Pacific Plate Biogeography, with Special Reference to Shorefishes
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S. Dillon Ripley Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 367
Pacific Plate Biogeography, with Special Reference to Shorefishes
Victor G. Springer
SMITHSONIAN INSTITUTION PRESS City of Washington 1982 ABSTRACT Springer, Victor G. Pacific Plate Biogeography, with Special Reference to Shorefishes. Smithsonian Contributions to Zoology, number 367, 182 pages, 65 figures, 5 tables, 1982.—It is the thesis of this study that the Pacific lithospheric plate (Pacific Plate) forms a major subunit of the Indo-Pacific biogeographic region. This thesis is defended on two primary grounds: the sharp decrease in numbers of taxa as one proceeds eastward across the western margin of the Pacific Plate, and the presence of a high degree of endemism on the Plate. The complex geotectonic history of the Plate and adjacent plate margins is discussed and the vicariance scenarios that produced the three main types of endemism (widely distributed Plate endemism, endemism restricted to a limited region on the Plate, and endemism restricted to single islands or island groups) are hypothesized. Anecdotal discussions of the distributions of 179 shorefish family groups are presented, as well as discussions of the distributions of selected terrestrial and marine taxa of other organisms. Particular attention is given to presumed erroneous or exceptional distributions of fishes on the Pacific Plate, including, but not limited to, north-south disjunct distributions, Hawaiian Islands' exceptional distributions, and distributions involving the Caroline Islands, which serve as stepping stones for dispersal onto the Pacific Plate.
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavemosa (Linnaeus). Library of Congress Cataloging in Publication Data Springer, Victor Gruschka, 1928- Pacific Plate biogeography, with special reference to shorefishes. (Smithsonian contributions to zoology ; no. 367) Bibliography: p. Supt. of Docs, no.: SI 1.27:367 1. Fishes—Indo-Pacific Region—Geographical distribution. 2. Biogeography—Indo-Pacific Region. 3. Fishes—Pacific Ocean—Geographical distribution. 4. Geology—Pacific Ocean. I. Title. II. Title: Shorefishes. III. Series. QL1.S54 no. 367 [QL623.98] 591s [591.92'5] 82-600146 AACR2 Contents
Page Introduction 1 Acknowledgments 7 Methods 7 Distribution Maps 7 Terminology 8 Pacific Plate Distributions 8 Family-Group Names 9 Classification 9 Abbreviations 9 Organization of the Study 9 Distributions of Indo-Pacific Organisms 9 Shorefishes 9 Other Organisms 105 Phylum MOLLUSCA 105 Class GASTROPODA 105 Class PELECYPODA 108 Class CEPHALOPODA 108 Phylum ARTHROPODA 109 Class CRUSTACEA 109 Class INSECTA 113 Phylum CHORDATA 114 Class MAMMALIA 114 Plants 115 Discussion 117 Numbers of Shorefish Families 117 Numbers of Shorefish Genera and Species 119 Boundary Effect between Pacific Plate and Indo-West Pacific 120 Reality Hypothesis 121 Extinction Hypothesis 124 Endemism 127 Endemicity of Shorefish Families and Genera 127 Endemicity of Shorefish Species 128 Hawaiian Islands and North-South Disjunct Distributions on the Pacific Plate 130 Easter Island 136 Marquesas Islands 139 Geological History of the Pacific and Philippine Plates and Adja- cent Margins as Related to Formation of Types 1 and 2 Endemics 143 Epilogue 148 iii SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Addendum 149 Appendix 1: The Andesite Line 150 Appendix 2: Tables 153 Table A 153 Table B 157 Table C 157 Literature Cited 162 S FOR PACIFIC PC.A'l'F. HIGCKOCNAPH\
Page 10, Fig. 5. Arantrhurua arhil)e.i has been reported from th* southern tip oi ba)u Calltornla based on a aight record (D. A. Thomson, et al., Iy7d, Reoy Fi ahea o£ the S«?a_ o£ Corte?)
Page 15, col. 1, line 14. replace tarnycrphalua with
Page 25. Replace fcranchioategicUe with Latllidae
Page 27. Under Curacanthidae. Mizuno and Tooinaga were in errori carac&nchlda are not present at the Galapagos (M. and T, In lite, to W.N. Eschmeyer, pers. coum.) Page 28, col. 1, penultimate line. Replace carupodid with carapid Page 60. Replace Monocentrldae with Honocentrididae
Page 63. Under Nomeldae, 7th and Uth lines iron bottom delete the three words "eastern Atlantic and"
Page 88, Figure 39 caption. Delete sentence, "While this study was in press..." Sarrta is not known from Fili.
Page 91. Under Scorpididae. Delete Parapaetno Lbelongs in EphlppldldaeJ Page 100. Under Torpedlnlfornes and figure 44. 1 missed a report or the occurrence of electric rays on the Nazca Hidqe, Nazca Plate. Page 149, right column, 4th line, replace corroroborare with corroborate. Page 165. Under Chubb, replace 1*74 with
Pacific Plate Biogeography, with Special Reference to Shorefishes
Victor G. Springer
Introduction The Pacific Plate (Figures 1,2) is the largest of effects for the distribution of Pacific Basin orga- all the earth's lithospheric plates and occupies nisms. most of the area that has been referred to as the My study is concerned primarily with the dis- Pacific Basin (Figures 3, 4). Although the Pacific tributions of the shorefishes of the Indo-Pacific as Basin has been the subject of numerous biogeo- these distributions relate to the Pacific Plate. I graphic studies, there are only two studies (Sprin- propose to demonstrate that the Pacific Plate ger, 1981a; Rotondo et al., 1981) that have al- defines a biogeographic region of major signifi- luded to the Pacific Plate as a separate biogeo- cance, and I will hypothesize the biological and graphic unit. I will not recount the Pacific Basin geological factors that produced the region. I literature here, but refer the reader to two studies believe that study of the Pacific Plate provides that together summarize the subject admirably. new insights into the patterns of organismal dis- Van Balgooy (1971) gave a comprehensive and tributions in the Indo-Pacific, both aquatic and scholarly rendering of the general literature on terrestrial. From this it follows that many of the Pacific Basin biogeography in his analysis of Pa- constraints to dispersal ordinarily invoked to ex- cific plant distributions. The main deficiencies of plain only the distributions of endemic terrestrial van Balgooy's treatise are the absence of reference and freshwater organisms are equally important to the importance of plate tectonics (a subject in explaining the distributions of endemic, shal- probably too new for him to have incorporated) low-dwelling marine organisms. and the use of a solely phenetic approach to The day I began to write this introduction, I relationships. Zoogeographers seem to be una- received a new issue of American Scientist that ware of van Balgooy's work, which merits wider included an article on chance dispersal by attention than it has received. Kay (1980), who Carlquist (1981). In that article (p. 415) was the did not cite van Balgooy, published an excellent following statement, "Those who investigate or- essay on the development of ideas concerning ganisms capable of long-distance dispersal have Pacific Basin biogeography and included a brief not adopted the methods of vicariance biogeog- discussion of the importance of plate tectonic raphy." If this is true, my study is an exception; many shorefishes appear to be so capable. The
Victor G. Springer, Department of Vertebrate Zoology, National land masses of the Pacific Plate are islands (Figure Museum of Natural History, Washington, D.C. 20560. 2), many of which are highly isolated, and I will
1 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
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rely heavily on vicariant events to develop hy- potheses to explain endemism among these is- lands. Carlquist apparently thought he was pointing out a weakness in the methodology of vicariance biogeography (Nelson and Platnick, 1981; hence- forth, I refer to the subject as vicariography), but he failed to realize that dispersal is not the ma- terial of that discipline. The materials of vicariog- raphy are the geological and physical factors that contributed to the geographical isolation of biotic populations and permitted those populations to speciate. The dispersal that expands the range of a species becomes of interest to a vicariographer when the dispersal is interrupted. The interrup- tion is the vicariant event, and the cause of that interruption needs to be explained. It is a premise of vicariography that a vicariant event affects a spectrum of organisms that are not necessarily related phyletically and do not necessarily have the same general ecological requirements. If an island exhibits high endemism in both its land snails and shorefishes, for instance, it is more informative and economical to search first for a general explanation of the isolation of the island's biota than it is to search for the possible factors that might have pertained to a single species or ecological grouping of species. Birds might have populated an island with land snails, but they certainly did not populate it with shorefishes. It is possible, of course, that the endemism of the land snails and shorefishes might have entirely different explanations, and if they do, the vica- riographer is more likely to determine this than is the dispersalist, because the methodology of vicariography allows for the falsification of its hypotheses, and even defines the evidence for dispersal (sympatry of sister taxa). I know of no methodology of dispersalism that might lead to falsification of its hypotheses or alternate (vicari- ance) interpretations. Knowledge of sister-group relationships and
FIGURE 3.—Topography of the Pacific Basin (from B.C. Heezen and M. Tharp, World Ocean Floor [chart]. United States Navy, Office of Naval Research, 1977). SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
AGE OF THE PACIFIC OCEAN BASIN (Based on information contained in B.C Heezen and D.J. Fornari, IS7S. Geological Map of ttw Pacific Ocean.) LEGEND I Pleistocene 5. Eocene 2. Pliocene 6 Paleocene 3 Miocene 7. Upper Cretaceoui 4 Oligocene 8. Lower Cretaceoui 9 Upper Juraeeic
[Continental shelf 8k slope
FIGURE 4.—Age of the Pacific Ocean Basin. distributions are of the utmost importance for structing my hypotheses I have had to rely on the vicariographic studies. If one knows the cladisti- inferences and intuitive assessments of relation- cally determined closest relatives (sister groups) ships as given by authors, and on some verbal of endemic taxa, and the distributions of those assessments made at my request. That these state- relatives, one can narrow the geographic and ments of relationships tend to support my find- temporal search for the pertinent vicariant events. ings based solely on areas of endemism is not Unfortunately, few cladistic studies of Indo-Pa- surprising to me, but I recognize the deficiences cific organisms are available, and for species of of my study and look forward to its correction, Indo-Pacific fishes, none (however, see Vari, 1978, elaboration, falsification, or corroboration—by for an Indo-West Pacific fish family). In con- others. NUMBER 367
Acknowledgments Tyler, Richard P. Van, Gilbert L. Voss, Richard C. Wass, George E. Watson, Alwyne C. Wheeler, Numerous individuals have provided informa- Peter J. Whitehead, Austin B. Williams, Jeffrey tion or other assistance that has been incorpo- T. Williams, Richard Winterbottom, George R. rated into my study; others listened patiently to Zug. my discussions and constructively challenged my The base map used for most of the distributions ideas or offered suggestions for improvement of was prepared by August E. Spreitzer. Jack R. certain sections of the manuscript. To all of them Schroeder masterfully rendered Figure 3, and I here extend my sincere appreciation: Gerald R. Penelope K. Hollingsworth, Figures 61 and 63. Allen, K. Amaoka, William D. Anderson, Jr., Complete drafts of the manuscript were gen- Albert H. Banner, Hans Bath, Marie Louise erously reviewed and constructively criticized by Bauchot, Ross T. Bell, Adam Ben-Tuvia, Mari- Gareth Nelson, John E. Randall, and Richard H. nus Boeseman, James E. Bohlke (deceased). Kurt Rosenblatt. Especial appreciation goes to my ed- Bruwelheide, Warren E. Burgess, Bruce A. Carl- itor, Barbara T. Spann, who found innumerable son, John H. Caruso, E.H. Chave, Barry Cher- ways to improve the manuscript and bring it to noff, Daniel M. Cohen, Bruce B. Collette, Leon- a publishable state. ard J.V. Compagno, Roger F. Cressey, William Acknowledgment is accorded Elsevier Publish- P. Davis, Charles E. Dawson, Guido Dingerkus, ing Company for permission to reproduce the Terry Erwin, William F. Eschmeyer, Isabel P. information in Figure 62. Farfante, Norma Feinberg, Ross M. Feltes, Wil- Grants for fieldwork from the Max and Victo- liam L. Fink, Thomas H. Fraser, Warren C. ria Dreyfus Foundation, the Lewis and Rosa Freihofer, John A.F. Garrick, Ray W. George, Strauss Memorial Fund, and the Smithsonian Robert H. Gibbs, Jr., J. Gill, Janet R. Gomon, Scholarly Studies Program enabled me to obtain Martin F. Gomon, Patricia H. Hansen, Phillip C. important distributional data on fishes. Heemstra, John B. Heiser, Dannie A. Hensley, Above all, the contribution of John E. Randall Robert W. Hiatt, Douglas F. Hoese, Walter stands out. His incomparable knowledge of Indo- Ivantsoff, E. Dale Jackson (deceased), Susan Jew- Pacific fishes, his publications, and his collections, ett, G. David Johnson, Robert K.Johnson, Harry together with his unstinted imparting of infor- T. Kami, Robert Kanazawa, E. Allison Kay, mation, permeate my study. To him I dedicate Leslie K. Knapp, Alan J. Kohn, Karl F. Koop- this study. Thank you, Jack, even though you man, Louis S. Kornicker, Ernest A. Lachner, continually encouraged me to shelve it and return Harry S. Ladd, Raymond S. Manning, John D. to revising genera. You may well have been cor- McEachran, R.J. Mackay, John E. McKosker, rect. William G. Melson, Henry W. Menard, Robert Rush Miller, Robert F. Myers, Izumi Nakamura, Methods Gareth Nelson, Dan Nicholson, James N. Norris, DISTRIBUTION MAPS.—The distribution maps N.V. Parin, David L. Pawson, Theodore W. include delineations of the major lithospheric Pietsch, Winston Ponder, Stuart G. Poss, Chris- plates and the distributions are based on infor- topher Powell, John E. Randall, Harald A. Reh- mation taken from the pre-1982 literature (in- der, Clyde F. Roper, Donn E. Rosen, Richard H. cluding several studies in press), specimens I ex- Rosenblatt, Joseph Rosewater, W.C. Ruark, amined in museum collections (particularly the Barry C. Russell; Joseph L. Russo, Seymour O. National Museum of Natural History (USNM), Schlanger, James B. Shaklee, Takeshi Shimuzu, Australian Museum, California Academy of Sci- Eli Silver, William F. Smith-Vaniz, August E. ences, Academy of Natural Sciences of Philadel- Spreitzer, Michael J. Sweeney, Leighton R. Tay- phia, and Bishop Museum), and information sup- lor, W. Ralph Taylor, Thomas B. Thorson, En- plied by colleagues based on their unpublished rico Tortonese, Ethelwyn Trewavas, James C. studies or examination of collections under their 8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY care. In most instances restricted (exact) localities lagigedos, Clipperton, Galapagos, San Felix, San were plotted, but it will be recognized that the Ambrosio, Juan Fernandez, etc.) of the Western diameter of the symbols used to indicate localities Hemisphere. cover a radius of 50-100 km on the maps. In some Eastern Pacific: The eastern portion of the instances I have had to indicate generalized dis- Pacific Ocean not included in the Indo-Pacific. tributions, but I prefer, and strongly recommend Western Pacific: The Pacific ocean west of to others, plots of specific localities in compiling the western margin of the Pacific Plate (mostly distribution maps. Usually, no attempt was made composed of inland seas: South China Sea, Ara- to verify identifications or locality records taken fura Sea, Coral Sea, etc.); depending on the from the literature unless there was reason to context used, may also include the area along the doubt their validity (for instance, a conspicuous western margin of the Pacific Plate (see "Pacific separation of a locality record for a taxon from Plate Distributions"). all other records for the taxon, especially if the Indo-West Pacific: Indian Ocean plus "west- record was on the Pacific Plate). After investiga- ern Pacific," not synonymous with Indo-Pacific. tion, many of the questionable locality records The area of the Indo-Pacific that is always ex- proved to be erroneous or suspicious on additional cluded from the Indo-West Pacific is the non- grounds. A number of these emanated from spec- marginal portion of the Pacific Plate. imens that were procured for the now defunct PACIFIC PLATE DISTRIBUTIONS.—Except as Museum Godeffroy of Hamburg and were re- noted, I consider any taxon that occurs on the ported on by A. Giinther and R. Kner in the late nonmarginal portions of the Pacific Plate to have 19th Century. If I did not accept a locality record a Pacific Plate distribution. If such a taxon is I give my reasons in the discussion under the limited to the Pacific Plate or to the Pacific Plate pertinent family account. Although I attempted and localities on other plate margins adjacent to to make the distribution maps exhaustive, I rec- the Pacific Plate margin, it is considered to be ognize that I probably missed numerous records. endemic to the Pacific Plate. On the other hand, I personally plotted and compiled the informa- Indo-West Pacific taxa that reach their eastern tion for most of the maps, but the data for a few distributional limits at localities on the western were furnished by colleagues, and a few others margin of the Pacific Plate are not considered to were compiled by assistants under my close su- have Pacific Plate distributions. The Pacific Plate pervision. is subducting the Indian-Australian Plate, and TERMINOLOGY.—Because my usage of certain some portions of the latter plate are situated well terms varies somewhat from traditional usage, east of the Pacific Plate margin; for instance, the these terms are defined below. continental shelf of New Zealand, including the Shorefishes: In general, warm-temperate to Chatham Islands. Taxa occurring on the New tropical fishes dwelling at depths to about 100 m Zealand continental shelf do not have a Pacific close to shore. Shallow-dwelling fishes that are Plate component in their distribution unless they typically restricted to the offshore oceanic (epi- occur otherwise, and nonmarginally, on the Pa- pelagic) realm (for instance, Molidae) are ex- cific Plate. cluded, as are some typically deep-dwelling forms Easter Island is located on the Nasca Plate (for instance Myctophidae) that often occur at close to that plate's western margin, which is shallow depths near shore. being formed at the crest of the East Pacific Indo-Pacific (not synonymous with Indo-West Ridge (as is also the eastern margin of the Pacific Pacific): The Indian Ocean, including contig- Plate). Because most Easter Island genera occur uous seas, and the Pacific Ocean as far east as otherwise only in the Indo-Pacific, and are on the Easter Island, but excluding the area occupied by Pacific Plate, I recognize Easter Island taxa as the coast and offshore islands (Guadalupe, Revil- part of the Pacific Plate assemblage, with one NUMBER 367 exception: eastern Pacific taxa (for instance, the ognized in the ichthyological literature, and fish family Girellidae) that reach their western where I differ, I cite references, colleagues, or distributional limits at Easter Island, are not other reasons for my usage. considered to be Pacific Plate taxa. Some Pacific ABBREVIATIONS.—The following abbreviations Plate taxa that have their western distributional are employed: FL = fork length; m.y. = million limits along the western margin of the Philippine years; SL = standard length; TL = total length; Plate, usually the northern portion of that mar- NMNH = National Museum of Natural History, gin, are considered to be Pacific Plate restricted. Smithsonian Institution; USNM = United States The reasons for this consideration are explained National Museum collections, Division of Fishes, in the discussions of Types 1 and 2 endemism. NMNH. FAMILY-GROUP NAMES.—With the exception of ORGANIZATION OF THE STUDY.—The study is Carapidae, I follow Steyskal (1980) for the spell- presented as follows: (1) consideration of shorefish ing of family-group names of fishes. Steyskal in- distribution patterns within each family and of forms me that his use of Carapodidae for this the distributions of other organisms; (2) a general family was based on an erroneous treatment of descriptive synthesis of shorefish distributions on the stem. and off the Pacific Plate, with discussion of hy- CLASSIFICATION.—I follow Compagno (1973a) potheses for explaining those distributions; and for the classification and composition of the elas- (3) discussions of patterns of endemism on the mobranchs (sharks and rays). Most of the other Pacific Plate and the formation of hypotheses to shorefish family groupings are those usually rec- explain those patterns.
Distributions of Indo-Pacific Organisms
Shorefishes examination of the records on other grounds tends to confirm the suspicion of error. In the discussions that follow, families known By treating the Pacific Plate as the "hole in a to occur within the warm shallow waters of the doughnut," one can define the Plate just as surely Indo-Pacific region are presented. Their arrange- by what is absent as by what is present. For this ment is alphabetical to facilitate locating them, reason I have presented distribution maps of nu- and follows the listing in Table A (Appendix 2). merous taxa (orders, families, genera, species) The information given focuses on how the family that reach the margins of the Plate but do not distributions, and those of the taxa within the extend beyond. In some instances, ichthyologists families, relate to the Pacific Plate. Attention is have stated that the absence of a taxon from given to presumed erroneous Pacific Plate distri- Oceania is a result of lack of collecting. I believe butions that surfaced in the course of my inves- that the discussions and distribution maps will tigations. These presumed erroneous distributions overwhelmingly demonstrate that such assump- are tests of the validity of my proposition that the tions are often unwarranted. Pacific Plate is a biogeographic unit. The weight One locality, the Hawaiian Islands, frequently of evidence for this proposition was so heavy that represents an area where valid exceptional occur- I was able to presume distributions might be rences of taxa appear nonmarginally on the erroneous only because they represented excep- Pacific Plate. I term each such occurrence an tional occurrences of a taxon that was otherwise "Hawaiian exception." Discussion of the Ha- distributed only on or off the Plate. Subsequent waiian Islands will be found in the section on 10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY endemism. Another exceptional area for the oc- donia (Calliblennius rubescens Aoyagi, 1954, new currence of taxa on the Pacific Plate is in the genus and species, from the Ryukyus, is a junior Eastern Caroline Islands chain, which I treat as synonym of B. fasciolatum)K See "Addendum." the "Caroline Islands conduit." The conduit is discussed in the section on boundary effect. ACANTHURIDAE The surgeonfishes comprise six genera (Ran- ACANTHOCLINIDAE dall, 1955a and pers. comm.) and about 70 species This family is restricted to the Indo-Pacific, of small- to moderate-sized (to about 700 mm where it comprises three genera and about five TL), free-swimming, reef-inhabiting fishes with species of small (largest species about 300 mm planktonic larvae. SL), rock or reef inhabiting forms. Acanthoclinus, Acanthurus contains about 35 species (Randall, with two species, occurs only in New Zealand, 1956, 1960, and in litt.; Shen and Lim, 1973, and possibly Australia (Glover, 1979:151). Acan- described three additional species, not recognized thoplesiops contains two species and ranges from here, based on what appear to be juvenile speci- southern India eastwards to the Marshall Islands. mens of described species) and is the only acan- Acanthoplesiops is poorly known, and only one thurid genus that is found in all warm seas. species, the diminutive A. hiatti, occurs nonmar- Twenty-eight species of Acanthurus occur in the ginally on the Pacific Plate, where it is recorded Indo-Pacific, of which three range into the eastern only from the Marshall Islands. Belonepterygion is Pacific as well. Of the 28 Indo-Pacific species, monotypic (B. fasciolatum) and ranges from south- eight are restricted to the Indo-West Pacific, and ern Japan and Taiwan (USNM collections) 20 occur nonmarginally on the Pacific Plate. One through the Philippines (USNM) to southern species, A. dussumieri, otherwise widely distributed Queensland, Lord Howe Island, and New Cale- in the Indo-West Pacific, has been reported on
FIGURE 5.—Distribution of two widely distributed, Pacific Plate species of surgeonfishes, Acanthurus achilles (solid circles) and A. leucopareius (stars). NUMBER 367 11 the Plate only from the Hawaiian Islands, but nonmarginally (eight ranging from the east coast J.E. Randall informs me he sighted a specimen of Africa, and one restricted to Japan and the in the Line Islands in 1968. The species can Hawaiian Islands). The other seven species are probably be considered as an Hawaiian exception restricted to various parts of the Indo-West Pa- with occasional waifs appearing elsewhere on the cific, and some are of questionable validity. Pacific Plate. Two species, A. achilles and A. leu- Prionurus (including Xesurus) has six species, of copareius (Figure 5), should possibly be considered which two are limited to the eastern Pacific, one as widely distributed Pacific Plate endemics that to the eastern Atlantic, and three to the western have undergone a relatively recent westward dis- Pacific (Smith, 1966a). Prionurus is the only acan- persal which has enabled them to reach the west- thurid genus that is absent from the nonmarginal ern margin of the Philippine Plate (see section on portions of the Pacific Plate. geology of Pacific and Philippine plates). Acanthu- Paracanthurus is monotypic (Randall, 1955b) rus achilles is known nonmarginally off the Pacific and ranges throughout the Indo-Pacific, from the Plate only from Cebu, Philippines (Rau and Rau, east coast of Africa to at least the Line Islands 1980), where it and A. leucopareius appear uncom- (Chave and Eckert, 1974), but it is absent from monly in the fish market (N. Rau, in litt.). The the Hawaiian Islands. distribution of A. leucopareius is also noteworthy Zebrasoma contains seven species. Randall because of its disjunct, antiequatorial nature (1955b) recognized six species, but now (in litt.) (Randall's 1976 report of A. leucopareius from Lord recognizes a seventh, Z. desjardinii, for the Indian Howe Island was in error; Marcus Island—on the Ocean populations of the species he treated as Z. Pacific Plate—was the intended locality; Ran- veliferum from the Indo-Pacific. Two other species dall, in litt.). (Z gemmatum, Z. xanthurum) are restricted to the Acanthurus nigroris also appears to be a widely Indian Ocean, and one species (Z. scopas) ranges distributed Pacific Plate endemic, although Ran- from the western Indian Ocean to the Society dall (in litt.) suggests that it may occur west of Islands. Zebrasoma rostratus is a Pacific Plate en- the Plate as a straggler (see Whitley, 1958, for a demic known positively only from the Tuamotu, record from the Great Barrier Reef). There is one Marquesas, Line, and Rapa islands. Zebrasoma other Pacific Plate endemic species of Acanthurus, Jlavescens should possibly be considered a Plate A. chronixis, known only from Kapingamarangi, endemic; it has been reported from Taiwan and Eastern Caroline Islands (Randall, 1960, and the Ryukyu Islands, along the western margin of pers. comm.; misidentified from New Guinea by the Philippine Plate, where it appears to be un- Burgess and Axelrod, 1975, fig. 418 = A. pyroferus). common, east to the Bonin, Mariana, Marshall, The intrageneric relationships of the Plate en- Wake, Marcus, Hawaiian, and Johnston islands. demic species of Acanthurus are unknown. In summary, there are five genera and 64 Indo- Ctenochaetus comprises six species (Randall, Pacific species of acanthurids, of which 39 species 1955c, 1960) and ranges throughout the Indo- occur on the Pacific Plate nonmarginally, and Pacific, one species also reaching the eastern Pa- possibly seven of these 39 are Plate endemics. cific. At least five of the species occur on the Pacific Plate nonmarginally, and one, C. hawaiien- ALBULIDAE sis, is a widely distributed Pacific Plate endemic, known from the Mariana, Marcus, Marshall, Ha- The bonefishes comprise two genera (Forey, waiian, Line, Austral, Society, Samoa, Tuamotu, 1973): Pterothrissus (sometimes placed in its own and Pitcairn Group islands. family, Pterothrissidae) with two, moderately Naso (including Axinurus and Callicanthus) con- deepwater species (one known from off Japan, the tains about 16 species (Srm'th, 1966a; Randall other from off west Africa), and Albula, with and Struhsaker, 1981; J.E. Randall, in litt.), of perhaps as many as six species, several of which which at least nine extend onto the Pacific Plate have been confused with one of the two currently 12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY recognized species, A. vulpes (Shaklee and Ta- of including the Ammodytidae among the warm- maru, 1981, and Shaklee, pers. comm.). water shorefish families is questionable. Albula is distributed circumglobally (as are, perhaps, two of its species), and occurs close to ANGUILLIDAE shore, where it is avidly sought by sport fishermen; larvae are taken pelagically. The freshwater eels consist of a single genus, Anguilla, with 16 species (Ege, 1939) that spawn in deepwater well out to sea, and have long-lived AMBASSIDAE planktonic larvae. The young ascend rivers and attain maturity in freshwaters. The largest species attains a length of about 2 m. The anguillids are The ambassids are a family of small fishes in widely distributed in the tropical and temperate need of revision. There are 40 nominal species waters of the Indo-Pacific and north Atlantic, but (R.P. Vari, pers. comm.), but perhaps only 20 are absent from the eastern Pacific, most of the valid species, which usually occur in estuarine west coast of Africa, and almost all of the Atlantic and mangrove areas. The ambassids have been coast of South America (for a world distribution reported to range from the east coast of Africa to map see Schmidt, 1925; for distribution maps of the Philippines, southern Japan, Guam, and Sa- the Indo-Pacific species see Ege, 1939). moa, on the margin of the Pacific Plate, and are known to occur on the Pacific Plate nonmargin- Fourteen of the Anguilla species are limited to ally only at Ponape (Ambassis; USNM collec- the Indo-Pacific, and of these, only three (none tions). The presence of ambassids at Ponape is endemic) occur on the Pacific Plate nonmargin- evidence favoring the operation of the Caroline ally. Although Anguilla is widely distributed on Islands conduit. the Pacific Plate, the genus is notably absent from the Hawaiian Islands.
AMMODYTIDAE ANOMALOPIDAE The sand lances are a small family of small The flashlight fishes comprise four monotypic fishes that are antitropically distributed: widely genera of small (to about 300 mm TL), reef- distributed in the warm-temperate marine waters inhabiting fishes. Two genera are restricted to the of the northern hemisphere, but known only in Indo-Pacific, one to the eastern Pacific, and one the Indo-Pacific in the southern hemisphere, to the Caribbean. Records of flashlight fishes are where they occur in southern Africa, southern relatively few because, until recently, the habits Australia (as far north as southern Queensland), of the species, which reside in deep crevices in the and Pitcairn Island (where they were collected daytime and enter shallow waters only on moon- along shore; J.E. Randall, in litt.). Aside from less nights, make them generally unavailable to Pitcairn, they are known on the Pacific Plate collectors. On the Pacific Plate, nonmarginally, nonmarginally only from the Hawaiian Islands, they are known only from Kwajalein (Marshall where a few specimens of Bleekeria gilli have been Islands; Photoblepharon; J.E. McCosker, pers. obtained from tuna spewings (Gosline and Brock, comm.), Rarotonga (Cook Islands; Anomalops; 1960); hence, the species is probably pelagic off- Seale, 1906), and the Tuamotus (Anomalops; shore. Bleekeria gilh also is reported from the Giinther, 1876). Both Anomalops and Photoblepharon Galapagos (Seale, 1940) and Baja Calfornia occur in the Banda Islands, Moluccas, and I have (Duncker and Mohr, 1939). Fowler (1928) be- seen and participated in collections of both genera lieved the unknown type locality of B. gilli was on the same night on the same reef at Mactan probably the Bonin Islands. The appropriateness Island, Philippine Islands. Anomalops has not been NUMBER 367 13 reported from the Indian Ocean, but Photoble- ing in the Society Islands, did not report Histrio pharon ranges westward to the Red Sea (which from those islands. T. Pietsch, who has revised has an endemic subspecies of Photoblepharon) and the antennariids, saw no material of Histrio from Comoro Islands. the Societies. Without reconfirmation, I consider the presence of Histrio in the Society Islands to be unsubstantiated. ANTENNARIIDAE It is noteworthy also that R.C. Wass, who has The frogfishes comprise 12 genera and 40 spe- spent several years in the Samoan Islands and cies of small- to moderate-sized fishes (to about concentrated on obtaining records and specimens 360 mm SL) that are (with one exception) of all Samoan fishes, has not obtained Histrio in benthic, shallow to moderately deep dwelling, Samoa. Histrio is known at Guam (on the eastern and occur primarily on rocky and coral reefs. A margin of the Philippine Plate) from only two few species may occasionally enter brackish wa- specimens taken about ten years apart (R.S. ter. Ten genera are restricted to the Indo-Pacific Myers, in litt.) in spite of many years of intensive and about 30 species occur in that region, but collecting at that island. There are no records of only nine species occur on the Pacific Plate non- Histrio from Palau. marginally, and of these, only Antennarius duescus, Eydoux and Souleyet (1842) described Chiro- known from two specimens from the Hawaiian nectes barbatulus, a synonym of//, histrio, and stated Islands, is apparently endemic to the Plate. (T. that they were unable to give the locality of origin Pietsch, in litt., and pers. comm.) for their specimen. Le Danois (1964), without One antennariid, Histrio histrio, occurs only in explanation, listed the type locality of Chironectes floating Sargassum, and has the potential for wide- barbatulus as Hawaii. Dr. M.L. Bauchot (in litt.) spread distribution. Histrio histrio is often cited as reconfirmed the lack of associated locality data cosmopolitan in warm seas, but its distribution with the holotype of C. barbatulus. Schultz (1957) appears to be restricted to the Atlantic and Indo- reported H. histrio from the Hawaiian islands West Pacific. There are four reports of H. histrio based on specimens labelled as having been taken from the Pacific Plate and one from the eastern by the Wilkes Expedition of 1838-1842 at Maui. Pacific (Galapagos). I consider all five suspect. There are no subsequent reports of Histrio from Fowler (1928, 1931) reported H. histrio from Raia- the Hawaiian Islands in spite of many years of tea and Huahine, both Society Islands, on the intensive collecting in those islands, even though Pacific Plate, based on listings of Antennarius mar- Sargassum is present. If Histrio occurs in the Ha- moratus and A. multiocellatus in Shmeltz (1877, waiian Islands it probably does so as a straggler 1879; references examined for me by W.F. Smith- brought in by ocean currents (Kuroshio Exten- Vaniz). Shmeltz's papers are catalogs of the Mu- sion) from Japan. seum Godeffroy collections, and the catalogs con- Schultz (1957) also reported Histrio from the tain no descriptive material or illustrations. Fishes Galapagos based on specimens first cataloged at in the Godeffroy collections were obtained by USNM in 1877, and listed as having been col- commercial collectors and boat captains, and lo- lected by a Captain Heerendeen (or Herrendea, calities assigned many Godeffroy specimens have inscription unclear) of Woods Hole, Massachu- proved to be well outside the established ranges setts. There are no other records of Histrio for the for the species and not confirmed by prior or eastern Pacific. In accord with R.H. Rosenblatt subsequent collecting. The location of the God- (pers. comm.), I do not believe that the species effroy specimens supposedly representing Histrio occurs in the Galapagos or elsewhere in the east- is not known. Fowler gave no reasons for recog- ern Pacific. nizing the Godeffroy specimens as H. histrio, and It is of interest that Sargassum is reported from Randall (1973), who spent many months collect- many of the high islands (including at least Ha- 14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY waii, Tahiti, Ponape) on the Pacific Plate, but Plate from the west. He notes that there are with the exception of Laysan Atoll (Hawaiian endemic apogonids in various Pacific Plate island Islands) is noticeably absent from low islands groups (Hawaiian, Marquesas, Easter), but that (Doty, 1954; Tsuda, 1976). he knows of no widely distributed Pacific Plate endemics. There are probably no more than 50 APLOACTINIDAE species of Apogonidae on the Plate. The apogonids demonstrate that mouth brood- The velvetfishes comprise about 15 genera and ing is obviously not a factor limiting distributions, about 25 species that appear to live on rocky or although the absence of the mouth-brooding fam- coral rubble bottoms, bottoms with coralline al- ilies Ariidae and Opistognathidae from the Pa- gae, or among vegetation (Poss and Eschmeyer, cific Plate might be ascribed to such a habit. 1978, 1979, 1980; Eschmeyer and Allen, 1978; Eschmeyer and Dor, 1978) at depths from near ARACANIDAE shore to 510 m, and are found only in the Indo- West Pacific; none occurs on the Pacific Plate. The aracanid trunkfishes comprise eight genera and about 12 species of small (to about 30 APOGONIDAE mm TL), often moderately deep-dwelling, benth- ically associated fishes. The family has a primarily The cardinalfishes comprise about 20 genera antitropical distribution, is restricted to the Indo- and over 200 species of small (to about 553 mm West Pacific, except for its exceptional occurrence SL for Epigonus, a pelagic genus, but most reef in the Hawaiian Islands, and is questionably forms are well under 200 mm), predominantly included here as a shorefish group. Aracanostracion coral-reef fishes, with some deep-dwelling or pe- is known only from southernmost Africa; Aracana, lagic species and some estuarine and freshwater Anoplocapros, Caprichthys, and Capropygia are lim- species. A revision of the most speciose genus, ited to the cooler coasts of Australia; Polyplacapros Apogon, is greatly needed. As far as known, the is known from a seamount east of New South species are all mouth brooders, and larvae are Wales (132 m deep) and along the Norfolk Ridge restricted to shelf waters (T.H. Fraser, pers. (ca. 167°33'E, 28°-33°S; 79-161 m); Strophiu- comm.). The family is distributed circumglobally, richthys occurs in Queensland (at depths to about with most species occurring on the reefs of the 64 m) and has been reported erroneously from Indo-Pacific. Many of the species are wide rang- the Solomon Islands (Randall, in press b); and ing; for instance, Apogon kallopterus, a known Kentrocapros (presently monotypic) is known from mouthbrooder, ranges from the east coast of Af- Japan (Sagami Bay south to Kyushu), Korea rica to the Hawaiian, Marquesas, and Tuamotu (Pusan), Taiwan (20-30 m depth), the Hawaiian islands (Fraser, pers. comm.). Even more spec- islands (from at least 104 m depth), and presum- tacular, Apogon evermanni, a moderately deep- ably, from off eastern Australia (Fujii and Uyeno, dwelling reef form (available to SCUBA- 1979). equipped divers), is known from the Hawaiian The distribution of the Aracanidae, which is Islands (Hawaii, Oahu), the Marquesas (Nuku often considered as a subfamily of the Ostraci- Hiva, Fatu Hiva), Kwajalein (Marshalls), Florida idae, contrasts strongly with the distribution of Island (Solomons), Ambon (Moluccas), Maldives, the Ostraciidae (sensu stricto), which occur cir- Chagos, Cocos-Keeling, and Mauritius (all In- cumglobally in primarily warm, shallow seas. dian Ocean), and Cozumel, Curacao, and Baha- mas (all western Atlantic; Randall and Bohlke, 1981). T.E. Fraser informs me that there is a ARIIDAE rapid dropoff in the number of apogonid species Thhe marine catfishes comprise about 140 spe- as one crosses the western margin of the Pacific cies of moderate size that are distributed circum- NUMBER 367 15 globally, except that none occurs on the Pacific species of Aulostomus are slender, less than 1 m Plate other than marginally. The eastern limits TL, free swimming, and reef inhabiting. of the family in the Indo-Pacific are almost, if not entirely, identical with those of the croakers, BALISTIDAE Sciaenidae (Figure 38). As far as known, all the (not including Monacanthidae) species have large eggs and are mouthbrooders (see discussion under Apogonidae), and most The triggerfishes comprise 12 genera (Mat- spend some portion of their life history in estu- suura, 1980; 1981) and about 30 moderately large aries. Several species are restricted to freshwater, (to about 750 mm TL), free-swimming species and it is probable that the family is derived from with pelagic young. All the genera, except Xeno- a freshwater ancestor (W.R. Taylor, pers. comm.). balistes, and most of the species occur in the Indo- There are three, notably erroneous, distribu- Pacific; however, Batistes, otherwise restricted to tion records for ariids in the Indo-Pacific. Arius the Atlantic and eastern Pacific, occurs in the tachycephalus was described by Giinther (1864) Indo-Pacific only in the Hawaiian Islands and from Hawaii. This species is a common one lim- only as a rare straggler (the common eastern ited to the eastern Pacific. Kner and Steindachner Pacific species Balistes polylepis, which has usually (1866) described Arius graeffi from Samoa, based been misidentified as B. fuscus in Hawaiian ref- on a Museum Godeffroy specimen, and Schmeltz erences; J.E. Randall, pers. comm.; I have not (1879, reference examined for me by W.F. Smith- included Balistes in Appendix 2, Table C). All Vaniz) listed Arius thalassinus from the Tonga but one of the native Indo-Pacific genera, Aba- Islands, also based on a Museum Godeffroy spec- listes, monotypic, occur on the Pacific Plate, but imen. No other records of ariids are known for not all the Indo-Pacific species in these genera Samoa and Tonga, and there is none for the occur on the Plate. nearby, and larger, Fiji Islands. Odonus and Balistapus are monotypic, widely distributed in, and restricted to, the Indo-Pacific. Xenobalistes, also monotypic, is known from a ATHERINIDAE single specimen from the stomach of a marlin The silversides are a large, circumglobal family caught off the Mariana Islands. of small, free-swimming, close-shore fishes. Most Canthidermis contains five species (Fedoryako, of the species are marine, but some are strictly 1981), one (C. maculatus) present on the Pacific freshwater (Celebes, North America) inhabitants. Plate and distributed in all oceans but the eastern There are four marine genera with about 18 Pacific; one (C villosus) known only from the Gulf species in the Indo-Pacific. All the genera, but of Aden, one (C. rotundatus) limited to the Indo- only six of the species, occur nonmarginally on Pacific), one (C. willughbeii) restricted to the east- the Pacific Plate, and one of the species is endemic ern Pacific, and one (C sufflamen) restricted to the to the Hawaiian Islands. (W. Ivantsoff, pers. western Atlantic. comm.). Pseudobalistes contains three species, two (P. fus- cus, P. flavimarginatus) occurring on the Plate as
AULOSTOMIDAE well as widely distributed in, and restricted to, the Indo-Pacific, and one (P. naufragium) restricted The trumpetfishes comprise a single genus, Au- to the eastern Pacific. lostomus, and three species (Wheeler, 1955): one Melichthys contains three species (the first two limited to the western Atlantic, one to the eastern being present on the Plate): one is circum- Atlantic, and one to the Indo-Pacific (including tropically distributed, one is limited to the the Pacific Plate nonmarginally) and eastern Pa- Indo-Pacific and eastern Pacific, and one is lim- cific. In the latter area it is known only from ited to the Indian Ocean (Randall and Klausew- offshore islands (Rosenblatt et al., 1972). The itz, 1973). 16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Abalistes stellatus is a common, widely distrib- uted in, the Indo-Pacific, and one restricted to uted triggerfish that occurs in a wide variety of the eastern Pacific. marine habitats, including coral reefs and over Rhineacanthus is restricted to the Indo-Pacific sandy and muddy bottoms. It ranges from the and comprises about six species (one undescribed, Red Sea and southeastern South Africa eastwards J.E. Randall, in litt.), of which at least three to Japan (Honshu), New Guinea, New Ireland, occur on the Pacific Plate nonmarginally. and Fiji, but is absent from the Pacific Plate nonmarginally. BATRACHOIDIDAE Xanthichthys comprises five species (Randall, Matsuura, and Zama, 1978). One species (X. The toadfishes comprise 19 genera and 64 spe- ringens) is restricted to the western Atlantic, and cies of small to moderate size, few, if any, attain- four are found in the Indo-Pacific, but one of the ing a standard length of 400 mm (B.B. Collette, four (X. lineopunctatus) has not been taken on the pers. comm.). The species are all primarily Pacific Plate nonmarginally. Another (X. mento) benthic in habit and most, if not all, occur at is distributed from the eastern Pacific only as far shallow depths during some period of their life west as the Ryukyu and Izu islands along the history. Gilbert (1968) quotes a responsible report western margin of the Philippine Plate. The dis- of an unidentified species of Porichthys that was tribution of these last two species emphasizes the seen in numbers at the surface of waters at least biogeographic boundary between the continental 1000 fathoms (1829 m) deep 30 miles (50 km) off and oceanic plates. the Pacific coast of Panama. Gilbert may have Balistoides contains two species, both present on been referring to the same incident as mentioned the Plate and widely distributed in and restricted in Walters and Robins (1961) of Porichthys being to the Indo-Pacific. rafted on logs 25 miles (40 km) from land in the Sufflamen comprises four species, three present Gulf of Panama. Adult Porichthys may also occur on the Plate and restricted to, and widely distrib- at very shallow depths. Variously, the species of
FIGURE 6.—Distribution of the toadfish family Batrachoididae. NUMBER 367 17 batrachoidids occur in, or are restricted to, a sey and Collette's figure 177, which might other- variety of habitats: freshwater rivers, estuaries, wise appear to indicate an example of an Ha- coral reefs, rocky or shelly marine bottoms, waiian exception distribution. muddy sand, and in the intertidal, where they Strongylura, the most speciose genus of needle- may even be stranded out of water for several fishes, comprises 13 species, of which two are hours without harm. Eggs are deposited in, and strictly freshwater, one estuarine, and ten marine, attached to, almost any type compartment: jars, many of which ascend freshwaters. Five species shells, holes, cans, crevices. Nests are guarded and of Strongylura (four marine, one estuarine) occur the young, upon hatching, are demersal. in (and are limited to) the Indo-Pacific, but only Mead (1970) believed that the toadfishes are one, a marine species {Strongylura incisa), extends distributed worldwide. Toadfishes are widely dis- nonmarginally onto the Pacific Plate (Cressey tributed latitudinally and longitudinally, but are and Collette, 1971). Strongylura incisa, together notably absent from the Pacific Plate (Figure 6). with the four worldwide species (Ablennes hians, The largest numbers of genera and species of Platybelone argalus, Tylosurus acus, T. crocodilus), toadfishes are found in the western Atlantic make a total of five species of needlefishes present (seven genera, 25 species, plus three more species on the Pacific Plate. in Atlantic freshwater drainage systems). Only Aside from the two worldwide species of Tylo- three genera and about nine species occur in the surus, there are three other species of Tylosurus: T. Indo-West Pacific, excluding the highly restricted choram, known only from the Red Sea and north- five endemic genera from southernmost Africa western Indian Ocean, T. gavialoides from north- (Collette and Russo, 1981; Hutchins, 1976). The ern Australia, and T. punctulatus, from the Ma- apparent absence of toadfishes from most of the layan and Philippine area. The five species of Indian Ocean coast of Africa, and from the coasts Tylosurus, the five marine and estuarine species of of Madagascar and mainland China is surprising. Strongylura, and the monotypic Ablennes and Pla- tybelone comprise the 12 species of needlefishes BELONIDAE found in the Indo-West Pacific.
