1981: Haig, 1981. Petrolisthes fimbriatus Borradaile, 1898: Highsmith, 1981. * Petrolisthes lamarckil (Leach, 1820). “t+ Petrolisthes masakii Miyake, 1943. “+ Petrolisthes penicillatus (Heller, 1862). “+ Petrolisthes pubescens Stimpson, 1858. “+t Petrolisthes [undescribed sp. 1, R. K. Kropp MS]. “+ Petrolisthes (undescribed sp. 2, J. Haig and R. K. Kropp MS). {| Petrolisthes decacanthus Ortmann, 1897 (in part). Infraorder BRACHYURA Section DROMIACEA Superfamily DROMIOIDEA Family DROMIIDAE “+ Cryptodromia canaliculata Stimpson, 1858. *+Cryptodromia sp. Family DYNOMENIDAE Dynomene hispida Desmarest, 1825: Highsmith, 1981. “+t Dynomene pilumnoides Alcock, 1899. * Dynomiene praedator A. Milne Edwards, 1879. *Dynomene spinosa Rathbun, 1911. Section OXYSTOMATA Superfamily CALAPPOIDEA Family LEUCOSIIDAE *+ Cryptocnemus haddoni Calman, 1900. *+ Ebalia woodmasoni Alcock, 1986. Ebaliopsis erosa (A. Milne Edwards, 1873): Garth, 1964. * Heterolithadia sp. “+ Heteronucia venusta Nobili, 1906. “+ Merocryptus durand! Seréne, 1955. “+ Myra fugax coalita Hilgendorf, 1878. “+ Nucia ingens (Rathbun, 1911). “+ Nucia speciosa Dana, 1852. “t+ Oreophorus (Oreotlos) latus Borradaile, 1903. “+ Species incertae sedis. Family CALAPPIDAE “+ Calappa calappa (Linnaeus, 1758). “t+ Calappa gallus (Herbst, 1803). *Calappa hepatica (Linnaeus, 1758):+ Roberts, 1964. Section OXYRHYNCHA Superfamily MAJOIDEA Family MAJIDAE “+ Camposcia retusa Latreille, 1825. “t+ Camposcia sp. *Cyclax suborbicularis (Stimpson, 1858). “+ Huenia brevifrons Ward, 1941. Huenia proteus De Haan, 1839: Garth, 1964. “+ Hyastenus irami (Laurie, 1906). “+ Hyastenus uncifer Calman, 1900. “+ Hyastenus verrucosipes (Adams and White, 1848). “+ Hyastenus sp. “New Enewetak record. +New Marshall Islands record. +Fossil record. fJ. S. Garth manuscript lists. 242 GARTH, HAIG, AND KNUDSEN Menaethius monoceros (Latreille, 1825): Garth, 1964. “t Micippa margaritifera Henderson, 1893. Micippa philyra (Herbst, 1803): Garth, 1964. “t Micippa platypes Riippell, 1830. “t Micippa thalia (Herbst, 1803). “Tt Naxiodes spinigera Borradaile, 1903. “+ Paratymolus bituberculatus Miers, 1882. Paratymolus sexspinosus Miers, 1884: Garth, 1964. “} Parazewa bocki Balss, 1938. Perinea tumida Dana, 1851: Garth, 1964. Schizophrys aspera (H. Milne Edwards, 1834): Garth, 1964. “t Trigonothir obtusirostris Miers, 1879. “t Tylocarcinus ?gracilis Miers, 1879. Tylocarcinus styx (Herbst, 1803): Garth, 1964. Superfamily PARTHENOPOIDEA Family PARTHENOPIDAE “+ Actaeomorpha sp., nr. erosa (Miers, 1878). “+ Cryptopodia ?pan Laurie, 1905. *Daldorfia horrida (Linnaeus, 1758). “+ Daldorfia (or Parthenope) sp. Harrovia elegans De Man, 1888: Garth, 1964. “t Heterocrypta? sp. “t Parthenope (Aulacolambrus) curvispinis (Miers, 1879). “} Parthenope (Aulacolambrus) hoplonotus (Adams and White, 1848). “} Parthenope (Aulacolambrus) sp. “} Parthenope (Pseudolambrus) sp. Parthenope sp.: Garth, 1964. “} Thyrolambrus erosus (Miers, 1879). “+ Thyrolambrus sp. Section CANCRIDEA Superfamily CORYSTOIDEA Family ATELECYCLIDAE “} Kraussia integra Rathbun, 1906. “+ Kraussia sp., cf. marquesa Serene, 1972. “t Kraussia nitida Stimpson, 1858. “+ Kraussia rastripes F. Miiller, 1857. “Tt Kraussia rugulosa (Krauss, 1843). Section BRACHYRHYNCHA Superfamily PORTUNOIDEA Family PORTUNIDAE *Carupa tenuipes Dana, 1851. {Carupa laeviuscula Heller, 1861. “+ Catoptrus inaequalis (Rathbun, 1906). *Catoptrus nitidus A. Milne Edwards, 1870. *+ Catoptrus rathbunae Serene, 1965. “+ Catoptrus ?truncatifrons De Man, 1887. *Charybdis (Goniosupradens) erythrodactylus (Lamarck, 1818). “+ Coelocarcinus foliatus Edmondson, 1930. “+ Libistes villosus Rathbun, 1924. *+ Lissocarcinus holothuricola Streets, 1877. *+ Lissocarcinus orbicularis Dana, 1852. Portunus (Achelous) granulatus (H. Milne Edwards, 1834). Portunus granulatus: Garth, 1964. Portunus (Achelous) sp., nr. orbicularis (Richters, 1880). Portunus orbicularis (Richters): Garth, 1964. Portunus (Hellenus) longispinosus Stephenson and Campbell, 1959. Portunus longispinosus (Dana, 1852): Garth, 1964. Thalamita admete (Herbst, 1803): Garth, 1964; Stephenson and Rees, 1967. Thalamita bouvieri Nobili, 1906: Stephenson and Rees, 1967. “New Enewetak record. +New Marshall Islands record. J. S. Garth manuscript lists. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 243 *t Thalamita chaptalii (Audouin, 1826). “+ Thalamita coeruleipes Jacquinot, 1852. “+ Thalamita corrugata Stephenson and Rees, 1961. Thalamita cooperi (Borradaile, 1902). *}Thalamita dakini Montgomery, 1931. Thalamita medipacifica Edmondson, 1954. Thalamita gloriensis Crosnier, 1962: Stephenson and Rees, 1967. Thalamita gracilipes (A. Milne Edwards, 1873). Thalamonyx gracilipes: Garth, 1964. Thalamita integra Dana, 1852: Pomeroy and Kuenzler, 1967. *t Thalamita oculea Alcock, 1899. Thalamita picta Stimpson, 1858: Garth, 1964; Stephenson and Rees, 1967. Thalamita pilumnoides Borradaile, 1902: Garth, 1964. *Thalamita prymna (Herbst, 1803). “ft Thalamita quadrilobata Miers, 1884. *+ Thalamita sexlobata Miers, 1886. “t+ Thalamita sima H. Milne Edwards, 1834. Thalamita spiceri Edmondson, 1954: Highsmith, 1981. “+ Thalamita spinimana Dana, 1852. IThalamita danae Stimpson, 1858. “t Thalamita stimpsoni A. Milne Edwards, 1861. “+ Thalamita wakensis Edmondson, 1925. “Tt Thalamita yoronensis Sakai, 1969. *} Thalamita sp., nr. auauensis Rathbun, 1906. Thalamitoides quadridens A. Milne Edwards, 1869: Garth, 1964;+ Roberts, 1964. Superfamily XANTHOIDEA Family XANTHIDAE “+ Actaea sp., nr. bocki Odhner, 1925. “+ Actaea(?) cavipes (Dana, 1852). *+ Actaea margaritifera Odhner, 1925. “+ Actaea pulchella modesta (De Man, 1888). “+ Actaea quadriareolata Takeda and Miyake, 1968. “+ Actaea sp. * Actaeodes consobrinus (A. Milne Edwards, 1873). 1 Actaea consobrina. Actaeodes hirsutissimus (Riippell, 1830). + Actaea hirsutissima: Roberts, 1964. *+ Actumnus antelmei Ward, 1942. “+ Actumnus asper (Riippell, 1830). “+ Actumnus setifer (De Haan, 1835). {Actumnus tomentosus Dana, 1852. “+ Actumnus sp. “+ Actumnus (or Pilumnus) sp. “+ Atergatis ?dilitatus De Haan, 1835. * Atergatis floridus (Linnaeus, 1767). “+ Atergatopsis signata (Adams and White, 1848). * Banareia nobilii (Odhner, 1925). 1 Actaea nobilii. *+ Banareia parvula (Krauss, 1843). {Actaea parvula. “+ Carpilius convexus (Forsskal, 1775). “+ Carpilius maculatus (Linnaeus, 1758). *+ Chlorodiella corallicola Miyake and Takeda, 1968. “+ Chlorodiella cytherea (Dana, 1852). Chlorodiella laevissima (Dana, 1852): Garth, 1964. Chlorodiella nigra (Forsskal, 1775): Garth, 1964. °+ Cycloxanthops cavatus Rathbun, 1907. Cymo andreossyi (Audouin, 1826): Garth, 1964. *New Enewetak record. +New Marshall Islands record. +Fossil record. WJ. S. Garth manuscript lists. 244 GARTH, HAIG, AND KNUDSEN Cymo deplanatus A. Milne Edwards, 1873: Garth, 1964. Cymo melanodactylus De Haan, 1835; Garth, 1964. *Cymo quadrilobatus Miers, 1884. Dacryopilumnus eremita Nobili, 1906: Danforth, 1970, *+ Dacryopilumnus rathbunae Balss, 1932. *Daira perlata (Herbst, 1790). Domecia glabra Alcock, 1899: Garth, 1964. Domecia hispida Eydoux and Souleyet, 1842: Garth, 1964. *Eriphia scabricula Dana, 1852. Eriphia sebana (Shaw and Nodder, 1803): Reese, 1969; Danforth, 1970. J Eriphia laevimana Guérin, 1838. “+ Etisus bifrontalis (Edmondson, 1935). “+ Etisus demani Odhner, 1925. “t Etisus sp., nr. demani Odhner, 1925. “Etisus dentatus (Herbst, 1785). Etisus electra (Herbst, 1801): Garth, 1964. “t Etisus frontalis Dana, 1852. +Etisus laevimanus (Randall, 1839): Roberts, 1964. “} Etisus molokaiensis (Rathbun, 1906). “t Etisus splendidus Rathbun, 1906:+ Roberts, 1964, “tT Etisus sp. 1. “t Etisus sp. 2. “t Etisus sp. 3. Euxanthus exsculptus (Herbst, 1790): Garth, 1964. “} Euxanthus (or Hypocolpus) sp. Gaillardiellus rueppellii (Krauss, 1843). Actaea reppellii [sic]: Garth, 1964. Gaillardiellus superciliaris (Odhner, 1925). Actaea superciliaris: Garth, 1964. *Globopilumnus globosus (Dana, 1852). *t Heteropilumnus sp., cf. longipes (Stimpson, 1858). *Lachnopodus ponapensis (Rathbun, 1907). {/Paraxanthias haematostictus Ward, 1930. *+ Lachnopodus subacutus (Stimpson, 1858). *Lachnopodus tahitensis De Man, 1889. *Leptodius exaratus (H. Milne Edwards, 1834). *Leptodius gracilis (Dana, 1852). “t+ Leptodius davaoensis Ward, 1941. *“Leptodius nudipes (Dana, 1852). IXantho danae Odhner, 1925. *Leptodius sanguineus (H. Milne Edwards, 1834). *“t Leptodius waialuanus Rathbun, 1906. Liocarpilodes armiger pacificus Balss, 1938: Garth, 1964; Highsmith, 1981. Liocarpilodes biunguis (Rathbun, 1906): Highsmith, 1981. Zozymodes biunguis: Garth, 1964. { Xanthodius biunguis. Liocarpilodes integerrimus (Dana, 1852): Garth, 1964. Liocarpilodes pumilus (Jacquinot, 1852): Garth, 1964. {Zozymodes pumilus. 1 Zozymodes cristatus (Borradaile, 1902). Liomera bella (Dana, 1852). Carpilodes bellus: Garth, 1964; Highsmith, 1981. Liomera coelata (Odhner, 1825). Carpilodes coelatus: Garth, 1964. *+Liomera loevis (A. Milne Edwards, 1873). ICarpilodes loevis. *tLiomera monticulosa (A. Milne Edwards, 1873). |Carpilodes monticulosus. “New Enewetak record. +New Marshall Islands record. }Fossil record. TJ. S. Garth manuscript lists. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 245 *Liomera pallida (Borradaile, 1900). 1Carpilodes pallidus. *+ Liomera rugata (H. Milne Edwards, 1834). 1Carpilodes rugatus. “+ Liomera stimpsoni (A. Milne Edwards, 1865). {Carpilodes stimpsoni. *Liomera tristis (Dana, 1852). 1Carpilodes tristis. “+ Liomera sp. 1Carpilodes sp. *t Lophozozymus dodone (Herbst, 1801). *T Lophozozymus incisus (H. Milne Edwards, 1834). *Lophozozymus pulchellus A. Milne Edwards, 1867. “+ Lybia caestifera (Alcock, 1898). “+ Lybia tessellata (Latreille, 1812). * Lydia annulipes (H. Milne Edwards, 1834). *+ Macromedaeus nudipes (A. Milne Edwards, 1867). 1 Xantho nudipes. Maldivia palmyrensis Rathbun, 1923: Highsmith, 1981. Maldivia triunguiculata (Borradaile, 1902): Highsmith, 1981. “+ Medaeus elegans A. Milne Edwards, 1867. *+ Medaeus ornatus Dana, 1852. *+ Neoxanthias impressus (Lamarck, 1818). {1 Xantho impressus. “+ Paractaea retusa (Nobili, 1905). 1 Actaea retusa. Paractaea rufopunctata (H. Milne Edwards, 1834). Actaea rufopunctata: Garth, 1964. *+ Paractaea rufopunctata f. plumosa Guinot, 1969. *+ Paractaea tumulosa (Odhner, 1925). 1 Actaea tumulosa. *+ Paramedaeus simplex (A. Milne Edwards, 1873). 1 Medaeus simplex. *+ Parapilumnus coralliophilus Takeda and Miyake, 1968. Parapilumnus verrucosipes (Stimpson): Garth, 1964. “+ Parapilumnus ?incertus Takeda and Miyake, 1969. 1 Heteropilumnus sp., nr. quadrispinosus (Zehntner, 1894). Paraxanthias notatus (Dana, 1852): Garth, 1964; Highsmith, 1981. *+ Paraxanthias pachydactylus (A. Milne Edwards, 1873). *Phymodius ?granulatus (Targioni-Tozzetti, 1877). *Phymodius laysani Rathbun, 1906. *+ Phymodius monticulosus (Dana, 1852). Phymodius nitidus (Dana, 1852): Garth, 1964. Phymodius ungulatus (H. Milne Edwards, 1834): Garth, 1964; Danforth, 1967, 1970. Pilodius areolatus (H. Milne Edwards, 1834): Highsmith, 1981. Chlorodopsis areolata: Garth, 1964. Pilodius flavus Rathbun, 1906: Garth, 1964. “+ Pilodius melanodactylus (A. Milne Edwards, 1873). 1Chlorodopsis melanodactylus. Pilodius pilumnoides (White, 1847): Garth, 1964. .Chlorodopsis pilumnoides. Pilodius pugil Dana, 1852: Garth, 1964. +Chlorodopsis pugil: Roberts, 1964. *+ Pilodius scabriculus Dana, 1852. Pilodius spinipes Heller, 1861: Garth, 1964. 1Chlorodopsis spinipes. *+ Pilumnus andersoni De Man, 1887. *Pilumnus caerulescens A. Milne Edwards, 1873. “New Enewetak record. +New Marshall Islands record. +Fossil record. J. S. Garth manuscript lists. 246 GARTH, HAIG, AND KNUDSEN “+ Pilumnus ?elegans De Man, 1888. Pilumnus longicornis Hilgendorf, 1878: Garth, 1964. Pilumnus sp.: Danforth, 1970. “+ Pilumnus ransoni Forest and Guinot, 1961. “+ Pilumnus rotumanus Borradaile, 1900. * Pilumnus tahitensis De Man, 1890. *Pilumnus vespertilio (Fabricius, 1793). “+ Pilumnus sp. Planopilumnus vermiculatus (A. Milne Edwards, 1873): Garth, 1974. Polydectus cupulifer (Latreille, 1825): Garth, 1964. “} Pseudoliomera granosimanus (A. Milne Edwards, 1865). “t Pseudoliomera helleri (A. Milne Edwards, 1865). 1 Actaea helleri. *t+Pseudoliomera sp., nr. helleri (A. Milne Edwards, 1865). \Actaea sp., nr. helleri. *Pseudoliomera lata (Borradaile, 1902). 1Actaea lata. “t Pseudoliomera sp., nr. lata (Borradaile, 1902). VActaea sp., nr. lata. “t Pseudoliomera rueppellioides (Odhner, 1925). 1 Actaea rueppellioides. Pseudoliomera speciosa (Dana, 1852). Actaea speciosa: Garth, 1964. * Pseudozius caystrus (Adams and White, 1848). * Pseudozius pacificus Balss, 1938. “t+ Ralumia dahli Balss, 1933. Tetralia glaberrima (Herbst, 1799): Garth, 1964; Knudsen, 1967. *Tetralia glaberrima rubridactylus Patton, 1966. Tetraloides nigrifrons (Dana, 1852). Tetralia heterodacytyla Heller, 1861: Garth, 1964; Knudsen, 1967. {Tetralia heterodacytyla fusca Serene, 1959. {Tetralia ?nigrifrons Dana, 1852. Trapezia cymodoce (Herbst, 1801): Garth, 1964; Knudsen, 1967. “t Trapezia dentata Macleay, 1838. Trapezia digitalis Latreille, 1825: Garth, 1964; Knudsen, 1967. “+ Trapezia digitalis bella Dana, 1852. Trapezia sp., digitalis group: Garth, 1964. Trapezia ferruginea Latreille, 1825: Garth, 1964; Knudsen, 1967. “t Trapezia guttata Riippell, 1830. Trapezia rufopunctata (Herbst, 1799): Garth, 1964; Knudsen, 1967. “} Trapezia rufopunctata flavopunctata Eydoux and Souleyet, 1841. “t Trapezia rufopunctata maculata Macleay, 1838. Trapezia speciosa Dana, 1852: Garth, 1964; Danforth, 1970. Trapezia tigrina Eydoux and Souleyet, 1842. Trapezia danai Ward, 1939: Garth, 1964; Knudsen, 1967. Trapezia wardi Serene, 1970. al Trapezia sp. 1. : Trapezia sp. 2. a Xanthias canaliculatus Rathbun, 1906. T Xanthias gilbertensis Balss, 1938. ., Xanthias lamarcki (H. Milne Edwards, 1834): Highsmith, 1981. “T Xanthias lividus Lamarck, 1808. rl Xanthias punctatus (H. Milne Edwards, 1834). a Xantho sp. a Zozymodes cavipes (Dana, 1852). t Zozymus actaeoides (A. Milne Edwards, 1867). {| Platypodia actaeoides. Zozymus sp., nr. actaeoides (A. Milne Edwards, 1867). {|Platypodia sp., nr. actaeoides. “New Enewetak record. +New Marshall Islands record. J. S. Garth manuscript lists. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) *Zozymus aeneus (Linnaeus, 1758). “+ Zozymus gemmula Dana, 1852. “+ Zozymus hawaiiensis (Rathbun, 1907). 1 Platypodia hawaiiensis. “t+ Zozymus kuekenthali De Man, 1902. Family GONEPLACIDAE “+ Ceratoplax sp. “+ Genus and species incertae sedis. Family PALICIDAE *+ Palicus jukesi (White, 1847). *+ Palicus whitei (Miers, 1879). *+ Palicus sp., nr. oahuensis Rathbun, 1906. Superfamily GRAPSOIDEA Family GRAPSIDAE * Cyclograpsus integer H. Milne Edwards, 1837. 1. Cyclograpsus parvulus De Man, 1896. *+ Cyclograpsus longipes Stimpson, 1858. *+ Cyclograpsus sanctaecrucis Griffin, 1968. Geograpsus crinipes (Dana, 1851): Page and Willason, 1982. *Geograpsus grayi (H. Milne Edwards, 1853). *+ Grapsus intermedius De Man, 1888. *Grapsus longitarsus Dana, 1851. *Grapsus tenuicrustatus (Herbst, 1783): Page and Willason, 1982. * Metasesarma rousseauxi H. Milne Edwards, 1853. * Metopograpsus thukuhar (Owen, 1839). Pachygrapsus minutus A. Milne Edwards, 1873: Highsmith, 1981. “+ Pachygrapsus planifrons De Man, 1888. * Pachygrapsus plicatus H. Milne Edwards, 1837. *Percnon abbreviatum (Dana, 1851). *+Percnon pilimanus (A. Milne Edwards, 1873). Percnon planissimum (Herbst, 1804): Highsmith, 1981. “+ Plagusia depressa tuberculata Lamarck, 1818. “+ Plagusia immaculata Lamarck, 1818. *+ Plagusia speciosa Dana, 1852. *Pseudograpsus albus Stimpson, 1858. “+ Sesarma (Holometopus) sp. Family GECARCINIDAE *Gecarcoidea lalandii H. Milne Edwards, 1837. Superfamily PINNOTHEROIDEA Family PINNOTHERIDAE * Xanthasia murigera White, 1846. Superfamily OCYPODOIDEA Family OCYPODIDAE “+ Macrophthalmus (Macrophthalmus) telescopicus (Owen, 1839) var. *Macrophthalmus (Mopsocarcinus) bosci Audouin and Savigny, 1825. Ocypode ceratophthalma (Pallas, 1872): Page and Willason, 1982. Ocypode cordimana Desmarest, 1825: Page and Willason, 1982. “+ Paracleistostoma (or Cleistostoma) sp. *t Uca tetragonon (Herbst, 1790). *+ Genus and species incertae sedis. Superfamily HAPALOCARCINOIDEA Family HAPALOCARCINIDAE “+ Cryptochirus coralliodytes Heller, 1861. “+ Hapalocarcinus marsupialis Stimpson, 1858. “t+ Neotroglocarcinus dawydoffi (Fize and Serene, 1955). Troglocarcinus viridis Hiro: Garth, 1964. Pseudocryptochirus viridis (Hiro): Garth and Hopkins, 1968. “+ Pseudocryptochirus crescentus (Edmondson, 1925). “t Species 1, incertae sedis. “Tt Species 2, inceriae sedis. “New Enewetak record. +New Marshall Islands record. 247 248 GARTH, HAIG, AND KNUDSEN SUMMARY The Anomura presently known from Enewetak Atoll comprise 76 species, representing 29 genera and 10 fami- lies. Of this number, 48 species are new to Enewetak, and 43 are new to the Marshall Islands as well. The family Diogenidae is best represented, with 27 species in seven genera; the family Porcellanidae has 17 species in three genera. Expressed in percentages, of the 76 species listed, 63.15% are new to Enewetak, and 56.57% are new to the Marshall Islands. The Diogenidae contain 35.52% of the species and 24.13% of the genera; the Porcellanidae con- tain 22.36% of the species and 10.34% of the genera. The Brachyura presently known from Enewetak Atoll comprise 291 species, representing 115 genera and 16 families. Of this number, 218 species are new to Enewetak, and 170 are new to the Marshall Islands as well. The family Xanthidae is best represented, with 155 species in 49 genera; the family Portunidae by 36 species in nine genera; the family Majidae by 23 species in 13 gen- era; and the family Grapsidae by 21 species in 10 genera. Expressed in percentages, 74.91% of the 291 species are new to Enewetak and 58.42% are new to the Marshall Islands as well. The Xanthidae contain 53.26% of the spe- cies and 42.60% of the genera; the Portunidae contain 12.37% of the species but only 7.82% of the genera; the Majidae contains 7.90% of the species and 11.3% of the genera; and the Grapsidae contain 7.22% of the species and 8.70% of the genera reported. (The number of species and genera new to Enewetak and the Marshall Islands would be even greater had not many been reported in pub- lications in which the crabs, often identified by the first writer, were not the primary interest of the investigator who reported them but incidental as the hosts of isopod or copepod parasites or as agents of bioerosion of corals.) Because so large a number of brachyuran crabs remain identified to genus (35) or even to family (3) only, no meaningful comparison with other crab faunas is possible at this time. This shortcoming will be rectified when the Enewetak crabs are elaborated a family at a time and the new or obscure species described and illustrated. The defi- ciency is particularly apparent in the family Parthenopidae, of which no comprehensive review has been made since the Siboga report (Flipse, 1930). ACKNOWLEDGMENTS The generous assistance in the identification of rare or obscure species of Brachyura by specialists who visited the Hancock Foundation while the first author’s studies were in progress, or to whom specimens were later sent, is gratefully acknowledged. These include R. S. K. Barnes, the genus Macrophthalmus; I. Gordon, the single pinnoth- erid; D. J. G. Griffin, the genus Cyclograpsus and the fam- ily Majidae; M. Takeda, the Pilumninae; T. Sakai, the Oxystomata; R. Serene, the Trapeziinae; and W. Stephen- son, the Portunidae. The second author’s studies on the Anomura were facilitated by Susan L. Brunenmeister, who provided information on her Enewetak collections; Patsy A. McLaughlin, who sent a portion of this collection for the verification of a few identifications; and Philip Helfrich, who provided a copy of the Brunenmeister report (Brunen- meister, 1975). For the thalassinoids, Michele de Saint Laurent and L. B. Holthuis helped with identifications and R. B. Manning with information. The authors are most grateful to the late Dennis M. Devaney and to Ann Fielding for providing information concerning Enewetak specimens in Bishop Museum collections, and most of all, to the many collectors whose records have been used with their permis- sion in cases where it was possible to contact them. REFERENCES Baba, K., 1977, Biological Results of the Snellius Expedition XXVIII, The Galatheid Crustacea of the Snellius Expedition, Zool. Meded. (Leiden), 50: 243-259. __, 1979, Expédition Rumphius II (1975), Crustacés Parasites, Commensaux, etc., (Th. Monod et R. Serene, Eds.), VII, Galatheid Crustaceans (Decapoda, Anomura), Bull. Mus. Natl. Hist. Nat. 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Forest, J., 1984, Revision du Genre Aniculus: Decapoda Diogeni- dae, Crustacea, Crustaceana (Leiden) Suppl., No. 8, pp. ix- 91, frontispiece. Fusaro, C., 1978, Food Availability and Egg Production: a Field Experiment with Hippa pacifica Dana (Decapoda; Hippidae), Pac. Sci., 32: 17-23. Galil, B., 1985, Tetraloides, A New Genus of Coral-Inhabiting Crabs, Crustaceana (Leiden), 50: 68-77. __., and C. Lewinsohn, 1984, On the Taxonomic Status of Trapezia tigrina Eydoux and Souleyet, 1842 (Decapoda, Brachyura), Crustaceana (Leiden), 46: 166-175. Garth, J. S., 1964, The Crustacea Decapoda (Brachyura and Anomura) of Eniwetok Atoll, Marshall Islands, with Special Reference to the Obligate Commensals of Branching Corals, Micronesica, 1: 137-144. __, and A. Alcala, 1977, Poisonous Crabs of Indo-West Pacific Coral Reefs, with Special Reference to the Genus Demania Laurie, Proceedings of the Third International Coral Reef Sym- posium, Miami, Florida, 1977, 1: 645-651. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 249 __, and T. S. Hopkins, 1968, Pseudocryptochirus crescentus (Edmondson), a Second Crab of the Corallicolous Family Hapalocarcinidae (Crustacea, Decapoda) from the Eastern Pacific with Remarks on Phragmosis, Host Specificity, and Distribution, Bull. South, Calif, Acad, Sci., 67: 40-48. Gross, W., 1964, Water Balance in Anomuran Land Crabs on a Dry Atoll, Biol, Bull., 126: 54-68. Haig, J., 1981, Three New Species of Petrolisthes (Decapoda, Anomura, Porcellanidae) from the Indo-West Pacific, J. Crust. Biol., 1: 265-271. Havens, A. D., 1974, The Ecology of Eight Species of Intertidal Crabs of the Family Xanthidae in the Marshall Islands, Ph.D. dissertation, University of Southern California, Los Angeles. Held, E. E., 1960, Land Crabs and Fission Products at Eniwetok Atoll, Pac. Sci., 14: 18-27. Highsmith, R. C., 1981, Coral Bioerosion at Enewetak: Agents and Dynamics, Int. Revue Gesamten. Hydrobiol., 66: 335-375. Holthuis, L. B., 1953, Enumeration of the Decapod and Stomato- pod Crustacea from Pacific Coral Islands, Atoll Res. Bull. No. 24. ___, 1983, Notes on the Genus Enoplometopus, with Descriptions of a New Subgenus and Two New Species (Crustacea Decapoda, Axiidae), Zool, Meded. (Leiden), 56; 281-298. Humes, A. G., 1971, Sunaristes (Copepoda, Harpacticoida) Asso- ciated with Hermit Crabs at Eniwetok Atoll, Pac. Sci., 25: 529-532. Knudsen, J. W., 1967, Trapezia and Tetralia (Decapoda, Brachyura, Xanthidae) as Obligate Ectoparasites of Pocillo- porid and Acroporid Corals, Pac. Sci., 21: 51-57. Kropp, R. K., 1984, Three New Species of Porcellanidae (Crus- tacea; Anomura) from the Mariana Islands and a Discussion of Borradaile’s Petrolisthes lamarckii Complex, Micronesica, 19: 91-106. Lamberts, A. E., and J. S. Garth, 1977, Coral-Crab Commensal- ism in Xanthids, Pac. Sci., 31: 245-247. Lawrence, J. M., 1976, Organic Composition and Energy Content of the Hepatopancreas of Hermit Crabs (Coenobita) from Eniwetok Atoll, Marshall Islands (Decapoda, Paguridea), Crustaceana, 31: 113-118. McLaughlin, P. A., and J. Haig, 1984, A Review of Pagurixus (Decapoda, Anomura, Paguridae) and Descriptions of New Species, Crustaceana (Leiden), 47: 121-148. Miyake, S., 1939, Notes on Crustacea Brachyura Collected by Professor Teiso Esaki’s Micronesia Expeditions of 1937-1938 Together with a Check List of Micronesian Brachyura, Rec. Oceanogr. Works Jpn., 10: 168-247. —, 1943, Studies on the CrabShaped Anomura of Nippon and Adjacent Waters, J. Dep. Agric. Kyushu Imp. Univ., 7: 49-158. Orians, G. H., and C. E. King, 1964, Shell Selection and Invasion Rates of Some Pacific Hermit Crabs, Pac. Sci., 18: 297-306. Page, H. M., and S. W. Willason, 1982, Distribution Patterns of Terrestrial Hermit Crabs at Enewetak Atoll, Marshall Islands, Pac. Sci., 36: 107-117. Patton, W. K., 1967, Studies on Domecia acanthophora, a Com- mensal Crab from Puerto Rico, with Particular Reference to Modifications of the Coral Host and Feeding Habits, Biol. Bull., 132: 56-67. Pomeroy, L. R., and E. T. Kuensler, 1967, Phosphorus Turnover by Coral Reef Animals, Symposium on Radioecology, D. J. Nelson and F. C. Evans (Eds.), Ann Arbor, Michigan, pp. 474-482. Provenzano, A. J., Jdr., 1963, The Glaucothoes of Petrochirus diogenes (L.) and Two Species of Dardanus (Decapoda: Diogenidae), Bull. Mar. Sci. Gulf Caribb., 13: 242-261. Reese, E. S., 1962, Submissive Posture as an Adaptation to Aggressive Behavior in Hermit Crabs, Z. Tierpsychol., 19: 645-651. __, 1968a, Shell Use: an Adaptation for Emigration from the Sea by the Coconut Crab, Science, 161: 385-386. —_, 1968b, Annual Breeding Seasons of Three Sympatric Species of Tropical Intertidal Hermit Crabs, with a Discussion of Factors Controlling Breeding, J. Exp. Mar. Biol. Ecol., 2: 308-318. ——_, 1969, Behavioral Adaptations of Intertidal Hermit Crabs, Am. Zool., 9: 343-355. __, and R. A. Kinzie Ill, 1968, The Larval Development of the Coconut or Robber Crab Birgus latro (L.) in the Laboratory (Anomura, Paguridea). Crustaceana, Suppl. 2: 117-144. Roberts, H. B., 1964, Fossil Decapod Crustaceans from the Marshall Islands, Geol. Surv. Prof. Pap. 260-HH: 1127-1131. Stephenson, W., and M. Rees, 1967, Some Portunid Crabs from the Pacific and Indian Oceans in the Collections of the Smith- sonian Institution, Proc. U. S. Natl. Mus., 120: 1-114. Wenner, A. M., 1977, Food Supply, Feeding Habits, and Egg Production in Pacific Mole Crabs (Hippa pacifica Dana), Pac. Sci., 31: 39-47. —, and C. Fusaro, 1979, An Analysis of Population Structure in Pacific Mole Crabs (Hippa pacifica Dana), Biol. Bull. (Woods Hole), 157: 205-229. —_, and S. R. Haley, 1981, On the Question of Sex Reversal in Mole Crabs (Crustacea, Hippidae), J. Crust. Biol., 1: 506-517. Wooster, D. S., 1984, The Genus Calcinus (Paguridea, Diogeni- dae) from the Mariana Islands Including Three New Species, Micronesica, 18: 121-162. Appendix COLLECTION RECORDS Anomuran Crustacea Taxa are listed in the same order as in the checklist under their respective families. Higher categories may be found in the checklist. For most species reported below, the records are abbreviated, giving only the islet, year, and collector; following these records is a brief statement on habitat. Where only one or two records are available for a species, more detailed information is included for each. A summary of collectors, years of collection, identifiers, and deposition of material is given in Table 1. Family CALLIANASSIDAE Callianassa sp. Enewetak: 1980, collected from the lagoon by airlift, P. Colin and D. M. Devaney. Calliax sp., aff. novaebritanniae (Borradaile) Enewetak: 1980, collected from the lagoon by airlift, P. Colin and D. M. Devaney. Callichirus vigilax (De Man) Enewetak: 1980, collected from the lagoon by airlift, P. Colin and D. M. Devaney. 250 GARTH, HAIG, AND KNUDSEN Thomassinia sp. Enewetak: 1980, collected from the lagoon by airlift, P. Colin and D. M. Devaney Family CALLIANIDEIDAE Callianidea [undescribed sp.] Engebi: 1959, shore collecting, +0.5-foot tide, from coral, F. C. Ziesenhenne. Enyu: 1959, shore collecting at north end of island, 0.8-foot tide, in or under coral, F. C. Ziesenhenne. New genus [undescribed sp. 1] Off Rigili: 1957, ocean reef flat, inner edge about 100 feet out, from Acropora or Pocillopora in 1.5 feet of water, A. H. Banner. New genus [undescribed sp. 2] Engebi: 1959, shore collecting, +0.5-foot tide, from coral, F. C. Ziesenhenne. Family AXIIDAE Enoplometopus (Enoplometopus) sp. Enewetak: 1969, in surge channel, 0 to 3 feet, C. E. Dawson. Enewetak: 1969, in surge channel, 2 to 10 feet, C. A. Child. Enoplometopus (Holometopus) holthuisi Gordon Enewetak: 1981, pinnacle, 10 m in small cave, frag- ments of both claws, S. Johnson. Enoplometopus sp. Off Enewetak: 1946, juvenile in E. longirostris stage, L. P. Schultz. Rigili: 1946, 2 miles south, leeward side of reef, light at night, juvenile in E. longirostris stage, L. P. Schultz. New genus [?undescribed sp.] Aaraanbiru: 1959, shore collecting, +0.6-foot tide, from coral, F. C. Ziesenhenne. Family HIPPIDAE Hippa adactyla Fabricius Enewetak: 1975, lagoon side southwest end of islet, 10 to 20 m northeast cargo pier, in sand at water’s edge, C. Fusaro. Hippa pacifica (Dana) Bogallua: 1968, J. W. Knudsen. Enewetak: 1968, A. Havens. Igurin: 1967, C. V. MacCoy. Rigili: 1956, student collector. On sandy beach, and in coarse sand and gravel on reef. Family ALBUNEIDAE Paralbunea dayriti (Serene and Umaali) Enewetak: 1966, dredged in lagoon at about 11°21.5’N, 162°20’E in 45 m of water, J. W. Knudsen. Enewetak: 1980, collected from the lagoon by airlift, P. Colin and D. M. Devaney. Family COENOBITIDAE Birgus latro (Linnaeus) Igurin: 1960, E. S. Reese and R. A. Boolootian. Mui: 1957, L. Donaldson and E. Held. Rigili: 1957, J. S. Garth. On land. Coenobita brevimanus Dana Aaraanbiru: 1957, J. S. Garth. Igurin: 1960, E. S. Reese and R. A. Boolootian. Mui: 1957, L. Donaldson and E. Held. Rigili: 1957, J. S. Garth. On land. Coenobita cavipes Stimpson Igurin: 1960, E. S. Reese and R. A. Boolootian. Mui: 1957, L. Donaldson and E. Held. Mui: 1966, J. W. Knudsen. Rigili: 1960, B. Sather and R. Stevenson. Inland and on beach. Coenobita perlatus H. Milne Edwards Aomon, Bijile, and Rojoa: 1959, F. C. Ziesenhenne. Chinimi: 1965, G. Bakus. Igurin: 1956, student collec- tor. Igurin: 1959, A. Smith and J. Coatsworth. Igurin: 1960, E. S. Reese and R. A. Boolootian. Medren: 1960, E. S. Reese and R. A. Boolootian. Medren: 1965, G. Bakus. Muzin, Kirinian, and Bokonaarappu: 1959, F. C. Ziesenhenne. Rigili: 1956, student collector. Rigili: 1957, J. S. Garth. Rigili: 1965, G. Bakus. Inland and on beach; on sandy bottom, outer reef flat; dredged in lagoon, 1.8 to 7.2 m. Coenobita rugosus H. Milne Edwards Aaraanbiru: 1959, F. C. Ziesenhenne. Aniyaanii: 1956, student collector. Aomon, Bijile, and Rojoa: 1959, F. C. Ziesenhenne. Igurin: 1959, A. Smith and J. Coatsworth. Igurin: 1960, E. S. Reese and R. A. Boolootian. Medren: 1959, F. C. Ziesenhenne. Medren: 1960, E. S. Reese. Muzin, Kirinian, and Bokonaarappu: 1959, F. C. Ziesenhenne. Rigili: 1956, student collectors. Inland and on beach; lagoon and seaward sides; on sandy bottom to 7.2 m. Family DIOGENIDAE Aniculus aniculus (Fabricius) Aaraanbiru: 1959, F. C. Ziesenhenne. Enewetak: 1967, J. W. Knudsen. Between Enewetak and Bokandretok: 1966, J. W. Knudsen. Igurin: 1960, E. S. Reese and R. A. Boolootian. On reef flat and algal ridge. Aniculus sp. Off Bokandretok: 1974, 2.4 to 3 m on sandy bottom, S. L. Brunenmeister. Calcinus elegans (H. Milne Edwards) Aniyaanii: 1957, J. S. Garth. Bogen: 1957, J. S. Garth. Enewetak: 1961, E. S. Reese. Enewetak: 1965, 1967, J. W. Knudsen. Between Enewetak and Bokandretok: 1966, J. W. Knudsen. Engebi: 1957, J. S. Garth. Engebi: 1959, F. C. Ziesenhenne. Igurin: 1957, J. S. Garth. Medren: 1959, J. S. Garth. Medren: 1960, E. S. Reese. Medren: 1974 or 1975, S. L. Brunenmeister. Rigili: 1957, 1959, J. S. Garth. Runit: 1959, J. S. Garth. Enewetak Atoll, no further locality data: 1946, F. C. Ziesenhenne. Lagoon side of reef; seaward reef flat and algal ridge. Under rocks and dead coral, on dead coral heads, on live Acropora and Pocillopora. Calcinus gaimardii (H. Milne Edwards) Enewetak: 1968, J. W. Knudsen. Igurin: 1957, J. S. Garth. Medren: 1959, J. Roberts and/or F. C. Ziesen- henne. Rigili: 1956, student collector. On reef and on sandy beach; under and around Porites colonies. Calcinus guamensis Wooster Enewetak: 1966, 1967, J. W. Knudsen. Between Enewetak and Bokandretok: 1966, J. W. Knudsen. On reef, from Porites, Favia, live Acropora, and live and dead Pocillopora. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 251 Calcinus imperialis Whitelegge Enewetak: 1967, J. W. Knudsen. Between Enewetak and Bokandretok: 1966, J. W. Knudsen. Reef flat and algal ridge, from Acropora and Pocillopora corals. Calcinus laevimanus (Randall) Aniyaanii: 1956, student collectors. Aniyaanii: 1957, J. S. Garth. Bokandretok: 1967, J. W. Knudsen. Enewetak: 1961, E. S. Reese. Enewetak: 1965, 1967, 1968, J. W. Knudsen. Between Enewetak and Bokan- dretok: 1966, J. W. Knudsen. Engebi: 1956, student collector. Engebi: 1957, 1959, J. S. Garth. Engebi: 1967, J. W. Knudsen. Igurin: 1957, 1959, J. S. Garth. Medren: 1956, student collectors. Medren: 1957, 1959, J. S. Garth. Medren: 1960, E. S. Reese. Medren: 1974 or 1975, S. L. Brunenmeister. Muti: 1965, G. Bakus. Muti: 1967, J. W. Knudsen. Rigili: 1959, J. S. Garth. Runit: 1959, J. S. Garth. Enewetak Atoll, no further locality data: 1946, F. C. Ziesenhenne. Seaward reef flat and beach on lagoon side; under rock, coral rubble, and dead corals, and in live Porites colonies. Calcinus latens (Randall) Aniyaanii: 1966, J. W. Knudsen. Aomon, Bijile, and Rojoa: 1959, F. C. Ziesenhenne. Bogallua: 1968, J. W. Knudsen. Bokandretok: 1967, J. W. Knudsen. Enewetak: 1961, E. S. Reese. Enewetak: 1966 to 1968, J. W. Knudsen. Between Enewetak and Bokan- dretok: 1966, J. W. Knudsen. Engebi: 1959, F. C. Ziesenhenne. Engebi: 1967, J. W. Knudsen. Igu- rin: 1959, J. S. Garth. Medren: 1957, A. H. Banner. Medren: 1957, T. Goreau. Medren: 1957, 1959, J. S. Garth. Medren: 1959, J. Coatsworth. Medren: 1960, E. S. Reese. Medren: 1974 or 1975, S. L. Brunenmeister. Between Medren and Muti: 1960, E. S. Reese. Muti: 1966, J. W. Knudsen. Rigili: 1959, J. S. Garth. Rojoa: 1966 to 1968, J. W. Knudsen. Runit: 1959, J. S. Garth. Sandy bottom and on corals in lagoon, to 9 m; seaward reef, from live and dead Acropora and Porites corals. Calcinus seurati Forest Enewetak: 1961, E. S. Reese. Medren: 1956, student collector. Medren: 1957, 1959, J. S. Garth. Medren: 1960, E. S. Reese. Medren: 1974 or 1975, S. L. Brunenmeister. Muzin, Kirinian, and Bokonaarappu: 1959, F. C. Ziesenhenne. On reef. Calcinus sp., aff. spicatus Forest Enewetak: 1967, from Porites, J. W. Knudsen. Rojoa: 1968, ocean reef three-fourths way out, green algal turf and burrows, J. W. Knudsen. Clibanarius sp., aff. boschmai Buitendijk Enewetak: 1975, collected intertidally from crevice in the benchrock at lagoon margin, northern end of island, S. L. Brunenmeister. Clibanarius corallinus (H. Milne Edwards) Aaraanbiru: 1959, F. C. Ziesenhenne. Aniyaanii: 1956, student collector. Aniyaanii: 1957, J. S. Garth. Enewetak: 1961, E. S. Reese. Enewetak: 1965, 1967, 1968, J. W. Knudsen. Enewetak: 1974, 1975, S. L. Brunenmeister. Engebi: 1959, F. C. Ziesenhenne. Igurin: 1959, J. S. Garth. Medren: 1959, J. S. Garth. Medren: 1960, E. S. Reese. Between Medren and Muti: 1960, E. S. Reese. Rigili: 1959, J. S. Garth. Runit: 1959, J. S. Garth. Intertidal on reef flat. Clibanarius eurysternus (Hilgendorf) Between Aomon and Bijile: 1975, in shallow of blind channel, lagoon side, E. Chave. Enewetak: 1975, inter- tidally on benchrock, lagoon side, at northern end of island, S. L. Brunenmeister. Clibanarius zebra rhabdodactylus Forest Enewetak: 1967, from under cemented slabs at the highest place the slabs occur on the beach, J. W. Knud- sen. Enewetak: 1975, collected intertidally from crevice in the benchrock at lagoon margin, northern end of island, S. L. Brunenmeister. Clibanarius sp. Enewetak: 1975, under intertidal on seaward reef flat under large rock, in large aggregation; also intertidally on benchrock at lagoon margin, northern end of island, S. L. Brunenmeister. (A species with antennules, anten- nae, and eyestalks orange, chelipeds brown with light orange fingers, legs banded orange and black.) Dardanus crassimanus (H. Milne Edwards) Off Bokandretok, just across interisland channel from Enewetak: 1974, collected at 2.4 to 3 m on sandy bot- tom around coral heads, S. L. Brunenmeister. Dardanus deformis (H. Milne Edwards) Off Bokandretok: 1974, S. L. Brunenmeister. Medren: 1959, F. C. Ziesenhenne. Between Medren and Muti: 1960, E. S. Reese. Muti: 1956, student collector. Enewetak Atoll, no further locality data: 1946, F. C. Ziesenhenne. Reef, ocean side; lagoon side; sublittoral, 0.6 to 9 m on sand and coral rubble. Dardanus gemmatus (H. Milne Edwards) Between Enewetak and Bokandretok: 1974, collected at low tide in shallows of interisland channel on sandy bot- tom with coral rubble, crabs carrying anemones on shells, S. L. Brunenmeister. Dardanus guttatus (Olivier) Between Aomon and Bijile: 1975, E. Chave. Off Enewetak: 1967, 1968, J. W. Knudsen. Off Enewetak: 1974 or 1975, S. L. Brunenmeister. Between Medren and Muti: 1960, E. S. Reese. Off Muti: 1967, J. W. Knudsen. Rigili: 1959, J. S. Garth. On reef, in channels, and in lagoon; littoral and to depths of 30 m. Dardanus lagopodes (Forsskal) Aaraanbiru: 1959, J. S. Garth. Off Bokandretok: 1974, S. L. Brunenmeister. Enewetak: 1967, J. W. Knudsen. Between Medren and Muti: 1960, E. S. Reese. Muzin, Kirinian, and Bokanaarappu: 1959, F. C. Ziesenhenne. Rigili: 1959, J. S. Garth. Enewetak Atoll, 11°25/N, 162°13’E: 1967, J. W. Knudsen. In lagoon, littoral, div- ing to 9 m on and around corals, and dredged at 45 m. Dardanus megistos (Herbst) Muti: 1968, collected in 6 m of water at night, J. W. Knudsen. Rigili: 1959, southeast side between ocean and lagoon, +0.8 foot tide, from rock and dead coral, J. S. Garth. 