The needlefishes comprise nine genera and 30 BLENNHDAE species of epipelagic fishes found in freshwaters, estuaries, and marine habitats, both inshore and The blennies are a circumglobal family that offshore. Needlefishes have a maximum body comprises five tribes, 54 genera, and about 300 length ranging from 42 mm (two South American species of small (maximum adult sizes ranging freshwater species) to 950 mm for one worldwide from about 15 to 532 mm SL, but few exceed 150 marine species, Tylosurus acus, which consists of mm), predominently near-shore, benthic fishes. five well-differentiated subspecies (one western Most of the species are restricted to warm marine Atlantic, one Mediterranean, one eastern Atlan- waters, but a few species occur in temperate tic, one eastern Pacific, and one Indo-Pacific; the marine or warm freshwaters. As far as known, all Indo-Pacific subspecies extends eastwards to the the species are nest builders, and although pre- offshore eastern Pacific islands of Cocos and Re- juveniles of several species may occur planktoni- villagigedos; Collette and Parin, 1970). Needle- cally a short distance offshore, the unmetamor- fish eggs bear entangling filaments and are de- phosed young are generally taken with the adults. mersal unless attached to floating objects (sum- Individuals of a few species have been taken marized in Breder and Rosen, 1966). around floating sargassum or other objects, and Cressey and Collette (1971) give distribution one Indo-West Pacific species {Omobranchus punc- maps for many of the needlefish species. Collette tatus), of an otherwise Indo-Pacific restricted ge- (pers. comm.) has many more Indo-Pacific rec- nus, occurs in the Caribbean, possibly as a result ords for Tylosurus acus than are indicated on Cres- of an introduction (Springer and Gomon, 1975). 18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.—Distribution of numbers of genera and species in the tribes of the fish family Blenniidae in the Indo-Pacifica
Number of Genera in Genera on Genera endemic Tribe genera I ndo-Pacific11 Pacific Plate0 to Pacific Plate"
Blenniini 16 3 1 Salariini 26 24 14 Omobranchini 6 6 4 Phenablenniini 1 1 0 Nemophini 5 5 5
Totals 54 39 24
Number of Species in Species on Species endemic Tribe species Indo-Pacifich Pacific Plate' to Pacific Plate1 Blenniini 70" 1 0 Salariini 150a 141" 45* 25' Omobranchini 31 31 5 0 Phenablenniini 1 1 0 0 Nemophini 41 40 11 2
Totals 293 226b 62 27d
a Estimated. b Includes Easter Island. c Other than only marginally present, but includes Easter Island. d Twenty-one known: seven only at Hawaiian Islands, four only at Marquesas, two only at Easter Island, one only in Tuamotus, and seven more widely distributed. All of the tribes, 72 percent of the genera, and there is no specialization that unites its compo- 77 percent of the species occur in the Indo-Pacific nents. The presence of an unidentified species of (Table 1), and all but one of the tribes, Blenniini, Parablennius in the Hawaiian Islands (reported have their centers of taxonomic abundance in here for the first time based on specimens from that area. The Blenniini, in contrast, has its larg- Oahu I examined at the Hawaii Institute of est representation in the Western Hemisphere Marine Biology many years ago) constitutes an and Mediterranean, where the other tribes are example of an Hawaiian exception, pertaining, poorly represented or absent. The Salariini is in this case, to a tribe, genus, and perhaps species represented in the Western Hemisphere and east- that are otherwise absent from the Pacific Plate. ern Atlantic by only three genera: Entomacrodus, A revision of Parablennius (which includes Picti- which is circumtropical; Ophioblennius, which is blennius; H. Bath, in litt.) will be necessary before restricted to the Western Hemisphere and eastern it will be possible to identify the Hawaiian spe- Atlantic; and Scarticthys, which is restricted to the cies. eastern Pacific. The Nemophini is represented Springer (1967b) and Strasburg (1956) dis- outside the Indo-Pacific by only one of the 11 cussed the erroneous reports from the Hawaiian species of Plagiotremus, P. azaleus, which is endemic Islands of the eastern Pacific species Hypsoblennius to the eastern Pacific. The Omobranchini and sordidus and H. brevipinnis. Phenablenniini occur only in the Indo-Pacific Salariini: Alloblennius contains four species (except for the presumably introduced species of and is restricted to the western Indian Ocean and Omobranchus in the Caribbean). Red Sea (Springer and Spreitzer, 1978; H. Bath, Blenniini: This is the least specialized tribe of ms.). blenniids, and probably is not monophyletic, as Alticus is in need of revision, but ranges over NUMBER 367 19 much of the Indo-Pacific. An endemic species populations (McKinney and Springer, 1976). Ec- occurs in the Marquesas Islands on the Pacific senius species are known from depths of less than Plate (Smith-Vaniz and Springer, 1971). one to about 45 m. Only three species occur other Andamia is in need of revision. It ranges from than marginally on the Pacific Plate, and two of the eastern Indian Ocean to the western margin these, E. midas and E. bicolor, have the greatest of the Pacific Plate in the New Guinea region overall ranges of any of the species of Ecsenius. (Smith-Vaniz and Springer, 1971; Springer, un- Ecsenius midas (east coast of Africa to the Marque- published data). sas) is one of the few species of Ecsenius that rises Antennablennius, which has been revised by H. above the bottom and becomes free swimming; Bath (ms.), contains seven species and is restricted E. bicolor (Maldives to Marshall and Samoan to the Red Sea and Indian Ocean (Smith-Vaniz islands), however, is closely tied to the substrate. and Springer, 1971). The third Plate species, E. opsifrontalis (not free Atrosalarias is monotypic, but with two allopa- swimming) consists of several color-pattern pop- tric subspecies: one restricted to the Indian Ocean ulations. The population of E. opsifrontalis on the and Red Sea, the other ranging from Western Pacific Plate consists of a color-pattern type that Australia to well out on the Pacific Plate is restricted to the Plate and its margins (Springer (Springer and Smith-Vaniz, 1968; Smith-Vaniz and Bruwelheide, unpublished data). The other and Springer, 1971). color-pattern populations of E. opsifrontalis are Cirripectes, which ranges throughout the Indo- restricted to Pacific areas west of the Plate or on Pacific, is being revised by J.T. Williams, who the Plate margin. informs me that two (of the approximately 25) Entomacrodus comprises 22 species (several spe- species are widely distributed Pacific Plate en- cies consist of unnamed, but recognizably distinct demics: Cirripectes jenningsi from the Gilbert, To- populations; Springer, 1967a, 1972a, and unpub- kelau, Phoenix, Line and Tuamotu islands; Cir- lished data) that live in the shallows adjacent to ripectes variolosus, with representation in every ma- shore and in the intertidal. Of the 17 Indo-Pacific jor nonmarginal island group on the Plate (except species, 13 occur nonmarginally on the Pacific the Pitcairn Group and Hawaiian Islands) and Plate, and I consider nine of these to be endemic from Guam, the Palaus, and the Tonga Islands, to the Plate (including margins and Philippine all adjacent to the margin. Plate; Figures 7, 8). Two of the endemics, E. Cirrisalarias is monotypic and ranges from the cymatobiotus and E. sealei, are widely distributed. Comoro Islands, Indian Ocean, to Samoa, on the Springer (1972a) noted that a specimen of E. margin of the Pacific Plate (Springer and sealei, the only one known from New Guinea, had Spreitzer, 1978:1). See Medusablennius below. a different color pattern from all other available Crossosalarias is monotypic and is known from specimens of the species, and questioned if the the Ryukyu Islands south to the Moluccas and color pattern was normal. A second specimen, eastward to New Guinea and southeast to the exhibiting the same peculiarities of color pattern, Tonga Islands; hence, it is absent from the Pacific from Indonesia (Ternate, Halmahera; USNM Plate nonmarginally (Smith-Vaniz and Springer, collections) was recently obtained. Entomacrodus 1971; USNM). sealei, thus consists of two allopatric color-pattern Dodekablennos is monotypic and known only types that are separated along the western margin from Mauritius (Springer and Spreitzer, 1978). of the Pacific Plate. Entomacrodus caudofasciatus, Ecsenius comprises about 30 species (including which I do not include among the Plate endemics, undescribed forms) of predominently benthic embraces five color-pattern types (Figure 7), of coral-reef inhabitants (Springer, 1971, 1972a; which one is widely distributed along the south- McKinney and Springer, 1976). Each of two ern island chains on the Pacific Plate, and a species (E. opsifrontalis, E. oculus) consists of several second, which I reported (1967a) from two widely recognizably distinct, allopatric color-pattern separated groups of localities, might also prove to 20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 367 21 be Plate restricted if a reappraisal shows my Springer and Spreitzer, 1978, fig. 6; a record for lumping of the color patterns from the separated the Philippine Islands can now be added; localities to be in error. USNM). Only one species reaches the Plate mar- Springer (1967a) considered three species of gin. Entomacrodus to form a closely related group, Nannosalarias is monotypic and ranges from which is restricted to the Pacific Plate: E. cymato- Christmas Island, eastern Indian Ocean, east- biotus (widely distributed, but absent from the wards to New Guinea and the Tonga Islands, Hawaiian Islands and Easter Island), E. strasburgi adjacent to the margin of the Pacific Plate (known only from Oahu), and E. chapmani (known (Smith-Vaniz and Springer, 1971), and is known only from Easter Island). The group was not nonmarginally on the Plate only from Ponape defined on the basis of a shared specialization (USNM), which I consider evidence for operation and the sister species group was not proposed. of the Caroline Islands conduit. The three species are quite similar and I intui- Pereulixia is monotypic and known only from tively believe they form a monophyletic group. the southeastern coast of Africa (Smith-Vaniz Exallias is monotypic and ranges throughout and Springer, 1971). most of the Indo-Pacific (Smith-Vaniz and Sprin- Praealticus is in need of revision; it ranges from ger, 1971). the eastern Indian Ocean to well out on the Glyptoparus is monotypic (Smith-Vaniz and Pacific Plate. Springer, 1971) and ranges from the western In- Rhabdoblennius is in need of revision: it ranges dian Ocean to the western margin of the Pacific from the Philippines to well out on the Pacific Plate, and is known from the Pacific Plate non- Plate (Smith-Vaniz and Springer, 1971). marginally only at Ponape (USNM collections), Salarias is in need of revision; it ranges from the Eastern Caroline Islands, where I consider its western Indian Ocean to well out on the Pacific presence as evidence favoring operation of the Plate. Caroline Islands conduit. Stanulus contains two species. One species is Hirculops is monotypic and known only from restricted to the Great Barrier Reef and Lord the Indian Ocean (Smith-Vaniz and Springer, Howe Island; the other ranges from the western 1971). Indian Ocean to well out on the Pacific Plate Istiblennius is in need of revision; it is distributed (Springer, 1968; Smith-Vaniz and Springer, widely in the Indo-Pacific (Smith-Vaniz and 1971). Springer, 1971). Omobranchini: Omobranchus comprises 20 Litobranchus is monotypic and known only from species of close-shore inhabitants, rarely occurring the Philippines and New Guinea, along the west- at depths greater than 0.5 m (Springer and Go- ern margin of the Pacific Plate, and possibly from mon, 1975; Springer, 1981b). Larvae and adults Gebe Island (between Halmahera and New are generally taken together, but there is at least Guinea; Smith-Vaniz and Springer, 1971). one record of a larva having been dipnetted at Medusablennius, which is the smallest blenniid, the surface offshore (Springer and Gomon, 1975). is monotypic and known only from Raroia, Tua- Several species are restricted to estuaries or man- motus, on the Pacific Plate (Figure 60). I consider grove habitats; others occur on rocky shores in the monotypic Cirrsalarias, which is restricted to undiluted sea water. One otherwise marine and the Indo-West Pacific, to be the sister genus of estuarine species has been taken in a freshwater Medusablennius (Springer, 1976; Springer and lake in the Philippines. I collected two species Spreitzer, 1978). (not together) at the mouths of freshwater streams Mimoblennius contains four allopatric species in Ambon Bay, Indonesia, and both have been and ranges from the Red Sea and western Indian taken also in strictly marine waters at other lo- Ocean to the Bonin Islands on the eastern margin calities. One of the two species, O. rotundiceps, is of the Philippine Plate (for a distribution map see the only species of Omobranchus that occurs other 22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
J. S
(combined distribution \ of 19 of the 20 species ©distribution of the 20th species
FIGURE 9.—Distribution of the blenniid fish genus Omobranchus; no more than five species are known from a single locality. than marginally on the Pacific Plate; Ponape, east to the Tuamotus and Rapa; E. kraussi ranges Oahu, and Palmyra (Figure 9). Springer and from the Red Sea and Comores east to the Mar- Gomon (1975) proposed that Omobranchus evolved ianas and New Guinea, on the margin of the in continental waters and that time has been Pacific Plate; E. petersi is known only from the insufficient for more species of the genus to colo- northern Red Sea; and E. flavipes is known only nize Pacific oceanic islands. They believed that from the Indo-Malayan area. Enchelyurus kraussi, there must be more than one satisfactory niche E. ater, and E. brunneolus form a closely related available for Omobranchus in Oceania. One species, species group that can be defined by synapomor- O. punctatus, which ranges from the Red Sea and phies (most restricted gill openings in Enchelyurus; east coast of Africa to the Solomon Islands and males with pattern of stripes or vermiculations on Fiji, along the margin of the Pacific Plate, is also head, as opposed to uniformly colored). A clad- known from Trinidad and the Caribbean en- istic classification of the three species has not been trance to the Panama Canal (Springer and Go- possible, but based on their distributions it seems mon, 1975, fig. 7). These Caribbean occurrences probable that one of the vicariant events (at least were ascribed by Springer and Gomon to an two are required to give rise to three species) accidental introduction. Omobranchus punctatus is occurred along the western margin of the Pacific the only close-shore (adjacent to shoreline) fish Plate and the other may have occurred on the species with an Indo-West Pacific and Caribbean Plate. distribution (see Apogonidae: Apogon evermanni). Parenchelyurus contains two species, one ranging Enchelyurus, comprises five species (Springer, from the western Indian Ocean to the Marshall 1972b) inhabiting shallow, rocky, fully marine Islands, on the Pacific Plate; the other known areas (Figure 10). Two species, E. brunneolus and only from the Philippines, New Guinea, and E. ater, occur on the Pacific Plate nonmarginally. northern Australia (Springer, 1972b; Springer Enchelyurus brunneolus is endemic to the Hawaiian and Gomon, 1975). Islands; E. ater ranges from Lord Howe Island Laiphognathus is monotypic, a coral-reef species, NUMBER 367 23
OBRUNNEOLUS •FLAVIPES • KRAUSSI •PETERSI
FIGURE 10.—Distributions of the five species of the blenniid fish genus Enchelyurus.
and ranges from the east coast of Africa to the M. anema, which inhabits freshwater streams and Bismarck Archipelago and Solomon Islands on estuaries. The species are often free swimming, the margin of the Pacific Plate (Springer, 1972b, just above the substrate, but most reside in worm 1981b). tubes. Only one species, M. atrodorsalis, is known Omox, a mangrove inhabitant, contains two unequivocally to occur on the Pacific Plate non- species (Springer, 1981b) and ranges from the marginally (Figure 11), a situation comparable to South China Sea to Palau, and the Bismarck that described above for Omobranchus (Figure 9). Archipelago, and is known nonmarginally on the Smith-Vaniz questioned the Tahitian locality ac- Pacific Plate only from Ponape (O. biporos), which corded a specimen of M. procne, which is otherwise provides evidence favoring operation of the Car- known only from the northern Tonga Islands, oline Islands conduit. collected in 1931 by Zane Grey, a novelist and Haptogenys is monotypic (Springer, 1972b), an game fisherman (see also Chaetodon plebeius, family estuarine inhabitant, and is known only from Chaetodontidae, for another questionable Zane Calicut, India, and Thailand (Springer and Go- Grey locality). mon, 1975). I have seen three species of Meiacanthus (M. Phenablenniini: This tribe is monotypic atrodorsalis, M. grammistes, and M. vittatus) swim- {Phenablennius hyeligeri) and restricted to freshwa- ming together on the same coral reef in the ter and estuaries in Borneo, Sumatra, and Cam- vicinity of Port Moresby, Papua-New Guinea. bodia (Springer and Smith-Vaniz, 1972; Smith- Meiacanthus atrodorsalis was the least common of Vaniz, 1975). the three species. I would expect that M. gram- Nemophini: This tribe was monographed by mistes, which ranges over much of the western Smith-Vaniz (1976), who provided distribution Pacific (Okinawa south to Australia, and Mari- maps for all included taxa. Most of the informa- anas), would be a successful colonizer of the tion I report on the Nemophini is derived from Pacific Plate if it were able to disperse to the his study. Plate. Aspidontus comprises two free-swimming, coral- Petroscirtes comprises 10 species, most of which reef species, and both range from the east coast of are benthic and occur around vegetation; how- Africa to at least the Society Islands. ever, P. fallax, which mimics Meiacanthus lineatus, Meiacanthus comprises 16 species, all but one of swims freely in the water column. Petroscirtes lupus which occur in coral-reef areas; the exception is uses dead mollusk shells as nests. Its larvae, based 24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
i combined distribution of 15 of the 16 species
©distribution of the 16th species
FIGURE 11.—Distribution of the blenniid fish genus Meiacanthus; no more than four species known from a single locality. on aquarium observations, congregate at the sur- from the Philippines east to the Gilbert Islands, face on hatching, and after one week begin to with one of its two subspecies restricted to Fiji. feed on the bottom. Young Petroscirtes often occur One species, P. azaleus, occurs only in the eastern near floating Sargassum. Only two species, P. mi- Pacific. The other five species are variously re- tratus and P. xestus, occur other than marginally stricted to the Indo-West Pacific and do not reach on the Pacific Plate. Both are wide ranging, the western margin of the Pacific Plate. extending from the east coast of Africa to the Xiphasia comprises two species of tube-dwelling Marshall or Line and Society islands, but neither (in soft bottoms), eel-like blennies. Xiphasia setifer is present in Hawaii. Three other species of Pe- ranges from the east coast of Africa to the western troscirtes have the eastern limits of their distribu- margin of the Pacific Plate; X. matsubarai also tions at the western margin of the Pacific Plate, ranges from the east coast of Africa to the western but only one of these, P. breviceps, ranges as far margin of the Pacific Plate, but one collection, west as the east coast of Africa. from ll°40'S, 175°10'E, is just north of the Pa- Plagiotremus comprises 11 species of free-swim- cific Plate margin. While I consider X. matsubarai ming fishes that use abandoned worm tubes in to occur nonmarginally on the Pacific Plate, the reefs as shelter. Five species occur on the Pacific species is essentially absent from the Plate, and Plate, of which two, P. ewaensis and P. goslinei, are the exceptional collection may represent a waif known only from the Hawaiian Islands, and two (see Smith-Vaniz, 1976, fig. 38). others, P. rhinorhynchos and P. tapeinosoma, range Summary: While reasonably complete infor- from the east coast of Africa through much of mation on Indo-Pacific blenniid distributions is Oceania, but are absent from the Hawaiian Is- far from available, what is known is probably lands (each of the two Hawaiian species appears indicative, and can be summarized as follows. to have a different one of the wide-ranging species Although it is unknown how many species of as its sister species). The fifth, P. laudandus, occurs Blenniini reach the western margin of the Pacific NUMBER 367 25
Plate, only one species occurs on the Plate non- is reported from localities ranging from the east marginally. coast of Africa eastward into the eastern Pacific, Of the 19 genera of Salariini that reach the and another, Bothus pantherinus, ranges from the western margin of the Pacific Plate from the west, Red Sea to the Society Islands. Several deep- only 13 extend onto the Plate nonmarginally. Of dwelling genera {Chascanopsetta, Parabothus, Pele- the 12 species of Ecsenius that reach the western canichthys, Taeniopsetta) not included in Table C margin of the Pacific Plate from the west, only (Appendix 2) are present nonmarginally on the three extend onto the Plate nonmarginally, and Pacific Plate only in the Hawaiian Islands. one of the three is represented by allopatric color- pattern populations that are restricted to one side BRACHAELURIDAE or the other of the Plate margin. Of the six species of Entomacrodus that similarly reach the western This family is discussed under the Orectolobi- margin of the Plate, only four extend nonmargin- dae. ally onto the Plate. Of the six genera and seven or eight species of BRANCHIOSTEGIDAE Omobranchini that reach the western margin of (not including Malacanthidae) the Pacific Plate from the west, only four genera and four species extend onto the Plate nonmar- The tilefishes comprise three genera and 23 ginally. species (two questionable) of moderately large Of the five genera and 19 species of Nemophini (230-867 mm SL), moderately deep-dwelling that reach the western margin of the Pacific Plate (20-600 m), commercially important, epibenthic from the west, all the genera but only nine species or benthic fishes with pelagic larvae (Dooley, occur on the Plate nonmarginally, and one of the 1978; 1981; Berry, 1978). nine (Xiphasia matsubarai) has been reported only Dooley (1978:14), who revised the family, de- once from the Plate, and not very far from the scribed its distribution as "worldwide in tropical, margin. warm and cold temperate waters along the edges It can be surmised from these data that the of continental shelves and the upper slopes of western margin of the Pacific Plate delineates a continental margins; occasionally found around sharp break in the distribution of blenniid fishes. continental or oceanic islands." I have plotted the The margin, as a feature of biogeographical im- distribution of the Branchiostegidae (Figure 12) portance, is further emphasized by the presence based on information in the studies of Dooley and of widely distributed blenniids endemic to the Berry and a record for Fiji (Carlson, ms.), and Plate. find that these fishes are well represented in the New World tropics, but they are absent from BOTHIDAE most of the Old World tropics. Indeed, they appear to be distributed antiequatorially in the The lefteye flounders are a large group (Nelson, Indo-West Pacific. Except for a Red Sea-South 1974, estimates that there are 36 genera and African species, none of the Indo-West Pacific about 212 species) of shallow- to deep-dwelling, species occur both north and south of the equator. benthic, and small- to moderate-sized fishes with The family is absent from the nonmarginal por- planktonic larvae. The family occurs circumglob- tions of the Pacific Plate. ally in tropical and temperate seas, and consists of two or three subfamilies, only one of which, BREGMACEROTIDAE Bothinae, the most speciose, occurs on the Pacific Plate. It is represented there by three genera with The codlets consist of a single genus (Bregma- probably five species. The Indo-Pacific species ceros) and about five species (Belyanina, 1974) of are not well known, but one species, Bothus mancus, small fishes. The codlets are distributed circum- 26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 12.—Distribution of the tilefishes, Branchiostegidae. globally (see D'Ancona and Cavinato, 1965, and CAESIONIDAE Belyanina, 1974, for distribution maps), and oc- cur at depths from the surface to perhaps 1000 m The caesios comprise four genera (Johnson, or more. The adults of some species are known 1980) and perhaps 25 species of small (to about only from shallow coastal waters. Most specimens 400 mm TL), free-swimming fishes that occur in collections are larvae and juveniles taken in only in the Indo-Pacific. Gymnocaesio and Diptery- plankton nets. Although I have collected speci- gonotus, each with two species, are restricted to the mens in a shell dredge close to shore in Indonesia, Indo-West Pacific, variously occurring from the depth unknown, but probably less than 50 m, I Gulf of Aden east to Melanesia, north to (south- only tentatively characterize the codlets as ern?) China, and south to the East Indies. As shorefishes. many as ten species of Caesio and Pterocaesio may be present nonmarginally on the Pacific Plate. These genera are widely distributed in the Indo- BYTHITIDAE Pacific and in need of revision; some of the species The bythitids are a speciose group of small, are restricted to the Indo-West Pacific; but it is viviparous fishes distributed circumglobally. doubtful that any of the others are Pacific Plate Cohen and Nielsen (1978) have reviewed the endemics. genera and provided general information on dis- tributions; the species are poorly known taxonom- CALLIONYMIDAE ically. They occur at depths of less than a meter down into the bathypelagic; at least one species The dragonets comprise several genera and occurs in freshwater. Four genera (probably ten about 130 species (Fricke, 1981c) of small (maxi- species) are reported from nonmarginal areas of mum about 300 mm TL), benthic fishes occurring the Pacific Plate. almost circumglobally in shallow to moderately NUMBER 367 27 deep temperate and tropical marine waters. Most Tonga, which are on plate margins adjacent to species live on sandy or muddy bottoms, but two that of the Pacific Plate, and to S. stellatus, which species enter and live in freshwater (Fricke, is an Indian Ocean endemic. He related S. laddi 1981c). to S. postulus and S. minutulus, both of which are Diplogrammus comprises five allopatric species Indian Ocean endemics. (Fricke, 1981a): one restricted to the Red Sea, The kaianus species group of Callionymus was one to the Arabian Sea, one to the western Indian revised by Fricke (1981c). This group comprises Ocean, one to Japan, and one distributed from 13 deep-dwelling species (most taken at over 100 the Ryukyu Islands south to Queensland and east m) that are confined to the Indo-West Pacific to the Marshall, Fiji, and Samoan islands. (east coast of Africa east to Japan, Philippines, Pogonymus comprises four species, each known New Ireland, and New Caledonia). Probably no from a single area (W.P. Davis, pers. comm.): more than three other species of Callionymus occur Hawaiian Islands, Japan, India, Nigeria. The on the Pacific Plate. Hawaiian species apparently constitutes an ex- ample of an Hawaiian exception (for a genus) on CARACANTHIDAE the Pacific Plate. Synchiropus was revised by Fricke (1981b). The The caracanthids comprise a single genus, Car- genus contains 26 species and is found in all acanthus, with four small (to about 50 mm TL) warm-temperate and tropical seas, from shallow species of coral-dwelling fishes. The family is to moderately great depths. The Mediter- widely distributed in the Indo-Pacific, and ranges ranean-eastern Atlantic, western Atlantic, and from the east coast of Africa eastwards to the eastern Pacific regions each harbor one endemic Galapagos Islands (Mizuno and Tomiaga, 1980, species. The remaining 23 species are variously fig. 3) in the eastern Pacific. One species is re- distributed in the Indo-Pacific, with six species stricted to the western Indian Ocean, one to the restricted to the Indian Ocean. Five species occur Hawaiian and Johnston islands, one widely dis- nonmarginally on the Pacific Plate, of which one tributed in the western and central Pacific Ocean (S. rubrovinctus) is an Hawaiian endemic, two are (but not at Hawaiian and Johnston islands), and widely distributed endemics (S. laddi: Palau Is- one species ranges from the east coast of Africa to lands, Bikini, Rongelap, Enewetak, and Raroia, the Society and Galapagos islands (W.N. Esch- and an additional locality, Cuyo Islands, Philip- meyer, pers. comm.). pines, noted by Fricke, in litt.; S. morrisoni: Ulithi, Bikini, Tutuila), one (S. altivelis) is an example of CARANGIDAE an Hawaiian exception (otherwise ranging from Indonesia and the Philippines north to Japan), The jacks comprise about 140 species (W.F. and one (S. splendidus) occurs widely in the western Smith-Vaniz, pers. comm.) of small to large fishes Pacific, but nonmarginally on the Pacific Plate (maximum about 1.75 m TL) with planktonic only in the Eastern Caroline Islands (thus provid- larvae. The species are free-swimming and some ing evidence favoring operation of the Caroline pelagic species are circumglobally distributed Islands conduit). Fricke did not treat the relation- (Briggs, 1960). Many species limited to the Indo- ships of Synchiropus cladistically, but his stated Pacific appear to be widely distributed; however, relationships of the two widely distributed en- some genera and species are notably absent from demics are of interest as both were related to the Pacific Plate. In all, 29 species in 14 genera species that do not occur on the Pacific Plate are present nonmarginally on the Plate. nonmarginally. He related S. morrisoni to S. ocel- Berry and Cohen (1974, fig. 1) gave the distri- latus, which is restricted to the Indo-West Pacific bution of the 13 species of Trachurus (reduced to as far west as Vietnam and as far east as Palau to 12 species by Stephenson and Robertson, 1977, 28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY and to 11 species with several allopatric subspe- CARCHARHINIDAE cies—for a total of 14 taxa—by Shaboneeve, The requiem sharks are a circumglobal family 1980, who provides a more detailed distribution comprising 11-12 genera and about 45 species of map). Trachurus occurs along coasts bordering all small to very large fishes (maximum adult size oceans, but is noticeably absent from the Pacific ranging from about 0.6-5.5 m). The species are Plate. While the Atlantic species are continuously free swimming, shallow dwelling, carnivorous, distributed from about 30°N-30°S, the Indo- live bearing, marine (a few species enter fresh or West Pacific species are essentially antitropical in brackish water), and may be restricted essentially distribution. to littoral areas or the open ocean. The Indo-Pacific genus Scomberoides contains four species. All are widely distributed in the Carcharhinus has been revised by Garrick (in Indo-West Pacific, but only one (S. laysari) extends press), who later (in prep.) included five addi- nonmarginally onto the Pacific Plate (to Hawaii, tional species. Carcharhinus is the most speciose Marquesas, and Tuamotus; Smith-Vaniz and genus in the family, with 30 species, ranging in Staiger, 1973). adult size from 0.95-3.62 m TL. Twenty-six of the species occur in the Indo-Pacific, but only Megalaspis (monotypic) is widely distributed nine (34.6%) of these occur on the Pacific Plate and very common in the Indo-West Pacific. It nonmarginally, and none of these is endemic to has been reported (Jordan et al., 1927) only once the Plate. Of the nine species on the Plate, four from the Pacific Plate, from a single specimen large species (2.6-3.0 m) are worldwide in distri- that was seen at the Bishop Museum in Honolulu, bution and common in all areas (two of these and presumed to have been taken in Hawaiian species are oceanic and the only ones so restricted waters. It seems doubtful that this specimen ac- in the genus); two species, C. plumbeus and C. tually came from the Hawaiian Islands, which altimus (2.4 and 2.8 m), are distributed circum- have been credited with numerous dubious or globally, but are known nonmarginally on the erroneous records of fishes (as well as valid excep- Pacific Plate only from the Hawaiian Islands tional occurrences for the Pacific Plate). (examples of Hawaiian exceptions), and three species (1.8-2.5 m) are widely distributed in the CARAPIDAE Indo-Pacific. Carcharhinus melanopterus, the smallest of the three widely distributed Indo-Pacific spe- The pearlfishes comprise six genera and about cies, ranges from the Red Sea and east coast of 30 species (Cohen and Nielsen, 1978) of small (to Africa to the Tuamotus and Marquesas, and has about 309 mm TL) coastal fishes. Most, if not all, been reported recently from the Mediterranean, have planktonic larvae, and most are internal possibly as a migrant through the Suez Canal. associates of echinoderms and mollusks. The fam- Carcharhinus albimarginatus, the largest of the three ily is almost circumglobally distributed, being widely distributed Indo-Pacific species, ranges absent only from the islands (Ducie, Eastef) near- into the eastern Pacific as well. est the eastern margin of the Pacific Plate- Most Of the 17 Indo-West Pacific species of Carchar- of the species live in shallow water, but the free- hinus that do not occur on the Pacific Plate, four living Snyderia canina has been taken in a dredge (C. brevipinna, C. brachyurus, C. leucas, C. obscurus) haul made at a depth of over 2000 feet (610 m) are among the largest species of Carcharhinus (2.78, off the Hawaiian Islands. About six shallow- 2.92, 3.24, 3.62 m) and are circumglobally dis- dwelling species of pearlfishes have beert taken tributed except for their noticeable absence from nonmarginally on the Pacific Plate. The total the Pacific Plate. The other 10 Indo-West Pacific number of Indo-Pacific carapodid species proba- species of Carcharhinus are distributed as follows: bly does not exceed 10. four (0.95-1.66 m) widely distributed but re- NUMBER 367 29 stricted to the Indo-West Pacific; one (2.23 m) in Bombay, India, east and south to Java and north the eastern Atlantic and Indo-West Pacific; three to Tinghai, Chekiang Province, China, and Ja- (1.0-1.5 m) only in the western Pacific; one (1.72 pan (and perhaps to the Bonin Islands; see Zama m) only in the western Indian Ocean; and one and Fujita, 1977, as S. wahlbeemi). It is also re- (1.34 m) in the western Atlantic, eastern Pacific, corded from Sarawak, but not the Philippines. and western Pacific. Loxodon is also monotypic, but with a wider Indo- From the information presented above, it is West Pacific distribution (Figure 13) than Scolio- apparent that size alone is insufficient to account don. Loxodon is the only one of the three genera for the presence or absence of Carcharhinus species that occurs on islands (Seychelles, Mascarene Pla- on the Pacific Plate (although all four of the teau, including, perhaps, Mauritius) remote from worldwide species are large). The large propor- continents. Springer (1964) reported extreme and tion of the species that reach the margin of, but discontinuous variation in numbers of precaudal do not extend out on, the Plate, agrees with my vertebrae in different populations of Loxodon, in- findings on other elasmobranch groups, which, in dicating that a species complex might be in- general, tend to be absent from the Pacific Plate. volved. The Seychelles and Mascarene Plateau To avoid the possible criticism that there are are continental fragments (Scrutton, 1976), and probably inadequate data available for large it is possible that the populations of Loxodon were sharks on the Pacific Plate, it is noted that Stras- left behind to diverge (vertebrally) when the con- burg (1958) reported on longline collections that tinent, or continents (India, Africa), that were covered much of the central Pacific during a four- associated with the fragments disappeared or year period and produced over 6000 specimens of moved away. sharks, but only 12 species, of which only three Rhizoprionodon, with seven species (three Indo- were Carcharhinus. West Pacific, of which one also occurs in the Triaenodon contains two species. Triaenodon obtu- eastern Atlantic; one eastern Pacific; and three sus is known only from the holotype from India western Atlantic) is more widespread (Figure 13) (Taniuchi, 1975). Triaenodon obesus, which attainsthan Loxodon, occurring more or less circumtrop- a length of 2.13 m, is found only near coral reefs, ically, but absent from the Pacific Plate. often resting in caves, yet it ranges from the Red It is not hard to visualize the distributions of Sea and east coast of Africa eastward to Panama Loxodon and Scoliodon as representative of levels of (Randall, 1977). Randall (1977) tagged 124 T. increasing restriction in distribution from what obesus at Johnston Island and recovered seven were previously widespread distributions similar after 16 days to two years; none had moved more to that found for Rhizoprionodon; especially as all than 2.9 km from their tagging site (tagging three genera occur together over much of their information in the abstract to Randall's paper Indo-West Pacific distribution. Bearing on this expands on tagging data reported in the body of are the distributions of the two subgenera of the paper). One may wonder, as did Randall, Rhizoprionodon. The type subgenus has the same how such a sluggish, reef-associated shark was overall distribution as the genus, whereas the able to cross the Pacific. A possible answer to this other subgenus, Protozygaena, has a disjunct distri- question is discussed in the section on the geolog- bution, occurring from India to Queensland in ical history of the Pacific and Philippine plates. the western Pacific, and from Panama to southern Scoliodon, Loxodon, and Rhizoprionodon comprise Brazil in the western Atlantic. If the distribution small, shallow-dwelling sharks with maximum of the subgenus Protozygaena is segregated, the total lengths of about 350-1000 mm. The species order of increasing distributional restriction is as are of commercial importance in Asia. The genera follows: Rhizoprionodon, Protozygaena, Loxodon, Sco- were revised by Springer (1964). Scoliodon is mon- liodon. otypic and its distribution is coastal, from about Galeocerdo (tiger shark), monotypic, is a large 30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 13.—Distributions of the carcharhinid shark genera Rhizoprionodon (solid circles) and Loxodon (stars).