252 GARTH, HAIG, AND KNUDSEN Dardanus scutellatus (H. Milne Edwards) Aomon, Bijile, and Rojoa: 1959, F. C. Ziesenhenne. Off Bokandretok: 1974, S. L. Brunenmeister. Chinieero: 1960, E. S. Reese. Medren: 1957, 1959, J. S. Garth. Medren: 1960, E. S. Reese. Muti: 1956, student collec- tor. In lagoon, littoral to 12 m, on sandy bottoms around and on coral heads and among coral rubble. Dardanus woodmasoni (Alcock) Aomon, Bijile, and Rojoa: 1959, 1.8 to 7.2 m on sandy bottom, lagoon side, F. C. Ziesenhenne. Diogenes gardineri Alcock Aomon, Bijile, and Rojoa: 1959, F. C. Ziesenhenne. Enewetak: 1966, 1967, J. W. Knudsen. Medren: 1959, J. S. Garth. Between Piiraai and Runit: 1967, J. W. Knudsen. On sandy bottoms in lagoon, 1.8 to 24 m. Diogenes pallescens Whitelegge Enewetak Atoll: 1966, in lagoon 100 yds west of marker, ran 250 yds due south, shell-sponge bottom at 60 m, J. W. Knudsen. Enewetak Atoll: 1968, in lagoon at 11°26’N, 162°17.5’E, dredging near coral mound, bottom of shell and some algae at 60 m, J. W. Knudsen. Paguristes sp. Medren: 1959, J. Coatsworth, J. S. Garth. Rojoa: 1967, J. W. Knudsen. Enewetak Atoll, 11°26’N, 162°17.5’E: 1968, J. W. Knudsen. Behind reef on coral, 1.5 m; in lagoon, to 45 m. Trizopagurus strigatus (Herbst) Enewetak: 1974 or 1975, in interisland channel at northern end of island, low tide at about 0.3 m depth, S. L. Brunenmeister. Medren: 1959, reef between island and L. C. T. wreck, intertidal at + 0.8-foot tide, J. Rob- erts and/or F. C. Ziesenhenne. Family PAGURIDAE Catapagurus sp. Enewetak Atoll: 1966, dredging in lagoon on sponge and sand bottom, depth not indicated, J. W. Knudsen. Enewetak Atoll: 1968, dredging in lagoon at 11°26’N, 162°17.5’E, in 30 m depth on bottom of broken coral and shell, J. W. Knudsen. Pagurixus anceps (Forest) Bogallua: 1968, J. W. Knudsen. Bokandretok: 1967, J. W. Knudsen. Enewetak: 1975, S. L. Brunenmeister. Between Enewetak and Bokandretok: 1966, J. W. Knudsen. Igurin: 1959, J. S. Garth. Medren: 1957, T. Goreau and R. Neshida. Rojoa: 1967, J. W. Knudsen. Intertidal under rocks and to 15 ft, lagoon side; more frequently on reef flat at low water, from Porites, Acropora, and Favia, some live. Pagurixus boninensis (Melin) Medren: 1957, about 8 ft, from Pocillopora and Acropora, A. H. Banner. Pagurixus maorus (Nobili) Aniyaanii: 1966, in mostly dead Acropora, J. W. Knud- sen. Between Piiraai and Runit: 1967, from 95 to 105 ft in lagoon, J. W. Knudsen. Rojoa: 1966, northwest side on outer reef channel, from dead Acropora, J. W. Knudsen. Enewetak Atoll: 1968, dredging in lagoon at 11°26’N, 162°17.5’E, 100 ft, broken coral and shell with much live coral, J. W. Knudsen. New genus, sp. Muti: 1966, collected at pier from dead and heavily encrusted pocilloporid coral, J. W. Knudsen. Family GALATHEIDAE Allogalathea elegans (Adams and White) Near Jierorv, at about 11°26’N, 162°21’E: 1967, J. W. Knudsen. Off Medren: 1968, 1971, J. W. Knudsen. Enewetak Atoll, 11°29’N, 162°19’E: J. W. Knudsen. Enewetak Atoll: 1976 or 1978, A. Fielding. In lagoon, to 9 m; all specimens from crinoids. Coralliogalathea humilis (Nobili) Bogombogo: 1957, D. Reish. Enewetak: 1967, J. W. Knudsen. Enewetak: 1967, S. Swerdloff. Rigili: 1965, J. W. Knudsen. Enewetak Atoll, 11°25’N, 162°13’E: 1967, J. W. Knudsen. On reef from dead corals and live Acropora; dredged in lagoon in 45 m on bottom of sponge, dead shell, coral, and algae. Galathea aegyptiaca Paulson Aniyaanii: 1960, north lagoon side, from coral head, E. S. Reese. Galathea affinis Ortmann Aniyaanii: 1957, A. H. Banner. Aniyaanii: 1966, 1967, J. W. Knudsen. Between Billee and Bokonaarappu: 1967, J. W. Knudsen. Bogallua: 1968, J. W. Knudsen. Bogombogo: 1957, D. Reish. Bokandretok: 1965, J. W. Knudsen. Enewetak: 1965, 1967, 1968, J. W. Knud- sen. Between Enewetak and Bokandretok: 1966, J. W. Knudsen. Engebi: 1967, J. W. Knudsen. Igurin: 1965, J. W. Knudsen. Between Kirinian and Muzin: 1965, J. W. Knudsen. Medren: 1957, T. Goreau and R. Neshida. Medren: 1959, J. Coatsworth. Rigili: 1965, 1968, J. W. Knudsen. Rojoa: 1966, 1967, J. W. Knud- sen. On reef to 4.5 m in live and dead corals, including Acropora and Porites; dredged in lagoon at 30 m. Galathea amboinensis De Man Enewetak Atoll: 1976 or 1978, from crinoids, A. Field- ing. Galathea sp., aff. tanegashimae Baba Enewetak: 1967, S. Swerdloff. Jierorv: 1967, 1968, J. W. Knudsen. Medren: 1957, A. H. Banner. Rojoa: 1966, J. W. Knudsen. On reef and in lagoon, to 9 m depth; on corals including live and dead Acropora, and from crinoids. Phylladiorhynchus serrirostris (Melin) Enewetak: 1967, from coral head in lagoon about 300 yds. from laboratory site, J. W. Knudsen. Medren: 1957, diving in 3 to 4.5 m, coral heads and dead coral, T. Goreau and R. Neshida. Family PORCELLANIDAE Neopetrolisthes maculatus (H. Milne Edwards) Bokandretok: 1968, J. W. Knudsen. Medren: 1971,. J. W. Knudsen. Between Medren and Enewetak: 1968, J. W. Knudsen. Rigili: 1968, J. W. Knudsen. To 3 m depth in channels and lagoon, from anemones. Pachycheles johnsoni Haig Bijile: 1968, A. Havens. Bokandretok: 1967, J. W. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 253 Knudsen. Bokandretok: 1968, A. Havens. Enewetak: 1965 to 1967, J. W. Knudsen. Enewetak: 1968, A. Ha- vens. Enewetak: 1969, C. A. Child. Between Enewetak and Bokandretok: 1966, J. W. Knudsen. Engebi: 1959, F. C. Ziesenhenne. Engebi: 1967, J. W. Knudsen. Mui: 1967, J. W. Knudsen. Muti: 1966, 1968, J. W. Knud- sen. Rigili: 1957, A. H. Banner. Rigili: 1966, J. W. Knudsen. On reef, intertidal or shallow subtidal, fre- quently in or under live and dead Porites, Pocillopora, and Acropora; occasionally under rocks. Pachycheles pisoides (Heller) Enewetak: 1969, surge channel and blow hole about 20 ft back from outer reef edge, C. A. Child. Between Enewetak and Bokandretok: 1966, on reef inside algal ridge, from dead encrusted algae or from Pocillopora, J. W. Knudsen. Pachycheles spinipes (A. Milne Edwards) Chinieero: 1965, J. W. Knudsen. Enewetak: 1965, 1967, J. W. Knudsen. Between Enewetak and Bokan- dretok: 1966, J. W. Knudsen. On reef, from corals. Petrolisthes asiaticus (Leach) Bogen: 1957, J. S. Garth. Bokandretok: 1967, J. W. Knudsen. Bokandretok: 1968, A. Havens. Enewetak: 1965, J. W. Knudsen. Enewetak: 1968, A. Havens. Enewetak or Gugegwe [Kwajalein Atoll]: 1946, F. C. Ziesenhenne. Engebi: 1957, J. S. Garth. Igurin: 1959, J. S. Garth. Jieruro: 1946, M. W. Johnson. Medren: 1957, J. S. Garth. Medren: 1965, J. W. Knudsen. Muti: 1957, J. S. Garth. Muti: 1965, G. Bakus. Muti: 1967, 1968, J. W. Knudsen. Rigili: 1956, student collector. Rigili: 1959, J. S. Garth. Runit: 1959, J. S. Garth. Runit: 1965, J. W. Knudsen. Runit: 1968, A. Havens. On reef, usually under rocks and coral rubble. Petrolisthes bispinosus Borradaile Bokandretok: 1967, J. W. Knudsen. Enewetak: 1965, 1967, J. W. Knudsen. Enewetak: 1969, C. E. Dawson, C. A. Child. Between Enewetak and Bokandretok: 1966, J. W. Knudsen. Mui: 1967, T. Smith. Muti: 1968, J. W. Knudsen. On seaward reef flat and algal ridge, in corals including Pocillopora. Petrolisthes borradailei Kropp Aniyaanii: 1956, student collector. Bogen: 1957, J. S. Garth. Bokandretok: 1967, J. W. Knudsen. Enewetak: 1965, 1967, J. W. Knudsen. Engebi: 1957, J. S. Garth. Igurin: 1959, J. S. Garth. Muti: 1965, 1968, J. W. Knudsen. Muti: 1965, G. Bakus. Muti: 1968, A. Ha- vens. Rigili: 1957, J. S. Garth. Runit: 1959, J. S. Garth. On reef, usually under rocks and in coral rubble. Petrolisthes coccineus (Owen) Enewetak: 1969, surge channel and blow hole about 20 ft back from outer reef edge, C. A. Child. Engebi: 1967, algal ridge, J. W. Knudsen. Petrolisthes decacanthus Ortmann Enewetak: 1967, algal ridge, J. W. Knudsen. Enewetak: 1969, outer reef rim and surge channels along north one-third of island, C. E. Dawson and C. A. Child. Enewetak: 1969, surge channel and blow hole about 20 ft back from outer reef edge, C. A. Child. Petrolisthes elegans Haig Enewetak: 1965, 1967, J. W. Knudsen. Enewetak: 1969, C. A. Child. Between Enewetak and Bokandre- tok: 1966, J. W. Knudsen. Medren: 1965, J. W. Knud- sen. Muti: 1968, J. W. Knudsen. Rujoru: 1946, M. W. Johnson. Reef flat and algal ridge, under rocks and from corals, including Pocillopora. Petrolisthes fimbriatus Borradaile Aniyaanii: 1957, J. S. Garth. Aniyaanii: 1967, J. W. Knudsen. Bogen: 1966, J. W. Knudsen. Bokandretok: 1967, J. W. Knudsen. Bokandretok: 1968, A. Havens. Enewetak: 1957, J. S. Garth. Enewetak: 1965 to 1968, J. W. Knudsen. Enewetak: 1968, A. Havens. Enewetak or Gugegwe [Kwajalein Atoll]: 1946, F. C. Ziesenhenne. Between Enewetak and Bokandretok: 1966, J. W. Knudsen. Engebi: 1956, student collector. Engebi: 1957, J. S. Garth. Igurin: 1957, 1959, J. S. Garth. Igu- rin: 1965, J. W. Knudsen. Jeiroju: 1946, J. P. E. Morri- son. Jieruro: 1946, M. W. Johnson. Medren: 1957, 1959, J. S. Garth. Medren: 1965, J. W. Knudsen. Muti: 1957, J. S. Garth. Muti: 1965, 1968, J. W. Knudsen. Muti: 1965, G. Bakus. Muti: 1968, A. Havens. Piiraai: 1968, A. Havens. Rigili: 1956, D. Reish. Rigili: 1959, J. S. Garth. Rigili: 1966, 1968, J. W. Knudsen. Runit: 1959, J. S. Garth. Enewetak Atoll, no further locality data: 1944, W. A. Bartos. Enewetak Atoll, no further locality data: 1946, F. C. Ziesenhenne. On reef, under rocks and from rubble; occasionally from corals. Petrolisthes lamarckii (Leach) Aniyaanii: 1957, J. S. Garth. Bokandretok: 1967, J. W. Knudsen. Bokandretok: 1968, A. Havens. Enewetak: 1965, 1967, 1968, J. W. Knudsen. Enewetak: 1968, A. Havens. Engebi: 1957, J. S. Garth. Igurin: 1957, 1959, J. S. Garth. Medren: 1959, J. S. Garth. Medren: 1965, J. W. Knudsen. Mui: 1968, A. Havens. Muti: 1957, J. S. Garth. Muti: 1965, G. Bakus. Muti: 1967, 1968, J. W. Knudsen. Muti: 1968, A. Havens. Piiraai: 1968, A. Havens. Rigili: 1957, 1959, J. S. Garth. Rigili: 1966, J. W. Knudsen. Runit: 1959, J. S. Garth. Runit: 1965, J. W. Knudsen. Enewetak Atoll, no further locality data: 1946, F. C. Ziesenhenne. On reef, usually under rocks and in coral rubble. Petrolisthes masakii Miyake Enewetak: 1969, surge channel and blow hole about 20 ft back from outer reef edge, C. A. Child. Petrolisthes penicillatus (Heller) Off Bokandretok: 1968, A. Havens. Enewetak: 1968, J. W. Knudsen. Enewetak: 1968, A. Havens. Enewetak: 1969, C. A. Child. Between Enewetak and Bokandre- tok: 1966, J. W. Knudsen. Medren: 1965, 1968, J. W. Knudsen. Rigili: 1965, J. W. Knudsen. On reef, to 0.9 m depth, under rocks and rubble, and in Porites, Acropora, and Pocillopora. Petrolisthes pubescens Stimpson Bokandretok: 1968, channel southwest of island, under rock, A. Havens. Petrolisthes [undescribed sp. 1] Rigili: 1965, north end of island from live Pocillopora elegans, J. W. Knudsen. 254 GARTH, HAIG, AND KNUDSEN Petrolisthes [undescribed sp. 3] 1967, J. W. Knudsen. Reef flat and ridge, some speci- Enewetak: 1967, J. W. Knudsen. Enewetak: 1968, mens from corals. A. Havens. Enewetak: 1969, C. A. Child. Engebi: TABLE 1 Data on Collections of Anomura* Collector(s) Date Identifier Depository Bakus, G. 1965 J. Haig AHF Banner, A.H. 1957 J. Haig, AHF M. de Saint Laurent, P. McLaughlin Bartos, W. A. 1944 J. Haig USNM Brunenmeister, S. L. 1974, 1975 S. Brunenmeister, P. McLaughlin, J. Haig Chave, E. 1975 P. McLaughlin Child, C. A. 1969 J. Haig, USNM, RMNH L. B. Holthuis Coatsworth, J. 1959 J. Haig AHF, MPRL Colin, P., and D. M. Devaney 1980 M. de Saint Laurent, BPBM J. Haig Dawson, C. E. 1969 J. Haig, USNM, RMNH L. B. Holthuis Donaldson, L., and E. Held 1957 J. S. Garth AHF, MPRL Fielding, A. 1976, 1978 K. Baba BPBM Fusaro, C. 1975 C. Fusaro BPBM Garth, J. S. 1957, 1959 J. S. Garth, AHF, MPRL J. Haig P. McLaughlin Goreau, T., and R. Neshida 1957 J. Haig, AHF, MPRL P. McLaughlin Havens, A. 1968 J. Haig AHF Johnson, M. W. 1946 J. Haig USNM Johnson, S. 1981 L. B. Holthuis BPBM, MPRL, USNM Knudsen, J. W. 1965 to 1968 J. Haig AHF 1971 P. McLaughlin AHF MacCoy, C. V. 1967 J. Haig AHF Morrison, J. P. E. 1946 J. Haig USNM Reese, E. S. 1960, 1961 E. S. Reese, J. Haig AHF Reese, E. S., and R. A. Boolootian 1960 E. S. Reese, J. Haig AHF Reish, D. 1956, 1957 J. Haig AHF Roberts, J. 1959 J. Haig AHF Sather, B., and R. Stevenson 1960 J. Haig AHF Schultz, L. P. 1946 L. B. Holthuis USNM Smith, A., and J. Coatsworth 1959 J. Haig AHF Smith, T. 1967 J. Haig AHF Student collectors 1956 J. S. Garth MPRL Swerdloff, S. 1967 J. Haig AHF Ziesenhenne, F. C. 1946, 1959 J. S. Garth, J. Haig, AHF M. de Saint Laurent *AHF, Allan Hancock Foundation; BPBM, Bernice P. Bishop Museum; MPRL, Mid-Pacific Research Laboratory, Enewetak; RMNH, Rijksmuseum van Natuurlijke Historie, Leiden; USNM, National Museum of Natural History, Smithsonian Institution. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 255 Brachyuran Crustacea Taxa are listed in the same order as in the checklist under their respective families. Higher categories and also synonyms may be consulted in the checklist. Because plans are to give complete data—including islet, substrate, depth, month, and year of collecting—in future publica- tions, only collectors’ surnames in chronological order are given here. Their respective periods of activity at Enewetak Atoll and the present depository of their collec- tions will be found in Table 2. Family DROMIIDAE Cryptodromia canaliculata Stimpson EMBL, Reish, Garth, Ziesenhenne, Bakus, Knudsen. Cryptodromia sp. Knudsen. Family DYNOMENIDAE Dynomene hispida Desmarest Garth, Reese, Knudsen, Havens, Child, Highsmith. Dynomene pilumnoides Alcock Knudsen. Dynomene praedator A. Milne Edwards Knudsen. Dynomene spinosa Rathbun Garth, Knudsen, Havens. Family LEUCOSIIDAE Cryptocnemus haddoni Calman Havens. Ebalia woodmasoni Alcock Knudsen. Ebaliopsis erosa (A. Milne Edwards) Garth, Ziesenhenne, Knudsen. Heterolithadia sp. Knudsen. Heteronucia venusta Nobili Knudsen, Havens. Merocryptus durandi Serene Knudsen. Myra fugax coalita Hilgendorf Knudsen. Nucia ingens (Rathbun) Knudsen. Nucia speciosa Dana Knudsen. Oreophorus (Oreotlos) latus Borradaile Knudsen, Havens. Species incertae sedis Knudsen. Family CALAPPIDAE Calappa calappa (Linnaeus) Knudsen, Havens. Calappa gallus (Herbst) Garth, Knudsen, Fielding. Calappa hepatica (Linnaeus) EMBL, Reish, Garth, Ziesenhenne, Knudsen, Havens, Child, USGS (fossil). Family MAJIDAE Camposcia retusa Latreille Garth. Camposcia sp. Knudsen. Cyclax suborbiculatus (Stimpson) Garth, Ziesenhenne, Knudsen, Havens. Huenia brevifrons Ward Knudsen. Huenia proteus De Haan Ziesenhenne, Knudsen. Hyastenus irami (Laurie) Banner, Knudsen, Child. Hyastenus uncifer Calman Knudsen, Child. Hyastenus verrucosipes (Adams and White) Knudsen. Hyastenus sp. Knudsen. Menaethius monoceros (Latreille) Reish, Banner, Garth, Ziesenhenne, Reese, Knudsen, Havens, Child. Micippa margaritifera Henderson Knudsen. Micippa philyra (Herbst) Garth, Ziesenhenne, Knudsen. Micippa platypes Riippell Reish, Ziesenhenne, Stokes, Knudsen, Havens. Micippa thalia (Herbst) Knudsen. Naxioides spinigera Borradaile Knudsen. Paratymolus bituberculatus Miers Knudsen. Paratymolus sexspinosus Miers Garth. Parazewa bocki Balss Knudsen. Perinea tumida Dana Reish, Banner, Garth, Ziesenhenne, Reese, Knudsen, Havens. Schizophrys aspera (H. Milne Edwards) Garth, Reese, Knudsen, Child. Trigonothir obtusirostris Miers Palumbo, Knudsen. Tylocarcinus ?gracilis Miers Knudsen. Tylocarcinus styx (Herbst) Banner, Garth, Reese, Shoup, Knudsen, Havens, Child. Family PARTHENOPIDAE Actaeomorpha sp., nr. erosa (Miers) Knudsen. Cryptopodia ?pan Laurie Knudsen. Daldorfia horrida (Linnaeus) EMBL, Garth, Knudsen, Havens. 256 Daldorfia (or Parthenope) sp. Knudsen. Harrovia elegans De Man Garth. ?Heterocrypta sp. Knudsen, Havens. Parthenope (Aulacolambrus) curvispinis (Miers) Knudsen. Parthenope (Aulacolambrus) hoplonotus (Adams and White) Knudsen. Parthenope (Aulacolambrus) sp. Knudsen, Havens. Parthenope (Pseudolambrus) sp. Knudsen. Parthenope sp. Knudsen. Thyrolambrus erosus (Miers) Knudsen. Thyrolambrus sp. Knudsen. Family ATELECYCLIDAE Kraussia integra Rathbun Knudsen, Havens. Kraussia sp., cf. marquesa Serene Knudsen. Kraussia nitida Stimpson Garth. Kraussia rastripes F. Miiller Havens. Kraussia rugulosa (Krauss) EMBL, Garth, Knudsen, Havens, Child. Family PORTUNIDAE Carupa tenuipes Dana Ziesenhenne, Knudsen, Havens. Catoptrus inaequalis (Rathbun) Knudsen. Catoptrus nitidus A. Milne Edwards Garth, Ziesenhenne. Catoptrus rathbunae Serene Ziesenhenne. Catoptrus ?truncatifrons De Man Havens. Charybdis (Goniosupradens) erythrodactylus (Lamarck) Havens, Fielding. Coelocarcinus foliatus Edmondson Havens. Libistes villosus Rathbun Knudsen, Havens. Lissocarcinus holothuricola Streets Knudsen. Lissocarcinus orbicularis Dana Garth, Shoup, Knudsen, Havens. Portunus (Achelous) granulatus (H. Milne Edwards) Ziesenhenne, Knudsen, Havens. Portunus (Achelous) sp., nr. orbicularis (Richters) Garth, Ziesenhenne. Portunus (Hellenus) longispinosus Stephenson and Campbell GARTH, HAIG, AND KNUDSEN Reish, Garth, Ziesenhenne, Knudsen, Havens. Thalamita admete (Herbst) EMBL, Reish, Garth, Ziesenhenne, Knudsen, Havens, Child. Thalamita bouvieri Nobili Knudsen. Thalamita chaptalii (Audouin) Knudsen. Thalamita coeruleipes Jacquinot Knudsen, Havens. Thalamita corrugata Stephenson and Rees Knudsen. Thalamita dakini Montgomery Knudsen, Havens, Child. Thalamita gloriensis Crosnier Morrison. Thalamita gracilipes (A. Milne Edwards) Garth, Ziesenhenne, Knudsen, Havens. Thalamita integra Dana Pomeroy and Kuenzler. Thalamita oculea Alcock Knudsen. Thalamita picta Stimpson EMBL, Garth, Ziesenhenne, Knudsen, Havens, Child. Thalamita pilumnoides Borradaile Garth, Knudsen. Thalamita prymna (Herbst) Knudsen, Havens, Child, Fielding. Thalamita quadrilobata Miers Knudsen. Thalamita sexlobata Miers Knudsen. Thalamita sima H. Milne Edwards Knudsen. Thalamita spiceri Edmondson Garth, Knudsen, Havens, Highsmith. Thalamita spinimana Dana EMBL, Garth. Thalamita stimpsoni A. Milne Edwards Knudsen. Thalamita wakensis Edmondson Knudsen, Havens. Thalamita yoronensis Sakai Knudsen. Thalamita sp., nr. auauensis Rathbun Knudsen. Thalamitoides quadridens A. Milne Edwards Banner, Garth, Reese, Knudsen, USGS (fossil). Family XANTHIDAE Actaea sp., nr. bocki Odhner Knudsen. Actaea(?) cavipes Dana Knudsen, Havens. Actaea margaritifera Odhner Knudsen. Actaea pulchella modesta (De Man) Knudsen. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 257 Actaea quadriareolata Takeda and Miyake Banner, Garth, Knudsen. Actaea sp. Knudsen, Havens. Actaeodes consobrinus (A. Milne Edwards) Garth, Reese, Knudsen, Havens. Actaeodes hirsutissimus (Riippell) USGS (fossil). Actumnus antelmei Ward Knudsen. Actumnus asper (Riippell) Knudsen. Actumnus setifer (De Haan) Knudsen. Actumnus sp. Knudsen. Actumnus (or Pilumnus) sp. Knudsen. Atergatis ?dilitatus De Haan de Gruy. Atergatis floridus (Linnaeus) Knudsen. Atergatopsis signata (Adams and White) Knudsen, Burke. Banareia nobilii (Odhner) Knudsen. Banareia parvula (Krauss) Knudsen. Carpilius convexus (Forsskal) Johnson, Ziesenhenne, Knudsen, Havens, Fielding. Carpilius maculatus (Linnaeus) Knudsen, Fielding. Chlorodiella corallicola Miyake and Takeda Knudsen. Chlorodiella cytherea (Dana) Ziesenhenne, Knudsen, Havens. Chlorodiella laevissima (Dana) Reish, Garth, Ziesenhenne, Reese, Knudsen, Havens. Chlorodiella nigra (Forsskal) Reish, Banner, Garth, Shoup, Knudsen, Havens. Cycloxanthops cavatus Rathbun Garth, Knudsen, Havens. Cymo andreossyi (Audouin) Banner, Reese, Knudsen. Cymo deplanatus A. Milne Edwards Ziesenhenne, Reese, Shoup, Knudsen, Child. Cymo melanodactylus De Haan Garth, Reese, Knudsen. Cymo quadrilobatus Miers Knudsen. Dacryopilumnus eremita Nobili Knudsen, Havens. Dacryopilumnus rathbunae Balss Knudsen, Havens. Daira perlata (Herbst) Havens. Domecia glabra Alcock Banner, Garth, Ziesenhenne, Reese, Shoup, Knudsen, Havens, Child. Domecia hispida Eydoux and Souleyet Reese, Shoup, Knudsen, Havens. Eriphia scabricula Dana Morrison, EMBL, Garth, Ziesenhenne, Knudsen, Havens, Child. Eriphia sebana (Shaw and Nodder) Morrison, EMBL, Garth, Ziesenhenne, Bakus, Knud- sen, Havens, Child. Etisus bifrontalis (Edmondson) Knudsen. Etisus demani Odhner Johnson, Knudsen, Garth, Havens. Etisus sp., nr. demani Odhner Knudsen, Havens. Etisus dentatus (Herbst) Garth, Ziesenhenne, Knudsen, Havens, Child. Etisus electra (Herbst) Reish, Knudsen. Etisus frontalis Dana Knudsen. Etisus laevimanus Randall USGS (fossil). Etisus molokaiensis (Rathbun) Garth, Havens. Etisus splendidus Rathbun Fielding, USGS (fossil). Etisus sp. 1 Knudsen, Havens. Etisus sp. 2 Knudsen, Havens. Etisus sp. 3 Havens. Euxanthus exsculptus (Herbst) Garth. Euxanthus (or Hypocolpus) sp. Knudsen. Gaillardiellus rueppellii (Krauss) Ziesenhenne, Knudsen. Gaillardiellus superciliaris (Odhner) Reish, Banner, Garth, Ziesenhenne, Reese, Shoup, Knudsen, Havens, Child. Globopilumnus globosus (Dana) Knudsen, Havens. Heteropilumnus sp., cf. longipes (Stimpson) Knudsen, Havens. Lachnopodus ponapensis (Rathbun) Knudsen, Havens. Lachnopodus subacutus (Stimpson) Knudsen, Havens, Fielding. Lachnopodus tahitensis De Man EMBL, Garth, Knudsen, Havens, Child, Burke. Leptodius davaoensis Ward Havens, Child. Leptodius exaratus (H. Milne Edwards) Reish, Garth, Bussing, Knudsen, Child. Leptodius gracilis (Dana) Ziesenhenne, Bakus, Knudsen, Havens. Leptodius nudipes (Dana) Knudsen, Havens. 258 Leptodius sanguineus (H. Milne Edwards) Ziesenhenne, Morrison, Johnson, EMBL, Garth, Bakus, Knudsen, Havens, Child. Leptodius waialuanus Rathbun Knudsen, Havens, Child. Liocarpilodes armiger pacificus Balss Ladd, Banner, Garth, Reese, Shoup, Knudsen, Havens, Highsmith. Liocarpilodes biunguis (Rathbun) Ladd, EMBL, Reish, Garth, Knudsen, Havens, Child, Highsmith. Liocarpilodes integerrimus (Dana) Reish, Knudsen, Havens. Liocarpilodes pumilus (Jacquinot) Ladd, EMBL, Knudsen. Liomera bella (Dana) Bartos, Morrison, Banner, Garth, Ziesenhenne, Shoup, Bakus, Knudsen, Havens, Highsmith. Liomera coelata (Odhner) Banner. Liomera loevis (A. Milne Edwards) Garth. Liomera monticulosa (A. Milne Edwards) Reese, Knudsen. Liomera pallida (Borradaile) Knudsen, Havens. Liomera rugata (H. Milne Edwards) Ziesenhenne, Garth, Knudsen, Havens. Liomera stimpsoni (A. Milne Edwards) Knudsen. Liomera tristis (Dana) Garth. Liomera sp. Knudsen. Lophozozymus dodone (Herbst) Knudsen, Child. Lophozozymus incisus (H. Milne Edwards) Johnson. Lophozozymus pulchellus A. Milne Edwards Shoup, Knudsen, Havens. Lubia caestifera (Alcock) Knudsen. Lybia tessellata (Latreille) Ziesenhenne, Stokes, Knudsen, Havens. Ludia annulipes (H. Milne Edwards) Knudsen, Havens, Fielding. Macromedaeus nudipes (A. Milne Edwards) Garth, Knudsen, Havens, Child. Maldivia palmyrensis Rathbun Knudsen, Highsmith. Maldivia triunguiculata (Borradaile) Knudsen, Havens, Highsmith. Medaeus elegans A. Milne Edwards Knudsen, Havens. Medaeus ornatus Dana Knudsen, Havens. GARTH, HAIG, AND KNUDSEN Neoxanthias impressus (Lamarck) Garth, Ziesenhenne, Knudsen, Burke. Paractaea retusa (Nobili) Knudsen, Havens. Paractaea rufopunctata (H. Milne Edwards) Banner, Garth, Knudsen, Havens. Paractaea rufopunctata f. plumosa Guinot Knudsen. Paractaea tumulosa (Odhner) Knudsen. Paramedaeus simplex (A. Milne Edwards) Knudsen, Havens, Child. Parapilumnus coralliophilus Takeda and Miyake Shoup, Knudsen, Havens. Parapilumnus ?incertus Takeda and Miyake Knudsen. Paraxanthias notatus (Dana) Ladd, Reish, EMBL, Banner, Garth, Ziesenhenne, Knudsen, Havens, Highsmith. Paraxanthias pachydactylus (A. Milne Edwards) Knudsen, Havens. Phymodius ?granulatus (Targioni-Tozzetti) Knudsen. Phymodius laysani Rathbun Reese, Knudsen, Havens. Phymodius monticulosus (Dana) Garth, Knudsen. Phymodius nitidus (Dana) Garth, Knudsen, Havens. Phymodius ungulatus (H. Milne Edwards) Banner, Reish, Garth, Reese, Shoup, Knudsen, Havens. Pilodius areolatus (H. Milne Edwards) Ziesenhenne, Johnson, Ladd, EMBL, Garth, Ziesenhenne, Shoup, Bakus, Knudsen, Havens, Highsmith. Pilodius flavus Rathbun Reese, Knudsen, Havens. Pilodius melanodactylus (A. Milne Edwards) Knudsen. Pilodius pilumnoides (White) Morrison, Reish, Garth, Knudsen, Havens. Pilodius pugil Dana Johnson, Reish, Banner, Ziesenhenne, USGS (fossil). Pilodius scabriculus Dana Knudsen, Havens. Pilodius spinipes Heller Reish, Banner, Garth, Knudsen, Havens. Pilumnus andersoni De Man Knudsen, Havens. Pilumnus caerulescens A. Milne Edwards Knudsen, Havens, Child. Pilumnus ?elegans De Man Knudsen. Pilumnus longicornis Hilgendorf Banner, Garth, Knudsen, Havens, Child. Pilumnus ransoni Forest and Guinot Knudsen. CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) 259 Pilumnus rotumanus Borradaile Knudsen. Pilumnus tahitensis De Man Knudsen, Havens. Pilumnus vespertilio (Fabricius) Knudsen. Pilumnus sp. Havens. Planopilumnus vermiculatus (A. Milne Edwards) Banner, Knudsen, Havens. Polydectus cupulifer (Latreille) Banner, Knudsen, Havens, Child. Pseudoliomera granosimanus (A. Milne Edwards) Knudsen, Havens. Pseudoliomera helleri (A. Milne Edwards) Knudsen. Pseudoliomera sp., nr. helleri (A. Milne Edwards) Knudsen. Pseudoliomera lata (Borradaile) Knudsen. Pseudoliomera sp. nr. lata (Borradaile) Knudsen. Pseudoliomera rueppellioides (Odhner) Knudsen. Pseudoliomera speciosa (Dana) Reish, Reese and Stevenson, Shoup, Knudsen, Havens. Pseudozius caystrus (Adams and White) Ziesenhenne, Johnson, EMBL, Garth, Knudsen, Havens, Child. Pseudozius pacificus Balss Garth, Knudsen. Ralumia dahli Balss Knudsen. Tetralia glaberrima (Herbst) Morrison, Ladd, Reish, EMBL, Banner, Garth, Ziesenhenne, Reese, Shoup, Knudsen, Havens, Child. Tetralia glaberrima rubridactylus Patton Reese, Child Tetraloides nigrifrons (Dana) Garth, Ziesenhenne, Reese, Shoup, Knudsen, Havens. Trapezia cymodoce (Herbst) Morrison, EMBL, Garth, Reese and Stevenson, Shoup, Knudsen, Havens, Child. Trapezia dentata Macleay Knudsen, Child. Trapezia digitalis Latreille Garth, Ziesenhenne, Shoup, Knudsen, Havens. Trapezia digitalis bella Dana Knudsen. Trapezia sp., digitalis group Reese, Knudsen. Trapezia ferruginea Latreille Morrison, Banner, Reish, Garth, Ziesenhenne, Reese and Stevenson, Shoup, Knudsen, Havens, Child. Trapezia guttata Riippell Ladd, Reish?, Reese, Shoup, Bussing, Knudsen, Havens. Trapezia rufopunctata (Herbst) Knudsen. Trapezia rufopunctata flavopunctata Eydoux and Souleyet Knudsen. Trapezia rufopunctata maculata Macleay Garth, Reese and Stevenson, Shoup, Knudsen. Trapezia speciosa Dana Ladd, Ziesenhenne, Knudsen, Havens. Trapezia tigrina Eydoux and Souleyet Reish?, Garth, Ziesenhenne, Reese and Stevenson, Shoup, Knudsen, Havens, Child. Trapezia sp. 1 Knudsen, Havens. Trapezia sp. 2 Havens. Xanthias canaliculatus Rathbun Knudsen, Havens. Xanthias gilbertensis Balss Knudsen. Xanthias lamarcki (H. Milne Edwards) Ziesenhenne, Morrison, Johnson, Ladd, EMBL, Garth, Ziesenhenne, Knudsen, Havens, Highsmith. Xanthias lividus Lamarck Knudsen. Xanthias punctatus (H. Milne Edwards) Knudsen, Havens. Xantho sp. Knudsen. Zozymodes cavipes (Dana) Ziesenhenne. Zozymus actaeoides (A. Milne Edwards) Knudsen. Zozymus sp., nr. actaeoides (A. Milne Edwards) Knudsen. Zozymus aeneus (Linnaeus) Morrison, Ladd, Garth, Ziesenhenne, Knudsen, Havens. Zozymus gemmula Dana Knudsen. Zozymus hawaiiensis (Rathbun) Knudsen. Zozymus kuekenthalii De Man Knudsen, Havens. Family GONEPLACIDAE Ceratoplax sp. Knudsen. Genus and species incertae sedis Knudsen. Family PALICIDAE Palicus jukesi (White) Knudsen. Palicus whitei (Miers) Knudsen. Palicus sp., nr. oahuensis Rathbun Ziesenhenne. Family GRAPSIDAE Cyclograpsus integer H. Milne Edwards Knudsen, Child. 260 Cyclograpsus longipes Stimpson Knudsen, Havens. Cyclograpsus sanctaecrucis Griffin Knudsen. Geograpsus crinipes (Dana) Held, Knudsen, Havens, Fielding. Geograpsus grayi (H. Milne Edwards) Held, Knudsen, Havens, Fielding. Grapsus intermedius De Man Knudsen. Grapsus longitarsus Dana EMBL, Held, Child, Havens. Grapsus tenuicrustatus (Herbst) EMBL, Garth, Ziesenhenne, Reese and Boolootian, Knudsen, Havens. Metasesarma rousseauxi H. Milne Edwards Knudsen. Metopograpsus thukuhar (Owen) Knudsen, Havens. Pachygrapsus minutus A. Milne Edwards EMBL, Reish, Garth, Ziesenhenne, Bakus, Knudsen, Havens, Child, Highsmith. Pachygrapsus planifrons De Man Garth, Bakus, Knudsen, Havens, Child. Pachygrapsus plicatus H. Milne Edwards EMBL, Bakus, Knudsen, Havens, Child. Percnon abbreviatum (Dana) Knudsen, Havens, Child. Percnon pilimanus (A. Milne Edwards) Knudsen, Havens, Child, Fielding. Percnon planissimum (Herbst) EMBL, Bakus, Knudsen, Havens, Child, Highsmith. Plagusia depressa tuberculata Lamarck Knudsen, Havens. Plagusia immaculata Lamarck Fielding. Plagusia speciosa Dana Knudsen, Havens, Child, Fielding. Pseudograpsus albus Stimpson EMBL, Knudsen, Havens. Collector(s) Bakus, G. J., and B. H. Bussing Banner, A. H. Bartos, W. A. Burke, R. Child, C. A. Coatsworth, J. GARTH, HAIG, AND KNUDSEN Sesarma (Holometopus) sp. Knudsen. Family GECARCINIDAE Gecarcoidea lalandii H. Milne Edwards Knudsen, Fielding. Family PINNOTHERIDAE Xanthasia murigera White Garth. Family OCYPODIDAE Macrophthalmus (Macrophthalmus) telescopicus (Owen) Knudsen. Macrophthalmus (Mopsocarcinus) bosci Audouin and Savigny Garth, Knudsen. Ocypode ceratophthalma (Pallas) EMBL, Garth, Ziesenhenne, Reese and Boolootian, Bakus, Child. Ocypode cordimana Desmarest Knudsen, Fielding. Paracleistosoma (or Cleistosoma) sp. Knudsen. Uca tetragonon (Herbst) Knudsen. Genus and species incertae sedis Knudsen. Family HAPALOCARCINIDAE Cryptochirus coralliodytes Heller Bakus, Knudsen, Wijsman. Hapalocarcinus marsupialis Stimpson Reese, Shoup, Knudsen, Havens. Neotroglocarcinus dawydoffi (Fize and Serene) Reese and Stevenson, BPBM. Pseudocryptochirus crescentus Edmondson Pichon. Species 1, incertae sedis Knudsen. Species 2, incertae sedis Knudsen. TABLE 2 Data on Collections of Brachyura* Date 1965 1957 1944 1978 1969 1959 Identifier Depository J. Garth AHF J. Garth AHF, MPRL tJ. Garth USNM J. Garth BPBM J. Garth USNM J. Garth AHF, MPRL “AHF, Allan Hancock Foundation; BPBM, Bernice P. Bishop Museum; MPRL, Mid- Pacific Research Laboratory, Enewetak; USNM, National Museum of Natural History, Smithsonian Institution. +Xanthidae only; the Portunidae were identified by Stephenson and Rees (1967). (This table continued on next page.) CRUSTACEA DECAPODA (BRACHYURA AND ANOMURA) TABLE 2 (cont'd) Collector(s) Date Identifier Depository de Gruy 1978 J. Garth BPBM Fielding, A. 1976, 1978 J. Garth BPBM Garth, J. S. 1957, 1959 J. Garth AHF, MPRL and others Goreau, T., and R. Neshida 1957 J. Garth AHF, MPRL Havens, A. D. 1968, 1969, A. Havens, AHF 1970 J. Garth Held, E. 1957 J. Garth AHF Johnson, M. W. 1946 tJ. Garth USNM Knudsen, J. W. 1965 to 1968, J. Garth AHF 1971, 1972 and others Ladd, H. S. 1952 tJ. Garth USNM Morrison, J. P. E. 1946 +J. Garth USNM Palumbo, R. 1957 J. Garth AHF Pichon, M. 1976 M. Takeda BPBM Reese, E. S., and R. A. Boolootian 1960 J. Garth AHF Reese, E. S., and R. A. Stevenson 1961 J. Garth AHF Reish, D. 1956, 1957 J. Garth AHF, MPRL Ryan, E. 1957 J. Garth AHF Shoup, J. 1961 J. Garth AHF Stokes, D. 1965 J. Garth AHF Wijsman, M. 1976 M. Takeda BPBM Ziesenhenne, F. C. 1946, + J. Garth AHF, USNM 1957, 1959 AHF, MPRL +Xanthidae only; the Portunidae were identified by Stephenson and Rees (1967). 261 Chapter 24 Holothurians of Enewetak Atoll BERTHA M. CUTRESS* and FRANCIS W. E. ROWET “Department of Marine Sciences, University of Puerto Rico, Mayagiiez, Puerto Rico 00708 +The Australian Museum Sydney, N.S.W., 2000 Australia INTRODUCTION Most of the species of holothurians recorded in the checklist (Table 1) that follows were included in an unpublished systematic report by B. M. Cutress prepared for the Mid-Pacific Research Laboratory in 1956. Collec- tion records based on that study are presented in the Appendix. Except for a short paper on Labidodemas sem- perianum by Cherbonnier (1970) and another on Thelenota anax by Lamberson (1978), the scant published literature on holothurians of Enewetak is nonsystematic. Hartman (1954) reported the polynoid, Gastrolepida clavigera, as a symbiont on Stichopus horrens and Holothuria gyrifer (H. hilla) at Enewetak Atoll. Pomeroy and Kuenzler (1967) included H. (Halodeima) atra and H. (Platyperona) difficilis among faunistic components at Enewetak examined for the excretion and turnover time of phosphorus. Reish (1968) noted that Gastrolepida clavigera was found on H. atra, H. gyrifer (H. hilla), Actinopyga TABLE 1 Checklist of Enewetak Holothurians Phylum ECHINODERMATA Class HOLOTHUROIDEA Order DENDROCHIROTIDA Family COCUMARIIDAE Afrocucumis africana (Semper): Lawrence, 1980; Lawrence and Guille, 1982. Order ASPIDOCHIROTIDA Family HOLOTHURIIDAE Actinopyga mauritiana (Quoy and Gaimard): Bakus, 1968, 1973; Lawrence, 1980; Lawrence and Guille, 1982. *+Actinopyga ?mauritiana juvenile. *Bohadschia argus Jaeger, 1883. *Holothuria (Thymiosycia) arenicola Semper, 1868. Holothuria (Halodeima) atra Jaeger: Pomeroy and Kuenzler, 1967; Bakus, 1968, 1973; Webb et al., 1977; Ebert, 1978; Lawrence, 1980; Lawrence and Guille, 1982. Holothuria (Platyperona) difficilis Semper: Pomeroy and Kuenzler, 1967; Bakus 1968, 1973; Lawrence, 1980; Lawrence and Guille, 1982. Holothuria (Thymiosycia) hilla Lesson: Bakus, 1968; Lawrence, 1980; Lawrence and Guille, 1982. *Holothuria (Thymiosycia) impatiens (Forsskal), 1775. Holothuria (Mertensiothuria) leucospilota (Brandt): Bakus, 1968, 1973; Smith et al., 1973; Lawrence, 1980; Lawrence and Guille, 1982. *Holothuria (Mertensiothuria) pervicax Selenka, 1867. Labidodemas semperianum Selenka: Cherbonnier, 1970. *Labidodemas ?semperianum, juvenile. *New published Enewetak record. +Very small specimen. (This table continued on next page.) 264 CUTRESS AND ROWE TABLE 1 (cont'd) Family STICHOPODIDAE Stichopus chloronotus Brandt: Lawrence, 1980. °Stichopus horrens Selenka, 1867. *Stichopus variegatus Semper, 1868. *+Stichopus ?variegatus juvenile. Thelenota ananas (Jaeger): Humes, 1973; Lamberson, 1978. Thelenota anax H. L. Clark: Rowe and Doty, 1977; Lamberson, 1978. Order APOPIDA Family CHIRIDOTIDAE Chiridota rigida Semper: Lawrence, 1980; Lawrence and Guille, 1982. Family SYNAPTIDAE *Euapta godeffroyi (Semper), 1868. *Pendekaplectana nigra (Semper), 1868. Synapta maculata (Chamisso and Eysenhardt): Bakus, 1968, 1973. *New published Enewetak record. +Very small specimen. mauritiana, and Stichopus horrens at Enewetak. Bakus (1968, 1973) reported on defense mechanisms and the ecology of several species. Enewetak was one site for a study (Bakus and Green, 1974) on changes in toxicity of holothurians with relation to various latitudes, water tem- peratures, and habitats. Presence of a tumor in one of 10 specimens of H. (Mertensiothuria) leucospilota was reported by Smith et al. (1973). Humes (1973) recorded a parasitic copepod on Thelenota ananas. In their report on Guam holothurians, Rowe and Doty (1977) included Enewetak in the general distribution of T. anax. Some aspects of the biology of H. (H.) atra were studied by Webb et al. (1977) and Ebert (1978). Lawrence (1980) recorded numbers and biomass of eight species of holothurians on the Enewetak reef flat. The organic com- position of the body wall of six species of Enewetak holothurians was analyzed by Lawrence and Guille (1982). BIOGEOGRAPHY According to distribution summaries for various Indo- Pacific species (H. L. Clark, 1921, 1946; A. M. Clark and Rowe, 1971), 18 of the 20 holothurian species known at present from Enewetak Atoll are widely distributed throughout the Indo-West Pacific. The other two, Thelenota anax and Chiridota rigida, range from northern Australia northward to the Philippines and eastward to the West and Central Pacific. Of the 18 Indo-West Pacific species, those most widely distributed are Holothuria (Platyperona) difficilis (Red Sea to the Hawaiian and Easter islands), H. (Thymiosycia) aren- icola (Red Sea to the Hawaiian, Galapagos, and Cocos Islands; Caribbean; Bermuda) and H. (T.) impatiens (Red Sea to Hawaii, Caribbean, Mediterranean). Twelve other species range from the shores of East Africa or the Red Sea to the Central Pacific: Actinopyga mauritiana, H. (Halodeima) atra, H. (T.) hilla, H. (Mertensiothuria) per- vicax, H. (M.) leucospilota, Stichopus chloronotus, S. horrens, and Euapta godeffroyi to the Hawaiian Islands; Stichopus variegatus, Thelenota ananas, and Synaptula maculata to the Line and (or) Society islands; and Pendaplectana nigra to the Marshall Islands. The other three have not been reported from Africa or the Red Sea. Afrocucumis africanus ranges from Mauritius to Fiji; Bohadschia argus, from the Seychelles islands to the Marshall Islands, and Labidodemas semperianum, from the Maldives to the Hawaiian and Society islands. Of the 78 holothurian species recorded from Pacific islands (A. M. Clark and Rowe, 1971), only 20 (26%) are represented at Enewetak. On the other hand, a vast major- ity (90%) of the Enewetak species also ranges into the Indian Ocean, compared with 68% of the total Pacific species recorded by Clark and Rowe. With the possible exception of the surprisingly low number of species found, the composition of the holothurian fauna at Enewetak could be predicted from the geographical position of that atoll and may be indicative of the reproductive strategies of the widespread species. Unfortunately, little is known of larval development of the species or length of larval life. Incidentally, it may be noted that Oshima (1916) reported brood-care in Afrocucumis africana in Japan, which is unusual for a widespread, tropical echinoderm. | Juvenile specimens of three species (Actinopyga ?mau- ritiana, Labidodemas ?semperianum, and _ Stichopus ?variegatus) have been collected at Enewetak, and figures of their spicules have been included. Due to the element of doubt regarding their identification and the distinctiveness of their color and spicules, all three forms have been included in the key that follows. HOLOTHURIANS 265 KEY TO SPECIES 1, 10. . Spicules in body wall minute grains and slender, dichotomously branched rods (Fig. 1f) Body sausage-shaped; podia present, as pedicels (tube feet) and papillae; body wall sometimes thick and muscular, not sticky to touch; spicules tables, rods or plates, never anchors, anchor plates, or wheels 2 Body worm-like; podia absent; body wall thin, usually sticky to touch; spicules anchors, anchor plates, and wheels 20 . Tentacles dendritic (richly branched); pedicels in double rows along each ambulacrum; spicules in body wall large knobbed, lenticular, perforated plates and smooth, perforated plates and rods (Fig. 3a); color uniformly deep violet, almost black in life, gray-brown in alcohol; body length up to 7 cm (Fig. 6a) Afrocucumis africana (Semper) Tentacles peltate or peltato-digitate (leaf-shaped); podia usually clearly differentiated into ventral pedi- cels and dorsal papillae 3 . Gonads in two tufts, one on either side of dorsal mesentery; body stout, flattened ventrally into dis- tinct sole, squarish in cross section or arched dorsally, with large dorsal and lateral papillae; among spicules either C-shaped rods or dichotomously branched rods 4 Gonads in single tuft to left of dorsal mesentery; body either cylindrical or arched dorsally and flat- tened into sole ventrally, with numerous, small, dorsal papillae; spicules tables, flat buttons, and/or branched rods, never C-shaped rods nu Spicules tables, C- or S-shaped rods, perforated rods and, in some forms, rosettes . Dorsal papillae united into groups of two to eight, the largest groups forming stellate figures; color of specimens at Enewetak yellowish red, reddish brown, or light brown; in some specimens, transverse reddish black lines between dorsal papillae; body length up to 70 cm (Fig. 5c) Thelenota ananas (Jaeger) Dorsal surface with numerous small and some large papillae, several papillae often joined basally but not forming stellate figures; color light gray or yellowish gray with spots of tan, brown, or red and, in some specimens, a broken network of reddish black between large papillae; body length up to 72.5 cm (Fig. 5a, b) Thelenota anax H. L. Clark . Rosettes absent or, at most, very rare; tables small with small disk; no large tack-like tables (Fig. 1d); dorsal papillae limited to ambulacra; color in life usually uniformly very dark green except dorsal papillae tipped with orange or scarlet; at Enewetak, some specimens dark olive to blue-black; body length up to about 30 cm (Fig. 5d) Stichopus chloronotus Brandt Rosettes common; tack-like tables present or absent 7 . In the papillae, large robust tables with conical, usually single-pointed spire and large disk with numerous perforations (Fig. 1c); color dark olive green mottled with deep brownish green in life, dull ocher in alcohol; body length up to about 16 cm (Fig. 5e) Stichopus horrens Selenka No large tack-like tables in papillae 8 . Tables (in specimen examined) small, 35 to 42 um high and wide; C- and S-shaped spicules 35 to 55 um long, rosettes 19 to 46 um long (Fig. 1a); color of specimen examined uniformly dark brown, almost black; colors reported in literature either variegated white or gray with brown and yellow, some specimens showing tinges of pink, lavender, or red, or uniformly dark or light brown; body length commonly 30 to 40 cm but apparently reaching more than 100 cm Stichopus ?variegatus Semper Tables (in specimen examined) 39 to 70 um high, 39 to 62 um across disk; C-shaped spicules up to 70 um long; rosettes to 54 um long (Fig. 1b); color mottled brown and orange-yellow dorsally, uni- formly dirty orange-yellow ventrally; body length 4.6 cm (Fig. 5f) Stichopus ?variegatus juvenile . Body wall with dichotomously branched X-shaped spicules (rosettes) and rods, never tables 10 Tables always among spicules 12 No large calcified anal teeth but five groups of anal papillae; spicules laterally smooth rods and rosettes (Fig. 3b); color brown-gray with golden brown or black-ringed darker blotches; body length up to about 30 cm (Fig. 6c) Bohadschia argus Jaeger Anus surrounded by five conspicuous, calcified teeth 11 (Key to Species continued on page 267.) 