species that appears to be circumglobally distrib- bottoms near mangroves. The family ranges from uted along warm, shallow, marine coasts, includ- the Red Sea and eastern Indian Ocean to along ing the Pacific Plate. The species purportedly much of the western margin of the Pacific Plate attains about 9 m TL, but there are few, if any, (however, unknown from Samoa). I know of only authenticated records of specimens over 5.5 m three records from the Pacific Plate nonmargin- TL. ally: one for Honolulu (Fowler, 1928), one in the Dirrhizodon (often referred to as Hemipristis), USNM collections supposedly collected by Alex- monotypic, attains about 2.5 m TL, and is re- ander Agassiz in 1899 from Makemo, Tuamotus, stricted to the Indo-West Pacific, from the Red and one at the Bishop Museum from Ponape. I Sea and western Indian Ocean to the Gulf of consider the first two of these records as question- Thailand. able, as does J.E. Randall (in litt.). On the other hand, the Ponape record (Aeoliscus) is undoubt- CENTRISCIDAE edly valid, and R.A. Croft, Fisheries Officer in Ponape and a marine fish collector, informs me (excluding Macrorhamphosidae) that he has obtained shrimpfish in Ponape. The The shrimpfishes consist of two genera {Aeolis- presence of shrimpfish in the Eastern Caroline cus, Centriscus) and four species of small (to about Islands chain, which extends westward to the Yap 150 mm TL), shallow-dwelling, suprabenthic and Palau islands (where centriscids are common; fishes presently found only in the Indo-Pacific California Academy of Sciences collections) on (Macfarlane, 1923:243, reported a fossil species of the margin of the Philippine Plate, is evidence for centriscid as common in the Oligocene rocks from operation of the Caroline Islands conduit, which Alsace to Baku, central Europe). They occur in a is allowing dispersal onto the Pacific Plate from variety of habitats, from coral-reefs to muddy the continental areas to the west. NUMBER 367 31
CENTROPOMIDAE In the Caribbean, snooks are seen occasionally The snooks, according to Greenwood (1976), in coral-reef areas adjacent to mangroves (Ran- comprise three extant genera (Centropomus, Psam-dall, 1968). moperca, Lates) and about 18 species of moderately large (to about 1.5 m), free-swimming fishes. CEPOLIDAE Seven of the species are in African freshwaters. (including Owstoniidae) The marine species frequently enter brackish wa- ter, and some species that live primarily in fresh- The bandfishes comprise four or five genera water spawn in estuaries (Lake, 1971). There are and about 10-15 species of small (to perhaps 700 three Indo-West Pacific marine species with an mm TL), benthic or suprabenthic fishes that overall range from the Persian Gulf to Taiwan, usually occur in moderately deep water (up to the Philippines, and northern Australia. The re- 400 m), but are often taken at much shallower maining species are in the New World tropics depths (almost on shore after storms in the Med- (Greenwood, 1976, fig. 36, presents a world dis- iterranean-Atlantic area). The species occur on tribution map for the family). A western Atlantic rocky or muddy bottom. Little is known about species of the western hemisphere genus Centropo- their reproductive biology. mus has been reported (Fraser-Brunner, 1931) at The bandfishes occur from the eastern Atlantic least once from West Africa (Nigeria). The indi- eastward to the western margins of the Philippine vidual was probably a waif. The fossil record for and Pacific plates (Figure 14). There is a single the family ranges from the Eocene of Monte western Atlantic record (Kanazawa, 1952) of a Bolca, northern Italy, to the Holocene, and in- cepolid (Cepola), which is based on a specimen cludes also the Paris Basin, Austria, Portugal, and taken from the stomach of a grouper (Serranidae) Croatia to Egypt, the Sahara, and eastern Africa caught at Bermuda. The specific identification of (Greenwood, 1976). the specimen was not possible because of its poor
FIGURE 14.—Distribution of the fishes of the family Cepolidae (stars indicate unspecified localities in general area). 32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY condition. Bandfishes are either adventitiously C. Birkeland, in litt.); C. smithi (= C. hemichrysus; present in the western Atlantic, or deep-dwelling Pitcairn, Rapa); and C. tinkeri (Hawaiian Islands in that region in areas inaccessible to normal and Enewetak, Marshall Islands, J.E. Randall, in fishing gear. * litt.; also mentioned in Myers, 1981). There are also two species restricted to the GHAETODONTIDAE margins adjacent to the Pacific Plate: Chaetodon The butterflyfishes comprise ten genera and flavocoronatus from Guam (Myers, 1981) and C. 116 species (total derived from the following: litus from Easter Island (both of which I have Allen, 1980; Allen and Kuiter, 1978; Burgess, included as Pacific Plate species for the purposes 1979; Lubbockand Edwards, 1980; Myers, 1981). of Table C, Appendix 2). The species are small (to about 300 mm TL), Of the 59 Indo-West Pacific species of chaeto- suprabenthic, predominantly coral-reef dwelling, dontids that do not occur on the Pacific Plate and conspicuously colored, with planktonic lar- nonmarginally, or are restricted only to the mar- vae. The family is circumglobally distributed, gins, 34 have their eastern distributional limits and only one genus and 17 species (in three well west of the Plate's western margin and 25 genera) do not occur in the Indo-Pacific. Johnran- reach to, or close to, that margin. Therefore, there dallia (= Pseudochaetodon) is limited to the easternare a total of 55 species of chaetodontids that Pacific. Only one of the Indo-Pacific species, For- reach the western margin of the Pacific Plate cipiger flauissimus,range s out of that region (to the from the west, but, as noted above, only 28 eastern Pacific). (50.9%) of these continue onto the Plate. The Five of the nine Indo-Pacific chaetodontid gen- western margin of the Plate thus delineates an era (Chelmon, Chelmonops, Coriadon, Parachaetodon, effective barrier to the eastward dispersal of chae- Amphichaetodon, including nine species) are widely todontids. distributed in the Indo-West Pacific but are ab- One of the species, Chaetodon plebeius, that is sent from the Pacific Plate, except marginally. included among the 25 that reach only to the Amphichaetodon is also represented by an endemic western margin of the Pacific Plate was reported species in the San Felix Islands, off Chile. by Burgess (1979) as occurring from the western Of the 99 Indo-Pacific chaetodontid species, 38 Indian Ocean to Fiji, and disjunctly, Tahiti, occur nonmarginally on the Pacific Plate. Of the which is the only nonmarginal Pacific Plate rec- 38, ten occur only on the Plate and 28 also occur ord for the species. The Tahitian record is based in the Indo-West Pacific. Five of the ten endemics on a specimen collected in April 1931 by Zane are known from single islands or island groups: Grey, a sport fisherman and writer, who, during three at the Hawaiian Islands (including John- May 1931, also collected C. plebeius in Fiji, where ston Island), one in the Marquesas, and one at the species has been taken by other collectors as Pitcairn. The other Plate endemics are more well. Randall (1973) and Allen (1980) did not widely distributed: Hemitaurichtys thompsoni (Ha- report C. plebeius from Tahiti, and R.G. Wass waiian, Johnston, and Tokelau islands, Burgess, (ms.) does not list it for Samoa. I believe the Zane 1979; Samoa, R.C. Wass, ms.; Guam and Ran- Grey record of C. plebeius to be an error in label- giroa, J.E. Randall, in litt.); Chaetodon trichrousling. Smith-Vaniz (1976, fig. 48) also questioned (Society, Tuamotu, Marquesas and possibly Gil- a Zane Grey Tahitian record for Meiacanthus procne bert islands; Allen, 1980:232, cited a questionable (Blenniidae). record from Palmyra, Line Islands); C. quadrima- There is only one example of an Hawaiian culatus (Hawaiian, Line, and Tuamotu islands, exception among the Chaetodontidae. Heniochus Burgess, 1979; southern Japan, Society, Pitcairn, diphreutes occurs from the western Indian Ocean Phoenix, Marshall, Wake, and Marquesas is- to Japan and Australia, and then appears in lands, Allen, 1980; Samoa, Guam, and Saipan, Hawaii (Allen and Kuiter, 1978, figs. 1 and 2). NUMBER 367 33
Its occurrence in Hawaii is possibly the result of I have traced only as far back as Bleeker dispersal from Japan via the Kuroshio extension. (1869-1875). The next largest size that I have been able to locate for wolf herrings is 4.5 ft (1.4 CHANIDAE m, Grant, 1978). Reports over 1 m are uncom- mon. The milkfish, Chanos chanos, is the sole member The wolf herrings are absent from the Pacific of the family. It attains a length of more than a Plate, but are widely distributed in the Indo-West meter, spawns in the sea, and the fry may enter Pacific, from the eastern Indian Ocean to Fiji, freshwater, where they remain for several years adjacent to the western margin of the Pacific until they are about 600 mm or more in length; Plate. thereafter, they return to the sea (Herre and Mendoza, 1929). The milkfish is widely distrib- uted in the tropical Indo-Pacific. It ranges from ClRRHITIDAE the east coast of Africa eastward to the eastern The hawkfishes comprise 9 or 10 genera and Pacific. about 35 species (Randall, 1963, and pers. comm.; Lavenberg and Yanez, 1972; Lubbock, 1978) of CHEILODACTYLIDAE small- to moderate-sized (maximum, 550 mm SL), benthic, reef-dwelling fishes. One species is The morwongs are a family of moderate-sized limited to the Caribbean and St. Helena, in the fishes (to about 400 mm FL) with pelagic eggs eastern Atlantic; one species is endemic to Ascen- (Tong and Vooren, 1972). The family is in need sion, in the eastern Atlantic; one species is en- of revision, but probably contains fewer than 20 demic to the west African coast; and one is en- species. demic in the eastern Pacific. The remaining 31 The general distribution of the morwongs can species are restricted to the Indo-Pacific, except be described as antiequatorial. They are distrib- that two species (Oxycirrhitus typus, Cirrhitichthys uted in the southern hemisphere on the warm- oxycephalus) range from the western Indian Ocean, temperate, rocky coasts of the continents and and one species (Cirrhitichthys serratus) ranges continental islands, ranging as far north as south- from Guam (but see Myers and Shepard, ern Queensland (Grant, 1978), in Australia. In 1981:343-344) and the Ryukyu Islands, into the the northern hemisphere they range from Japan eastern Pacific. Of the 31 Indo-Pacific species, 14 and Korea south to about Hong Kong. On the do not occur on the Pacific Plate nonmarginally Pacific Plate there are only two species: Cheilodac- and 17 do. Nine of the 17 are widely distributed tylus vittatus, with a north-south disjunct distribu- in the Indo-Pacific, seven occur only at Pacific tion (Hawaiian Islands, Lord Howe Island, New Plate localities (two also occur at Philippine Plate Caledonia), and an undescribed species known localities, but all seven are considered here as only from Rapa and Easter Island (Randall, In Pacific Plate endemics), and one species, Cirrhiti- press b, c). chthys serratus, has a remarkable distribution: Guam (Kami et al., 1968; but see Myers and Shepard, 1981:343-344), Maug (northern Mar- CHIROCENTRIDAE ianas, J.E. Randall, in litt.), Samoa (R.C. Wass, The wolf herrings consist of a single genus, ms.), questionably, Oahu (Randall, 1963, and in Chirocentrus, and two species of free-swimming, litt.), and Gorgona Island (Colombia, eastern Pa- voracious, near-shore fishes with pelagic eggs. The cific; Randall, 1963). Cirrhitichthys serratus is the wolf herrings are often reported (on hearsay) to only sedentary reef species I know that appears reach a length of 12 ft (3.6 m). I have been unable to be limited to the Pacific Plate and the eastern to locate the initial report listing this size, which Pacific. Randall (in litt.) suspects that it may be 34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY synonymous with the widely distributed western ably synonymous. The last two named species Pacific species C. falco, which occurs as far east as then, are very close, if not identical. the western margin of the Philippine and Pacific Lavenberg and Yanez (1972) stated that Cir- Plates, between the Marianas and Fiji. rhitus wilhelmi (Easter Island, Rapa, Hots du Bass, Four of the endemic Pacific Plate species are Pitcairn Island group; Randall, pers. comm.) and widely distributed on the Plate (Figure 60), and C. splendens (Lord Howe Island) form a "group" include one monotypic, endemic Plate genus, within Cirrhitus. If true, the endemicity of the Isocirrhitus (I. sexfasciatus, Marshall and Tuamotu "possibly new genus" on the Pacific Plate (Figure islands), and possibly a second, undescribed (for 60) would be eliminated; however, Randall (in Cirrhitus wilhelmi; J.E. Randall, pers. comm.; Cres-litt.) does not consider these two species to be sey and Randall, 1978, assigned C. wilhelmi to congeneric. Arablycirrhitus without comment). The proportion The evidence, although weak, seems to indicate of cirrhitids restricted to the Pacific Plate is, that the Pacific Plate endemic cirrhitids tend to perhaps, the highest among all the Indo-Pacific have allopatric Indo-West Pacific sister species fish families. (or slightly differentiated populations); however, Relationships among the cirrhitid species have a new revision seems desirable. not been treated cladistically. If, however, Ran- dall's (1963) comments to the effect that certain ClTHARIDAE species are more closely related can be extrapo- lated to indicate sister-species groups, then the The Citharidae (= Eucitharidae) is a small closest relative of the Pacific Plate endemic, Par- family consisting of five monotypic genera acirrhites xanthus (Tuamotu, Society, and Marque-(Hubbs, 1945) of small, bottom-dwelling flat- sas islands, Caroline Atoll; Randall, 1963, and in fishes (probably not attaining 300 mm TL) that litt.) is the allopatric western Pacific species P. ordinarily occur at depths of 100-200 m. The amblycephalus (Sangi Islands). Paracirrhites nisus family is distributed from the Mediterranean (Tuamotu, Oeno, and Phoenix islands; Randall, south to about 16°S latitude on the west African 1963, and in litt.) and P. bicolor (Caroline Atoll coast, and from the Red Sea south to southeast and Tuamotus; Randall, 1963, and in litt.), both Africa and eastward to Japan, the Philippines, Plate endemics, were considered to be closely and Arafura Sea. One species, Brachypleura nova- related, and are sympatric in at least part of their zeelandiae, was described from New Zealand, but ranges. It is not possible to decide the sister species Chabanud (1954) believes that locality to be of this pair from Randall's study. The two species erroneous, and cites the distribution of this species of Cirrhitops are allopatric: C. fasciatus from theas Maldives to the Arafura Sea. Indo-West Pacific (Japan, Madagascar, Mauri- tius) and the Hawaiian Islands (Hawaiian excep- tion) and C. hubbardi from the Bonin, Phoenix, CLINIDAE Gilbert, and Tuamotu islands. As implied above, The blennioid fishes of the family Clinidae (as Randall (1963) considered Cirrhitichthys senatus todefined by George and Springer, 1980, who ex- be "very closely related" to C. falco. The western cluded the Tripterygiidae, Labrisomidae, and margin of the Pacific Plate separates the distri- Chaenopsidae, which have often been treated as butions of the two species. Randall noted that the clinids) comprises about 20 genera and 70 species, Plate endemic Paracirrhites typee (Marquesas andmost of which are restricted to Australia and Guam; Kami, 1971) was "morphologically simi- Africa. The species are small (few, if any, reaching lar" to P. forsteri and P. hemistictus, both widely 300 mm SL) and benthic, although some species distributed Indo-Pacific species, and Randall inhabit kelp forests off California. The western (1973) stated that P. typee and P. forsteri are prob-hemisphere and Mediterranean species are all NUMBER 367 35
•--.....--'
FIGURE 15.—Distribution of the fishes of the family Clinidae sensu George and Springer, 1980 (star indicates unspecified New Guinea locality). oviparous, hence unspecialized, relative to the nonmarginally (Whitehead, in litt.). The western Indo-Pacific species, which are all viviparous. All Plate margin, therefore, represents a strong bar- but four or five (tropical) species are limited to rier to the eastward dispersal of the clupeids. warm-temperate waters. The family appears to Randall (1973) mentions several records of clu- have an antiequatorial distribution (Figure 15), peids from the Society Islands that Whitehead except for the Indo-Malayan region, and is absent (in Randall, 1973) considers questionable because from the Pacific Plate. the species are not otherwise reported from Oceania. For instance, Randall considered the CLUPEIDAE record of Sardinetla gibbosa from the Society Islands (derived from Giinther's 1909 report of Clupea The herring family comprises about 64 genera gibbosa) plausible because the species was also and 196 species (P.J. Whitehead, in litt.) of small reported (by Giinther) from Fiji, Tonga, and (usually under 500 mm SL), free-swimming fishes Samoa. On only this basis I would question the with planktonic young. The herrings are usually plausibility of S. gibbosa's occurring in the Society marine coastal fishes, but several species are re- Islands, because Fiji, Tonga, and Samoa are on stricted to freshwater, particularly in the western or close to the margin of the Pacific Plate, and hemisphere. many otherwise continental plate species have The herrings are distributed circumglobally in their eastern limits in these islands. However, tropical and temperate waters. Because of a lack Randall did not mention Giinther's inclusion of of recent comprehensive revisions within the fam- Ponape (Eastern Caroline Islands) in the range ily, it is not possible to detail distributions, but it of S. gibbosa, and with this added locality I am appears that no more than about six of the 104 more inclined to accept the Society Islands record Indo-Pacific species occur on the Pacific Plate of S. gibbosa. On the other hand, there have been 36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY so many suspect records of species occurring on tirely restricted to freshwater. The absence of the Pacific Plate based only on Museum Godef- gizzard shads from the Pacific Plate (Figure 16) froy specimens, as were Giinther's, that I remain is possibly the result of lack of habitat, and this skeptical of the Society Islands record of & gibbosa. absence is duplicated by most of the Indo-Pacific If the Society Islands record is in error, the pres- genera and species of clupeids. Based on the ence of the species in Ponape may possibly indi- successful introduction of Sardinella marquesensis in cate operation of the Caroline Islands conduit. Hawaii, it would appear that more clupeids could Sardinella is represented by about 17 species exist on the Pacific Plate if they could invade the (Chan, 1965; Whitehead, 1967) in the Indo-Pa- area. cific, of which only two or three appear to be Whitehead (1963) revised the subfamily Dus- present nonmarginally on the Pacific Plate. Only sumieriinae (which he recognized as a family), in one species appears to be endemic to the Plate, S. which he recognized six genera and ten species. marquesensis from the Marquesas. Sardinella marque- Five genera and eight species were recorded from sensis has been introduced artificially into the the Indo-Pacific, but only Spratelloides delicatulus Hawaiian Islands, where it has become estab- was reported to occur nonmarginally on the Pa- lished (Berry and Whitehead, 1968; Randall and cific Plate, and then only at Hawaii. Whitehead's Kanayama, 1972). There are no species of Sardi- distribution reports are misleading as he made no nella native to the Hawaiian Islands. mention of the numerous and undoubtedly valid The clupeid subfamily Dorosomatinae, gizzard reports of dussumieriins from Plate localities, in- shads, comprises about 17 species (Nelson and cluding, at least, the Marshall, Caroline, Gilbert, Rothman, 1973) of large clupeids (to about 520 Phoenix, and Tuamotu islands (Herre, 1939; mm SL). They occur in habitats ranging from Schultz and Welander, 1953; Randall, 1955d; freshwater lakes and rivers to fully marine bays. Schultz, 1943; Harry, 1953). Most of these reports The western hemisphere species are almost en- refer only to Spratelloides, but Dussumieria has been
FIGURE 16.—Distribution of the fishes of the clupeid subfamily Dorosomatinae, gizzard shads (dark areas in North and Central America indicate generalized freshwater distribution). NUMBER 367 37 reported from the Marshalls and Hawaii, and G. ippines), and A. bowersi, which is probably en- Nelson (in litt.) informs me that Etrumeus (E. demic to the Hawaiian Islands, where it is com- micropus) occurs in the Hawaiian Islands (Ha- mon. Giinther (1910) reported on a specimen (as waiian exception; the same species also occurs in Congromuraena bowersii) procured by the Godeffroy Japan). Whitehead (in litt.), however, doubts that Museum from south of the Gilbert Islands. The Dussumieria occurs in the Hawaiian Islands. specimen is in the British Museum collections Whitehead (1973) essentially duplicated the dis- and its identification should be verified. tribution data of his earlier paper; however, most of the dussumierins do appear to be restricted to CONGROGADIDAE continental plate coastal areas. The congrogadids comprise about ten genera CONGRIDAE and 20 species (Winterbottom, 1980) of small (maximum size about 300 mm SL), elongate The conger eels are probably distributed cir- fishes restricted to the reefs and rocky bottoms of cumglobally and consist of at least 21 genera and the Indo-West Pacific (Figure 17). Castle (1980) 80 species (R. Kanazawa, pers. comm.) of mod- has shown that a report of Congrogadus from the erate to very large (to at least 2.7 m TL) fishes. Hawaiian Islands was based on a congrid eel. They occur in waters from a few centimeters to several hundred meters depth, but most known species are shallow dwelling. Breeding probably CORACINIDAE takes place at great depths; the larvae are plank- The coracinids consist of a single genus, Cora- tonic. cinus, with two species. They are of moderate size There are few shallow-dwelling species of con- (to over 500 mm TL and 7 kg), free swimming, grids on the Pacific Plate. Conger cinereus ranges and limited to the southern and southeastern throughout the Indo-Pacific (the Hawaiian pop- coasts of Africa and Madagascar (Smith, 1953). ulation has been recognized as an endemic sub- They are here questionably considered to be species), and there is one other species of Conger warm-water fishes. on the Plate, an Hawaiian endemic. The monotypic Poeciloconger is known only from CORYPHAENIDAE Mauritius (J.E. Bohlke, pers. comm.), Madagas- car, Celebes, and Tahiti (Klausewitz, 1971). The The dolphins consist of one genus and two Tahitian record is based on a Museum Godeffroy species {Coryphaena hippurus, C. equisetis), approxi- specimen (Giinther, 1910), and for this reason mately 1500 and 570 mm maximum TL, that are alone might be suspect, but the rarity of this eel circumglobally distributed in warm waters leaves open the question of the Tahitian record. (Gibbs and Collette, 1959; Collette, Gibbs, and The garden eels (subfamily Heterocongrinae) Clipper, 1969). The species are primarily pelagic, are represented on the Pacific Plate by three but the young are often taken inshore. Coryphaena species, of which two are endemics: Heteroconger equisetis has been reported on the Pacific Plate lentiginosus from the Marquesas and Societies only from the Hawaiian Islands, but J.E. Randall (Bohlke and Randall, 1981), and Gorgasia ha- (in litt.) has obtained specimens from numerous waiiensis from the Hawaiian Islands (Randall and island groups on the Plate. Chess, 1980). The other Plate species, Heteroconger hassi, ranges from the Maldives east to the Mar- CREEDIIDAE shalls (B6hlke and Randall, 1981). There are two species of Ariosoma reported from The Creediidae was reviewed by Nelson (1978, the Pacific Plate: A. obud from the Marshall Is- 1979), who recognized seven genera and 12 spe- lands (Schultz, 1953a; also known from the Phil- cies. The identifications of several specimens he 38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY examined were left in doubt; hence, the species diate in dorsal- and anal-fin ray counts (which distributions and/or numbers of species as ex- Nelson considered important for distinguishing tracted here will undoubtedly change. The family the two species) between C. chameleontoculis and C. consists of very small fishes (maximum SL rang- tauensis. It seems possible, therefore, that the two ing from 29-72 mm) that live on and in sandy species may only represent geographic popula- and gravelly bottoms, and are restricted to the tions of a single species. However, the extensive Indo-Pacific. geographic gap between the two groups of pop- Apocreedia (monotypic) is restricted to southeast ulations, from which no specimens of Chalixodytes Africa; Creedia and Shizochirus (both monotypic) are known, may be indicative of a barrier that is are restricted to Australia; and Tewara (mono- effectively isolating the populations and allowing typic) is restricted to New Zealand. them to diverge. Interestingly, Allen and Steene According to Nelson (1978), Chalixodytes con- (1979), who did not refer to Nelson's paper, re- tains two species, one (C. chameleontoculis) re- ported C. tauensis from Christmas Island (Indian stricted to the Indian Ocean—Mauritius, Alda- Ocean), which is relatively close to Cocos-Keel- bra, Denis Island (Seychelles), southeast coast of ing. At my suggestion, Allen re-examined his Africa, and Cocos-Keeling Island—and the other Christmas Island specimens and informed me (in (C. tauensis), a widely distributed Pacific Plate litt.) that they also were intermediate in dorsal endemic—Tau and Tutuila islands (Samoa), Bi- and anal-fin ray counts between the two species kini, Rongerik, Rongelap, and Enewetak (Mar- of Chalixodytes. shalls), Pitcairn, Saipan, and Palau (the last two Crystallodytes (monotypic: C. cookei) is reported localities on the Philippine-Pacific Plate margin). from the Hawaiian Islands (an endemic subspe- Nelson's specimens of C. chameleontoculis from Co- cies) and Enderbury (Phoenix) and Tau (Sa- cos-Keeling, which is intermediate in location moan) islands (with a common endemic subspe- between the areas where the other populations of cies). Randall (1973) reported Crystallodytes sp. the two species of Chalixodytes occur, are interme- from Tahiti, and there are specimens in the
FIGURE 17.—Distribution of the fish family Congrogadidae (compiled by R. Winterbottom). NUMBER 367 39
USNM collections from McKean (Phoenix Is- sized (up to about 500 mm TL), benthic fishes lands), Vostok, Jarvis, and Maiden (all Line Is- that are found on sandy and muddy bottoms at lands). Randall (in litt.) informs me that he col- depths ranging from a few centimeters (particu- lected Crystallodytes at Easter Island; the identity larly some juveniles) to almost 2000 m. Some of his species has not been established. Crystallo- species enter freshwater, others are common in dytes appears to be a widely distributed Pacific brackish or marine waters. Eggs and larvae are Plate endemic genus (Figure 60). Its closest sister pelagic. group is undecided. The distribution of the two Cynoglossus (49 species, restricted to the Indo- subspecies of Crystallodytes cookei parallels the dis- West Pacific and upper west coast of Africa) and tribution of the two subspecies of Hyporhamphus Paraplagusia (three species, restricted to the Indo- acutus (Hemirhamphidae, q.v.). West Pacific) were revised recently (Menon, 1977, Limnichthys contains five species: two are re- 1980). Symphurus is distributed circumglobally in stricted to New Zealand and the Chatham Is- tropical and warm-temperate waters, but, other lands; one is widely distributed in, and limited than marginally, is known on the Pacific Plate to, the Indian Ocean; one (L. donaldsoni) is widely only from the Hawaiian Islands (Hawaiian ex- distributed on the Pacific Plate (but not in the ception; Figure 18), where two endemic species Gilbert Islands) and perhaps also occurs at Tai- occur at depths of about 100 m or more. wan; and one (L. fasciatus) is widely distributed The inability of cynoglossids to colonize the in the western Pacific with a lone occurrence in littoral portions of many islands of the Pacific the Gilbert Islands, on the Pacific Plate. Plate may be a consequence of improper ecologi- cal conditions. On the other hand, the true soles CYNOGLOSSIDAE (Soleidae), which appear to occupy habitats sim- ilar to those of the tongue soles, and may even be The tongue soles comprise three genera and collected together with them, are found at various perhaps over 100 species of small- to moderate- islands on the Pacific Plate. I have obtained
FIGURE 18.—Distribution of the fishes of the family Cynoglossidae (tongue soles). 40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY tongue soles in habitats on the coast of Florida with planktonic eggs and young. Only one genus, that were seemingly very similar to habitats on Diodon, has been revised in recent times (Leis, the coast of Ponape, where I obtained none. I, 1978; see also Kotthaus, 1979, for an additional, therefore, doubt that ecological conditions are probably invalid, species). Diodon comprises five responsible for the general absence of cynoglossids species, four of which occur nonmarginally on the from the Pacific Plate. Pacific Plate (Leis, 1978, figs. 8 and 14 are world distribution maps) as well as in the Indo-West DACTYLOPTERIDAE Pacific. The fifth species is restricted to southern The flying gurnards comprise two to four gen- Australia. era and five species of moderate-sized, benthic Chilomycterus, which has a circumglobal distri- fishes (to about 500 mm SL) with planktonic bution, has been reported nonmarginally from larvae and young. The family is distributed al- the Pacific Plate only from the Hawaiian Islands most circumglobally. One species is restricted to (C. affinis; Hawaiian exception). Chilomycterus af- the Mediterranean, Atlantic, and eastern Pacific, finis is also reported from Japan and the eastern and four are found in the Indo-Pacific. Only one Pacific, and Randall and Cea Egana (in press) of the Indo-Pacific species, Dactyloptena orientahs, report it from Easter Island. occurs on the Pacific Plate, and in that area has been reported nonmarginally only from the Ha- waiian, Society, and Tuamotu islands (Pacific DREPANIDAE Plate distributions similar to this one often char- The Drepanidae (sometimes considered a acterize endemic species; reexamination of speci- subfamily of the Ephippididae) consists of one mens from these localities might be of interest). genus and one or two species of moderate-sized Two of the remaining three species of flying (to about 400 mm TL), free-swimming fishes that gunards are widely distributed in the Indo-West range from the tropical west African coast east- Pacific, and have their eastern distributional lim- ward to Japan, New Guinea, and Samoa on the its along the western margin of the Philippine western margin of the Pacific Plate. The absence Plate. The third species (undescribed; W.N. Esch- of drepanids from the nonmarginal portions of meyer, pers. comm.) is known only from the the Pacific Plate is perhaps unexpected. Philippine Islands.
DASYATIDIDAE ECHENEIDIDAE The stingrays comprise four or five genera and There are four genera and eight species of a large number of viviparous, benthic species, remoras. These vary in maximum size from about some of which attain great size (breadth about 2 300-1000 mm TL. The species are usually found m, weight about 135 kg). There are few records attached by their sucking disks to free-swimming of stingrays from the Pacific Plate, possibly be- vertebrates, but some are also free swimming. All cause collectors avoid the group, most of the but one species (Echeneis naucratoides), which is species of which are extremely venomous. Prob- restricted to the Atlantic, are reported to be ably five species in four genera occur nonmargin- worldwide in distribution (Lachner, 1978), but I ally on the Plate. John E. Randall (pers. comm.) have been informed by E.A. Lachner (pers. recently obtained a large specimen of an uniden- comm.) that Remora australis has not been tified species of Urogymnus from Enewetak. This recorded from anywhere in the Indo-Pacific (ex- constitutes the only record of Urogymnus from the cept southeasternmost South Africa) and that Pacific Plate. Echeneis naucrates is either rare or absent in the DlODONTIDAE eastern Pacific. Probably six species of Echenei- The porcupinefishes comprise two or three gen- didae are present nonmarginally on the Pacific era and about 15 moderate-sized species of fishes Plate. NUMBER 367 41
ELEOTRIDIDAE lands on the Pacific Plate. Elops contains three to eight species (a revision is needed) of moderate The sleepers are a speciose family of small- to size (to about 1 m SL). Two of these occur moderate-sized (maximum about 500 mm TL) nonmarginally on the Pacific Plate. Tarpon is fishes occurring circumglobally in habitats rang- monotypic, very large (to about 2.4 m TL and ing from freshwater to coral reefs. The Indo-Pa- 160 kg), and restricted to the warm waters of the cific genera are in need of revision. Many marine Atlantic, both sides. Tarpon reaches the Pacific Indo-Pacific genera treated as eleotridids in the terminus of the Panama Canal from the Atlantic, literature belong in the family Gobiidae. Accord- but apparently has not established a breeding ing to Larson and Hoese (1980), there are only population in the eastern Pacific. Megalops is four genera of eleotridids confined to coral reefs monotypic, of moderate size (to about 550 mm in the Indo-Pacific {A Homier ode smus, Xenisthmus, TL), and ranges from the east coast of Africa to Calumia, and one undescribed); the other Indo- the Society Islands. Interestingly, Megalops has Pacific genera inhabit fresh and brackish water. not been reported from the Eastern Caroline, A Homierodesmus was described from a single Marshall, Gilbert, Ellice, Line, Tuamotu, or Ha- specimen from the Marshall Islands and placed waiian islands. Possibly, Megalops occurs only as in the Microdesmidae by Schultz (1966a). D.F. a straggler on the Pacific Plate, although Randall Hoese (in litt.) believes Aliomicrodesmus is a highly (in litt.) does not believe it to be a straggler in the aberrant eleotridid, and states that he has a spec- Society Islands. More records from Plate locali- imen from the northern Great Barrier Reef. I ties, including the occurrence of young, would be know of no other records for this genus. desirable. Calumia contains two reef-dwelling species (Lar- son and Hoese, 1980): C. godeffroyi, which ranges from the east coast of Africa to Tahiti, and C. EMMELICHTHYIDAE profunda, known from the Philippines, Moluccas The emmelichthyids comprise three genera (Ceram), western Admiralty, New Britain, Solo- and ten species of small (to about 500 mm SL), mon, and New Hebrides islands (Larson and moderately deep-dwelling fishes (Heemstra and Hoese, 1980; E.A. Lachner, unpublished data); Randall, 1977), which, as adults, are supra- hence, absent from the Pacific Plate nonmargin- benthic at depths of 100-400 m. The only records ally. for the Pacific Plate are either marginal or for the The widely distributed Indo-West Pacific ge- Hawaiian Islands, where three species occur (Fig- nus Butis, which occurs in rivers and estuaries, ure 19), Hawaiian exceptions. Of the three Ha- and mangroves, is known from the Pacific Plate waiian species, one also occurs at Japan, Aus- nonmarginally only at Ponape (USNM collec- tralia, and the Philippines (Randall, in press b), tions), which provides evidence for operation one also at Easter Island and Guam (Myers and of the Caroline Islands conduit. See also Shepard, 1981), and which may be limited to the "Addendum." Pacific Plate and its margins, and the third is either an Hawaiian endemic or also occurs at ELOPIDAE Okinawa (Heemstra and Randall, 1977). Because of the difficulty of collecting emmelichthyids, (including Megalopidae) their occurrences as Hawaiian exceptions may be The tarpons and ladyfishes comprise three gen- a collecting artifact. The family seems to be re- era of free-swimming fishes with planktonic stricted somewhat to subtropical and temperate young that enter and develop in shallow coastal areas in the Indo-West Pacific, which may ex- areas and estuaries. The adults may ascend rivers. plain its general absence from the Pacific Plate Elops is, perhaps, circumglobally distributed, but islands. Nonmarginal Pacific Plate records are has not been reported from the easternmost is- worth reporting. 42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 19.—Distribution of the fishes of the family Emmelichthyidae (stars indicate unspecified localities in certain areas of the southeast Brazilian (Zavala-Camin, 1981) and west African coasts.
ENGRAULIDIDAE Norman's (1966) erroneous synonymy of Pterapo- gon kauderni Koumans (1933) with Enoplosus arma- The anchovies are a circumglobally distributed tus. Pterapogon kauderni bears a striking, but super- group of small, free-swimming fishes (largest spe- ficial, similarity (see Koumans, 1933, pi. 1: fig. 1) cies attaining about 350 mm TL) that inhabit to the way one might expect the young of E. coastal areas. Many species appear to live pri- armatus to appear, but Koumans was correct in marily in fresh or low salinity waters. There are considering Pterapogon to be an apogonid (see also 15 genera and 128 species of anchovies, and eight Fraser, 1972). genera and 62 species occur in the Indo-Pacific (PJ. Whitehead, in litt.), but only two genera (Stolephorus, Encrasicholind) and four species occur EPHIPPIDIDAE nonmarginally on the Pacific Plate (G. Nelson, in (including Platacidae) litt.). The spadefishes are perhaps a circumglobal ENOPLOSIDAE family, but they are unreported from the eastern- most islands on the Pacific Plate. The family There is but one species of enoplosid, Enoplosus includes six genera and about ten species of mod- armatus, a small fish attaining about 250 mm TL, erate-sized (to about 9 kg), free-swimming fishes. and primarily restricted to the cooler waters of Four genera and about seven species occur in southern Australia, but occasionally reaching (and are restricted to) the Indo-Pacific, but only southern Queensland and entering freshwater one genus, Platax, and only one or two of its (Lake, 1971). Nelson (1974) included Celebes in approximately five species, occur nonmarginally the range of Enoplosus, based undoubtedly on on the Pacific Plate. NUMBER 367 43
EXOCOETIDAE wara, Islands, adjacent to the margin of the Pa- (not including Hemiramphidae) cific Plate, and Dor (1970), reported it from the Red Sea. Reports of the occurrence of Apolectus in There are about eight genera and perhaps as Hawaii all appear to emanate from Matsubara many as 100 species-group taxa of flyingfishes, of (1955:565). Matsubara's listing is probably in which the Indo-Pacific forms are most poorly error (see also Menidae). I know of no other known systematically. The family is circumglob- nonmarginal Pacific Plate record for Apolectus. ally distributed and the species are epipelagic and, in general, widely distributed. The young GERREIDAE are often taken inshore. Possibly five genera and 15 species occur nonmarginally on the Pacific The mojarras are a family of small, free-swim- Plate. ming, epibenthic fishes (maximum size about 400 mm TL) that are poorly known taxonomically. There are about six genera and perhaps 40 spe- FlSTULARIIDAE cies. The species are primarily marine but often The cornetfishes consist of a single, circumglob- enter brackish or freshwater, and are found on ally distributed genus, Fistularia, and four species. the warm coasts of all oceans; however, they are The species are free swimming and attain a max- reported nonmarginally from the Pacific Plate imum length of 1-2 m. Fritzsche (1976) revised only from the Eastern Caroline, Gilbert, Ellice, Fistularia and presented a distribution map for and Marshall islands (Fowler, 1928, 1931, 1934). the species. One species is restricted to the tropical These records probably involve only species of eastern Pacific and one to the tropical Atlantic Genes. Randall (1973) discounts Jordan and (both sides). One species occurs from the east Seale's (1906) report of Genes argyreus (as Xystaema African coast to the tropical coasts of the eastern argyreum) from Tahiti. Pacific. The fourth species, F. petimba, is almost circumglobal and has an Hawaiian exception GlNGLYMOSTOMATIDAE distribution: both sides of the tropical Atlantic; Indo-West Pacific from Madagascar to Japan, This family is discussed under the Orectolobi- Philippines, and Australia; and Hawaiian Islands dae. (only nonmarginal Pacific Plate occurrence). Al- though Fistularia is not present in the Mediterra- GlRELLIDAE nean, it is reported from the Miocene of that region (Arambourg, 1925). The girellids are a small family containing one or two genera and about 15 species of moderate- sized fishes (to about 625 mm TL and 8 kg) that FORMIONIDAE (= APOLECTIDAE) are absent from the Pacific Plate. These fishes are There is but a single, monotypic genus in the questionably included here as a warm-water Formionidae, Apolectus (= Formio = Parastro- group. The family is antiequatorially distributed, mateus), a free-swimming fish probably not attain- and occurs on the coasts of Japan, China, south- ing more than 300 mm TL, but occasionally ern Australia (as far north as southern Queens- stated to reach 600 mm TL. Witzell (1978, fig. 8) land), Lord Howe and Norfolk islands, New Zea- reported on the distribution of Apolectus, which land, Easter Island, California to, and including, extends from eastern Africa eastward to Japan, the Gulf of California, Galapagos, Chile and its Philippines, and eastern Australia. Additionally, offshore islands, Peru, and the Cape Verde and Zama and Fujita (1977) give citations to the Canary islands in the eastern Atlantic (D.F. occurrence of Apolectus in the Bonin, or Ogasa- Hoese, in litt.). Lee and Chang (1981) reported 44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Girella punctata from Batan Island, which lies mid- from several Indonesian localities and from Ra- way between Taiwan and Luzon, and is politi- roia, Tuamotus (Briggs, 1955). Liobranchia stria is cally part of the Philippines. I have not included reported only from Saipan, Marianas, and Bikini, the Girellidae as part of the Philippine fauna for Marshalls (Briggs, 1955; Schultz, 1966b). Liobran- the purposes of Tables A and B (Appendix 2). chia, is possibly a Pacific Plate endemic genus (Figure 60). Randall (in press b) obtained Lepad- GLAUCOSOMATIDAE ichthys frenatus from the Pitcairn Island group; this species is otherwise known only from widely scat- This family contains a single genus, Glaucosoma, tered localities in the western Pacific. Briggs and four species (Arai and Fujii, 1981) of free- (1955:215, map) inadvertently indicated the pres- swimming fishes that attain a maximum total ence of Lepadichthys \minor\ in the Ellice Islands, length of about 450 mm. The species have been on the Pacific Plate; Samoa was intended, as reported only from the coasts of Japan, China, indicated in his materials list. Philippines, and Australia. In Queensland, it is Most species of tropical Indo-Pacific clingfishes known only from the offshore southern portions are tiny, and more will probably be found to of the coast; therefore, the overall distribution of occur on the Pacific Plate. Glaucosoma appears to be antiequatorial.
GOBIIDAE GOBIESOCIDAE The gobies are probably the largest family of The clingfishes comprise about 40 genera and fishes, perhaps containing as many as 2000 species 110 species of small (largest species is about 300 (Hoese and Allen, 1977). Most of the species are mm TL, but most are under 75 mm), benthic, small, and among these are the smallest verte- nest-building fishes with planktonic larvae. The brates (some species of Eviota mature at less than species occur in a variety of habitats, including 10 mm SL; E.A. Lachner, pers. comra.). Among freshwater rivers and coral reefs. Several species the largest gobies is the Japanese genus Synecho- live commensally with crinoids. Clingfishes occur gobius, which attains a total length of 500 mm in all oceans, as well as the Mediterranean and (Takita, 1975). Gobies are essentially benthic or Black Sea, and are generally widely distributed suprabenthic in habit. Some species are restricted along continental coastlines. Most of the genera, to freshwater, some are estuarine, but most are but only about one-third of the species, occur in reef or near-reef inhabitants. Hoese and Allen the Indo-Pacific. Only four species are known (1977) estimate that 400-600 species of gobiids from the nonmarginal areas on the Pacific Plate, occur on the coral reefs of the Indo-Pacific. and on the Plate each is known from a single The taxonomic limits of gobiid genera are locality. One of these species, Trachelochismus pin- poorly known and very few of the Indo-Pacific nulatus (otherwise restricted to New Zealand), genera have been revised in recent years. The list questionably occurs at the Chatham Islands, of Pacific Plate genera (estimated 150 species; which, although nonmarginal on the Pacific Table C, Appendix 2) was compiled from the Plate, are on the continental shelf of New Zea- literature, USNM collections, and in consultation land. Hence, T. pinnulatus is excluded from further 1 with D.F. Hoese (see also "Addendum"). Among consideration here. Pherallodus indicus is reported the genera, Lentipes (adults of which inhabit fresh- water in Hawaii), with one species (Maciolek, Hutton (1896) described Crepidogaster simus (a synonym of T. pinnulatus) from "Lyttleton Harbor [New Zealand] and elochismus pinnulatus has been collected at New Zealand lo- Chatham Islands." Information in the description appears calities many times, but has not been reported to have been to refer to a single specimen. If only one specimen was collected at the Chatham Islands subsequent to the initial involved, it could not have come from both localities. Trach- report. NUMBER 367 45
1978), and Vitraria (coral-reef inhabitant), with One of three species of Gobiopsis, not included in one species, appear to be Pacific Plate endemic Gobiopsis sensu stricto, reaches from the Indo-West genera (Figure 60). The presence ofLentipes in the Pacific to the Gilbert Islands (Lachner and Ryukyus is based on a species described by Sakai McKinney, 1979). Signigobius, monotypic, also a and Nakamura (1979); its generic allocation reef inhabitant (Hoese and Allen, 1977), is dis- should be verified. tributed from the Philippine and Palau islands to Kelloggella was revised by Hoese (1975) and a the Great Barrier Reef and Solomon Islands, and new record added by Kuramochi (1980). Hoese is, thus, limited in the east by the margin of the (in litt.) informed me of an additional species Pacific Plate. Oxuderces, monotypic, a muddy bot- (undescribed) and of changes (pers. comm.) in tom inhabitant, ranges from India to the Gulf of the reported distributions of two described spe- Carpentaria, Australia (Springer, 1978, and Aus- cies. The distribution of the five Kelloggella species tralian Museum collections). is given in Figure 20. Except for endemic genera, Feia was discussed by Lachner and McKinney Kelloggella is unique among fishes in having more (1979). They recognized one species in the genus, Pacific Plate species than Indo-West Pacific spe- but noted that their Indo-West Pacific specimens cies. (Mozambique, Seychelles, Banda, and Great Bar- Among the reasonably well-known Indo-Pa- rier Reef) were separable from their Rapa Island cific goby genera that are excluded from the (Pacific Plate) specimens, based on meristics and Pacific Plate is Gobiopsis (sensu stricto), with ten pigmentation, indicating that two taxa are pos- species (Lachner and McKinney, 1978, 1979), sibly involved. which are often collected on coral reefs. Gobiopsis Eviota comprises about 60 species of dimin- sensu stricto is distributed from southeastern Af- utive coral-reef gobies, all restricted to the Indo- rica to the western margin of the Pacific Plate. Pacific (S. Jewett, pers. comm.). Lachner and
• CARDINALIS
U CENTRAL IS
* NEW SPECIES
AOLIGOLEPIS
O TRICUSPIDATA
FIGURE 20.—Distributions of the species of the gobiid fish genus Kelloggella ("new species" pet D.F. Hoese, in litt.). 46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Karnella (1980) revised 31 of the species, and Randall and Allen's study whether N. decora and Karnella and Lachner (1981) revised three others. N. helfrichi are sister species, but these two species Of the 34 species revised, 23 reach the western were keyed out together and show greater mor- margin of the Pacific Plate from the west, but phological similarity to each other than either only 11 of the 23 (48%) extend nonmarginally does to N. magnifica. The evidence is slight, but onto the Plate. One of the 11, £ lachdeberei, is there is a good chance, in the present case, that known on the Pacific Plate only from Ponape an Indo-West Pacific species is the sister species (USNM collections) and Truk, Eastern Caroline of a Pacific Plate endemic, and that the two Islands, providing evidence favoring operation of species have undergone slight dispersal since their the Caroline Islands conduit. Two species are divergence, which would account for their sym- Pacific Plate endemics: E. epiphanes, from Ha- patry along the Pacific Plate margin. waiian, Johnston, and Christmas (Line) islands; Periophthalmus, a mangrove inhabiting genus in E. disrupta, from the Tonga, Samoan, Society, and need of revision, occurs on the tropical west Af- Tuamotu islands. rican coast as far south as Angola. It reappears Nemateleotris contains three species of coral-reef on the east African coast and ranges westward to gobies (Randall and Allen, 1973): N. decora, which the western margin of the Pacific Plate, and is ranges from the western Indian Ocean to the known nonmarginally on the Plate only from western margin of the Pacific Plate; N. helfrichi, Ponape, Eastern Caroline Islands, which provides which is a widely distributed Pacific Plate en- evidence for the operation of the Caroline Island demic; and N. magnijica, which completely over- conduit. Disjunct west African distributions of laps the range of the other two species (Figure otherwise Indo-Pacific fishes is a repeated distri- 21). The three species are sympatric in the Palau bution pattern (for some additional examples see and Samoan islands, both along the margin of Lethrinidae). the Pacific Plate. It is not possible to decide from Discordipinna is monotypic, diminutive (to 20
o DECORA •HELFRICHI * MAGNIFICA
FIGURE 21.—Distributions of the three species of the gobiid fish genus Nemateleotris. NUMBER 367 47 mm SL), and a reef inhabitant. It ranges from ing that the Scripps Institution of Oceanography the Red Sea to the Hawaiian and Marquesas marine vertebrate collection has a specimen from islands (Randall, 1981a). the Juan Fernandez and San Felix Ridge, off Chile. Some authors believe that there is only one GOBIOIDIDAE species of gonorynchid, but the north-south dis- (including Taenioididae) junct distribution of the family suggests the pos- sibility that two species or more are involved. The gobioidids are small, elongate fishes (max- imum size about 470 mm SL) living benthically GRAMMATIDAE on muddy bottoms, typically in brackish bays, The monophyly of the Grammatidae is open although the same species may be taken offshore to serious question. For instance, Nelson (1974) in marine waters and well up freshwater rivers. included the following in the Grammatidae: Pseu- There are several genera, but probably fewer than dochromichthys, which is a synonym of the plesiopid 20 species, and these occur, variously, along the genus Plesiops (Aoyagi, 1943; Schultz, 1943); Frau- warm coastal areas of the eastern Pacific, eastern della, which is a plesiopid (G.D. Johnson, pers. and western Atlantic, and Indo-West Pacific from comm.; incidentally, the Plesiopidae also appears Africa to Guam and New Guinea. Prior to my to be polyphyletic); and Pseudocrenilabrus, which is 1980 collecting trip to Ponape, Eastern Caroline a cichlid (Trewavas, 1973). Even without these Islands, gobioidids were unknown on the Pacific three genera there is little evidence to unite the Plate nonmarginally. I obtained two genera remaining five genera, and perhaps 20 species (Table C, Appendix 2) in a single collection at that could be considered in the family (as most Ponape, in a coarse muddy area adjacent to a recently defined by Bohlke, 1960). The western river mouth and mangrove shore. The Caroline Atlantic genera {Gramma, Lipogramma) are coral- Islands chain apparently serves as a conduit per- reef forms of shallow to moderate depths. Stig- mitting gobioidids to disperse to the Pacific Plate. matonotus is known only from coastal Western I doubt that they occur further east than Kosrae Australia. Grammatonotus is known from deep wa- (Kusaie) at the end of the chain. See also ter off Hawaii (Hawaiian exception; only non- "Addendum." marginal Pacific Plate occurrence of Grammati- dae), the western Indian Ocean (based on the GONORYNCHIDAE type species of Parabarossia Kotthaus, 1976, which The Gonorynchidae consists of a single genus, Kotthaus assigned to the Plesiopidae), Japan, Gonorynchus, and perhaps as many as six species Philippines, New Caledonia (G.D. Johnson, pers. (the genus has not been revised). The gonoryn- comm.), and the Kei Islands, Indonesia. The chids attain a maximum size of about 600 mm nominal species of Callanthias all appear to belong TL and are suprabenthic on sandy bottoms. The in the Grammatidae, as considered here. They family occurs nonmarginally on the Pacific Plate have been reported, variously, from southern Af- only at the Hawaiian islands (Hawaiian excep- rica, southern Australia, New Zealand, Japan, tion); it is otherwise antitropically distributed. the Mediterranean-eastern Atlantic, and the Gonorynchids have been reported most often Nasca Ridge (G.D. Johnson, pers. comm.) and from the Indo-West Pacific: Japan and Korea San Felix and Juan Fernandez islands in the south to the Pescadores Islands, southern Aus- eastern Pacific, and are, thus, bitemperately dis- tralia (as far north as southern Queensland), New tributed. Zealand, Reunion, and southeastern Africa, but they also occur at St. Helena in the eastern GYMNURIDAE Atlantic (Penrith, 1976), and Mead (1970) and The butterfly rays probably consist of a single McCosker (1971) quote R.H. Rosenblatt as stat- genus, Gymnura, and fewer than ten, viviparous, 48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY species. The largest species attain over 2 m Plate (absent from the Pacific Plate nonmargin- breadth. The family is circumglobally distrib- ally). Eleven of the haemulid genera are restricted uted, except that it is apparently absent from the to the western hemisphere; two occur only in the Pacific Plate other than marginally, a distri- eastern Atlantic, one (Diagramma) only in the bution pattern duplicated by most elasmobranch Indo-West Pacific (absent from the Pacific Plate), groups. one {Parapristipoma) only in the eastern Atlantic Fowler (1928, 1941) placed Raia pastinaca vari- and Indo-West Pacific (absent from the Pacific ety altavela Forster (not Linnaeus), from Tana Plate), and one (Plectorhynchus) in the Mediterra- (New Hebrides), marginally off the Pacific Plate, nean, eastern Atlantic, and throughout much of into the synonymy of Gymnurapoecilura (Shaw), an the Indo-Pacific (but absent from the Hawaiian Indo-West Pacific species. Fowler did not give his Islands), where it is represented nonmarginally reasons for the synonymy, which I believe was in on the Pacific Plate by five species at most. The error, and also mistakenly placed Tana in the family has, hence, only token representation on Society Islands, which are well out on the Pacific the Pacific Plate. Plate. Forster's (1844) description was based on an observed specimen that, apparently, was not HARPADONTIDAE retained. Forster included in his description a statement that there was a single row of small The Bombay ducks comprise a single genus, spines on the middle of the dorsum of his speci- Harpadon, and four species of small- to moderate- men. Gymnurids lack such a feature. Forster's sized fishes (to about 710 mm TL) of commercial specimen was probably a dasyatid. In the Indo- importance. These fishes are found in close-shore West Pacific, gymnurids range from the Red Sea waters, estuaries, and relatively deep continental and east coast of Africa eastward to Japan, and shelf waters (R.K. Johnson, pers. comm.) of the south to the Philippines and eastern Australia. Indo-West Pacific, but are absent nonmarginally There are no records of gymnurids from New from the Pacific Plate. Sulak (1977) placed Guinea or any other islands along the southwest- Saurida (which in my study is included in the ern margin of the Pacific Plate. Based on present Synodontidae) and Harpadon in a subfamily, Har- information, I do not believe that gymnurids padontinae, of the Synodontidae. The constitu- occur in the New Hebrides or on the Pacific Plate. tion of these family-group taxa in my study re- Bigelow and Schroeder (1953:398) probably fol- flects only common usage. lowed Fowler (1928, 1941, 1944) in including Harpadon has a spotty distribution. It appears Polynesia in the range of gymnurids. to be present along the northeast and northwest coasts of India, but absent from around southern India and Ceylon, and ranges eastward through HAEMULIDAE the Bay of Bengal to Sumatra, Java, Borneo, New There are 17 genera (Johnson, 1980) and about Guinea, northern Australia, Philippines (Rau 100 species of grunts. The species are of small to and Rau, 1980), and northward to the Yellow moderate size (to about 610 mm TL), free swim- Sea as far as Korea. A number of authors list ming, suprabenthic, and usually associated with Zanzibar in the range of Harpadon, the earliest of coral reefs or rocks. Although the family is cir- which that I can locate is Bleeker (1866-1872). cumglobally distributed, no genus is that widely These reports appear to be second hand and, I distributed. Pomadasys, the most widely distrib- believe, erroneous. Playfair and Giinther (1866) uted genus of grunts, is found in the eastern did not report Harpadon from Zanzibar, nor did Pacific, western Atlantic, Mediterranean, eastern they in their subsequent papers on additions to Atlantic, and throughout the tropical Indo-West the Zanzibar fish fauna. Harpadon is frequently Pacific up to the western margin of the Pacific dried for commercial purposes. In view of the NUMBER 367 49 extensive dhow movements around the Indian genera with 40 species, three freshwater genera Ocean it could be expected that dried Harpadon with eight species, and one genus {Zenarchopterus) would find its way to Zanzibar markets (and with about 15 species, some of which are restricted other ports of call) and be mistakenly believed to to freshwater. Only four of the marine genera, have originated in that area. represented by about eight species, and one eu- ryhaline species of Zenarchopterus (Z. dispar) occur on the Pacific Plate nonmarginally. Two of the HEMIRAMPHIDAE eight marine species (Hyporhamphus acutus, Hemi- The halfbeaks are a family of small- to mod- ramphus depauperatus) are widely distributed Pacific erate-sized, nearshore, epipelagic fishes present in Plate endemics. all tropical and warm-temperate seas. Some spe- Hyporhamphus acutus consists of two subspecies: cies enter brackish and freshwater. The family one at Johnston and Hawaiian islands, the other consists of 12 genera and 75 species (B.B. Collette, widely distributed on the Pacific Plate, but absent pers. comm.). Most species are oviparous, but from Johnston and Hawaiian islands (Figure 22; some are viviparous. Eggs of marine forms bear Collette, 1974). The taxa Collette (pers. comm.) numerous filaments and are often found attached considers to be most closely related to H. acutus to floating objects. N.V. Parin, B.B. Collette, and are essentially absent from the Pacific Plate, al- Yu. N. Shcherbachev (1980) revised the Indo- lopatric, and consist of a group of three species Pacific halfbeaks and presented distribution maps restricted to southern Australia, Lord Howe, Nor- for the species (see also Collette, 1976, for a folk, New Zealand, and the Chatham Islands. distribution map of Rhynchorhamphus, four species, Specimens of Hyporhamphus a. acutus reported by which ranges from the Gulf of Aden to New Collette (1974) for the Tonga Islands, which are Guinea, on the margin of the Pacific Plate). marginally off the Pacific Plate, may not have In the Indo-Pacific there are seven marine come from Tonga as the Tongan locality is en-
FICURE 22.—Distributions of the two subspecies of the hemiramphid fish, Hyporhamphus acutus. 50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY tered in the USNM catalog as questionable; how- from the Eastern Caroline Islands. I questioned ever, the occurrence of H. a. acutus at Tonga Parin about this exceptional record, and after would not be unexpected. reexamination he wrote that the specimens were Hemiramphus depauperatus is known only from misidentified and belonged in 0. micropterus. the Hawaiian, Line, Society, and Marquesas is- I collected Zenarchopterus dispar in the rivers of lands (Parin et al., 1980, fig. 34). Its closest rela- Ponape, eastern Caroline Islands, above the level tive is allopatric, either H. saltator, of the eastern of tidal influence, as well as in the mangrove Pacific, or H. lutkei (B.B. Collette, pers. comm.), areas along seashore. These specimens represent which is widely distributed in the Indian and the only nonmarginal Pacific Plate records of western Pacific oceans as far east on the Pacific Zenarchopterus and offer evidence favoring opera- Plate as the Gilbert, Marcus, and Kapingamar- tion of the Caroline Islands conduit. angi islands.