266 CUTRESS AND ROWE re 5 Ce es - ee S&S a SS — 100 ym 6 By : ae? LP exp ata IN 3 GE Sa: a b E F Fig. 1A, Stichopus ?variegatus Semper. a, tables, C-, S- and X-shaped spicules and rosettes, dorsal integument; b, table disk and supporting rods, tentacles. B, Stichopus ?variegatus, juvenile. a, tables, dorsal and ventral integument; b, C-shaped spicules and rosettes, dorsal integument; c, perforated rods, ventral pedicels; d, rods, tentacles. C, Stichopus horrens Selenka. a, large table, side view and disk of large table, dorsal papilla; b, smaller tables, side views, disks and tops of spires, dorsal integument; c, C-shaped spicules, dorsal and ventral integument; d, rosettes, dorsal integument; e, sup- porting rod, ventral pedicel (redrawn from Fisher, 1907, Pl. 70, Fig. 1). D, Stichopus chloronotus Brandt. a, tables, side view and tops of spires, dorsal and ventral integument; b, disks of tables, ventrum; c, d, rosette and C-shaped spicules, dorsal and ventral integument; e, rod, ventral pedicel (redrawn from Mitsukuri, 1912, text Fig. 27 and Theel, 1886, Pl. 7, Fig. 6). E, Holothuria pervicax Selenka. a, tables, side view and disk; b, table with rudimentary spire; c, rods of various shapes; d, larger rod; all from dorsal and ventral integument (redrawn from Fisher, 1907, Pl. 68, Fig. 2). F, Thelenota ananas (Jaeger). a, dichotomously branched rods, dorsal and ventral integument; b, minute, smooth granules, dorsal and ventral integument; c, spinous rods and perforate rods, ventral pedicels; d, rod, tentacle. HOLOTHURIANS 267 (Key to Species cont'd.) ile 12. 13. 14. 15. 16. 72 18. 19. Dorsal spicules laterally branched rods up to 120 um long and rosettes 38 to 54 yum long; in ventral body wall, small oval grains and rods without lateral projections (Fig. 3d); color light chocolate brown, usually mottled with grayish white; body length up to about 20 cm (Fig. 7d) Actinopyga mauritiana (Quoy and Gaimard) Dorsally, in single specimen examined, laterally branched rods up to 102 um long, rosettes to 77 um long; ventral spicules rods with lateral edges smooth, roughened, or finely branched, no oval grains (Fig. 3c); dorsal color yellowish white with three broad, irregular, transverse bands of brownish black, ventral surface uniformly yellowish white with bright yellow pedicels; body length 5.5 cm (Fig. 7e) Actinopyga ?mauritiana juvenile Calcareous ring ribbon-like, with radial pieces shorter than broad and interradials similar but tending to be curved; podia comparatively few, in double rows confined to ambulacra or, dorsally, more or less scattered 13 Calcareous ring stout and relatively strong, not ribbon-like, radial plates either as long as broad or longer than broad, not curved; podia crowded ventrally, numerous, and scattered dorsally 14 Spicules in body wall tables with spire ending in four to six long, simple or bifurcate, horizontal spines or four short, vertical spines, together with smooth rods with simple, branched, or perforated ends (Fig. 3e); 20 tentacles; color translucent pinkish white in life, opaque white in alcohol; body length up to 25 cm (Fig. 6d) Labidodemas semperianum Selenka Spicules in body wall incomplete or complete tables, the latter with spires of four (usually) or three pil- lars crowned with three or more forked spines or simple circlet of fine spines, also spiny or smooth dichotomously branched, X-shaped rods (Fig. 7f); up to 18 tentacles; color in life bright, translucent canary yellow with very fine, closely spaced brown specks and scattered small brown spots, in alcohol tannish yellow; body length up to 2.5 cm (Fig. 6b) Labidodemas ?semperianum Selenka Spicules in body wall tables with reduced disk, rosettes and small perforated plates (Fig. 2a); color uniformly purplish black; body length up to about 30 cm (Fig. 6g) Holothuria (Halodeima) atra Jaeger Spicules in body wall tables with well-developed, smooth or spinous disk and buttons, complete or incomplete 15 Buttons typically complete and regular; disk of tables with smooth rim 16 Buttons commonly incomplete and asymmetrical or reduced to knobbed bars; disk of tables with smooth or spinous rim 19 Buttons flat, thin, oval, with four or more pairs of small holes and with distinct median optical discon- tinuity; perforate plates in pedicels (Fig. 2d); color uniformly rich olive brown or reddish brown, lighter on ventral side; body length up to 10 cm (Fig. 6f) Holothuria (Platyperona) difficilis Semper Buttons not flat and thin; color not uniformly brown 7 Buttons 75 to 93 um long, with three pairs of holes; disk of tables squarish, 78 to 95 um wide; per- forated rods in pedicels up to 233 um long (Fig. 2f); color yellowish brown, with irregular, transverse bands or blotches of purplish brown dorsally; body length up to 15 cm (Fig. 7c) Holothuria (Thymiosycia) impatiens (Forsskal) Buttons, tables, and rods smaller 18 Buttons up to 78 um long, with three (usually) or four pairs of holes; disk of tables 62 to 68 um wide, spire ending in 20 or more teeth; pedicel rods up to 27 ym long (Fig. 2e); color white to tan with two longitudinal rows of brown spots dorsally and very fine, reddish brown spots scattered over all surfaces; body jength up to about 20 cm (Fig. 7a, b) Holothuria (Thymiosycia) arenicola Semper Buttons up to 78 um long, with three to five pairs of holes, disk of tables 62 to 75 um wide, spire ending in fewer than 20 teeth; pedicel rods up to about 190 um long (Fig. 2c); color in life yellowish brown, with yellow-tan dorsal papillae; body length up to 30 cm (Fig. 6e) Holothuria (Thymiosycia) hilla Lesson Tables with smooth but sometimes uneven rim, spire reduced; buttons reduced, often as knobbed bars (Fig. le); color mottled cream and brown or gray, with some yellow, base of dorsal papillae with red ring; body length to about 20 cm (Fig. 6h) Holothuria (Mertensiothuria) pervicax Selenka Tables well-developed, top of spire with eight to 12 spines, disk usually spinous; some buttons regular with three pairs of holes, others incomplete and asymmetrical (Fig. 2b); color purplish brown to almost black; body length up to about 20 cm (Fig. 6i) Holothuria (Mertensiothuria) leucospilota (Brandt) (Key to Species continued on page 271.) 268 CUTRESS AND ROWE "pre AMES : : ; mg Ih oC 5G SO pO b ene we © d . oo ba ® LS { \7 CO £7 : i A c (ok) A & fH) Bd SK SOO 00 00 0o 09) \0v) & wes a b Fig. 2A, Holothuria (Halodeima) atra Jaeger. a, tables, side views, disks, and a crown; b, X-shaped and irregular rod-like spicules, ventral integument; c, small granule-like deposits, tentacles. B, Holothuria (Mertensiothuria) leucospilota (Brandt). a, tables; b, buttons and rods, dorsal and ventral integument; c, rods, tentacles. C, Holothuria (Thymiosycia) hilla Lesson. a, b, tables and buttons, dorsal and ventral integument, and perforated rod, dorsal papilla; c, rods, tentacles. D, Holothuria (Platyperona) difficilis Semper. a, b, tables and buttons, dorsal and ventral integument; c, perforated plate, dorsal papilla; d, rods, tentacles. E, Holothuria (Thymiosycia) arenicola Semper. a, b, c, tables, buttons and perforated rods, dorsal and ventral integument; d, rods, tentacles. F, Holothuria (Thymiosycia) impatiens (Forskal). a, b, c, tables, buttons, perforated rods, dorsal and ventral integument; d, rods, tentacles. HOLOTHURIANS 269 Sh Os & GR, sy Be a a AK ~s9 LG ee 2 Say co [= we oes) e—— 309 NF we Be greece 5 AN oD 3 ANGLO 0 EN AG SS BS rah atevad BER 2 Mivisctce of Feo en CRS ENTE b x 3 (93 Re ¢ b LASYoro ose? PLE 9) “Cop TOR c oye a) Fa WaiGc2 ADSENSE, c Bons A ea7 Sy aes PL pe XY 100 pm _, B B-Da,c,d,E,G-H OSy i S ‘0, & <5 mM ae BU a o eat Gt mw A Baty “TEAS Bi, SSA % ) « os q “GN Savy, NEE || Gry oo Den as , Ys BW fh dd ee se a \. g fe | YD “> < {ww AES : mere 2 a Ss ae Al wy) b s \} et b Y c f¢ A Be Cc C " a D 200 um A,Fec Db ,Fa,b Fig. 3A, Afrocucumis africana (Semper). a, thick, perforated and spined disks, dorsal and ventral integument; b, rods, dorsal integument; c, rods, tentacles. B, Bohadschia argus Jaeger. a, rosettes and rod, dorsal integument and papilla; b, rod and rosettes, ventral integument; c, rods, tentacles. C, Actinopyga ?mauritiana (Quoy and Gaimard). a, rods and rosettes, dorsal integument; b, rods, ventral integument; c, rods, tentacles. D, Actinopyga mauritiana (Quoy and Gaimard). a, rods and rosettes, dorsal integument; b, rods and small grains, ventral integu- ment; c, rods, tentacles. E, Labidodemas semperianum Selenka. a, tables, disks, side view and from crowns, dorsal and ventral integument; b, rods, ventral integument; c, rods, tentacles. F, Labidodemas ?semperianum, juvenile. a, tables, complete and incomplete, and X-shaped spicules, spined and smooth, dorsal and ventral integument; b, rods, tentacles. 270 CUTRESS AND ROWE ) a) s © 6 @ Aki ° ® b 7 is) = VS oe G b Cc a A B 100 um o,7 S gy Cc @ (G Cc b ae Ne Yo oy of og d - D ig. 4A, Chiridota rigida Semper. a, b, wheels, rods and grains, body wall and integument; c, rods, tentacles. B, Euapta godeffroyi (Semper). a, b, c, anchor, anchor plate and rosettes, body wall and integument; d, rod, tentacles (as reported and figured by Fisher, 1907, p. 722, Pl. 81, Fig. 3b; not found in Enewetak specimen nor mentioned by Semper). C, Pendekaplectana nigra (Semper). a, b, c, anchor, anchor plate and rosettes, body wall and integument. D, Synapta maculata (Chamisso and Eysenhardt). a, b, c, d, anchor, young anchors, anchor plate and rosettes, body wall and integument. HOLOTHURIANS 271 Fig. 5A, B, Thelenota anax H. L. Clark. Live specimen on reef at Enewetak, about 55 cm long (photographed by Janet Lamberson). C, Thelenota ananas (Jaeger). Live specimen from Enewetak, about 55 cm long, partially contracted (photographed by Janet Lamberson). D, Stichopus chloronotus Brandt. Live specimen in situ, Enewetak (photographed by Scott Johnson). E, Stichopus horrens Selenka. Live specimen from Hawaii, 20 cm long. F, Stichopus ?variegatus Semper, juvenile. Preserved specimen from Enewetak, 5 cm long. (Key to Species cont'd.) 20. Spicules in body wall wheels and small, curved rods and smooth grains; 11 to 12 tentacles (Fig. 4a); 21. color translucent brownish red, with scattered, raised patches of white (accumulations of wheels) dorsally; body length up to 5 cm (Fig. 7f) Chiridota rigida Semper Spicules in body wall anchors and perforated anchor plates, never wheels; 15 tentacles 21 Stock of anchors with six to eight minutely spined branches, rosettes granular, subcircular, with hole in center (Fig. 4b); color yellowish grayish white, with greenish or brown stripes or blotches dorsally; body length up to about 40 cm (Fig. 7h) Euapta godeffroyi (Semper) Stock of anchors unbranched 22 272 CUTRESS AND ROWE 22. Anchors very large, 85 to 100 um long and 60 to 70 um across flukes; anchor plates also large, up to 75 um long; rosettes small dichotomously branched rods (Fig. 4d); color yellow-brown to grayish white with or without pattern of dark gray spots; body length up to about 2 m (Fig. 7i) Synapta maculata (Chamisso and Eysenhardt) Anchors and plates not massive, up to 30 um and 22 um long, respectively; rosettes small, irregular, perforate buttons (Fig. 4c); color translucent reddish brown, with many small, diffuse white spots; body length to about 20 cm (Fig. 7g) APPENDIX Collection Records These records are only of holothurians collected for the reference collections at the Enewetak Marine Biological Laboratory (EMBL) (later the Mid-Pacific Marine Labora- tory [MPML]). Most of the specimens were collected in 1955 by Charles E. Cutress at snorkeling depths and were identified by Bertha Cutress. During subsequent years, a few specimens were added to the collections by Robert W. Hiatt, J. J. Naughton, J. S. Pearse, G. A. Hiatt, and Janet Lamberson. These were identified by F. W. E. Rowe, B. Cutress, and J. Lamberson. Records of Enewetak holothurians mentioned in published literature (Table 1) are not included here. Afrocucumis africana Aniyaanii (Ananij): May 8, 1955, inner reef, attached to under surfaces of loose coral or rock. Enewetak: under rocks around quarry. Actinopyga mauritiana Medren: May 10, 1955, inner reef, between coral in tide pools and surge channels. Two commensal worms the same color as the ground color of the holothurian were found among the tube feet of one specimen and were preserved separately. Seen but not collected in 1955 at Aniyaanii (Ananij), Igurin, Runit, and Japtan. Enewetak: ocean reef off runway. Pendekaplectana nigra (Semper) Actinipyga ?mauritiana juvenile Aniyaanii (Ananij): May 11, 1955, inner reef, under rub- ble. Bohadschia argus Nancy (Elle): lagoon side, 2.7 m. Holothuria (Thymiosycia) arenicola Runit: June 20, 1955, buried in sand under loose coral on causeway. Sand Island: channel. Holothuria (Halodeima) atra Runit: June 20, 1955, in open on sandy or coral bottom of tide pools. Seen but not collected in 1955 at Igurin, Aniyaanii (Ananij), Medren, and Japtan. Enewetak: ocean reef north of island. Holothuria (Platyperona) difficilis Aniyaani (Ananij): May 8 and June 21, 1955, inner reef attached to under surfaces of coral or rubble. Enewetak: under rocks around quarry. Holothuria (Thymiosycia) hilla Aniyaanii (Ananij): June 21, 1955, attached to under surfaces of coral or rubble. Enewetak: ocean side, north end, under rocks just below beach; Sand Island; Enjebi: ocean side. Holothuria (Thymiosycia) impatiens Runit: June 20, 1955, under rock on causeway. Holothuria (Mertensiothuria) leucospilota Runit: June 20, 1955, under loose coral in pools on causeway. Igurin: May 11, 1955, inner reef, under rub- ble. Seen but not collected in 1955 at Aniyaanii (Ananij), Medren, and Japtan. Enewetak: ocean side, north end. (Collection records continued on page 275.) (Fig. 6 on page 273.) ———» Fig. 6A, Afrocucumis africana (Semper). Preserved specimen from Enewetak, 4.7 cm long. B, Labidodemas ?semperianum, juvenile. Preserved Enewetak specimens, 2.25 and 2.5 cm long. C, Bohadschia argus Jaeger. Live specimen in situ; Cocos Lagoon, Guam (photographed by Charles Birkeland). D, Labidodemas semperianum Selenka. Preserved Enewetak specimen, 20 cm long. E, Holothuria (Thymiosycia) hilla Lesson. Live specimen from Hawaii, 19 cm long. F, Holothuria (Platyperona) difficilis Semper. Preserved Enewetak specimen, 6 cm long. G, Holothuria (Halodeima) atra Jaeger. Live specimen from Hawaii, 30 cm long. H, Holothuria (Mertensiothuria) pervicax Selenka. Live specimen from Hawaii, 18 cm long. I, Holothuria (Mertensiothuria) leucospilota (Brandt). Live specimen from Hawaii, 20 cm long. (Fig. 7 on page 274.) ———» Fig. 7A, B, Holothuria (Thymiosycia) arenicola Semper. a, preserved specimen from Enewetak, 8 cm long; b, live specimen from Hawaii, 19 cm long. C, Holothuria (Thymiosycia) impatiens (Forskal). Preserved Enewetak specimen, 17.5 cm long. D, Actinopyga mauritiana, juvenile. Preserved specimen from Hawaii, 20 cm long. E, Actinopyga ?mauritiana, juvenile. Preserved Enewetak specimen, 5.5 cm long. F, Chiridota rigida Semper. Preserved Enewetak specimen, 5 cm long. G, Pendekaplectana nigra (Semper). Preserved Enewetak specimen, 12 cm long. H, Euapta godeffroyi (Semper). Preserved Enewetak specimen, 21 cm long. I, Synaptula maculata (Chamisso and Eysenhardt). Small preserved specimen from Enewetak, 21 cm long. HOLOTHURIANS 273 274 CUTRESS AND ROWE Fig. 7 HOLOTHURIANS 275 Holothuria (Mertensiothuria) pervicax Runit: June 20, 1955, under loose coral in pools on ocean side of causeway. Labidodemas semperianum Aniyaanii (Ananij): June 21, 1955, buried in sand under rock near causeway. Aaraanbiru (Alembel): in sand on sea reef. Labidodemas ?semperianum juvenile Aniyaanii (Ananij): June 21, 1955, attached to under- side of coral rock. Stichopus chloronotus Enjebi: northwest ocean reef between Bokoluo and Oak craters, in open among coral heads and rubble, also in shallow quarries, intertidal to 2 m. Lojwa (Ursula): June 21, 1963, extremely abundant on reef flat, usually in open alongside Holothuria atra; heavily infested with snails in body wall; one Carapus found out of four speci- mens. Stichopus horrens Enewetak: July 19, 1963, quarry, under live coral. Stichopus ?variegatus Aaraabiru (Alembel): sea reef near shore. Stichopus ?variegatus juvenile Medren: June 21, 1955, inside piece of coral. Thelenota ananas Medren: September 1956, sea reef channel. On rubble at top or base of pinnacles, also near concrete ship 10 m depth. Japtan: Oct. 5, 1975, near concrete ship, 10 m depth; also around pinnacles and patch reefs in lagoon. Enewetak: top of sunken barge in lagoon, 1 m depth; one specimen had spotted commensal crab, col- lected as it tried to enter holothurian’s mouth. Thelenota anax Enewetak: lagoon side; Sand Island; Medren; Japtan: on top or bottom of rock pinnacles, 5 to 30 m. Chiridota rigida Aniyaanii (Ananij): June 21, 1955, in groups buried in sand beneath coral or rubble or between coral in tide pools. Sand Island. Euapta godeffroyi Aniyaanii (Ananij): May 8, 1955, inner reef, in or on sand under loose coral or rubble. Runit: June 20, 1955; Enewetak: quarry. Pendekaplectana nigra Aniyaani (Anaij): May 8, 1955, inner reef, under rubble. Runit: June 20, 1955, ocean side of causeway, under loose coral. Synapta maculata Runit: June 20, 1955, ocean side of causeway, under loose coral in pool. Enjebi: Cactus Crater. REFERENCES Bakus, G. J., 1968, Defensive Mechanisms and Ecology of Some Tropical Holothurians, Mar. Biol., Berlin, 2: 23-32. ___, 1973, The Biology and Ecology of Tropical Holothurians, Biology and Geology of Coral Reefs, Vol. Il, Biology 1, O. A. Jones and R. Endean (Eds.), Academic Press, New York and London, pp. 325-367. , and G. Green, 1974, Toxicity in Sponges and Holothurians, a Geographic Pattern, Science, 185: 951-953. Cherbonnier, G., 1970, Note sur |’Holothurie Aspidochirote Labidodemas semperianum Selenka, Bull. Mus. Natl., Hist. Nat., Paris, Ser. 2, 42: 566-569, Figs. A-P. Clark, A. M., and F. W. E. Rowe, 1971, Monograph of Shallow- Water Indo-West Pacific Echinoderms, British Museum (Natural History), London, pp. i-vii, 1-238. Clark, H. L., 1921, The Echinoderm Fauna of Torres Strait; Its Composition and Its Origin, Pap. Dep. Mar. Biol., Carnegie Inst. Wash., 10: 1-223. —_, 1946, The Echinoderm Fauna of Australia; Its Composition and Its Origin, Carnegie Inst. Wash. Pub., 566: 1-567. Ebert, T. A., 1978, Growth and Size of the Tropical Sea Cucumber Holothuria (Halodeima) atra Jaeger at Enewetak Atoll, Marshall Islands, Pac. Sci., 32: 183-191. Hartman, O., 1954, Marine Annelids from the Northern Marshall Islands, Geol. Surv. Prof. Pap. 260-Q, pp. 619-644. Humes, A. G., 1973, Nanaspis (Copepoda: Cyclopoida) Parasitic on the Holothurian Thelenota ananas (Jaeger) at Eniwetok Atoll, J. Parasit., 59: 384-395. Lamberson, J. O., 1978, Notes on the Morphology, Ecology, and Distribution of Thelenota anax H. L. Clark (Holo- thuroidea: Stichopodidae), Micronesica, 14: 115-122. Lawrence, J. M., 1980, Numbers and Biomass of Common Holothuroids on the Windward Reef Flat at Enewetak Atoll, Marshall Islands, in Echinoderms: Past and Present, Proceed- ings of European Colloquium on Echinoderms, Brussels, September 3-9, 1979, M. Jangoux (Ed.), A. A. Balkema. Rotterdam, pp. 201-204. Lawrence, J. M., and A. Guille, 1982, Organic Composition of Tropical, Polar and Temperate-Water Echinoderms, Comp. Biochem. Physiol., 72B: 283-287. Pomeroy, L. R., and E. J. Kuenzler, 1967, Phosphorus Turnover by Coral Reef Animals, in Symposium on Radioecology, D. J. Nelson and F. C. Evans (Eds.), Ann Arbor, Michigan, pp. 474-482. Reish, D. J., 1968, The Polychaetous Annelids of the Marshall Islands, Pac. Sci., 22 : 208-231. Rowe, F. W. E., and J. E. Doty, 1977, The Shallow-Water Holothurians of Guam, Micronesica, 13: 217-250. Smith, A. C., R. L. Taylor, H. Chun-Akana, and F. Ramos, 1973, Intestinal Tumor in the Sea Cucumber, Holothuria leu- cospilota, J. Invertebr. Pathol., 22: 305-307. Webb, K. L., W. D. Du Paul, and C. F. D’Elia, 1977, Biomass and Nutrient Flux Measurements of Holothuria atra Popula- tions on Windward Reef Flat at Enewetak, Marshall Islands, Proceedings of the Third International Coral Reef Symposium, Miami, Florida, 1977, pp. 409-415. Chapter 25 Echinodermata Other than Holothuroidea of Enewetak Atoll DENNIS M. DEVANEY (deceased) Bernice P. Bishop Museum Honolulu, Hawaii 96817 INTRODUCTION The echinoderm fauna at Enewetak became known through publications by A. H. Clark (1952, 1954) that recorded nonholothurian species from several of the Marshall Islands. These two publications included nine Asteroidea, nine Ophiuroidea, and 19 Echinoidea from Enewetak. The Crinoidea and Holothuroidea were not recorded from the atoll. In the late 1950s and early 1960s, Fred C. Ziesen- henne, then with the Allan Hancock Foundation (AHF), was asked by the Mid-Pacific Research Laboratory (MPRL) to provide an identified reference collection of nonho- lothurian echinoderms. His identifications, based on speci- mens that he collected and those collected by students and researchers working at MPRL, provided not only a valu- able reference collection but also increased the number of taxa known from Enewetak. An unpublished report, which includes habitat information and keys to many of the species, was prepared by FF. C. Ziesenhenne and R. A. Boolootian and was left at the laboratory for use by visiting researchers. The holothurians were being studied separately by Bertha Cutress, and her work has been updated in a separate chapter for the present volume (see Cutress and Rowe, Chapter 24). Since these early efforts, new studies and additional records of echinoderms have been compiled by MPRL per- sonnel and other researchers working at Enewetak. This chapter brings together these numerous records (Table 1). Although the majority of Enewetak’s shallow-water (to about 100 m) echinoderms is reasonably well known, fauna from deeper waters, especially in passes and off the sea- ward reef face, is still incompletely known. Less than 1% of the species so far recorded have been taken in water deeper than 100 m. In comparison with the Marshall Islands as a whole, knowledge of the Enewetak echinoderms is considerable. 277 At this one atoll, 86% of all Marshall Islands species are now known to occur at Enewetak. CRINOIDEA Although crinoids had been reported from the Marshall Islands by Gislén (1940) and A. H. Clark (1952, 1954), no species from Enewetak were included. Crinoids collected in the late 1950s and identified by F. C. Ziesenhenne included Comanthus bennetti and Comaster gracilis, species recorded previously in the Marshall Islands from Rongelap and Bikini by A. H. Clark (1952) and from Arno by A. H. Clark (1954). Banner and Banner (1968) reported the alpheid shrimp, Synalpheus demani, from C. bennetti col- lected at Enewetak. Zmarzly (personal communication) doubts the association of alpheids with this crinoid and, based on her studies, feels that the host was a different species. One of the Enewetak specimens of C. bennetti examined by the author and Ann Fielding had a single alpheid, S. stimpsoni, associated; two other alpheids were also associated with other crinoid species at Enewetak (Devaney and Bruce, Chapter 22, this volume). One of these, S. carinatus, was also noted in association with “cri- noids” at Enewetak by Banner and Banner (1968). Humes (1972) reported a new species of copepod from C. ben- netti, and Bartolini et al. (1973) examined pigments from Enewetak specimens of this crinoid. In 1976, Devaney and Fielding examined Enewetak cri- noids for their symbionts; crustaceans, polychaetes, myzos- tomes, and other invertebrates were noted and collected. It was thought that only Comanthus bennetti and Comaster gracilis were represented in the material examined. Subse- quently, however, due to a more intensive study on Enewetak crinoid ecology and their symbionts by Zmarzly in 1980, two additional species (Comanthina schlegeli and Comanthus parvicirrus) were recognized among the 1976 material. The crustaceans associated with these crinoids have been tabulated in the present volume (Devaney and Bruce, pp. 225-226), and more information was given in Zmarzly (1984). The small 10-armed antedonid, Dorometra nana, was collected in 1976 off one of the southern wind- ward islets (Ananij) beneath coral in limestone rubble at 18 278 DEVANEY TABLE 1 Checklist of Echinodermata Other than Holothuroidea from Enewetak Atoll Phylum ECHINODERMATA Subphylum CRINOZOA Class CRINOIDEA Family COMASTERIDAE Comanthina schlegeli (Carpenter, 1881): Zmarzly, 1984. Comanthus bennetti (Miiller, 1841): Banner and Banner, 1968; Humes, 1972; Bartolini et al., 1973; Zmarzly, 1984. Comanthus parvicirrus (Miiller, 1841): Zmarzly, 1984. Comaster gracilis (Hartlaub, 1890); Zmarzly, 1984. Family EUDIOCRINIDAE Eudiocrinus tenuissimus Gislén, 1940: Zmarzly, 1984. Family ANTEDONIDAE Dorometra nana (Hartlaub, 1890): Zmarzly, 1984. Subphylum ASTEROZOA Class STELLEROIDEA Subclass ASTEROIDEA Family ASTROPECTINIDAE Astropecten polyacanthus Miiller and Troschel, 1842: A. H. Clark, 1952. Family OREASTERIDAE *Choriaster granulatus Liitken, 1869. Culcita novaeguineae Miiller and Troschel, 1842: A. H. Clark, 1952; Humes, 1971; Bruce, 1979. Family OPHIDIASTERIDAE *+Fromia hemiopla Fisher, 1913. Fromia milleporella (Lamarck, 1816). +Fromia balansae Perrier, 1875: A. H. Clark, 1952. * Fromia monilis Perrier, 1875. * Gomophia egyptiaca Gray, 1840. * Leiaster speciosus von Martens, 1866. Linckia laevigata (Linnaeus, 1758): A. H. Clark, 1952. Linckia multifora (Lamarck, 1816): A. H. Clark, 1952; 1954. * Neoferdina cumingi Gray, 1840. *Ophidiaster granifer Liitken, 1871: A. H. Clark, 1952. Ophidiaster robillardi de Loriol, 1885: Marsh, 1977. Ophidiaster lorioli Fisher, 1906: A. H. Clark, 1952. Family ASTEROPSEIDAE *Asteropsis carinifera (Lamarck, 1816). Family ASTERINIDAE *§Asterina anomala H. L. Clark, 1921. Family ASTERINIDAE Asterina burtoni Gray, 1840: A. M. Clark and Rowe, 1971. Asterina cephea (Miiller and Troschel, 1842): A. H. Clark, 1952. Family ACANTHASTERIDAE Acanthaster planci (Linnaeus, 1758): Allen, 1972; Highsmith, 1980. Family MITHRODIDAE *Mithrodia clavigera (Lamarck, 1816). Family ECHINASTERIDAE Echinaster luzonicus (Gray, 1840). Othilia luzonica Gray: A. H. Clark, 1952; Lawrence and Guille, 1982. Subclass OPHIUROIDEA Family OPHIOMY XIDAE * Neoplax crassipes Koehler, 1922. * Ophiomyxa australis Liitken, 1869. Family GORGONOCEPHALIDAE *Astroboa nuda (Lyman, 1874). *New record at Enewetak. tEnewetak specimens determined by F. C. Ziesenhenne at AFH (1 dry) and MPRL; those reported as E. hemiopla from Bikini by A. H. Clark (1952) probable juveniles of other species (L. M. Marsh, personal communication.) +Considered probably referrable to F. milleporella after reexamination by L. M.-Marsh (personal communication). §Considered as possible syn. of Asterina burtoni by A. M. Clark and Rowe, 1971, p. 68. (This table continued on next page.) ECHINODERMATA OTHER THAN HOLOTHUROIDEA 279 TABLE 1 (cont’d) Family AMPHIURIDAE * Amphiodia sp. * Amphipholis squamata (Delle Chiaje, 1829). Amphiura luetkeni Duncan, 1879: Highsmith, 1981. Family OPHIACTIDAE Ophiactis savignyi (Miiller and Troschel, 1842): A. H. Clark, 1952; Highsmith, 1981. Family OPHIOTRICHIDAE * Macrophiothrix longipeda (Lamarck, 1816). * Macrophiothrix rhabdota H. L. Clark, 1915. Macrophiothrix sp.: Highsmith, 1981. * Ophiothela danae Verrill, 1869. Ophiothrix (Ophiothrix) exigua Lyman, 1874: A. H. Clark, 1952. * Ophiothrix (Acanthophiothrix) proteus Koehler, 1905. * Ophiothrix (Acanthophiothrix) purpurea von Martens, 1867. * Ophiothrix (Ophiothrix) trilineata (Liitken, 1869). Family OPHIOCOMIDAE *Ophiarthrum elegans Peters, 1851. Ophiocoma anaglyptica Ely, 1944: A. H. Clark, 1952; Devaney, 1967, 1970; Chartock, 1972; Sloan et al., 1979. Ophiocoma brevipes Peters, 1851: A. H. Clark, 1952; Devaney, 1970; Chartock, 1972; Sloan et al., 1979. Ophiocoma dentata Miiller and Troschel, 1842: Devaney, 1970; Chartock, 1972. Ophiocoma doederleini Loriol, 1899: Pomeroy and Kuenzler, 1967; Devaney, 1970; Chartock, 1972; Sloan et al., 1979. Ophiocoma erinaceus Miiller and Troschel, 1842: A. H. Clark, 1952; Chartock, 1972; Sloan et al., 1979. Ophiocoma pica Miiller and Troschel, 1842: A. H. Clark, 1952; Chartock, 1972; Sloan et al., 1979. Ophiocoma pusilla (Brock, 1888): Devaney, 1970. Ophiocoma scolopendrina (Lamarck, 1816): A. H. Clark, 1952; Chartock, 1972; Sloan et al., 1979. Ophiocomella sexradia (Duncan, 1887): Highsmith, 1981. Ophiocomella clippertoni A. H. Clark, 1939: A. H. Clark, 1952. Ophiomastix caryophyllata Liitken, 1869: Devaney, 1978. Ophiomastix marshallensis Devaney, 1978: Devaney, 1978. Ophiomastix mixta Liitken, 1869: A. H. Clark, 1952; Devaney, 1978. Ophiomastix palaoensis Murakami, 1943: Devaney, 1978. Ophiomastix variabilis Koehler, 1905: Devaney, 1978; Highsmith, 1981. Ophiopsila sp.: Highsmith, 1981. Family OPHIONEREIDIDAE Ophionereis porrecta Lyman, 1860: Highsmith, 1981. *Ophionereis sp. Family OPHIODERMATIDAE * Ophiarachna affinis Liitken, 1869. * Ophiarachnella gorgonia (Miiller and Troschel, 1842). Ophiarachnella parvispina H. L. Clark, 1925: Devaney, 1974. * Ophioconis permixta Koehler, 1905. Ophiopeza spinosa (Ljungman, 1867): Devaney, 1974. Family OPHIURIDAE *Ophiolepis superba H. L. Clark, 1915. JOphiolepis sp. Ophioplocus imbricatus (Miiller and Troschel, 1842): Lawrence and Guille, 1982. *Ophiura kinbergi (Ljungman, 1867). Subphylum ECHINOZOA Class ECHINOIDEA Family CIDARIDAE *Chondrocidaris gigantea A. Agassiz, 1863. Eucidaris metularia (Lamarck, 1816): A. H. Clark, 1952. Family ECHINOMETRIDAE Echinometra mathaei (de Blainville, 1825): A. H. Clark; 1952; Pomeroy and Kuenzler, 1967; Lawrence, 1970; Russo, 1980; Ebert, 1982. “New record at Enewetak. {Within the Ophiolepis cincta complex of species under investigation by the author. (This table continued on next page.) 280 DEVANEY TABLE 1 (cont’d) Family ECHINOMETRIDAE (cont'd) Echinometra oblonga (de Blainville, 1825). Echinometra mathaei var. oblonga (de Blainville): A. H. Clark, 1952. Echinostrephus aciculatus A. Agassiz, 1863: A. H. Clark, 1952; Pomeroy and Kuenzler, 1967; Russo, 1980; Lawrence, and Guille, 1982; Ebert, 1982. *Heterocentrotus mammillatus (Linnaeus, 1758). Heterocentrotus trigonarius (Lamarck, 1816): A. H. Clark, 1952; Odum and Odum, 1955; Lawrence and Dawes, 1969; Humes, 1970; Ebert, 1982. Family DIADEMATIDAE *Diadema savignyi Michelin, 1845. Echinothrix calamaris (Pallas, 1774): A. H. Clark, 1952; Humes, 1977. Echinothrix diadema (Linnaeus, 1758): A. H. Clark, 1952; Lawrence and Guille, 1982; Ebert, 1982. Family PARASALENIIDAE Parasalenia gratiosa A. Agassiz, 1863: A. H. Clark, 1952; Highsmith, 1981. Family TEMNOPLEURIDAE Mespilia globulus (Linnaeus, 1758): A. H. Clark, 1952. Family TOXOPNEUSTIDAE Cyrtechinus verruculatus (Liitken, 1864): Highsmith, 1981. *Toxopneustes pileolus (Lamarck, 1816). Tripneustes gratilla (Linnaeus, 1758): A. H. Clark, 1952. Family ECHINONEIDAE Echinoneus abnormalis de Loriol, 1883: A. H. Clark, 1952. Echinoneus cyclostomus Leske, 1778: A. H. Clark, 1952. Family SCHIZASTERIDAE *Diploporaster savignyi (Fourtau, 1904). Family FIBULARIIDAE Echinocyamus australis (Desmoulins, 1835). Fibularia australis Desmoulins: A. H. Clark, 1952. *Echinocyamus crispus Mazetti, 1893. Echinocyamus megapetalus H. L. Clark, 1914: A. H. Clark, 1952. Fibularia ovuulum Lamarck, 1788:.A. H. Clark, 1952. Fibularia volva L. Agassiz and Desor, 1847: A. H. Clark, 1952. Family SPATANGIDAE Maretia planulata (Lamarck, 1816). Maretia ovata (Leske, 1778): A. H. Clark, 1952. Family BRISSIDAE Brissus latecarinatus (Leske, 1778): Pomeroy and Kuenzler, 1967. Metalia dicrana H. L. Clark, 1917: A. H. Clark, 1952. *Metalia spatagus (Linnaeus, 1758). Rhynobrissus hemiasteroides A. Agassiz, 1879: Pomeroy and Kuenzler, 1967. Family CLYPEASTERIDAE *Clypeaster reticulatus (Linnaeus, 1758). Family LAGANIDAE Laganum depressum L. Agassiz, 1841: A. H. Clark, 1952. “New record at Enewetak. to 21 m and was identified by the author. Additional speci- mens of this species have been found under coral rubble in the lagoon. Dorometra nana had been recorded from Ebon and Jaluit by Gislén (1940). In 1980 at Enewetak, a five-armed Eudiocrinus, tentatively identified as E. tenuis- simus, was found in recesses on the steep seaward dropoff near Rigili islet. Gislén (1940) recorded this species origi- nally from Jaluit Atoll in the Marshalls. Six crinoid species are now known from Enewetak shal- low waters; all but one (E. tenuissimus) occur in other Indo-West Pacific areas outside the Marshall Islands (A. M. Clark and Rowe, 1971). Additional crinoids recorded from other Marshall Islands include: Cenometra bella var. magnifica from Jaluit (Gislén, 1940), as well as Stephanometra indica (as S. protectus) from Ebon and Jaluit (Gislén, 1940) and (as S. indica protectus) from Rongelap and Rongerik (A. H. Clark, 1952). Six of the nine crinoids known from the Marshall Islands have been found at Enewetak. Seven species are known from Guam and 21 from Palau (Meyer and Macurda, 1980); Enewetak shares four species with Guam and five with Palau. ECHINODERMATA OTHER THAN HOLOTHUROIDEA 281 OPHIUROIDEA Gorgonocephalidae A basket star, Astroboa nuda, is the first species in this family to be recorded from the Marshall Islands. One speci- men was found at a depth of 13 m in a cave on the sea- ward side just north of Kidrenen Islet on the southwest side of Enewetak. Ophiomyxidae The Ophiomyxidae are represented by two species at Enewetak. One, Ophiomyxa australis, is known from Bikini and Rongerik (A. H. Clark, 1952) and Jaluit (as O. brev- ispina in Koehler, 1927). The other is a single specimen of Neoplax crassipes, known from the Philippines and was collected in shallow water. Amphiuridae Three amphiurids are now known from Enewetak. These include an undetermined Amphiodia collected from lagoon sediment. Amphipholis squamata, a cosmopolitan species, is common in shallow water around Enewetak; two specimens have been collected at Bikini (AHF). _Amphiura luetkeni is the only amphiurid previously recorded from the Marshall Islands (Highsmith, 1981). It is | often associated with live coral. Outside of the Marshall Islands, A. luetkeni is known from Ceylon to Polynesia. | Ophiactidae One ophiactid, the cosmopolitan species Ophiactis savignyi, has been recorded from Enewetak (A. H. Clark, 1952; Highsmith, 1981). It is often found in sponges. - Ophiotrichidae At least three genera of Ophiotrichidae are known from Enewetak. Macrophiothrix is represented by M. longipeda, originally reported from the Marshall