Oxyporhamphus is a circumglobal, oceanic, epi- HEMISCYLLIIDAE pelagic genus that contains two species. Oxypor- hamphus micropterus consists of two subspecies, one This family is discussed under the Orectolobi- in the Atlantic (both sides and centrally) and one dae. in the Indo-Pacific and eastern Pacific. Oxypor- hampus convexus also contains two subspecies: 0. c. HETERODONTIDAE bruuni, which occurs only in the Red Sea and northwest Indian Ocean, and 0. c. convexus, which The horn, or Port Jackson, sharks comprise a ranges (allopatrically) from the east coast of Af- single extant genus (Heterodontus) with eight spe- rica to the western margin of the Pacific Plate. cies variously restricted to the Indo-West Pacific Parin et al. (1980, fig. 47) included a single and Eastern Pacific (Figure 23). These sharks are nonmarginal Pacific Plate record for O. c. convexus, relatively small (less than 1.5 m TL), sluggish,
FIGURE 23.—Distribution of the shark family Heterodontidae (horn sharks) (closed circles indicate recent records; solid stars indicate fossil records; open star represents record based on an egg case). NUMBER 367 51 oviparous, and benthic in shallow marine waters 600 mm, but specimens 300-400 mm are not (to a maximum depth of about 170 m). The common), suprabenthic, live primarily around combined geographic distribution of the extant coral reefs and rocks, and have planktonic larvae. species is primarily antiequatorial. A revision (which I have not seen) of Ostichthys Heterodontids (including Heterodontus) first ap- and description of a new genus is in press (Ran- pear in the Jurassic of Europe (Casier, 1954), and dall, Shimuzu, and Yamakawa, in press). the fossil record, so far as I have been able to Among the squirrelfishes, the species of Myri- extract it (Figure 23), indicates the presence of pristis have received the most attention systemat- heterodontids in eastern North America and west- ically (Greenfield, 1965, 1968, 1974; Randall and ern Europe (Atlantic) from the Mesozoic to the Gueze, 1981). Even so, there is much useful work Eocene or Oligocene. Romer (1945, 1966) lists to be done on the classification, relationships, and heterodontids as occurring during the Miocene in distribution of the species. There are about 19 South America, but I have been unable to locate species of Myripristis in the Indo-Pacific. Most of the origin of Romer's report or the specific local- these species are widely distributed, and include ity. one species that ranges into the eastern Pacific. J.A.F. Garrick (in litt.) states that there is only Four species appear to be restricted to areas west one firm (nonfossil) record of Heterodontus from of the Pacific and Philippine plates, and four New Zealand, from a gill net in shallow water appear to be restricted to the Pacific Plate (Figure near Makara, Wellington, in 1954. He considers 24), of which, three are widely distributed Pacific the occurrence to have been the result of strag- and Philippine plate endemics (Table 2). Twenty- gling. The same situation probably explains five other species are present nonmarginally on Bleeker's (1856, 1860) unrepeated records of Het- the Pacific Plate. Greenfield (1968, 1974) dis- erodontus zebra from Amboina and Celebes, if thesecussed relationships of the species in noncladistic records are valid; however, Fourmanoir and Nhu- terms. He related the widely distributed Pacific Nhung (1965) reported H. zebra to be common Plate endemics most closely to other widely dis- during the summer at Nha-Trang, Viet Nam. tributed Pacific Plate endemics or to widely dis- The egg cases of some heterodontids have tendrils tributed Indo-Pacific species. that may become entangled in marine plants, which in turn may spread about during storms. ISONIDAE Heterodontid egg cases have been reported from The isonids are a small, poorly known family areas where no free-living heterodontids are comprising two genera and probably fewer than known. For instance, Whitley (1940) reported an ten species of small (to about 50 mm SL) fishes egg case from Moa Island, north Queensland that are often taken in the heavy surf zone (but (Figure 23), whereas the sharks themselves have I have collected them in protected areas in Tai- not been taken farther north than southern wan). They have been collected along the coasts Queensland on the Australian coast. Hatching of southern and southeastern Africa, West Paki- time has been reported variously to take from five stan, northwestern India, southern Japan to to nine months, and an egg case would have southern Taiwan, the Admiralties, southern New considerable time to be dispersed before hatching. South Wales (a doubtful record also from Free- The narrow distributions of the living heterodon- mantle, Western Australia), Samoa, Hawaii, Pit- tids, however, lead me to believe that the species cairn, Rapa, and both sides of the southern end have not dispersed widely in recent times. of South America (W. Ivantsoff, B. Chernoff, and J.E. Randall, pers. comm.; R.C. Wass, ms.). HOLOCENTRIDAE The squirrelfishes are a circumglobally distrib- ISTIOPHORIDAE uted family comprising about 50-75 species. The The billfishes, including the sailfishes and mar- species are small (reported to attain an SL of over lins, are large, primarily open ocean forms with 52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
planktonic larvae and widespread distributions. They are included here because they are caught occasionally close to shore. Such records for two istiophorid species have been reported for the Pacific Plate.
KRAEMERIIDAE The kraemeriids, a poorly known group taxo- nomically, are small (not exceeding 40 mm SL), benthic fishes consisting of three genera (Krae- meria, Gobiotrichonotus, Parkraemeria) and about nine species. They occur in sandy areas close to shore, and although primarily marine, also occur in freshwater. They are distributed from the east coast of Africa (Winterbottom, 1976) eastward to Japan, Australia, New Guinea (USNM collec- tions), Hawaiian Islands, and Tahiti.
KUHLIIDAE The kuhliids comprise a single genus (Kuhlia), and about six species of small fishes (to about 560 mm TL). Most authors include Nannoperca, Nan- natherina, and Edelia in the Kuhliidae, but these three Australian freshwater genera belong in the Percichthyidae (G.D.Johnson, pers. comm.). The kuhliids are free swimming and suprabenthic in habit. Both young and adults may enter fresh- water. The kuhliids range throughout the warm portions of the entire Indian and Pacific oceans. A revision of the family is needed.
KURTIDAE The kurtids contain a single genus, Kurtus, and two species of small fishes (maximum size about 600 mm TL). The kurtids are primarily fresh and brackish water inhabitants that range hundreds of kilometers upstream (Lake, 1971). Males carry the eggs attached to a special process on top of the head. The kurtids, which are absent from the Pacific Plate nonmarginally, are distributed from
FIGURE 24.—Distributions of certain species of the holocen- trid fish genus Myripristis. NUMBER 367 53
Ceylon and the east coast of India eastward which island is marginally tropical, and one other through Indonesia, as far as Borneo and New questionable record, labracoglossids are unknown Guinea. In Australia they range from northern from the Pacific Plate nonmarginally. Lindberg Queensland northward around Australia to and Krasnyukova (1971) stated that labracoglos- northern Western Australia. They are often listed sids were "indicated for the Vostok Islands [sic] as occurring in "China," but I find no verifiable in the South Pacific" [translation]. Vostok Island reference for China, and doubt that they occur is one of the Line Islands, which are tropical. I even as far north as southern China. Kurtids are have been unable to locate the source of the not reported from South Viet Nam or the Phil- Vostok record, which does not appear to have ippines. A fishery for kurtids exists in the Bay of originated with Lindberg and Krasnyukova. The Bengal (Kuthalingam, 1967). possibility comes to mind that the "Vostok Is- lands" are those islands that exist off Vladivostok, KYPHOSIDAE Russia, which is possibly in the range of labra- coglossids. The chubs comprise three genera and fewer than ten species of moderate-sized (to over 600 LABRIDAE mm TL), free-swimming fishes. Hermosilla (mon- otypic) is restricted to the eastern Pacific, and The wrasses comprise over 50 genera and per- Kyphosus, with several species, is circumglobally haps as many as 500 species of small-to-large, distributed on warm coasts. Randall (1961) and free-swimming, epibenthic, or benthic fishes. The Rosenblatt et al. (1972) reported on the peculiar family is distributed circumglobally in tropical distribution of the monotypic Sectator, which is and warm-temperate marine coastal waters. known only from the Hawaiian and Society is- There are no freshwater species and only a few lands in the central Pacific, and from Cabo San species occur under estuarine conditions. The Lucas, Baja California, south to Isla La Plata, wrasses form a conspicuous element of the coral- Ecuador, in the eastern Pacific. reef fish fauna. The following discussion treats only those genera for which the Indo-Pacific spe- cies are reasonably well known. LABRACOGLOSSIDAE Anampses is restricted to the Indo-Pacific and The labracoglossids are a poorly known family consists of 12 species (Randall, 1972a, 1974). Only consisting of two genera and perhaps fewer than six species occur on the Pacific Plate nonmargin- five species. The species probably do not much ally, and two of these are endemic to the Ha- exceed 250 mm TL, are free swimming, and are waiian Islands, including Johnston Island; the primarily antiequatorially distributed. They are other four species occur also in the Indo-West common in Japan (Honshu; also Bonin Islands Pacific. Six species are limited to various parts of on margin of Pacific Plate), and occur as far the Indo-West Pacific. south as the Philippines. They are absent from Bodianus, a circumglobal genus, consists of 29 the Indo-Malayan area but reappear to the south species (Gomon, 1979; Lobel, 1981b), of which 22 in New South Wales and New Zealand, and occur in and are limited to the Indo-Pacific (in- Norfolk, Lord Howe, Rapa (Randall and Sinoto, cludes one species known only from a specimen 1978), and Easter islands. Bruce A. Carlson (ms.) obtained in the Tokyo fish market, and seven questionably assigned this family to a specimen species restricted to the Indian Ocean). Of the he obtained in Fiji. As labracoglossids might be Indo-Pacific species, only seven occur nonmargin- expected to occur in Fiji, I have listed the family ally on the Pacific Plate, and of the seven, two as present in Fiji in Table A (Appendix 2); cor- species, two of the three subspecies of a third roboration of the family in Fiji is desirable. species, and one of the two subspecies of a fourth With the exception of Bathystethus at Rapa, species are endemic to the Plate. The endemics 54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY are distributed as follows: one species (B. sangui- Epibulus contains two species, one of which is neus) is endemic to the Hawaiian Islands and one widely distributed in, and restricted to, the Indo- (B. prognathus) is known only from Fanning Island, Pacific, and the other (undescribed) is known Line Islands; one subspecies (B. bilunulatus albo- only from the Philippine and Palau islands (J.E. taeniatus) is endemic to the Hawaiian and John- Randall, in litt.). ston islands, and a conspecific subspecies (B. bi- Labrichthys is monotypic and is distributed from lunulatus busellatus) is limited to the Marquesas, eastern Africa to Samoa and the Marshall Islands, Henderson, and Ducie islands (the third conspe- on the Pacific Plate (Randall and Springer, 1973). cific subspecies, B. b. bilunulatus, is distributed Randall and Springer considered a report (Fow- from the east coast of Africa to Japan, Philip- ler, 1931) of Labrichthys from Tahiti to be ques- pines, Moluccas, New Guinea, and New Cale- tionable. donia, hence, limited to continental plates); one Labroides consists of five small species of cleaner subspecies (B. loxozonus trotteri) is distributed in fishes and is limited to the Indo-Pacific from the the Line, Marquesas, Tuamotu, and Austral is- east coast of Africa to the Pitcairn Island group lands (the other conspecific subspecies, B. I. loxo- (Randall, 1958; Randall and Springer, 1975). zonus, ranges from Vietnam to Japan, Taiwan, The distributions of two species, L. dimidiatus (well and the Philippines, is absent from these areas known as the model in a mimetic association with eastward to the Marshall Islands, where it occurs, the blenniid fish Aspidontus taeniatus) and L. bicolor, and southward to New Caledonia and the Tonga cover, in general, the entire range of the genus Islands, where it also occurs, and is, surprisingly, (both species, however, are absent from the Ha- also present in the Tuamotus together with the waiian Islands). The three remaining species of conspecific B. I. loxozonus). The distribution of one Labroides are more localized and allopatric with species, B. vulpinus, appears to be antiequatorial, regard to each other (Figure 25), but may occur and is an Hawaiian exception as well: Ryukyu sympatrically with either or both of the two and Hawaiian islands in the north, southeast and widely distributed species. Labroides rubrolabiatus is southwest Australia, New Zealand, Lord Howe, a widely distributed endemic on the Pacific Plate, and Easter Island in the south. but is absent from Plate localities where its pre- Lobel (1981b) appears to have related the Pa- sumed closest relative, L. pectoralis (Randall and cific Plate restricted B. prognathus most closely to Springer, 1975) occurs. Labroides phthirophagus is B. diana, which ranges from the east coast of an Hawaiian endemic and appears to be the sister Africa eastward to the Palau and Solomon islands species of L. pectoralis and L. rubrolabiatus. along the margin of the Pacific Plate. If Lobel's Labropsis was revised by Randall (in press a), assessment of relationships is true, the Plate mar- who recognized six species. Labropsis is restricted gin separates the distributions of the two species, to the reefs of the Indo-Pacific. Two species (L. albeit there is an apparent distributional gap manabei, L. australis) are restricted to the western between the Plate margin and the most western Pacific, one (L. xanthonota) ranges from the east occurrence of B. prognathodes. coast of Africa eastward to the Marshall Islands, Choerodon, with 25 species, is limited to the Indo- one (L. alleni) ranges from the Moluccas eastward West Pacific, from the Red Sea and eastern Africa to the Marshall Islands, and two are endemic to to the western margins of the Philippine and the Pacific Plate: L. polynesica from Rarotonga Pacific plates (Gomon, 1979). (Cook Islands), Tahiti, Moorea, and Tetiaroa Diproctacanthus is monotypic and is limited to (Society Islands), Rangiroa, Takaroa, and Tika- the Indo-West Pacific, from Java east to the hau (Tuamotus), and Temoe (Gambier Group of Palau and Admiralty islands, on the western Tuamotus), and L. micronesica from Enewetak and margin of the Pacific Plate (Randall and Sprin- Kwajalein (Marshalls), Guam (Marianas), and ger, 1973, and museum collections). Palau and Ifaluk (Carolines). Randall stated that NUMBER 367 55
it Labroides pectoral Is Labroides phthirophagus Labroides rubrolabiatus
FIGURE 25.—Distributions of three of the five species of the labrid fish genus Labroides; the other two species of Labroides are widely distributed in the Indo-Pacific. the closest relative of L. polynesica is the widely from eastern Australia and New Caledonia. distributed, but allopatric L. xanthonota, and the There are three species on the Pacific Plate: one closest relative of L. micronesica is the allopatric, is endemic to the Hawaiian Islands; one ranges western Pacific L. australis, which has been taken from the Philippine Islands and New South along the marginal Plate islands from the Solo- Wales to the Pitcairn Island group; and the third mons to Fiji, Samoa, and Tonga. Thus, for one of ranges from the Philippines and Okinawa east to the Plate endemics, the Plate margin is the the Marshall Islands. boundary separating it from its proposed closest Paracheilinus, with seven species (Randall and relative. Lubbock, 1981b), is limited to the Indo-West Leptojulis comprises three species and is re- Pacific, from the Red Sea and western Indian stricted to the Indo-West Pacific. One species is Ocean to the Philipines, New Guinea, and Solo- known only from the Philippines, one only from mon Islands, on the western margin of the Pacific Thailand (Indian Ocean coast) and Ceylon, and Plate. one from the Gulf of Oman eastward to the Polylepion is a moderately deep-dwelling genus Solomon Islands, along the margin of the Pacific with two species. The genus is known only from Plate (Randall and Ferraris, 1981). the Ryukyus, Hawaiian Islands (Hawaiian excep- Macropharyngodon, with ten species, is limited totion), and the tropical eastern Pacific (Gomon, the Indo-Pacific, from the Red Sea to the Pitcairn 1979). Island group (Randall, 1978b; Shepard and Psendojuloides, an Indo-Pacific genus, was re- Meyer, 1978). Most of the species are restricted vised by Randall and Randall (1981), who rec- in distribution, with three known only from the ognized eight species: three restricted to the west- western Indian Ocean, one only from eastern ern Indian Ocean (two to Mauritius, one to Zan- Australia, one only from southern Japan, and one zibar and Aldabra), two restricted to the western 56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Pacific (one in Japan, Australia, and New Zea- the Indo-Malayan region and Palau Islands (Go- land, one in the Philippines), one widely distrib- mon, 1979). uted in the Indo-Pacific (from the east coast of Of the 17 labrid genera discussed above, 12 Africa to the Society Islands), and two endemic occur on the Pacific Plate nonmarginally; none to the Pacific Plate: one, P. pyrius, to the Marque- of the 12 is endemic. The 17 genera are repre- sas, the other, P. atavai, to several island groups in sented by about 125 Indo-Pacific species, but only southeast Oceania and Guam (J.E. Randall, in 45 (36%) of these occur nonmarginally on the litt.; hence, can be considered to be a widely Pacific Plate, and 14 (31 %) of the 45 are endemics. distributed Plate endemic). Of the 45 Plate species, 11 are found on the Plate Stethojulis was synopsized by Randall and Kay only in the Hawaiian Islands (and for a few, (1974), who recognized nine species. Shepard and Johnston Island also). The relatively few species Randall (1976) tentatively treated two of the nine of labrids found on the Pacific Plate and the species as subspecies of a single species. Three of relatively high number of these that are endemics, the species are widely distributed in the Indo- emphasize the distinction of the Plate as a bio- Pacific including the Pacific Plate (one of which geographical unit within the Indo-Pacific region. reaches the eastern Pacific as well); one is endemic to the Hawaiian Islands; one is known from LACTARIIDAE southern Japan south to Okinawa and the Bonin Islands; one is known only from the Indian This family consists of a single genus and spe- Ocean; and the species with two subspecies is cies {Lactarius lactarius) of moderate-sized (to about reported from Japan, on the one hand, and the 500 mm TL), free-swimming coastal fish distrib- Indo-Malayan region on the other. uted from the Persian Gulf eastward to Taiwan and the Admiralty Islands, and south to northern Suezichthys comprises about eight species (B.C. Australia and Fiji (Carlson, ms.). It has not been Russell, pers. comm.) and is widely distributed reported from the Pacific Plate nonmarginally. in, and limited to, the Indo-West Pacific, except for the occurrence of one species in the Hawaiian Islands (Hawaiian exception). LAMNIDAE Thalassoma, which is distributed circumglob- The mackerel sharks, including the great white ally, has been revised by Heiser (1981), who shark and makos, are large, widely distributed, recognized 21 species (including one undes- viviparous species that are usually taken offshore. cribed): two restricted to the western Atlantic, The makos are commonly taken by fisherman at two restricted to the eastern Atlantic (including Pacific Plate localities. The great white shark, one endemic to Ascension and St. Helena), three Carcharodon carcharias, is recorded from all oceans, restricted to the western Indian Ocean and/or but on the Pacific Plate has been taken only in Red Sea, three restricted to the eastern Pacific, the Hawaiian Islands, where it is uncommon, and one restricted to the western Pacific (southern from six miles (10 km) off Bikini, Marshall Is- Japan), one restricted to the Indo-West Pacific, lands, based on a single specimen (Schultz, one ranging from the western Pacific to the So- 1953b). Strasburg (1958) did not record Carchar- corro Islands in the eastern Pacific, five widely odon during longline collections that covered distributed in the Indo-Pacific, and three endemic much of the central Pacific (see his fig. 1) during to the Pacific Plate (two to the Hawaiian Islands, a four-year period that produced over 6000 spec- and one, undescribed, to Pitcairn and Ducie). imens of sharks (Carcharodon is known to be taken There are, therefore, nine species of Thalassoma on longlines—I obtained a specimen during long- present on the nonmarginal portions of the Pacific line fishing off the New Jersey coast). Carcharodon Plate. appears to be primarily antiequatorially distrib- Xiphocheilus is a monotypic genus restricted to uted in the Pacific. NUMBER 367 57
LEIOGNATHIDAE coasts of Western Australia, Queensland, North- ern Territory, and the Gulf of Papua (P. Heem- The Leiognathidae were reviewed by James stra, in litt.). In Australia, they occur mainly (1978). He recognized four genera and 17-20 along beaches and in river mouths in brackish species, the largest of which attains 242 mm TL. water (Lake, 1971), but have been taken 50 km The species are epibenthic, schooling fishes of above the mouth of the Aird River, which empties commercial importance. They are primarily ma- into the Gulf of Papua. rine but often occur in brackish or mangrove areas, and occasionally enter freshwater. Leiog- nathids have not been reported to occur outside LETHRINIDAE the Indo-Pacific except for an immigrant species The Lethrinidae comprises five genera (John- of Red Sea origin in the eastern Mediterranean son, 1980) and about 30 species of commercially (Ben-Tuvia, 1966). Leiognathids are widely dis- important, moderate-sized (to about 1 m TL), tributed and common in the Indo-West Pacific free-swimming, suprabenthic fishes. Monotaxis from the Red Sea and east coast of Africa to (monotypic) and Gnathodentex (monotypic) are Japan, the Mariana, Palau, and Bonin islands, limited to, and widely distributed in, the Indo- New Guinea, Fiji, and Samoa. I collected Gazza Pacific. Gymnocranius (at least two species) is lim- minuta and an unidentified species of Leiognathus ited to the Indo-West Pacific from the Red Sea at Ponape, Eastern Caroline Islands; otherwise, and eastern Africa to Japan, New Guinea, Solo- there have been only two reports of leiognathids mons, Samoa (Wass, ms.), and Australia (south from the Pacific Plate nonmarginally, both based to southern Queensland); and Wattsia (mono- on specimens collected over 100 years ago. In typic) is limited to the Indo-West Pacific, from these reports, four species (in three genera) were the Red Sea east to at least the South China Sea. recorded from Tahiti. Randall (1973) discussed Lethrinus has 26 species (Sato, 1978) of which 25 these reports, noted that he had not collected occur in the Indo-Pacific and one is limited to the leiognathids in all Polynesia, and questioned the tropical eastern Atlantic from Cape Verde south validity of the Tahitian records, especially as to Gabon (several other primarily Indo-Pacific three genera and three species emmanated from fish families have disjunct representation along one collection (Kner, 1865). Fowler (1928) listed the tropical west African coast, for example: Mu- two species of leiognathids from Ebon Atoll (Mar- giloididae, Platycephalidae, Psettodidae, Mono- shall Islands), which is near Kosrae (easternmost dactylidae). Of the 25 Indo-Pacific species, only of the Eastern Caroline Islands), where leiognath- five occur nonmarginally on the Pacific Plate ids might be expected to occur. In spite of consid- (four of these range as far west as the western erable, relatively recent collecting in the Marshall Indian Ocean and Red Sea, and one only as far Islands, however, there are no other records of west as the East Indies. Of the 20 species restricted leiognathids from those islands. I believe that to the Indo-West Pacific continental areas, at leiognathids on the nonmarginal portions of the least 10 reach the western margin of the Pacific Pacific Plate are restricted to the Eastern Caroline Plate. Islands chain, which has acted as a conduit per- In summary, three of the four genera of leth- mitting leiognathids to disperse onto the Pacific rinids, but only seven of the 29 Indo-Pacific spe- Plate. cies, occur nonmarginally on the Pacific Plate; no species is endemic to the Plate. LEPTOBRAMIDAE This family consists of a single genus, Lepto- LOBOTIDAE brama, and two species of small (to about 430 mm The tripletails consist of two genera, each with TL), free-swimming fishes that occur on the two species. Datnoides is a genus of small fishes(t o 58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY about 350 mm TL) that inhabits freshwater and prises 13 species) are known from the Pacific Plate may enter brackish water. The genus ranges from nonmarginally, and both are endemic to the Ha- northeast India (Ganges) to Borneo and New waiian Islands (Hawaiian exceptions) and known Guinea. only from depths of 274-576 m. Lophiodes is other- Lobotes attains a moderately large size (to about wise restricted to the eastern and western Atlantic 1 m and 14 kg) and is often, erroneously, reported and eastern and Indo-West Pacific. to be worldwide (circumtropical) in distribution. The genus is present in the eastern Pacific (an endemic species), on both sides of the tropical LUTJANIDAE Atlantic, although infrequently reported from the The snappers are a circumglobally distributed west African coast, and in the Mediterranean. In family comprising 17 genera (G.D. Johnson, the Indo-West Pacific, it is recorded from the east 1980) and about 75 species of small to moderately coast of Africa east to New Guinea and New large, free-swimming, suprabenthic fishes, many Britain, north to Japan and Korea, and south to of commercial importance. All but three lutjanid Fiji (Carlson, ms.) and southern Queensland. Lo- genera occur in the Indo-Pacific, and of the 14 botes has been reported from only three localities that do, seven are monotypic. Four of the mono- nonmarginally on the Pacific Plate: Ponape, typic genera (Pinjalo, Symphorichthys, and Sympho- where R.A. Croft, local fisheries officer, informed rus, which are shallow dwelling, and Lipocheilus, me that he had seen (collected and discarded) which occurs at depths of 94 m or more) do not only one individual in his six years of residence occur on the Pacific Plate nonmarginally. Macolor on that island; the Hawaiian Islands, where it is and Aprion, both monotypic and shallow dwelling, rare (J.E. Randall, pers. comm.) and was first are widely distributed in the Indo-Pacific, as is taken in 1957 (Gosline and Brock, 1960); and the deep-dwelling, monotypic Randallichthys, Tahiti, where it has been reported only once which is known only from the Hawaiian Islands, (Gosline and Brock, 1960). There are, however, New Caledonia, and Okinawa at depths of 152- no specimens or subsequent reports of Lobotes 293 m (Anderson et al., 1977). Parapristipomoides, from Tahiti, and its presence there remains to be monotypic, is a deep-dwelling (160-430 m) Pa- verified. Of interest here is that R.C. Wass (ms.) cific Plate endemic known only from Rapa and does not record Lobotes despite several years resi- Easter islands. Kami (1973) described Parapristi- dence and collecting fishes in Samoa, which is pomoides as a subgenus of Pristipomoides, but G.D. much closer to well documented areas of occur- Johnson (1980) recognizes both as genera. Of the rence of Lobotes than is Tahiti. Young Lobotes often seven polytypic Indo-Pacific genera (all in need occur around floating seaweed, and it is possible of revision), Lutjanus (the only circumglobally that Hawaiian specimens emanate from Japan distributed genus of lutjanids) and Aprion (both via ocean currents. I do not believe, however, that shallow dwelling), and Aphareus and Pristipomoides there is a breeding population of Lobotes present (both deep dwelling) are widely distributed in the on the Pacific Plate nonmarginally. Indo-Pacific, and Etelis (deep dwelling), Apsilus, and Paracaesio (both shallow dwelling) are re- stricted to the Indo-West Pacific. LOPHIIDAE Several species of Lutjanus have been intro- The goosefishes are primarily a benthic, deep- duced into the Hawaiian Islands, where the genus water, shelf group. They are included here only did not occur naturally (Randall and Kanayama, for the sake of completeness as a few species occur 1972). Although some of the introduced species at shallow depths in northern, temperate waters appeared to survive for several years, only two of the western hemisphere. There are four genera species, L. kasmira (introduced in 1955, 1956, and and 25 species of goosefishes (Caruso, 1981), of 1958) and Lfulvus (introduced in 1956 and 1958), which only two species (of Lophiodes, which com- established breeding populations, and L. kasmira NUMBER 367 59 has become so abundant that fisherman "believe" pending on the species. The species build burrows it is replacing more desirable native species, par- in rubbly or sandy bottom. Malacanthus is often ticularly goatfishes (Tabata, 1981; J.E. Randall, seen swimming well up in the water column, but in litt.). The two introduced species of snappers Hoplolatilus appears to remain close to its burrows. are common and widely distributed in the Indo- Larvae or prejuveniles are planktonic. Pacific, and it was presumed when they were Hoplolatilus is restricted to the Indo-Pacific, and introduced that they would compete only with the distributions of its species are poorly known labrids, holocentrids, synodontids, cirrhitids, scor- because the species are difficult to collect. Only paenids, and deep-water lutjanids, which have three species of Hoplolatilus occur nonmarginally similar feeding habits to those of the snappers. on the Pacific Plate (Dooley, 1978, Randall, All these families and Lutjanus (and the success- 1981b), and all three occur also in the western fully introduced serranid, Cephalopholis guttatus, aPacific or farther west. senior synonym of C. argus) occur together com- Malacanthus has a wider distribution (Dooley, monly at many Pacific Plate localities, and I 1978, fig.37 ) than Hoplolatilus. It is almost circum- presume that the success of the introduced snap- tropically distributed, but is known from only one pers (and Cephalopholis) is due to the lack of native locality in the eastern Atlantic, Ascension. The predators. The mugilid Chelon engeli, introduced two Indo-Pacific species of Malacanthus range from into Hawaii in 1955, also appears to be competing the east coast of Africa well out onto the Pacific with native Mugil cephalus (Randall and Kanay- Plate, and one of these, M. brevirostris, reaches the ama, 1972), although M. cephalus is still one of the eastern Pacific. most abundant Hawaiian fishes (Hawaii Division of Fish and Game, 1979). MENIDAE It may be argued that a complex ecosystem, as must exist in the Hawaiian Islands, may take This family consists of a single genus and spe- many years before reaching equilibrium when a cies (Mene maculata) of small (to about 200 mm relatively few species are added to the system. In TL) coastal fish that may enter estuaries. It is other words, the replacement (extinction) of a distributed from the east coast of Africa eastward native (or introduced) species will not become to Japan, New Britain, northern Queensland, and apparent for many more years than the 20+ years Fiji (Carlson, ms.). It is absent from the Pacific that have passed since the successful introductions Plate nonmarginally. Reports (all apparently em- were made. On the other hand, several species of anating from Matsubara, 1955; see also Formion- Lutjanus and Epinephelus (Serranidae) that were idae) of Mene\ occurring in the Hawaiian Islands introduced in quantity into the Hawaiian Islands are probably erroneous. There are no specimens simultaneously with the successful species did not from these islands, nor has it been seen there by succeed in colonization (Randall and Kanayama, J.E. Randall or W.A. Gosline (pers. comms.). 1972). Many reasons for the failures can be pos- tulated, but the important point for the present is that there is no evidence that a native species MlCRODESMIDAE has been replaced. The wormfishes comprise five genera and about 35 species (some undescribed) of small (to MALACANTHIDAE about 250 mm SL) fishes that are found in all warm seas. The species are indwellers in shallow (not including Branchiostegidae) sandy, muddy, or coral rubble bottoms, and oc- The sand tilefishes consist of two genera, Ma- casionally in estuaries. The postlarvae are plank- lacanthus, with three species (Dooley, 1978), and tonic. The two genera that occur in the Indo- Hoplolatilus, with eight (Randall, 1981b). Maxi- Pacific are restricted to that region. Gunnellichthys mum adult size is about 100-700 mm SL, de- is distributed from eastern Africa to the Society 60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Islands and Paragunnellichthys, from the Red Sea Hawaiian Islands, one (C. rapanui) is endemic to to the Eastern Caroline Islands; none of the spe- Easter Island, one (C longicaudus) is restricted to cies are Pacific Plate endemics (C.E. Dawson, southeast Oceania (Tahiti and Rarotonga), two 1974, and pers. comm.). (C. multilineatus, C. fronticinctus) are restricted to the Indo-West Pacific (Randall, 1964; Kyushin MOBULIDAE et al., 1977; reaching the western margin of the Pacific Plate), one (C. pardalis) ranges throughout The manta rays comprise two to four genera of most of the Indo-Pacific, but is absent from the large to extremely large fishes. The taxonomy of Hawaiian Islands, and one (C. dumerili) ranges the family is poorly known, probably as result of throughout the Indo-Pacific (including the Ha- the large size of the species and concomitant lack waiian Islands) to the Revillagigedo Islands in of specimens for study. Only Manta and Mobula the eastern Pacific. The distribution patterns of are generally accepted as valid genera in the the various species of Cantherhines seem to reflect family; the number of species in each is question- most of the main patterns exhibited by Indo- able. Manta is circumglobal in distribution; Mob- Pacific fish taxa, with the notable lack of Indian ula is, perhaps, circumglobal, but has been re- Ocean and widely distributed Pacific Plate en- ported only from the Hawaiian Islands nonmar- demics. ginally on the Pacific Plate, and the Hawaiian Oxymonacanthus and Paraluteres are both mono- record is considered questionable (Gosline and typic and range from the east coast of Africa to Brock, 1960). at least the Marshall Islands; however, the Red Sea population of Oxymonacanthus has been rec- MONACANTHIDAE ognized as a separate species (Marshall, 1952). Pseudomonacanthus contains five species (Fraser- There are 28 genera and about 75 species of Brunner, 1940; Randall, 1975a). Four of the spe- filefishes (Hutchins, 1977; J.C. Tyler, pers. cies are known only from the region comprising comm.). The species are of small to moderate size northern Australia, East Indies, Philippines, and (to about 1 m TL), free swimming, or associated Singapore. The fifth species, P. garretti, is known with vegetation or soft and hard corals; some only from the Hawaiian Islands and would con- species have pelagic young, often associated with stitute an Hawaiian exception (for the genus), floating Sargassum. All the genera, and 54 of the but I am informed (J.B. Hutchins in J.E. Ran- species (Hutchins, 1977), occur in Australia, the dall, in litt.) that P. garretti is a species of Tham- only relatively limited area where all the genera naconus. occur. Only nine genera are present on the Pacific Plate nonmarginally. Randall (1964) revised Amanses and Canther- MONOCENTRIDAE hines. Amanses is monotypic and ranges from the There are two genera (Monocentris, Cleidopus) Red Sea to the Tuamotus. Randall recognized 11 and three species of pinecone fishes. The species species in Cantherhines, but Hutchins (1977) syn- are small (to about 230 mm TL), usually deep onymized one of these (C. melanoides with C. par- dwelling, free swimming, and predominantly re- dalis) and referred another (C. longipinnis) to Can- stricted to the Indo-West Pacific. The family is theschenia (with two species; genus known only distributed from the east coast of Africa to Japan, from Lord Howe Island and Great Barrier Reef). the Philippines, and Australia; there are no rec- Hutchins and Randall (in press) have described ords from the Pacific Plate. Monocentris has been an additional species of Cantherhines. Of the ten reported from the Juan Fernandez Islands, off the species of Cantherhines, two (C. macroceros, C. pullus)coast of Chile, based on a single dried specimen are restricted to the western Atlantic, two (C (Fowler, 1955; Schultz, 1956), but R.H. Rosen- verecundus, C. sandwichiensis) are endemic to the blatt (pers. comm.) believes that the species, NUMBER 367 61
which is otherwise known only from San Felix been collected on the Pacific Plate nonmargin- Island (Scripps Institution collections), probably ally, and only at Rapa (Lotella sp., J.E. Randall, came from San Felix, which is also off the coast in litt.). Lotella is antiequatorially distributed: of Chile. South Australia, Lord Howe Island, New Zea- land, Rapa, Juan Fernandez Islands, and Japan MONODACTYLIDAE south to 13°40'N (D.E. Cohen, pers. comm.). The monodactylids comprise one genus {Mon- MORINGUIDAE odactylus) and two or three species of small fishes (to about 200 mm SL). Schuettea, a South Austra- The moringuid eels are indwellers of sandy lian genus, was excluded from the family by bottom, but become free swimming when sex- Tominaga (1968). The species are free swimming ually mature. They have planktonic larvae and and enter freshwater. The family is distributed attain a total length of up to 1200 mm. The from the tropical east coast of Africa to the family contains two genera, Moringua, with fewer Eastern Caroline Islands, Fiji, and Samoa (Wass, than ten species and in need of revision, and ms.); there is, in addition, a disjunct representa- Neoconger, with three species. Moringua is distrib- tion of Monodactylus on the tropical west coast of uted throughout the Indo-Pacific from the east Africa (however, absent from the coast of An- coast of Africa to Easter Island, and is also present gola), a distribution pattern similar to that of in the western Atlantic. Neoconger is represented several other primarily Indo-Pacific fish families by a different species in each of the following (see also Lethrinidae). Monodactylus has been re- areas: western Atlantic, eastern Pacific, and Aus- ported only from Ponape and Suvarow, Eastern tralia (only the larval stage is known of the Aus- Caroline Islands, on the Pacific Plate nonmargin- tralian species; Smith and Castle, 1972). ally (Herre, 1939; Fowler, 1928), and the presence of Monodactylus at these two islands is evidence for MUGILIDAE the operation of the Caroline Islands conduit. The essential absence of Monodactylus from the The mullets are a circumglobal family of epi- Plate may be a consequence of its requirement pelagic shorefishes of commercial importance. A for brackish water for the proper development of few species appear to be confined to freshwater, its eggs and larvae (Ogasawara et al., 1978; Ak- but many marine species may enter fresh and atsu et al., 1977). The high island of Ponape has brackish water. Spawning occurs offshore. Maxi- river runoff but Suvarow is an atoll and should mum size attained is about 1 m TL (Smith, lack freshwater runoff. Sterba's (1962, fig. 1022) 1953:323). The mullets are poorly known taxo- distribution map for the family fails to include nomically. Possibly ten species in seven genera areas (Fiji, Samoa, Ponape) to the east of Aus- occur nonmarginally on the Pacific Plate. For a tralia and New Guinea as part of the range of discussion of the successful introduction of Chelon Monodactylus. engeli into the Hawaiian Islands see Lutjanidae.
MORIDAE MUGILOIDIDAE The morids are a circumglobally distributed The mugiloidids are a benthic group of small group of primarily deep-dwelling, cool-water fishes (less than 500 mm TL) and are poorly fishes that are poorly known taxonomically. known taxonomically. Several species are undes- There are perhaps 15 genera and 75 species (D.M. cribed, and some described species are possibly Cohen, pers. comm.). A few species have been representative of different sexes of the same spe- collected at depths of under 100 m in the Indo- cies, although some species are hermaphroditic Pacific, but only one shallow-dwelling species has (Marshall, 1950). There are about three genera 62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
(not including the New Zealand freshwater genus currently known only from the Marquesas and Cheimarrichthys, which has been referred to the Line Islands (Randall and Gueze, 1981). Its pre- Mugiloididae on occasion). The most speciose sumed closest relative, M. vanicolensis, is widely genus, Parapercis, contains perhaps 50 species, and distributed in the Indo-Pacific, including the Line is distributed from the Red Sea and east coast of and Marquesas islands. Africa to New Zealand and the Pitcairn Island Randall (1981a) reported the first records (ob- Group (J.E. Randall, in litt.), and also is present tained in 1976 and 1977) of Upeneus vittatus from off the southwest and southeast coasts of South the Hawaiian Islands. He believes that the occur- America, and along the central west coast of rence of this species in Hawaii is the result of an Africa. introduction in 1955 from the Marquesas Islands; In examining the literature on Indo-Pacific however, the possibility that the species is a strag- Parapercis species (see Gomon, 1981, for compila- gler from Japan, where the species is common, tion), I find that only seven of the 36 currently should be considered. recognized species occur nonmarginally on the Pacific Plate. Five of these seven are widely dis- tributed in the Indo-Pacific, one (P. multifasciata) MURAENESOCIDAE is known only from Japan and Hawaii (Hawaiian The muraenesocids comprise eight genera and exception), and one (P. hexophthalma) is widely 18 species of eels (Castle and Williamson, 1975) distributed in the Indo-West Pacific with a single with pelagic larvae. The largest muraenesocids report (Fowler, 1928) from the Pacific Plate, Ta- attain lengths as great as 2.1 m, and are of hiti. The Tahitian record is based on a specimen commercial importance, at least in Japan. The obtained by Andrew Garrett and deposited at the group is primarily marine, but some species may Museum of Comparative Zoology, Harvard. I enter brackish water. The muraenesocids are dis- consider the Tahitian record to be highly suspect, tributed cirrcumglobally in the warm shelf and especially in view of the numerous suspect local- shore waters of the continental plates, but are ities assigned to many specimens obtained by absent from the Pacific Plate nonmarginally Garrett for the Museum Godeffroy. (Castle and Williamson, 1975, provide distribu- tion maps for the seven species included in Mu- MULLIDAE raencsox, Congeresox, and Cynoponticus). The goat fishes are a circumglobal group of small (to about 520 mm TL), commercially im- MURAENIDAE portant fishes with planktonic larvae. The family The moray eels include about ten genera and consists of six genera and, perhaps, as many as 50 over 100 species of short to very long (to 3.8 m species. Mullus occurs only in the Atlantic and TL) fishes with planktonic larvae. The family is Mediterranean; Pseudupeneus occurs only in the distributed circumglobally (nine genera nonmar- eastern Pacific and Atlantic; Upeneichthys occurs ginally on the Pacific Plate), but outside the only in Australia and New Zealand; Paraupeneus western Atlantic is very poorly known taxonomi- occurs only in the Indo-Pacific; and Upeneus and cally. The presence of Muraena on the Pacific Mulloides (replaces Mulloidichthys, P.C. Heemstra, Plate is based on the widely distributed Muraena pers. comm.) occur only in the Indo-Pacific and pardalis, which possibly merits ? different (new) western Atlantic. The last three genera are pres- genus (J.E. Randall, in litt.). ent nonmarginally on the Pacific Plate. Although Thomas (1969) reviewed all the Indo-Pacific gen- era, except Upeneichthys, a modern revision is still MYLIOBATIDIDAE needed. The eagle rays comprise four genera and fewer Mulloides mimicus is a Pacific Plate endemic than 20 species of viviparous, free-swimming NUMBER 367 63 fishes. They are poorly known systematically Caroline Islands (Ponape; Herre, 1939; I have probably because of their large size (maximum also collected Scolopsis from Ponape), Marshall disc width about 2.5 m) and concomitant lack of Islands (Fowler, 1928; Hiatt and Strasburg, specimens in collections. Two of the four genera, 1960), and Gilbert Islands (Fowler, 1928). Scolopsis and most of the species, are unreported from the has also been reported from the Hawaiian Islands Pacific Plate nonmarginally, a common circum- (Cuvier and Valenciennes, 1830), but the lack of stance for many elasmobranchs. subsequent records (J.E. Randall, in litt.) for Aetobatus is circumglobally distributed. Hawaii raises doubts as to its presence there. Aetomylaeus is distributed from the Red Sea to (Gnathodentex aureolineatus, Lethrinidae, has often Japan, the Philippines, and Australia, but is ab- been reported from Pacific Plate localities as Pen- sent from the Pacific Plate. tapodus aureolineatus.) Myliobatis, with about 11 species, is the most speciose genus in the family. It is almost circum- globally distributed, but does not occur on the NEOSTETHIDAE nonmarginal portions of the Pacific Plate. The neostethids comprise six to eight genera Pteromylaeus is reported from the eastern Atlan- and about 15 species of tiny (to about 25 mm tic and Indian oceans, and the Pacific Ocean SL), oviparous, free-swimming fishes that are re- from the east coast of Australia north to the stricted primarily to the fresh and brackish waters Admiralty Islands and east to the Gilbert Islands, of the Philippines, Borneo, Malay Peninsula, and on the Pacific Plate. northwest Sumatra. They occasionally occur in fully marine waters, probably adventitiously, and NEMIPTERIDAE for that reason only are included here. The family is sometimes considered to be a subfamily within the Phallostethidae, which includes primarily The Nemipteridae comprises three genera freshwater species that are occasionally taken in {Nemipterus, Pentapodus, Scolopsis; Johnson, 1980) estuarine waters. of small (maximum TL, 764 mm; most species under 400 mm), free-swimming, reef-inhabiting fishes that are confined to the Indo-Pacific. The NOMEIDAE three genera are in need of revision, but probably include fewer than 50 species. Rao and Rao's The nomeids comprise three genera (Cubiceps, (1981) putative revision of Scolopsis is vastly in- Nomeus, Psenes) and about 14 species of oceanic complete. fishes, which are occasionally taken close to shore. Nemipterus ranges over most of the tropical The species are of small to large size (largest, over Indo-West Pacific, but is absent other than mar- 1 m TL), have planktonic larvae (Ahlstrom et al., ginally from the Pacific Plate, except for a speci- 1976), and many of the species are widely distrib- men purportedly obtained by the Albatross at uted. Probably five species are present nonmar- Erben Guyot at a depth of about 350 m. Erben ginally on the Pacific Plate. Guyot is about 1100 km west of Los Angeles, Nomeus, the man-of-war fish, is monotypic, oc- California. Barry C. Russell (pers. comra.), who curs in all warm oceans except the eastern Atlan- is revising Nemipterus, considers the Erben Guyot tic and Mediterranean, and its young stages are record suspect. Pentapodus and Scolopsis are distrib- often associated with the pelagic coelenterate uted similarly to Nemipterus, but there are a few Physalia (Haedrich, 1967). The distribution of reports of Pentapodus caninus from the Gilbert and Nomeus (and Physalia, the world distribution of Marshall islands (see Fowler, 1928, for citations), which I have been unable to ascertain) in the P. microdon from the Gilbert Islands (Weber and central Pacific is worthy of further study. Physalia de Beaufort, 1936), and Scolopsis from the Eastern occurs irregularly at Guam, but Nomeus has not 64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY been taken there (R.F. Myers, pers. comm.), or at frequently present in shallow depths along shore, Fiji and Samoa. Nomeus on the central and eastern as well as at moderate depths offshore, but Odon- Pacific Plate is known only from three areas: the taspis seems to frequent deep water. Eugomphodus Hawaiian Islands, where it (and Physalia; Wood- is widely distributed on the tropical and warm- cock, 1956) is common; a specimen from 06°N, temperate coasts of the Western Hemisphere, 162°W (near northern Line Islands; reported by eastern Atlantic, western Indian Ocean, Aus- Fowler, 1928); and some specimens from off the tralia, and eastern Pacific, but has not been re- Marquesas, 10°S, 130°W and more easterly ported from the Pacific Plate. Odontaspis is known (Scripps Institution of Oceanography collections). from the Mediterranean, eastern Atlantic, eastern If the distribution of Nomeus on the nonmarginal Pacific, Indian Ocean, Japan, southern Australia, Pacific Plate is reasonably well reflected by the New Zealand, and the Hawaiian Islands, where information given here, the distribution is similar two specimens were taken from gill nets set at to that of some endemic Pacific Plate species of depths of 185-310 m (Clarke, 1972). According fishes: Hemiramphus depauperatus (Hemiramphi- to Tinker (1978), 0. kamoharai occurs in the Ha- dae) and Ostracion whitleyi (Ostraciidae) and, per- waiian Islands, but L.V.J. Compagno informs me haps, Eviota epiphanes (Gobiidae) and Scorpaena that this species belongs in Pseudocarcharias (Pseu- coniorta (Scorpaenidae), which two species are, docarchariidae, an epipelagic group not included however, unreported from the Marquesas. It is in my study). One may question whether the possible that the presence of Nomeus on the Pacific occurrence of Odontaspis in the Hawaiian Islands Plate is adventitious and the result of two differ- is a case of an Hawaiian exception, representative ent ocean current regimes: the Kuroshio exten- of stragglers (from Japan), or an indication that sion flowing from Japan (where Nomeus occurs) Odontaspis is distributed more widely on the Pa- toward the Hawaiian Islands and the westward cific Plate, but is less available because it is flowing North and South Equatorial currents relatively deep dwelling. With regard to the last from the warm latitudes of the west coast of the mentioned possibility, it is noteworthy that Stras- Western Hemisphere (where Nomeus also occurs). burg (1958) failed to record Odontaspis among the Butler (1979) gives detailed distribution maps 6000+ sharks taken during several years of long- for the seven species of Cubiceps. line operations over much of the central Pacific.