Islands by A. H. Clark (1952) from Bikini; Devaney considers Koehler’s (1927) Ophiothrix galatheae from Jaluit as prob- ably referrable to this species. Another species is con- _sidered as M. rhabdota and is a new record for the Marshall Islands. Highsmith (1981) reported and identified _Macrophiothrix sp. Ophiothela danae is an associate of 'alcyonacean corals. At least four species of Ophiothrix -occur at Enewetak; only O. exigua was previously | recorded from the atoll. Opiothrix (Acanthophiothrix) pur- | purea was found on a gorgonian coral; A. H. Clark (1952) ) also reported this species (as O. lepida) from a gorgonian at Bikini Atoll. At Enewetak, O. trilineata is frequently associated with branching coral such as Pocillopora; A. H. Clark (1952) reported it previously from Bikini. The two specimens found at Enewetak were collected at the base of coral (Goniastrea) in the lagoon on a pinnacle (South | Medren) at a depth of 8.1 to 13.3 m. Ophiocomidae The ophiocomids are diverse at Enewetak represented by 16 species in five genera with a new record, Ophiarthrum elegans. One specimen of Ophiopsila remains to be identified to species. Ophionereididae Two species of the ophionereids are recognized: Ophionereis porrecta and Ophionereis sp., which resemble those recorded by A. H. Clark (1952) from Bikini Atoll as Ophionereis degeneri. Ophiodermatiidae Five ophiodermatids occur at Enewetak. Ophiarachna affinis, Ophiarachnella gorgonia, and Ophioconis permixta are new records for the Marshall Islands. Ophiopeza spi- nosa was recorded (as Ophiopezella spinosa by A. H. Clark, 1952) from Bikini and Rongerik Atolls. The only other ophiodermatid recorded from the Marshall Islands is Ophiopezella (Ophiopeza) dubiosa from Jaluit (Koehler, 1927). An additional specimen of the uncommon Ophiarachnella parvispina, reported from Enewetak by Devaney (1974), was located in the Allan Hancock Foun- dation from the intertidal lagoon, July 17, 1960. Ophiuridae Of the four ophiurids known from Enewetak only one, Ophioplocus imbricatus, a widespread warm water Indo- West Pacific species has been previously reported from this atoll. All four ophiurids have been reported from else- where in the Marshall Islands (A. H. Clark, 1952). The Collection Records list the habitats of the newly reported species of Ophiuroidea. Table 2 lists comments on ophiuroid color, anatomy, disc and arm length size, and repository. ASTEROIDEA Astropectinidae The family Astropectinidae is represented by a single widespread Indo-West Pacific species, Astropecten poly- acanthus, collected in the lagoon (A. H. Clark, 1952). Two small specimens were collected at Bikini Atoll in July 1946 and identified by F. C. Ziesenhenne (AHF). Oreasteridae The family Oreasteridae is represented by two species. Culcita novaeguineae has been collected on the ocean reef at Enewetak (A. H. Clark, 1952) and from the lagoon and channel. Symbiotic crustaceans were recorded from this species by Humes (1971) and Bruce (1979). A second oreasterid, Choriaster granulatus hitherto unknown from the Marshall Islands, was discovered during the Hawaii 282 DEVANEY TABLE 2 . Comments on Color, Disc and Arm Length, Size, and Repository Neoplax crassipes, color deep brown with white markings, one specimen dry with disc diameter (d.d) 13 mm, AHF. Ophiomyxa australis, color in alc. dull straw, arm spines tipped with red-brown, disc and mouth light gray, 1 alc. MPRL. Astroboa nuda, color above disc purplish black also main arm axes except lighter belts of hooklets, a darker band between each row, broader above and narrowly curved distally around sides of arms; narrow depressed longitudinal dark line down center of arms, more distal arm branches quite light in color; lower surface of arms lighter than above without dark bands or longitudinal line, a few distal branches with some dark color not uniform, 1 dry, d.d. 85 mm, MPRL. Ampbhiura luetkeni, red and white disc with reddish arms. Amphiodia sp., white disc, white arms, 1 dry, disc 2.5 to 3 mm, arm 60 mm, BPBM. Macrophiothrix longipeda, from Araanbiru, 1 alc. d.d. 23.5 mm, arm 300 mm, MPRL; from Engebi, 1 alc, d.d. 5.5 mm, arm 132 mm, MPRL; from Engebi, 1 spec., disc 13 mm, arm 155 mm, MPRL. Macrophiothrix rhabdota, 1 dry, d.d. 8 mm, arms broken, MPRL; 1 alc. d.d. 8 mm, arms broken, AHF. Ophiothela danae, disc blue or green, arms annulated with narrow band of light or dark across, 10+ dry, disc 0.7 to 4 mm usually with six arms or in fissiparous stage with only three arms, BPBM. Ophiothrix (Acanthophiothrix) proteus, 1 dry, d.d. 9 mm, arms 100 to 110 mm, BPBM. Ophiothrix (Acanthophiothrix) purpurea, color of wide, dark stripe or along middle or narrow dark stripe bordered by light lines on dorsal side of arms; arm spines banded dark and light; radial shields bare, but between, radially and interradially, either granular stumps or stumps and a few long spines, marginally and on ventral side, disc with stumps and spines widely spaced, 1 alc., d.d. 7 mm, BPBM. Ophiothrix (Ophiothrix) trilineata, from Enewetak Islet, 1 spec., gonads ripe; from Japtan Islet, 1 spec. gonads ripe; from Medren, 2 spec., gonads ripe. Ophianthrum elegans, living color dark black, arms red and whitish band with some small dark speckling between arm spines with irregular dark and light marks, from Enewetak, 1 alc., d.d. 12 mm, BPBM; Sand Islet, 2 alc., d.d. 10, d.d. 13 mm, BPBM; Lagoon, color dark black with red arms, 1 dry, BPBM. Ophionereis sp., from Enewetak Islet, 2 dry, d.d. 5, d.d. 6, BPBM. Ophiarachna affinis, 1 alc. d.d. 12 mm, MPRL. Ophiarachnella gorgonia, from Enewetak Islet, 1 dry, d.d. 12 mm, BPBM; from Rabaion Islet, 3 dry, d.d. 14 to 16 mm, AHF. Ophioconis permixta, 1 alc., d.d. 2 mm, BPBM. Ophiolepis superba, from Enewetak Islet, 1 alc., d.d. 24 mm, AHF. Ophiolepis sp. after O. cinta, from Enewetak Islet, 1 dry d.d. 13 mm, 1 dry, d.d. 11.5 mm, BPBM; from Lagoon, 1 spec. d.d. 7 mm, AHF. Ophiura kinbergi, from Engebi Islet, 1 juv., alc., d.d. 3 mm, MPRL. Undersea Research Laboratory dives in 1981 at a depth of 122 to 130 m. Ophidiasteridae This family has the largest number of species represented at Enewetak, with at least 10 now recorded. Four have been reported previously from the Marshall Islands; all are widespread Indo-West Pacific species. Asteropseidae The Asteropseidae is recorded for the first time in the Marshall Islands in the form of Asteropsis carinifera, a species which ranges in the Indo-West Pacific from Hawaii through Oceania and into the Indian Ocean. Asterinidae The Asterinidae at Enewetak are represented by two species and possibly a third (Asterina coronata) determined by F. C. Ziesenhenne. All three have been previously recorded from the Marshall Islands ‘A. H. Clark, 1952). Acanthasteridae Acanthaster planci, the crown-of-thorns sea star, is known at Enewetak from studies of its predatory activity on scleractinian corals (Highsmith, 1980, 1982) and a fish symbiont (Allan, 1972). ECHINODERMATA OTHER THAN HOLOTHUROIDEA 283 Mithrodiidae The family Mithrodiidae is newly recorded from Enewetak and the Marshall Islands, on the basis of a widespread Indo-West Pacific species, Mithrodia clavigera. Echinasteridae Echinaster luzonicus is known from Enewetak and else- where in the Marshall Islands (A. H. Clark, 1952). Lawrence and Guille (1982) recorded the dry weight organic composition of this sea star and other echinoderms from Enewetak. ECHINOIDEA Cidaridae The cidarids are represented by two species. Eucidaris metularia has been found in shallow water at Enewetak and Rongerik and to a depth of 88 m at Bikini (A. H. Clark, 1952). Specimens of Chondrocidaris gigantea were collected from approximately 130 m and confirm an earlier record from Bikini based on one spine (A. H. Clark, 1952). This species has been observed at Kwajalein by divers (S. Johnson, personal communication). Additional cidarid species were observed and photographed in deeper waters (to 360 m depth) off the seaward side of Enewetak during submersible dives in 1981. Echinometridae Three genera and species of Echinometridae occur at Enewetak. Four of the species have been reported from Enewetak previously, and these range widely in the Indo- West Pacific. The fifth species, Heterocentrotus mammil- latus, is now known from the atoll on the basis of one specimen. This species ranges from the Philippines to Hawaii and the Tuamotus. Banner and Banner (1960, 1968) reported collecting the caridean shrimp Athanas dor- salis from large Heterocentrotus sea urchins at Enewetak and elsewhere. They also reported another alpheid, A. indicus, associated with burrowing sea urchins in the genus Echinometra. Diadematidae The Diadematidae includes Diadema savignyi, a widely distributed Indo-West Pacific species, newly recorded from the Marshall Islands. Echinothrix calamaris and E. diadema are also known from Enewetak and others of the Marshall Islands (A. H. Clark, 1952, 1954). Parasaleniidae and Temnopleuridae One representative of both the Parasalenidae and Tem- nopleuridae (Parasalenia gratiosa and Mespilia globulus, respectively) has been reported at Enewetak and elsewhere in the Marshall Islands (A. H. Clark, 1952). These species are widespread in the Indo-West Pacific. Toxopneustidae The Toxopneustidae has three Enewetak species; Crytechinus verruculatus and Tripneustes gratilla have been previously reported from Rongelap, Bikini (A. H. Clark, 1952), and Enewetak (Highsmith, 1981); Tripneustes gra- tilla is recorded only from Enewetak by A. H. Clark (1952). Toxopneustes pileolus is newly recorded from the atoll and the Marshall Islands. Schizasteridae Among the irregular echinoid families, there is evi- dence that at least one, possibly two, members of the Schizasteridae occur in the lagoon substrata (V. Fry, per- sonal communication). Fibulariidae The Fibulariidae were known at Enewetak by two species each of Echinocyamus and Fibularia reported by A. H. Clark (1952). A third species of Echinocyamus, E. crispus, is a new record. This may be a senior synonym for E. elongatus, a species recorded from Bikini by A. H. Clark (1952). Spatangidae The only spatangid known from Enewetak is Maretia planulata (as M. ovata) from Enewetak and Bikini by A. H. Clark (1952). This is one of the most common echinoids in certain areas of the Enewetak Lagoon (V. Fry, personal communication). Brissidae Three genera and four species of Brissidae, Brissus, Metalia, and Rhynobrissus are found at Enewetak. Three of the species have been reported previously from the atoll. A. H. Clark (1952, 1954) recorded Brissus latecarinatus from Enewetak and from elsewhere in the Marshall Islands. Rhynobrissus hemiasteroides is recorded by Pomeroy and Kuenzler (1967), and Metalia spatagus is newly recorded from Enewetak and the Marshall Islands. Clypeasteridae Clypeaster reticulatus, the only representative of the Clypeasteridae known from the Marshall Islands: (A. H. Clark, 1952, Bikini; 1954, Taka and Ujae) is newly recorded from Enewetak. Laganidae The family Laganidae is represented by Laganum depressum at Enewetak and other Marshall Islands (A. H. Clark, 1952, 1954). COLLECTION RECORDS Ophiomyxidae Neoplax crassipes Enewetak Islet: July 16, 1960, shallow coral head, J. H. 284 DEVANEY Roberts, orig. det. F. C. Ziesenhenne, verified by D. M. Devaney (AHF). Ophiomyxa australis Enewetak Islet: August 1957, 7 to 8 m, Richard Nishioka, orig. det. F. C. Ziesenhenne, verified by D. M. Devaney (MPRL). Gorgonocephalidae Astroboa nuda Kidrenen Islet: May 26, 1976, ocean side, depth of 13 m in a cave on the seaward side just north of Kidrenen Islet, det. D. M. Devaney (MPRL). Amphiuridae Amphipholis squamata Lagoon: July 1960, shallow, J. H. Roberts (Stn. 252-B), id. F. C. Ziesenhenne; 1 dry (AHF). Enewetak Islet: July 25, 1957, “found abundantly” in the sludge area of the sewage plant discharge line; the only area where silt and sediment were found on the reef; coll. and id. F. C. Ziesenhenne; 80 alc. (30 MPRL; 55 AHF). Runit Islet: July 8, 1959, intertidal, 0.8 ft tide, coll. J. S. Garth and F. C. Ziesenhenne, id. F. C. Ziesenhenne; 1 alc. (AHF). Japtan Islet: July 19, 1957, seaward reef; coll. and id. F. C. Ziesenhenne; 20 to 30 alc. (MPRL). Engebi Islet: July 21, 1959, intertidal, 0.5 ft tide, coll. J. S. Garth and F. C. Ziesenhenne, id. F. C. Ziesenhenne; 4 alc. (AHF). Igurin Islet: July 11, 1959, intertidal, 1.1 ft tide, coll. by J. S. Garth and F. C. Ziesenhenne, id. F. C. Ziesenhenne, 1 alc. (AHF). Enewetak Islet: Aug. 9, 1965, under coral, rock, quarry, D. R. Stokes, det. D. M. Devaney, 1 spec. (BPBM). Amphiodia sp. Lagoon: Sept. 24, 1980, airlift sample of lagoon bot- tom, 35 ft off Enewetak Islet, coll. and id. D. M. Deva- ney, 1 dry (BPBM). Ophiotrichidae Macrophiothrix longipeda Araanbiru Islet: July 22, 1959, id. F. C. Ziesenhenne (MPRL). Engebi Islet: July 21, 1959; id. F. C. Ziesenhenne; 2 spec. alc. (MPRL). Igurin Islet: September 1956, outer reef, 2 to 3 ft depth, coll. D. Watson, id. F. C. Ziesenhenne, 1 spec. (MPRL). daptan Islet: July 18, 1959, id. F. C. Ziesenhenne; 4 spec. (MPRL). (Note, at least 10 additional lots at AHF [not examined by D.M. Devaney, det. by F. C. Ziesenhenne)). Macrophiothrix rhabdota Enewetak Islet: Feb. 24, 1957, coll. A. H. Banner, id. F. C. Ziesenhenne, 1 spec. dry (MPRL). Enewetak Islet: Feb. 27, 1957, lagoon side about % of length of south- ern tip of islet, sandy bottom ca. 2 m deep, heads of Pocillopora and branching Acropora, A. H. Banner (Stn. B. E. 11), id. F. C. Ziesenhenne, verified (insofar as pos- sible) by D. M. Devaney, 1 spec. alc. (AHF). Ophiothela danae Enewetak Islet: May 21, 1975, depth 8 ft, lagoon, cling- ing to soft nephtheid coral colony that was attached to chunks of dead coral buried in sandy bottom, coll. L. Ciereszko, det. D. M. Devaney, 17 spec. (6 MPRL, 11 BPBM). Biken Islet: Aug. 1, 1975, ocean side on reddish orange gorgonian coral, 30 m coll. J. Lamber- son, det. D. M. Devaney, 10+ spec. dry (BPBM). Ophiothrix (Acanthophiothrix) proteus Enewetak Islet: Dec. 13, 1976, lagoon pinnacle, north end of islet, depth 60 to 70 ft, extracted from live coral, Porites andrewsi, coll. and det. D. M. Devaney, 1 spec. dry (BPBM). Ophiothrix (Acanthophiothrix) purpurea Deep Pass: July 1963, between Medren and Japtan, depth 18 m, coll. R. Grigg, det. D. M. Devaney, 1 spec. alc. (BPBM); Rigili and Giriinien Islets: Feb. 5, 1973, entwined in branches of gorgonian coral, 30 m depth, coll. and det. D. M. Devaney, 1 spec. (BPBM). Ophiothrix (Ophiothrix) trilineata Enewetak Islet: June 26, 1965, in coral Pocillopora sp., depth 1.6 m, coll. D. Stokes, det. D. M. Devaney, 1 spec. Japtan Islet: July 22, 1965, coll. D. Stokes, det. D. M. Devaney, 1 spec. Medren Islet: Aug. 8, 1965, quarry, from Pocillopora sp., coll. D. Stokes, det. D. M. Devaney, 2 spec. Ophiocomidae Ophianthrum elegans Enewetak Islet: Sept. 3, 1965, lagoon, just off shore under rocks, 1.7 m depth, coll. D. R. Stokes, 1 spec., alc. (BPBM). Sand Islet: Sept. 7, 1966, off northwest end of islet, 1 m depth, coll. M. Youngbluth, det. D. M. Devaney, 2 spec., alc. (BPBM). Lagoon: Sept. 13, 1980, pinnacle across from Medren Islet under coral slab on side of pinnacle with rubble beneath, 8 m depth, coll. by C. Arneson, det. D. M. Devaney, 1 spec. (BPBM). Ophionereidae Ophionereis sp. Enewetak Islet: Sept. 3, 1965, lagoon, just off shore under rocks, 1.8 m depth, coll. D. R. Stokes, det. D. M. Devaney, 2 spec., dry (BPBM). Ophiodermatidae Ophiarachna affinis Rigili Islet: Sept. 3, 1957, 7 m depth, coll. Roberts, det. D. M. Devaney, 1 spec. alc. (MPRL). Ophiarachnella gorgonia Enewetak Islet: July 19, 1959, reef south end of islet, coll. and det. F. C. Ziesenhenne, 3 spec. (MPRL); Aug. 14, 1965, quarry, under coral rocks, 1 m depth, coll. D. R. Stokes, det. D. M. Devaney, 1 spec., dry (BPBM). Rabaion Islet: July 12, 1959, northwest end of islet, coll. and det. F. C. Ziesenhenne, 3 spec. dry (AHF). Ophioconis permixta Medren Islet: Feb. 7, 1976, lagoon outcrop, shallow 3 to 4 m, with (?o0n) crinoid Comanthus bennetti, coll. D. M. Devaney and A. Fielding, det. D. M. Devaney, 1 spec. alc. (BPBM). Ophiuridae Ophiolepis superba Enewetak Islet: July 15, 1961, in tide pools on seaward ECHINODERMATA OTHER THAN HOLOTHUROIDEA 285 reef, 0.3 to 1.0 m depth, coll. C. E. King, det. F. C. Ziesenhenne, 1 spec. alc. (AHF). Ophiolepis sp. aff. O. cinta Enewetak Islet: Aug. 14, 1965, under coral rocks, 1 m depth, S. R. Stokes, det. D. M. Devaney, 1 spec. dry (BPBM); 0.7 to 1.0 m depth, coll. J. H. Roberts, Jr., det. D. M. Devaney, 1 spec. dry (AHF). Lagoon: from coral head, coll. and det. D. M. Devaney, 1 spec. (AHF). Ophiura kinbergi Engebi Islet: July 21, 1959, intertidally, sand under boulder, coll. and det. F. C. Ziesenhenne; 1 juv. spec., alc., (MPRL). Biliiri Islet: lagoon, July 24, 1959, dredged at 3.5 to 5.2 depth, coll. and det. F. C. Ziesenhenne; 4 spec., dry (AHF). REFERENCES Allen, G. R., 1972, Observations on a Commensal Relationship Between Siphamia fuscolineata (Apogonidae) and the Crown- of-Thorns Starfish, Acanthaster planci, Copeia, 1972: 595. Bartolini, G. L., T. R. Erdman, and P. J. Scheuer, 1973, Anthra- quinone Pigments from the Crinoid Comanthus bennetti, Tetrahedron, 29: 3699-3702. Bruce, A. J., 1979, A Report on a Small Collection of Pon- toniine Shrimps from Eniwetok Atoll, Crustaceana, Suppl., 5, pp. 209-230. Chartock, M. A., 1972, The Role of Detritus in a Tropical Marine Ecosystem: Niche Separation in Congeneric Ophiuroids, Food Partitioning in Cryptic Invertebrates and Herbivore Detritus Production at Eniwetok, Marshall Islands, Ph.D. dissertation, University of Southern California, Los Angeles, California. Clark, A. H., 1952, Echinoderms from the Marshall Islands, Proc. U. S. Nat. Mus., 102 (3303): 265-303. —_, 1954, Records of Indo-Pacific Echinoderms, Pac. Sci., 8: 243-263. Clark, A. M., and F. W. E. Rowe, 1971, Monograph of Shallow- Water Indo-West Pacific Echinoderms, Brit. Mus. (Nat. Hist.), London. Devaney, D. M., 1970, Studies on Ophiocomid Brittlestars, I, A New Genus (Clarkcoma) of Ophiocominae with a Re- evaluation of the Genus Ophiocoma, Smithson. Contrib. Zool. No. 51, pp. 1-44. __, 1974, Shallow-water Asterozoans of Southeastern Polynesia II, Ophiuroidea, Micronesica, 10 (1): 105-204. __., 1978, A Review of the Genus Ophiomastix (Ophiuroidea: Ophiocomidea), Micronesica, 14 (2): 273-359. Eberth, T. A., 1982, Longevity, Life History, and Body Wall Size in Sea Urchins, Ecol. Monogr., 52 (4): 353-394. Gislen, T., 1940, A Collection of Crinoids from the South Sea Islands, K. svenska Vetensk Akad. Handl. Series 3, 18 (10): 3-16. Highsmith, R. C., 1980, Burrowing by the Bivalve Mollusc Lithophaga curta in the Living Reef Coral Montipora berryi and a Hypothesis of Reciprocal Larval Recruitment, Mar. Biol. (Berlin), 56: 155-162. __, 1981, Coral Bioerosions at Enewetak: Agents and Dynamics, Int. Rev. Gesamten. Hydrobiol., 66: 335-375. Humes, A. G., 1970, Clavisodalis heterocentroti gen. et sp. n., a Cyclopoid Copepod Parasitic on an Echinoid at Eniwetok Atoll, Jour. Parasit., 56: 575-583. , 1971, Cyclopoid Copepods (Stellicomitidae) Parasitic on Sea Stars from Madagascar and Eniwetok Atoll, J. Parasitol., 57: 1330-1343. _—_, 1972, Pseudanthessius comanthi n. sp. (Copepoda, Cyclo- poida) Associated with a Crinoid at Eniwetok Atoll, Pac. Sci., 26: 373-380. __., 1977, Pseudanthessid Copepods (Cyclopoida) Associated with Crinoids and Echinoids (Echinodermata) in the Tropical Western Pacific Ocean, Smithson. Contrib. Zool. No. 243, pp. 1-43. King, C. E., 1964, Relative Abundance of Species and MacArthur’s Model, Ecology, 45: 716-727. Lawrence, J. M., 1970, The Effects of Starvation on the Lipid and Carbohydrate Levels of the Tropical Sea Urchin Echinometra mathaei (de Blainville), Pac. Sci., 24{4): 487-489. __., and C. J. Davies, 1969, Algal Growth over the Epidermis of Sea Urchin Spines, J. Phycol., 5 (3): 269. , and A. Guille, 1982, Organic Composition of Tropical, Polar and Temperate-Water Echinoderms, Comp. Biochem. Physiol., 72B (2): 283-287. Marsh, L. M., 1977, Coral Reef Asteroids of Palau, Caroline Islands, Micronesica, 13 (2): 251-281. Meyer D. L., and D. B. Macurda, Jr., 1980, Ecology and Distribu- tion of the Shallow-Water Crinoids of Palau and Guam, Micronesica, 16 (1): 59-99. Pomeroy, L. R., and E. J. Kuenzler, 1967, Phosphorus Turnover by Coral Reef Animals, Symposium on Radioecology, D. J. Nelson, and F. C. Evans (Eds.), Ann Arbor, Michigan, pp. 474-482. Russo, A. R., 1980, Bioerosion by Two Rock Boring Echinoids (Echinometra mathaei and Echinostrephus aciculatus) on Enewetak Atoll, Marshall Islands, J. Mar. Res., 38: 99-110. Sloan, N. A., A. M. Clark, and J. D. Taylor, 1979, The Echino- derms of Aldabra and Their Habitats, Bull. Br. Mus. (Nat. Hist.) Zool., 37: 81-128. Zmarzly, D. L., 1984, Distribution and Ecology of Shallow-Water Crinoids at Enewetak Atoll, Marshall Islands, with an Anno- tated Checklist of Their Symbionts, Pac. Sci., 38: 105-122. Chapter 26 Protochordates of Enewetak Atoll LUCIUS G. ELDREDGE University of Guam, The Marine Laboratory, UOG Station, Mangilao, Guam 96913 INTRODUCTION Protochordates are members of two _ acraniate subphyla—Tunicata and Cephalochordata. Tunicates include the sessile ascidians and the pelagic larvaceans and thaliaceans. Few of either subphyla have been recorded from Enewetak. TUNICATA Even though ascidians are known to be abundant and diverse at Enewetak, only 10 species have been reported in the literature (Eldredge, 1967). These names are pro- vided in the checklist (Table 1). All of the forms known to occur at the atoll belong to the family Didemnidae. Of these reported in 1967, two name changes were executed by Kott (1980). The original specimens of Diplosoma virens should now be named Diplosoma similis, and the unidentified Trididemnum sp. should be named T. clinides. Enewetak is the type locality and the only known location of Didemnum gintonicum from a coral knoll at Kinedrol (Chinieero) and Diplosoma hiatti from the reef between Japtan and Jinimi (Chinimi). The remaining species have widespread distributions. Diplosoma macdonaldi is found in the tropical and subtropi- cal waters of the Atlantic and Pacific, and Diplosoma (Lis- soclinum) fragile is known from Bermuda and the West Indies as well as from Japan, Malaysia, and other Pacific islands. These two species and Didemnum moseleyi and Diplosoma similis are found in Hawaiian waters. Didemnum moseleyi is also known from Japan, Philippine Islands, Malaysia, Australia, Tasmania, New Caledonia, south Ara- bia, and from several other Pacific island locations. The TABLE 1 Checklist of Ascidians and Cephalochordates of Enewetak Atoll Phylum CHORDATA Subphylum TUNICATA Class ACIDIACEA Order ENTEROGONA Suborder APLOUSOBRANCHIATA Family DIDEMNIDAE Didemnum gintonicum Eldredge: Eldredge, 1967. Didemnum grande (Herdman): Eldredge, 1967. Didemnum moseleyi (Herdman): Eldredge, 1967. Didemnum quincuciale Michaelson: Eldredge, 1967. Diplosoma similis (Sluiter): Kott, 1980. Diplosoma (Diplosoma) virens Hartmeyer: Eldredge, 1967. Diplosoma (Diplosoma) hiatti Eldredge: Eldredge, 1967. Diplosoma (Diplosoma) macdonaldi Herdman: Eldredge, 1967. Diplosoma (Lissoclinum) fragile (Van Name): Eldredge, 1967. Trididemnum clinides Kott: Kott, 1980. Trididemnum sp.: Eldredge, 1967. Trididemnum cyclops Michaelson: Eldredge, 1967. Subphylum CEPHALOCHORDATA Family ASYMMETRONTIDAE Notasymmetron caudatum (Willey): Schultz (1953). 288 ELDREDGE known distribution of Diplosoma similis (as separated from Diplosoma virens) includes Indonesia, the Great Barrier Reef, Fiji, Japan, and questionably the Philippine Islands. Didemnum grande, a widespread species, is found in Japan, Philippine Islands, Malaysia, Australia, Tasmania, south Arabia, and Palmyra in the Line Islands. Didemnum quincunciale has been known previously only from Zanzi- bar and Trididemnum cyclops from Australia and Madagas- car. Tokioka (1967) studied the ascidians in the U. S. National Museum and recorded four species from other atolls in the Marshall Islands. These are: Diplosoma virens Hartmeyer, 1909 (Leptoclinum virens)—Ebon Ascidia aperta Sluiter, 1904—Rongelap Polycarpa cryptocarpa (Sluiter, 1885)—Bikini Polycarpa iwayamae Tokioka, 1950—Arno No specimens were found from Enewetak. Didemnids show an increase in diversity toward the Indo-Malayan region. The didemnid fauna of Enewetak contains a number of widespread species, showing its strong faunistic affinity to the Indo-West Pacific. Seven species of larvaceans (Gerber, 1981) and two species of thaliaceans (salps) (Gilmartin, 1958) have been reported from plankton samples taken at Enewetak. These species are discussed in Chapter 20 in this volume. CEPHALOCHORDATA Cephalochordata include the lancelets commonly known as Amphioxus. A small cephalochordate specimen dredged from the lagoon at Bikini was identified provision- ally as Notasymmetron caudatum (Willey) by Schultz (1953). At least one unidentified specimen was seen in the Mid-Pacific Marine Laboratory collections. REFERENCES Eldredge, L. G., 1967, A Taxonomic Review of Indo-Pacific Didemnid Ascidians and Descriptions of Twenty-Three Central Pacific Species, Micronesica, 2: 161-261. Gerber, R. P., 1981, Species Composition and Abundance of Lagoon Zooplankton at Eniwetok Atoll, Marshall Islands, Atoll Res. Bull., 247: 1-22. Gilmartin, M., 1958, Some Observations on the Lagoon Plankton of Eniwetok Atoll, Pac. Sci., 12: 313-316. Kott, P., 1980, Algal-Bearing Didemnid Ascidians in the Indo- West Pacific, Mem. Queensland Mus., 20: 1-47. Schultz, L. P., 1953, Fishes of the Marshall and Marianas Islands, Bull. U. S. Nat. Mus., 202: 1-685. Tokioka, T., 1967, Pacific Tunicata of the United States National Museum, Bull. U. S. Nat. Mus., 251: 1-247. Chapter 27 Annotated Checklist of the Fishes of Enewetak Atoll and Other Marshall Islands JOHN E. RANDALL and HELEN A. RANDALL Bernice P. Bishop Museum Honolulu, Hawaii 96817 INTRODUCTION Intensive fish collecting was carried out in the Marshall Islands in 1946 and 1947 in connection with and following the atom bomb tests of Operation Crossroads. These col- lections were deposited in the National Museum of Natural History of the Smithsonian Institution in Washington, D. C. Additional Marshall Islands fishes were taken in 1948 and 1949 during expeditions of the Applied Fisheries Laboratory of the University of Washington. All these fishes, plus collections from the Mariana Islands, were studied by Leonard P. Schultz, then Curator of Fishes of the National Museum of Natural History, and his collaborators, Wilbert M. Chapman, Earl S. Herald, Ernest A. Lachner, Arthur D. Welander, and Loren P. Woods. The result was the valuable three-volume work on the fishes of the Marshall and Mariana Islands (1953 to 1966). A total of 543 species of fishes was reported from the Marshall Islands in these three volumes. Unfortunately, a study of the family with the largest number of species, the Gobiidae, was not completed. Very few fishes were reported from the Marshall Islands prior to 1953. Giinther (1873 to 1910) listed 31 species from the atolls of Ebon, Jaluit, and Majuro [though his Anguilla mauritiana and Monacanthus ayraudi are prob- able locality errors, and his Echeneis scutata (Remora aus- tralis) is suspect]. Kendall and Goldsborough (1911) recorded 58 species. In a study of poisonous fishes of the Marshalls, Hiyama (1943) illustrated 71 species in color. The text (translated from the Japanese by Van Campen, 1950) deals more with the toxicity of these fishes than their classification, but the high quality of most of the fig- ures enables one to make positive identifications to species. Donald W. Strasburg (1953) submitted a report to the 289 Office of Naval Research entitled Fishes of the Southern Marshall Islands. He treated 345 species in this report, largely from collections made at Arno. He worked closely with Schultz et al. and had access to their manuscripts on Marshall Islands fishes. He listed 26 species of fishes that were not included in the manuscripts of Schultz et al. at that time (later a few of these fishes were reported in Vols. 2 and 3 of Fishes of the Marshall and Marianas Islands). Ichthyological activity in the Marshall Islands continued in the years following the research of Schultz et al., largely as a result of the establishment of the Mid-Pacific Research Laboratory on Enewetak (initially named Eniwetok Marine Biological Laboratory). Also of importance has been the presence of a number of scuba divers at Kwajalein with an interest in fishes. Some of these are expert underwater photographers, and their efforts to capture the rare or unusual species on film have provided new records of fishes from the archipelago. Randall (1986) reviewed all the new records and new species of fishes of the Marshall Islands that have been documented since Schultz et al., including those of Strasburg’s Office of Naval Research report (since few cop- ies were distributed). A total of 86 such fishes was found in the literature, and another 106 new records were added by Randall. Also there are about 45 species of fishes from the Marshalls which are still undescribed. In addition to new records and new species, many changes in the names of fishes as listed by Schultz et al. have been made in recent publications. The present paper provides a checklist (Table 1) of the reef, shore, and epipelagic fishes known from the Marshall Islands. We record a total of 817 species in 338 genera and 92 families. Deep-water fishes have been omitted from 290 RANDALL AND RANDALL TABLE 1 Checklist of the Fishes of the Marshall Islands Phylum CHORDATA Class CHONDRICHTHYS Subclass ELASMOBRANCHII Order HEXANCHIFORMES Family HEXANCHIDAE Hexanchus griseus (Bonnaterre, 1788): Randall, 1986. Order LAMNIFORMES Family GINGLYMOSTOMATIDAE Nebrius concolor Riippell, 1837 Ginglymostoma ferrugineum Schultz, 1953. Nebrius ferrugineus Randall, 1980a. Family ISURIDAE Carcharodon carcharias (Linnaeus, 1758): Schultz, 1953. Family CARCHARHINIDAE Carcharhinus albimarginatus (Riippell, 1837): Schultz, 1953. Carcharhinus amblyrhynchos (Bleeker, 1856). Carcharhinus menisorrah Schultz, 1953. Garrick (1982) concluded that the true C. menisorrah (Valenciennes in Miiller and Henle) is a junior synonym of C. falciformis (Bibron in Miiller and Henle). Carcharhinus falciformis (Bibron, 1841): Garrick, 1982. Carcharhinus galapagensis (Snodgrass and Heller, 1905): Randall, 1980a. Carcharhinus limbatus (Valenciennes, 1841): Randall, 1980a. Carcharhinus longimanus (Poey, 1861). Carcharhinus brachyurus Schultz, 1953. Garrick (1982) stated that the specimens Schultz identified as C. brachyurus are C. longimanus. Garrick pointed out that C. maou (Lesson) is an older name for longimanus, but he gave reasons for retaining the long-established longimanus. Carcharhinus melanopterus (Quoy and Gaimard, 1824): Schultz, 1953. Galeocerdo cuvier (Peron and Lesueur, 1822): Schultz, 1953. Negaprion acutidens (Riippell, 1837). Hemigaleops fosteri Schultz and Welander, 1953. Garrick and Schultz in Gilbert (1963) referred Hemigaleops fosteri to the synonymy of Negaprion acutidens. Triaenodon obesus (Riippell, 1837). Schultz (1953) classified this shark in the Triakidae; recent authors place it in Carcharhinidae; however, A. Baranes and G. Dingerkus (MS) plan to reclassify it in the Hemigaleidae. Order RAJIFORMES Family DASYATIDIDAE Dasyatis sp. Sighted by the senior author at Enewetak. Taeniura melanospilos Bleeker, 1853: Randall, 1980a. Taeniura brocki Schultz, 1953. Urogymnus sp. A specimen collected in the Enewetak Lagoon is under study by the senior author. Family MYLIOBATIDAE Aetobatus narinari (Euphrasen, 1790): Randall, 1986 (after Strasburg). Family MOBULIDAE Manta alfredi (Krefft, 1868): Schultz, 1953. Class OSTEICHTHYES Subclass ACTINOPTERYGII Order ELOPIFORMES Family ALBULIDAE Albula glossodonta (Forsskal, 1775): Randall, 1986. Shaklee and Tamaru (1981) have shown that there are two species of Albula in the Hawaiian Islands (and elsewhere in the Indo-Pacific region), neither of which is A. vulpes (Linnaeus), now known to be restricted to the Atlantic. Order ANGUILLIFORMES Family MORINGUIDAE Moringua ferruginea Bliss, 1883. Moringua macrocephala Schultz, 1953. Moringua macrochir Schultz, 1953. Gosline and Strasburg (1956) showed the remarkable differences of immature Hawaiian Moringua from adults and between the sexes (including vertebral counts), as have Castle and Bohlke (1976) for the western Atlantic Moringua edwardsi (Jordan and Bollman). Gosline and Strasburg identified their Hawaiian material as M. macrochir Bleeker. Castle (1968a) believes the Hawaiian species (and thus presumably the specimens from the Marshall Islands identified by Schultz as macrochir) is M. ferruginea. Also, Castle regards M. macrocephala (Bleeker) as the immature form of macrochir. (This table continued on next page.) ANNOTATED CHECKLIST OF FISHES 291 TABLE 1 (cont’d) Family MORINGUIDAE (cont'd) Moringua javanica (Kaup, 1856): Schultz, 1953. Moringua bicolor Schultz, 1953. Castle (1968a) stated that M. bicolor Kaup is a possible mature male of javanica. Moringua linearis Gray, 1831: Schultz, 1953. Schultz’s identification is questionable. Moringua microchir Bleeker, 1853. Moringua abbreviata Schultz, 1953. Castle (1968a) is followed in regarding M. abbreviata (Bleeker) as a probable synonym of microchir. Family XENOCONGRIDAE Kaupichthys atronasus Schultz, 1953. Kaupichthys brachychirus Schultz, 1953. Kaupichthys hyoproroides (Stromann, 1896). Kaupichthys diodontus Schultz, 1953. Bohlke and Smith (1968) showed that K. diodontus Schultz (1943) and K. atlanticus Bohlke are junior synonyms of K. hyoproroides, named from a leptocephalus larva. Family MURAENIDAE Anarchias allardicei Jordan and Starks, 1906: Schultz, 1953. Anarchias cantonensis (Schultz, 1943): Schultz, 1953. Anarchias seychellensis Smith, 1962. Anarchias leucurus Schultz, 1953. Randall and McCosker (1975) reidentified Marshall Islands A. leucurus as seychellensis. Echidna leucotaenia Schultz, 1943: Schultz, 1953. Echidna nebulosa (Ahl, 1789): Schultz, 1953. Echidna polyzona (Richardson, 1844): Schultz, 1953. Echidna unicolor Schultz, 1953. Enchelycore bikiniensis (Schultz, 1953). Gymnothorax bikiniensis Schultz, 1953. Enchelycore bayeri (Schultz, 1953). Gymnothorax bayeri Schultz, 1953. Enchelynassa canina (Quoy and Gaimard, 1824): Schultz, 1953. Gymnomuraena zebra (Shaw, 1797). Echidna zebra Schultz, 1953. Gymnothorax buroensis (Bleeker, 1857): Schultz, 1953. Gymnothorax enigmaticus McCosker and Randall, 1982. Gymnothorax rupelli Schultz, 1953—an unjustified emendation of G. rueppelliae (McClelland). The latter is a valid species often misidentified as G. petelli (Bleeker), a junior synonym. Gymnothorax fimbriatus (Bennett, 1831): Schultz, 1953. Gymnothorax favagineus var. isingteenus Hiyama, 1943. Gymnothorax flavimarginatus (Riippell, 1828): Schultz, 1953. Gymnothorax fuscomaculatus (Schultz, 1953). McCosker and Rosenblatt (1975) have determined that the genus Rabula (in which Schultz described both fuscomaculata and marshallensis) is based on an aberrant specimen of the temperate eastern Pacific Gymnothorax mordax (Ayres). Gymnothorax gracilicaudus Jenkins, 1903: Schultz, 1953. Gymnothorax javanicus (Bleeker, 1859): Schultz, 1953. Gymnothorax margaritophorus Bleeker, 1864: Schultz, 1953 Gymnothorax marshallensis (Schultz, 1953). Gymnothorax melatremus Schultz, 1953. Gymnothorax meleagris (Shaw and Nodder, 1795): Schultz, 1953. Gymnothorax monochrous Bleeker, 1864: Schultz, 1953. Gymnothorax monostigma (Regan, 1909): Schultz, 1953. Gymnothorax pindae Smith, 1962. Gymnothorax moluccensis Schultz, 1953. Randall and McCosker (1975) determined that Schultz’s material is G. pindae, not G. moluccensis (Bleeker). Gymnothorax rueppelliae (McClelland, 1845). Gymnothorax petelli Schultz, 1953. Randall (1973) and McCosker and Rosenblatt (1975) have shown that G. petelli of Schultz and other authors is the true G. rueppelliae. Gymnothorax undulatus (Lacepede, 1803): Schultz, 1953. Gymnothorax zonipectis Seale, 1906: Schultz, 1953. Rhinomuraena quaesita Garman, 1888: Giinther, 1910; Schultz, 1953. Siderea picta (Ahl, 1789). Gymnothorax pictus Schultz, 1953. Siderea prosopeion (Bleeker, 1853). Gymnothorax thrysoideus (non-Richardson) Hiyama, 1943; Schultz, 1953. Giniednblercontneedioninestipace) 292 RANDALL AND RANDALL TABLE 1 (cont’d) Family MURAENIDAE (cont’d) Uropterygius concolor Riippell, 1837: Schultz, 1953. Schultz’s identification should be confirmed. Uropterygius fuscoguttatus Schultz, 1953. Uropterygius kamar McCosker and Randall, 1977: Randall, 1986. Uropterygius macrocephalus (Bleeker, 1865). Uropterygius reidi Schultz, 1953. Gosline (1958) placed U. reidei Schultz (1943) in the synonymy of U. knighti (Jordan and Starks). Uropterygius knighti Schultz, 1953. McCosker et al. (1984) have referred U. knighti and U. necturus (Jordan and Gilbert) to the synonymy of U. macrocephalus, now known to be a trans-Pacific species. Uropterygius marmoratus (Lacepede, 1803): Schultz, 1953. Uropterygius supraforatus (Regan, 1909): Schultz, 1953. Uropterygius dentatus Schultz, 1953, from Johnston Island, was synonymized by Gosline (1958). Uropterygius xanthopterus Bleeker, 1859: Schultz, 1953. Family CONGRIDAE Ariosoma scheelei (Stromman, 1896). Ariosoma obud Schultz, 1953. P. H. J. Castle (personal communication) wrote us, “it seems highly likely” that A. obud Herre is a junior synonym of A. scheelei, a species with low vertebral counts. Three Bishop Museum specimens from Enewetak have 116 to 120 vertebrae (low for the genus—see Castle, 1968b). Conger cinereus cinereus Riippell, 1828. Conger noordziekii Schultz, 1953. Kanazawa (1958) placed C. noordziekii Bleeker in the synonymy of cinereus. Gorgasia sp. An apparent undescribed species; 19 specimens from Enewetak, with 181 to 189 vertebrae. Heteroconger hassi (Klausewitz and Eibl-Eibesfeldt, 1959): Bohlke and Randall, 1981; Randall, 1986. Taenioconger hassi Matsuura, 1984. Poeciloconger fasciatus Giinther, 1871: Randall, 1986. Family OPHICHTHIDAE Apterichtus klazingai (Weber, 1913): Randall, 