NOTOGRAPTIDAE OGCOCEPHALIDAE The notograptids consist of single genus, Noto- The batfishes comprise nine genera and about graptus, and two or three species (Tyler and Smith, 60 species (Bradbury, 1967) of small (to about 1970; Whitley, 1964:54) of small (to about 130 300 mm TL), benthic fishes. The family occurs in mm TL), mangrove or reef inhabiting eel-like the warmer latitudes of all three major oceans, fishes that are restricted to the Northern Territory but is absent from the Pacific Plate nonmargin- and Queensland coasts of Australia and the Fly ally, except for the occurrence of five species in River area of southern Papau-New Guinea. the deep waters of the Hawaiian Islands (three of the Hawaiian species also occur in Japan). Bat- fishes occur in both deep and shallow depths in ODONTASPIDIDAE the tropical western Atlantic and in some portions of the eastern Pacific, but are rarely reported The sand-tiger sharks consist of two genera, from less than 100 m depth in the Indo-Pacific. each with one or two species (L.V.J. Compagno, pers. comm.). The family is in need of revision. These sharks are viviparous, of medium to large OPHICHTHIDAE size (reported to attain 6 m TL, but probably The snake eels comprise about 50 genera and rarely, if ever, exceeding 3.5 m). Eugomphodus is 200 species of small to moderately long (less than NUMBER 367 65
200 mm to over 2 m TL) fishes that are predom- pines (McCosker, in litt.). Schismorhynchus labialis inantly shallow-water, benthic indwellers (but is sympatric with both species of Schultzidia in there are two monotypic midwater genera, one Palau and with Schultzidia johnstonensis at Johnston undescribed) with pelagic larvae (McCosker, and the Samoan islands. Dispersal has, thus, ob- 1977, and pers. comm.). scured the vicariant event that initiated the di- The family is circumglobally distributed, with vergence of Schultzidia and Schizmorhynchus from the largest number of genera (25) occurring in their common ancestor. the Indo-Pacific. Seventeen of the genera are McCosker (1979) reported on the ophichthids reported from the Pacific Plate, and at least two of the Hawaiian Islands and found five species to of these (Cirricaecula, Schismorhynchus) are restricted be endemic; however, four were rare, deepwater to the Plate or its margins (Figures 26 and 60). species or belonged to genera in need of revision. An additional genus, Evips, is known only from McCosker noted that there are recognizably dis- Palau, on the eastern margin of the Philippine tinct populations of several other ophichthid spe- Plate (Figure 60) and conceivably may occur cies in the Hawaiian Islands, but declined to otherwise only on the Pacific Plate. Cirricaecula accord names to these populations. and Evips are known from only a few specimens, Castle (1972) discussed the distribution of the but the monotypic Schismorhynchus is a widelymonotypic, deep-dwelling, pelagic ophichthid ge- distributed Plate endemic. McCosker (1977) con- nus Benthenchelys, and provided a distribution sidered Schultzidia, with two species, to be the map for its three allopatric subspecies. One sub- closest related genus to Schismorhynchus. Schultzidia species ranges from the northeast Indian Ocean johnstonensis is known from the Palau, Johnston, to New Caledonia, one from the Sulu Sea to Hawaiian, Samoa (Wass, ms.), and Christmas Palau (margin of the Pacific Plate), and one is (Pacific) islands, and Schulzidia retropinnis is knownrestricted to the area of the Phoenix Islands (Pa- from Kapingamarangi, Palau, and .the Philip- cific Plate). Although ophichthids appear to have
O SCORPAENOPSIS FOWLERI - SCORPAENIDAE • SCHISMORHYNCHUS LABIALIS - OPHICHTHIDAE * CANTHIGASTER EPILAMPRA - TETRAODONTIDAE
FIGURE 26.—Distributions of three widely distributed Pacific Plate endemic fishes, including a monotypic genus, Schismorhynchus (for the other Pacific Plate endemic genera see Figure 60). 66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY long pelagic stages (10-12 months; Castle, 1965), and, as far as known, mouth brooders. They occur the formation of narrowly restricted endemic pop- at depths as shallow as a few centimeters to as ulations is not precluded, and the western margin great as 375-384 m. Aquarium-reared larvae of of the Pacific Plate holds promise as defining the one species were free swimming for 18 days and vicariant event that permitted divergence of con- then entered burrows (Colin, 1972). Dispersal by specific populations on either side of the margin. currents is, therefore, feasible. Jawfishes are found The fact that pelagic ophichthid larvae do not only in the waters of the Western Hemisphere necessarily disperse great distances is further dem- and the Indo-West Pacific; they are absent from onstrated by the benthic genus Neenchelys. Castle the Pacific Plate except marginally (Figure 27). (1981) plotted the distributions of the larvae and adults of the two species of Neenchelys, which genus OPLEGNATHIDAE is restricted to the Indo-West Pacific. The larvae of both species of Neenchelys reach the western This family, which is included here only for the margin of the Pacific Plate, but do not occur on sake of completeness, consists of one genus (Opleg- the Plate. Most of the larvae were collected by nathus) and about six species of moderate-sized (to the Dana expeditions, which also made collections about 1 m TL), free-swimming fishes with plank- in Polynesia on the nonmarginal portions of the tonic larvae. The family is antitropically distrib- Pacific Plate (Greve, 1934, pi. 1). uted, and occurs only in the Pacific Ocean: Vla- divostok south to southern Japan, Hawaiian Is- lands (Hawaiian exception), southeastern coast of OPHIDIIDAE Africa, southern coast of Australia, Galapagos Islands, Peru, and Chile. (including Brotulidae) Oplegnathus (0. fasciatus, a Japanese species) has The cuskeels are a speciose group of small to been reported in the literature only once from the large fishes that are circumglobally distributed, Hawaiian Islands (Steindachner, 1893), and this and occur at depths of less than 1 m to the report has been considered of questionable valid- greatest depths where a fish has been caught. ity (Gosline and Brock, 1969; Tinker, 1978). Only two shallow-dwelling species have been re- There are valid recent records of Oplegnathus from ported nonmarginally on the Pacific Plate: Brotula the Hawaiian Islands, however. One record, multibarbata, which is widely distributed in the based on a photograph of a specimen taken at the Indo-Pacific, and B. townsendi, which is reported island of Hawaii in 1978 (specimen discarded), only from the Marshall, Hawaiian, and Johnston was shown me by a resident. A recent mimeo- islands, and may be endemic to the Pacific Plate graphed project report (Hawaii Division of Fish (and is undoubtedly more widely distributed than and Game, 1979) records captures of Oplegnathus is indicated, as there are many unidentified col- punctatus (also a Japanese species) from Midway lections of Brotula from Plate localities). Ophidian, and Kure, northernmost Hawaiian Islands. The another genus of shallow-dwelling cuskeels, oc- presence of Oplegnathus in the Hawaiian Islands is curs at Easter Island, and represents an eastern probably the result of dispersal from Japan via Pacific component in the Easter Island fish fauna. the Kuroshio extension. It would be interesting to determine if breeding populations of Oplegna- OPISTOGNATHIDAE thus have become established in the Hawaiian Islands. The jawfishes comprise three or four genera and about 60 species of small (25-600 mm SL) fishes (W.F. Smith-Vaniz, pers. comm.). The spe- ORECTOLOBIDAE cies are all marine, with a few occasionally occur- The order Orectolobiformes comprises seven ring in estuaries. They are all burrow dwellers families and 12 genera of sharks (Compagno, NUMBER 367 67
FIGURE 27.—Distribution of the jawfishes, family Opistognathidae (compiled by W.F. Smith- Vaniz); while this study was in press I collected Opistognathus in the Fiji Islands.
1973a, and pers. comm.), all of generally sluggish forms known only from the western Pacific. behavior, and most, benthic in habit. The num- The Ginglymostomatidae, which are shallow ber of species included is not large (probably less dwelling, include Ginglymostoma, a viviparous ge- than 30), but has not been established for most of nus (with two or three species), which is distrib- the families. The Rhiniodontidae, or whale uted on the warm coasts of the Atlantic, eastern sharks, comprises a single monotypic genus, Rhi- Pacific, and western Indian oceans, and Nebrius niodon, which is an egg layer, the largest of all (two species, but only one on Pacific Plate non- fishes (to 18 m TL or more), pelagic in habit, and marginally), which is oviparous and widely dis- circumglobally distributed. The other six families tributed in the IndoPacific, at least as far east as in the order have usually been combined in the Samoa and the Marshall Islands. Nebrius ferrugi- literature and treated as one family, Orectolobi- neus has frequently been assigned to Ginglymostoma, dae. I recognize all six families, but treat them accounting for the supposed occurrence of that together here for the sake of convenience. genus in Oceania (LJ.V. Compagno, pers. The Orectolobidae (sensu stricto: Eucrossorhinus,comm.). Orectolobus) are viviparous and restricted to the The Stegostomatidae {Stegostoma) are ovipa- western Pacific; they do not occur nonmarginally rous, shallow dwelling, and widely distributed in on the Pacific Plate. Bigelow and Schroeder the Indo-West Pacific, but absent from the Pacific (1948) list the eastern Pacific within the range of Plate nonmarginally. Orectolobus, but neither I nor LJ.V. Compagno The Hemiscylliidae (Hemiscyllium, Chiloscyllium) (pers. comm.) know of an eastern Pacific record are shallow-dwelling, oviparous sharks that are for the genus. restricted to the Indo-West Pacific, from the east- The Parascylliidae (Parascyllium, Cirrhoscyllium) ern Indian Ocean to the Philippines, Japan, New are moderately deep-dwelling, outer shelf or slope Guinea and northern Australia. 68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
The Brachaeluridae (Brachaelurus, Heteroscyl- (to about 100 mm TL), benthic inhabitants of lium) are viviparous, shallow dwelling, and re- shallow to moderately deep water (to about 150 stricted to Australia. m), but juveniles of 10 mm have been taken at Thus, all of the 12 genera of the Orectolobi- the surface in the Hawaiian Islands (USNM col- formes occur in (and nine are restricted to) the lections). The family is restricted to the Indo- Indo-Pacific area, but only two genera (and two Pacific (Figure 28), but is represented on the species) occur nonmarginally on the Pacific Plate, Pacific Plate only marginally and by an endemic, and one of these genera (Rhiniodon, monotypic, moderately deep-dwelling (to about 150 m as pelagic) is circumglobally distributed. The pat- adults) Hawaiian species, Eurypegasus papilio (Ha- tern exhibited by the Orectolobiformes is the waiian exception). There is only one other species same shown by many elamosbranch groups, of Eurypegasus, E. draconis, which ranges from the which tend to be restricted to continental plates. east African coast to southern Japan and Aus- tralia; hence, the distributions of these two sister species are separated by the margin of the Pacific OSTRACIIDAE Plate. (not including Aracanidae) A specimen of pegasid in the marine vertbrate collection at Scripps Institution of Oceanography The ostraciid trunkfishes comprise up to six was taken from the stomach of a tuna reportedly genera and, perhaps, as many as 25 species of caught in the vicinity of Tahiti. I have not entered small (maximum size about 460 mm TL), shal- this record on the distribution map of the family, low-dwelling, benthically associated fishes. The nor two others, one in the Scripps collection, family is distributed circumglobally in warm wa- supposedly from California, that was obtained by ters. The Indo-Pacific species are particularly in a diver, and one in the Charleston (South Caro- need of revision, but one species, Ostracion whitleyi, lina) Museum, purportedly taken off Charleston appears to be a widely distributed Pacific Plate and provided by a commercial seafood dealer. endemic: Marquesas, Tuamotus, Society, Ha- These last two specimens are identifiable as Indo- waiian, and Johnston islands (Randall, 1972b). West Pacific species (T. Pietsch, in litt.). Randall (1975c) considered Ostracion tracys, which is known only from Mauritius, to be the closest relative of 0. whitleyi. If the distributions and PEMPHERIDIDAE relationships are as Randall has reported, the The sweepers comprise three genera and prob- distributions are relict, and extinction must have ably fewer than 25 species of small (to about 200 occurred over the great distance separating the mm TL), free-swimming fishes that usually occur nearest populations of the two species. about rocks and reefs, and most often in caves. The genera are greatly in need of revision. Liopem- PATAECIDAE pheris is known only from Australia; Parapriacan- thus is broadly distributed from the Red Sea and The prowfishes comprise three genera and east coast of Africa eastward to at least the Mar- about six species of small (to 230 mm TL), rare shall and Phoenix islands; Pempheris is distributed fishes restricted to the more southern coasts of similarly to Parapriacanthus, but is also disjunctly Australia, but reaching as far north as southern present along the warmer coasts of the western Queensland. They are questionably included here Atlantic. There are no sweepers in the eastern as warm-water inhabitants. Pacific, eastern Atlantic, or Mediterranean.
PEGASIDAE PENTACEROTIDAE There are four genera and five species of sea The boarfishes were revised by J.L.B. Smith moths (T. Pietsch, in litt.). The species are small (1964), who recognized five genera and 12 species. NUMBER 367 69
FIGURE 28.—Distribution of the fishes of the family Pegasidae (star indicates unspecified locality on the coast of South Australia).
The boarfishes tend to be deep-dwelling inhabit- reason, and because many Museum Godeffroy ants of temperate coastal waters, but are taken specimens seem to be accompanied by question- occasionally in tropical areas and at depths ac- able locality data, I am doubtful that the locality cessible to SCUBA divers. The young (and rarely accorded Giinther's specimen is correct. (2) P. the adults) of some species have been taken at the capensis from southern and southeastern Africa. surface. Most species are known from specimens (3) P. japonicus from Japan, southern Australia, under 500 mm TL, but one species attains 900 and the following localities, which have been mm. The family is included here for the sake of reported since Smith's revision: Bonin (Ogasa- completeness and because the reported distribu- wara) Islands (Zama et al., 1977) and Taiwan tion of one genus (Histiopterus) provided a test of (Burgess and Axelrod, 1974). (4) P. richardsoni, my observations on Pacific Plate biogeography. Tristan da Cunha, southernmost Africa, Japan, Three of the genera of boarfishes (Zanclistius, New Zealand, Leeward Hawaiian Islands and Paristiopterus, Pentaceropsis), including five species, seamounts (Randall, 1981a), Alaska to Califor- are found only in the waters of southern Australia. nia, and southernmost South America (hence, Pentaceros includes four deep-dwelling species. antiequatorially distributed). Penrith (1967) be- (1) P. decacanthus, known only from two juveniles lieves that P. richardsoni is distributed throughout collected over 100 years ago, one without locality the world in warm-temperate waters and that the data, the other supposedly from the southeast known distribution only reflects fishing intensity. Atlantic at 10°25'S, 30°W (a Museum Godeffroy The distribution of Histiopterus, with three spe- specimen; Giinther, 1873). The western Atlantic cies was mapped by J.L.B. Smith (1964, fig. 1), has been well and continuously sampled ich- and is reproduced here as Figure 29A, but with thyologically. It is, therefore, surprising that no the addition of three recent locality records: off other pentacerotid specimens have been taken in the southwest coast of India (Kotthaus, 1975), the western Atlantic for over 100 years. For this Lord Howe Island (Allen et al., 1976), and the 70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 29.—Distributions of the species of the pentacerotid fish genus Histiopterus: A, uncor- rectcd, after J.L.B. Smith (1964, fig. 1) with additional records entered for southwest India, Lord Howe Island, and the Red Sea; B, corrected (see page 69).
Red Sea (Klausewitz, 1980). My examination of Duke of York specimen had been examined for Smith's distribution map of Histiopterus, even be- him at the British Museum by G. Palmer, and as fore the addition of the recent records, aroused there is a Duke of York Island at about the my suspicions that one of Smith's records might coordinates Smith gave (in the Tokelau Islands), be in error: "Duke of York Is. (9°S X 172°W)." I was inclined to drop my suspicions and ascribe Except for the Duke of York record, the map the lack of reports of Histiopterus from Oceania to appeared to me to be a normal Hawaiian excep- collecting deficiencies. Smith, however, men- tion type distribution. Smith mentioned that the tioned that the Duke of York specimen had been NUMBER 367 71 reported originally by Giinther (1877), and al- 1975:1429). Burgess and Axelrod did not report though Smith did not cite the reference more Pholidichthys from New Caledonia, but in their fully, I decided to find it and determine if photograph of an aggregation of Plotosus anguil- Giinther might have been in error. Giinther, how- laris, they call attention to two nonplotosids in ever, had not made an error, but Smith had! The the aggregation. These nonplotosids are juvenile specimen did come from the Duke of York Is- Pholidichthys (brought to my attention by J.R. land—not the one in the Tokelaus, but the one Gomon). between New Britain and New Ireland, on the western margin of the Pacific Plate (see corrected PLATYCEPHALIDAE distribution map, Figure 29B), which is consistent with an Hawaiian exception distribution pattern. The flat heads comprise about 12 genera and The three recently reported records of Histiopterus about 60 species of small to large (0.23-1.20 m are congruent with this pattern. This is a small maximum TL), mostly shallow-dwelling, benthic but didactic example of the usefulness of a Pacific fishes that occur on sandy, muddy, and rubbly Plate biogeographic hypothesis. Had I not been bottoms in habitats as diverse as coral reefs and aware of the Hawaiian exception type of distri- estuaries. The family is limited to the Indo-Pa- bution, I would have had no reason to question cific, from the Red Sea and east coast of Africa Smith's distribution map. Similar incidents in- eastward to the Pitcairn Island group, except for volving other fish groups occurred often during a disjunct, endemic species, Grammoplites gruveli, the course of my study. on the tropical west coast of Africa (for some In summary, the Pentacerotidae are primarily other Indo-Pacific taxa with disjunct west African antiequatorially distributed, but are absent in the representation see Lethrinidae). Grammoplites is Mediterranean and Atlantic, except for occur- otherwise restricted to the Indian Ocean and a rences at Tristan da Gunha and around the south- recent immigration into the eastern Mediterra- ern tips of Africa and South America (recorded nean via the Suez Canal. from Cape Horn), and Giinther's (1873) question- Only three genera and five species of flatheads able record of a juvenile from off Brazil. The occur nonmarginally on the Pacific Plate, and pentacerotids are also unreported in the eastern none of these taxa is endemic to the Plate. Nu- Pacific south of California (except for Cape Horn) merous species are widely distributed in the Indo- or from the Pacific Plate, except marginally and West Pacific and find their easternmost limits at the Hawaiian Islands. along the eastern margin of the Philippine Plate or western margin of the Pacific Plate; other species are restricted to the Indian Ocean. No PHOLIDICHTHYIDAE flatheads have been reported from the Hawaiian The western Pacific family Pholidichthyidae Islands (the Bembridae is here recognized as a includes a single genus and species, Pholidichthys separate, deep-dwelling family). (Most of the in- leucotaenia, which is a small (to 337 mm TL) formation on this family was supplied by L.W. inhabitant of shallow rocky and coral-reef areas, Knapp.) even when young (at least as small as 15 mm). Springer and Freihofer (1976, fig. 1) give a distri- PLESIOPIDAE bution map for the species, which was known to them only from the Solomon Islands, southeast The Plesiopidae consists of small fishes (under New Guinea, Ambon, northeast Borneo, and Pa- 300 mm TL) that inhabit rocky and coral-reef lawan (Philippine Islands). To these localities areas of the tropical and warm-temperate Indo- may be added Negros Oriental, Philippine Is- Pacific. The family includes six genera and about lands, based on a collection I made in 1978, and 20 species. I do not believe the family is a mono- New Caledonia (Burgess and Axelrod, phyletic group. 72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Fraudclla (monotypic), Paraplesiops (about fiveI will remain skeptical until all specimens in Reg. species), and Trachinops (four species; Allen, 1977) #67 are accounted for and shown to provide for are restricted to Australia, and all but Fraudella inclusion of the Calloplesiops specimen, or Callople- are warm-temperate water inhabitants. siops' presence in the Tuamotus, or some neigh- Assessor (three species), a mouth brooder, is boring island group (such as Societies, Line Is- known only from the Great Barrier Reef, the lands, or Marquesas), is corroborated. Ryukyus, northern Philippines, and a visual re- port from New Caledonia (Allen and Kuiter, 1976; Randall, in press b); its distribution is PLEURONECTIDAE antiequatorial. The flounders are a moderately large family of Plesiops (about six species; last revised by Inger, taxonomically poorly known, small to large, 1955) ranges from the Red Sea and east coast of benthic fishes that are primarily inhabitants of Africa to the Marshall, Gilbert, and Phoenix deep or temperate waters. Only two genera are islands. Only one or two species occur on the reported nonmarginally from the Pacific Plate. Pacific Plate nonmarginally, and these are not Samariscus is a common, widely distributed Indo- endemics. Pacific genus of small fishes, and Poecillopsetta is McCosker (1978) gave the distribution of Cal- a deep-dwelling genus of small fishes found in the loplesiops (monotypic) as "Tuamotus, Palau, Phil- western Atlantic and Indo-Pacific, but known ippines, Indonesia, Maldives, Grande Comore, nonmarginally on the Pacific Plate only at the Seychelles, Zanzibar, Mozambique, Pemba, sev- Hawaiian Islands. eral Red Sea localities, Ryukyus Islands, and
Southern Japan." To these may be added Guam PLOTOSIDAE (Kami, 1975), One Tree Island (Great Barrier Reef, in Australian Museum collections), and The plotosid catfishes comprise about ten gen- Hermit Islands (Bismarck Archipelago), Fiji, and era and 30 species of small to moderately large Ponape (all USNM collections). The only non- (to 1.3 m TL) fishes that are restricted to the marginal Pacific Plate records for Calloplesiops are Indo-Pacific, probably to the Indo-West Pacific Ponape and Tuamotus, an unusual distribution (Figure 30). More than half the species are essen- for an area that has been so well collected in tially confined to freshwater, although some of recent years. Without the Tuamotus record, I these enter brackish water. The other species are would expect that Calloplesiops was an Indo-West found in brackish and marine waters, some oc- Pacific taxon that dispersed onto the Pacific Plate curring normally on coral reefs. Unlike many via the Caroline Islands conduit. I, therefore, catfishes, the plotosids are not mouth brooders. question the Tuamotus record, which is suppos- As far as known, they lay demersal eggs in nests edly based on the relatively modern collections (freshwater) or rock crevices (marine). Young ma- made at Raroia by the George Vanderbilt Foun- rine plotosids are not planktonic, but occur in dation's 1952 expedition. The field data sheet dense spherical aggregations in shallow water, (GVF Reg. #67) for the collection that pertains sometimes almost at the shoreline. Adults are to the Calloplesiops specimen (California Academy usually suprabenthic, but may occur at shallow of Sciences 40448) mentions that 21 species and depths around pilings. Few, if any, of the marine 39 specimens were taken in the collection. The forms occur at depths greater than about 20 m. list of species accompanying the field data sheet The Plotosidae have not been revised, but J.R. refers to only 18 species and 36 specimens and Gomon and W.R. Taylor are studying the marine does not include mention of a plesiopid. While it genera and G. Nelson and N. Feinberg, the fresh- is possible that the additional three specimens water genera. All four of these investigators con- listed on the field data sheet include Calloplesiops, sider the three nonmarginal Pacific Plate records NUMBER 367 73
FIGURE 30.—Distribution of the fishes of the family Plotosidae (compiled by N. Feinberg, J. Gomon, G. Nelson, and W.R. Taylor; see discussion under "Plotosidae" for questionable locality records). of plotosids (Plotosus lineatus, Figure 30) as suspect:occurrence at that island, despite his six years Kingsmill Group (Gilbert Islands), Society Is- residence there and active reef-fish collecting. lands, and Jaluit (Marshall Islands). The first two localities are based on old Museum Godeffroy POLYNEMIDAE specimens, which have been the source of many puzzling distribution records (Randall, 1955b, The threadfins comprise seven genera (Myers, 1973, did not record plotosids from the Gilbert or 1936) and about 35 species of small to moderately Society Islands). The Jaluit record is based on a large (to 2 m TL), commercially important, su- relatively recently collected specimen in the Paris prabenthic fishes that are almost circumglobally Museum (the collector and details of the collec- distributed on warm, marine shores. Some species tion are unknown to the current curator, M.L. enter freshwater. Bauchot, in litt.). Because Plotosus lineatus usually Pentanemus (monotypic) is restricted to the trop- occurs in large numbers, is easy to collect, and ical west African coast, with a single, questionable there have been no other reports from the heavily record from Cuba (R.M. Feltes, who is revising collected Marshall, Society, and Gilbert islands, the family, pers. comm.). Galeoides is represented I believe the Pacific Plate locality records are by a common species on the west African coast. either in error or the result of straggling. Further- A second species of Galeoides was described by more, I did not see or collect plotosids at Ponape Steindachner (1869) from a single specimen from (where I would expect them to occur if anywhere "China," and subsequent specimens have not on the Pacific Plate), and the local fishery officer, been reported. There is also no indication that R.A. Croft, at Ponape had no knowledge of their the holotype has been re-examined since its de- 74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY scription, and I suspect that the generic allocation restricted region (C. hotumatua), and two (C.Jisheri, or locality of the specimen is erroneous. Of the C. potteri) occur only in the Hawaiian Islands. remaining five genera, Filimanus (monotypic), One of the endemics, C. loriculus, is represented Eleutheronema (two species), Polynemus (seven spe- by a slightly differentiated population in the cies), and Polistonemus (monotypic) occur only in Marquesas (Randall, 1980a). Another species, C. the Indo-West Pacific, variably from India or the shepardi (Figure 31), appears to be endemic to the Malay Archipelago east to the western margins eastern margin of the Philippine Plate, and an- of the Philippine and/or Pacific plates. Polydacty- other, C. interruptus (Figure 31), which was be- lus, the largest genus of polynemids, with about lieved to be endemic to southern Japan, was 20 species, occurs in all oceans, and in the Indo- recently collected at Reunion (Randall, in press Pacific ranges from eastern Africa to the Pitcairn b), in the Indian Ocean, and at Kure, northern- Island group. The species of Polydactylus art in most of the Hawaiian Islands (Ralston, 1981), need of revision, but of the 10-15 Indo-Pacific possibly carried there by the Kuroshio extension. species, probably no more than two or three occur It is doubtful that breeding populations of C. nonmarginally on the Pacific Plate. interruptus are present at Kure. The large gap in It is clear from this information that the west- the distribution of C. interruptus is probably a ern margin of the Pacific Plate delineates a strong collecting artifact, but seems unusual in view of barrier to the eastward dispersal of polynemids. the large amount of collecting in the area of the gap in recent years (see also Pomachromis richard- POMACANTHIDAE soni, Pomacentridae). There are seven to nine genera and about 78 The remaining six genera of pomacanthids species of marine angelfishes (Allen, 1980; W.E. occur only in the Indo-Pacific. Arusetta, mono- Burgess, pers. comm.). The species are moderately typic, does not occur on the Pacific Plate non- small (maximum size varying from 75 to 600 mm marginally. TL) and free swimming, but are obligatory reef Euxiphipops, with three species, is restricted to inhabitants. The family is distributed circum- the Indo-West Pacific, except that one species, E. globally in warm, shallow seas. xanthometopon, is present on the Pacific Plate non- Holacanthus, with eight species (one undes- marginally at Ponape (personal observation) and cribed, W.E. Burgess, pers. comm.) occurs only in Kosrae (Eldredge et al., 1979), which I consider the eastern Pacific and western Atlantic. evidence for operation of the Caroline Islands Pomacanthus (Allen, 1980, includes Arusetta and conduit. Euxiphipops in Pomacanthus; I treat them sepa- Chaetodontoplus, with nine to 11 species, is prob- rately) has nine species, of which two are re- ably absent from the Pacific Plate, but there is a stricted to the Atlantic and one to the eastern report of C. mesoleucus that requires mention. Pacific. All the remaining six species occur in the Fraser-Brunner (1933) and Fowler (1934) consid- Indo-West Pacific, but only Pomacanthus imperator ered Borodin's (1930) Holacanthus bicolor variety ranges nonmarginally onto the Pacific Plate. oahuensis to be a synonym of Chaetodontoplus meso- Centropyge has about 30 species (including at leucus (Bloch), and Fowler considered the Ha- least two undescribed, W.E. Burgess, pers. waiian locality accorded Borodin's specimen to comm.). Two species are restricted to the western be in error (see also discussion of Sparidae for Atlantic, one to Ascension in the eastern Atlantic, another suspicious Borodin locality). Fraser- and all the others to the Indo-Pacific. Of the 25 Brunner did not discuss Borodin's locality but described Indo-Pacific species, 15 occur nonmar- gave the distribution of C. mesoleucus as "East ginally on the Pacific Plate, and of the 15, six are Indies," indicating that he did not consider the endemic to the Plate. Of the six endemics, three Hawaiian Islands to be within the range of the are widely distributed C. loriculus, C. multicolor, species. and C. nigriocellus, Figure 31), one is limited to a Apolemichthys contains seven species (Carlson NUMBER 367 75
•AT HOTUMATUA
• LOR1CUIUS
O NIGRIOCELLUS
• SHEPARDI
FIGURE 31.—Distributions of certain species of the pomacanthid fish genus Centropyge. and Taylor, 1981), of which three occur on the In summary, the Indo-Pacific pomacanthids Pacific Plate nonmarginally: A. arcuatus, endemic consist of eight genera and 60-63 species, of which to the Hawaiian Islands, A. xanthopunctatus, widely six genera and 25 species (11 endemic) occur distributed Plate endemic recorded from Fanning nonmarginally on the Pacific Plate. Sister species (Line Islands) and Kapingamarangi (Eastern of the Plate endemics are not known, but it is Caroline Islands), and A. grtffisi, recorded from inevitable that one or all the sister species of the Fanning and Christmas (Line Islands), Canton Plate endemic species of Apolemichthys are Indo- (Phoenix Islands), and the Gilbert Islands. Three West Pacific species that do not occur on the species of Apolemichthys are variously restricted to Pacific Plate. This fact in addition to the large the Red Sea and western and central Indian number of endemic Plate species of Pomacanthids Ocean, and one species (A. trimaculatus) ranges indicates that the western margin of the Pacific from the east coast of Africa to at least New Plate delineates an important boundary for the Guinea (Steene, 1978; Allen, 1980). I question distribution and speciation of the Pomacanthi- Allen's (1980, table 4) unsubstantiated listing of dae. A. trimaculatus from the Marshall and Gilbert Islands. POMACENTRIDAE Pygoplites, monotypic, is widely distributed in the Indo-Pacific. The damsel fishes consist of 27 genera and Genicanthus, with nine species, is represented on about 300 species of small (maximum size about the Pacific Plate by four species, of which two are 350 mm TL) fishes. The family is circumglobally endemic, one to the Hawaiian Islands, the other distributed and primarily restricted to warm ma- to Pitcairn and the Austral Islands (Randall, rine waters and reef habitats, but a few species 1975b), and two have their eastern distributional occur on temperate coasts or in freshwater in the limits along the western margin of the Pacific tropics. Few genera have been revised in recent Plate (Allen, 1980). years, but much information, particularly on the 76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Indo-Pacific forms, can be found in Allen (1975) nonmarginally onto the Pacific Plate (none en- and several of his subsequent studies. demic), and four that do not. Only five genera (Azurina, Hypsypops, Nexilosus, Amphiprion, the clownfish genus, contains 27 Ncxilarius, Microspathodori) that occur in the Atlan- species (Allen, 1972, 1973, 1975; Burgess, 1981), tic and/or eastern Pacific, do not occur in the all obligatorily associated with sea anemones. Indo-Pacific. Three genera (Chromis, Abudefduf, Only five species occur nonmarginally on the Stegastes) are circumglobally distributed. The re- Pacific Plate. One of the five, A. clarkii, is known maining 19 genera are confined to the Indo-Pa- nonmarginally on the Plate only at Truk (Figure cific, and of these, 11 genera with about 45 species 32), Eastern Caroline Islands, and a second spe- reach the western margin of the Pacific Plate, but cies, A. tricinctus, is known only from the Marshall do not occur nonmarginally on the Plate: Acan- Islands, an unusual area for endemism in fishes. thochromis (monotypic), Amblypomacentrxis (mono-A third species, A. chrysopterus, is a widely distrib- typic), Cheiloprion (monotypic), Dischistodus (6 spe- uted Pacific Plate endemic (Figure 32). Allen cies), Hemiglyphidodon (monotypic), Neopomacentrus(1972) considered A. chrysopterus and A. tricinctus (about 15 species; Allen and Randall, 1980), Par- to be most closely related to A. clarkii. If Allen's aglyphidodon (6 species), Parma (9 species), Premnasassessment of the relationships of these three spe- (monotypic), Pristotis (3 species), and Teixeirichthys cies is correct it would appear that a vicariant (monotypic). event occurred along the western margin of the The following discussion treats the 11 genera Pacific Plate that divided a widely distributed with nonmarginal Pacific Plate representation. ancestral population of Amphiprion into two pop- Abudefduf contains 13 species, one of which is ulations: one, which evolved into A. clarkii, on the represented by two subspecies (D.A. Hensley, west side of the margin, and the other, which pers. comm.). The Atlantic has only one species evolved into the population ancestral to A. chry- (endemic subspecies; the other subspecies occurs sopterus and A. tricinctus, on the east side of the throughout the Indo-Pacific) and the eastern Pa- margin. The location of the vicariant event that cific has only one species, an endemic. The re- resulted in the formation of A. tricinctus and A. maining 11 species (and the subspecies—of a 12th chrysopterus is not hypothesized here. Subsequent species—mentioned above) are confined to the breakdown of the barrier separating the Pacific Indo-Pacific, but only six of these taxa occur Plate populations from those to the west has nonrnarginally on the Pacific Plate. One of these allowed dispersal and sympatry to develop be- six, A. notatus, has been reported on the Plate only tween A. clarkii and A. chrysopterus-tricinctus. I be- from the Phoenix and Marquesas islands (Fowler, lieve that the Caroline Islands conduit provides 1928; Borodin, 1932, as Glyphidodon), but Hensley the means for this dispersal. Allen included seven believes that Borodin's Marquesas report is based other species in his "Clarkii Complex," all of on a misidentification or erroneous locality as- which are restricted to areas well west of the signment (see Pomacanthidae and Sparidae for Pacific Plate margin. Amphiprion melanopus and A. other questionable Borodin reports). One of the peraderaion, which are not members of the Clarkii six Pacific Plate taxa of Abudefduf is endemic to Complex, are widely distributed both on and off the Hawaiian Islands; the other five range from the Pacific Plate, but do not occur in the Indian the Indian Ocean eastward to well out on the Ocean. Plate. Hensley and Allen (1977, fig. 4) mapped Chromis is the most speciose genus of damsel- the distribution of two Abudefduf species restricted fishes. Chromis contains about 65 species (includ- to the Indo-West Pacific that they considered to ing several undescribed species). About 15 species be a sister pair. are confined variously to the eastern Pacific and Amblygliphidodon contains seven species (Allen, Atlantic-Mediterranean. Of approximately 50 1975; Allen and Randall, 1980), three that extend Indo-Pacific species, about half (24) occur non- NUMBER 367 77
FIGURE 32.—Distributions of three species of the pomacentrid fish genus Amphiprion that Allen (1972) considered as forming a closely related group. marginally on the Pacific Plate (the Easter Island Egum Atoll, Samoan, Line, Marshall, Mariana, species, an endemic, is included in this group). New Hebrides, Johnston, Oahu, Gilbert, and So- Eleven or 12 of the Plate species are endemics, ciety. Allen's (1975) tables 1 and 2 also list New variously restricted to the Hawaiian, Marquesas, Guinea and Pitcairn Group within the range of Pitcairn, and Easter islands. One of these species C. acares. Randall and Swerdloff (1973) men- ranges from Rapa and the Gambier Islands tioned that Allen had informed them that he had Group to Pitcairn, and another, C. opercularis, is observed C. acares at Osprey Reef off northern listed in Allen (1975, table 2) as occurring at the Queensland, a nonmarginal, continental plate Tuamotus, Marquesas, Tahiti, and Pitcairn is- locality; Allen (1975) did not mention Osprey lands. Allen does not discuss this species in his Reef within the range of C. acares. Even if Allen's text, although he mentions that there are 36 observation is valid, the presence of C. acares at species of Chromis in the South Seas, and he Osprey Reef may only indicate a relatively recent discusses 36. Allen and Steene (1979) reported westward range expansion of what appears to be that C. opercularis was a widely distributed Indian a common, widely distributed Pacific Plate en- Ocean species (hence my reference to "eleven or demic. Possible supporting evidence favoring re- 12" endemic species of Chromis on the Pacific cency of westward range expansions might be Plate). The other 12 Plate species are more widely Davies and Kinsey's (1977) finding that Great distributed on the Plate, and one of these, C. Barrier Reef coral reefs probably date from only acares, is possibly a widely distributed Plate en- 9000 years ago. Randall and Swerdloff considered demic. Chromis acares is reported (Randall and C. acares to be most closely related to C. lineata, Swerdloff, 1973; Allen, 1975) to occur at the which ranges from Christmas Island (Indian following islands, which are either on the Pacific Ocean) to Palau, New Britain, and the Solomons, Plate or marginally off the Plate: Cook, Austral, and is, thus, only marginally present on the Pa- 78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY cific Plate, and to C. vanderbilti, which ranges from eastern Indian Ocean to just west of the Society the Great Barrier Reef eastward to the Pitcairn Islands, which are the western distribution limits Group, and is, thus, sympatric with C. acares over of D. flavicaudus. The vicariant event, and possibly much of its range. dispersal events, that resulted in the distributions Chrysiptera (= Glyphidodontops) contains aboutof these last two species, if they are sister species, 25 species, of which only six occur nonmarginally is difficult to propose. Dascyllus melanurus, which on the Pacific Plate. At least two of the six Plate is widely distributed in the western Pacific, is species are endemic: C. galbus, from the Cook, known on the Pacific Plate nonmarginally only Austral, Tuamotu, Rapa, and Pitcairn islands from Ponape (my collections) and Kosrae (Eld- (Allen and Randall, 1974; Allen, 1975), and C. redge et al., 1979), Eastern Caroline Islands, al- rapanui, endemic to Easter Island (just east of the though it probably occurs in other, more western, Plate margin). These two species were considered islands of the chain, which have been little col- (Allen and Randall, 1974) to form a closely re- lected. I believe the presence of D. melanurus in lated group with C. notialis, which is known from the Eastern Caroline Islands chain to be evi- New South Wales, Lord Howe Island, and New dence for the operation of the Caroline Islands Caledonia, all off the Pacific Plate, but reaching conduit. close to its margin, and in line with the southern Lepidozygus is monotypic and ranges over much island chains in which C. galbus and C. notialis of the Indo-Pacific. occur. The eastern and western margins of the Plectroglyphidodon contains eight described and Pacific Plate, thus, are boundaries along which at least two undescribed species (Allen, 1975). probable vicariant events occurred that separated Apparently, only the eight described species occur more widely distributed ancestral species into nonmarginally on the Pacific Plate. Two species, populations and allowed them to diverge. The P. flaviventris, from the Tuamotus, and P. sindonis, vicariance mechanism resulting in Easter Island from the Hawaiian Islands, are localized endem- endemism is discussed on pages 136-139. ics. Plectroglyphidodon imparipennis, which should Dascyllus comprises nine species, of which seven probably be considered a widely distributed Plate occur nonmarginally on the Pacific Plate (H.A. endemic, is listed by Allen (1975) as occurring at Randall and Allen, 1977). Of the seven, three are the Palau, Marshall, Line, Fiji, Tahiti, Tuamotu, Plate endemics: D. albisella (Hawaiian Islands), Marquesas, Rapa, Pitcairn, Line, Hawaiian, and D. strasburgi (Marquesas Islands), D. flavicaudus New Hebrides islands, which are either on, or (Society, Pitcairn, Tuamotu, and Rapa islands). marginally off, the Plate, and New Caledonia Randall and Allen reported that D. albisella and and the Loyalty Islands, which are just west of D. strasburgi are most closely related to D. trima- the marginal Plate areas (R.C. Wass, ms., reports culatus, which ranges from the Red Sea and east P. imparipennis from Samoa). Allen (1975) listed coast of Africa to Pitcairn, but is allopatric to the P. phoenixensis as present in the Marshall, Gilbert, other two species. The origin of D. albisella, which Fiji, Samoan, Society, Tuamotu, Marquesas, Randall and Allen considered to be more differ- Phoenix, Wake, and Marcus islands, which are entiated from D. trimaculatus than is D. strasburgi, either on, or marginally off, the Plate, and from probably antedates the origin of D. strasburgi, and Kenn Reef in the Coral Sea, which is somewhat may be the result of island integration (Rotondo west of the Plate margin. The last locality may et al., 1981; see also discussion about Hawaiian indicate a recent westward dispersal of P. phoenix- Islands, page 132). The origin of Marquesas Is- ensis, which I have included as a type I plate lands endemics is discussed in the section dealing endemic in Table 2. Allen also mentioned Maur- with endemism in those islands. Dascyllus flavicau- itius as a possible locality for P. phoenixensis, but dus was reported to be most closely related to the the Mauritius specimens are at least a "color allopatric D. reticulatus, which ranges from the variant" of the typical form. The relationships of NUMBER 367 79
P. imparipennis and P. phoenixensis have not been Pitcairn Island Group (Allen, 1975). proposed. In summary, there are 22 genera and about Pomacentrus comprises 60 species (Allen and 225 species of pomacentrids in the Indo-Pacific. Woods, 1980), but only four species, all widely Eleven genera (50%) and about 72 species (32%) distributed, occur nonmarginally on the Pacific occur nonmarginally on the Pacific Plate, and of Plate; however, one of these four, P. melanopterus, the 72, about 28 (39%) are endemic to the Plate. is known nonmarginally on the Pacific Plate from Overall, the Plate endemics account for more a single specimen collected at Rongelap in the than 12% of the Indo-Pacific pomacentrids. Marshall Islands (Woods and Schultz, 1960); re- confirmation of the presence of P. melanopterus on the Pacific Plate is desirable. At least 18 species POMATOMIDAE of Pomacentrus have their eastern distribution lim- This family consists of a single, monotypic its along the western margin of the Pacific Plate. genus, Pomatomus saltatrix, the bluefish or tailor It is remarkable that so speciose a coral-reef genus (the deepwater genus Scombrops is sometimes in- as Pomacentrus is so poorly represented on the cluded in the Pomatomidae, but is not considered Pacific Plate. here). The bluefish is a commercially and recre- Pomachromis contains four species. Three of ationally important species. It is a strong swim- these are restricted to the Pacific Plate and its mer and ferocious predator, attaining a moder- margin: P. guamensis (Guam), P. exilis (Truk and ately large size (to perhaps 1.5 m TL, although Enewetak), P. fuscidorsalis (Society, Tuamotu, Pit- rarely exceeding 1 m). The bluefish has been cairn Group islands). The fourth species, P. ri- reported to occur in vast shoals extending over chardsoni, ranges from Mauritius (only Indian several hundred square kilometers (Lund and Ocean record; see also Centropyge interruptus, Po- Maltezos, 1970). In the western Atlantic, bluefish macanthidae) eastward to the Ryukyus and Sa- spawn between 55 and 148 km offshore. The eggs moa, on the Plate margin. Although there are and larvae concentrate at the surface along the several possible cladograms that can be con- edge of the continental shelf, but the juveniles structed for these species, it is clear that one of apparently descend to deepwater before moving the vicariant events resulting in the present dis- inshore (Norcross et al., 1974; for additional in- tributions occurred along the margin of the Pa- formation on spawning, and distribution of ju- cific Plate. veniles, see Kendall and Walford, 1979). Both Emery and Allen (1980) estimate that there are young and adults may occur in close shore waters about 35 species of Stegastes (— Eupomacentrus), anads well as estuaries. Adults are known to make Allen and Randall (1980) and Allen and Woods extensive longitudinal migrations. (1980) estimate that there are about 30 species, of The distribution of the bluefish (Figure 33) is which eight occur in the Indo-Pacific. Of these, not approximated by any other fish I know. The five occur nonmarginally on the Pacific Plate, species is discussed here primarily because it is and two of the five are Plate endemics. Stegastes often reported as occurring in the warm waters of aureus is a widely distributed endemic: Tuamotus, all seas. My findings do not support such claims. Marquesas, Line, Gilbert, Phoenix, and Samoan Some distributional reports of bluefish appear to islands. Stegastes emeryi is endemic to southeast be erroneous, while others are probably so, al- Oceania: Tuamotus and Pitcairn Island Group. though the possibility of strays cannot be dis- The relationships of S. aureus were not proposed, counted. There are numerous reports of bluefish but Allen and Randall (1974) related S. emeryi occurring at Valparaiso, Chile, but all of these most closely to S. jenkinsi (a junior synonym of S. emanate from Gay (1848), who reported on a fasciolatus), which is widely distributed in the specimen in the collections of the Paris Museum. Indo-Pacific and sympatric with S. emeryi in the Attempts by myself and C.E. Dawson, on my 80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 33.—Distribution of the monotypic fish family Pomatomidae {Pomatomus saltatrix) (star denotes unspecified locality in Gulf of Guinea).
behalf, to verify the presence of bluefish in Chile species for many years, and J.S. Scott (1959) a have failed, and I do not accept the Chilean Malaysian fisheries biologist, did not list bluefish record (see also Kendall and Walford, 1979), in his Sea Fishes of Malaya. although one must wonder at the absence of Given their general robustness, strong swim- bluefish in the eastern Pacific. ming ability, and pelagic eggs and young, it seems Bleeker's (1844) record of bluefish (as Temnodon, surprising that bluefish are not more widely dis- a junior synonym) from Djakarta (as Batavia), tributed than they are. On the other hand, it is Java, is also doubtful. The record was reported in probably worth examining whether the Atlantic Bleeker's first ichthyological publication (of sev- and Indo-Pacific populations are truly conspe- eral hundred) at a time "when still an amateur, cific. without sufficient education or literature, and without any experience in this discipline [ichth- PRIACANTHIDAE yology]" (M. Boeseman, in litt.). The material on The bigeyes consist of three genera (Priacanthus, which Bleeker based his identification is not Pristigenys, Cookeolus) and probably fewer than 20 known to exist, and I know of no other specimens species of small (probably rarely, if ever, attaining of bluefish from the Indo-Malayan area. Further- 600 mm TL), suprabenthic fishes. They inhabit more, Bleeker never again mentioned the bluefish reef and shelf areas and have planktonic larvae. in any of his numerous compilations of Indone- Some species are reported to have very broad sian fishes. Records of bluefish for the Malay distributions, and one, Priacanthus cruentatus, is Peninsula (for example, La Monte, 1952) and circumtropically distributed. New Zealand (for example, Liem and Scott, 1966) All three genera of priacanthids are reported are also doubtful. Colleagues (J.A.F. Garrick and from the Indo-West Pacific, but only Priacanthus J. Moreland, in litt.) in New Zealand have not and Cookeolus are reported nonmarginally from encountered bluefish although on alert for the the Pacific Plate. NUMBER 367 81
PRISTIDAE which is limited to continental plate areas and is repeated by many other elasmobranch groups. There are two genera (Pristis, Anoxypristis) and Giinther's (1870) record of Pristis from the Cape five or six species of sawfishes. Some of the species of Good Hope is included in Figure 34; however, rank among the largest fishes and are reputed to Penrith (1978) reported that Pristis is absent from attain lengths up to 35 ft (10.7 m), but it is the African coast between Mocamedes, Angola, doubtful that many, if any, exceed 8 m. The and Natal, southeast Africa. species are viviparous and usually benthic in habit in shallow water (to depths of about 10 m), but often are seen swimming when attacking prey. Although sawfishes usually occur in marine PSETTODIDAE waters, they may ascend great distances up rivers and are even found in freshwater lakes (Thorson, The Psettodidae contains a single genus, Pset- 1978). todes, and three species (Stauch and Cadenat, In view of the wide distribution of sawfishes 1965) of moderate-sized (to about 640 mm TL, 9 (Figure 34), including such isolated islands as kg) shallow-dwelling, benthic flatfishes.Th e fam- Bermuda, Bigelow and Schroeder (1953) stated, ily has a disjunct distribution, two species occur- "It is probable that the failure of scientific liter- ring on the tropical west African coast and one in ature to report their presence for any of the island the Indo-West Pacific from tropical east Africa groups of the western tropical Pacific chiefly re- eastward to southern Japan, Queensland, and flects the imperfection of the published record." New Guinea (see Lethrinidae for other families In contrast, I believe that the available distribu- with similar disjunct distributions; also Gobiidae tion records for sawfishes gives a reasonably good for Periophthalmus). The Psettodidae do not occur indication of the group's overall distribution, on the Pacific Plate nonmarginally.