1986. McCosker (1977) has shown that this generic name replaces Verma. Also some species of Apterichtus have been mistakenly described in Sphagebranchus and Caecula. Brachysomophis sauropsis Schultz, 1953. Callechelys marmoratus (Bleeker, 1853): Schultz, 1953. Callechelys melanotaenia Bleeker, 1864: Schultz, 1953. Leptenchelys pinnaceps Schultz, 1953. McCosker (1970) stated that this taxon is a junior synonym of C. melanotaenia. Cirricaecula johnsoni Schultz, 1953. Leiuranus semicinctus (Lay and Bennett, 1839): Schultz, 1953. Muraenichthys gymnotus Bleeker, 1864: Schultz, 1953. Muraenichthys laticaudata (Ogilby, 1897): Schultz, 1953. Muraenichthys macropterus Bleeker, 1857: Schultz, 1953. Muraenichthys schultzei Bleeker, 1857: Schultz, 1953. Muraenichthys sibogae Weber and de Beaufort, 1916: Schultz, 1953. Myrichthys bleekeri Gosline, 1951. Myrichthys semicinctus Schultz, 1953. Gosline (1951) provided a new name for Ophisurus fasciatus var. semicinctus Bleeker, preoccupied by O. semicinctus Lay and Bennett. Myrichthys colubrinus (Boddaert, 1781): Schultz, 1953. Myrichthys maculosus (Cuvier, 1817): Schultz, 1953. Myrophis uropterus (Temminck and Schlegel, 1842): Schultz, 1953. Ophichthus sp. Under study by John E. McCosker. Phyllophichthus xenodontus Gosline, 1951: Randall, 1986 (after Strasburg). Schismorhynchus labialis (Seale, 1917). Leptenchelys labialis Schultz, 1953. McCosker (1970) described a new genus, Schismorhynchus, for labialis. Schultzidia johnstonensis Schultz and Woods, 1949: Schultz, 1953. Schultzidia retropinnis (Fowler, 1933): Schultz, 1953. Order CLUPEIFORMES Family CLUPEIDAE Dussumieria hasselti Bleeker, 1850: Schultz and Welander, 1953. Herklotsichthys quadrimaculatus (Riippell, 1837): Randall, 1986 [after Strasburg, who identified this clupeid from Arno as Harengula kunzei (Bleeker)). Spratelloides atrofasciatus Schultz, 1953. Spratelloides delicatulus (Bennett, 1831): Schultz and Welander, 1953. Order SALMONIFORMES Family SYNODONTIDAE Saurida gracilis (Quoy and Gaimard, 1824): Schultz, 1953. Soned ie pinotats: Schultz, 1253. (This table continued on next page.) ANNOTATED CHECKLIST OF FISHES TABLE 1 (cont'd) 293 Family SYNODONTIDAE (cont'd) Synodus englemani Schultz, 1953. Cressey (1981) revised Synodus; he recognized this species and S. binotatus as valid. Synodus variegatus (Lacepede, 1803): Schultz, 1953. Synodus sp. Order GONORYNCHIFORMES Family CHANIDAE Chanos chanos (Forsskal, 1775): Randall, 1986. Order GOBIESOCIFORMES Family GOBIESOCIDAE Liobranchia stria Briggs, 1955: Schultz, 1966. Order LOPHIIFORMES Family ANTENNARIIDAE Antennarius analis (Gosline, 1957): Randall, 1986. Antennarius coccineus (Lesson, 1831): Schultz, 1966. Antennarius pictus (Shaw and Nodder, 1794): Randall, 1986. Theodore W. Pietsch (personal communication) informed us that A. pictus is a senior synonym of A. chironectes Lacepede (the name used by most recent authors for this species). Antennarius randalli Allen, 1970: Randall, 1986. Antennarius rosaceus (Smith and Radcliffe, 1912). Trichophryne rosaceus Schultz, 1966. Antennatus tuberosus (Cuvier, 1817): Randall, 1986. Antennarius bigibbus Fowler, 1928. Order GADIFORMES Family BREGMACEROTIDAE Bregmaceros nectabanus Whitley, 1941. Bregmaceros mcclellandi Schultz, 1953. Marshall Islands material (including a Bishop Museum specimen from the Enewetak Lagoon) was reidentified as B. nectabanus, following D’Ancona and Cavinato (1965). Order OPHIDIIFORMES Family OPHIDIIDAE Brotula townsendi Fowler, 1900: Schultz, 1960. Family BYTHITIDAE Brosmophyciops pautzkei Schultz, 1960. Cohen and Nielson (1978) are followed in considering Bythitidae a family for this and related fishes. Dinematichthys iluocoeteoides Bleeker, 1855: Schultz, 1960. A badly needed revision of the genus Dinematichthys and allies has just been commenced by Allegra N. Sedor of the University of Southern California. More than one species of the genus occurs in the Marshall Islands; it is not clear that the name iluocoeteoides can be confidently applied to any of them. Family CARAPIDAE Carapus homei (Richardson, 1846): Schultz, 1960. Carapus mourlani (Petit, 1934): Schultz, 1960. Schultz identified his material as C. mourlani “with considerable uncertainty,” though pointed out that his specimens are clearly distinct from his C. homei (which Arnold, 1956, listed as a senior synonym of mourlani). Jordanicus gracilis (Bleeker, 1856): Schultz, 1960. Order ATHERINIFORMES Family BELONIDAE Ablennes hians (Valenciennes, 1846): Schultz, 1953. Platybelone argalus platyura (Bennett, 1832). Belone platyura Schultz, 1953. Strongylura incisa (Valenciennes, 1846): Schultz, 1953. Rhaphiobelone robusta Schultz, 1953. Mees (1962) and Parin (1967) placed this taxon in the synonymy of S. incisa. Tylosurus crocodilus crocodilus (Peron and Lesueur, 1821). Family HEMIRAMPHIDAE Euleptorhamphus viridis (van Hasselt, 1824): Schultz, 1953. Hyporhamphus acutus acutus (Giinther, 1871): Collette, 1974. Hyporhamphus acutus Schultz, 1953. Hyporhamphus affinis (Giinther, 1866): Parin et al., 1980. Hyporhamphus dussumieri Schultz, 1953. Hyporhamphus dussumieri (Valenciennes, 1846): Parin et al., 1980. Hyporhamphus laticeps Schultz, 1953. Oxyporhamphus micropterus micropterus Parin et al., 1980. (This table continued on next page.) 294 RANDALL AND RANDALL TABLE 1 (cont'd) “Family EXOCOETIDAE Cypselurus antoncichi Woods and Schultz, 1953. Cypselurus spilonotopterus (Bleeker, 1866): Woods and Schultz, 1953. Cypselurus unicolor (Valenciennes, 1846): Woods and Schultz, 1953. Exocoetus volitans Linnaeus, 1758: Woods and Schultz, 1953. Parexocoetus mento mento (Valenciennes, 1864). Parexocoetus mento Woods and Schultz, 1953. Prognichthys sp. Woods and Schultz, 1953. Family ATHERINIDAE Atherinomorus lacunosus (Bloch and Schneider, 1801). Pranesus pinguis Schultz, 1953. Whitehead and Ivantsoff (1983) placed Atherina pinguis Lacepede in the synonymy of Atherinomorus lacunosus. Atherion elymus Jordan and Starks, 1901. Atherion elymus asper Schultz, 1953. Walter Ivantsoff (personal communication) does not recognize Schultz’s subspecies of A. elymus. Hypoatherina barnesi Schultz, 1953. Hypoatherina ovalaua (Herre, 1935). Allanetta ovalaua Schultz, 1953. Generic change from Whitehead and Ivantsoff, 1983. Stenatherina panatela (Jordan and Richardson, 1908). Stenatherina temminckii Schultz, 1953. Walter Ivantsoff (personal communication) has corrected this misidentification by Schultz. Family ISONIDAE Iso hawaiiensis Gosline, 1952: Randall, 1986. Order BERYCIFORMES Family ANOMALOPIDAE Photoblepharon palpebratus (Boddaert, 1781): Randall, 1986. Family HOLOCENTRIDAE Myripristis adusta Bleeker, 1853: Randall, 1986. Myripristis amaena (Castelnau, 1873). Myripristis argyromus Woods, 1953. Greenfield (1974) placed M. argyromus Jordan and Evermann in the synonymy of M. amaenus (Castelnau). Myripristis berndti Jordan and Evermann, 1903: Woods, 1953. Myripristis kuntee Cuvier, 1831. Myripristis multiradiatus Woods, 1953. Greenfield (1974) synonymized M. multiradiatus Giinther with M. kuntee. Myripristis murdjan (Forsskal, 1775): Woods, 1953. Myripristis bowditchae Woods, 1953. Randall and Guézé (1981) placed this species, M. parvidens Cuvier, and M. axillaris Valenciennes in the synonymy of M. murdjan. Myripristis pralinia Cuvier, 1829: Woods, 1953. Myripristis violacea Bleeker, 1851. Myripristis microphthalmus Woods, 1953. Greenfield (1974) synonymized M. microphthalmus Bleeker with violacea. Myripristis vittata Cuvier, 1831: Randall, 1986. Neoniphon argenteus (Valenciennes, 1831). Holocentrus laevis Woods, 1953. Shimizu and Yamakawa (1979) placed Holocentrus laeve Giinther in the synonymy of argenteus. Randall and Heemstra (1985) have shown that Neoniphon Castelnau replaces the generic name Flammeo Jordan and Evermann. Neoniphon opercularis (Valenciennes, 1831). Holocentrus opercularis Woods, 1953. Neoniphon sammara (Forsskal, 1775). Holocentrus sammara Woods, 1953. Plectrypops lima (Valenciennes, 1831). Holotrachys lima Woods, 1953. Woods and Sonoda (1973) showed that Holotrachys Giinther is a generic synonym of Plectrypops Gill. Sargocentron caudimaculatum (Riippell, 1838): Randall, 1986. Matsuura and Shimizu (1982) noted that Sargocentron Fowler is an older name than Adioryx Starks. Sargocentron diadema (Lacepede, 1802). Holocentrus diadema Woods, 1953. *Parin (1960; translation 1963) listed nine species of Cypselurus, in addition to the three included here, three of Prognichthys, and Parexocoetus brachypterus brachypterus from Oceania. Possibly some of these species will eventually be found in the Marshalls area. Ran- dall (1955a) reported Cypselurus suttoni and Prognichthys albimaculatus from the nearby Gilbert Islands. (This table continued on next page.) ANNOTATED CHECKLIST OF FISHES TABLE 1 (cont'd) Family HOLOCENTRIDAE (cont'd) Sargocentron melanospilos (Bleeker, 1858): Randall, 1986. Sargocentron microstoma (Giinther, 1859). Holocentrus microstomus Woods, 1953. Sargocentron punctatissimum (Cuvier, 1829). Holocentrus lacteoguttatus Woods, 1953. Randall and Heemstra (1985) pointed out that S. punctatissimum has priority over S. lacteoguttatum (Cuvier). Sargocentron praslin (Lacepede, 1802): Randall, 1986. Sargocentron spiniferum (Forsskal, 1775). Holocentrus spinifer Woods, 1953. Sargocentron tiere (Cuvier, 1829). Holocentrus tiere Woods, 1953. Sargocentron tiereoides (Bleeker, 1853). Holocentrus tieroides Woods, 1953. Sargocentron violaceum (Bleeker, 1853): Shimizu and Yamakawa, 1979. Order GASTEROSTEIFORMES Family AULOSTOMIDAE Aulostomus chinensis (Linnaeus, 1766): Schultz, 1953. Family FISTULARIIDAE Fistularia commersonii Riippell, 1838. Fistularia petimba Schultz, 1953. Fritzsche (1976) showed that shallow-water Indo-Pacific specimens of Fistularia that most authors have called petimba are F. commersonii. The name petimba is correct for the deep-water tropical Atlantic and Indo-West-Pacific species with small spines posteriorly on the lateral line. Family SOLENOSTOMIDAE Solenostomus paradoxus (Pallas, 1870). Solenostomus armatus Schultz, 1953. Ronald A. Fritzsche (personal communication) has reidentified Schultz’s specimen (27.5 mm SL from a plankton tow off Bikini) as a juvenile of S. paradoxus. Family SYNGNATHIDAE Choeroichthys sculptus (Giinther, 1870): Dawson (1976); Randall, 1986 (after Strasburg). Corythoichthys flavofasciatus (Riippell, 1838). Corythoichthys flavofasciatus conspicillatus Herald, 1953. Dawson (1977a) could find no substantial basis for Herald’s division of this species to subspecies. Corythoichthys intestinalis (Ramsay, 1881). Corythoichthys intestinalis waitei Herald, 1953. Corythoichthys nigripectus Herald, 1953. Corythoichthys schultzi Herald, 1953. Cosmocampus banneri (Herald and Randall, 1972). Cosmocampus maxweberi (Whitley, 1933). Syngnathus maxweberi Herald, 1953. Dawson (1980) referred S. maxweberi to the genus Cosmocampus Dawson. Doryrhamphus excisus excisus Kaup, 1956. Doryrhamphus melanopleura melanopleura Herald, 1953. Dawson (1981) placed D. melanopleura (Bleeker) in the synonymy of D. excisus Kaup. Dunckerocampus dactyliophorus (Bleeker, 1853): Herald, 1953. Halicampus brocki (Herald, 1953). Halicampus mataafae (Jordan and Seale, 1906): Herald, 1953. Ichthyocampus bikiniensis Herald, 1953. Dawson (19776) wrote that he is unable to refer this species, which Herald described from two planktonic postlarvae, to any genus, but the origin of the dorsal fin on the trunk rules out its placement in Ichthyocampus. It is probably the young of Halicampus, Phoxocampus, or Corythoichthys (C. E. Dawson, personal communication). Micrognathus brevirostris pygmaeus Fritzsche, 1981. Micrognathus brevirostris Herald, 1953. Phoxocampus diacanthus (Schultz, 1943). Ichthyocampus diacanthus Herald, 1953. Dawson (1977b) erected Phoxocampus for this species and two other pipefishes. Order SCORPAENIFORMES Family SCORPAENIDAE Dendrochirus biocellatus (Fowler, 1938): Schultz, 1966. Dendrochirus zebra (Cuvier, 1829): Randall, 1986. Parascorpaena mcadamsi (Fowler, 1938): Randall, 1986. Parascorpaena mossambica (Peters, 1855). Sebastapistes mcadamsi Schultz, 1966. 295 (This table continued on next page.) 296 RANDALL AND RANDALL TABLE 1 (cont'd) Family SCORPAENIDAE (cont’d) Pterois antennata (Bloch, 1787): Schultz, 1966. Pterois radiata Cuvier, 1829: Schultz, 1966. Pterois volitans (Linnaeus, 1758): Kendall and Goldsborough, 1911; Allen and Eschmeyer, 1973. Scorpaenodes brocki (Schultz, 1966). Hypomacrus brocki Schultz, 1966. Scorpaenodes guamensis (Quoy and Gaimard, 1824): Schultz, 1966. Scorpaenodes hirsutus Smith, 1957: Randall, 1986. Scorpaenodes kelloggi (Jenkins, 1903): Schultz, 1966. Scerpaenodes parvipinnis (Garrett, 1863): Schultz, 1966. Scorpaenopsis diabolus (Cuvier, 1829). Scorpaenopsis gibbosa Schultz, 1966. Eschmeyer and Randall (1975) stated that the true S. gibbosa appears to be restricted to the Indian Ocean. Scorpaenopsis fowleri (Pietschmann, 1934): Eschmeyer and Randall, 1975. This small species is only tentatively placed in the genus Scorpaenopsis. Scorpaenopsis sp. A new species which will be described by William N. Eschmeyer and Kaza Rama Rao. Sebastapistes cyanostigma (Bleeker, 1856). Sebastapistes albobrunnea Schultz, 1966. Eschmeyer and Rama Rao (MS) have shown that Scorpaena albobrunnea Giinther is a junior synonym of cyanostigma. Sebastapistes mauritiana (Cuvier, 1829). Scorpaena corallicola Schultz, 1966. Eschmeyer and Randall (1975) placed S. corallicola (Jenkins) in the synonymy of S. ballieui (Sauvage). Eschmeyer and Rama Rao (MS) regard Sebastapistes ballieui as a Hawaiian endemic. The closely related S. mauritiana is a wide-ranging Indo-Pacific species that does not occur in Hawaii. Sebastapistes strongia (Cuvier, 1829). Sebastapistes bynoensis Schultz, 1966. Sebastapistes bynoensis Richardson is a junior synonym of Parascorpaena picta (Cuvier) (Eschmeyer and Rama Rao, MS). The Bikini specimen identified as S. bynoensis by Schultz is strongia. Synanceia verrucosa Bloch and Schneider, 1801: Schultz, 1966. Taenianotus triacanthus Lacepede, 1802: Schultz, 1966. Family CARACANTHIDAE Caracanthus maculatus (Gray, 1831): Schultz, 1966. Caracanthus unipinna (Gray, 1831). Caracanthus unipinnus Schultz, 1966. Family APLOACTINIDAE Cocotropus sp. An undescribed species which will be named by Stuart G. Poss. Family PLAT YCEPHALIDAE Thysanophrys arenicola Schultz, 1966. Thysanophrys chiltonae Schultz, 1966. Thysanophrys malayanus (Bleeker, 1853). Thysanophrys papillolabium Schultz, 1966. Leslie W. Knapp (personal communication) regards this species as a junior synonym of malayanus. Wakiyus welanderi Schultz, 1966. Order PEGASIFORMES Family PEGASIDAE Eurypegasus draconis (Linnaeus, 1776): Randall, 1986. Order PERCIFORMES Family SERRANIDAE Anthias bartlettorum Randall and Lubbock, 1981. Anthias bicolor Randall, 1979. Anthias dispar (Herre, 1955): Randall and Lubbock, 1981. Anthias pascalus (Jordan and Tanaka, 1927): Randall and Lubbock, 1981. Anthias pleurotaenia Bleeker, 1857: Randall, 1986. Anthias randalli Lubbock and Allen, 1978. Anthias smithvanizi Randall and Lubbock, 1981. Anthias ventralis ventralis Randall, 1979. Anyperodon leucogrammicus (Valenciennes, 1828): Schultz, 1953. Belonoperca chabanaudi Fowler and Bean, 1930: Schultz, 1953. Cephalopholis argus (Bloch and Schneider, 1801): Schultz, 1953. Randall and Ben-Tuvia (1983) have shown that Cephalopholis guttatus (Bloch) is an earlier name for this species, but they have petitioned the International Commission for Zoological Nomenclature to suppress C. guttatus (Bloch) in order to retain the long-established C. argus. Cephalopholis leopardus (Lacepede, 1801): Schultz, 1953. Cephalopholis miniatus (Forsskal, 1775): Schultz, 1953. (This table continued on next pagel ANNOTATED CHECKLIST OF FISHES 297 TABLE 1 (cont'd) Family SERRANIDAE (cont’d) Cephalopholis sexmaculata (Riippell, 1830): Randall and Ben-Tuvia, 1983. Cephalopholis sonnerati (Valenciennes, 1828): Randall, 1986. Cephalopholis spiloparaea (Valenciennes, 1828). Cephalopholis analis Randall, 1986. Cephalopholis urodeta (Bloch and Schneider, 1801). Cephalopholis urodelus Schultz, 1953. Epinephelus caeruleopunctatus (Bloch, 1790): Kendall and Goldsborough, 1911; Randall, 1986. Epinephelus cyanopodus (Richardson, 1846). Serranus flavocaeruleus Hiyama, 1943. Many authors have mistakenly regarded E. flavocaeruleus (Lacepede) as a synonym of E. hoedtii (cyanopodus). Epinephelus kohleri Schultz, 1953. Katayama (1960) and Randall (1980a) placed this species in the synonymy of E. hoedtii. Epinephelus cyanopodus is a still older name for the species (Randall and Whitehead, 1985). Epinephelus fasciatus (Forsskal, 1775). Epinephelus emoryi Schultz, 1953. Katayama (1960) correctly referred emoryi to the synonymy of fasciatus. Epinephelus fuscoguttatus (Forsskal, 1775). Schultz (1953) confused two species under the heading E. fuscoguttatus. His illustration (Plate 26 A) is E. microdon. Randall (1964a) distinguished these two related species. Epinephelus hexagonatus (Bloch and Schneider, 1801): Schultz, 1953. Epinephelus lanceolatus (Bloch, 1790): Schultz, 1966. Epinephelus macrospilos (Bleeker, 1855). Epinephelus macrospilos Schultz, 1953. Serranus howlandi Giinther, 1883 and Epinephelus spilotus Schultz, 1953 are synonyms. Randall (1955a) erroneously regarded E. spilotus as a synonym of E. corallicola (Valenciennes). Epinephelus maculatus (Bloch, 1790). Epinephelus medurensis Schultz, 1953. Randall (1980a) regarded E. medurensis (Giinther) as a synonym of E. maculatus. x Epinephelus melanostigma Schultz, 1953. Randall (1955a) provisionally placed this species in the synonymy of E. hexagonatus. It is now known to be valid. Epinephelus merra Bloch, 1793: Schultz, 1953. Epinephelus microdon (Bleeker, 1856). Epinephelus fuscoguttatus Schultz, 1953 (in part). Epinephelus horridus Schultz, 1966. Serranus horridus Valenciennes is a junior synonym of E. fuscoguttatus (Forsskal). Epinephelus ongus (Bloch, 1790). Epinephelus summana Schultz, 1953. Randall and Ben-Tuvia (1983) have shown that E. summana (Forsskal) is endemic to the Red Sea. The related E. ongus occurs elsewhere in the Indo-Pacific. Epinephelus socialis (Giinther, 1873): Schultz, 1953. Randall (1980a) mistakenly used a photo of E. ongus from Ponape to illustrate E. socialis. Epinephelus spilotoceps Schultz, 1953. Randall (1955a) was in error referring this species to the synonymy of E. hexagonatus. Epinephelus tauvina (Forsskal, 1775). Epinephelus elongatus Schultz, 1953. Katayama (1960) and Randall (1964a) regarded this species as a synonym of E. tauvina. Gracila albomarginata (Fowler and Bean, 1930): Randall, 1986. Grammistes sexlineatus (Thunberg, 1792): Schultz, 1953. Grammistops ocellatus Schultz, 1953. Liopropoma pallidum (Fowler, 1938). Ypsigramma pallida Schultz, 1953. Liopropoma susumi (Jordan and Seale, 1906). Ypsigramma lineata Schultz, 1953. Randall (1955a) referred this species to the synonymy of Liopropoma susumi. Ypsigramma brocki Schultz, 1953. Randall (1955a) stated that Y. brocki is a probable synonym of Liopropoma susumi. This fish is also a possible hybrid of Liopropoma susumi and L. pallidum; it is still under study. Liopropoma sp. A small species with two broad red stripes on side of body which extend into caudal fin; it will be described by Randall and Taylor who are revising the genus. Luzonichthys waitei (Fowler, 1931). Luzonichthys robustus Fourmanoir, 1977. Randall (1981b) regarded L. robustus as a probable synonym of L. waitei. Plectranthias fourmanoiri Randall, 1980. Plectranthias nanus Randall, 1980. Plectropomus laevis (Lacepede, 1801). Plectropomus leopardus Schultz, 1953 (non-Lacepede). Plectropomus melanoleucus Randall, 1980a. Hoese et al., 1981, suspected that P. laevis (Lacepede) would have priority over P. melanoleucus (Lacepede); this has been confirmed (Randall and Hoese, in press). What Schultz called P. leopardus is the most common color phase of this species. liven CISReentinfiecominere pace) 298 RANDALL AND RANDALL TABLE 1 (cont’d) Family SERRANIDAE (cont'd) Plectropomus oligacanthus (Bleeker, 1854): Hiyama, 1943; Randall, 1986. Plectropomus areolatus (Riippell, 1830). Plectropomus truncatus Schultz. Randall and Hoese (in press) have shown that P. truncatus Fowler is a junior synonym of P. areolatus. Pogonoperca punctata (Valenciennes, 1830): Randall, 1986. Pseudogramma bilinearis (Schultz, 1943). Aporops bilinearis Schultz, 1953. Randall (1955a) synonymized the genus Aporops with Pseudogramma. Pseudogramma polyacantha (Bleeker, 1856): Schultz, 1953. Pseudogramma sp. A new species to be described by J. Randall who plans a revision of the genus. Variola louti (Forsskal, 1775): Schultz, 1953. Family PSEUDOCHROMIDAE Pseudochromis aurea marshallensis Schultz, 1953. Pseudochromis tapeinosoma Bleeker, 1853: Schultz, 1953. Pseudoplesiops revellei Schultz, 1953. Pseudoplesiops rosae Schultz, 1953. Pseudoplesiops sargenti Schultz, 1953. Pseudoplesiops sp. One specimen from Kwajalein. Family ACANTHOCLINIDAE Acanthoplesiops hiatti Schultz, 1953. Family PLESIOPIDAE Calloplesiops altivelis (Steindachner, 1903): Randall, 1986. Plesiops coeruleolineatus Riippell, 1835. Plesiops melas Schultz, 1953. Schultz (1966, addendum) (after Inger, 1955) reidentified his Marshall Islands material as P. coeruleolineatus. Plesiops corallicola Bleeker, 1853. Plesiops nigricans Schultz, 1953. Inger (1955) has shown, from material available to him for his revision of Plesiops, that the true P. nigricans (Riippell) is endemic to the Red Sea. Family CIRRHITIDAE Amblycirrhitus bimacula (Jenkins, 1903). Cirrhitoidea bimacula Schultz, 1960. Cirrhitichthys oxycephalus (Bleeker, 1855): Giinther, 1873; Randall, 1963a. Cirrhitus pinnulatus (Bloch and Schneider, 1801): Schultz, 1960. Isocirrhitus sexfasciatus (Schultz, 1960). Cirrhitoidea sexfasciata Schultz, 1960. Randall (1963a) created the monotypic genus Isocirrhitus for this species. Oxycirrhites typus Bleeker, 1857: Randall, 1986. Paracirrhites arcatus (Cuvier, 1829). Gymnocirrhites arcatus Schultz, 1960. Paracirrhites forsteri (Bloch and Schneider, 1801): Schultz, 1960. Paracirrhites hemistictus (Giinther, 1874): Schultz, 1960. Family KUHLIIDAE Kuhlia marginata (Cuvier, 1829): Schultz, 1953. Kuhlia mugil (Bloch and Schneider, 1801). Kuhlia taeniura Schultz, 1953. Randall (1973) showed that K. mugil is an earlier name for K. taeniura (Cuvier). Family PRIACANTHIDAE Heteropriacanthus cruentatus (Lacepede, 1801). Priacanthus cruentatus Schultz, 1953. Priacanthus hamrur (Forsskal, 1775): Schultz, 1953. Family APOGONIDAE Apogon coccineus Riippell, 1838. Apogon erythrinus Lachner, 1953. Smith (1961) could find no reason for maintaining A. erythrinus Snyder as a distinct species. Apogon cyanosoma Bleeker, 1853. Apogon novae-guineae Lachner, 1953. The author and Ernest A. Lachner have determined that the specimens tentatively identified by Lachner as A. novaequineae appear to be A. cyanosoma. Marshall Islands specimens differ from typical cyanosoma in having yellow instead of orange stripes. Apogon doryssa (Jordan and Seale, 1906): Randall, 1986. Apogon ellioti Day, 1875: Randall, 1986 (after Strasburg). Apogon evermanni Jordan and Snyder, 1904: Randall and Bohlke, 1981. (This table continued on next page.) ANNOTATED CHECKLIST OF FISHES 299 TABLE 1 (cont'd) Family APOGONIDAE (cont'd) Apogon exostigma (Jordan and Starks, 1906): Lachner, 1953. Apogon fraenatus Lachner, 1953. Randall (1955a) reidentified Lachner’s Marshall Islands specimens as A. exostigma. Apogon fraenatus Valenciennes is a valid species but apparently does not occur in the Marshall Islands. Apogon fuscus Quoy and Gaimard, 1824. Apogon savayensis Lachner, 1953. Apogon savayensis Giinther is a junior synonym of A. fuscus. Apogon guamensis Valenciennes, 1832. Apogon nubilus Lachner, 1953. Thomas H. Fraser (personal communication) informed us that A. guamensis is an earlier name for this species than A. nubilus Garman. Apogon kallopterus Bleeker, 1856. Apogon snyderi Jordan and Evermann, 1904. Apogon leptacanthus Bleeker, 1856: Strasburg, 1966. Apogon graeffii Giinther, 1873. Apogon nigrofasciatus Lachner, 1953. Apogon novemfasciatus Cuvier, 1828: Lachner, 1953. Apogon taeniophorus Regan, 1908. Apogon robustus Lachner, 1953. Randall and Lachner (1986) reidentified Marshall Islands specimens as A. taeniophorus. Apogon taeniopterus Bennett, 1835. Apogon menesemops Lachner, 1953. The authors agree with Smith (1961) in considering this species a synonym of A. taeniopterus. Apogon trimaculatus Cuvier 1828: Lachner, 1953. Apogonichthys ocellatus (Weber, 1913). Apogon ocellatus Lachner, 1953. Archamia fucata (Cantor, 1850): Lachner, 1953. Cheilodipterus isostigma (Schultz, 1940). Cheilodipterops isostigma Lachner, 1953. Randall (1955a) and Fraser (1972) placed the genera Cheilodipterops and Paramia in the synonymy of Cheilodipterus. Cheilodipterus macrodon Lacepede, 1802: Lachner, 1953. Cheilodipterus quinquelineatus Cuvier, 1828. Paramia quinquelineata Lachner, 1953. Cheilodipterus truncatus Giinther, 1873: Lachner, 1953. Foa sp. Two specimens (BPBM 28761, 28 to 29 mm) from a Halimeda bed in 19 to 27 m in the Kwajalein Lagoon may represent an undescribed species. Fowleria isostigma (Jordan and Seale, 1906). Apogon isostigma Lachner, 1953. Fowleria marmorata (Alleyne and Macleay, 1876). Apogon marmoratus Lachner, 1953. Fraser (1972) wrote that the nominal species of Fowleria should be restudied because of the contrasting treatments by Lachner (1953) and Smith (1961). Gymnapogon gracilicauda Lachner, 1953. Gymnapogon urospilotus Lachner, 1953. Rhabdamia cypselurus Weber, 1909: Fraser, 1972. Apogon cypselurus Lachner, 1953. Rhabdamia gracilis (Bleeker, 1856): Fraser, 1972. Apogon gracilis, Lachner, 1953. Siphamia fuscolineata Lachner, 1953. Family MALACANTHIDAE Hoplolatilus cuniculus Randall and Dooley, 1974: Randall, 1986. Hoplolatilus starcki Randall and Dooley, 1974. Malacanthus brevirostris Guichenot, 1858: Randall, 1986. Malacanthus latovittatus (Lacepede, 1801): Randall, 1986. Family ECHENEIDIDAE Echeneis naucrates Linnaeus, 1758: Lachner, 1966. Remora australis (Bennett, 1840). Echeneis scutata Giinther, 1876. Lachner (1966) treated this species as a synonym of R. australis. Giinther’s locality, “Maduro,” should be confirmed. Remora remora (Linnaeus, 1758): Lachner, 1966. Family CARANGIDAE Alectis ciliaris (Bloch, 1787): Randall, 1986. Atule mate (Cuvier, 1833): Randall, 1986. Selaroides leptolepis Strasburg, 1953. (This table continued on next page.) RANDALL AND RANDALL TABLE 1 (cont’d) Family CARANGIDAE (cont’d) Carangoides ferdau (Forsskal, 1775). Carangoides gilberti Woods, 1953. William F. Smith-Vaniz (personal communication) has determined that the true ferdau is the species some recent authors such as Woods have labelled gilberti, and the species usually identified as C. ferdau jordani Nichols is C. orthogrammus (Jordan and Gilbert). Carangoides orthogrammus (Jordan and Gilbert, 1881). Carangoides ferdau jordani Woods, 1953. Carangoides plagiotaenia Bleeker, 1857: Randall, 1986. Caranx ignobilis (Forsskal, 1775): Randall, 1980a. Caranx lugubris Poey, 1861: Woods, 1953. Caranx melampygus Cuvier, 1833: Woods, 1953. Caranx sexfasciatus Quoy and Gaimard, 1825: Woods, 1953. Decapterus muroadsi (Temminck and Schlegel, 1844): Woods, 1953. Elagatis bipinnulata (Quoy and Gaimard, 1825): Woods, 1953. Gnathanodon speciosus (Forsskal, 1775): Schultz, 1966, addendum; Randall, 1986 (after Strasburg). Scomberoides lysan (Forsskal, 1775). Scomberoides sancti-petri Woods, 1953. Smith-Vaniz and Staiger (1973) have shown that S. lysan is an earlier name for this species than S. sanctipetri (Cuvier). Selar crumenophthalmus (Bloch, 1793). Trachurops crumenophthalmus Woods, 1953. Trachinotus bailloni (Lacepede, 1801): Woods, 1953. Trachinotus blochii (Lacepede, 1801): Randall, 1986. Family CORYPHAENIDAE Coryphaena hippurus Linnaeus, 1758: Schultz, 1953. Family LUTJANIDAE Aphareus furca (Lacepede, 1801). Aphareus furcatus Schultz, 1953. Aprion virescens Valenciennes, 1830: Schultz, 1953. Lutjanus bohar (Forsskal, 1775): Schultz, 1953. Lutjanus fulvuus (Bloch and Schneider, 1801): Randall, 1986 (after Strasburg). Lutjanus marginatus Kendall and Goldsborough, 1911. Lutjanus flavipes Hiyama, 1943. Randall (1973) noted that L. fulvus predates L. vaigiensis (Quoy and Gaimard) and L. marginatus (Cuvier), the names usually given to this common lutjanid. Lutjanus gibbus (Forsskal, 1775): Schultz, 1953. Anthias heraldi Schultz, 1953. Heemstra (1972) showed that this taxon is a juvenile L. gibbus. Lutjanus kasmira (Forsskal, 1775): Schultz, 1953. Lutjanus monostigma (Cuvier, 1828). Lutjanus monostigmus Schultz, 1953. Lutjanus semicinctus Quoy and Gaimard, 1824: Hiyama, 1943; Randall, 1986. Lutjanus vitta (Quoy and Gaimard, 1824): Randall, 1986 (after Strasburg). Lutjanus semicinctus Quoy and Gaimard, 1824: Hiyama, 1943; Randall, 1986. Lutjanus vitta (Quoy and Gaimard, 1824): Randall, 1986 (after Strasburg). Macolor niger (Forsskal, 1775): Schultz, 1953. Family CAESIONIDAE Caesio caerulaurea Lacepede, 1801: Randall, 1986. Caesio teres Seale, 1906. Caesio xanthonotus Schultz, 1953. Pterocaesio marri Schultz, 1953. Pterocaesio kohleri Schultz, 1953. Kent E. Carpenter, who is revising the family, believes this is the same species as P. marri. Pterocaesio tile (Cuvier, 1830): Schultz, 1953. Pterocaesio sp. Family HAEMULIDAE Plectorhinchus obscurus (Giinther, 1871): Randall, 1986. Plectorhinchus picus (Cuvier, 1830): Randall, 1986. Family LETHRINIDAE Gnathodentex aureolineatus (Lacepede, 1802): Schultz, 1953. Gymnocranius microdon (Bleeker, 1851): Randall, 1986 (after Strasburg) used the name microdon for the Marshall Islands Gymnocranius with reservations. The genus Gymnocranius needs revision. Lethrinus amboinensis Bleeker, 1854: Randall, 1980a (after Sato, 1978). Lethrinus variegatus Schultz, 1953. (This table continued on next ANNOTATED CHECKLIST OF FISHES 301 TABLE 1 (cont'd) Family LETHRINIDAE (cont'd) ?Lethrinus reticulatus Schultz, 1953. Schultz provided no illustrations of any of the five species of Lethrinus he reported from the Marshall Islands, and Sato (1978) made reference to only two of them, kallopterus and miniatus (elongatus). The small specimens identified as reticulatus by Schultz are only tentatively regarded as amboinensis here. Lethrinus elongatus Valenciennes, 1830. Lethrinus miniatus Schultz, 1953. Randall and Wheeler (MS) have determined that the true L. miniatus (Bloch and Schneider) is a species most authors have identified as L. chrysostomus Richardson. Lethrinus harak (Forsskal, 1775): Schultz, 1953. Lethrinus kallopterus Bleeker, 1856: Schultz, 1953. Lethrinus ramak (Forsskal, 1775): Randall, 1986. Lethrinus xanthochilus Klunzinger, 1870: Randall, 1980a. Lethrinus microdon Schultz, 1953. Monotaxis grandoculis (Forsskal, 1775). Family NEMIPTERIDAE Pentapodus caninus (Cuvier, 1830): Randall, 1986. Scolopsis lineatus Quoy and Gaimard, 1824. Scolopsis cancellatus Strasburg, 1953 (mimeo report to ONR); Hiatt and Strasburg, 1960. Barry C. Russell (personal communication) has informed us that S. lineatus is an older name for this species than S. cancellatus (Cuvier). Family GERREIDAE Gerres argyreus (Bloch and Schneider, 1801): Schultz, 1953. Family MULLIDAE Mulloides flavolineatus (Lacepede, 1801). Mulloidichthys samoensis Lachner, 1960. Mulloides vanicolensis (Valenciennes, 1831). Mulloidichthys auriflamma Lachner, 1960. Mulloidichthys vanicolensis Lachner, 1960. Mulloides pflugeri Steindachner, 1901: Randall, 1986. Parupeneus barberinoides (Bleeker, 1852): Randall, 1986. Upeneus atrocingulatus Kner, 1870. Lachner (1960) erred in considering U. atrocingulatus as a synonym of P. trifasciatus. Parupeneus barberinus (Lacepede, 1801): Lachner, 1960. Parupeneus bifasciatus (Lacepede, 1801): Randall, 1986. Parupeneus crassilabrus Lachner, 1960. Parupeneus cyclostomus (Lacepede, 1801): Lachner, 1960. Parupeneus heptacanthus (Lacepede, 1802): Randall, 1986. Parupeneus multifasciatus (Quoy and Gaimard, 1824). Parupeneus trifasciatus Lachner, 1960. Randall (MS) regards P. trifasciatus (Lacepede) as a synonym of P. bifasciatus (Lacepede). The species in the central and western Pacific identified by most authors as trifasciatus should be called P. multifasciatus. It is not known from the western Indian Ocean. Parupeneus pleurostigma (Bennett, 1831): Lachner, 1960. Upeneus taeniopterus Cuvier, 1829: Randall, 1986. Family PEMPHERIDIDAE Parapriacanthus beryciformes Franz, 1910: Schultz, 1953. Pempheris oualensis Cuvier, 1831: Schultz, 1953. Family KYPHOSIDAE Kyphosus cinerascens (Forsskal, 1775): Schultz, 1953. Kyphosus vaigiensis (Quoy and Gaimard, 1825): Randall, 1986. Family EPHIPPIDIDAE Platax orbicularis (Forsskal, 1775): Randall, 1986 (after Strasburg). Family CHAETODONTIDAE Chaetodon auriga Forsskal, 1775: Woods, 1953. Chaetodon bennetti Cuvier, 1831: Burgess, 1978. Chaetodon citrinellus Cuvier, 1831: Woods, 1953. Chaetodon ephippium Cuvier, 1831: Woods, 1953. Chaetodon kleinii Bloch, 1790: Woods, 1953. Chaetodon lineolatus Cuvier, 1831: Burgess, 1978. Chaetodon lunula (Lacepede, 1802): Woods, 1953. Chaetodon melannotus Bloch and Schneider, 1801: Woods, 1953. Chaetodon mertensii Cuvier, 1831: Woods, 1953. Chaetodon meyeri Bloch and Schneider, 1801: Woods, 1953. Chaetodon ornatissimus Cuvier 1831: Woods, 1953. : : (This table continued on next page.) 302 RANDALL AND RANDALL TABLE 1 (cont'd) Family CHAETODONTIDAE (cont’d) Chaetodon punctatofasciatus Cuvier, 1831: Woods, 1953. Chaetodon quadrimaculatus Gray, 1833: Woods, 1953. Chaetodon rafflesi Bennett, 1830: Burgess, 1978 (from underwater photo by Scott Johnson, but locality Kwajalein, not Enewetak). Chaetodon reticulatus Cuvier, 1831: Woods, 1953. Chaetodon semeion Bleeker, 1855: Woods, 1953. Chaetodon tinkeri Schultz, 1951: Randall, 1986. Chaetodon trifascialis Quoy and Gaimard, 1825. Megaprotodon strigangulus Woods, 1953. Chaetodon trifascialis Mungo Park, 1797: Woods, 1953. Chaetodon ulietensis Cuvier, 1831. Chaetodon falcula Woods, 1953. Burgess (1978) wrote that C. falcula Bloch is an Indian Ocean species that is often not distinguished from the Pacific C. ulietensis. Chaetodon unimaculatus Bloch, 1787: Woods, 1953. Chaetodon vagabundus Linnaeus, 1758: Randall, 1986 (after Strasburg). Forcipiger flavissimus Jordan and McGregor, 1898. Forcipiger longirostris Woods, 1953. Randall and Caldwell (1970) showed that most authors have used the name longirostris for the most common of the two species of the genus (the one with the shorter snout); the correct name for this butterflyfish is F. flavissimus. Forcipiger longirostris (Broussonet, 1782): Randall and Caldwell, 1970. Hemitaurichthys polylepis (Bleeker, 1857): Burgess, 1978. Heniochus acuminatus (Linnaeus, 1758): Fowler, 1928. Heniochus chrysostomus Cuvier, 1831. Heniochus permutatus Woods, 1953. Burgess (1978) reidentified Woods’ material of this species from the Marshall Islands as H. chrysostomus. Heniochus monoceros Cuvier, 1831: Woods, 1953. Heniochus varius (Cuvier, 1829): Randall, 1986. Family POMACANTHIDAE Centropyge bicolor (Bloch, 1787): Randall, 1986. Centropyge bispinosus (Giinther, 1860): Woods and Schultz, 1953. Centropyge flavissimus (Cuvier, 1831): Woods and Schultz, 1953. Centropyge heraldi Woods and Schultz, 1953. Centropyge loriculus (Giinther, 1874): Randall, 1986. Centropyge multicolor Randall and Wass, 1974. Centropyge multifasciatus (Smith and Radcliffe, 1911): Randall, 1986. Centropyge urolikii (Bleeker, 1853): Randall, 1986 (after Strasburg). Genicanthus bellus Randall, 1975: Randall, 1986. Genicanthus watanabei (Yasuda and Tominaga, 1970): Randall, 1986. Pomacanthus imperator (Bloch, 1787): Woods and Schultz, 1953. Pygoplites diacanthus (Boddaert, 1772): Woods and Schultz, 1953. Family POMACENTRIDAE Abudefduf septemfasciatus (Cuvier, 1830): Woods and Schultz, 1953. Abudefduf sexfasciatus (Lacepede, 1801). Abudefduf coelestinus Allen, 1975. Allen, Bauchot, and Desoutter (1978) placed A. coelestinus (Cuvier) in the synonymy of A. sexfasciatus. Abudefduf sordidus (Forsskal, 1775): Woods and Schultz, 1960. Amblyglyphidodon aureus (Cuvier, 1830). Abudefduf aureus Woods and Schultz, 1960. Amblyglyphidodon curacao (Bloch, 1787). Abudefduf curacao Woods and Schultz, 1960. Amblyglyphidodon leucogaster (Bleeker, 1847): Randall, 1986. Amphiprion chrysopterus Cuvier, 1830. Ampbhiprion bicinctus Woods and Shultz, 1960. Allen (1972a) found specimens of the true A. bicinctus Riippell only from the Red Sea. Amphiprion melanopus Bleeker, 1852: Woods and Schultz, 1960. Amphiprion perideraion Bleeker, 1955: Woods and Schultz, 1960. Ampbhiprion tricinctus Schultz and Welander, 1953. Chromis acares Randall and Swerdloff, 1973. Chromis agilis Smith, 1960: Randall and Swerdloff, 1973. Chromis leucurus Woods and Schultz, 1960. (This table continued on next page.) ANNOTATED CHECKLIST OF FISHES 303 TABLE 1 (cont’d) Family POMACENTRIDAE (cont'd) Chromis amboinensis (Bleeker, 1873): Allen, 1975. Chromis atripectoralis Welander and Schultz, 1951. Chromis atripes Fowler and Bean, 1928: Randall, 1986. Chromis elerae Fowler and Bean, 1928: Randall, 1986. Chromis lepidolepis Bleeker, 1877: Woods and Schultz, 1960. Chromis margaritifer Fowler, 1946. Chromis dimidiatus Woods and Schultz, 1960. Randall, Ida, and Moyer (1981) stated that C. dimidiata (Klunzinger) is an Indian Ocean species. Chromis ternatensis (Bleeker, 1956): Woods and Schultz, 1960. Recent examination of the syntypes of Chromis caerulea (Cuvier, 1830) revealed them to be the damselfish widely known as C. ternatensis. A petition has been prepared for the International Commission on Zoological Nomenclature to suppress C. caerulea in favor of C. ternatensis. The next available name for the species known as C. caerulea is C. viridis (Cuvier, 1830) (Randall, Bauchot, and Desoutter, 1985). Chromis vanderbilti (Fowler, 1941): Randall, 1986. Chremis viridis (Cuvier, 1830). Chromis caeruleus Woods and Schultz, 1960. See remarks under C. ternatensis above. Chromis xanthura (Bleeker, 1854): Allen, 1975. Chromis sp. Chromis sp. A of Allen, 1975. Chrysiptera biocellata (Quoy and Gaimard, 1825). Abudefduf biocellatus Woods and Schultz, 1960. Chrysiptera caeruleolineata (Allen, 1973): Randall, 1986. Chrysiptera glauca (Cuvier, 1830). Abudefduf glaucus Woods and Schultz, 1960. Chrysiptera leucopoma (Lesson, 1830). Abudefduf leucopomus Woods and Schultz, 1960. Abudefduf amabilis Woods and Schultz, 1960. Allen (1975) discovered that amabilis and leucopomus are two very different color forms of the same species associated with different reef flat zones. Chrysiptera traceyi Woods and Schultz, 1960. Dascyllus aruanus (Linnaeus, 1758): Woods and Schultz, 1960. Dascyllus reticulatus (Richardson, 1846): Woods and Schultz, 1960. Dascyllus trimaculatus (Riippell, 1828): Kendall and Goldsborough, 1911; Woods and Schultz, 1960. Lepidozygus tapeinosoma (Bleeker, 1856): Randall, 1986. Plectroglyphidodon dickii (Liénard, 1839). Abudefduf dicki Woods and Schultz, 1960. Plectroglyphidodon imparipennis (Vaillant and Sauvage, 1875). Abudefduf imparipennis Woods and Schultz, 1960. Plectroglyphidodon johnstonianus (Fowler and Ball, 1924). Abudefduf johnstonianus Woods and Schultz, 1960. Plectroglyphidodon lacrymatus (Quoy and Gaimard, 1825). Abudefduf lacrymatus Woods and Schultz, 1960. Plectroglyphidodon leucozona (Bleeker, 1859). Abudefduf leucozona Woods and Schultz, 1960. Plectroglyphidodon phoenixensis (Schultz, 1943). Abudefduf phoenixensis Woods and Schultz, 1960. Pomacentrus amboinensis Bleeker, 1868: Randall, 1986 (after Strasburg). Pomacentrus brachialis (Cuvier, 1830). Pomacentrus melanopterus Woods and Schultz, 1960. Gerald R. Allen (personal communication) has replaced the name P. melanopterus Bleeker with P. brachialis. Pomacentrus coelestis Jordan and Starks, 1901: Woods and Schultz, 1960. Pomacentrus pavo (Bloch, 1787): Woods and Schultz, 1960. Pomacentrus vaiuli Jordan and Seale, 1906: Woods and Schultz, 1960. Pomachromis exilis (Allen and Emery, 1973). Stegastes fasciolatus (Ogilby, 1889). Pomacentrus jenkinsi Woods and Schultz, 1960. Allen (1975) found that fasciolatus is an earlier name for P. jenkinsi Jordan and Evermann. At that time he classified it and the following species in the genus Eupomacentrus. Emery and Allen (1980) determined that Stegastes is a senior generic synonym for Eupomacentrus. Stegastes nigricans (Lacepede, 1803). Pomacentrus nigricans Woods and Schultz, 1960. Stegastes lividus (Bloch and Schneider, 1801). Pomacentrus lividus Giinther, 1881. Giiverable confinted’on nextapage) RANDALL AND RANDALL TABLE 1 (cont'd) Family MUGILIDAE Chaenomugil leuciscus (Giinther, 1871). Neomyxus chaptalii Schultz, 1953. James M. Thomson (personal communication) stated that Chaenomugil leuciscus is the correct name for the specimens Schultz identified as N. chaptalii (Eydoux and Souleyet). This and the following mugilid generic placements were made with his counsel. Chelon labiosus (Valenciennes, 1836). Plicomugil labiosus Schultz, 1953. Crenimugil crenilabis (Forsskal, 1775): Schultz, 1953. Liza vaigiensis (Quoy and Gaimard, 1825). Chelon vaigiensis Schultz, 1953. Valamugil engeli (Bleeker, 1858). Chelon engeli Schultz, 1953. Family SPHYRAENIDAE Sphyraena barracuda (Walbaum, 1792): Schultz, 1953. Sphyraena forsteri Cuvier, 1829: Schultz, 1953. Sphyraena helleri Jenkins, 1901: Schultz, 1953. Sphyraena genie Klunzinger, 1870: Schultz, 1953. Family POLYNEMIDAE Polydactylus sexfilis (Valenciennes, 1831): Schultz, 1953. Family LABRIDAE Anampses caeruleopunctatus Riippell, 1828: Schultz, 1960. Anampses melanurus Bleeker, 1857: Randall, 1986. Anampses meleagrides Valenciennes, 1839: Randall, 1986. Anampses twistii Bleeker, 1856: Schultz, 1960. Bodianus anthioides (Bennett, 1831): Randall, 1986. Bodianus axillaris (Bennett, 1831): Randall, 1986. Bodianus diana (Lacepede, 1801): Randall, 1986. Bodianus loxozonus (Snyder, 1908): Randall, 1986. Cheilinus arenatus Valenciennes, 1840: Randall, 1981c. Cheilinus bimaculatus Valenciennes, 1840: Randall, 1986. Cheilinus celebicus Bleeker, 1853: Schultz, 1960. Cheilinus chlorourus (Bloch, 1791): Schultz, 1960. Cheilinus digrammus (Lacepede, 1801): Schultz, 1960. Cheilinus fasciatus (Bloch, 1791): Schultz, 1960. Cheilinus orientalis Giinther, 1862: Randall, 1986. Cheilinus oxycephalus Bleeker, 1853: Schultz, 1960. Cheilinus trilobatus Lacepede, 1801: Schultz, 1960. Cheilinus undulatus Riippell, 1835: Randall, Head, and Sanders, 1978. Cheilinus unifasciatus Streets, 1877. Cheilinus rhodochrous Woods, 1960. Cheilio inermis (Forsskal, 1775): Randall, 1986. Cirrhilabrus exquisitus Smith, 1957: Randall, 1986. Cirrhilabrus sp. Cirrhilabrus temmincki Schultz, 1960. This is an undescribed species allied to C. temminckii Bleeker (Randall, MS). Cirrhilabrus sp. A new species, found on dense beds of benthic algae in lagoons of Enewetak and Kwajalein in 18.5 to 27.5 m (Randall, MS). Cirrhilabrus sp. A new species related to C. cyanopleura (Bleeker) (Randall, MS). Cirrhilabrus sp. A new species found in outer reef areas in 30 m or more (Randall, MS). Coris aygula Lacepede, 1801: Schultz, 1960. Coris gaimard (Quoy and Gaimard, 1824): Schultz, 1960. Coris variegata (Riippell, 1835): Schultz, 1960. Cymolutes praetextatus (Quoy and Gaimard, 1834): Schultz, 1960. Cymolutes torquatus (Valenciennes, 1839): Randall, 1986. Epibulus insidiator (Pallas, 1770): Schultz, 1960. Gomphosus varius Lacepede, 1801: Schultz, 1960. Gomphosus tricolor Schultz, 1960. Strasburg and Hiatt (1957) discovered that G. tricolor Quoy and Gaimard is the male of G. varius. Halichoeres biocellatus Schultz, 1960. Halichoeres chrysus Randall, 1980. Halichoeres hortulanus (Lacepede, 1801): Schultz, 1960. (This table continued on next page.) ANNOTATED CHECKLIST OF FISHES 305 TABLE 1 (cont’d) Family LABRIDAE (cont’d) Halichoeres margaritaceus (Valenciennes, 1839): Schultz, 1960. Halichoeres marginatus Riippell, 1835: Schultz, 1960. Halichoeres melanurus (Bleeker, 1851). Halichoeres hoeveni Schultz, 1960. Halichoeres kallochroma Schultz, 1960. Randall (1980b) concluded that Schultz misidentified the female of H. melanurus as H. hoevenii Bleeker [actually hoevenii is the female of H. vrolikii (Bleeker)] and the male as H. kallochroma (Bleeker). Halichoeres melasmapomus Randall, 1980. Halichoeres richmondi Fowler and Bean, 1928: Randall, 1986. Halichoeres trimaculatus (Quoy and Gaimard, 1834): Schultz, 1960. Halichoeres sp. Probably undescribed; close to H. hartzfeldii (Bleeker). Hemigymnus fasciatus (Bloch, 1792): Schultz, 1960. Hemigymnus melapterus (Bloch, 1791): Schultz, 1960. Hologymnosus annulatus (Lacepede, 1801): Randall, 1982. Labrichthys unilineatus (Guichenot, 1847). Labrichthys cyanotaenia Schultz, 1960. Randall and Springer (1973) showed that L. unilineatus has priority over L. cyanotaenia Bleeker. Labroides bicolor Fowler and Bean, 1928: Schultz, 1960. Labroides dimidiatus (Valenciennes, 1839): Schultz, 1960. Labroides pectoralis Randall and Springer, He: Labropsis alleni Randall, 1981. Labropsis micronesica Randall, 1981. Labropsis xanthonota Randall, 1981. Macropharyngodon meleagris (Valenciennes, 1839): Schultz, 1960. Macropharyngodon pardalis Schultz, 1960. Randall (1978b) determined that M. pardalis (Kner) is the female of M. meleagris. Macropharyngodon negrosensis Herre, 1932: Randall, 1978b. Novaculichthys taeniourus (Lacepede, 1801). Xyrichthys taeniourus Schultz, 1960. Paracheilinus sp. An undescribed species related to P. filamentosus Allen (Randall, MS). Pseudocheilinus evanidus Jordan and Evermann, 1903: Randall, 1986 (after Strasburg). Pseudocheilinus hexataenia (Bleeker, 1857): Schultz, 1960. Pseudocheilinus octotaenia Jenkins, 1900: Schultz, 1960. Pseudocheilinus tetrataenia Schultz, 1960. Pseudocheilinus sp. A new species with a large ocellated black spot on side of caudal peduncle. Pseudocoris yamashiroi (Schmidt, 1930): Smith-Vaniz and Randall (MS) discuss this species in their revision of the genus. Pseudodax moluccanus (Valenciennes, 1839): Randall, 1986. Pseudojuloides cerasinus (Snyder, 1904): Randall, 1986. Pteragogus cryptus Randall, 1981. Pteragogus guttatus Schultz, 1960. Randall (1981c) reidentified the specimen from Rongelap which Schultz called P. guttatus (Fowler and Bean). Stethojulis bandanensis (Bleeker, 1851). Stethojulis axillaris Schultz, 1960. Stethojulis linearis Schultz, 1960. Randall and Kay (1974) showed that S. axillaris (Quoy and Gaimard) is the initial phase of the Hawaiian endemic species S. balteata (Quoy and Gaimard), and S. linearis Schultz is the terminal male of S. bandanensis. Stethojulis strigiventer (Bennett, 1832): Schultz, 1960. Thalassoma amblycephalum (Bleeker, 1856). Thalassoma amblycephalus Schultz, 1960. Thalassoma hardwicke (Bennett, 1830). Thalassoma hardwickei Schultz, 1960. Thalassoma lunare (Linnaeus, 1758): Schultz, 1960. Thalassoma lutescens (Lay and Bennett 1839): Schultz, 1960. Thalassoma purpureum (Forsskal, 1775): Schultz, 1960. Thalassoma umbrostygma Schultz, 1960 (in part). Randall and Edwards (1984) have shown that T. umbrostygma (Riippell) is a junior synonym of T. purpureum based on the initial phase (holotype of umbrostygma examined in Senckenberg Museum). The initial phase of purpureum is nearly identical in color to that of T. trilobatum. Thalassoma quinquevittatum (Lay and Bennett, 1839). Thalassoma quinquevittatus Schultz, 1960. (This table continued on next page.) RANDALL AND RANDALL TABLE 1 (cont’d) Family LABRIDAE (cont'd) Thalassoma trilobatum (Lacepede, 1801). Thalassoma fuscum Schultz, 1960. Thalassoma fuscum (Lacepede), described in Labrus, is a junior homonym of L. fuscus Gmelin, as pointed out by Randall, 1955a. Thalassoma umbrostygma Schultz, 1960 (in part). Wetmorella albofasciata Schultz and Marshall, 1954: Randall, 1983. Wetmorella nigropinnata (Seale, 1900). Wetmorella ocellata Schultz and Marshall, 1954. Randall (1983) determined that W. ocellata is a junior synonym of W. nigropinnata. Xyrichtys aneitensis (Giinther, 1862). Hemipteronotus aneitensis Schultz, 1960. Xyrichtys celebicus (Bleeker, 1856). Hemipteronotus celebicus Schultz, 1960. Xyrichtys pavo Valenciennes, 1839: Randall, 1986. Family SCARIDAE Calotomus carolinus (Valenciennes, 1839). Calotomus spinidens Schultz, 1958; Schultz, 1960. Bruce and Randall (1985) showed that Schultz and some other authors have misidentified C. carolinus as C. spinidens (Quoy and Gaimard). Calotomus spinidens (Quoy and Gaimard, 1824): Randall, 1986. Cetoscarus bicolor (Riippell, 1829). Chlorurus bicolor Schultz, 1958; Schultz, 1960. Chlorurus pulchellus Schultz, 1958; Schultz, 1960. Randall (1963b) found that Cetoscarus pulchellus (Rtippell) is the terminal male of C. bicolor. Hipposcarus longiceps (Valenciennes, 1839). Scarus harid Schultz, 1958; Schultz, 1960. Randall and Bruce (1983) showed that Hipposcarus harid (Forsskal) is an Indian Ocean species distinct from the Pacific H. longiceps. Scarus altipinnis (Steindachner, 1879). Scarus brevifilis Schultz, 1958; Schultz, 1960. Scarus chlorodon Schultz, 1958; Schultz, 1960. Randall and Choat (1980) pointed out that S. chlorodon Jenyns is the terminal male (and a junior synonym) of S. prasiognathos Valenciennes. Scarus brevifilis (Giinther) is a junior synonym of S. altipinnis based on the initial phase. Scarus atropectoralis Schultz, 1958; Randall, 1986. Scarus bleekeri (de Beaufort, 1940): Randall and Choat, 1980. Scarus dimidiatus Bleeker, 1859: Randall, 1986. Scarus festivus Valenciennes, 1840. Scarus lunula Schultz, 1958; Schultz 1960. Randall and Bruce (1983) determined that S. festivus is an older name for S. lunula (Snyder). Scarus flavipectoralis Schultz, 1958: Randall and Choat, 1980. Scarus forsteni (Bleeker, 1861): Randall, 1986. Schultz (1958; 1969) confused two scarids under the name S. lepidus (non-Jenys). Randall and Choat (1980) followed but used the name S. tricolor Bleeker. Randall (1986) has shown that the true S. tricolor apparently does not occur in the Marshall Islands, but the related S. forsteni does. Scarus frenatus Lacepede, 1802: Randall, 1986. Scarus frontalis Valenciennes, 1839. Scarus jonesi Schultz, 1958; Schultz, 1960. Randall and Bruce (1983) mentioned that S. frontalis has priority over S. jonesi (Streets) in their account of the related S. enneacanthus Lacepede. Scarus ghobban Forsskal, 1775: Randall, 1986. Scarus gibbus Riippell, 1828. Scarus microrhinos Schultz, 1958; Schultz, 1960. Smith (1959) showed that Schultz (1958) was in error in applying the name Chlorurus gibbus (Riippell) to the largest of the parrotfishes, Bolbometopon muricatum (Valenciennes). Bolbometopon muricatum is not known from the Marshall Islands, but it occurs in the Gilberts and the Line Islands so it might eventually be found in the southern Marshalls. Scarus microrhinos Bleeker is regarded as a junior synonym of S. gibbus by Randall and Bruce (1983), following Schultz (1969). Scarus globiceps Valenciennes, 1840: Schultz, 1958; Schultz, 1960. Scarus aeruginosus Schultz, 1958, 1960 (in part). Randall and Bruce (1983) documented this misidentification. The true S. aeruginosus Valenciennes is a junior synonym of the Red Sea endemic S. ferrugineus Forsskal (Randall and Ormond, 1978). Scarus niger Forsskal, 1775: Randall, 1986. Scarus oviceps Valenciennes, 1839: Randall, 1986. Scarus psittacus Forsskal, 1775. Scarus forsteri Schultz, 1958; Schultz, 1960. Randall and Ormond (1978) demonstrated that S. psittacus (the type species of Scarus) is a senior synonym of S. forsteri Valenciennes; they described a neotype of S. psittacus. (This table continued on next page.) ANNOTATED CHECKLIST OF FISHES 307 TABLE 1 (cont'd) Family SCARIDAE (cont'd) Scarus taeniurus Schultz, 1958; Schultz, 1960. Scarus taeniurus Valenciennes is also a junior synonym of S. psittacus (Randall and Bruce, 1983). Scarus rubroviolaceus Bleeker, 1849: Randall, 1980a. Scarus schlegeli (Bleeker, 1861): Schultz, 1958; Schultz, 1960. Scarus sordidus Forsskal, 1775: Schultz, 1958; Schultz, 1960. Scarus spinus (Kner, 1868). Scarus formosus Schultz, 1958; Schultz, 1960. Randall and Choat (1980) showed that the true S. formosus Valenciennes is a junior synonym of the Hawaiian endemic species S. dubius Bennett. Family CREEDIIDAE Chalixodytes tauensis Schultz, 1943: Schultz, 1960. Limnichthys donaldsoni Schultz, 1960. Family TRICHONOTIDAE Trichonotus sp. Closely related to T. elegans Shimada and Yoshino, 1984. Family MUGILOIDIDAE Parapercis millipunctata (Giinther, 1860). Parapercis cephalopunctata Schultz, 1960. Randall (MS) has shown that Percis cephalopunctatus Seale is a junior synonym of Parapercis millipunctata. Parapercis clathrata Ogilby, 1910: Schultz, 1960. Parapercis cylindrica (Bloch, 1792): Schultz, 1960. Family BLENNIIDAE Aspidontus dussumieri (Valenciennes, 1836). Petroscirtes quadrimaculatus Kendall and Goldsborough, 1911. Petroscirtes fluctuans Schultz, 1960. Smith-Vaniz and Randall (1973) found that Aspidontus dussumieri is an older name for P. fluctuans Weber. Aspidontus taeniatus Quoy and Gaimard, 1834: Schultz, 1960. Astrosalarias fuscus holomelas (Giinther, 1866): Springer and Smith-Vaniz (1968); Smith-Vaniz and Springer (1971). Cirripectus fuscoguttatus Strasburg and Schultz, 1953: Schultz and Chapman, 1960. Cirripectes polyzona (Bleeker, 1868). Cirripectes sebae Schultz and Chapman, 1960. Jeffrey T. Williams (MS) has determined that C. sebae (Valenciennes) is a junior synonym of C. castaneus (Valenciennes); thus C. polyzona is the correct name for the Schultz and Chapman specimens. Cirripectes quagga (Fowler and Ball, 1924): Schultz and Chapman, 1960. Cirripectes stigmaticus Strasburg and Schultz, 1953: Schultz and Chapman, 1960. Cirripectus variolosus (Valenciennes, 1836): Schultz and Chapman, 1960. Ecsenius bicolor (Day, 1888): Strasburg, 1967. Ecsenius opsifrontalis Chapman and Schultz, 1952; Schultz and Chapman, 1960. Entomacrodus caudofasciatus (Regan, 1909): Springer, 1982; Randall, 1986. Entomacrodus cymatobiotus Schultz and Chapman, 1960. Entomacrodus sealei Bryan and Herre, 1903. Entomacrodus incisolabilatus Schultz and Chapman, 1960. Springer (1967) referred this taxon to the synonymy of E. sealei. Entomacrodus striatus (Valenciennes, 1836). Salarias marmoratus Giinther, 1877 (non-Blennius marmoratus Bennett). Entomacrodus plurifilis marshallensis Schultz and Chapman, 1960. Springer (1967) placed this nominal subspecies in the synonymy of E. striatus. Entomacrodus thalassinus thalassinus (Jordan and Seale, 1906): Springer, 1967. Entomacrodus thalassinus Schultz and Chapman, 1960. Exallias brevis (Kner, 1868): Schultz and Chapman, 1960. Istiblennius coronatus (Giinther, 1872): Schultz and Chapman, 1960. These authors stated that J. nitidus (Giinther, 1877) (non-nitidus Giinther, 1861) is the male form. Giinther recorded it from Jaluit. Istiblennius edentulus (Bloch and Schneider, 1801): Schultz and Chapman, 1960. Istiblennius gibbifrons (Quoy and Gaimard, 1824). Istiblennius rodenbaughi Schultz and Chapman, 1960. Doubtfully distinct at the species level from gibbifrons. Istiblennius lineatus (Valenciennes, 1836): Schultz and Chapman, 1960. Istiblennius paulus (Bryan and Herre, 1903): Schultz and Chapman, 1960. Meiacanthus atrodorsalis atrodorsalis (Giinther, 1877): Smith-Vaniz, 1976. Meiacanthus atrodorsalis Schultz, 1960. Parenchelyurus hepburni (Snyder, 1908): Springer, 1972. Petroscirtes mitratus Riippell, 1830: Schultz, 1960. Petroscirtes xestus Jordan and Seale, 1906: Smith-Vaniz, 1976. Aihisableeontinued on rextiaace) 308 RANDALL AND RANDALL TABLE 1 (cont’d) Family BLENNIIDAE (cont'd) Plagiotremus laudandus laudandus (Whitley, 1961): Smith-Vaniz, 1976. Plagiotremus rhinorhynchos (Bleeker, 1852): Randall, 1986. Plagiotremus tapeinosoma (Bleeker, 1857): Schultz, 1960. Praealticus sp. An unidentified species from an underwater photograph taken in Kwajalein. Rhabdoblennius rhabdotrachelus (Fowler and Ball, 1924): Schultz and Chapman, 1960. Rhabdoblennius snowi (Fowler, 1928): Schultz and Chapman, 1960. Salarias fasciatus (Bloch, 1786): Randall, 1986 (after Strasburg). Stanulus seychellensis Smith, 1959. Fallacirripectes minutus Schultz and Chapman, 1960 and F. wellsi Schultz and Chapman, 1960. Smith-Vaniz and Springer (1971) relegated F. minutus and F. wellsi to the synonymy of Stanulus seychellensis. Family TRIPTERYGIIDAE Enneapterygius capidata (Rosenblatt in Schultz, 1960). Enneapterygius chica (Rosenblatt in Schultz, 1960). Enneapterygius hemimelas (Kner and Steindachner, 1866): Schultz, 1960. Enneapterygius minutus (Giinther, 1877): Schultz, 1960. Enneapterygius nanus Schultz, 1960. Helcogramma hudsoni (Jordan and Seale, 1906): Hansen, 1982. Norfolkia brachylepis (Schultz, 1960). Tripterygion brachylepis Schultz, 1960. Family SCHINDLERIIDAE Schindleria praematurus (Schlindler, 1930): Schultz, 1960. Family CALLIONYMIDAE Callionymus simplicicornis Valenciennes, 1837: Fricke, 1982. Diplogrammus goramensis (Bleeker, 1858): Schultz, 1960. Synchiropus laddi Schultz, 1960. Synchiropus morrisoni Schultz, 1960. Synchiropus ocellatus (Pallas, 1770): Randall, 1986. Family GOBIIDAE Allomicrodesmus dorotheae Schultz, 1966. Dawson (1974) showed that this species is not a microdesmid; Victor G. Springer (personal communication) classifies it as a goby of the subfamily Xenisthminae. Amblveleotris fasciata (Herre, 1953). Amblveleotris guttata (Fowler, 1938). Amblveleotris steinitzi (Klausewitz, 1974). Amblyeleotris wheeleri Polunin and Lubbock, 1977. Amblygobius decussatus (Bleeker, 1855). Amblygobius phalaena (Valenciennes, 1837). Amblygobius rainfordi (Whitley, 1940). Asterropteryx ensiferus (Bleeker, 1865). Asterropteryx semipunctatus Riippell, 1830. Bathygobius cocosensis (Bleeker, 1854). Bathygobius fuscus fuscus (Riippell, 1830). _Cabillus tongarevae (Fowler, 1927). Callogobius bauchotae Goren, 1979. Callogobius centrolepis Weber, 1909. Callogobius hasselti (Bleeker, 1851). Callogobius maculipinnis (Fowler, 1918). Callogobius okinawae (Snyder, 1980). Callogobius sclateri (Steindachner, 1880). Callogobius sp. Cryptocentrus strigilliceps (Jordan and Seale, 1906). Ctenogobiops aurocingulus (Herre, 1935). Ctenogobiops sp. Eviota afelei Jordan and Seale, 1906. Eviota distigma Jordan and Seale, 1906. Eviota fasciola Karnella and Lachner, 1981. Eviota infulata (Smith, 1956). Eviota melasma Lachner and Karnella, 1980. Eviota nebulosa Smith, 1958. Eviota sebreei Jordan and Seale, 1906. Eviota smaragdus Jordan and Seale, 1906. (This table continued on next page.) ANNOTATED CHECKLIST OF FISHES TABLE 1 (cont’d) Family GOBIIDAE (cont'd) Eviota zonura Jordan and Seale, 1906. Eviota spp. (2 species). Fusigobius neophytus (Giinther, 1877). Fusigobius spp. (2 species). Gnatholepis anjerensis (Bleeker, 1850). Gnatholepis scapulostigma Herre, 1953. Gobiodon citrinus (Riippell, 1838). Gobiodon erythrospilus Bleeker, 1875. Gobiodon okinawae Sawada, Arai, and Abe, 1972. Gobiodon rivulatus (Riippell, 1830): Giinther, 1877. Hetereleotris sp. Istigobius decoratus (Herre, 1927). Istigobius ornatus (Riippell, 1930). Istigobius rigilius (Herre, 1953). Kelloggella centralis Hoese, 1975. Lotilia graciliosa Klausewitz, 1960. Macrodontogobius wilburi Herre, 1936. Oplopomops diacanthus (Schultz, 1943). Oplopomus oplopomus (Valenciennes, 1837). Opua nephodes E. K. Jordan, 1925. Pandaka pruinosa (Jordan and Seale, 1906). Paragobiodon echinocephalus (Riippell, 1830): Giinther, 1877. Paragobiodon lacunicolus Kendall and Goldsborough, 1911. Paragobiodon melanosomus (Bleeker, 1852). Paragobiodon modestus (Regan, 1908). Paragobiodon xanthosomus (Bleeker, 1852). Pleurosicya bilobatus (Koumans, 1941). Pleurosicya muscarum (Jordan and Seale, 1906). Priolepis farcimen (Jordan and Evermann, 1903). Silhouettea sp. Stiphodon elegans (Steindachner, 1879). Trimma caesiura Jordan and Seale, 1906. Trimma eviotops Schultz, 1943. Trimma naudei Smith, 1956. Trimma okinawae (Aoyagi, 1949). Trimma spp. (6 species). Valenciennea puellaris Tomiyama, 1956. Valenciennea sexguttatus (Valenciennes, 1837). Valenciennea strigatus (Broussonet, 1782). Valenciennea sp. Vanderhorstia ambanoro (Fourmanoir, 1957). Vanderhorstia spp. (2 species). Xenisthmus chapmani (Schultz, 1966). Kraemericus chapmani Schultz, 1966. Victor G. Springer (personal communication) advised us that K. chapmani is not a kraemeriid but a goby of the genus Xenisthmus. Xenisthmus sp. Family ELEOTRIDIDAE Ophieleotris aporos (Bleeker, 1854). Family KRAEMERIIDAE Kraemeria bryani Schultz, 1941: Schultz, 1966. Kraemeria samoensis Steindachner, 1906: Schultz, 1966. Family MICRODESMIDAE Gunnellichthys monostigma Smith, 1958: Strasburg, 1967. Gunnellichthys pleurotaenia Bleeker, 1858: Schultz, 1966. Paragobioides grandoculis Kendall and Goldsborough, 1911. Nemateleotris helfrichi Randall and Allen, 1973. Nemateleotris magnifica Fowler, 1938: Randall and Allen, 1973. Ptereleotris evides (Jordan and Hubbs, 1925): Randall and Hoese, 1985. Ptereleotris hanae (Jordan and Snyder, 1901): Randall and Hoese, 1985. Ptereleotris heteroptera (Bleeker, 1855): Randall and Hoese, 1985. (This table continued on next page.) 310 RANDALL AND RANDALL TABLE 1 (cont’d) Family MICRODESMIDAE (cont'd) Ptereleotris microlepis (Bleeker, 1856): Randall and Hoese, 1985. Ptereleotris zebra (Fowler, 1938): Randall and Hoese, 1985. Family ACANTHURIDAE Acanthurus achilles Shaw, 1803: Schultz and Woods, 1953. Acanthurus blochii Valenciennes, 1835. Acanthurus mata Schultz and Woods, 1953. Randall (MS) has shown that many authors have been mistaken in applying the name A. mata to this fish. It is the earliest name for the species that most authors have identified as A. bleekeri Ginther, 1861. Acanthurus guttatus Bloch and Schneider, 1801: Schultz and Woods, 1953. Acanthurus lineatus (Linnaeus, 1758): Schultz and Woods, 1953. Acanthurus maculiceps (Ahl, 1923): Randall, 1986. Acanthurus mata (Cuvier, 1829). Acanthurus bleekeri Schultz and Woods, 1953. See discussion under A. blochii. Acanthurus nigricans (Linnaeus, 1758). Acanthurus aliala Woods and Schultz, 1953. Randall (1956) placed A. aliala Lesson in the synonymy of A. glaucopareius Cuvier; both, however, are synonyms of A. nigricans (Linnaeus) (Randall, MS). Acanthurus nigricauda Duncker and Mohr, 1929. Acanthurus nigricans Schultz and Woods, 1953. Acanthurus nigricans (Linnaeus) is the species most authors have identified as A. glaucopareius. Acanthurus gahhm Randall, 1956. Acanthurus gahhm (Forsskal) is an endemic Red Sea species. Acanthurus nigrofuscus (Forsskal, 1775). Acanthurus elongatus Schultz and Woods, 1953 (in part). Randall (1956) indicated that nigrofuscus is the oldest name for this species; A. elongatus (Lacepede) is not the same fish. Acanthurus nigroris Valenciennes, 1835. Acanthurus elongatus Schultz and Woods, 1953 (in part). Acanthurus olivaceus Bloch and Schneider, 1801: Schultz and Woods, 1953. Acanthurus pyroferus Kittlitz, 1834. Acanthurus leucosternon Schultz and Woods, 1953. Randall (1956) showed that this is a misidentification; A. leucosternon Bennett is an Indian Ocean species. Acanthurus thompsoni (Fowler, 1923). Acanthurus philippinus Schultz and Woods, 1953. Randall (1956) referred A. philippinus Herre to the synonymy of A. thompsoni. Acanthurus triostegus triostegus (Linnaeus, 1758): Woods and Schultz, 1953. Acanthurus xanthopterus Valenciennes, 1835: Randall, 1980a. Ctenochaetus binotatus Randall, 1955b: Randall, 1986. Ctenochaetus hawaiiensis Randall, 1955b: Randall, 1986. Ctenochaetus marginatus (Valenciennes, 1835). Ctenochaetus sp. Hiyama, 1943. Ctenochaetus cyanoguttatus Randall, 1955b. Ctenochaetus striatus (Quoy and Gaimard, 1825): Schultz and Woods, 1953. Ctenochaetus strigosus (Bennett, 1828): Randall, 1986. Naso annulatus (Quoy and Gaimard, 1825): Schultz and Woods, 1953. Naso brevirostris (Valenciennes, 1835): Schultz and Woods, 1953. Naso hexacanthus (Bleeker, 1855): Schultz and Woods, 1953. Naso lituratus (Bloch and Schneider, 1801): Schultz and Woods, 1953. Naso tuberosus Lacepede, 1801: Randall, 1986. Naso unicornis (Forsskal, 1775): Schultz and Woods, 1953. Naso vlamingii (Valenciennes, 1835): Schultz and Woods, 1953. Paracanthurus hepatus (Linnaeus, 1766): Randall, 1981d. Zebrasoma flavescens (Bennett, 1828): Schultz and Woods, 1953 (in part). Zebrasoma scopas (Cuvier, 1829). Zebrasoma flavescens Schultz and Woods, 1953 (in part). Randall (1955c) showed that Z. flavescens and Z. scopas are distinct species. Zebrasoma veliferum (Bloch, 1795): Schultz and Woods, 1953. Family ZANCLIDAE Zanclus cornutus (Linnaeus, 1758): Woods, 1953. Zanclus canescens Woods, 1953. Randall (1955a) demonstrated that Z. canescens (Linnaeus) is the late postlarval stage of Z. cornutus. The latter has priority due to decision of the first revisor, Cuvier, 1831. (This table continued on next page.) ANNOTATED CHECKLIST OF FISHES 311 TABLE 1 (cont'd) Family SIGANIDAE Siganus argenteus (Quoy and Gaimard, 1825): Woods, 1953. Siganus rostratus Woods, 1953. Woodland (in press) has shown that argenteus, named for the late postlarval stage of this species, has priority over S. rostratus (Valenciennes). Siganus puellus (Schlegel, 1852): Randall, 1986. Siganus punctatus (Bloch and Schneider, 1801): Woods, 1953. Siganus vulpinus (Schlegel and Miiller, 1844): Woods, 1953. Family SCOMBRIDAE Acanthocybium solandri (Cuvier, 1831): Schultz, 1960. Euthynnus affinis (Cantor, 1849). Euthynnus affinis yaito Schultz, 1960. Grammatorcynus bilineatus (Riippell, 1836): Schultz, 1960. Gymnosarda unicolor (Riippell, 1836). Gymnosarda nuda Schultz, 1960. Collette and Gibbs (1963) and Collette and Chao (1975) are followed in using the name G. unicolor for the Dogtooth Tuna. Lewis (1968) identified parasitic copepods from a scombrid at Enewetak he identified as Sarda orientalis. This species is not known from the Marshall Islands. From the parasites found he probably had Gymnosarda unicolor (Bruce B. Collette and Roger F. Cressey, personal commmunication). Katsuwonus pelamis (Linnaeus, 1758): Schultz, 1960. Thunnus albacares (Bonnaterre, 1788). Neothunnus albacora macropterus Schultz, 1960. Collette and Gibbs (1963) and Gibbs and Collette (1967) stated that Neothunnus macropterus (Temminck and Schlegel) is a synonym of T. albacares. Family NOMEIDAE Psenes sp. Schultz, 1960. Family BOTHIDAE Arnoglossus intermedius (Bleeker, 1866): Woods, 1966. Bothus mancus (Broussonet, 1782): Woods, 1966. Bothus pantherinus (Riippell, 1830): Woods, 1966. Order PLEURONECTIFORMES Family PLEURONECTIDAE Samariscus triocellatus Woods, 1966. Family SOLEIDAE Aesopia heterorhinos (Bleeker, 1856): Woods, 1966. Aseraggodes melanostictus (Peters, 1876): Woods, 1966. Aseraggodes smithi Woods, 1966. Aseraggodes whitakeri Woods, 1966. Order TETRAODONTIFORMES Family BALISTIDAE Balistapus undulatus (Mungo Park, 1797): Woods, 1966. Balistoides conspicillum (Bloch and Schneider, 1801). Balistes niger Kendall and Goldsborough, 1911. Balistoides niger Woods, 1966. Randall and Klausewitz (1973) showed that Balistes niger Bonnaterre is a homonym; therefore, B. conspicillum becomes the valid name. Balistoides viridescens (Bloch and Schneider, 1801): Woods, 1966. Canthidermis senticosus (Richardson, 1848): Woods, 1966. Melichthys niger (Bloch, 1786): Randall and Klausewitz, 1973. Melichthys vidua (Solander, 1844): Woods, 1966. Odonus niger (Riippell, 1837): Hiyama, 1943; Randall, 1986 (after Strasburg). Pseudobalistes flavimarginatus (Riippell, 1829): Randall, 1986 (after Strasburg). Balistes flavimarginatus Hiyama, 1943. Pseudobalistes fuscus (Bloch and Schneider, 1801): Woods, 1966. Rhinecanthus aculeatus (Linnaeus, 1758): Woods, 1966. Rhinecanthus rectangulus (Bloch and Schneider, 1801): Woods, 1966. Sufflamen bursa (Bloch and Schneider, 1801): Woods, 1966. Sufflamen chrysoptera (Bloch and Schneider, 1801): Woods, 1966. Sufflamen fraenatus (Latreille, 1804). Sufflamen capistratus Woods, 1966. Sufflamen fraenatus is the oldest name for this species (Frederick H. Berry, personal communication). Xanthichthys auromarginatus (Bennett, 1831): Randall, Matsuura, and Zama, 1978. Xanthichthys caeruleolineatus Randall, Matsuura, and Zama, 1978: Randall, 1986. (This table continued on next page.) 312 RANDALL AND RANDALL TABLE 1 (cont'd) Family MONACANTHIDAE Aluterus scriptus (Osbeck, 1765). Alutera scripta Woods, 1966. Amanses scopas (Cuvier, 1829): Woods, 1966. Brachaluteres taylori Woods, 1966. Cantherhines dumerilii (Hollard, 1854). Amanses carolae Woods, 1966. Randall (1964b) determined that Cantherhines carolae (Jordan and McGregor) is a junior synonym of C. dumerilii. Cantherhines fronticinctus (Playfair and Giinther, 1867): Randall, 1986. Cantherhines pardalis (Rtippell, 1837). Amanses sandwichiensis Woods, 1966. Randall (1964b) showed that C. sandwichiensis (Quoy and Gaimard) is endemic to the Hawaiian Islands. The correct name for the wide-ranging species from elsewhere in the Indo-Pacific is C. pardalis. Oxymonacanthus longirostris (Bloch and Schneider, 1801): Woods, 1966. Paraluteres prionurus (Bleeker, 1851): Woods, 1966. Paramonacanthus cryptodon (Bleeker, 1855): Woods, 1966. Paramonacanthus oblongus (Temminck and Schlegel, 1846): Woods, 1966. Pervagor alternans (Ogilby, 1899). Pervagor melanocephalus marshallensis Woods, 1966. Synonymy from Hutchins, 1986. Pervagor aspricaudatus Hollard, 1854. Pervagor melanocephalus johnstonensis Woods, 1966. Pervagor melanocephalus (Bleeker, 1853). This record based on one specimen reported by Hutchins, 1986. Family OSTRACIIDAE Lactoria fornasini (Bianconi, 1846): Woods, 1966. Ostracion cubicus Linnaeus, 1758: Woods, 1966. Ostracion meleagris Shaw, 1796: Woods, 1966. Family TETRAODONTIDAE Amblyrhynchotes honckenii (Bloch, 1785): Randall, 1986 (after Strasburg). Arothron hispidus (Linnaeus, 1758): Woods and Schultz, 1966. Arothron manilensis (Procé, 1822): Randall (1985) has shown that the strongly striped A. manilensis is a distinct species from the related unmarked A. immaculatus (Bloch and Schneider). Arothron mappa (Lesson, 1826): Randall, 1986 (after Strasburg). Arothron meleagris (Lacepede, 1798): Woods and Schultz, 1966. Arothron nigropunctatus (Bloch and Schneider, 1801): Woods and Schultz, 1966. Arothron stellatus (Bloch and Schneider, 1801). Arothron alboreticulatus Woods and Schultz, 1966. Canthigaster amboinensis (Bleeker, 1865): Woods, 1966. Canthigaster bennetti (Bleeker, 1854): Randall, 1986. Canthigaster coronata (Vaillant and Sauvage, 1875): Randall, 1986. Canthigaster epilampra (Jenkins, 1903): Randall, 1986. Canthigaster janthinoptera (Bleeker, 1855). Canthigaster jactator Woods. Allen and Randall (1977) regarded C. jactator (Jenkins) as a Hawaiian endemic. The related widespread species from elsewhere in the Indo-Pacific region is C. janthinoptera. Canthigaster solandri (Richardson, 1844): Woods, 1966. Canthigaster valentini (Bleeker, 1953): Randall, 1986. Family DIODONTIDAE Diodon hystrix Linnaeus, 1758: Woods, 1966. Diodon liturosus Shaw, 1804: Randall, 1986. this list. Only a few such fishes have been reported from the area. Schultz in Schultz et al. (1953), for example, listed only two myctophids, Myctophum brachygnathos (Bleeker) and Diaphus schmidti Taning, both by name only. Of the Gonostomatidae, Grey (1960) recorded only Diplophos sp. (taenia complex), Gonostoma atlanticum Nor- man, and G. ebelingi Grey from the Marshalls region. R. K. Johnson recently identified a giganturid (BPBM 26320, 62.3 mm SL) taken with a midwater trawl at 11°20’N, 162°07’E at a depth of 0 to 200 m as Rosaura indica (Brauer). There has not been adequate sampling of deep-water fishes from the Marshall Islands or, for that matter, from the islands of Oceania in general (except the Hawaiian Islands). Even species of moderate depths (100 to 300 m), such as snappers of the genera Etelis and Pristipomoides have not been reported from the Marshalls, though some surely occur there. If complete collections were made of mesopelagic, bathypelagic, and deep benthic fishes, these would have to be treated differently from a zoogeographic standpoint than shore fishes. In his excellent review of the biogeography of the Pacific Plate with emphasis on fishes, Springer (1982) discussed shore fishes (those from depths less than 100 m), not deeper water species (except some from the Hawaiian Islands). | We solicited from Ernest A. Lachner and Christine Baer a preliminary list of the Gobiidae from the Marshall Islands to include with the present checklist. We have added a few species to their list from Bishop Museum specimens, and Douglass F. Hoese has added some from a collection of about 150 lots of gobies sent him by Ronald S. Nolan, who conducted his doctoral thesis research on reef fishes at Enewetak. Hoese and E. O. Murdy also advised on some gobiid name changes. Ran- dall and Hoese (1985) have removed the gobioid genera Nemateleotris and Ptereleotris from the family Gobiidae and provisionally placed them in the subfamily Ptereleo- trinae of the family Microdesmidae. Although we have endeavored to make the present checklist as current as possible, we wish to emphasize that it is far from definitive. Species of shore fishes surely remain to be collected from the Marshall Islands (though we believe we have more than the 95% level in this respect). The estimated 45 new species which have been collected need to be described. Most important, many of the taxonomic groups of Indo-Pacific fishes need to be revised. Inevitably, such studies will result in changes in scientific names now in widespread use. Among those groups most in need of revision are the Mobulidae; Dasya- tididae; all of the eel families; the Bythitidae; the scor- paenid genera Scorpaena (sensu lato), Scorpaenodes, and Scorpaenopsis; the serranid genera Anthias, Cephalopholis, Epinephelus, Plectropomus, and Pseudogramma; Pseudo- chromidae; Kuhliidae; Priacanthidae; Apogonidae; the carangid genera Carangoides, Decapterus, and Trachinotus; the Caesionidae; the haemulid genus Plectorhinchus; the lethrinid genera Gymnocranius and Lethrinus; the Nemi- pteridae; Gerreidae; Mullidae; Pempherididae; Kyphosidae; the pomacentrid genera Chromis, Chrysiptera, and Pomacentrus; the Mugilidae; Sphyraenidae; Polynemidae; the labrid genera Cheilinus, Cirrhilabrus, _ Coris, Halichoeres, Pseudocheilinus, Pteragogus, and Xyrichtys; the scarid genus Scarus; the blenniid genera Cirripectes, Istiblennius, Praealticus, Rhabdoblennius, and Salarias, the Tripterygiidae; the callionymid genera Callionymus and Diplogrammus; the Gobiidae; the acanthurid genus Naso; the soleid genus Aseraggodes; and the tetraodontid genus Arothron. Fortunately, many of these groups are under study, and clarification of their taxonomy can be expected reasonably soon. The present checklist is tied closely to the work of Schultz et al. Anyone working on Marshall Islands fishes will surely begin with these three volumes. This list will, therefore, serve as a supplement to provide additions and name changes. We have cited all the fish names of Schultz et al. When these names have been changed, they are ANNOTATED CHECKLIST OF FISHES 313 listed as synonyms below the correct name and are indented three spaces. When needed, an annotation is pro- vided to explain the name change and give the authority. Only those synonyms are listed that were used as valid names in Schultz et al. The author given for each of these synonyms is the one responsible for the Marshall Islands record, not the author of the species (unless named as new in Schultz et al. or in other papers on the Marshall Islands fish fauna). We have eliminated the listing of Schultz et al. each time Schultz or one of his collaborators is cited in the checklist. Thus Schultz, 1953, implies Schultz in Schultz et al., 1953. Similarly, we have shortened Cuvier (or Valenci- ennes) in Cuvier and Valenciennes to just Cuvier (or Valen- ciennes). Subgeneric categories have not been included in the checklist. The order of presentation is phylogenetic, approximat- ing that of Greenwood et al. (1966). We have not made a documentation of the different atolls at which the various species of fishes have been col- lected. Such listing is provided by Schultz et al. and by Randall (1986). Little difference in the fish fauna can be expected from one atoll to the next within the Marshall Islands, particularly those of about the same latitude. There are