FIGURE 34.—Distribution of the sawfishes, family Pristidae (stars represent unspecified locality records for Madagascar and Vietnam). 82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PSEUDOCHROMIDAE caught pelagically offshore over the continental shelf. Eggs are planktonic offshore (Joseph et al., This Indo-Pacific family includes the formerly 1964), and juveniles may occur as far as 111 km recognized Pseudoplesiopidae and Anisochromi- from shore (Dawson, 1971), or almost at the dae (Springer et al., 1977). The Pseudochromidae shoreline (Springer and Woodburn, 1960). Rachy- contains about five or six genera and 100 species centron appears to be attracted to boats and off- (including undescribed species) of small (to about shore oil-drilling rigs. 200 mm TL), colorful, reef-associated fishes. Lub- Rachycentron is often reported to occur in "all bock (1975) described the spawning behavior of warm seas." Its distribution does not warrant Pseudochromis in aquaria, but was unsuccessful in such claims, which in part, have been fostered by rearing the larvae. The genera, except Aniso- reports of its occurrence in the Juan Fernandez chromis, which contains only two species and is Islands (De Buen, 1957) or Chile (for example, restricted to the western Indian Ocean, and La- Mann, 1954). Rachycentron, however, has not been bracinus, which contains four species (Schultz, reported elsewhere in the eastern Pacific, or other 1967) and is restricted to the western Pacific than marginally on the Pacific Plate (Figure 35). (except for the Indian Ocean coast of Australia) There are no specimens that bear eastern Pacific from Japan south to Australia, are in need of locality data, and my attempts and those of C.E. revision. The most speciose genus, Pseudochromis, Dawson, on my behalf, to confirm the existence contains more than half the species in the family, of Rachycentron in Chile have failed. I believe that but only the Indian Ocean species are well known. all reports of Chilean Rachycentron stem from Del- All but one or two of the Indian Ocean and Red fin (1899), who reported Elacate chilensis [nomen Sea species of Pseudochromis, and all the Indian nudum], based on an identification by Philippi Ocean and Red Sea species of Chlidichthys, appear (in Delfin), from the Juan Fernandez Islands. to be endemic to various parts of those areas Delfin, however, believed the species to be the (Lubbock, 1975, 1976, 1977, 1980). Most of the same as the species called "dorado" at Iquiqui. other species of pseudochromids also appear to be Elacate is a junior synonym of Rachycentron, highly localized, and on this basis Pacific Plate whereas dorado (or dorado de alta mar) is the endemics in the family can reasonably be ex- Ecuadorian, Peruvian, and Chilean common pected. Pseudochromids have not been reported name for the unrelated Coryphaena hippurus, the east of the Marshall and Gilbert islands, except dolphin, which is common in the eastern Pacific. for the Pitcairn Island Group (Randall, in litt.). Delfin should actually be credited with disposing of Chilean records of Rachycentron, but his actions PSEUDOTRICHONOTIDAE obviously have been misinterpreted. It seems re- markable to me, nevertheless, that Rachycentron This family contains a single monotypic genus, does not occur in the eastern Pacific, or, for that Pseudotrichonotus, of small fishes (to 88 mm SL) matter, other than marginally on the Pacific known only from sandy bottom in southern Japan Plate. (Masuda et al., 1975). Adam Ben-Tuvia (in litt.) recently obtained a specimen of Rachycentron from Haifa, Mediterra- RACHYCENTRIDAE nean coast of Israel. This is the only record of The family Rachycentridae (cobias) consists of Rachycentron for the Mediterranean and undoubt- a single genus and species, Rachycentron canadwn, edly represents a recent introduction from the of moderately large size (to about 2 m TL and 50 Red Sea through the Suez Canal. kg). It is an excellent food and game fish, but Suborder RAJOIDEI does not occur in sufficient concentrations to support a commercial fishery. Little is known of The Rajoidei comprises four families (Anacan- the biology of Rachycentron. The adults are usually thobatidae, Arhynchobatidae, Pseudorajidae, NUMBER 367 83
FIGURE 35.—Distribution of the monotypic fish family Rachycentridae (Rachycentron Canadian) (star represents unspecified locality for Argentina; Markle et al., 1980, reported a larval specimen from off Nova Scotia that has not been plotted here).
Rajidae; Compagno, 1973a), 11 genera, and tion records of otherwise continentally restricted about 200 species, of rays, of which about half are taxa, have failed to produce a single record of a in the genus Raja (J. McEachran, pers. comm.). rajoid (P. Struhsaker, in litt.). A partial search of The species are oviparous and benthic in habit, several thousand Pacific deep-sea bottom photo- of moderate to large size (to about 2 m TL), and graphs have also failed to give evidence of rajoids occur at depths from less than 1 m to over 2900 on the Pacific Plate (David Pawson, pers. comm.). m. The four families are treated together here for The order Rajiformes comprises the suborders the sake of convenience. Rajoidei and Rhinobatoidei. The distribution of The distribution map (Figure 36) for the Ra- the entire order parallels the distribution of many joidei does not include records from the Antartic other elasmobranch taxa, which appear to be Plate or for northern regions not shown on the absent from the Pacific Plate, except marginally. map. Most of the records of rajoids from the Pacific Plate are from locations on the New Zea- Suborder RHINOBATOIDEI land Plateau or Chatham Rise, which are part of the continental shelf of New Zealand. Eastern The Rhinobatoidei comprises four families Pacific records of rajoids on the Pacific Plate are (Platyrhinidae, Rhinidae, Rhinobatidae, Rhyn- also associated with juxtaposed continental areas. chobatidae; Compagno, 1973a), and about 50 While it is possible that the deeper, cooler waters species (most in the genus Rhinobatos) of moder- overlying nonmarginal portions of the Pacific ately large to large fishes (up to about 3 m TL) Plate may harbor rajoids, and, thus, de-empha- that are benthic and viviparous in habit, and size the Plate boundaries as limits to rajoid dis- occur at depths ranging from less than 1 m to tribution, it is noteworthy that deep bottom col- about 100 m. The species are primarily marine lections in the vicinity of the Hawaiian Islands, but some enter freshwater. The four families are which have provided many exceptional distribu- treated together here for the sake of convenience. 84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 36.—Distribution of the elasmobranch fishes of the suborder Rajoidei, excluding localities on the Antarctic Plate and in the northernmost portions of the Northern Hemisphere; the Rajoidei and Rhinobatoidei (Figure 37) compose the Order Rajiformes (stars denote records based on egg cases; gray areas denote generalized range).
The rhinobatoids are absent from the Pacific width to about 2.2 m), free-swimming, viviparous Plate, except marginally (Figure 37; records for fishes. The family is present in all tropical and Samoa and Saipan are based on photographs: warm-temperate seas, but is absent from the Pa- J.E. Randall, in litt., Myers and Shepard, 1981). cific Plate. In the Indo-Pacific it occurs as far east The order Rajiformes comprises the suborders as Australia and Okinawa, but has not been Rajoidei and Rhinobatoidei. The distribution of reported from New Guinea or other marginal the entire order parallels the distributions of Pacific Plate localities. The distribution of cow- many elamsobranch groups, which appear to be nose rays conforms to a pattern exhibited by absent from the Pacific Plate, except marginally. numerous other elasmobranch groups, which are absent from the Pacific Plate, except, perhaps, RHINOPRENIDAE marginally. This family contains a single genus and species, Rhinoprenes pentanemus, of small fish (to about 160 SCARIDAE mm TL) that is known only from the Gulf of The parrotfishes are a circumtropically distrib- Papua, New Guinea, and Western Australia (G.R. Allen, pers. comm.). It is apparently re- uted group consisting of about ten genera and 70 stricted to muddy bottoms and areas influenced species. Although numerous species have been by river runoff. revised recently (e.g., Randall and Choat, 1980), none of the genera have been completely and critically revised recently. The parrotfishes are RHINOPTERIDAE among the more conspicuous inhabitants of coral The cownose rays comprise a single genus, reefs. They are free swimming and range in size Rhinoptera, and less than ten species of large (disc from about 0.3 to 1.3 m TL. Thirty-one species NUMBER 367 85
FIGURE 37.—Distribution of the elasmobranch fishes of the suborder Rhinobatoidei; the Rhinobatoidei and Rajoidei (Figure 36) compose the order Rajiformes. in six genera are present nonmarginally on the (specimens at California Academy of Sciences), Pacific Plate (J.E. Randall, in litt.). and Ponape.
SCATOPHAGIDAE SCHINDLERIIDAE The scats consist of two genera and three spe- This family contains a single genus, Schindlena, cies of small (to about 400 mm TL), shallow- and two species of very small (to about 25 mm dwelling fishes that occur primarily in brackish TL) pelagic fishes. Apparently, the species are water, but are not uncommon in freshwater or common (Gosline and Brock, 1960, considered the sea. Until now, they have been reported to them to be possibly among the commonest fishes occur only in the Indo-West Pacific, from the east in the Hawaiian Islands), but because of their coast of Africa eastward to Japan, Palau, Yap, small size and transparent bodies, have been New Guinea, Fiji (Carlson, ms.), and New Cale- reported only from widely scattered localities donia. I collected Scatophagus argus in Ponape,(Madagascar and intermediate localities east to Eastern Caroline Islands, where it is common in Hawaii and Tahiti; Sardou, 1974). Specimens are freshwater streams and river mouths. The pres- usually taken under lights at night with fine-mesh ence of scats in Ponape is evidence for the oper- dip nets, but I obtained a specimen (see ation of the Caroline Islands conduit, which prob- "Addendum") in a rotenone collection on a coral ably enabled the scats to penetrate the Pacific reef at depths of about 9.5-12.5 m in Fiji. Plate. Sterba (1962, fig. 1024; reproduced in Nelson, SCIAENIDAE 1974, map 39) gives a distribution map for the Scatophagidae, but omits the western Indian The Sciaenidae (croakers and drums) com- Ocean portion of the distribution, of which Sterba prises about 40 genera and 150-175 species of was aware (see his page 656), and Palau and Yap small to large (weights up to 66 kg have been 86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY reported for the western Atlantic species, Pogomas cific and the New World have more species in the cromis; Jordan and Evermann, 1905), shallow- former area than in the latter (the Batrachoididae dwelling, suprabenthic fishes. Many of the species are similarly exceptional). are of commercial or sport-fishery importance. The species are oviparous and the larvae, at least SCOMBRIDAE of the marine forms, are planktonic. Most of the species are coastal marine or estaurine, but a few The Scombridae, or mackerels and tunas, com- New World species are restricted to freshwater (a prises 15 genera and 48 species (Collette, 1979) of pattern duplicated by the subfamily Dorosoma- small to very large (to over 450 kg), strong-swim- tinae of the Clupeidae) or have become adapted ming, epiplelagic marine fishes of great commer- exclusively to coral reefs. Surprisingly, there do cial and recreational importance. A few species not appear to be any coral-reef adapted species enter estuaries and ascend rivers, and all of the of sciaenids in the Indo-Pacific (see also discussion species appear to have pelagic young. In the of Sparidae for a similar pattern of reef exclusion). following discussion, an asterisk marks genera I The world distribution of sciaenids is given in consider to be high-seas forms. Figure 38. The absence of sciaenids from the * Acanthocybium is monotypic and found primar- nonmarginal areas of the Pacific Plate and from ily in the high seas around the world in tropical the islands of the Indian Ocean is noteworthy, in and subtropical waters (Collette 1979); however, view of their presence on the islands of the Carib- John E. Randall (in litt.) informs me that he has bean. Another aspect of sciaenid distribution is seen Acanthocybium near coral reefs. unusual: less than haH* the species occur in the * Allothunnus is monotypic and found around Indo-Pacific (Trewavas, 1977; L. Chao, pers. the world in the southern oceans south of 35°S comm.). Most families common to the Indo-Pa- (Collette and Chao, 1975, fig. 69), with one highly
FIGURE 38.—Distribution of the Fishes of the family Sciaenidae (solid dark areas represent generalized distributions; star represents Salton Sea introduction). NUMBER 367 87 unusual record from the southern coast of Cali- neither known nor suspected to occur at these fornia (Collette, 1979). localities (B.B. Collette, pers. comm.). Rastrelliger There are two species of * Auxis, both of which kanagurta appears to have used the island stepping are oceanic and at least one of which, A. thazard, stones of the Caroline Islands conduit to penetrate is circumglobally distributed in warm waters the Pacific Plate, but the other two species of (Collette, 1979; and various authors). Rastrelliger, which reach the western margins of Cybiosarda is monotypic and is distributed along the Philippine and Pacific plates, are still un- the northern three-quarters of the coast of Aus- known from the nonmarginal portions of the tralia and the Gulf of Papua (Collette, 1979). Pacific Plate. See also "Addendum." There are three, essentially allopatric, species Sarda (bonitos) comprises four allopatric species of Euthynnus: E. alletteratus, in the Atlantic; E. (Collette and Chao, 1975) that are usually found qffinis, found throughout the Indo-Pacific, with a in bays and close to shore, although reported to single, valid record from the eastern Pacific; and spawn up to 80 miles (153 km) offshore (Walford, E. lineatus, from the eastern Pacific, with two valid 1937). Their eggs are pelagic, but little is known records from the Hawaiian Islands (Collette, about their larvae and juveniles, which have been 1979). taken pelagically both offshore and nearshore The monotypic genus Gasterochisma, which (Klawe, 1961). The species undergo migrations probably should be placed in a separate family, and during some periods are absent from certain occurs only in the temperate waters of the south- areas where they are otherwise known to occur ern hemisphere (Collette, 1979). (Demir, 1962; Pinkas, 1961). Sarda is restricted Grammatorcynus is monotypic and restricted to, almost entirely to continental plates (Figure 39), and widely distributed in, the Indo-Pacific (in- with the Hawaiian islands a noteworthy excep- cluding the Pacific Plate nonmarginally), espe- tion (see also Scomber above). The Hawaiian spe- cially near coral reefs (Collette, 1979). cies, S. oreintalis, is the same as that which occurs Gymnosarda is monotypic and widely distributed in Japan, Philippines, Indian Ocean, and the throughout the tropical Indo-Pacific, including tropical eastern Pacific. The other Indo-West Pa- the Pacific Plate nonmarginally (Collette and cific species, S. australis, is restricted to New Ca- Chao, 1975, fig. 69). ledonia and the southeast coast of Australia. It is * Katsuwonus is monotypic and cosmopolitan in interesting that the distribution of S. orientalis in tropical and subtropical seas (Collette, 1979). the eastern Pacific separates the distribution of Orcynopsis is monotypic and restricted to the the other eastern Pacific species of Sarda, S. chilen- eastern Atlantic and Mediterranean (Collette and sis, into two discontinuous areas, north and south Chao, 1975, fig. 69). of, and allopatric to, the distribution of S. orientalis Rastrelliger comprises three species (Matsui, (Collette and Chao, 1975, fig. 70). 1967) and is restricted to the Indo-Pacific, except Scomber comprises three species: S. scombrus, from for a recent immigration from the Red Sea the north Atlantic; S. australasicus, in the western through the Suez Canal (Collette, 1979). Prior to Pacific from Japan south to southern Australia, my 1980 collecting trip to Ponape, Eastern Car- and east to the Hawaiian Islands (Hawaiian ex- oline Islands, where I obtained specimens of R. ception), and Socorro Island off Mexico; S.japon- kanagurta, Rastrelliger was not known to occur on icus, antiequatorially distributed worldwide (Col- the Pacific Plate nonmarginally. Through a for- lette, 1979). While it is possible that S. japonicus tuitous error, a distribution map of R. kanagurta occurs nonmarginally on the Pacific Plate, only (Fischer and Whitehead, 1974) indicated the 5. australasicus is definitely so reported (Matsui, presence of the species in the Eastern Caroline 1967). Scomber japonicus has been reported often Islands and other nonmarginal localities on the from the Hawaiian Islands, but these reports Pacific Plate at a time when R. kanagurta was predate Matsui's revision and may be misidenti- 88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY fications. The distribution of Scomber is similar to moness of Scomberomorus in coral-reef habitats else- that of Sarda (Figure 39). where, including the Caribbean. There are 18 species of Scomberomorus (Collette There are seven species of tuna, * Thunnus: T. and Russo, 1979; 1980). The genus is distributed atlanticus of the western Atlantic; T. tonggol of the throughout the warm shore waters of all oceans, Indo-West Pacific (absent from the Pacific Plate but is noticeably absent from the Pacific and nonmarginally); T. albacares, pantropical; T. thyn- Philippine plates nonmarginally (Figure 40). Ten nus (with different subspecies in the Atlantic and of the 18 species occur in the Indo-West Pacific. Pacific), T. obesus, and T. alalunga, all worldwide Nine of the Indo-West Pacific species are rela- in tropical and temperate waters; and T. maccoyi tively restricted in their distributions (see Collette worldwide in southern temperate waters. and Russo, 1979, figs. 9-11 for distribution maps In summary, all but one (Orcynopsis) of the 15 of all species ofScomberomorus), but the distribution genera of scombrids occur in the Indo-Pacific; of the tenth, S. commerson, extending from the Red two of the Indo-Pacific genera {Scomberomorus, Sea and east coast of Africa eastward to Fiji, Cybiosarda) are absent from the Pacific Plate, and almost covers the entire range of the other nine. three genera (Rastrelliger, Sarda, Scomber) have been Scomberomorus commerson the largest Scomberomorus reported on the Plate nonmarginally only from species (to 2.2 m fork length) has also recently the Eastern Caroline or Hawaiian islands. Of the invaded the eastern Mediterranean via the Suez other nine genera of scombrids, the monotypic Canal. In contrast, S. multiradiatus, which is the Grammatorcynus is restricted to, and widely distrib- smallest species of Scomberomorus (to about 300 mm uted in, the Indo-Pacific, and the other eight FL), has the most restricted distribution of the genera are circumglobal in distribution, although species: Gulf of Papua. I find it surprising that some species are restricted to the cooler waters of Scomberomorus does not occur at the nonmarginal the southern hemisphere. Of the 28 tropical spe- islands of the Pacific Plate, in view of the com- cies of scombrids in the Indo-Pacific, only 11-13
FIGURE 39.—Distribution of the fishes of the scombrid genus Sarda. While this study was in press, A.D. Lewis informed me that he had obtained Sarda in the Fiji Islands. NUMBER 367 89
FIGURE 40.—Generalized distribution of the scombrid fish genus Scomberomorus (compiled from Collette and Russo, 1979, and Kong, 1978; dark area in eastern Mediterranean Sea represents recent dispersal of widely distributed Indo-West Pacific S. commerson through the Suez Canal).
(including high seas species) occur nonmarginally family is circumglobally distributed in tropical on the Pacific Plate. The six genera I consider to and temperate waters. Most of the cool-water be shorefishes (Appendix 2, Table C) are repre- species are viviparous, and all the shallow-dwell- sented on the Pacific Plate by only seven species. ing tropical species are oviparous; however, little Collette and Russo (1979, fig. 68) have pre- is known of their reproductive biology. The spe- sented a phylogenetic tree for the scombrid gen- cies are predominantly sedentary (most tropical era. If Gasterochisma, which they consider to be the species) or free-swimming suprabenthic (most most primitive genus, and which is questionably cool-water species), and occur in a wide variety included in the Scombridae, is excluded, the most of marine habitats, with a few tropical species primitive genera tend to be restricted to continen- entering freshwater streams. Much revisionary tal plates and the most specialized genera tend to work remains to be done in the family. be circumglobally distributed. Minous, sole genus of the subfamily Minoinae, contains ten species (Eschymeyer, Hallacher, and SCORPAENIDAE Rama-Rao, 1979; Amaoka and Kanayama, 1981), which are found on soft bottoms in depths (including Synanceidae) of 10-240 m. The genus ranges from the Red Sea There are about 60 genera and 400 species of and east African coast to Japan and Australia, scorpionfishes (W.N. Eschmeyer, pers. comm.). but is absent from the Pacific Plate nonmargin- The species vary in maximum size from about 40 ally. Distribution maps for nine of the species of mm to over 1 m TL (most tropical species are Minous are given in Eschmeyer, Hallacher, and under 400 mm) and are found at depths from Rama-Rao (1979, fig. 1). under I m to several thousand meters (most Choridactylns (two species) and Inimicus (eight tropical species occur at depths under 50 m). The species) form the subfamily Choridactylinae. 90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
These genera are found on silty or open-sand cluding Brachirus), Parapterois, and Brachypterois, bottoms from near shore to 90 m depth, but most which are found only in the Indo-Pacific. Only of the species occur at less than 40 m. The two Ebosia, with two species, has been revised recently, genera are widely distributed in the Indo-West and only Pterois and Dendrochirus, widely distrib- Pacific (Eschmeyer, Rama-Rao, and Hallacher, uted in the Indo-Pacific, are present on the Pacific 1979, fig. 8), but are absent from the Pacific Plate Plate nonmarginally. The single species, each, of nonmarginally. According to Eschmeyer, Rama- Pterois and Dendrochirus that occur in the Ha- Rao, and Hallacher (1979), these fishes waiian Islands, are endemics (Eschmeyer and "seemingly could, but [do] not cross the Andesite Randall, 1975). Line." The subfamily Pteroidichthyinae includes The subfamily Synanceiinae (stonefishes) con- Rhinopias, Pogonoscorpius, Hipposcorpaena, and Pter- sists of six genera: Erosa, Dampierosa, Pseudosynan- opelor (Eschmeyer, Hirosaki, and Abe, 1973), ceia, Leptosynanceia, and Trachicephalus, all mono- which are restricted to the Indo-Pacific. Rhinopias, typic, and Synanceia, with four or five species with four species, is the only one of these genera (Eschmeyer and Rama-Rao, 1973). These genera that is reasonably well known and that occurs on are restricted to the Indo-Pacific and occur in the Pacific Plate (Hawaiian Islands and Condor shallow water in habitats varying from coral reefs Reef, Eastern Caroline Islands). The other genera to muddy bottoms in estuaries. Only one of the are rare and probably monotypic. All the genera species, Synanceia verrucosa, occurs on the Pacific are moderately deep dwelling, although species Plate nonmarginally, and it ranges from the Red of Rhinopias occur at depths available to SCUBA Sea and east coast of Africa to the Tuamotus. divers. Giinther (1860) reported a common Indo-West The genera that follow are discussed without Pacific species of stonefish (S. horrida) from several reference to subfamily. Pacific localities and from St. Helena in the Six deep-dwelling genera, two represented by Atlantic. Eschmeyer (1971:503) believed the St. Hawaiian endemics, are reported from the Ha- Helena locality was probably in error, and this waiian Islands (Eschmeyer and Randall, 1975), was reaffirmed by Eschmeyer and Rama-Rao but are otherwise unknown on the Pacific Plate: (1973:3). Briggs (1974:93) unfortunately (but un- Ectreposebasetes (circumglobal), Neomerinthe (Ha- derstandably) overlooked Eschmeyer's references waiian endemic), Phenacoscorpius (also western At- and accepted the presence of Synanceia at St. lantic, western Pacific), Plectrogenium (monotypic, Helena. Japan and Hawaiian Islands), Pontinus (circum- The subfamily Tetraroginae was tentatively global with Hawaiian endemic species), Setarches recognized by Poss and Eschmeyer (1975), in (monotypic, Atlantic and Indo-Pacific). The cir- which they included Amblyapistus, Hypodytes, Lio- cumglobal genus Helicolenus, also deep dwelling, cranium, Ocasia, Paracentropogon, Ptarmus, Snyderina, has been reported (Abe and Eschmeyer, 1972) on Tetraroge, and "several other genera of uncertain the Pacific Plate nonmarginally only from the status." The Tetraroginae are confined to the Emperor Seamount chain, which is continuous Indo-West Pacific (absent from the Pacific Plate) with the Hawaiian Island chain. and occur in shallow water out to depths of about Taenianotus is monotypic and widely distributed 300 m. Only Ocosia, with five species, has been in the Indo-Pacific. William N. Eschmeyer in- revised in recent years, although information on forms me that Taenianotus has been photographed some of the genera and species can be found in in the eastern Pacific (Galapagos Islands), but J.L.B. Smith (1958). that no specimens from that area are available. The subfamily Pteroinae can be inferred from Scorpaena and Scorpaenodes are speciose genera Eschmeyer and Rama-Rao's (1977) study of that are circumglobally distributed; Scorpaenopsis Ebosia to include Ebosia, Pterois, Dendrochirus (in- is also speciose, but restricted to the Indo-Pacific. NUMBER 367 91
Most of the species of these three genera are and it occurs only in the Hawaiian Islands (Ha- shallow-dwelling reef inhabitants. Many of the waiian exception). species are island endemics, and at least two may Microcanthus is a small, free-swimming, supra- be widely distributed Pacific Plate endemics: Scor- benthic species, probably not attaining 200 mm paena coniorta, reported only from the Hawaiian,TL, that is found close to shore, even in tide pools Johnston, Wake, and Line islands (Eschmeyer as juveniles, in rocky and coral-reef areas. The and Randall, 1975), but present also at Marcus species is antiequatorially distributed: Hawaiian and Makatea islands (J.E. Randall, in litt.), and Islands; Japan south to Hong Kong and Taiwan; Scorpaenopsis fowleri, known only from the Tua- southern Queensland; New South Wales; south- motu, Society, Marshall, and Hawaiian islands western Western Australia; Lord Howe Island; (Figure 26; Eschmeyer and Randall, 1975). The and New Caledonia (see also Aulacocephalus under closest relative (sister species) of Scorpaena coniortaSerranidae). Reports by various authors that M. has not been proposed, but Eschmeyer and Ran- strigatus occurs in the Philippines appear to stem dall (1975) and Eschmeyer (pers. comm.) state from Weber and de Beaufort's (1936) statement that S. coniorta belongs to a group (Sebastapistes)that the species "probably" occurs in the northern containing three other species: S. albobrunnea Philippines. That may be, but there is no firm (widely distributed in the Indo-Pacific), S. tink- record from the Philippines. The presence of M. hami (Asian, poorly known), and an unnamed strigatus in the Hawaiian Islands is possibly the species from French Oceania (Pacific Plate local- result of dispersal from Japan via the Kuroshio ity). Thus, there is a one-in-three possibility that extension. the sister species is restricted to the Indo-West The other genera mentioned above are var- Pacific. The sister species of Scorpaenopsis fowleriiously found along the southern half of the coast has not been proposed. of Australia, Lord Howe Island, south coast of From the sample of scorpaenids discussed Africa, North Island of New Zealand, and tem- above, it is obvious that there are many more perate eastern Pacific. The distribution of the genera and species present in the Indo-West Pa- family is, therefore, antiequatorial. cific than are present on the Pacific Plate non- marginally. Many of the genera that are missing SCYLIORHINIDAE on the Plate are shallow reef-dwelling forms that might be expected to occur on the Plate. On the The catsharks comprise 17 genera and 86 spe- other hand, there are a few widely distributed cies of small- to moderate-sized (to about 1.5 m), Plate endemics (as well as some species restricted predominantly bottom or near-bottom dwelling to single island groups), and this number will fishes (S. Springer, 1979). The catsharks contain probably be increased when the species of un- viviparous and oviparous species, and shallow revised genera are better known. and deep-dwelling species. A few species are found in Indo-West Pacific reef-type habitats, but the distribution of the family is best characterized SCORPIDIDAE as antiequatorial in nature. As is true of most The composition of the Scorpididae appears to elasmobranchs, the catsharks, with one exception, be unsettled, but includes at least the following are restricted to continental plates. The only oc- nominal genera: Scorpis, Atypichthys, Medialuna, currence of a catshark on the Pacific Plate non- Microcanthus, Parascorpis, Parapsettus, Neatypus, andmarginally is that of Parapristiurus spongiceps, Neoscorpius. It does not appear that there are more which was collected in the Hawaiian Islands at a than about ten species in the family; only one of depth between 572 and 1462 m. The same species the genera, the monotypic Microcanthus (M. stri- has also been reported from the Banda Sea (We- gattis), occurs nonmarginally on the Pacific Plate, ber, 1913). 92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
SERRANIDAE Cromileptes in the Hawaiian Islands is the result of aquarium releases but wafting from Japan via (including Grammistidae and Pseudogrammidae) the Kuroshio Extension is another possibility. I The sea basses are a circumglobally distributed have not included Cromileptes in Appendix 2, family containing more than 50 genera and pos- Table C. sibly more than 400 species. Much revisionary Aulacocephalus, monotypic (A temmincki), is an work is needed in the family, and the Indo-Pacific Indo-Pacific genus inhabiting shallow, rocky forms are the poorest known. All the genera, and areas. It has an essentially antiequatorial distri- most of the species, that occur on the Pacific Plate bution in the Indo-Pacific. On the Pacific Plate, appear to range widely in the Indo-Pacific. (See it is known only from Rapa, where it was taken Lutjanidae for discussion of introduction of Ce- in relatively deep water (70 fathoms = 107 m; phalopholis into the Hawaiian Islands, where the Randall, 1978a). It is also known from middle genus is not native.) Japan and Cheju Island, Korea, south to Taiwan Serranids, for the most part, are shallow-dwell- and Thailand, from southern Africa, Mauritius, ing and benthic or suprabenthic in habit. They Kermadec Islands, New Zealand, and the Anda- occur primarily around rocky areas and coral man Islands (Randall, in press b). reefs. They vary in adult size from under 100 mm The subgenus Mirolabrichthys of Anthias was re- to about 2.5 m TL. Nineteen genera and about vised by Randall and Lubbock (1981a). Katay- 82 species are present nonmarginally on the Pa- ama and Masuda (1980) and Randall and cific Plate (J.E. Randall, pers. comm.). McCosker (in press) described two species not Diploprion, a coral-reef genus with two species treated by Randall and Lubbock. Mirolabrichthys (Randall et al., 1971), is restricted to the Indo- comprises 13 species of small, planktivorous, reef West Pacific. One species, 0. drachi, is endemic to inhabitants restricted to the Indo-Pacific (the the Red Sea, the other, D. bifasciatum, ranges from distributions of eight of the species are given on India to Ceylon eastward to the Solomon Islands Figure 41). Eight species occur on the Pacific and New Hebrides (both on the margin of the Plate nonmarginally, of which four are endemic Pacific Plate), southward to New Caledonia to the Plate: A (M.) regalis, endemic and common (Fourmanoir and Laboute, 1976) and the Great in the Marquesas; A. (M.) bartlettorum, from the Barrier Reef, and northward to southern Japan. Marshall, Phoenix, and Line Islands (Randall Belonoperca, monotypic (B. chabanaudi), is a and McCosker, in press, give records for the last coral-reef inhabitant that ranges from the east two island groups); A. (M.) aurulentus, from the coast of Africa eastward to Samoa and the Gilbert Line Islands; and A. (M.) pascalus, widely distrib- Islands, on the Pacific Plate. Randall et al. (1980, uted on the Plate and reaching the western mar- fig. 2) give a spot distribution map for Belono- gin of the Philippine Plate (Figure 41). The dis- perca, to which can be added Ponape and Ant tribution of the other species is as follows: two Atoll, Eastern Caroline Islands (my collections, species (4. (M.) evansi, A. (M.) ignitus) restricted USNM). to the Indian Ocean, three {A. (M.) pavirostris, A. Cromileptes, monotypic (C. altivclis), is also a (M.) flavoguttatus, A. (M.) tuka) limited to the coral-reef inhabitant, and ranges from the Nico- western Pacific, two {A. (M.) smithvanizi, A. (M.) bar Islands, eastern Indian Ocean, eastward to bicolor) widely distributed in the Indo-Pacific, at southern Japan, Guam, Admiralty Islands, and least as far west as Cocos-Keeling, one {A. (M.) New Caledonia. It is known nonmarginally on dispar) widely distributed on the Pacific Plate and the Pacific Plate only from the Hawaiian Islands, in the western Pacific, and one {A. (M.) lori) where only a single specimen has been obtained limited to the western Pacific and Pacific Plate. (Randall, 1981a) and another observed, both re- Prior to Randall and Lubbock's revision, A. cently. Randall believes that the occurence of (M.) tuka and A. (M.) pascalus were considered to NUMBER 367 93
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A REGALIS
•A-TUKA
FIGURE 41.—Distributions of eight of the 13 species forming the subgenus Mirolabrichthys of the serranid fish genus Anthias. be conspecific subspecies, and even Randall and islands); hence, none of the species ranges across Lubbock included them in the same key couplet, the margin of the Pacific Plate. indicating the possibility that they are sister spe- Plectranthias was revised by Randall (1980b); cies. These two species are sympatric in the Palau an additional species was described from Japan Islands, on the eastern margin of the Philippine by Katayama and Masuda (1980) and another Plate (see also discussion of Nemateleotris, Gobi- from the Coral Sea by Fourmanoir and Rivaton idae), but their distributions, otherwise, are allo- (1980). The genus contains 32 small species, most patric and defined by the joint margin of the of which occur at depths of over 100 m; however, Philippine and Pacific plates (Figure 41). If A. several species are shallow dwelling. One species (M.) tuka and A. (M.) pascalus prove to be sister is limited to the tropical western Atlantic, but all species, the vicariant event that divided the range the others are restricted to the Indo-Pacific. Seven of their common ancestor would appear to have species, including two that are deep dwelling, occurred along the joint margin of the plates. occur nonmarginally on the Pacific Plate, and Sympatry of the two species in the Palaus indi- three of these are each endemic to single islands: cates that subsequent dispersal has occurred, with P. htlenae (Oahu, deep-dwelling), P. taylori (Can- the Caroline Islands conduit permitting the Pa- ton, deep dwelling), P. cirritoides (Rapa, shallow cific Plate species to disperse westward. dwelling). Interestingly, two shallow-dwelling Odontanthias comprises nine species and occurs species, P. nanus and P. fourmanoiri, are widely only in the Indo-Pacific (Mauge, 1980). One distributed on the Pacific Plate, but are otherwise species is confined to the western Indian Ocean, known only from Christmas and/or Cocos-Keel- five to the western Pacific (from Celebes to ing islands, both eastern Indian Ocean. If the two Guam), and three to the Pacific Plate (two in the species are actually absent from the area between Hawaiian Islands; one in the Cook and Society the Indian Ocean and Pacific Plate, extinction of 94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY populations from the intermediate area is indi- and Nelson (1974) 117 species, of soles. About cated. half the genera and species are restricted to the John E. Randall informs me (in litt.) that Indo-Pacific, although the family is almost cir- Epinephelus socialis is a widely distributed Pacificcumglobally distributed in shallow tropical and Plate endemic. The Bishop Museum has speci- warm-temperate marine waters. Some species oc- mens from the Society Islands, Tuamotus, Pit- cur well up rivers, and some in deep coastal cairn Group, and Rapa. The USNM collections waters, but all appear to be inhabitants of sandy have specimens additionally from the Tokelau, and muddy bottoms. Most of the species are Samoan, Phoenix, Line, and Marshall islands. under 300 mm TL, but some soles in European waters may attain over 500 mm TL. The larvae SIGANIDAE are planktonic. Soles are poorly represented on the Pacific The Siganidae contains a single genus (Siganus) Plate, where only two genera, Soleichthys and As- and 26 species of small (to about 425 mm TL), eraggodes, are known. Both are in need of revision. free-swimming, shallow-dwelling fishes. A revi- Pietschmann's (1938) report of Achirus thepassi sion of the family, which I have not seen, by D.J. from Honolulu appears to be a misidentification Woodland is nearing completion. The species are of a damaged specimen of Aseraggodes kobensis (see abundant in reef areas, particularly on the grassy also Clark and George, 1979:106). flats around reefs. Pelagic young are often taken close to shore under a light at night. The family Of the Indo-West Pacific genera, only Parda- is widely distributed in the Indo-Pacific, from the chirus has been revised recently. Clark and George Red Sea and east coast of Africa to the Pitcairn (1979) recognized four species, with the overall Island group. Two species have become well es- range for the genus extending eastward from the tablished in the eastern Mediterranean, having Red Sea and east coast of Africa to southern invaded that area via the Suez Canal (Ben-Tuvia, Japan and Palau, Solomon, New Hebrides, 1966). The fact that an endemic siganid species Tonga, and Samoan islands, along the margin of occurs in the Fiji Islands (Gawel and Woodland, the Pacific Plate (see their fig. 1 for a distribution 1974) suggests the possibility that other siganids map). One species inhabits estaurine areas, others may be endemic to Pacific Plate island groups. occur in sandy areas about reefs.