differences, however, from the northern to the southern Marshalls. More species of fishes are presently known from the northern atolls because these have been more heavily collected. This is especially true of Bikini and Enewetak. Bikini is the atoll best represented in the material at the National Museum of Natural History, but the extensive Bishop Museum collections made by the senior author and associates over the last 16 years at Enewetak have made it the locality with the most species. We believe, however, that the southern atolls would have a slightly richer fish fauna if the collecting effort were equal. A number of conspicuous species that have been found at Kwajalein or other southern atolls in the Marshalls have not been observed at Enewetak. Examples are Myripristis adustus, Sargocentron rubrum, Anthias bartlettorum, Epinephelus caeruleopunctatus, Plectropomus oligacanthus, Carangoides plagiotaenia, Pterocaesio sp., Scolopsis lineatus, Heniochus varius, Halichoeres richmondi, Scarus bleekeri, Amblygobius decussatus, Acanthurus maculiceps, Siganus vulpinus, and Balistoides conspicillum. This, of course, is not to say that they are truly absent from Enewetak. How does the fish fauna of the Marshall Islands com- pare with the rest of the Indo-West-Pacific region? Of the estimated total of 20,000 species of fishes in the world, about 8000 are shallow, warm-water marine species (Cohen, 1970). The majority of these warm-water fishes occurs in the vast Indo-West-Pacific region. Using two dif- ferent methods to arrive at an estimate, Springer (1982) concluded that there are about 4000 species of fishes in the tropical Indo-West-Pacific region. Cohen (1973), how- ever, estimated 3000 to 4000 for the Indian Ocean alone. If Cohen is correct, Springer admitted his estimate for the 314 RANDALL AND RANDALL Indo-West-Pacific as a whole is much too low. By analyzing the 111 shorefish families which occur nonmarginally on the Pacific Plate, Springer estimated that there are 461 genera and an estimated 1312 species. Thus the Marshall Islands, with 817 species, have 62% of the total Pacific Plate fish fauna. Proper comparisons between island groups of the Pacific are possible only when there has been a compa- rable collecting effort as well as an accurate compilation of species. This would seem to be the case only for the Hawaiian Islands, Society Islands, Samoa Islands, southern Mariana Islands, and Easter Island. Randall (1981a) stated that approximately 610 species of fishes occur in the Hawaiian Archipelago. By excluding those which are pelagic, deep sea, or freshwater species, he arrived at 420 which could be regarded as reef and shore fishes. This figure should be modified to 460 to include epipelagic species, recent new records, and new species (to make it equivalent to estimates herein from other island groups). The extreme isolation of the Hawaiian Islands, both geographically and hydrographi- cally, accounts for its somewhat impoverished fish fauna. Randall (1973) provided a preliminary checklist of 616 fishes from the Society Islands. By eliminating those which occur in deep water or fresh water but adding new records, the list is revised to 620. Wass (1984) has prepared a checklist of fishes of the Samoa Islands. His total for the list is 958, which includes 84 identified only to genus. If those which occur in deep water and in fresh water are eliminated, along with dubi- ous literature records, the total is lowered to 915. The Samoa Islands lie marginally on the Pacific Plate. There is a continental component to the fish fauna of Samoa due to its proximity to the continental plate. These islands are mostly high islands with a greater variety of habitats than one finds on atolls. Furthermore, they are less isolated from other island groups than are the Marshalls. There- fore, a greater number of species of fishes should be expected from Samoa. Shepard and Myers (1981) published a preliminary checklist of the fishes of Guam and other southern Mariana Islands. Their total is 801, from which 12 should be sub- tracted due to occurrence in deep water or fresh water. Myers (MS submitted to Micronesica) has raised the num- ber of fishes for the Marianas to 854 (not including fresh- water and deep-sea species). The Marianas are also high islands which lie on the margin of the Pacific Plate. Randall and Cea Egana (1984) made a checklist of the fishes of Easter Island. Only 130 shore and epipelagic species have been found at this small, remote South Pacific island. Recent collecting by Randall and colleagues has raised the total to 155 species. Springer (1982) has provided an analysis of endemic fishes of the Pacific Plate. He placed them in three categories. Type 1 endemics are those that are widely dis- tributed on the Plate, Type 2 are those that are limited to a few islands or island groups, and Type 3 are those that are found only at a single island or island group. The Marshall Islands have 28 of the 48 Type 1 Plate endemics tabulated by Springer. Of the Type 2 endemics, Springer listed only Pomachromis exilis and Labropsis micronesica from the Marshalls. Type 3 endemics (the most common type) are found mainly at high islands. Springer mentioned only three from the Marshalls, Acanthoplesiops _hiatti, Amphiprion tricinctus, and Cirricaecula johnsoni. To these one might add Hypoatherina barnesi, Pseudochromis aurea marshallensis, and Aseraggodes smithi. In the Pacific, the Hawaiian Archipelago has the highest percentage of endemism among its shore fishes, with 30%; Easter Island is next with 27.3%; Lord Howe TABLE 2 Summary of Nonsystematic Fish Research at Enewetak Type of study Fish taxa* Reference Ecology and Behavior Acoustics and Holocentrus spp., Myripristis spp. environment Carcharhinidae Artificial reefs Reef fishes Coral reef Reef fishes community Reef fishes Chaetodontidae, Pomacentridae Acanthurus guttatus, A. triostegus, Scarus jonesi Acanthuridae and Scaridae Reef fishes Fauna of nuclear test crater Horch and Salmon, 1973 Nelson and Johnson, 1972 Nolan, 1974 Hiatt and Strasburg, 1960 Odum and Odum, 1955 Reese, 1973 Webb and Wiebe, 1975 Weibe et al., 1975 Nolan et al., 1955 ‘The scientific names given in the table are those that appeared in the publication cited. Some of these names have been changed (see checklist herein). (This table continued on next page.) ANNOTATED CHECKLIST OF FISHES TABLE 2 (cont’d) Type of study 315 Fish taxa* Reference Feeding and food habits Reproduction Shark behavior and biology Social behavior Symbiosis Territoriality and home range Zooplankton composition Cell structure Phosphorus exchange Senses Tooth development Venom Ecology and Behavior Reef fishes Chromis caeruleus; plankton-feeding fishes Reef fishes Carcharhinus melanopterus, C. menisorrah, Triaenodon obesus Reef fishes Chaetodontidae Acanthurus triostegus, Holocentrus spinifer, Scarus gibbus, S. jonesi Chromis spp. Crenimugil crenilabis Chromis caeruleus Carcharhinus melanopterus, C. menisorrah, Triaenodon obesus Carcharhinus menisorrah Carcharhinus amblyrhynchos Triaenodon obesus Carcharhinus melanopterus Acanthuridae Pomacentrus jenkensi Siphamia fuscolineata Reef and pelagic fishes parasitized by copepods (for listing see Devaney, Chapter 19, this volume) Abudefduf abdominalis, Aspidontus filamentosus, A. taeniatus, Ecsenius bicolor, Labroides spp. and host fishes, Meiacanthus atrodorsalis, Runula laudandus Acanthurus spp., Amblygobius phalaena, Cryptocentroides maculosus, Ptereleotris microlepis, Valenciennea puellaris, V. sexguttatus, V. strigatus, Zebrasoma spp. Chromis spp. Chaetodontidae Chromis spp. Fish eggs, Monocanthus and other fish larvae Fish eggs Physiology and Histology Carcharhinus melanopterus, Negaprion acutidens Carcharhinus spp., rays Carcharhinus melanopterus Reef fishes Carcharhinus melanopterus Reef fishes Carcharhinus melanopterus, C. menisorrah Carcharhinus menisorrah, Triaenodon obesus Carcharhinus menisorrah Synanceja verrucosa Pterois volitans, Synanceja horrida, S. verrucosa Bakus, 1967 Gerber and Marshall, 1974a; 1974b Hiatt and Strasburg, 1960 Hobson, 1963 Hobson and Chess, 1978 Reese, 1977 Smith and Paulson, 1974 Swerdloff, 1970 Helfrich and Allen, 1975 Swerdloff, 1970 Hobson et al., 1961 Johnson and Nelson, 1973 Nelson, 1982 Randall, 1977 Randall and Helfman, 1973 Barlow, 1974a; 1974b Allen, 1972b Lewis, 1968 Losey, 1971; 1972; 1974; 1975 Paulson, 1978 Swerdloff, 1970 Reese, 1973; 1978 Swerdloff, 1970 Gilmartin, 1958 Johannes and Gerber, 1974 Bullock and Corwin, 1979 Corwin, 1977a; 1974b; 1978 Saland et al., 1974 Pomeroy and Kuenzler, 1967 Fay et al., 1974 Munz and McFarland, 1973 Tester, 1963 Kemp, 1974a, 1974b Kemp and Park, 1974 Saunders, 1959 Saunders and Taylor, 1959 316 RANDALL AND RANDALL Island, Middleton and Elizabeth Reefs, and Norfolk Island combined have about 12% endemism (Randall, 1976). Randall (1978a) made an approximation of 10% endemism for the fishes of the Marquesas. For the Marshall Islands the Type 3 endemism, based on species which have been named, is less than 1%. This figure will rise slightly when some new species currently known only from the Marshalls are described. However, when there has been a compa- rable collecting effort in the Central Pacific, and in particu- lar the Gilbert Islands and eastern Caroline Islands (which lie near the Marshalls and include atolls), it is expected that most of the species now regarded as endemic to the Marshall Islands will be found elsewhere. In addition to publications on the classification of fishes from the atoll, there has been a variety of nonsystematic studies dealing with the Enewetak ichthyofauna. Table 2 summarizes these and gives the references and taxa involved. The three plates (Figs. 1, 2, and 3) of underwater pho- tographs of fishes herein were taken by the senior author in the Marshall Islands. Plates 1-3 Fig. 1 a, The gray reef shark, Carcharhinus amblyrhynchos (Bleeker), one of the three most abundant sharks at Enewetak; responsible for four attacks on man at the atoll; b, Gymnothorax javanicus (Bleeker), the largest moray of the Indo-Pacific region; attains a length of at least 2 m; c, Poeciloconger fasciatus Ginther; known from only five specimens. This one emerged from the sand from the effect of ichthyocide at Enewetak; d, The squirrelfish Sargocentron microstoma (Ginther). Photograph taken at night; e, The soldierfish Myripristis berndti Jordan and Evermann being “cleaned” by the wrasse Labroides pectoralis Randall and Springer; f, The grouper Epinephelus microdon (Bleeker) being cleaned by the wrasse Labroides dimidiatus (Valenci- ennes); g, The bright violet and yellow Anthias bartlettorum Randall and Lubbock, named in honor of Nathan and Patricia Bartlett who discovered it at Kwajalein; h, The red snapper Lutjanus gibbus (Forsskal). Photograph taken at night. _—» Page 317 Fig. 2 a, A school of the fusilier Casio caerulaureus Lacepede from the Kwajalein lagoon. This photo was the basis for the first Marshall Islands record; b, The nocturnal lethrinid fish Monotaxis grandoculis (Forsskal); has well- developed molariform for crushing mollusks, echinoids, etc; c, The black jack, Caranx lugubris Poey. Circumtropical in distribution; d, The goatfish Parupeneus barberinoides (Bleeker). Recent collections provided the first record for the Marshall Islands; e, The colorful damselfish Pomacentrus pavo (Bloch). Blue in life with yellow on the caudal fin; f, The anemonefish Amphiprion chrysopterus Cuvier. Like others of the genus, it lives symbiotically with sea anemones; g, The wrasse Labropsis micronesica Randall. Enewetak is the type- locality. Adults feed on coral polyps; h, The terminal male of wrasse Halichoeres trimaculatus (Quoy and Gaimard). A com- mon lagoon species in sand and rubble areas around coral heads. —_ Page 318 Fig. 3 a, A group of the butterflyfish Heniochus chryso- stomus Cuvier. Usually seen in pairs or small aggregations; b, The yellow angelfish Centropyge heraldi Woods and Schultz; named for the late Earl S. Herald, who collected most of the type specimens in the Marshall Islands in 1946; c, The goby Amblyeleotris guttata (Fowler) and its symbiotic partner Alpheus ochrostiatus Miya. The shrimp is just removing a load of sand and gravel from the burrow; d, The colorful goby Nemateleotris helfrichi Randall and Allen, named for Philip Helfrich, the last director of the Mid-Pacific Research Laboratory at Enewetak; e, The surgeonfish Acanthurus nigri- cans Linnaeus, distinctive in the white spot under the eye and yellow band at the base of the dorsal and anal fins; f, The filefish Oxymonacanthus longirostris (Bloch and Schneider); blue-green with orange spots. Feeds on coral polyps; g, The female of the triggerfish Xanthichthys auromarginatus (Ben- nett); generally found at depths greater than 30 m; h, The clown triggerfish Balistoides conspicillum (Bloch and Schneider). Often identified as B. niger (Bonnaterre), but this name is a homonym, hence invalid. — Page 319 ANNOTATED CHECKLIST OF FISHES 317 Ree I. eae ee seeps CAGE seainosratrrgiae se RANDALL AND RANDALL 318 Fig. 2 319 ANNOTATED CHECKLIST OF FISHES : 5 i i Fig. 3 320 RANDALL AND RANDALL ACKNOWLEDGMENTS We thank the following persons for assisting us in obtaining the correct scientific names for the fishes in this checklist: Gerald R. Allen, Kent E. Carpenter, P. H. J. Castle, Bruce B. Collette, Charles E. Dawson, William N. Eschmeyer, Thomas H. Fraser, Ronald Fricke, Ronald A. Fritzsche, Douglass F. Hoese, Walter Ivantsoff, Susan Jewett, Robert Karl Johnson, Leslie W. Knapp, Ernest A. Lachner, Keiichi Matsuura, John E. McCosker, Edward O. Murdy, Theodore W. Pietsch, William F. Smith-Vaniz, Vic- tor G. Springer, James M. Thomson, Richard C. Wass, Jef- frey T. Williams, and David J. Woodland. The manuscript was reviewed by Lori J. Bell, Patrick L. Colin, Victor G. Springer, and Richard C. Wass. REFERENCES Allen, G. R., 1972a, The Anemonefishes, T. F. H. Publications, Neptune City, N. J. ___, 1972b, Observations on a Commensal Relationship Between Siphamia fuscolineata (Apogonidae) and the Crown-of-Thorns Starfish, Acanthaster planci, Copeia, 3: 595-597. __., 1975, Damselfishes of the South Seas, T. F. H. Publica- tions, Neptune City, N. J. __., M. L. Bauchot, and M. Desoutter, 1978, The Status of Abudefduf sexfasciatus (Lacepede), a Pomacentrid Fish from the Indo-West Pacific, Copeia, 2: 328-330. ——, and W. N. Eschmeyer, 1973, Turkeyfishes at Eniwetok, Pac. Discovery, 26(3): 3-11. __., and J. E. Randall, 1977, Review of the Sharpnose Puffer- fishes (Subfamily Canthigasterinae) of the Indo-Pacific, Rec. Austral. Mus., 30: 475-517. Arnold, D. C., 1956, A Systematic Revision of the Fishes of the Teleost Family Carapidae (Percomorphi, Blennioidea), with Descriptions of Two New Species, Bull. Brit. Mus. Nat. Hist., Zool., 4: 247-307. Bakus, G. J., 1967, The Feeding Habits of Fishes and Primary Production at Eniwetok, Marshall Islands, Micronesica, 3: 135-149. Barlow, G. W., 1974a, Extraspecific Imposition of Social Group- ing Among Surgeonfishes (Pisces: Acanthuridae), J. Zool., Lond., 174: 333-340. ___, 1974b, Contrasts in Social Behavior Between Central Ameri- can Cichlid Fishes and Coral-Reef Surgeonfishes, Am. Zool., 14: 9-34. Bohlke, J. E., and J. E. Randall, 1981, Four New Garden Eels (Congridae, Heterocongrinae) from the Pacific and Indian Oceans, Bull. Mar. Sci., 31: 366-382. __, and D. G. Smith, 1968, A New Xenocongrid Eel from the Bahamas, with Notes on Other Species in the Family, Proc. Acad. Nat. Sci. Philadelphia, 120: 25-43. Bruce, R. W., and J. E. 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Springer, 1971, Synopsis of the Tribe Salariini, with Description of Five New Genera and Three New Species (Pisces: Blenniidae), Smithson. Contr. Zool., No. 73, pp. 1-72. —_, and J. C. Staiger, 1973, Comparative Revision of Scomberoides, Oligoplites, Parona, and Hypacanthus with Comments on the Phylogenetic Position of Campogramma (Pisces: Carangidae), Proc. Calif. Acad. Sci., 4th Ser., 39: 185-256. Springer, V. G., 1967, Revision of the Circumtropical Shorefish Genus Entomacrodus (Blenniidae: Salariinae), Proc. U. S. Nat. Mus., 122: 1-150. ___, 1972, Synopsis of the Tribe Omobranchini with Descriptions of Three New Genera and Two New Species (Pisces: Blenni- idae), Smithson. Contr. Zool., No. 130, pp. 1-31. __, 1982, Pacific Plate Biogeography, with Special Reference to Shorefishes, Smithson. Contr. Zool., No. 367, pp. iv + 182. __, and W. F. Smith-Vaniz, 1968, Systematics and Distribution of the Monotypic Indo-Pacific Blenniid Fish Genus Atrosalarias, Proc. U. S. Nat. Mus., 124: 1-12. Strasburg, D. W., 1953, Fishes of the Southern Marshall Islands, Mimeo Report to Office of Naval Research, U. S. Navy, pp. ii + 267. —_, 1966, Observations on the Ecology of Four Apogonid Fishes, Pac. Sci., 20: 338-341. , 1967, Gunnellichthys monostigma and Ecsenius bicolor, New Fish Records from the Marshall Islands, Copeia, 4: 839-841. __, and R. W. Hiatt, 1957, Sexual Dimorphism in the Labrid Fish Genus, Gomphosus, Pac. Sci., 11: 133-134. Swerdloff, S. N., 1970, Behavioral Observations on Eniwetok Damselfishes (Pomacentridae: Chromis) with Special Refer- ence to Spawning of Chromis caeruleus, Copeia, 2: 371-374. Tester, A. L., 1963, Olfaction, Gustation and the Common Chemical Sense in Sharks, Sharks and Survival, P. W. Gilbert (Ed.), D. C. Heath & Co., Boston, Ch. 8, pp. 255-282. __., and S. Kato, 1966, Visual Target Discrimination in Black- tip Sharks (Carcharhinus melanopterus) and Grey Sharks (C. menisorrah), Pac. Sci., 20: 461-471. Van Campen, W. G., 1950, Poisonous Fishes of the South Seas, Spec. Sci. Rep., U. S. Dept. Interior, Fish & Wildl. Serv., Fisheries, No. 25, pp. 1-221 (Translated from the Japanese Language). Wass, R. C., 1984, An Annotated Checklist of the Fishes of Samoa, NOAA Tech. Rep., Nat. Mar. Fisheries Serv., Spec. Rep. Fish., No. 781, v + 43. Webb, K. L., and W. J. Wiebe, 1975, Nitrification on a Coral Reef, Can. J. Microbiol., 21: 1427-1431. Whitehead, P. J. P., and W. Ivantsoff, 1983, Atherina lacunosa and the Fishes Described by J. R. Forster, Japan. Jour. Ichth., 29: 355-364. Wiebe, W. J., R. E. Johannes, and K. L. Webb, 1975, Nitrogen Fixation in a Coral Reef Community, Science, 188: 257-259. Woodland, D. J., Revision of the Rabbitfishes, Family Siganidae, Indo-Pacific Fishes, in press. Woods, L. P., and P. M. Sonoda, 1973, Family Holocentridae, Fishes of the Western North Atlantic, Part 6, Mem. Sears Found. Mar. Res., No. 1, pp. 331-386. Reptiles of Enewetak Atoll JANET O. LAMBERSON U. S. Environmental Protection Agency Hatfield Marine Science Center, Newport, Oregon 97365 Seven species of lizards and one species of blind snake comprise the known terrestrial herpetofauna of Enewetak Atoll, based on specimens in the U. S. National Museum of Natural History, the B. P. Bishop Museum, the Ameri- can Museum of Natural History, the Museum of Compara- tive Zoology, and the Mid-Pacific Research Laboratory col- lections (Table 1). Species found on nearby atolls may also be present on Enewetak; however, no additional species have been collected from there. No amphibia or sea snakes are known from Enewetak. None of the species of reptiles known from Enewetak is endemic to Micronesia. Presumably, Gehyra oceanica, Lipinia noctua, and Emoia cyanura originated in Papua New Guinea, and Hemiphyllodactylus typus and Lepido- dactylus lugubris evolved in southern Asia (Burt and Burt, 1932; Brown, 1957). All are now widely distributed among the islands of the Pacific (McCoy, 1980). Hemidactylus frenatus and Ramphotyphlops bramina are known from tropical Asia, India, and Africa and have been introduced to Central America and the islands of the Pacific and Indian Oceans, including Hawaii (Hunsaker, 1966; Hunsaker and Breese, 1967; Bustard, 1970; McKeown, 1978; McCoy, 1980). Varanus indicus probably originated in the Indo-Australian archipelago or Papua New Guinea and has been intentionally introduced to a number of the Mariana, Caroline, and Marshall Islands (Fisher, 1948). This species is extensively distributed on Pacific Islands, including the eastern part of the Indo-Australian archipelago, Papua New Guinea, northern Queensland, Bismarck archipelago, Solomons, Trobriands, Carolines, Marianas, and the Marshalls (R. Crombie, USNM, personal communication). Keys for identification of the reptiles of Enewetak, distribution records, notes on natural history, and excellent color illustrations may be found in booklets by McKeown (1978) and McCoy (1980). The most often seen lizard on Enewetak is the house gecko, Hemidactylus frenatus, which frequents areas of human occupation. It is common on or inside buildings, especially near lights, or on windows at night where it 325 Chapter 28 stalks insects attracted by the light. It is also found under scrap metal, plant debris, and driftwood or other debris on the beach. Hemidactylus frenatus is an aggressive species that may replace other resident species of geckos when it is introduced to a new area. In Hawaii it is reported to have replaced Hemidactylus garnoti (McKeown, 1978) and possi- bly also Lepidodactylus lugubris to some extent (Hunsaker and Breese, 1967). McKeown (1978) reports that L. lugubris, which also frequents areas of human habitation, is still quite abundant in Hawaii. On Enewetak Atoll, L. lugubris is common on Enjebi Islet but is infrequently col- lected relative to the collection of H. frenatus on Enewetak Islet (Table 1). Hemidactylus frenatus is highly vocal, and in encounters with other geckos, it frequently emits a series of five to six chirping call notes. If attacked, it emits a squeak. An extensive discussion of vocalization in this spe- cies is given by Marcellini (1974). The female H. frenatus lays two round white eggs in crevices, and the eggs or adults may be introduced to new areas as stowaways among materials or equipment moved among _ islands (McKeown, 1978). The mourning gecko, Lepidodactylus lugubris, is found on buildings; trees and bushes; among coconut, pandanus, and other plant debris in areas occupied by people; in open forest; and among beach driftwood. This species is parthenogenic (McKeown, 1978), and its highly adhesive eggs are easily transported among goods carried by boat. This gecko has been widely distributed around the Pacific and is known from several islets on Enewetak Atoll (Table 1). Its call has been compared to the sound of two pebbles being hit together (Marshall, 1951). It sometimes emits a squeak when attacked by another gecko or captured by a collector. The tree gecko, Hemiphyllodactylus typus, is repre- sented from Enewetak by only two specimens in the USNM collection. This gecko, however, is reported to have a quick escape reaction (Oliver and Shaw, 1953) and to be wary and agile (Hunsaker and Breese, 1967). Therefore, it may be more abundant on Enewetak than indicated by the frequency of collection. In Hawaii it is found in low den- sity in forested areas, rock and wood piles, occasionally on the sides of buildings and on tree trunks at night, and under the bases of coconut palm fronds. It is not generally 326 LAMBERSON TABLE 1 Checklist of Terrestrial Reptiles of Enewetak Atoll* Phylum CHORDATA Class REPTILIA Order SQUAMATA Suborder SAURIA (Lizards) Family GEKKONIDAE “gecko” BPBM-Bokombako Islet: Gehyra oceanica (Lesson) USNM—Biken Islet: USNM—Elugelab Islet: USNM—lkuren Islet: BPBM—Ananij Islet: BPBM—lkuren Islet: MPRL—Japtan Islet: Hemidactylus frenatus Dumeril and Bibron USNM—Enewetak Islet: USNM—Medren Islet: USNM—Runit Islet: BPBM—Enewetak Islet: BPBM— Japtan Islet: BPBM—Medren Islet: BPBM—Runit Islet: MPRL—Enewetak Islet: Hemiphyllodactylus typus Bleeker USNM—Alembel Islet: Lepidodactylus lugubris (Dumeril and Bibron) USNM—Alembel Islet: USNM—Biken Islet: USNM—Billae Islet: USNM—Boken (Irene) Islet: Dec. 30, 1950, under loose canvas, Y. Oshiro, 1 spec., 0695. May 25, 1946, J. P. E. Morrison, 2 spec., 124076-77; 1948, Draeger, 7 eggs, 128000; Aug. 21, 1976, in Pisonia forest at night, P. B. Lamberson, 1 spec., 205527; Aug. 21, 1976, E. Reese, 1 spec., 205528. Dec. 25, 1965, under loose bark of fallen coconut tree, K. J. Frogner, 1 spec., 4033; June 28, 1964, in chela of crab, Geograpsus crinipes (still alive), in coconut grove, C. Berry, 1 spec., 4028; Dec. 29, 1965, K. J. Frogner, 1 spec., 4034; Jan. 25, 1967, K. J. Frogner, 20 spec., 4273, 4279-97; July 15, 1967, on Pisonia at night, E. S. Reese, 1 spec., 4001. Jan. 11, 1976, J. Lamberson, 2 eggs; March 7, 1976, around buildings near lagoon, F. C. Rabelais, 1 spec. March 5, 1977, trailer park area, J. Lamberson, 1 spec., 205529; March 12, 1977, on lawn furniture, trailer park area, P. B. Lamberson and J. Lamberson, 2 spec., 205530-31; March 13, 1977, on lawn under Coccoloba tree, trailer park area, P. B. Lamberson, 1 spec., 205532; March 17, 1977, under piece of corrugated metal, J. Lamberson, 1 spec., 205533; Aug. 1964, W. B. Jackson, 3 spec., 197841; duly 24, 1966, W. B. Jackson, 1 spec., 197842; Sept. 1, 1968, L. N. Huber, 2 spec., 197874. dan. 22, 1967, K. J. Frogner, 1 spec., 4247; Feb. 2, 1967, K. J. Frogner, 1 spec., 4253; dune 22, 1964, K. J. Frogner, 2 spec., 4046-47; Aug. 8, 1964, K. J. Frogner, 2 spec., 4031 and 4078; Aug. 8, 1964, in 10 ft coconut trees at edge of lagoon beach, K. J. Frogner, 3 spec., 4048-50; Dec. 30, 1965, large crumpled tarp on seaward beach, K. J. Frogner, 1 spec., 4064; Dec. 30, 1965, K. J. Frogner, 27 spec., 4065-67, 4079-4100, 4200-01; Dec. 30, 1965, under pile of sheet tin on cement tent base, K. J. Frogner, 3 eggs, 6948; dan. 22, 1967, K. J. Frogner, 21 spec., 4202 and 4227-46; duly 5, 1964, in small pile of rusted sheet metal, K. J. Frogner, 1 spec., 4021; Aug. 29, 1964, in wood and tarp pile on seaward beach, K. J. Frogner, 1 spec., 4044; Aug. 29, 1964, in driftwood on seaward beach, K. J. Frogner, 1 spec., 4045. March 7, 1976, around occupied buildings, F. C. Rabelais and J. Lamberson, 2 spec. April 17, 1977, in “roof,” P. B. Lamberson and J. Lamberson, 2 spec., 205534-35. July 30, 1966, W. B. Jackson, 1 spec., 197844; May 25, 1946, J. P. E. Morrison, 2 spec., 124078-79; Aug. 1, 1966, W. B. Jackson, 1 spec., 197843; Aug. 24, 1968, L. N. Huber, 3 spec., 197876; Aug. 24, 1968, L. N. Huber, 2 spec. with 17 eggs, 197875; Feb. 28, 1977, on bunker, J. Lamberson, 3 spec., 205538-40; ‘With collection data for specimens recorded in the collections of the U. S. National Museum of Natural History (USNM), the B. P. Bishop Museum (BPBM), the American Museum of Natural History (AMNH), and the Mid-Pacific Research Laboratory (MPRL). (This table continued on next page.) REPTILES 327 TABLE 1 (cont’d) Family GEKKONIDAE (cont’d.) USNM—Enewetak Islet: Enjebi Islet: USNM— _Japtan Islet: BPBM—Biken Islet: BPBM—Enewetak Islet: BPBM—lkuren Islet: BPBM—Medren Islet: AMNH—Runit Islet: MPRL—Biken Islet: Family SCINCIDAE Emoia cyanura (Lesson) USNM—Billae Islet: USNM—Japtan Islet: USNM—Lyjor Islet: USNM—Medren Islet: USNM—Mut Islet: BPBM—lkuren Islet: BPBM—Japtan Islet: BPBM—Medren Islet: AMNH—Runit Islet: MPRL—Medren Islet: Lipinia noctua (Lesson) USNM—lkuren Islet: USNM— _Japtan Islet: BPBM—lkuren Islet: BPBM—Japtan Islct: AMNH—Runit Islet: MPRL—Japtan Islet: March 12, 1977, on lawn furniture in trailer park area, P. B. Lamberson and J. Lamberson, 1 spec., 205536; March 12, 1977, on palm tree in trailer park area, J. Lamberson, 1 spec., 205537; Feb. 28, 1977, on LLL trailer, J. Lamberson, 2 spec., 205541—42; March 4, 1977, on LLL shack, P. Lamberson, 2 spec., 205543—-44; March 11, 1977, on rain gauge, 1 spec., 205545, on tide gauge, 4 spec., 205546-49, and on LLL shack, 3 spec., 205550-52, P. B. Lamberson and J. Lamberson; Aug. 3, 1965, W. B. Jackson, 1 spec., 197845. Aug. 1964, K. J. Frogner, 3 spec., 4022-24; Aug. 5, 1964, in tip of coconut “boat” in tree, K. J. Frogner, 1 spec., 4043; Jan. 22, 1967, K. J. Frogner, 5 spec., 4248-52; June 24, 1964, in driftwood along beach, K. J. Frogner, 2 spec., 4029-30; Dec. 29, 1965, K. J. Frogner, 2 spec., 4275-76; Jan. 25, 1967, K. J. Frogner, 4 spec., 4274, 4277-78, 4298; July 15, 1967, E. S. Reese, 1 spec., 4002; Aug. 8, 1964, under log just above seaward beach, K. J. Frogner, 1 spec., 4027; Dec. 30, 1965, large crumpled tarp on seaward beach, K. J. Frogner, 3 spec., 4061-63; Dec. 30, 1965, K. J. Frogner, 10 spec., 4068-77; Dec. 30, 1965, K. J. Frogner, 6 + eggs, 6949; Jan. 22, 1967, K. J. Frogner, 8 spec., 4219-26; Jan. 23, 1967, K. J. Frogner, 4 spec., 4215-18. 1 spec., 66570. March 11, 1976, in forest, J. Lamberson, 1 spec. Aug. 25, 1968, A. B. Amerson, Jr., 2 spec., 197918; June 6, 1946, J. P. E. Morrison, 1 spec., 124082; Aug. 7, 1966, W. B. Jackson, 1 spec., 197847; June 2, 1946, J. P. E. Morrison, 2 spec., 124080-81; Aug. 1, 1965, W. B. Jackson, 1 spec., 197848; May 28, 1946, J. P. E. Morrison, 5 spec., 124071-75. Dec. 29, 1965, K. J. Frogner, 5 spec., 4036-40; June 24, 1964, in coconut grove, K. J. Frogner, 2 spec., 4025-26; Dec. 28, 1965, K. J. Frogner, 1 spec., 4035; Jan. 25, 1965, K. J. Frogner, 2 spec., 4271-72; dan. 31, 1967, K. J. Frogner, 8 spec., 4009-16; Feb. 1, 1967, K. J. Frogner, 4 spec., 4261-64; Feb. 2, 1967, K. J. Frogner, 4 spec., 4257-60; Aug. 8, 1964, under board on sand on seaward beach, K. J. Frogner, 1 spec., 4051; Aug. 8, 1964, K. J. Frogner, 8 spec., 4052-59; Dec. 30, 1965, large crumpled tarp on seaward beach, K. J. Frogner, 1 spec., 4060; Jan. 23, 1967, K. J. Frogner, 12 spec., 4203-14. 4 spec., 66573-74, plus 2 untagged. March 29, 1977, inside redwood water tank, P. B. Lamberson, 1 spec. Aug. 1964, W. B. Jackson, 1 spec., 197850; Aug. 7, 1966, W. B. Jackson, 1 spec., 197849. June 24, 1964, in coconut grove in pile of fallen nuts at base of tree, K. J. Frogner, 1 spec., 4032; Dec. 29, 1965, K. J. Frogner, 2 spec., 4041-42; Jan. 25, 1967, K. J. Frogner, 3 spec., 4268-70; Jan. 31, 1967, K. J. Frogner, 10 spec., 4003-08, 4017-20; Feb. 1, 1967, K. J. Frogner, 3 spec., 4265-67; Feb. 2, 1967, K. J. Frogner, 3 spec., 4254-56. 6 spec., 66571-—72, plus 4 untagged. March 7, 1976, among coconut debris, F. C. Rabelais, 2 spec. (This table continued on next page.) 328 LAMBERSON TABLE 1 (cont'd) Family VARANIDAE Varanus indicus (Daudin) USNM—Japtan Islet: AMNH—4Japtan Islet: MPRL—Japtan Islet: Suborder SERPENTES (Snakes) Family TYPHLOPIDAE Ramphotyphlops bramina (Daudin) BPBM—Enewetak Islet: MCZ—Medren Islet: Order TESTUDINES (=CHELONIA) (Turtles) Suborder CRYPTODIRA Family CHELONIDAE Chelonia mydas (Linnaeus, 1758)—Green turtle. Eretmachelys imbricata (Linnaeus, 1766)—Hawksbill turtle. dune 7, 1947, J. P. E. Morrison, 2 spec., 124112-13. A. A. Vincenzi, 1 spec., 78994. Sept. 15, 1975, in open field, P. B. Lamberson and J. Lamberson, 1 spec. Sept. 13, 1980, MPRL lanai at night, S. Ige, 1 spec., 6953. June 1954, C. Okino and A. Souza, 1 spec., MCZ 53782. found in association with man (Hunsaker and Breese, 1967; Oliver and Shaw, 1953; McKeown, 1978). The female produces two eggs which adhere to surfaces (McKeown, 1978). The large Polynesian gecko, Gehyra oceanica, is found in the forest on the leaves or trunks of coconut, Pisonia, or Pandanus trees or under fallen logs or other debris on the ground (Heatwole, 1975). It can be spotted at night by the way its eyes shine red when touched by light. Its call is a loud “Kraaaaaaa” (Marshall, 1951). The blue-tailed skink, Emoia cyanura, is common under coconut or other plant debris, under driftwood, and among scrub vegetation at the top of island beaches, especially among the vines of the beach morning glory Ipomoea pes- caprae (Sachet, 1962). These skinks are reportedly poor climbers and are active primarily during the day (Moul, 1954; Marshall, 1951). The moth skink, Lipinia (Lygosoma or Leiolopisma) noctua, is fairly common on Enewetak Atoll among leaf litter and coconut debris. It is essentially diurnal and may be partly arboreal (McCoy, 1980). Oliver and Shaw (1953) reported that they had collected L. noctua on Runit Islet on Enewetak Atoll on the ground among “herbaceous vegetation along the beach dunes” and that specimens they collected had “two or more incomplete digits on the feet.” They speculated that the missing digits might have been sheared off by terrestrial hermit crabs which are com- mon on the islet or by other lizards during fights. Rats are also common on Runit and may have been responsible for some of the missing digits. Missing toes among specimens of L. noctua in Hawaii have been attributed to the lizards’ characteristic rolling escape behavior (McKeown, 1978). Lipinia noctua brings forth one to four young alive from eggs retained in the body (McKeown, 1978). The largest lizard on Enewetak Atoll is the monitor lizard, Varanus indicus, a native of Australia and Papua New Guinea which was introduced to Guam, the Mariana, Caroline, and Marshall Islands by the Japanese for rat con- trol or as a food source (Fisher, 1948; Marshall, 1975). This species has been collected on Enewetak Atoll only from Japtan Islet, and it apparently feeds on land crabs and some marine life. Specimens from Enewetak Atoll are commonly about 1 m in length. The Brahminy blind snake, Ramphotyphlops bramina (Daudin) (Typhlops or Typhlina braminus), is a small (<20 cm), secretive burrowing snake native to the Philip- pines and southeast Asia. The species has been introduced into new areas in the dirt accompanying transported plants and equipment and is now widely distributed among major island groups of the Pacific and Indian Oceans. It is found in Africa, Australia, India, and Mexico (Brown, 1957; McCoy, 1980). In Micronesia it is found on Guam, Tinian, Saipan, and Enewetak (Burt and Burt, 1932; Cagle, 1946; Knight, 1984). It may have been introduced to Enewetak with plantings from Hawaii, where it was initially observed in 1930 (Slevin, 1930; Fisher, 1948; Oliver and Shaw, 1953; Hunsaker and Breese, 1967). Blind snakes are secretive, nocturnal, and generally seen abroad only following a heavy rain. They are found in loose, moist soil in or under rotting logs and other debris and in gardens under stones, plastic cover, and potted plants (Cagle, 1946; McKeown, 1978; McCoy, 1980). Two specimens have been collected on Enewetak Atoll: one on Medren in 1954 (Knight, 1984) and one in 1980 on Enewetak Islet on the MPRL lanai at night. It remains to be seen whether any more specimens will be found on the atoll or whether the species can survive in the island’s rela- tively dry, sandy soil. Blind snakes are parthenogenic, and the single female parent lays two to eight elongate eggs in moist soil (McKeown, 1978). A population may thus be established from a single introduced individual. Blind snakes are beneficial to man because they eat termites and other soft-bodied insects and insect larvae (McKeown, 1978; McCoy, 1980). The relatively dry climate at Enewetak makes it a rather harsh environment for terrestrial reptiles. Some of the species found there are adapted to living where humans live and may have been transported there by man REPTILES 329 accidentally or intentionally (Hemidactylus, Lepidodactylus, Varanus, Ramphotyphlops). Some species inhabit beach areas and are found among driftwood (Lipinia, Emoia, Hemidactylus, Lepidodactylus). They may have been washed ashore on the atoll from other areas on floating debris or come as stowaways on the boats of early inter- island travelers. Gehyra and Hemiphyllodactylus are_pri- marily restricted to forest areas and may have been rafted to the atoll from other islands by storm waves which some- times wash over low-lying areas and can dislodge large trees and fallen debris. Enewetak Atoll has few species of reptiles compared to large, high islands such as Hawaii. Due to its dry climate, lack of extensive forest area, and relative isolation, it also has fewer species than some other atolls. Systematic reports on the terrestrial reptiles of atolls are rare and often are based on chance observation (e.g., Marshall, 1951; Moul, 1954; Sachet, 1962). Specimens discussed in these reports and many of the specimens collected from Enewetak were found by chance by people working on other projects. K. J. Frogner took advantage of limited opportunities to visit the various islands of Enewetak Atoll to collect as many specimens as possible, and much of what is known about the habitat of the lizards on the atoll is based on his work. However, a more thorough search, especially on the forested islets, could reveal more species present. Based on specimens in the USNM collection from nearby Pacific islands and other atolls in the Marshall Islands, the following species of lizards possibly could be found on Enewetak Atoll (R. Crombie, USNM, personal communication): Gehyra mutilata — widespread in Oceania Lamprolepis smaragdina — widespread in Oceania (found on Ailuk, Arno, Jaluit, Likiep, Taka, Ujae, Ujelang, Utirik, and Wotho Atolls in the Marshall Islands) Cryptoblepharus boutoni — widespread in Oceania (Pokak and Taongi in the Marshalls) Emoia baudini group — spotty in Oceania, identification doubtful E. arnoensis — Marshalls and possibly Caroline Islands (Arno, Bikini?, Lae, Jaluit, Rongerik?) E. boettgeri orientalis — spotty (Arno, Bikini?, Rongerik?) E. mivarti — spotty (Lae, Ujae, Wotho) Cyrtodactylus pelagicus — spotty (Arno Atoll) Perochirus ateles — spotty in Oceania but on Arno Atoll According to George Balazs (personal communication), only two species of sea turtles—the green turtle, Chelonia mydas, and the hawksbill turtle, | Eretmachelys imbricata—are known to Enewetak Atoll. Too few speci- mens have been examined to determine whether these two species belong to the subspecies C. mydas japonica of the western Pacific or E. imbricata squamata of the Indo- Pacific. At least one incidence of a possible sea turtle nest- ing on Ikuren Islet has been observed (P. Lamberson, per- sonal communication and photo). ACKNOWLEDGMENTS This report is based in part on observations made by the author during employment at the Mid-Pacific Research Laboratory, which is supported by the Division of Biomedi- cal and Environmental Research of the Department of Energy under contract No. AT(26-1)-628 to the University of Hawaii. Appreciation is extended to R. Crombie for his interest and help in identification of specimens and for his critique of the manuscript. Thanks are also extended to S. McKeown for many helpful suggestions and to G. Balazs for information on sea turtles. REFERENCES Brown, W. C., 1957, The Distribution of Terrestrial Reptiles in the Islands of the Pacific Basin, in Proceedings of the Eighth Pacific Science Congress, Vol. IIIA, University of the Philip- pines, Quezon City, Nov. 16-28, 1953, pp. 1479-1491, Pacific Science Association, Quezon City, Philippines. Burt, C. E., and M. D. Burt, 1932, Herpetological Results of the Whitney South Sea Expedition, VI, Pacific Island Amphibians and Reptiles in the Collection of the American Museum of Natural History, Bull. Amer. Mus. Nat. Hist., 63: 461-585. Bustard, H. R., 1970, Activity Cycle of the Tropical House Gecko, Hemidactylus frenatus, Copeia, 1970(1): 173-176. Cagle, F. R., 1946, Typhlops braminus in the Marianas Islands, Copeia, 1946(2): 101. Fisher, H. I., 1948, Locality Records of Pacific Island Reptiles and Amphibians, Copeia, 1948(1): 69. Heatwole, H., 1975, Biogeography of Reptiles on Some of the Islands and Cays of Eastern Papua—New Guinea, Atoll Research Bull. No. 180, Pacific Science Board, Washington, DEG Hunsaker, D., 1966, Notes on the Population Expansion of the House Gecko, Hemidactylus frenatus, Philippine J. Sci., 95: 121-122. ___, and P. Breese, 1967, Herpetofauna of the Hawaiian Islands, Pacific Sci., 21: 423-428. Knight, J. L., 1984, Typhlina bramina, Herp Review, 15: 115. Marcellini, Dale L., 1974, Acoustic Behavior of the Gekkonid Lizard, Hemidactylus frenatus, Herpetologica, 30: 44-52. Marshall, J. T., 1951, Vertebrate Ecology of Arno Atoll, Marshall Islands, Atoll Research Bull. No. 3, Pacific Science Board, Washington, D. C. Marshall, M., 1975, The Natural History of Namoluk Atoll, Eastern Caroline Islands, Atoll Research Bull. No. 189, Pacific Science Board, Washington, D. C. McCoy, M., 1980, Reptiles of the Solomon Islands, Wau Ecology Institute Handbook No. 7, Papua New Guinea. McKeown, S., 1978, Hawaiian Reptiles and Amphibians, The Oriental Publishing Company, Honolulu, Hawaii. Moul, E. T., 1954, Preliminary Report on Land Animals at Ono- toa Atoll, Gilbert Islands, Atoll Research Bull. No. 28, Pacific Science Board, Washington, D. C. Oliver, J. A., and C. E. Shaw, 1953, The Amphibians and Rep- tiles of the Hawaiian Islands, Zoologica, 38(5): 65-95. Sachet, M. H., 1962, Geography and Land Ecology of Clipperton Island, Atoll Research Bull. No. 86, Pacific Science Board, Washington, D. C. Slevin, J. R., 1930, A Note on the Discovery of the Genus Typhlops in the Hawaiian Islands, Copeia, 1930(4): 158. ® ‘ = ' , Py, Oe ee = | . eT ar . i ot 2 efi Gp ened ~ a > , .