SOLENOSTOMIDAE SlLLAGINIDAE The Solenostomidae consists of a single genus, The Sillaginidae comprises three genera and Solenostomus, and perhaps as many as five species about 25 species of small (to about 500 mm TL) of small (to about 160 mm TL), benthic, marine suprabenthic, shallow-dwelling fishes that occur fishes restricted to the Indo-Pacific. The family is in marine and estaurine areas, and occasionally widely distributed (but rarely collected) in the enter freshwater. They usually occur over sandy Indo-West Pacific, from the Red Sea and east bottom. The family is restricted to the Indo-West coast of Africa eastward to Japan, Guam, New Pacific (Figure 42), except for a recent introduc- Guinea, Australia, and Fiji (Carlson, ms.). The tion into the eastern Mediterranean via the Suez only nonmarginal Pacific Plate records for Sole- Canal (Mouneimne, 1977). A revision of the fam- nostomus are for a single specimen (25.7 mm SL) ily by R.J. McKay is nearly completed. taken off Bikini Atoll in an open plankton net lowered from the surface to 175 fathoms (320 m) SOLEIDAE and hauled open to the surface (Schultz, 1953c), and a photograph of an adult taken by a Kwa- (including Achiridae) jalein resident (J.E. Randall, in litt.). It seems The soles are poorly known taxonomically. possible that Solenostomus is wafted planktonically Norman (1966) reported that there are 31 genera, onto the Pacific Plate, but that breeding popu- NUMBER 367 95
FIGURE 42.—Distribution of the fishes of the family Sillaginidae (compiled mostly from information provided by R.J. McKay, with additions from the literature). lations do not exist in that area. I have, neverthe- limited to the Indo-West Pacific, except for one less, recognized the family as represented on the record of M. berda from Raiatea, based on a report Pacific Plate (Appendix 2: Tables A, B, C). by Borodin (1932). The distribution of M. berda, which was mapped by Munro (1949, fig. 3), SPARIDAE almost covers the combined distributions of the other species of Mylio. The Raiatea record is The porgies are a large family containing over prominent for its wide geographic separation 25 genera and 100 species of small to moderately from all other records of Mylio, which are from large, suprabenthic fishes, many of which are of continental plate areas. It would be most note- commercial importance. The species are typically worthy if Mylio occurred on the Pacific Plate marine, but some frequently enter estuaries. They nonmarginally, but I suspect that Borodin either occur over all types of bottom in habitats as misidentified or associated incorrect locality data diverse as mangrove swamps and coral reefs. The with his specimen (see Pomacanthidae and Po- family is distributed worldwide in tropical and macentridae for other questionable Borodin re- temperate shelf waters, but with one exception ports). Munro (1949, fig. 5) also figured the dis- (see below) has not been reported from the non- tribution of Rhabdosargus sarba, which closely par- marginal portions of the Philippine and Pacific allels the distribution of Mylio (less Raiatea), and Plates (Monotaxis, which is a lethrinid, has often which embraces the distribution of the other spe- been reported as a sparid from Pacific Plate cies of Rhabdosargus, which are confined to the localities). Sparids are common about the coral western Indian Ocean (Smith, 1979). reefs of the Caribbean, just as some sciaenids are, but neither of these two families occurs on the SPHYRAENIDAE coral reefs of the Indo-Pacific. Munro (1949) reported the distribution of the The barracudas consist of single genus, Sphy- genus Mylio, which comprises seven species and is raena, and about 20 species (de Sylva, 1974) of 96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY small to moderately large (to about 2 m TL), Reports of Squatina from tropical latitudes usually free-swimming fishes. The genus is almost circum- indicate depths of capture in excess of 100 m globally distributed in tropical and warm-tem- (Poll, 1951; Bullis and Thompson, 1965; Cervi- perate marine waters. There are about 11 species gon, 1966). Angel sharks, like most elasmo- in the Indo-Pacific, but their distributions within branchs, are absent from the nonmarginal por- the area are inadequately known (de Sylva, 1974). tions of the Pacific Plate. Their easternmost oc- At least two species occur nonmarginally on the currences in the Indo-West Pacific are in Japan Pacific Plate, and one of these, S. barracuda, essen- and Australia (New South Wales). tially duplicates the entire range of the genus, Kossmann and Rauber (1877) reported on a except that it is absent from the eastern Pacific Mediterranean species of angel shark, Squatina (de Sylva, 1963), where other species of Sphyraena squatina, that reputedly was taken in the Red Sea. occur. Kossmann's. collections were obtained from both the northern (including the Gulf of Suez) and southern portions of the Red Sea and were mixed. SPHYRNIDAE Possibly the Squatina specimen came from the The hammerhead sharks consist of a single Gulf of Suez and represents an incursion from the genus, Sphyrna, and nine viviparous species (Gil- Mediterranean. There are no other reports of bert, 1967) of moderately large to very large fishes angel sharks from the Red Sea (or northern por- (maximum TL varying from about 1.0 to more tions of the Indian Ocean) and I question the than 6.0 m). The family is circumglobally distrib- validity of Kossmann and Rauber's report. uted, as are two of the three largest species:
Sphyrna zygaena has a bitemperate distribution, STEGOSTOM ATI DAE essentially surrounding the distribution of S. lew- ini, which is circumtropical (both species occur in This family is discussed under the Orectolobi- Hawaiian waters). The third large species, S. dae. mokarran, which apparently occurs along all warm continental coasts, is absent from the Pacific Plate STROMATEIDAE nonmarginally. All the other, small, species, ex- cept S. blochii, are variously restricted to the The pomfrets comprise three genera (Pampus, coastal waters of the Atlantic and eastern Pacific. Peprilus, Stromateus) and about 15 species, of small Sphyrna blochii is limited in distribution to the (to about 500 mm TL), commercially important northern coasts of the Indian Ocean, including fishes that are widely distributed, but restricted the Red Sea and Persian Gulf, and to the coasts to tropical and warm-temperate continental plate of Indonesia and Malaya, eastward to the Phil- coastlines, where they may enter brackish water. ippines. The supposed occurrence of S. blochii in Haedrich (1967) gives a generalized distribution Australia has been discredited by Scott (1973). map for each of the genera, which are mostly allopatric, and notably absent from the coasts of eastern Africa, the Red Sea, Australia, and New SQUATINIDAE Guinea. Pampus has been reported from Hawaii The angel sharks comprise a single genus, Squa- by Fowler (1938), who questioned (as did Hae- tina, and about ten species of small to moderately drich, 1967) the correctness of the locality as- large (to about 2 m TL), benthic, viviparous signed his specimens. Pampus, which is restricted fishes. The species live in shallow to moderately to, and is the only genus of the family in, the deep coastal waters, and the genus is, for the most Indo-West Pacific, does not occur on the Pacific part, antiequatorially distributed on all continen- Plate. Pampus was reported from the Adriatic by tal plates (continuously distributed from the Soljan (1963; as Stromateus griseus), but this record Mediterranean to Angola in the eastern Atlantic). is also untenable. NUMBER 367 97
SYNGNATHIDAE Enchelyocampus is monotypic and is known only from Australia and Palau, on the margin of the The pipefishes and seahorses comprise about Pacific Plate (Dawson and Allen, 1978). 56 genera and 240 species (C.E. Dawson, pers. Festucalex contains five species and occurs only comm.) of small (to about 610 mm TL) fishes. in the Indo-Pacific (Dawson, 1977c, fig. 10). Only The species are usually shallow dwelling and one of the species occurs on the Pacific Plate benthic in marine waters, but may occur at mod- nonmarginally, and only at the Hawaiian Islands erate depths, in freshwater, and pelagically (usu- (Hawaiian exception). ally associated with seaweed). Young are carried Halicampus contains two species (Dawson and by the males until parturition, after which the Randall, 1975) and is restricted to the Indo-West young may become pelagic. Much revisionary Pacific (Red Sea to Solomon Islands; C.E. Daw- work remains to be done among the syngnathid son, pers. comm.). genera, but a number of the Indo-Pacific genera Hippichthys contains three species and is re- have been revised recently and these are dis- stricted to the Indo-Pacific (Dawson, 1978c, fig. cussed. 3). Two of the species range from the western Apterygocampus is monotypic and known only Indian Ocean to the margin of the Pacific Plate; from Ceram, Moluccas (Dawson and Allen, the third ranges from the western Indian Ocean 1978). to the Eastern Caroline and Gilbert islands, well Bhanotia contains two species that are restricted out on the Plate. to the Indo-West Pacific, from the eastern Indian Ichthyocampus is monotypic and known only Ocean eastward to the New Hebrides, on the from the Indo-West Pacific, from the west coast margin of the Pacific Plate (Dawson, 1978b, of India to Borneo and Java (Dawson, 1977c, fig. 4). fig. 4). Campichthys contains four species and occurs Lissocampus contains five species and is known only in the Indo-West Pacific, from the east coast only from Australia, New Zealand, Chatham Is- of Africa and Australia (Dawson, 1977c, fig. 19). lands (nonmarginal Pacific Plate locality, but on Choeroichthys contains five species and occurs the continental shelf of New Zealand, hence, not only in the Indo-Pacific (Dawson, 1976, 1978a). considered a Plate locality for my purposes), and Two species are widely distributed in the Indo- the Red Sea (Dawson, 1977a). Pacific, one is limited to the western Indian Mannarichthys is monotypic and is known only Ocean, one to the Moluccas, and one to Australia. from the Indo-West Pacific, from the east coast Corythoichthys contains ten species and occurs of Africa to the Great Barrier Reef (Dawson, only in the Indo-Pacific (Dawson, 1977b, figs. 6 1977c, fig. 4). and 11). Although eight species range onto the Nannocampus contains two species, one known margin of the Pacific Plate, only four continue only from the southeast coast of Africa, the other nonmarginally on to the Plate. from Australia and Indonesia (Dawson, 1979b). Doryrhamphus was revised by Dawson (1981), Notiocampus is monotypic and known only from who recognized five species (including eight taxa; Australia (Dawson 1979b). one species with three subspecies, another with Oostethus is monotypic and consists of four al- two subspecies). The genus is restricted to the lopatric subspecies (Dawson, 1979a). The genus Indo-Pacific except for D. e. excisus, which ranges occurs along all warm coastlines (including is- from the east coast of Africa to the Gulf of lands on the Pacific Plate), except those of the California and Galapagos Islands in the Eastern eastern Pacific, with one exception: the western Pacific. Four taxa reach the western margin of Atlantic subspecies is present on the Pacific coast the Pacific Plate, but of these only D. e. excisus of Panama, and Dawson believes it to have occurs nonmarginally on the Plate. Dawson pro- reached that area via the Panama Canal. Another vides distribution maps for all the taxa. subspecies is restricted to the Western Indian 98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Ocean, the third to the eastern Indian Ocean and ing 500 m. The family is circumglobally distrib- central and western Pacific Ocean, and the fourth uted, as is one of the three genera, Synodus. to the west coast of Africa. Trachinocephalus is monotypic. It is present in Penetopteryx contains two species (Dawson and the eastern and western Atlantic and throughout Allen, 1978). One species ranges from Mauritius the Indo-West Pacific. It is absent from the east- to the Line Islands; the other is known only from ern Pacific and has been obtained on the Pacific the Bahamas. Plate nonmarginally only in the Hawaiian and Phoxocampus contains three species and is found Society islands (Anderson et al., 1966, fig. 15). only in the Indo-Pacific (Dawson, 1977c, fig. 4). Trachinocephalus was reported to occur on the Pa- Only one of the species occurs on the Pacific Plate cific Plate at the Society Islands (Vaillant, 1887; nonmarginally. Fowler, 1938) and the Cook Islands (Seale, 1906). Siokunichthys contains six species and is re- Fowler (1928) reidentified Seale's Cook Islands stricted to the Indo-West Pacific, from the Red specimens as a Synodus species (Anderson et al. Sea to Fiji, on the margin of the Pacific Plate noted Seale's misidentification, but erroneously (C.E. Dawson, pers. comm.). entered the Cook Islands record on their fig. 15). Stipecampus is monotypic and known only from Randall (1973) cast doubt on Vaillant's Society southern Australia (Dawson, 1977c, fig. 10). Islands record, which Randall believed was prob- Urocampus contains two species and is known ably based on a species of Synodus. Fowler's (1938) only from the Indo-West Pacific: Japan south to Society Islands specimens are postlarvae; their Ryukyu Islands, Australia, and New Guinea identification has been verified by Randall (1973) (Dawson, 1980). and W. Saul (Academy of Natural Sciences of Yozia contains three species and is distributed Philadelphia, pers. comm.). It would be of interest from the Red Sea and east coast of Africa to to ascertain the presence of adult Trachinocephalus Japan and Australia (Dawson et al., 1979). on the Pacific Plate other than at the Hawaiian In summary, of the 22 genera and 54 species of Islands. Indo-Pacific pipefishes discussed, only eight gen- Saurida has a distribution similar to that of era and 12 species (22.2%) are present on the Trachinocephalus, except that Saurida is more com- Pacific Plate nonmarginally. No Pacific Plate monly distributed on the Pacific Plate. Shindo endemics are among these taxa. Five other genera and Yamada (1972) reported that there are nine (Appendix 2, Table C) and perhaps 13 more species of Saurida in the Indo-Pacific, but recent species of pipefishes also occur on the Pacific study of the genus indicates that there is at least Plate, but I expect that the species in these genera one undescribed species in the area and that a that are restricted to the Indo-West Pacific will revision is needed (R. Waples, in litt.). Sulak exhibit the same or greater ratio to the genera (1977) includes Saurida with the Harpadontidae and species occurring on the Pacific Plate as (q.v.). noted so far. There are about 30 species of Synodus, of which 21 occur in the Indo-Pacific.(R.F. Cressey, 1981, SYNODONTIDAE and pers. comm.). Only eight of the Indo-Pacific species occur on the Pacific Plate nonmarginally, (not including Harpodontidae) and one of these is a Plate endemic. Two species, The lizardfishes comprise three genera and 5. usitatus and S. ulae, are known only from Japan about 45 species of small- to moderate-sized (max- and the Hawaiian Islands. Synodus capricomis is imum about 635 mm TL), benthic fishes with known only from Easter and Pitcairn islands. pelagic larvae. The species occur on rocky, sandy, or muddy bottoms, often about coral reefs, and TERAPONTIDAE occasionally in estuaries. Most are shallow dwell- There are 15 genera and about 40 species of ing, but some species are taken at depths exceed- terapontids (Vari, 1978; Vari and Hutchins, NUMBER 367 99
1978, and Vari, pers. comm.). The species are of Tyler, pers. comm.) of small to moderate size small to moderate size (65-800 mm maximum (maximum TL ranging from about 50 to 760 SL). More than two-thirds of the species are mm). The species are mostly suprabenthic, shal- restricted to freshwater; the remaining species, low-dwelling, tropical and warm-temperate ma- although primarily marine, often enter fresh and rine forms, but some species are pelagic in the brackish water, but breed in the sea. Vari believes open sea; others enter or occur only in freshwater that the terapontids originated in freshwater, (tropically), and some occur only at moderate with a series of marine representatives having depths (to about 500 m). The Indo-Pacific species been derived from the freshwater forms, followed of only one tropical genus have been revised in by a secondary reinvasion of the freshwaters of recent years. Probably 20 species in four tetrao- Australia and New Guinea. The species are all dontid genera occur nonmarginally on the Pacific shallow dwelling and suprabenthic, although I Plate. have seen the young of a marine form bury Canthigaster contains 24, primarily reef-inhabit- themselves in sand ..close to shore. Little has been ing, species (Allen and Randall, 1977; Lubbock reported on the life history of terapontids, but and Allen, 1979). One species is restricted to the Munro (1945) reported that the postlarvae of one Atlantic, occurring in both the eastern and west- species are planktonic in shallow water. The fam- ern portions, and one species is restricted to the ily is restricted to the Indo-West Pacific (Figure tropical eastern Pacific. Among the 22 Indo-Pa- 43), except for a recent introduction into the cific species (one of which reaches the eastern eastern Mediterranean (Lourie and Ben-Tuvia, Pacific), four occur only in the Red Sea and/or 1970). Indian Ocean. The remaining 18 species are var- iously distributed in the Indo-Pacific, with 13 TETRAODONTIDAE ranging onto the Pacific Plate nonmarginally. Of The pufferfishes are a circumglobal family the 13, eight are widely distributed in the Indo- comprising about 14 genera and 110 species (J.C. Pacific, two are known only from the Hawaiian
FIGURE 43.—Distribution of the fishes of the family Terapontidae. 100 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Islands, one only from Rapa, one only from the possible to determine what family was involved). Marquesas, and one, C. epilampra, appears to be The only records of torpedos (Figure 44) from the a widely distributed Pacific Plate endemic (Ha- Pacific Plate, other than marginally, are for the waiian, Society, Rarotonga, Palau, and Solomon Chatham Islands, which are on the continental islands; Figure 26). Allen and Randall related C. shelf of New Zealand (hence not considered Pa- epilampra most closely to the Marquesas and Rapa cific Plate for the purposes of the present study) endemics, and these three species were related to and the deep sea off the Hawaiian Islands an Indian Ocean species and a widely distributed (Tinker, 1978). Indo-Pacific species. If these relationships are cladistically supportable, the vicariant event that TOXOTIDAE initiated formation of the three Plate endemic species of the C. epilampra group has been ob- The archerfishes consist of a single genus, Tox- scured by dispersal of the widely distributed spe- otes, and six small (to 400 mm SL) species (Allen, cies or by extinction of Canthigaster populations 1978). The species occur in freshwater and brack- between the Indian Ocean and the Pacific Plate. ish mangrove habitats, and are restricted to the Indo-West Pacific, ranging from the east coast of India to the western margin of the Pacific Plate: Order TORPEDINIFORMES Palau (Koror; Herre, 1939), Admiralty, Solomon, The electric rays, or torpedos, comprise four and New Hebrides islands. Whitley (1927) re- families, ten genera (Compagno, 1973a), and ported Toxotes from Fiji, based on a listing in a about 35 species of small to large (about 200-2000 Shmeltz catalogue (number 5, 1874, reference not mm TL), benthic, viviparous fishestha t occur at available to me) of the Museum Godeffroy; how- depths of from less than 1 m to about 1100 m. ever, neither B.A. Carlson (pers. comm.), who The families are combined here for the sake of collected fishes while resident in Fiji, P.A. Ryan convenience (while it was easy to identify reports (1980 and pers. comm.), nor I collected Toxotes in of torpedos in the literature, it was often not Fiji. All of us believe it does not occur there.
FIGURE 44.—Distribution of the elasmobranch fishes of the order Torpediniformes. NUMBER 367 101
In the Pacific, the archerfishes range north only pagno, pers. comm.; P. Heemstra, in litt.) of to the Philippines and south to northern Queens- viviparous, deep or shallow-dwelling, small (to land and New Hebrides. One species, T. jaculator, about 1.5 m TL) sharks. The family is distributed extends over the entire coastal range of the genus. primarily in the vicinity of continental coasts, It is surprising, perhaps, that the archerfishes and is absent from the Pacific Plate nonmargin- have not invaded the Pacific Plate nonmargin- ally, as are many other elasmobranch groups. ally, particularly the Eastern Caroline Islands, Triakids are mostly deep-dwelling or antiequa- where the proper habitat appears to be present. torially distributed in the Indo-West Pacific, but Sterba (1962, fig. 1015) gave a generalized mainly shallow-dwelling, tropical or subtropical distribution map for the Toxotidae (reproduced along the continental coasts in other areas where in Nelson, 1974, map 38) and included the north- they occur. ern Indian Ocean coast from southern India to the southern Red Sea within the range of the family. I know of no records of toxotids west of TRICHIURIDAE the southern tip of India (see also Allen, 1978, The limits of the cutlassfish family (for in- map 1, for a reasonably complete distribution stance, whether it should be included with the map). Gempylidae or divided into more than one fam- ily) are uncertain. In Tucker's (1956) revision of TRIACANTHIDAE the Trichiuridae, two apparently closely related The triacanthids consist of four genera and monotypic genera, Trichiurus and Lepturacanthus, seven small (to 272 mm SL) species of benthic were recognized as composing the subfamily Tri- fishes (Tyler, 1968, 1970) that are restricted to chiurinae, and only this taxon, which is almost the Indo-West Pacific, ranging from the Persian circumglobally distributed, but is absent from the Gulf eastward to New Guinea, northward to Ja- Pacific Plate nonmarginally, is considered here. pan, and southward to New South Wales (Tyler, In the literature, Trichiurus and lepturacanthus, 1968, fig 3, gives a generalized distribution map which have similar habits and are sympatric over for the family). Fossil genera and species of tria- part of their ranges, are often not clearly distin- canthids are known from the Upper Eocene of guished by authors. It is possible, therefore, that Italy, Oligocene of Switzerland, and Miocene of some of the localities marked for Trichiurus Upturns Celebes, indicating a much broader distribution (Figure 45), especially those in the area from in the past. Tyler believes that the triacanthids India to Australia, may represent Lepturacanthus evolved from the deep-dwelling Triacanthodidae, savala. The addition of the distribution of L. savala which are found in the western Atlantic, Indo- to Figure 45 would not change the general limits West Pacific, and Hawaiian Islands on the Pacific of the distribution as given for T. Upturns. Mainly, Plate; hence, are also restricted to continental the addition would be noticeable only by the plates except for the Hawaiian occurrence of an presence of more spots along the coasts of Aus- endemic species of Hollardia, which genus is other- tralia (from Northern Territory to New South wise known only from the western Atlantic. If the Wales), India, and the South China Sea. The ancestor of the Triacanthidae and Triacanthodi- records of Trichiurus from Palau and Fiji (Figure dae was a deep-dwelling form, one can only 45) are based on reports by Abe (1939) and wonder why shallow-water representatives did Carlson (ms.). not also evolve in the Pacific Plate area. Trichiurus Upturns is a ribbonshaped, free-swim- ming, predatory species that has planktonic lar- vae and attains a maximum size of about 1.5 m TRIAKIDAE TL. It occurs along the continental shelf to depths The Triakidae comprise ten genera (Com- of up to about 100 m; it also enters shallow pagno, 1973b) and about 37 species (L.VJ. Com- estuaries. 102 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 45.—Distribution of the trichiurid fish Trichiurus Upturns (stars indicate unspecified localities in general areas; shaded area is generalized distribution).
TRICHONOTIDAE Leseurina is known only from southern Aus- The Trichonotidae is a poorly known family of tralia. small (to about 250 mm TL), sand-dwelling fishes that occur in marine and estaurine areas. The TRIGLIDAE family consists of two genera, Trichonotus, with at least three species (at least one undescribed), and The triglids are poorly known taxonomically. Leseurina, monotypic, which questionably belongs There is a question as to whether the family in the family. should include the Peristediidae, a usually deep- Trichonotus is distributed from the Red Sea and dwelling, benthic group of moderately small east coast of Africa to Japan, Philippines, Queens- fishes found in continental coastal waters round land, Guam, Palau, New Hebrides, and Fiji (Carl- the world, but only at the Hawaiian Islands on son, ms.), and is known from a single locality, the Pacific Plate nonmarginally. The nonperis- Enewetak, on the nonmarginal portions of the tediid triglids are small to moderately large (up Pacific Plate (J.E. Randall, in litt.). Its distribu- to 1 m TL), benthic forms, usually shallow dwell- tion on the Plate is probably more extensive than ing in marine and brackish water, and exhibit presently indicated. Munro (1958) reported Tri- much the same general distribution as the peris- chonotus from Port Moresby, Papua-New Guinea, tediids, with the following differences: they ap- and Jaquinot Bay, New Britain. Although Tri- parently occur farther north in the Atlantic and chonotus should be expected to occur at these are absent from the Pacific Plate nonmarginally, localities, the specimens, which were larvae except for a report of a single specimen of the (Munro, 1967), and upon which Munro's reports shallow-dwelling Cheilodonichthys kumu (as Trigla are based, leaves doubt as to their correct identi- kumu; Pietschmann, 1930), from Hawaii. Cheilo- fication. donichthys kumu is otherwise widely distributed in NUMBER 367 103 the Indo-West Pacific. The absence of other re- TRYPAUCHENIDAE ports of nonperistediid triglids from the Hawaiian Islands leads me to question the validity of The trypauchenids comprise about four genera Pietschmann's report. and fewer than ten species of small (to about 220 mm TL), burrowing fishes that are found in muddy bottoms in salt and brackish waters from TRIODONTIDAE near shore to depths of about 100 m. The family is limited to the Indo-West Pacific, from the east The Triodontidae consists of a single genus and coast of Africa (Smith, 1959) to New Guinea species of small fish (to about 480 mm SL), Triodon (Munro, 1958, as Ctenotrypauchen under Taenioi- macropterus, which is questionably included here dae), on the margin of the Pacific Plate. In the as a shallow-dwelling form. According to Tyler western Pacific they occur as far south as Queens- (1967), the only depth records available for the land and as far north as Japan. species are 150 m and 200-300 m. Triodon ranges from the east coast of Africa eastward to Guam, Volcano, Fiji, and Samoan islands (Tyler, 1967; URANOSCOPIDAE Kami et al., 1968; Carlson, ms.; Wass, ms.); the The stargazers comprise about ten genera and last four localities are on the western margin of 30 species of small- to moderate-sized fishes (up the Pacific Plate. Although Triodon is generally to about 750 mm TL). As juveniles and adults considered to be a rare species, and few specimens the species are benthic in habit, burrowing into are available in museum collections, Fourmanoir sand and mud. They occur at depths varying and Laboute (1976) reported that they obtained from less than a meter down to about 550 m. 20 specimens from the stomachs of tunas caught Most species are marine, but some occur in brack- in the vicinity of New Caledonia and the New ish estuaries. Haast (1873) reported specimens Hebrides. obtained in freshwater as much as 40 miles (67 km) upstream from river mouths, and others buried in sand well above low-tide mark along TRIPTERYGIIDAE the New Zealand coast. Little is known about their life history, but the species are reported to The triplefins are a very poorly known family have planktonic larvae (Pearson, 1941; Fierstine taxonomically, with as many as one-third of its and Werner, 1963). estimated 20 genera and 150 species as yet un- described. The family is circumglobally distrib- The uranoscopids are restricted almost entirely uted in tropical and warm-temperate waters, and to continental plate coastlines, and occur only the species are predominantly benthic inhabit- marginally on the Pacific Plate (Figure 46). Bloch ants of rocky and coral-reef areas. The species are and Schneider (1801) reported a uranoscopid all small, ranging in maximum size from about from Tahiti based on information in an unpub- 15 to 200 mm SL (the largest species is in the lished manuscript by Forster. The same species, New Zealand genus Notoclinus), although most which occurs in New Zealand, was also reported species attain less than 75 mm. A revision of the by Bloch and Schneider from New Zealand. I southeastern Atlantic and Indo-Pacific genus Hel- consider the Tahitian record to be improbable cogramma by P.H. Hansen is nearing completion; and erroneous as did McCulloch (1929). there appears to be one widely distributed Pacific Plate endemic species among its approximately 12 included species. Probably nine other tripter- UROLOPHIDAE ygiid species are present nonmarginally on the The urolophids comprise two genera and 21 Pacific Plate. species (Chirichigno and McEachran, 1979) of 104 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 46.—Distribution of the fishes of the family Uranoscopidae (star represents unspecified locality in Black Sea).
benthic, viviparous rays of moderate to very large the sea, from near shore to depths of at least 355 size (to about 2.7 m TL and 119 kg). These rays m. As adults they are burrowers, but may come are found in the eastern Pacific (five species), to the surface (to spawn?). Their larvae are plank- western Atlantic (one species), and Indo-Pacific tonic. (15 species) on muddy or sandy bottoms, often in The family is probably circumglobally distrib- shallow waters, but are unknown in the eastern uted. The deep-dwelling forms tend to be widely Atlantic. Urotrygon daviesi is the only urolophid distributed with wide gaps separating collections; known from the Pacific Plate nonmarginally. It for example, Chlopsis dentatus (from about 126-355 is recorded from depths of 144-1800 feet (44-549 m) is known from the western Indian Ocean and m) and reported only from South Africa (Wal- Caribbean, and Robinsia catherinae (52-243 m) lace, 1967), the Bay of Bengal (Nair and Soun- from off Kenya and the Atlantic coast of Panama. dararajan, 1973), and Hawaii (Tinker, 1978). As One shallow-dwelling species, Kaupichthys hyopro- is true of most elasmobranch groups, the urolo- roides, occurs widely in the Caribbean and Indo- phids are essentially absent from the Pacific Plate. Pacific, but is absent from the eastern Pacific and eastern Atlantic. While some of the Indo-Pacific
XENOCONGRIDAE species are known only from the Indian Ocean, others are widely distributed. The monotypic ge- The xenocongrid eels consist of seven genera nus Powellichthys is known from a single specimen and 15 species (Bohlke and Smith, 1968), but (ripe female) taken at the surface over a depth of there are large unstudied collections available 500 fathoms (914 m; Smith, 1966b) off Cook that probably include several undescribed species Island (= Tarawa, Gilbert Islands) and could be (J.E. Bohlke, pers. comm.). These eels are all a deep-dwelling endemic on the Pacific Plate; small (largest species, 436 mm TL, but most however, I believe that the available information species not attaining 200 mm), and occur only in of xenocongrid species and their distributions is NUMBER 367 105 much too meager to be accepted as definitive. meaning to be inferred from my arrangement of Perhaps five species of xenocongrid eels are pres- the phyla, or the taxa within a phylum. ent nonmarginally on the Pacific Plate. Phylum MOLLUSCA ZANCLIDAE Class GASTROPODA The Zanclidae consist of a single genus and species, Zanclus cornutus, the Moorish idol. Zanclus Abbott (1960) revised the Indo-Pacific species is a free-swimming, coral-reef species that attains of the conch genus Strombus (Strombidae), which a maximum size of 250 mm TL. It is distributed is distributed circumglobally in the shallow por- throughout the tropical portions of the Indo-Pa- tions of warm seas; its larvae are planktonic. cific, from the east coast of Africa to at least the Strombus comprises 51 extant species and 60 spe- Tuamotus, and is also present in the eastern cies-group taxa (including subspecies; I have Pacific. made no attempt to determine if additional new taxa of Strombus have been described since Ab- Other Organisms bott's revision). Forty-nine species-group taxa of Strombus occur in the Indo-Pacific, and of these, The discussions of the taxa that follow are 12 have the eastern limits of their distributions treated in the same manner as those for fishes. I along the western margin of the Pacific Plate (see made no attempt to be inclusive, as I did for the Figure 47 for the distribution of one of these, S. shore fishes. The nonfish taxa were selected to labiatus), 13 range from the Indo-West Pacific well demonstrate that distribution patterns commonly out onto the Plate, and five are endemic to the exhibited by the fishes are also exhibited by a Plate, including its margins. Strombus helli and S. wide variety of other organisms. There is no vomer hawaiiensis occur only in the Hawaiian Is-
FIGURE 47.—Distributions of two species of the marine mollusk genus Strombus (Gastropoda), compiled from the literature and museum collections (stars = S. labiatus, closed circles = 5. maculatus). 106 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
lands (the other two subspecies of & vomer occur below the intertidal zone to high above the sea, as follows: S. v. vomer, Ryukyus and New Cale- where they are wet only occasionally by ocean donia; S. v. iredalei, western and northern Aus- spray (Rosewater, 1970). Certain species are able tralia; hence, the subspecies of S. vomer do not to survive for long periods without immersion in cross the margin of the Pacific Plate). Strombus seawater. Most of the species release pelagic egg variabilis athenius occurs in the Marshalls, Eastern capsules and only a few deposit egg masses on the Carolines, northwestern New Guinea, Samoa, substrate or are ovoviviparous. In the Indo-Pa- Fiji, and New Caledonia. The nominal, and only cific, only one species, Littorina scabra (subspecies other, subspecies of S. variabilis ranges from Su- scabra), is ovoviviparous. It has by far the broadest matra to Palau, Bougainville, and northern Aus- distribution of all the species in the genus in the tralia; hence, one subspecies of 5. variabilis ranges Indo-Pacific (from the east coast of Africa to the from the western Pacific to the western margin of Gambier Group of islands), between 28°N-S lat- the Pacific Plate, and the other subspecies ranges itudes (Rosewater, 1970, plate [= figure] 354), from the western margin of the Pacific Plate out but is notably absent from the Marquesas (Ro- onto the Plate. sewater recognizes three subspecies in S. scabra: S. Strombus maculatus is a widely distributed Pacific s. scabra, Indo-Pacific; S. s. aberrans, eastern Pa- Plate endemic (Figure 47). Abbott stated that S. cific; S. s. angulifera, eastern and western Atlantic). maculatus, S. microurceus, and S. mutabilis form a The subfamily Tectariinae (Littorinidae) con- natural group within the subgenus Canarium. tains a single genus, Tectarius, and three subgenera These three species have quite different distribu- (Rosewater, 1972). The subgenus Tectarius con- tions. Strombus microurceus ranges through much of tains five extant species and one fossil species the western Pacific, but occurs nonmarginally on (from Java). Of the extant species, two are re- the Pacific Plate only in the Eastern Caroline stricted to the Pacific west of the Pacific Plate Islands, where it is sympatric with S. maculatus margin, two have their eastern limits along the (Abbott, 1960, plate [= figure] 47). The presence western margin of the Pacific Plate, and one, T. of S. microurceus in the Eastern Carolines is evi- grandinatus, is a widely distributed Pacific Plate dence favoring operation of the Caroline Islands endemic, ranging from the Tonga Islands to the conduit. Strombus mutabilis ranges from the Red Gambier Group islands in the Tuamotus. The Sea and east coast of Africa to the Tuamotus, closest relative of T. grandinatus is, then, a conti- and thus overlaps most of the range of the other nental form, which Rosewater believes is T. rugo- two species (it is absent from the Hawaiian Is- sus, confined to the Philippines and Celebes, thus, lands; Abbott, 1960, plate 50). If S. microurceus is not quite reaching to the western margin of the the sister species of S. maculatus, and there is a Pacific Plate. good chance that it is, the Pacific Plate boundary Emerson and Cernohorsky (1973) revised the is probably the area where the vicariant event genus Drupa (Thaisidae) in the Indo-Pacific. occurred that divided the ancestral population Among the species, one, Drupa elegans, is a widely that gave rise to these two species. Subsequent distributed Pacific Plate endemic known from the dispersal would then have resulted in the partial following islands: Wake, Enewetak, Bikini, Flint, sympatry of these two species in the Eastern Caroline (Line Islands), Niue, Akamaru, Mani- Caroline Islands. The distributions of the three hiki, Marutea, Vahitahi, Nengonengo, Anau, Na- species of the subgenus Canarium closely parallel puka, Taenga, Fakahina, Gambier, Pitcairn. The the distributions of the three species of the gobiid sister species was not indicated. fish genus Nemateleotris (Figure 21). Cernohorsky (1976) revised Mitra (Mitridae). The members of the family Littorinidae are Among the species, Mitra testacea is a widely dis- found in most coastal regions of the world, and tributed Pacific Plate endemic, known from the occupy habitats from relatively shallow waters following islands: Kingsmill Group of the Gil- NUMBER 367 107 berts, Palmerston, Rarotonga, Tahiti, Anaa, Gar- tropods are also duplicated by terrestrial gastro- umaoa, Raroia, Aukena, Pitcairn. The sister spe- pods. The family Endodontidae comprises about cies was not indicated. 475 species of minute land snails (Solem, 1976; Rehder (1980) reported on the mollusks of Solem's delimitation of the family is not accepted Easter Island and provided overall distributions universally: see Sirgel, 1980). The endodontids of each species. The following gastropod species are restricted to the islands on the Pacific Plate are widely distributed Pacific Plate endemics and its margins (Figure 48). Solem (1981) states (families not indicated): Cerithium rubus (Line, To- that the Charopidae are the closest related family kelau, Cook, Samoan, Palmerston, and Easter to the Endontidae, and the charopids most inter- islands); Dendropoma platypus (Marshall, Ha- mediate [closely related to?] between the Charo- waiian, and Easter islands); Planaxis fasciatus (Ni- pidae and Endodontidae occur in southwestern uafoou and Samoan islands east to Easter Island); Australia; hence, somewhat west of the margin of Balcis aciculata (Hawaiian, Cook, Tuamotu, and the Pacific Plate. The charopids are distributed Easter islands); Natica ochrostigmata (Hawaiian, in North and South America, St. Helena, portions Cook, Samoan, and Easter islands); Trimusculus of southeast Africa, Australia, New Zealand, and odhneri (Micronesia, Polynesia, Niuafoou). the Indo-Malayan area as far north as the Phil- Many marine gastropods have Indo-West Pa- ippines (Solem, 1981, fig. 5.5), all localities where cific distributions with their eastern distribution endodontids do not occur. The charopids also limits along the western margin of the Pacific occur in several Pacific Plate island groups where Plate. One need only to leaf through the issues of endodontids are also known: Cook, Ellice, Samoa Indo-Pacific Mollusca to find them. (marginal), Tonga (marginal), Palau (marginal). Distribution patterns exhibited by marine gas- Solem (1976) reported that the Caroline, Mar-
FICURE 48.—Distribution of the land snail family Endodontidae (Mollusca: Gastropoda), compiled from Solem, 1976 (stars indicate localities where only fossils are known; the genera represented by fossils are extant on other islands). 108 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
iana, and main Fiji islands have charopids, but restricted in distribution and occur in the Indo- no endodontids, and (as is also true for the en- West Pacific (Voss, 1977). The species are small, dodontids) occur only as fossils at the Marshall the largest attaining a maximum total length of Islands. If the southwestern Australian charopids about 600 mm (G.L. Voss, pers. comm.). The are the sister group of the endodontids, the partial species are primarily benthic, but occur in a wide sympatry that presently exists with these two variety of habitats ranging from the open sea families would indicate that dispersal occurred surface down to depths of, perhaps, 1000 m subsequent to the evolutionary divergence of the (Adam and Rees, 1966; Roeleveld, 1972), along two families (Springer 1981a). shore, intertidally, and on coral reefs. At least one species occurs in shallow estuaries. The species Class PELECYPODA lay encapsulated eggs, which they attach to al- most anything on the substrate, including sea- Rosewater (1961) revised the Pinnidae of the weeds and phanerogams. The oldest known fossils Indo-Pacific. The distribution of one species, of Sepia are from the Eocene of continental areas. Pinna bicolor, is of interest here. Pinna bicolor ranges Only one species of Sepia has been obtained from the Red Sea and east coast of Africa east- other than marginally from a Pacific Plate local- wards to Japan, The Philippines, New Guinea, ity: Arno Atoll, Marshall Islands (Figure 49; Arno and New Caledonia (all on or west of the western specimens in collection of Division of Mollusks, margin of the Pacific Plate). Pinna bicolor also is NMNH). The identification of this species is un- present on the Pacific Plate nonmarginally, but certain. Sepia is obviously a continental plate only in the Hawaiian Islands. Rosewater genus and its exceptional occurrence on the Pa- (1961:195) stated that "the sporadic appearance cific Plate only at Arno Atoll is unexpected. Sepia of P. bicolor in Hawaii is probably the result of is often obtained in rotenone collections of fishes chance introduction." Rosewater was unaware, and its absence from Ponape, Eastern Caroline understandably, that an Hawaiian exception type Islands, and Bikini, Marshall Islands, both of of distribution is a common occurrence among which have been intensively collected with rote- marine organisms, and there is no need to rely on none emphasizes the unusual nature of its pres- chance introductions to account for such distri- ence at Arno. I have no explanation for its occur- butions. It is possible that the presence of P. bicolor rence at Arno and encourage others to be alert in the Hawaiian Islands is the result of dispersal for its possible occurrence elsewhere on the Pacific via the Kuroshio extension, which may also ac- Plate. count for numerous fish species common to Japan and only the Hawaiian Islands on the Pacific The living Nautilidae were revised by Saunders Plate. (1981), who recognizes a single genus and four species. The distribution of Nautilus, based on live Rehder (1980) listed three widely distributed specimens, includes the region from the Andaman Pacific Plate endemic species of pelecypods (fam- Islands east to Japan, south to the Philippines, ilies not indicated here): Malleus maculosus (Micro- through Palau, to southern Australia, and east to nesia, Tuamotus, and Easter islands); Lasaea ha- Fiji; hence, absent all but marginally from the waiensis (Hawaiian, Gambier Group, Easter, and Pacific Plate (Saunders, 1981, fig. 1). Nautilus is Rapa islands); Semele australis (Cook Island north moderately deep dwelling, and Saunders allows to Hawaiian Islands and east to Easter Island, that its distribution may be more extensive than with a single valve each from Bikini, Marshall presently indicated. Additionally, fresh, drifted Islands, and Lifu, Loyalty Islands). shells have been obtained from Majuro, Marshall Islands, and Kosrae, Eastern Caroline Islands, Class CEPHALOPODA localites upcurrent from potential sources of drift There are about 100 species in the cuttlefish shells (drift shells have been taken as far west as genus Sepia (Sepiidae), most of which are highly the coast of Africa; I am unaware of drift shells NUMBER 367 109
FIGURE 49.—Generalized distribution of the cuttlefish genus Sepia (Mollusca: Cephalopoda), compiled from Adam and Rees, 1966, Lubet, 1973, and G.L. Voss, in litt.
having been taken at other Pacific Plate localities, Its nearest relative, based on a shared specializa- with the exception of the Samoan Islands, cited tion (synapomorphy: presence of a dorsal spine by Toriyama et al., 1965, supposedly based on on the constricted anterior portion of the cara- literature, which Saunders, apparently did not pace) is L. valida (Williams, pers. comm.), which accept). The presence of Nautilus in the Eastern is restricted to the Indo-West Pacific, ranging Carolines, if verified, would be evidence favoring from the southeast coast of Africa to Japan, the operation of the Caroline Islands conduit. Majuro Philippines, and eastern Indonesia (two of the is close to, and down current from the Carolines, other three Indo-West Pacific species of Latreillia so that Nautilus drifts from the Carolines might have similar eastern distributional limits). A vi- be expected. cariant event occurring along the western margin of the Philippine and/or Pacific plates could have divided a widely distributed population ancestral Phylum ARTHROPODA to these two species, permitting speciation of the two populations, and resulting in the distribution Class CRUSTACEA patterns presently manifested. The crab genus Latreillia (Latreillidae) has been Banner and Banner (1975, 1981) reported on revised by A.B. Williams (in press). Latreillia con- the pistol shrimp Alpheus lobidens (Alpheidae). tains six small (maximum carapace length about This is a small (to 50 mm long), burrowing species 20 mm), benthic species that occur at depths of consisting of two subspecies: A. lobidens lobidens, 35-474 m. The genus is broadly distributed, but which ranges from the Mediterranean and Red noticeably absent from the eastern Pacific (Figure Sea to southern Japan, Yap, New South Wales, 50). One species. L. mrtanxa, is a widely distrib- and Lord Howe Island, and A. I. polynescia, which uted Pacific Plate endemic, and the only one of ranges widely on the Pacific Plate and its western the four Indo-Pacific species to occur on the Plate. margin (Figure 51). Banner and Banner (1975) 110 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
* ELEGANS
BMANNINei
• HETANESA
• species near HANNINGI
OPENNIFERA • VALIDA
FIGURE 50.—Distributions of the species of the crab genus Latreillia (Arthropoda: Crustacea), after Williams (in press).
FIGURE 51.—Distributions of the two subspecies of the alpheid shrimp Alphetis lobidens (Arthro- poda: Crustacea), after Banner and Banner, 1975, 1981 (closed circles = A. I. lobidens; stars = A. I. polyntsica). NUMBER 367 111
FIGURE 52.—Distribution of the approximately 75 benthic species of the ostracode family Cypridinidae (Arthropoda: Crustacea), based on data provided by L.S. Kornicker; records for Antarctic Plate are incomplete. were unable to identify their Philippine speci- other than marginally on the Pacific Plate (on mens, which were too small to exhibit the differ- the Chatham Rise, which is part of the continen- entiating character for the subspecies, but the tal shelf of New Zealand, hence excluded from nominal subspecies would most probably be ex- the Pacific Plate for my purposes; Figure 52). Of pected in the Philippines. While there is an area the planktonic genera, three (Codonocera, Melavar- between the distributions of the two subspecies gula, Monopia), with a total of 13 species, are from which no specimens of A. lobidens are avail- restricted to the Indo-West Pacific. Of the other able, the separation between the two subspecies three planktonic genera (Cypridina, Gigantocypris, appears to be delineated by the western margin Macrocypridina), nine of the ten included species of the Pacific Plate and the result of a scenario have Indian and/or Pacific ocean distributions, similar to that proposed above for Latreillia. but only five species occur on the Plate nonmar- Recent ostracods in the suborder Myodoco- ginally. In addition to the Cypridinidae, there are pina, especially the benthic species, generally tend four other families of rnyodocopid ostracodes, all to have localized distributions, and are generally primarily benthic. They comprise about 33 gen- absent from the Pacific Plate nonmarginally. era and more than 200 species, of which only two There are about 20 genera and 100 species in the species are known to occur other than marginally family Cypridinidae. Fourteen of these genera, on the Pacific Plate. For example, the family including about 75 species, are primarily benthic Cylindroleberididae comprises three subfamilies, inhabitants. The remaining genera and species in of which one contains both deep- and shallow- the family are planktonic. Of the 75 benthic dwelling species and is absent from the Pacific species, 40 occur in the Indo-Pacific, but only one Plate. The other two subfamilies (comprising of these (Hadacypridina bruuni) is known to occuabour t 60 species) are essentially restricted to 112 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 53.—Distributions of two of the three subfamilies of the ostracode family Cylindrole- berididae, based on data provided by L.S. Kornicker (closed circles = Asteropteroninae; stars = Cyclasteropinae). depths under 100 m. One of these last two There are about 300 species of mantis shrimps, subfamilies is absent from the Plate nonmargin- Stomatopoda, which as far as known, have plank- ally and the other is present on the Plate non- tonic larvae (R.B. Manning, pers. comm.). The marginally, but only at the Hawaiian Islands genus Harpiosquilla comprises some of the largest (Hawaiian exception; Figure 53). While the evi- species of mantis shrimps (up to 335 mm TL). dence tends to support a continental plate type Harpiosquilla, with seven species, is a benthic in- distribution for the Myodocopina, L.S. Kor- dweller of soft bottom in the tropical marine and nicker, who provided all my information on the brackish waters of the Indo-West Pacific (Figure Myodocopina, believes that a cautionary note is 54) from depths of less than 2 m to 206 m necessary: there have been few studies of myodo- (Manning, 1969). Manning (1969) wrote that the copid ostracodes in Pacific Plate areas. In contrast absence of Harpiosquilla from Oceania reflects a to Kornicker, I believe that the present data lack of suitable habitats. I doubt this explanation, provide a good indication of the overall distri- as shallow, soft bottoms in brackish areas exist at bution of the known benthic species. There is localities both on and off the Pacific Plate (New evidence that the other ostracode suborder, Po- Guinea, New Caledonia, Fiji, Palau, Eastern Car- docopina, may also contain taxa that duplicate oline Islands, Society Islands, etc.) where Harpios- distribution patterns found in the Myodocopina; quilla is not known. While the absence of Harpios- for instance, the Tertiary to Recent distribution quilla from these localities may be due in part to of the marine genus Caudites, whose distribution collecting deficiencies, I do not believe either was mapped by McKensie (1967, fig. 5). Caudites collecting deficiencies or lack of suitable habitat is distributed throughout much of the world's provide a satisfactory explanation for the absence warm seas, but is absent from the nonmarginal of Harpiosquilla from the Pacific Plate. There is a portions of the Pacific Plate in the Indo-Pacific. remarkable similarity between the distribution of NUMBER 367 113
FIGURE 54.—Distribution of the mantis shrimp genus Harpiosquilla (Arthropoda: Crustacea), based on Manning, 1969.
Harpiosquilla and the distributions of the fisheso f though most species are winged, there are no the family Congrogadidae (reef inhabiting; Fig- published flight records, and it is believed that if ure 17), the Indo-West Pacific distributions of the they fly at all, it is only as a means to reach toadfish family Batrachoididae (reef and estuary decayed areas in the upper branches of trees; inhabiting; Figure 6), and the shark genera Lox- dispersal by human transport of wood, or by odon and Rhizoprionodon (both free swimming; Fig-rafting logs seems feasible (Bell, 1979). Only three ure 13). The similar distribution of these different species of rhysodines occur other than marginally taxonomic groups (some with habits and habitat on the Pacific Plate, all in the Eastern Caroline preferences similar to, and others with habits and Islands. Two of the three species are endemic to habitat preferences dissimilar to, those of Harpios- the Plate and the third has been reported other- quilla) indicates to me that these distributions are wise only from Palau, based on a possibly misla- part of a common pattern involving groups of belled, old specimen. Three other species of rhy- continental origin that have been unable to dis- sodines are restricted to Palau, Yap, and Ulithi, perse onto the Pacific Plate. islands along the eastern margin of the Philippine Plate. There are no other species reported from these islands. Of the six species included in the Class INSECTA present discussion, three are found on more than There are about 300 species of rhysodine bee- one island (up to four islands), and it is believed tles (Carabidae; Rhysodini), all of which live in that each of the three originally occupied only dead wood, both as larvae and adults, and are one island, with human associated introductions distributed throughout much of the tropical and accounting for the spread to other islands (R.T. warm-temperate portions of the world (Figure Bell, in litt. and pers. comm.). The Pacific Plate 55). The species are gregarious, and up to 50 distribution of the Rysodini appears to be an individuals have been found in a single log. Al- example of the operation of the Caroline Islands 114 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 55.—Generalized distribution of the carabid beetles of the tribe Rhysodini (Arthropoda: Insecta), based on information provided by R.T. Bell.