- : = : a Sw ren See Yi ' é ‘ 4 (Os _ 504 x i , / 7 we : ows .* eee @ > nv @ a ie <= m hea hay? O74 \iewe) ton P. ; eae im ® Ng ' ¥ tre y ¥ A | as S = SS ea = Birds of Enewetak Atoll ANDREW J. BERGER University of Hawaii, Honolulu, Hawaii 96822 In Chapter 1, Roy Tsuda wrote that “our present knowledge of the floristics and ecology of the marine benthic algae on Pacific atolls is based primarily on studies conducted on Enewetak Atoll.” This has not been the case with birds: I know of no systematic studies that are based on the birds of Enewetak Atoll. Most of the seabirds that occur at Enewetak have a very wide range in the Pacific basin, and they have not been shown to demonstrate a tendency for subspeciation. Because of its geographical location, Enewetak pro- vides nesting habitat for some species whose primary breeding range is north of the atoll and for others whose primary breeding range is south of it. For example, species that nest on the Hawaiian Islands include the wedge-tailed shearwater (Puffinus pacificus), red-tailed and white-tailed tropic birds (Phaethon rubricauds, P. leturus), red-footed Chapter 29 and brown boobies (Sula sula rubripes, S. leucogaster), great frigate bird (Fregata minor), gray-backed tern (Sterna lunata), sooty tern (S. fuscata), brown, black, and blue-gray noddies (Anous stolidus, A. tenuirostris, Procelsterna ceru- lea), and the white tern (Gygis alba). Enewetak species that also occur as breeding birds south of the equator include the sooty and slender-billed shearwaters (Puffinus griseus, P. tenuirostris) and the black-naped and crested terns (Sterna sumatrana, Thalassepus bergii). The reef heron (Egretta sacra) is a permanent resident on Enewetak Atoll and has a wide range that includes Korea, Japan, Malaysia, Australia, Melanesia, Polynesia, and Micronesia. The long-tailed cuckoo (Eudynamis taitensis), a nesting bird in New Zealand, spends the nonbreeding season on Enewetak and many other Pacific Islands from the Bismarck Archipelago eastward to the Marquesas Islands. At least 17 species of shorebirds that nest in Alaska or Siberia have been recorded as winter visitants on the islets of Enewetak Atoll. Table 1 provides a checklist of Enewetak Atoll birds. TABLE 1 Checklist of Enewetak Atoll Birds Order PROCELLARIIFORMES Family PROCELLARIIDAE Puffinus pacificus Salvin: Woodbury, 1962; Carpenter et al., 1968; Hailman, 1979. Puffinus griseus (Gmelin): Pearson and Knudsen, 1967. Puffinus tenuirostris (Temminck): Pearson and Knudsen, 1967. Pterodroma hypoleuca nigripennis Salvin: Johnson and Kienholz, 1975. Order PELICANIFORMES Family PHAETHONTIDAE Phaethon rubricauda (Mathews): Carpenter et al., 1968; Amerson, 1969. Phaethon lepturus Mathews: Carpenter et al., 1968; Amerson, 1969. Family SULIDAE Sula sula Gould: Carpenter et al., 1968; Amerson, 1969. Sula leucogaster (Forster): Woodbury, 1962; Amerson, 1969. Family FREGATIDAE Fregata minor (Gmelin): Carpenter et al., 1968; Amerson, 1969. Order CICONIIFORMES Family ARDEIDAE Egretta sacra Gmelin: Pearson and Knudsen, 1967; Carpenter et al., 1968; Amerson, 1969. (This table continued on next page.) 332 BERGER TABLE 1 (cont'd) Order ANSERIFORMES Family ANATIDAE Anas acuta Linnaeus: Temme, MS; Hailman, 1979. Anas querquedula Linnaeus: Hailman, 1979. Order GALLIFORMES Family PHASIANIDAE Gallus gallus (Linnaeus): Amerson, 1969. Order CHARADRIIFORMES Family SCOLOPACIDAE Limosa lapponica baueri Naumann: Pearson and Knudsen, 1968; Carpenter et al., 1968. Numenius phaeopus (Scopoli): Woodbury, 1962; Pearson and Knudsen, 1967. Numenius tahitensis (Gmelin): Woodbury, 1962; Pearson and Knudsen, 1967. *Tringa glareola Linnaeus: Hailman, 1979. Heteroscelus brevipes (Vieillot): Pearson and Knudsen, 1967; Carpenter et al., 1968. Heteroscelus incanus (Gmelin): Woodbury, 1962; Pearson and Knudsen, 1967. Arenaria interpres (Linnaeus): Woodbury, 1962; Pearson and Knudsen, 1967. Gallinago hardwickii (Gray): Hailman, 1979. Calidris alba (Pallas): Woodbury, 1962; Pearson and Knudsen, 1967. Calidris acuminata (Horsfield): Woodbury, 1962; Pearson and Knudsen, 1967. Calidris ruficollis (Pallas): Johnson and Kienholz, 1975. Tryngites subruficollis (Vieillot): Pearson and Knudsen, 1967. *Philomachus pugnax (Linnaeus): Temme photograph; Hailman, 1979. Family CHARADRIIDAE Charadrius dubius curonicus Gmelin: Baker, 1951; Pearson and Knudsen, 1967. Charadrius mongolus Stresemann: Hailman, 1979. Pluvialis dominica fulva (Gmelin): Woodbury, 1962; Pearson and Knudsen, 1967. Pluvialis squatarola (Linnaeus): Baker, 1951; Pearson and Knudsen, 1967. Family LARIDAE Sterna paradisaea Pontoppidan: Woodbury, 1962. Sterna sumatrana Raffles: Woodbury, 1962; Pearson and Knudsen, 1967. Sterna lunata Peale: Woodbury, 1962; Amerson, 1969. Sterna fuscata Linnaeus: Woodbury, 1962; Pearson and Knudsen, 1967. Thalasseus bergii (Lichtenstein): Woodbury, 1962; Pearson and Knudsen, 1967. Procelsterna cerulea (Bennett): Woodbury, 1962; Pearson and Knudsen, 1967. Anous stolidus (Linnaeus): Woodbury, 1962; Pearson and Knudsen, 1967. Anous tenuirostris (Bryan): Woodbury, 1962; Pearson and Knudsen, 1967. Gygis alba (Sparrman): Woodbury, 1962; Pearson and Knudsen, 1967. Order CUCULIFORMES Family CUCULIDAE Eudynamis taitensis (Sparrman): Woodbury, 1962. Order STRIGIFORMES Family STRIGIDAE Asio flammeus (Pontoppidan): Johnson and Kienholz, 1975. “New record for Enewetak and northern Marshall Islands. REFERENCES Amerson, A. B., dr., 1969, Ornithology of the Marshall and Gil- bert Islands. Atoll Res. Bull. No. 127, Smithsonian Institu- tion, Washington, 348 pp. Baker, R. H., 1951, The Avifauna of Micronesia, its Origin, Evo- lution and Distribution, University of Kansas Pub., Mus. of Nat. Hist., 3: 1-359. Carpenter, M. L., W. B. Jackson, and M. W. Fall, 1968, Bird Populations at Enewetok Atoll, Micronesica, 4: 295-307. Hailman, J. P., 1979, Notes on Birds of Enewetak Atoll, Marshall Islands, Elepaio, 40: 87-90. Johnson, O. W., and R. J. Kienholz, 1975, New Avifaunal Records from Enewetak, Auk, 92: 592-594. Pearson, D. L., and J. W. Knudsen, 1967, Avifaunal Records from Enewetak Atoll, Marshall Islands, Condor, 69: 201-203. Woodbury, A. M., 1962, A Review of the Ecology of Enewetak Atoll, Pacific Ocean, University of Utah Press Inst. of Biol. Res., 123 pp. Mammals of Enewetak Atoll ERNST S. REESE Department of Zoology University of Hawaii Honolulu, Hawaii 96817 Man is the dominant mammal of Enewetak Atoll. Humans arrived at least 2000 years ago (Craib, 1983). In all probability the Polynesian rat, Rattus exulans, and the domestic pig, Sus scrofa, accompanied him. Domestic dogs, Canis familiaris, and cats, Felix catus, most likely did not arrive until European contact in the 19"" century. The ubiquitous common house mouse, Mus musculus, and the roof rat, Rattus rattus, apparently arrived early in the 20th century when foreign contacts were more fre- quent. There is no evidence of the Norway rat, Rattus nor- vegicus, on Enewetak, although it occurs elsewhere in the Marshall Islands (Jackson, Vessey, and Bastian, Chapter 12 of Volume 1). All of these species of rats and mice are considered as human commensals and have been spread widely throughout the world through the activities of man (Eisenberg, 1981; Nowak and Paradiso, 1983). Domestic cats, pigs, and goats, Capra hircus, were left on Otdia (Wotjii) Atoll of the Ratak, or sunrise chain, by Otto von Kotzebue in 1816 and 1817. There is, however, no evidence that goats ever lived on Enewetak Atoll, which lies far to the northwest in the Ralik, or sunset chain of .atolls. Kotzebue did not visit this chain until his second voyage in 1824 (Kotzebue, 1830). Marine mammals are occasional visitors to Enewetak Atoll. Unfortunately there are only two positive identifica- tions based on skeletal remains, beached specimens, and photographs collected by Philip Lamberson, former laboratory manager of the Mid-Pacific Marine Laboratory, 1974-1977, and sent to William F. Perrin, National Marine Fisheries Service, La Jolla, California, for identification. The spinner dolphin, Stenella longirostris, is identified from photographs (Fig. 1) and by observations of its dis- tinctive spinning breaching behavior (Perrin, 1972), while the striped dolphin, Stenella coeruleoalba, is identified from a skeleton. The occurrence of this species at Enewetak represents a new record for the central Pacific (Hubbs, Per- rin, and Balcomb, 1973; W. F. Perrin, personal communi- cation to P. Lamberson, 1977). 333 Chapter 30 A checklist of mammals of Enewetak Atoll is provided in Table 1. Twenty-seven species of whales, dolphins, and _por- poises may be expected to occur at Enewetak Atoll (W. F. Perrin, personal communication). These are listed below in the order they are presented in Leatherwood et al. (1982), which provides a guide to their identification. Large whales with a dorsal fin: Blue whale, Balaenoptera musculus Fin whale, Balaenoptera physalus Sei whale, Balaenoptera borealis Bryde’s whale, Balaenoptera edeni Humpback whale, Megaptera novaeangliae Sperm whale, Physeter macrocephalus Medium-sized whales with a dorsal fin: Minke whale, Balaenoptera acutorostrata (Southern ?) bottlenose whale, Hyperoodon sp.? Cuvier’s beaked whale, Ziphius cavirostris Beaked whales of the genus Mesoplodon Stejneger’s beaked whale, M. stejnegeri Blainville’s beaked whale, M. densirostris Ginkgo-toothed beaked whale, M. ginkgodens Killer whale, Orcinus orca False killer whale, Pseudorca crassidens Short-finned pilot whale, Globicephala macrorhynchus Risso’s dolphin, Grampus griseus Small whales, dolphins, and porpoises with a dorsal fin: Spotted dolphin, Stenella attenuata Spinner dolphin, Stenella longirostris Striped dolphin, Stenella coeruleoalba Common dolphin, Delphinus delphis Fraser’s dolphin, Lagenodelphis hosei Bottlenose dolphin, Tursiops truncatus Rough-toothed dolphin, Steno bredanensis Pygmy killer whale, Feresa attenuata Melon-headed whale, Peponocephala electra Pygmy sperm whale, Kogia breviceps Dwarf sperm whale, Kogia simus Other marine mammals, such as seals and sea lions, are not known to occur in the Marshall Islands. According to Lamberson, the Marshallese recognize two kinds of porpoises. One is small and travels in large 334 REESE (a) (b) Fig. 1 Two views of the spinner dolphin, Stenella longirostris, in the lagoon of Enewetak Atoll. [Photos by (a) Paul M. Allen and (b) Philip Lamberson.] MAMMALS 335 TABLE 1 Checklist of the Mammals of Enewetak Atoll* Phylum CHORDATA Class MAMMALIA Order ARTIODACTYLA Family SUIDAE Sus scrofa Linnaeus, 1758. Domestic pig. Order CARNIVORA Family CANIDAE Canis familiaris Linnaeus, 1758. Domestic dog. Family FELIDAE Felis catus Linnaeus, 1758. Domestic cat. Order CETACEA Family DELPHINIDAE +Stenella coeruleoalba (Meyen), 1833. Striped dolphin. +Stenella longirostris (Gray), 1828. Spinner dolphin. Order PRIMATE Family HOMINIDAE Homo sapiens Linnaeus, 1758. Humans. Order RODENTIA Family MURIDAE Mus musculus Linnaeus, 1766. House mouse. Rattus exulans (Peale), 1848. Polynesian rat. Rattus rattus (Linnaeus), 1758. Roof rat. "Classification modified from Nowak and Paradiso, 1983, and Tomich, 1969. +New Enewetak record. groups and is called “ke.” It is probably Stenella longirostris. The other is larger and swims in smaller groups. It may be S. coeruleoalba or one of the other species which should occur at Enewetak. Lamberson thinks the Marshallese call the larger animal “rak.” Occasionally Marshallese men harpoon dolphins for food which may explain the dolphins’ wariness of boats. They are also extremely wary of swimmers and divers at Enewetak. REFERENCES Craib, J. L., 1983, Micronesian Prehistory: An Archeological Overview, Science, 219: 922-927. Eisenberg, J. F., 1981, The Mammalian Radiations, University of Chicago Press, Chicago. Hubbs, C. L., W. F. Perrin, and K. C. Balcomb, 1973 Stenella coeruleoalba in the Eastern and Central Tropical Pacific, J. Mammalogy, 54: 549-552. Kotzebue, O. von, 1930, A New Voyage Round the World in the Years 1823, 24, 25, and 26, Henry Colburn and Richard Bentley, London. Leatherwood, S., R. R. Reeves, W. F. Perrin, and W. E. Evans, 1982, Whales, Dolphins, and Porpoises of the Eastern North Pacific and Adjacent Arctic Waters. A Guide to Their Identifi- cation, NOAA Technical Report NMFS Circular 444, pp. v, 245. Nowak, R. M., and J. L. Paradiso, 1983, Walker’s Mammals of the World, 4th edition, Johns Hopkins University Press, Baltimore. Perrin, W. F., 1972, Color Patterns of Spinner Porpoises (Stenella cf. s. longirostris) of the Eastern Pacific and Hawaii, with Comment on Delphinid Pigmentation, Fishery Bull., 70: 983-1003. Tomich, P. Q., 1969, Mammals in Hawaii, Bishop Museum Press, Honolulu. Chapter 31 Miscellaneous Species Records of Enewetak Atoll RICHARD H. TITGEN and BEATRICE L. BURCH Bernice P. Bishop Museum Honolulu, Hawaii 96817 INTRODUCTION The preceding chapters in Volume II are compilations of species records from Enewetak Atoll. For various reasons all systematic groups could not be covered in a separate chapter. Consequently, this final chapter includes miscellaneous records from the Mid-Pacific Research Laboratory at Enewetak Atoll and from the literature. Ideally, all Enewetak species records not contained in the previous chapters would be presented here. Unfor- tunately, the vastness of the biological literature and the time available would not permit such an undertaking. Therefore, the search was confined to the more obvious reference materials that might contain these records. We hope this small checklist (Table 1) will be of some help, and we apologize to those authors whose records we have missed. Some of the previous chapters listed only geologically recent material. For this reason some fossil material and/or material from drill cores is contained herein, even though the systematic group may have been more com- pletely discussed in previous chapters of this volume. TABLE 1 Checklist of Miscellaneous Species of Enewetak Atoll KINGDOM PLANTAE Division PYRRHOPHYTA Class DINOPHYCEAE Order PERIDINIALES Family HETERAULACACEAE Gambierdiscus toxicus Adachi and Fukuyo, 1979: see Colin, Vol. I. Division CHLOROPHYTA Class CHLOROPHYCEAE Order CLADOPHORALES Family CLADOPHORACEAE Cladophora sp. Cladophora hemisphaerica: Bailey-Brock et al., 1980. Order SIPHONOCLADALES Family VALONIACEAE Valonia trabeculata Egerod, 1952: Bailey-Brock et al., 1980. Valonia ventricosa J. Agardh, 1887. Valonia ventricava: Bailey-Brock et al., 1980 Order DASYCLADALES Family DASYCLADACEAE Acetabularia clavata Yamada, 1934: Bailey-Brock et al., 1980. Division PHAEOPHYTA Class PHAEOPHYCEAE Order DICTYOTALES Family DICTYOTACEAE Padina japonica Yamada, 1931: Bailey-Brock et al., 1980 (This table continued on next page.) 338 TITGEN AND BURCH TABLE 1 (cont’d) Division RHODOPHYTA Class FLORIDEOPHYCEAE Order RHODYMENIALES Family RHODYMENIACEAE Chylocladia sp.: Bailey-Brock et al., 1980. KINGDOM ANIMALIA Phylum SARCOMASTIGOPHORA Subphylum SARCODINA Class GRANULORETICULOSA Order FORAMINIFERIDA Suborder ROTALIINA Superfamily GLOBIGERINOIDEA Family GLOBIGERINIDAE *Globigerina sp. aff. G. apertura Cushman, 1918: Todd, 1964. *Globigerina conglomerata Schwager, 1866: Todd, 1964. *Globigerina (Beella) digitata Brady, 1884: Todd, 1964. *Globigerina (Globorotaloides) hexagona Natland, 1938: Todd, 1964. *Globigerina obesa (Bolli, 1957): Todd, 1964. *Globigerina rubescens Hofker, 1956: Todd, 1964. *Globigerinella adamsi (Banner and Blow, 1959): Todd, 1964. *Globigerinella aequilateralis (Brady, 1884): Todd, 1964. *Globigerinita glutinata (Egger, 1893): Todd, 1964. *Globigerinita humilis (Brady, 1884): Todd, 1964. *Globigerinoides elongatus (d’Orbigny, 1826): Todd, 1964. *Globigerinoides sacculifer fistulosa (Schubert, 1910): Todd, 1964. *Globoquadrina altispira (Cushman and Jarvis, 1936): Todd, 1964. *Globoquadrina altispira globosa Bolli, 1957: Todd, 1964. *Orbulina bilobata (d’Orbigny, 1846): Todd, 1964. *Orbulina suturalis Bronnimann, 1951: Todd, 1964. *Sphaeroidinella disjuncta Finlay, 1940: Todd, 1964. *Sphaeroidinella kochi (Caudri, 1934): Todd, 1964. Family GLOBOROTALIIDAE *Globorotalia (Turborotalia) acostaensis Blow, 1959: Todd, 1964. *Globorotalia fohsi robusta Bolli, 1950: Todd, 1964. *Globorotalia (Truncorotalia) punctulata (d’Orbigny, 1826): Todd, 1964. Family HASTIGERINIDAE *Hastigerina pelagica (d’Orbigny, 1839): Todd, 1964. Phylum CNIDARIA Class SCYPHOZOA Order CORONATAE Family LINUCHIDAE +Linuche sp. Class HYDROZOA Order HYDROIDA Suborder ANTHOMEDUSAE Family CLAVIDAE ?Rhizogeton sp.: Cooke, 1975. Family EUDENDRIIDAE Eudendrium ?breve Fraser, 1938: Cooke, 1975. Eudendrium capillare Alder, 1856: Cooke, 1975. Family HALOCORDYLIDAE Halocordyle disticha (Goldfuss, 1820): Cooke, 1975. Suborder LEPTOMEDUSAE Family CAMPANULARIIDAE Clytia hemisphaerica (Linnaeus, 1767): Cooke, 1975. Family HALECIIDAE Halecium beani (Johnson, 1847): Cooke, 1975. Halecium sp.: Cooke, 1975. *Fossil material and/or material from cores. +Records from the Enewetak Laboratory Card Catalogue. (This table continued on next page.) MISCELLANEOUS SPECIES RECORDS 339 TABLE 1 (cont'd) Family PLUMULARIIDAE +Aglaophenia lophocarpa Allman, 1877. Aglaophenia pluma (Linnaeus, 1758): Cooke, 1975. Halopteris diaphana (Heller, 1868): Cooke, 1975. Plumularia halecioides Adler, 1859: Cooke, 1975. Plumularia setacea (Linnaeus, 1758): Cooke, 1975. Family SERTULARIIDAE Dynamena cornicina McCrady, 1857: Cooke, 1975. Dynamena crisioides Lamouroux, 1824: Cooke, 1975. Sertularella miniscula Billard, 1925: Cooke, 1975. Sertularia subtilis Fraser, 1937: Cooke, 1975. Sertularia westindica Stechow, 1919: Cooke, 1975. Thyroscyphus vitiensis Marktanner-Turneretscher, 1890: Cooke, 1975. Order MILLEPORINA Family MILLEPORIDAE +Millepora murrayi Quelch, 1886. Millepora platyphylla Hemprich and Ehrenberg, 1834: Wells, 1954. Millepora tenera Boschma, 1949: Wittle and Wheeler, 1974; Wittle et al., 1974. Millepora dichotoma: Middlebrook et al., 1971. Order STYLASTERINA Family STYLASTERIDAE +Distichopora coccinea Gray, 1860. Distichopora fisheri Broch, 1942: Wells, 1954. +Distichopora violacea (Pallas, 1766). Stylaster elegans Verrill, 1864: Wells, 1954. Class ANTHOZOA Subclass ALCYONARIA Order ALCYONACEA Family NEPHTHEIDAE Nephthea sp.: Ciereszko et al., 1968; Schmitz et al., 1974; Vanderah et al., 1978. Subclass ZOANTHARIA Order CORALLIMORPHARIA Family ACTINODISCIDAE +Rhodactis inchoata Carlgren, 1943. Order SCLERACTINIA Suborder ASTROCOENIINA Family ACROPORIDAE *Dendracis pacificus Wells, 1964. Family ASTROCOENIIDAE *Actinastrea minutissima (Gerth, 1921): Wells, 1964. Family POCILLOPORIDAE *Seriatopora micrommata Felix, 1921: Wells, 1964. *Seriatopora ornata Felix, 1921: Wells, 1964. *Stylophora sp. cf. S. sokkohensis Gerth, 1921: Wells, 1964. *Stylophora stellata (von Fritsch, 1875): Wells, 1964. *Stylophora sp.: Wells, 1964. Suborder DENDROPHYLLIINA Family DENDROPHYLLIIDAE +Rhizopsammia minuta var. bikiniensis Wells, 1954. Suborder FAVIINA Family FAVIIDAE *Favia sp. cf. F. oligophylla (von Fritsch, 1875): Wells, 1964. Family MUSSIDAE *Acanthophyllia sp.: Wells, 1964. Family OCULINIDAE *Galaxea sp.: Wells, 1964. Suborder FUNGIINA Family AGARICIIDAE *Leptoseris sp. cf. L. floriformis Gerth, 1923: Wells, 1964. “Fossil material and/or material from cores. +Records from the Enewetak Laboratory Card Catalogue. (This table continued on next page.) 340 TITGEN AND BURCH TABLE 1 (cont'd) Family FUNGIIDAE *Discotrochus sp. cf. D. orbignyanus Milne Edwards and Haime, 1848: Wells, 1964. Family PORITIDAE *Alveopora polyacantha Reuss, 1867: Wells, 1964. *Dictyaraea micrantha Reuss, 1867: Wells, 1964. *Porites sp. cf. P. capricornis Rehberg, 1892: Wells, 1964. Order ZOANTHIDEA Family ZOANTHIDAE +Gemmaria sp. +Isaurus duchassaingi Andres, 1883. +Palythoa tuberculosa (Esper, 1791). +Zoanthus sp. Subclass CERIANTIPATHARIA Order CERIANTHARIA +Suborder PENICILLARIA +Family ARACHNACTIDAE +Arachnanthus australiae Carlgren, 1937. Phylum PLATYHELMINTHES Class CESTODA Subclass EUCESTODA Order CYCLOPHYLLIDEA Family HYMENOLEPIDIDAE Hymenolepis diminuta (Rudolphi, 1819): see Jackson et al., Vol. I., Chapter 12. Phylum NEMATODA Class SECERNENTEA Subclass SPIRURIA Order SPIRURIDA Suborder SPIRURINA Superfamily SPIRUROIDEA Family GONGYLONEMATIDAE Gongylonema neoplasticum Fibiger and Ditlevsen, 1914: see Jackson et al., Vol. I, Chapter 12. Phylum SIPUNCULA Family PHASCOLOSOMATIDAE §Phascolosoma albolineatum Baird, 1868. §Phascolosoma pacificum Keferstein, 1866. Family SIPUNCULIDAE §Siphonosoma rotumanum (Shipley, 1898). Phylum CHAETOGNATHA Class SAGITTOIDEA Order PHRAGMOPHORA Family SPADELLIDAE Spadella legazpichessi Alvarino, 1981. Phylum ARTHROPODA Subphylum CRUSTACEA Class MALACOSTRACA Subclass EUMALACOSTRACA Superorder PERACARIDA Order ISOPODA Suborder ONISCOIDEA Infraorder LIGIAMORPHA Section CRINOCHETA Superfamily ATRACHEATA Family ONISCIDAE 8Alloniscus oahuensis Budde-Lund, 1885. *Fossil material and/or material from cores. +Records from the Enewetak Laboratory Card Catalogue. See den Hartog (1977) on the systematic classification of the genus Arachnanthus. §New Enewetak records. (This table continued on next page.) MISCELLANEOUS SPECIES RECORDS TABLE 1 (cont’d) 341 Superfamily PSEUDOTRACHEATA Family PORCELLIONIDAE Porcellionides pruinosus (Brandt, 1833). Metapornorthrus pruinosus: Woodbury, 1962. Suborder EPICARIDEA Infraorder BOPYRINA Family BOPYRIDAE Bopyrella thomsoni Bonnier, 1900; Bourdon, 1980. Bopyrella thomsoni muiensis Danforth, 1970. Cancricepon garthi Danforth, 1970. Cancricepon? knudseni (Danforth, 1970). Merocepon knudseni Danforth, 1970. Gigantione pratti Danforth, 1967: Danforth, 1970. Grapsicepon sp.: Danforth, 1970. Merocepon knudseni Danforth, 1970. Scyracepon hawaiiensis Richardson, 1911: Danforth, 1970. Order AMPHIPODA Suborder GAMMARIDEA Superfamily GAMMAROIDEA Family GAMMARIDAE Beaudettia palmeri Barnard, 1965. Elasmopus pectenicrus (Bate, 1862): Barnard, 1965. Elasmopus pseudaffinis Schellenberg, 1938: Barnard, 1965. Elasmopus rapax Costa, 1853: Barnard, 1965. Elasmopus spinidactylus Chevreux, 1907: Barnard, 1965. Elasmopus sp.: Croker, 1971b. Jerbarnia mecochira Croker, 197 1a. Maera hamigera Haswell, 1879: Barnard, 1965. Maera inaequipes (Costa, 1851): Barnard, 1965. Maera othonopsis Schellenberg, 1938: Barnard, 1965. Meera pacifica Schellenberg, 1938: Barnard, 1965. Maera quadrimana (Dana, 1853): Barnard, 1965. Meera serrata Schellenberg, 1938: Barnard, 1965. Mallacoota insignis (Chevreux, 1901). Maera insignis: Barnard, 1965. Melita celericula Croker, 1971b. Superfamily LEUCOTHOIDEA Family LEUCOTHOIDAE Leucothoe hyhelia Barnard, 1965. Superfamily TALITROIDEA Family HYALIDAE Hyale chevreuxi K. H. Barnard, 1916: Barnard, 1965. Hyale dentifera Chevreux, 1907: Barnard, 1965. Hyale honoluluensis Schellenberg, 1938: Barnard, 1965. Hyale media (Dana, 1853): Barnard, 1965. Talorchestia spinipalma (Dana, 1853): Barnard, 1960; Woodbury, 1962. Superfamily COROPHIOIDEA Family AMPITHOIDAE Ampithoe ramondi Audouin, 1826: Barnard, 1965. Cymadusa brevidactyla (Chevreux, 1907): Croker, 1971b. Cymadusa filosa Savigny, 1816: Barnard, 1965. Paragrubia vorax Chevreux, 1901: Barnard, 1965. Family AORIDAE Lembos aequimanus Schellenberg, 1938: Barnard, 1965. Lembos bryopsis Barnard, 1965. Lembos sp. cf. L. francanni Reid, 1951: Barnard, 1965. Lembos sp. cf. L. intermedius Schellenberg, 1938: Barnard, 1965. Microdeutopus tridens Schellenberg, 1938: Barnard, 1965. Family ISAEIDAE Gammaropsis atlanticus (Stebbing, 1888). Eurystheus atlanticus: Barnard, 1965. (This table continued on next page.) 342 TITGEN AND BURCH TABLE 1 (cont’d) Family ISAEIDAE (cont'd) Gammaropsis digitatus Schellenberg, 1938. Eurystheus digitatus: Barnard, 1965. Gammaropsis pacificus Schellenberg, 1938. Eurystheus ?pacificus: Barnard, 1965. Megamphopus abbotti Barnard, 1965. Suborder CAPRELLIDEA Family CAPRELLIDAE §Metaprotella sandalensis Mayer, 1903. Phylum TARDIGRADA Order EUTARDIGRADA Family MACROBIOTIDAE Hypsibius (Isohypsibius) augusti Murray, 1907. Hypsibius (Isohypsibius) angusti: Mehlen, 1972. Hypsibius (Calohypsibius) truncatus Thulin, 1928: Mehlen, 1972. Macrobiotus allani Murray, 1913: Mehlen, 1972. Macrobiotus harmsworthi Murray, 1907: Mehlen, 1972. Macrobiotus richtersi Murray, 1911: Mehlen, 1972. Family MILNESIIDAE Milnesium tardigradum Doyere, 1840: Mehlen, 1972. Phylum ECHINODERMATA Subphylum ECHINOZOA Class ECHINOIDEA Subclass EDECHINOIDEA Superorder GNATHOSTOMATA Order CLYPEASTEROIDA Suborder LAGANINA Family FIBULARIIDAE *Echinocyamus parviporus Kier, 1964. *Echinocyamus cf. parviporus Kier, 1964. *Echinocyamus petalus Kier, 1964. Phylum HEMICHORDATA Class ENTEROPNEUSTA Family PT YCHODERIDAE +Ptychodera flava Eschscholtz, 1825. “Fossil material and/or material from cores. tRecords from the Enewetak Laboratory Card Catalogue. §New Enewetak records. ACKNOWLEDGMENTS The authors would like to thank those who helped sup- ply species records used in this chapter. J. L. Barnard supplied records for peracarid Crustacea, G. A. Samuel- son helped with the Tardigrada, and J. H. Bailey-Brock forwarded records of Sipuncula that had been identified by E. B. Cutler. J. Markham helped us update the bopyrid isopod records. I. Abbot helped with the algae, and A. Alvarinio helped with the chaetognatha. REFERENCES Alvarifio, A., 1981, Spadella legazipichessi, A New Benthic Chae- tognath from Enewetak, Marshall Islands, Proc. Biol. Soc. Wash., 94: 107-121. Bailey-Brock, J. H., J. K. White, and Ik, (A. Ward, 1980, Effects of Algal Turf and Depressions as Refuges on Polychaete Assemblages of a Windward Reef Bench at Enewetak Atoll, Micronesica, 16: 43-58. Barnard, J. L., 1960, Insects of Micronesia, Crustacea: Amphi- poda (Strand and Terrestrial Talitridae), Insects Micronesia, 4: 13-30. ——, 1965, Marine Amphipoda of Atolls in Micronesia, Proc. U. S. Natl. Mus., 117(3516): 459-552. Bourdon, R., 1980, Les especes du genre Bopyrella, J. Bonnier (Crustacea, Isopoda, Bopyridae), Bull. Mus. Natn. Hist. Nat. (Paris), (4) 2(A): 185-236. Ciereszko, L. S., M. A. Johnson, R. W. Schmidt, and C. B. Koons, 1968, Chemistry of Coelenterates—VI, Occurrence of Gorgosterol, a C39 Sterol, in Coelenterates and Their Zoo- xanthellae, Comp. Biochem. Physiol., 24: 899-904. Cooke, W. J., 1975, Shallow Water Hydroids from Enewetak Atoll, Marshall Islands, Micronesica, 11: 85-108. Croker, R. A., 1971a, A Remarkable New Amphipod Genus (Crustacea, Gammaridae) from Eniwetok Atoll Lagoon, Pac. Sci., 25: 382-386. MISCELLANEOUS SPECIES RECORDS 343 __, 1971b, A New Species of Melita (Amphipoda: Gammaridae) from the Marshall Islands, Micronesia, Pac. Sci., 25: 100-108. Danforth, C. G., 1967, Northern Pacific Gigantione (Isopoda), Biol. Bull. (Woods Hole), 132: 147-155. —_, 1970, Epicarids (Isopoda) of Eniwetok Atoll, Pac. Sci., 24: 462-471. den Hartog, J. C., 1977, Descriptions of Two New Ceriantharia from the Caribbean Region, Pachycerianthus curacaoensis n. sp. and Arachnanthus nocturnus n. sp., with a Discussion of the Cnidom and of the Classification of the Ceriantharia, Zool. Meded. (Leiden), 51: 211-242. Kier, P. M., 1964, Fossil Echinoids from the Marshall Islands, in Bikini and Nearby Atolls, Marshall Islands, Geol. Surv. Prof. Pap., 260-GG, pp. 1121-1126. Mehlen, R. H., 1972, Eutardigrada: Distribution at Eniwetok Atoll, Marshall Islands, Pac. Sci., 26: 223-225. Middlebrook, R. E., L. W. Wittle, E. D. Scura, and C. E. Lane, 1971, Isolation and Purification of a Toxin from Millepora dichotoma, Toxicon, 9: 333-336. Schmitz, F. J., D. J. Vanderah, and L. S. Ciereszko, 1974, Marine Natural Products: Nephthenol and Epoxynephthenol Acetate, Cembrene Derivatives from a Soft Coral, J. Chem. Soc. Chem. Commun., 1974: 407-408. Todd, R., 1964, Planktonic Foraminifera from Deep-Sea Cores off Eniwetok Atoll, in Bikini and Nearby Atolls, Marshall Islands, Geol. Surv. Prof. Pap., 260-CC, pp. 1067-1100. Vanderah, D. J., N. Rutledge, F. J. Schmitz, and L. S. Ciereszko, 1978, Marine Natural Products: Cembrene-A and Cembrene-C from a Soft Coral, Nephthea sp., J. Org. Chem., 43: 1614-1616. Wells, J. W., 1954, Recent Corals of the Marshall Islands, in Bikini and Nearby Atolls, Part 2, Oceanography (Biologic), Geol. Surv. Prof. Pap., 260-I, pp. 385-486. , 1964, Fossil Corals from Eniwetok Atoll, in Bikini and Nearby Atolls, Marshall Islands, Geol. Surv. Prof. Pap., 260-DD, pp. 1101-1111. Wittle, L. W., E. D. Scura, and R. E. Middlebrook, 1974, Sting- ing Coral (Millepora tenera) Toxin: A Comparison of Crude Extracts with Isolated Nematocyst Extracts, Toxicon, 12: 481-486. __, and C. A. Wheeler, 1974, Toxic and Immunological Proper- ties of Stinging Coral Toxin, Toxicon, 12: 487-493. Woodbury, A. M., 1962, A Review of the Ecology of Enewetok Atoll, Pacific Ocean, University of Utah Institute of Environ- mental Biological Research, Ecology and Epizoology Research, Salt Lake City. Author Index Arneson, A. C., 53 Bailey-Brock, J. H., 97 Berger, A. J., 331 Bruce, A. J., 221 Burch, B. L., 337 Chave, E. H., 37 Child; ‘©. A., 179 Cox, R. S., 85 Cuffey, R. J., 85 Cutress, B. M., 263 GutressnGa Es 53 Devaney, D. M., 37, 49, 63, 67, 93, 95, 97, 197, 221, 277 Dunn, P. H., 11 Eldredge, L. G., 287 Garth, J. S., 235 Gerber, R. P., 203 Grant, R. E., 77 Haig, J., 235 345 Johnson, S., 105 Kay, E. A., 105 Kornicker, L. S., 217 Knudsen, J. W., 235 Lamberson, J. O., 17, 325 Lang, J. C., 67 Manning, R. B., 181 Nishida, G. M., 147 Randall, H. A., 289 Randall, J. E., 289 Reaka, M. L., 181 Reese, E. S., 333 Reynolds, D., 11 Rowe, F. W. E., 263 Samuelson, G. A., 147 Titgen, R. H., 191, 337 Tsuda, R. T., 1 Subject Index The reader interested in a particular group of organisms should consult the table of contents at the front of the volume and turn to the appropriate chapter. Scientific names and authors’ names are not listed here as they would be redundant with the checklists included in each chapter. Vernacular names of organisms and certain scientific names used in the vernacular sense are included. Only Indo-Pacific geographic place names are included, with the exception of Enewetak Atoll and Marshall Islands, which are referred to in every chapter, and the names of islands within Enewetak Atoll. Ailinglapalap Atoll, 8 Alaska, 331 Algae, 1, 2, 8 Andaman Islands, 181 Arachnids, 158 Arno Atoll, 153, 154, 164, 166, 167, 168, 228, 235 Atomic Energy Commission, 25 Australia, 67, 80, 102, 103, 141, 158, 159, 161-169, 180, 181, 182, 194, 225, 228, 264, 287, 288, 328, 331 Barnacles, 193-195 Basket stars, 281 Behavior geckos, 325, 328 stomatopods, 181, 187 Bikini Atoll, 1, 24, 25, 63, 67, 77, 80, 82, 96, 97, 141, 154, 158, 162, 167, 169, 179, 199, 212, 213, 214, PPD, CI) PUT PAM, Gus} Bioerosion corals, 73 crabs, 248 polychaetes, 97, 102 sipunculans, 93 sponges, 49 Biogeography, 17, 23, 134, 141, 154-156, 313, 314 Birds, 331 Bismarck Archipelago, 325, 331 Burma, 180 Caroline Islands, 37, 105, 141, 155, 156, 158, 159, 160-169, 191, 194, 195, 199, 316, 325, 328 Cats, 333 Ciliates, 46 Coconut crabs, 29, 34, 236 Cocos Island, 264 Color patterns, stomatopods, 181, 182, 186, 187 Communities, soft bottom, 133 Copepods, 197 Crabs, 235-238 Crinoids, 277, 280 Crown-of-Thorns, 282 Dogs, 333 Dolphins, 333 Domestic animals, see Mammals Easter Island, 102, 264, 314 Ecological succession, 25, 27-29 Endemism fishes, 314, 316 insects, 154, 155, 158, 159, 160-169 mollusks, 133 stomatopods, 181, 182 Environmental effect of German occupation, 24, 34 of Japanese occupation, 24, 30 of man, 141 of nuclear testing, 17, 24, 25, 30, 34 of tropical storms, 23 of war, 17, 24, 30, 34 Extinctions, mollusks, 141, 142 Fanning Island, 133 Fiji, 141, 142, 163, 167, 194, 217, 288 Fishes, 289, 313-316 Flatworms, 95, 96 Flukes, 95, 96 Foraminifera, 37, 46 Fossil bryozoans, 85, 91 crustaceans, 237 mollusks, 133, 139, 141, 142 pollen grains, 17 French Polynesia, 54 Frigate birds, 34 Funafuti Atoll, 133 Fungi, 11, 14, 15 347 Galapagos Islands, 264 Geckos, 325-328 Genetic variability, mollusca, 134 Geological record, mollusks, 134 Gilbert Islands, 54, 105, 156, 158, 159, 160-169, 194, 316 Goats, 333 Guam, 8, 15, 141, 159, 161, 162, 163, 164, 168, 264, 280, 314, 328 Hawaii, 14, 15, 37, 53, 102, 133, 141, 153, 156, 158-169, 180, 182, 217, 264, 282, 283, 312, 313, 314, 325, 328, 329, 331 Hermit crabs, 235-237 Heron Island, 102 Ifaluk Atoll, Caroline islands, 199 Indian Ocean, 15, 181, 225, 227, 313, 325, 328 Indonesia, 181, 288 Jaluit Atoll, 8, 164, 225, 235, 280, 281, 289 Japan, 166, 168, 181, 194, 227, 228, 287, 288, 331 Korea, 331 Kwajalein Atoll, 37, 54, 134, 141, 153, 155, 156, 159-169, 226, 228, 235, 237, 283, 289-313 Land crabs, 236 Line Islands, 264 Lizards, 325, 328 Lobsters, 221, 228 Madagascar, 288 Majuro Atoll, 211, 212, 213, 214, 235, 289 Malaysia, 287, 288, 331 Maldives Islands, 264 348 Mammals, 333 Man, 333 Mangrove swamps, 17 Mantis shrimps, 181, 182, 187-189 Mariana Islands, 105, 134, 141, 159, 165, 167-169, 289, 314, 325, 328 Marquesas Islands, 331 Meiobenthos, polychaetes, 103 Melanesia, 331 Micronesia, 217, 331 Molucca Islands, 228 Mollusks, 105, 132, 133, 134 bivalves, 133, 134 cephalopods, 105, 134 gastropods, 132, 133, 134, 139 nonmarine, 133 opisthobranchs, 132 Monitor lizard, 328 Moorea, 15 Mortality, of stomatopods, 188 Mouse, 333 Natural products chemistry, of sponges, 63 Nematodes, 95, 96 New Caledonia, 15, 194, 288 New Guinea, 15, 225, 228 New Hebrides, 142 New Zealand, 15, 180, 331 Nonmarine Mollusks, 133 Oceania, 181 Okinawa 133, 158, 194 Operation Crossroads, 1, 37, 97, 105, 235, 289 Ostracods, 217 Palmyra, Line Islands, 288 Palau, 158, 159, 160, 162, 167, 194, 237, 280 Papua New Guinea, 325, 328 Parasites cirripedia, 193 copepods, 197, 199 crabs, 236, 237, 248 holothuria, 264 SUBJECT INDEX insects, 159, 167, 168 Pycnogonida, 179 worms, 95 Philippines, 141, 162, 163, 167, 169, 194, 225, 237, 264, 283, 287, 288, 328 Phoenix Islands, 161, 163 Phytoplankton, 203 Pigs, 333 Pisonia forest, 23, 28, 30, 34 Plankton, see Zooplankton Plants, 17, 23, 24, 25, 27, 28, 30, 34 breadfruit, 24, 25 coconut, 24, 25, 30, 34 pandanus, 17, 24, 30, 34 pisonia, 17, 23, 27, 28, 29, 30, 34 Polychaetes, 97, 102, 103 Polynesia, 228, 331 Porpoise, 333 Predation fishes on foraminifera, 37 nudibranchs on sponges, 49 stomatopods, 186, 187 Productivity algae, 1 corals, 73 phytoplankton, 203 Protozoans, 37, 46 Radiobiology algae 1, 24, 25, 30 corals, 73 sponges, 49 Rats, 333 Red Sea, 181 Rongelap Atoll, 211, 221, 227, 235, 277 Rongerik Atoll, 211, 280, 281 Ryukyu Islands, 225, 237 Samoa Islands, 159, 160, 161, 164, 167, 168, 181, 194, 314 Scorpions, 158 Sea anemones, 53, 54 Sea birds, 331 Sea cucumbers, 263, 264 Sea grass, 8, 14, 134, 141 Seychelles Islands, 264 Shearwaters, 30 Shorebirds, 331 Shrimps, 221, 225, 226 Singapore, 180 Skinks, 328 Snakes, 325, 328 Society Islands, 15, 264, 314 Solomon Islands, 37, 102, 325 South China Sea, 181 Spiders, 158 Sponges, 49, 51 Sri Lanka, 164, 165 Starfish, 277 Sumatra, 102 Symbiosis barnacles, 194, 195 copepods, 197 corals, 63, 73 crabs, 236, 237 foraminifera, 46 polychaetes, 97 pycnogonida, 179 shrimps, 225-228 Tahiti, 160 Tapeworms, 95 Tasmania, 80, 287, 288 Temperature adaptations, of scleractinia, 74 Thailand, 102, 181, 182, 217 Tonga, 15 Toxicity, crabs, 237 Trobriand Islands, 325 Tuamotus Islands, 283 Turtles, 329 Ujelang Atoll, 8, 25, 235 Vanuatu, 142 Whales, 333 Zanzibar, 288 Zooplankton, 203, 211-215 Zooxanthellae, corals, 63 -87079-580-5 ISBN O i 4 4 “ i , -