FIGURE 56.—Generalized distribution of the bats (Chordata: Vertebrata), based on information provided by K.F. Koopman (closed circles represent distribution of Pteropus tonganus, mostly after Wodzicki and Felten, 1981). NUMBER 367 115 conduit, which has similarly affected the distri- PLANTS butions of numerous other terrestrial, and marine, organisms. Data on plant distributions were more difficult for me to obtain and interpret than similar data on animals. Although distributions parallel to Phylum CHORDATA those described for animals exist, in some cases commonly, I have selected only a few to illustrate Class MAMMALIA the parallelisms, and most of these are from the pages of the volumes of Pacific Plant Areas. With the possible exception of rodents, bats Van Steenis and van Balgooy (1966) mapped (order Chiroptera) are the most widely distrib- the distribution of Korthalsella platycaula (Loran- uted group of mammals (Figure 56). Most bat thaceae), which is a widely distributed Pacific species are highly localized and, unlike rodents, Plate endemic that ranges from Samoa east to have not been subject to widespread introduc- Henderson Island, and north to the Hawaiian tions by humans (Koopman, 1970). There are Islands. The species is parasitic on woody dico- 169 genera and about 847 species of recent bats tyledonous trees and shrubs, mostly on high is- (Koopman and Knox Jones, 1970). Only three lands. The closest relative was not indicated. Van genera (Emballonura, Pteropus, Lasiurus) and seven Balgooy (1975) mapped the distribution of Hedy- species occur other than marginally on the Pacific otis romanzoffiensis (Rubiaceae), also a widely dis- Plate. Lasiurus cinereus is the only bat on the tributed Pacific Plate endemic, which is a low Hawaiian Islands, and the only bat species on the shrub restricted to coral islands. The closest rela- Pacific Plate (nonmarginally) that has its closest tive was not indicated. relationships in the Western Hemisphere. Lasiurus Van Balgooy (1975) mapped the distribution cinereus is widely distributed in the Western Hem- of Gunnera (Haloragaceae), which contains 30-50 isphere, where Lasiurus (with six species) is other- species of perennial, herbaceous plants up to 6 m wise restricted. Emballonura and Pteropus are re- in height that, in the tropics, occur only in mon- stricted to the Indo-Pacific area, and five of the tane areas. Gunnera is widely distributed in conti- six nonmarginal Plate species in these genera nental plate areas, but on the Pacific Plate non- occur on various of the Eastern Caroline Islands. marginally, it occurs only in the Hawaiian Islands Of the five species, four are endemic to the East- (Hawaiian exception distribution). ern Caroline Islands; the fifth also occurs to the The eastern portion of the distribution of Rhi- east of the Pacific Plate margin. The presence of zophora apiculata (Rhizophoraceae) was mapped bats in the Eastern Caroline Islands is evidence by van Steenis (1963). This mangrove attains a favoring support of the Caroline Islands conduit. height up to 30 m, occurs in muddy marine areas, The seventh Plate bat species, Pteropus tonganus, and has buoyant seeds. The eastern limits of the comprises two subspecies, which occur only on distribution of R. apiculata is generally defined by islands on and near the Plate margin (Figure 56). the western margin of the Pacific Plate between While the bat distribution map is not as "clean" the Marianas and Santa Cruz islands, and the as the other distribution maps given in my study, species is absent from the Pacific Plate nonmar- I believe that it basically conforms to a continen- ginally except for its occurrence in the Eastern tally restricted pattern, with indications of recent Caroline Islands, evidence favoring the operation dispersals and speciations in Pacific Plate areas of the Caroline Islands conduit. adjacent to the western margin of the Plate. The order Coniferales includes about 50 extant (Much of the information presented in this dis- genera and about 600 extant species of typically cussion was provided by K.F. Koopman, in litt. woody and evergreen plants, such as pines, and pers. comm.) spruces, hemlocks, firs, cypresses, junipers, se- 116 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY quoias, podocarps, araucarias, and larches. The ias do (F.R. Fosberg, pers. comm.). species vary from only 8 cm to 90 m in height at Van Balgooy (1971:143), who discussed the maturity. They occur in habitats ranging from relationship of dispersibility to plant distribution sea-level swamps to dry, rocky, mountain sides at was hard put to explain the absence of conifers 3400 m altitude. Species are found in the tropics (and other plants of low dispersibility) from Pa- as well as at the timberline within the Arctic cific islands east of the Andesite Line (essentially circle, where temperatures drop to —45° C. In the the same as the western margin of the Pacific warmer parts of their range, conifers usually pre- Plate; Figure 2). He noted the presence of conifers fer higher altitudes (Wiggins, 1974). on islands along and west of the Andesite Line The conifers first appear in the fossil record that were no more isolated from each other than during the Carboniferous of the Paleozoic, but they were from islands lacking conifers to the east the earliest known contemporary genera are from of the Line. In referring to these nonconifer-in- the middle Jurassic, with few contemporary gen- habited islands, van Balgooy (1971:146) stated, era known earlier than the Cretaceous (Miller, "The floristic character of [these islands] pleads 1978). for rather difficult accessibility. Yet under the Conifers are frequently reported to occur na- tively throughout the world, but it is obvious present day configuration of land and sea the from their distribution (Figure 57) that they are facts found are also difficult to explain. They can absent from the nonmarginal portions of the Pa- best be explained by archipelagic conditions." I cific Plate. Conifers have been introduced into am unable to determine from van Balgooy's study various Plate islands, particularly the Hawaiian what he was implying by archipelagic conditions Islands, mostly as ornamentals. Pines do not (perhaps he was ascribing unique ecological con- adapt well in the Hawaiian Islands, but araucar- ditions to archipelagos), but his understandable
FIGURE 57.—Combined generalized distribution of fossil and recent plants of the order Coni- ferales, based on Florin, 1963; there are some northern distributional records for areas not shown on map. NUMBER 367 117 loss at explanation pervades this portion of his and are bound to continental plates simply be- study. I find the similarities exhibited by the cause they have low dispersal abilities. Their distributions of the terrestrial conifers and the distributions are the result of fragmentations and marine rajoid rays (Figure 36), in so far as they movements of the continental lithosphere to can be compared, given the general habitat re- which they are attached; ecological factors affect quirements of the two groups, to be striking. only relatively minor areas within the overall These two groups are of continental plate origin ranges of their distributions.
Discussion
Numbers of Shorefish Families stricted in their distribution on the Pacific Plate: Moridae, Hawaiian and Rapa islands; Labracog- The warm, shallow-water fish fauna of the lossidae, Rapa and Easter islands; Cheilodactyli- Indo-Pacific is summarized in Appendix 2, Tables dae, Hawaiian, Rapa, and Easter islands; Isoni- A and B, and has been discussed under the family dae, Samoa, Hawaiian and Pitcairn islands. It is accounts. There are about 179 fish families (three significant, perhaps, that most of the shorefish orders of elasmobranchs are treated as families families that have limited distribution on the for convenience) with warm, shallow-water rep- Pacific Plate (including those occurring only at resentation in the Indo-Pacific (henceforth, these the Hawaiian or Eastern Caroline islands) are families will be termed shorefish families). Of confined to high islands. On the other hand, high these shorefish families, only 121 (68%) have non- island groups (Society, Cook, Marquesas, for in- marginal representation on the Pacific Plate, and stance) do not necessarily have exceptional shore- 15(12%) of the 121 have nonmarginal represen- fish family representation. tation only in the Hawaiian islands, 6 (5%) only The greatest number of shorefish families pres- in the Eastern Caroline Islands, and 3 (2%) only ent in any general area in the Indo-Pacific is in the Marshall Islands. Ten of the 15 families about 163 (Appendix 2, Tables A, B; Figure 58), that occur on the Plate only in the Hawaiian or about 90 percent of all such families included Islands, occur there only in deep water, effectively here; this occurs along the Queensland coast. The reducing the number of families with shorefish diversity of shorefish families in Queensland is representation on the Plate to 111 (or 62% of the partly explained by the presence in northernmost total number of shorefish families). All of the Queensland of some families (for instance, Noto- Pacific Plate shorefish families are widely distrib- graptidae, Kurtidae, Rhinoprenidae, Menidae, uted in the Indo-West Pacific, if not even more Lactariidae, etc.) that are otherwise restricted to widely. In contrast, several of the Indo-West Pa- areas of the western Pacific north of Queensland, cific shorefish families have highly restricted dis- and the presence in southernmost Queensland of tributions; for instance, Coracinidae, only in the some families (for instance, Scorpididae, Gono- western Indian Ocean; Pataecidae and Brachae- rynchidae, Enoplossidae, Glaucosomatidae, etc.) luridae, only in Australia; Leptobramidae, No- that are otherwise restricted to areas of Australia tograptidae, Rhinoprenidae, only in Australia south of Queensland (some of these last-named and New Guinea; Pseudotrichonotidae, only in families are antiequatorially distributed). Any southern Japan. A few shorefish families, aside Indo-West Pacific area, except for a few margin- from those limited on the Pacific Plate to its ally located islands, has more shorefish families margins or to the Eastern Caroline, Marshall, than the number (121) of such families that occur and Hawaiian islands, appear to be highly re- on the entire Pacific Plate nonmarginally. 118 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 58.—Numbers of shorefish families known and expected to occur in various areas in the Indo-Pacific, based on information in Appendix 2, Table B.
Closely similar to Queensland in numbers of ings, based on numbers of shorefish families, tend shorefish families present are the southeast coast to agree with those of Cohen's. Areas such as of Africa, with about 161 families, the Philippine Queensland and southeast Africa are consider- Islands, with about 158, and the Indo-Malayan ably removed from each other, yet have about region, with about 157. Although Queensland the same numbers of shorefish families, and have and southeast Africa have almost identical num- slightly more than the Philippine or Indo-Ma- bers of families, they do not share all families. layan regions, and only two or three fewer families Fourteen of the Queensland families do not occur than the combined Indo-Malayan and Philippine (or are not expected to occur) in southeast Africa, regions. While it is presently not feasible to deter- and 13 of the southeast African families do not mine if the distribution of numbers of families is occur (or are not expected to occur) in Queens- paralleled by numbers of genera and species, I land. would not be surprised to find that it is for genera, Some portion of the central Indo-West Pacific but is not for species. region is almost invariably considered by zoo- The problem is complicated by the subjective geographers to be the center of greatest taxo- choice of how large an area one selects to consti- nomic diversity, with a progressive decrease in tute the central Indo-West Pacific region. Abbott numbers of taxa occurring in Indo-Pacific areas (1960, pi. 19; reproduced here as Figure 59) pre- outside the region. Talbot (1970), Goldman and sented a distribution map for the numbers of Talbot (1976), and Sale (1980), all of whom Indo-Pacific species of the gastropod genus Strom- discussed numbers of fish species, believed that bus. Abbott's delimited areas are somewhat com- the decrease emanated in all directions. Cohen parable to mine. His findings, however, contrast (1973), however, based on a study of selected fish with mine—particularly for the southeast African groups, found no decrease in numbers between coast and Queensland, which have many fewer the eastern and western Indian Ocean. My find- species of Strombus than do the Indo-Malayan, NUMBER 367 119
FIGURE 59.—Numbers of taxa of the gastropod genus Strombus at various Indo-Pacific localities; total number of taxa in area, 60 (after Abbott, 1960, pi. 19).
Philippine, or New Guinea regions—and, thus, 10 percent. If the estimates are reasonably ap- support generally held opinions about relative proximate, the five most speciose families (Gobi- diversity of taxa in a central Indo-West Pacific idae, Labridae, Serranidae, Pomacentridae, Blen- area. Most attempts to explain species diversity niidae) together contain about 29 percent (136) in the Indo-West Pacific have been couched in of all shorefish genera and about 36 percent (466) ecological terms (warm temperatures, climatic of all shorefish species on the Plate nonmargin- stability), but I must agree with Sale (1980) who ally. If the genera and species of the next six most stated that the high diversity of the central Indo- speciose families (Apogonidae, Chaetodontidae, West Pacific may have historical causes that are Ophichthidae, Acanthuridae, Muraenidae, Scor- outside the scope of ecological investigation. Tax- paenidae—the last two families tied for tenth onomic diversity is to a large degree dependent place) are added, these figures are increased to on the number of vicariant events that have about 42 percent (194) of the genera and about occurred in an area, and there have probably 54 percent (705) of the species. Thus, 10 percent been more such events in Indonesia (as politically of the Plate families (111 with shorefish represen- defined today) than in any other equivalent sur- tation) account for over half of the species present. face area in the Indo-Pacific. Similar figures for these families were reported by Myers and Shepard (1981) for Guam alone, on Numbers of Shorefish Genera and Species the margin of the Pacific Plate. These same fam- ilies form an important part of the shorefish The 111 families with shorefish representation faunas in all parts of the world, but nowhere else, on the Pacific Plate nonmarginally include about including the Indo-West Pacific, are they so dom- 461 genera and an estimated 1312 species (Ap- inating a component. pendix 2, Table C). The list of genera was com- Using information developed from the analysis piled from the literature, museum collections, and of the Pacific Plate shorefish fauna, I attempted in consultation with numerous colleagues, and is to ascertain the total number of shorefish species probably reasonably complete. The number of for the entire Indo-Pacific. I did this in two ways: species was similarly compiled, but includes nu- one, by assigning known or estimated numbers of merous estimates that might affect the total by species to each family and, two, by estimating the 120 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY total number based on the proportion of the total sitional zone or whether the transition is gradual number of Indo-Pacific species represented on the and uniform." Hedley (1899), however, had al- Plate in certain reasonably well-known and spe- ready discussed the presence of a sharp break in ciose groups of fishes (for instance, Labridae, both the terrestrial and marine molluscan faunas Pomacentridae, Pomacanthidae, Syngnathidae). to the east of the continental islands ranging from The first method resulted in an estimate of 3966. New Guinea to Fiji. The phenomenon of a sharp Using the second method, percentage represen- break was reviewed by Thome (1963) for the tations in the Plate groups range from about 20 terrestrial biota and emphasized for plants by van to 50 percent, and average about 33 percent, Balgooy (1971), but recent general discussions of giving a total of 3936 (±10 percent) shorefish Indo-Pacific biogeography (Briggs, 1974; Kay, species for the Indo-Pacific. This latter figure is 1980), and the fish literature in general, do not probably low because it does not allow for the mention a sharp break. My findings on shorefish numerous families that have no Pacific Plate families support both a sharp break in numbers representation. It overcompensates, however, for of families and a gradual decrease in numbers of monotypic families with 100 percent representa- families as one proceeds across the western mar- tion. The two methods, nevertheless, are in re- gins of the Philippine and Pacific plates. The markable agreement, and an estimated total of decrease, whether sharp or gradual, depends on about 4000 shorefish species for the Indo-Pacific the path one follows from the continental areas is probably close to reality. Such a total is about and on the groups of fishes one selects. 30 percent greater than the only other recent If one proceeds southeast from New Guinea estimate of which I am aware, "—probably more along the continuous island chains to Fiji and than 3000 (Briggs, 1974:13)," which may have then east along the island chains to Easter Island, been based on the numbers included in the out- the decrease in number of shorefish families is dated 11 volumes of The Fishes of the Indo-Australian gradual (Figure 58). If one proceeds due east from Archipelago. Cohen (1973) offered a guess that the Philippine Islands or New Guinea, the de- there may be 3000-4000 species of tropical shore- crease is abrupt: an immediate 32-36 percent of fishes in the Indian Ocean, where there is consid- the families drop out as one reaches or leaves the erable endemism (the Red Sea alone may have western margin of the Pacific Plate. On a group as much as 15 percent endemic fishes; see Briggs, basis, the elasmobranchs are conspicuous for an 1974, for discussion). If Cohen's estimates are abrupt decrease in numbers as one enters the correct, my estimated total of 4000 for the entire Pacific Plate at any point along its eastern mar- Indo-Pacific is much too low. gin. The entire order Rajiformes (Figures 36 and 37) and the entire families Pristidae (Figure Boundary Effect between Pacific Plate 34), Gymnuridae, Rhinopteridae, Squantinidae, and Indo-West Pacific Triakidae, Stegastomatidae, Orectolobidae, Bra- chaeluridae, Hemiscylliidae, and Parascylliidae While there is reason to question whether there (all but the Parascylliidae with shorefish compo- is a decrease in taxonomic diversity as one moves nents), and certain genera of Carcharhinidae (at west from the Indo-Malayan region, there is un- least Loxodon, Scoliodon, Rhizoprionodon (Figure 13), questionably a decrease in diversity as one moves and Dirrhizodon) are all absent from the Pacific east (Figures 58, 59). The nature of this decrease, Plate. Other elasmobranch groups are known whether abrupt or gradual, is of interest here. nonmarginally on the Plate only from a single Ekman (1953:18) was uncertain where to draw species each, and only from deep water at the the boundary between the marine faunas of the Hawaiian Islands: Urolophidae, Scyliorhinidae, Indo-Malayan region and the oceanic islands of Odontaspididae, Torpediniformes (Figure 44). A the Pacific, and remarked that it was not known perusal of the distribution maps I present for whether there was "any definite border or tran- many teleost groups emphasises the sharpness of NUMBER 367 121 the break in the distributions of shorefishes along for each of the Indo-West Pacific taxa that are the Pacific Plate margin, regardless of the direc- missing from the Plate. In the case offish families, tion taken from the continental areas, indicating, there are no Plate endemic shorefish families. that absences from the Plate may be selective, With regard to fish genera, there are about ten rather than only random. endemic to the Plate (Figures 26 and 60). Sister- The sharp decrease in taxa as one proceeds group relationships for these genera have not eastward across the Pacific Plate's western margin been proposed formally, but for one of them, the has not been explained. Two hypotheses have minute Mednsablennius (Blenniidae), known only bearing on any proposed explanations. from the Tuamotus, the sister genus is probably Reality Hypothesis: Those taxa that are ab- Mimoblennius or Cirrisalarias (based on synapomor- sent from the Pacific Plate have never been pres- phies involving nostril size and distribution of ent on the Plate. nasal cirri), both of which are widely distributed, Extinction Hypothesis: Those taxa that are and restricted to the Indo-West Pacific. There are absent from the Pacific Plate were once present numerous endemic Plate species (Table 2 lists on the Plate, but have become extinct there. some of the more important ones for this discus- Although I believe that the Reality Hypothesis sion, the widely distributed Plate endemics). Un- is the more parsimonious and pertains to a large fortunately, the sister-group relationships of these majority of the taxa that are absent from the taxa also have not been proposed formally. Some Pacific Plate, there is merit in both hypotheses species distributions, however, are strongly and both will be discussed. suggestive of relationships, and for some species REALITY HYPOTHESIS.—The only direct evi- the specialists' intuitive statements of relationship dence that would confirm this hypothesis would are probably correct; a few of these distributions be the presence of a Plate endemic sister taxon and relationships are discussed here.
• Crystallodytes
O Evips
O Isocirrhitus
• Lentipes
•fo Liobranchia
S Hedusablennius
FIGURE 60.—Distributions of nine of the ten shorefish genera endemic to the Pacific Plate and its margins; for tenth genus see Figure 26; all endemic genera are monotypic, with possible exception of Lentipes (see "Gobiidae," page 44). 122 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 2.—List of probable Type 1 (widely distributed) POMACENTRIDAE Pacific Plate endemic fishes Amphiprion chrysopterus Figure 32 Plectroglyphidodon imparipennis P. phoenixensis Family/Species/Subspecies Distribution map Stegastes aureus ACANTHURIDAE SCORPAENIDAE Acanthurus achilles Figure 5 Scorpaena coniorta A. leucopareius Figure 5 Scorpaenopsis fowleri Figure 26 A. nigroris SERRANIDAE Ctenochaetus hawaiiensis Anthias bartlettorum Figure 41 Zebrasoma flavesccns A. pascalus Figure 41 BLENNIIDAE Epinephelus socialis Cirripecles jenningsi TETRAODONTIDAE C. variolosus Canthigaster epilampra Figure 26 Entomacrodus cymatobiotus Figure 8 * Also recorded off Pacific coast of Colombia in the E. sealei Figure 8 eastern Pacific. CALLIONYMIDAE Synchiropus laddi S. morrisoni Bruce B. Collette (in Springer, 1981a) believes CHAETODONTIDAE that the closest relatives of the widely distributed Chaetodon quadrimaculatus C. trichrous Plate endemic, Hyporhamphus acutus (Hemiram- Hemitaurichthys thompsoni phidae), is a group of species limited to the Aus- ClRRHITIDAE tralian-New Zealand Region. There are only two Cirrhitichthys serratus* species of Chalixodytes (Creediidae), one a widely Cirrhitops hubbardi distributed Plate endemic, the other endemic to Isocirrhitus sexfasciatus Figure 60 the Indian Ocean. Randall (in press a) believes Paracirrhites nisus CREEDIIDAE that the closest relative of the Plate endemic Chalixodytes tauensis labrid, Labropsis polynesica (ranging from the Cook Crystallodytes cookei Figure 60 to Gambier islands), is L. australis, which is known GOBIIDAE only from the Great Barrier Reef and islands Kelloggella centralis Figure 20 along the western margin of the Pacific Plate, K. oligolepis Figure 20 from the Solomons to Fiji, Samoa, and Tonga. Nemateleotris helfrichi Figure 21 HEMIRAMPHIDAE Perhaps not so clear-cut, but still suggestive, are Hemiramphns depauperatus the three species of Nemateleotris (Gobiidae): one, Hyporhamphus acutus Figure 22 a widely-distributed Plate endemic; one, widely HOLOCENTRIDAE distributed in, and restricted to, the Indo-West Myripristis amaenus Figure 24 Pacific; and one encompassing the range of the M. randalli Figure 24 M. woodsi Figure 24 other two species (Figure 21). LABRIDAE Several invertebrates provide evidence com- Bodianus loxozonus trotteri parable to that provided by fishes. Rosewater Labroides rubrolabiatus Figure 25 (1972) believed that the closest relative of the Pseudojuloides atavai widely distributed Pacific Plate endemic gastro- OPHICHTHIDAE Schismorhynchus labialis Figure 26 pod, Tectarius grandinatus, is T. rugosus, known only OSTRACIIDAE from the Philippine Islands and Celebes. Solem Ostracion whitleyi (1981; see also Springer, 1981a) seems to indicate POMACANTHIDAE that the closest relatives of the widely distributed Apolemichthys griffisi Pacific Plate endemic land snail family Endodon- A. xanthopunctatus Centropyge loriculus Figure 31 tidae are those members of the Charopidae con- C. multicolor Figure 31 fined to southwestern Australia. Banner and Ban- C. nigriocellus Figure 31 ner (1975) recognize two subspecies of the pistol NUMBER 367 123 shrimp, Alpheus lobidens, one a widely distributed Pacific Plate, a large proportion of the Plate- Plate endemic, the other widely distributed in the absent families, including several that are widely Indo-West Pacific, with a single, disjunct occur- distributed in the Indo-West Pacific, have com- rence in the Mediterranean (Figure 51). Austin ponents with a strong estuarine or freshwater B. Williams (pers. comm.) believes that the closest parameter in their ecology: Ariidae, Batrachoid- relative of the widely distributed Plate endemic idae, Centropomidae, Chirocentridae, Kurtidae, crab, Latreillia metansia, is L. valida, which ranges Plotosidae, Pristidae, Sciaenidae, Sillaginidae, from the east coast of Africa to Japan and Phil- Sparidae, Terapontidae, Toxotidae, Trichiuri- ippines (Figure 50). dae. One might assume therefrom that the pau- If one accepts these presumed sister taxa rela- city of estuarine and freshwater habitats on the tionships, then the reality hypothesis has been Pacific Plate provides an ecological basis for the corroborated. Certainly, more rigorous corrobo- inability of some taxa of these families to become ration is needed. established on the Plate. On the other hand, five It is desirable to explain why many taxa that families (Ambassidae, Gobioididae, Leiognathi- are widely distributed in the Indo-West Pacific dae, Monodactylidae, Scatophagidae) with (and, variously, throughout the world) have not strong estuarine and freshwater parameters in become established on the Pacific Plate. An ob- their ecology do occur, variously, in the three vious first premise would be that a physical bar- widely separated high islands (Truk, Ponape, and rier to dispersal is present along the western mar- Kosrae-Kusaie) of the Eastern Caroline chain gin of the Pacific Plate.2 Depending on when this where rivers and estuaries exist (none of the spe- barrier first appeared, it might also have served cies in these families is represented by endemics to isolate taxa on the Pacific Plate and permitted in the Carolines). One might wonder, therefore, them to speciate. why more such families are not present at these The western margin of the Pacific Plate is islands. outlined by a series of deep-sea trenches (Figure Springer (1981a), in calling attention to the 1), which probably represent a barrier that pre- existence of widely distributed Pacific Plate en- vents some shallow-benthic marine fishes and demics, noted that plate margins are character- other organisms from dispersing onto or off the ized by frequent and varied tectonic activity, and Plate, but great depths alone are not barriers to that physical evidence for a barrier on the western the dispersal of forms with pelagic life-history margin of the Pacific Plate may have been sub- stages, which many Plate-absent and Plate-en- ducted and lost forever. If so, one might expect demic shorefishes possess. Nor, obviously, are that recent dispersal would be obscuring the effect great depths a barrier to the dispersal of terrestrial of the former barrier, and there is reason to organisms. believe that it is. In contrast to the overwhelming majority of The east-west Caroline Islands chain is being the fish families that are well represented on the carried westward on the Pacific Plate as it moves toward the Philippine Plate margin (Figure 1), 2 The well-known East Pacific Barrier (Ekman, 1953; and islands on the eastern margin of the Philip- Briggs, 1961), a long expanse of ocean separating the islands pine Plate are moving eastward through the of the Pacific Plate from the coasts of the eastern Pacific, is, mechanism of back-arc spreading (Karig, 1971, with very few exceptions (only about 40 Indo-Pacific shore- 1974). If no islands of the relatively young Caro- fish species occur in the eastern Pacific; Rosenblatt et al., line Islands chain3 on the Pacific Plate have been 1972), effective in isolating the Plate biota from that of the eastern Pacific, but will not be considered further. The obvious relationships of the shallow marine forms of the 3 Recent unpublished radiometrically determined ages for western and central Plate areas are predominantly with the Kosrae, Ponape, and Truk, in order, are 4, 8, and 12-14 biota of the Indo-West Pacific, and it is the possibility of million years (B. Keating, Department of Geology and Geo- barriers along the western margin of the Plate that concern physics, University of Hawaii at Manoa, pers. comm.; also me. mentioned in Kerr, 1981). My extrapolation from the spac- 124 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY lost through subduction (Pacific Plate under Phil- Islands, New Hebrides, and Fiji, all along the ippine Plate), the expanse of open water separat- margin of the Pacific Plate. A 2.8 m long speci- ing the Caroline Islands on the Pacific Plate from men was trapped at Ponape in 1971, where it was the eastern margin of the Philippine Plate must believed to have been resident for at least six have been greater than it is today (see section on months, based on the disappearance of pigs teth- geological history of Pacific and Philippine ered at the shoreline. While no other crocodiles Plates). If such an expanse existed, it could have are known to have occurred at Ponape (infor- served as a barrier to dispersal. Today, however, mation obtained during my visit in 1980), the the westernmost islands of the Caroline chain on potential exists for crocodiles to colonize Ponape the Pacific Plate are proximate to the islands on (providing they avoid pigs and people). the eastern margin of the Philippine Plate and The western margin of the Pacific Plate need seem to be serving as a conduit (which I define as not be the site of a physical barrier to dispersal in an essentially unidirectional corridor; see Mc- order that it sharply delimit the distribution of Kenna, 1972) that permits both terrestrial and Indo-West Pacific organisms. It might simply be marine organisms to island hop (disperse) onto that these organisms lack the ability to actively the Pacific Plate. Some possible examples of dis- disperse. It is significant, therefore, to note that persals using the Caroline Islands conduit are: fishes, such as Scomberomorus (Figure 40), that have Ambassidae, Centriscidae, Gobioididae, Leiog- planktonic young and fairly large free-swimming nathidae, Monodactylidae, Scatophagidae, Per- adults, have distributions similar to those of iophthalmus (Gobiidae), Butis (Eleotrididae), Omox, fishes, such as the Batrachoididae (Figure 6), that Glyptopams, Nannosalarias (all Blenniidae), Synchi- are relatively small and benthic and have large, ropus splendidus (Callionymidae), Zenarchopterus demersal eggs and no pelagic life-history stages (Hemiramphidae), Calloplesiops (Plesiopidae), (at least in the Indo-West Pacific). Both groups Dascyllus melanurus (Pomacentridae), Euxiphipops are widely distributed along continental coasts navarchus (Pomacanthidae), Rastrelliger kanagurta and absent from the Pacific Plate. If there were (Scombridae), Belonoperca (Serranidae), all of no widely distributed endemic Pacific Plate fish which are widely distributed in the Indo-West species, I could accept that a physical barrier to Pacific and present on the Pacific Plate nonmar- dispersal along the western margin of the Pacific ginally only in the Eastern Caroline Islands. Ex- Plate is not needed to restrain the dispersal onto amples from other organisms include certain bee- the Plate of species presently restricted to the tles (Figure 55), certain bats (Figure 56), and a Indo-West Pacific. I can only conclude, under the mangrove, Rhizophora apiculata (van Steenis, circumstances, that a barrier (or barriers) must 1963:253). The conduit must be relatively recent exist, or have existed. because its influence seems not to have greatly EXTINCTION HYPOTHESIS.—There are no fish diluted the integrity of the Pacific Plate biota. fossils reported from the islands on the Pacific One can expect that it is merely a matter of time Plate, hence, no evidence that any of the families, before other fishes (and other organisms) cur- genera, or species of fishes currently limited to the rently missing from the Pacific Plate are able to Indo-West Pacific were once present on the Plate. disperse onto the Plate. Of interest here is a recent Fossils of other organisms are present on the report (Allen, 1974) of the occurrence of a marine Pacific Plate, and are best documented for the crocodile, Crocodylus porosus, at Ponape. Crocodylus Mollusca, although I found it difficult and un- porosus is distributed from India eastward only as feasible to assess all the possibly relevant data. far as Palau, Bismarck Archipelago, Solomon The most informative areas on the nonmar- ginal portions of the Pacific Plate that are re- ing of the dated islands would indicate that edifices of the ported to have marine molluscan fossils are: chain nearest the subduction zone could be about 16-20 Enewetak and Bikini, Marshall Islands; Funafuti, million years in age. Ellice Islands; and the Hawaiian Islands. All the NUMBER 367 125 fossils from these islands are from the Cenozoic. the Pacific Plate is possibly exemplified by what Information on older fossils is too sketchy to has occurred in the Hawaiian Islands. Kohn appraise (for instance, see Kauffman, 1976). (1980, based mostly on Kosuge, 1969) discussed Ladd (1966, 1972, 1977) reported on the abun- the presence of fossil marine mollusks in the dant chiton and gastropod molluscan fossils at Hawaiian Islands. He noted that of 163 species Bikini, Enewetak, and Funafuti (at least one ad- known as fossils there, only three endemic pele- ditional report in this series is in preparation). cypod species in two genera (Ostrea, Septifer; based These fossils range in age from about Eocene to on Kosuge, 1969) and one endemic gastropod (a Recent, and include 48 families, of which only species of Conus) are universally extinct. All three four (Melongenidae, Turritellidae, Potamididae, genera, however, are represented by extant spe- Iravadiidae, all gastropods) are not known to be cies in the Hawaiian Islands. Eight other species extant on the Pacific Plate, but are extant in the that are extinct in the Hawaiian Islands are Indo-West Pacific. Three of the four families that extant and common at other islands on the Pacific appear to be absent are inhabitants of estuarine Plate. The remaining 152 species of fossils are also or mangrove habitats, and the fourth, Turritelli- represented in the extant fauna of the Hawaiian dae, is a sand-dwelling group that has been found Islands. in harbors, and, thus, may have an estuarine A perusal of all the issues of the journals lndo- component in its ecology. Estuarine and man- Pacific Mollusca and Monographs of Marine Mollusca grove habitats are not common on the Plate, and evinced no examples of families, genera, or species are predominantly restricted to high islands. The that were extant on the Pacific Plate, but are now appearance of the four families in the Eocene to absent. Indeed, none of the larger-sized mollusks Pliocene strata of the Marshall Islands, which are (macromullusca) that are presently confined to atolls today, indicates that these islands were the Indo-West Pacific are known as fossils on the formerly high islands with brackish and man- Pacific Plate, leading one to the conclusion that grove environments. It is quite possible that some, the supposed absence of some extant Indo-West or all, of the four missing families will be found Pacific taxa known otherwise only as fossils on extant at high islands, such as Ponape, which has the Plate is a consequence of their having been extensive estuarine and mangrove habitats, but overlooked. The largest dimension of any of the whose molluscan fauna has not been well studied Plate fossils reported by Ladd (1962, 1972, 1977) or collected (H.A. Rehder, pers. comm.). is about 50 mm, but for many is less than 5 mm. Ladd reported on numerous fossils of genera in That Ladd reported on so many diminutive taxa families extant at both Pacific Plate and Indo- is the result of his detailed examination of core West Pacific localities. It was not feasible for me samples from a deep-drilling program. In the to determine which of these genera were extant same three studies, Ladd did record fossil mol- and, if so, where; however, the range of one of lusks of larger sizes, but only from andesitic lo- the genera, Cynisca (Turbidinidae), was discussed calities (Fiji, Marianas, etc.) along the western by Ladd (1966). Cynisca is known only from Mio- margin of the Pacific Plate. Similarly, the journals cene fossils at Bikini and several extant species mentioned above also include numerous listings from the seas off South Africa. If the known of large-sized molluscan fossils, but only from distribution is truly indicative of the present dis- continental areas and the Hawaiian Islands. tribution of this micromolluscan genus (maxi- Comparing mollusks with fishes, it might be mum dimension of Ladd's specimens, 4.5 mm), argued that mollusks are more susceptible to extinction of Cynisca would appear to have oc- extinction than fishes, because mollusks are pre- curred in all the extensive area between South dominantly sedentary and less able to actively Africa and Oceania. avoid conditions adverse to their existence than The manner in which extinctions have gener- are fishes. The larvae of mollusks, however, are ally affected marine molluscan distributions on predominantly planktonic and could recolonize 126 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY areas on the Plate that offered proper habitats4 if The land snail family Endodontidae (Gastro- they were ever able to colonize the Plate. Simi- poda), which is endemic to the Pacific Plate and larly, large, strong, free-swimming fishes (such as its margins, has been extensively studied (Solem, Scomberomorus, Rachycentron, and certain species of 1976, 1977). The endodontids are minute forms Carcharhinus) should be able to colonize the Plate, and are known as fossils only from Midway (Ha- but do not in spite of the fact that proper habitats waiian Islands) and Bikini (Marshall Islands). seem to be present, and some of these fisheshav e The fossil species at Bikini belong to the genera planktonic larvae. Furthermore, if a taxon were Cookeconcha (a lower Miocene species) and Mini- widely distributed on the Plate, extinction would donta (a Pliocene-Pleistocene species). Cookeconcha have to cover an extraordinarily broad area and is extant only on the principal Hawaiian Islands. innumerable islands, factors that would seem to Minidonta is extant on the Samoan, Cook, Austral, buffer whatever unfavorable effects were causing and Gambier islands, and Henderson Island. The extinctions on one part of the Plate. Cases in fossil species at Midway belong to Cookeconcha and point are the eight species of marine mollusks Protoendodonta (a Pleistocene species in each). Pro- that are represented only by fossils in the Ha- toendodonta is known only from Midway. waiian Islands, but are extant at other islands on The evidence favoring the selective extinction the Pacific Plate. To these cases may be added of Pacific Plate taxa as an explanation for the some terrestrial examples. restriction of the range of extant members of the same taxa to the Indo-West Pacific or continental 4 Evidence that proper habitats are available on the Pa- plate areas seems much weaker and less promising cific Plate for species that do not occur there naturally is to than that favoring the Reality Hypothesis. There be found in the cases of successful introductions. The com- are numerous opportunities for corroborating or mercially important gastropod, Trochus niloticus (Trochidae) falsifying the Reality Hypothesis, but these will is widely distributed in the Indo-West Pacific, from the Andaman Islands in the west through Indonesia to the require the difficult, although not impossible, Philippines, Yap, Palau, New Guinea, New Caledonia, work of establishing the sister species of the widely northern Australia, and the Admiralty, Solomon, New He- distributed Pacific Plate endemics. Nevertheless, brides, and Fiji islands (McGowan, 1958). Trochus niloticus it is instructive to examine the causes (particularly was unknown from Pacific Plate localities, but was intro- the geotectonic causes) and results of the limited duced by the Japanese into the Eastern Caroline and Mar- types of extinctions that may occur on the Pacific shall islands (includes high islands and atolls) in the late 1920s and 1930s (McGowan, 1958), and now forms an Plate. important food resource in islands where it was introduced. Based on the examples of extinctions discussed Trochus niloticus is a shallow-dwelling species that prefers the above, one might expect to find geographically ocean sides of reefs as its habitat. Although T. niloticus has patchy distributions of organisms relatively com- spread, apparently by larval dispersal (McGowan, 1958), to other locations on the immediate islands or atolls where it monly among the islands on the Pacific Plate. was introduced, it apparently has not spread to other islands Kay (1976, 1980) noted that patchy distributions or atolls. Heslinga (1981) ascribes the limited distribution of of mollusks are common among the islands of the T. niloticus to the short duration of its planktonic stage. Yet, central Pacific, and ascribed such distributions to its natural, widespread occurrence in the Indo-West Pacific, patchy ecology, at the same time excluding col- including many islands, might tend to discount this factor. lecting artifacts and chance dispersal as contrib- The cichlid fish, Sarotherodon mossambicus (tilapia) is a freshwater and estuarine species native to Africa. In 1958, uting factors. Although some patchy distributions tilapia were introduced into the lagoon at Fanning Atoll, may be ascribable directly to patchy ecology, I Line Islands (Lobel, 1981a). Tilapia are presently living and believe the major causes are extinctions resulting reproducing in the lagoon, which is fully marine. Of similar from physical processes, which, of course, directly nature to the Trochus and tilapia introductions are the intro- affect ecology. The presence of proper habitat, as ductions of Lutjanus (Lutjanidae, snappers), Cephalopholis (Serranidae, groupers), and Sardinella marquesensis (Clupeidae, shown above (footnote 4), is insufficient of itself sardines) into the Hawaiian Islands from the Society and to assure the presence of a species adapted for Marquesas islands (see discussions under family accounts). that habitat. Physical processes that affect extinc- NUMBER 367 127 tions include volcanism, tectonics, weathering, there is only one hot spot that is presently forming and glacioeustatic changes in sea level. These islands, and that is the Hawaiian hot spot (there processes may act alone or in concert to cause may be a second in the Eastern Caroline Islands, extinctions or modify ecology. To understand but this has not been clearly established, nor has how these processes operate on the Pacific Plate, it been effective for four million years). Other the following generalized scenario is presented. than the Hawaiian Islands, no islands have been Islands on the Plate arise by volcanism. At the forming on the Plate for about a million years completion of formation, an island is usually a (Pitcairn, which is about 0.9 m.y. old, appears to high island. High islands are more complex eco- be one of the youngest). If these circumstances logically than low islands or atolls and can sup- persist, all the islands on the Pacific Plate, except port many more species than low islands or atolls. the newly formed ones at the Hawaiian hot spot, After formation, island height is lowered by will gradually become low islands, then atolls, weathering, which destroys habitats. The Pacific drowned, and then be subducted, causing partial Plate is moving in a northwesterly direction and and then total extinction of their biotas. Inas- carries islands with it, moving them into different much as all islands are not synchronously or climatic regimes, which will cause extinctions if equally affected by the physical processes de- the island species do not evolve adaptations to scribed, patchy extinctions result, which are re- compensate for these climatic changes. The Pa- flected by patchy distributions. cific Ocean increases in depth from east to west and, thus, Plate movement carries islands into Endemism deeper water. While coral reef growth in tropical latitudes can offset some of the effects of deepen- ENDEMICITY OF SHOREFISH FAMILIES AND GEN- ing waters, an island is ultimately drowned and ERA.—Although several shorefish families are en- eliminated as it approaches subduction zones demic to the Indo-West Pacific, none is endemic along the Plate margins—its terrestrial and shal- to the Pacific Plate. The only possibility of family low-dwelling biota extinguished. Glacioeustatic level endemism among fishes on the Plate is that changes (see footnote 5), aesthenospheric bumps of an undescribed family, genus, and species of (Menard, 1973), and crustal loading (McKnutt large shark collected in 1976 at a depth of 150 m and Menard, 1978) can also temporarily increase off the Hawaiian Islands (Tinker, 1978). The 3.5 effective island height, or lower it, but, again, m long specimen was entangled in a sea anchor. these changes come to naught. If an island is of It seems probable that this shark species is more the solitary type that forms near a fault zone or widely distributed in the sea than present infor- on the rise formed by a seafloor spreading zone, mation indicates. its biota can survive only if the island integrates Among the shorefishes, I have found only ten (Rotondo et al., 1981) with a younger, higher genera (2.2 percent of the number of genera island, or if another island, by chance, appears present nonmarginally on the Pacific Plate; ap- nearby (see section on Easter Island). In contrast, pendix 2, Table C) that appear to be endemic to an island that forms as a result of the periodic the Plate (Figures 26 and 60). Among the endem- activity of a melting anomaly (hot spot), which ics I have included one genus that ranges, ques- by its intermittent activity gives rise to the rela- tionably, to the northwestern margin of the Phil- tively closely spaced islands of an island chain, ippine Plate (see Lentipes in Gobiidae), and an- such as the Hawaiian Islands, elements of the other, Evips, which is known only from the holo- island's biota can survive by dispersing back to a type of its type-species, collected at Palau on the recently formed (high), nearby island (in such an eastern margin of the Philippine Plate. All of the instance, the age of an island's biota is not nec- endemic genera, with the possible exception of essarily equal to that of the island, but to that of Lentipes and a possible new genus of cirrhitid (see the island chain). On the Pacific Plate today, Cirrhitus wilhelmi in Cirrhitidae) are monotypic. 128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
The endemic genera belong in the following fam- (Collette, 1974; Springer, 1981a) that attention ilies: Blenniidae (Medusablennius), Cirrhitidae (Is- has been called to Type 1 endemics. ocirrhitus and possible new genus), Creediidae A list of the 48 Type 1 endemics that I have {Crystallodytes), Gobiesocidae (Liobranchia), Gobi- been able to determine is given in Table 2, and idae (Lentipes, Vitraria), and Ophichthidae (Cirri- each is discussed or mentioned in the appropriate caecula, Evips, Schismorhynchus). In addition to thefamily account. There are undoubtedly many ten genera, there is one endemic, monotypic more Type 1 endemics, and colleagues actively subgenus, Acanthotaurichthys (proposed somewhat revising Indo-Pacific fish groups continually ap- cryptically by Burgess, 1979:218; Chaetodonti- prise me of them. Examples of Type 1 endemics dae, genus Hemitaurichthys), known only from Pit- among invertebrates have been mapped in Fig- cairn Island. ures 47, 48, 50, 51, and several others are men- It is possible that some of the genera now tioned in the discussion under "Other Orga- considered as endemics will be shown to be more nisms." widely distributed in the Indo-Pacific. For ex- Most Type 2 endemics are confined to the ample, during much of the time my study was in southern islands of the Plate in the area between progress I believed that the gobiid genus Kellog- the Cook and Society islands in the west, and gella was a Plate endemic, but a species (undes- Pitcairn and Easter in the east. A high proportion cribed, D.F. Hoese, in litt.) was obtained recently of these islands are high islands (Figure 2). Ex- in the Indian Ocean (Figure 20; see discussion in amples of Type 2 endemics are Heteroconger lenti- Gobiidae). I doubt, however, that many, if any ginosus (Congridae; Marquesas and Society Is- additional Plate endemic shorefish genera will be lands); Chaetodon hemichrysus (Chaetodontidae; recognized. Rapa and Pitcairn); Cirrhitus wilhelmi (Cirrhiti- Although only indirectly relevant to my study, dae; Rapa to Easter; see "possible new genus" on it appears that there may be numerous endemic Figure 60); Entomacrodus rofeni (Blenniidae; Tua- deep-slope fish genera on the Pacific Plate, for motu, Henderson, and Ducie islands; Figure 8); example: Paraprislipomoid.es (Lutjanidae), Neomer- a color-pattern type of E. caudofasciatus (Raro- inthe (Scorpaenidae), Stethopristes (Zeidae; see tonga to Ducie; Figure 7); Centropyge hotumatua Heemstra, 1980, for a world distribution map of (Pomacanthidae; Rapa to Easter; Figure 31); the species of this moderately deep-dwelling fam- four species of Pomacentridae: Stegastes emeryi ily), Pelecanichthys (Bothidae), and, possibly, the (Tuamotu and Pitcairn islands), Chrysiptera galbus shark mentioned above. (Cook to Pitcairn islands), Pomachromisfuscidorsalis ENDEMICITY OF SHOREFISH SPECIES.—The lack (Society Islands to Pitcairn), and P. exilis (Eastern of endemic shorefish families and paucity of en- Caroline and Marshall islands); Synodus capricornis demic shorefish genera on the Pacific Plate are (Synodontidae; Pitcairn and Easter); Cantherhines not duplicated at the species level. The commo- longicaudus (Monacanthidae; Rarotonga and Ta- ness of endemic shorefish species on the Plate, hiti); Eviota epiphanes (Gobiidae; Hawaiian, John- particularly the widely distributed endemics, con- ston, and Line islands); E. disrupta (Tonga, Sa- vinced me of the importance of the Pacific Plate moa, Society, and Tuamotu islands); Mulloides as a biogeographic region. I have arbitrarily di- mimicus (Mullidae; Line and Marquesas islands); vided the endemic species into three types: (1) Bodianus bilunulatus busellatus (Labridae; Ducie, widely distributed endemics occurring in numer- Henderson, and Marquesas islands; M.F. Gomon, ous island groups or in widely separated island 1979); B. b. albotaeniatus (Hawaiian and Johnston groups; (2) endemics limited to a few islands or islands; perhaps, more appropriately considered island groups within a limited area on the Plate; as a Type 3 endemic); B. loxozonus trotteri (Line, and (3) single island or island group endemics. Marquesas, Tuamotu, and Austral islands); La- Types 2 and 3 endemics have been recognized bropsis polynesica (Labridae; Rarotonga to Gam- often in the literature, but it is only recently bier Islands); L. micronesica (Guam, Palau, Ifaluk, NUMBER 367 129
Enewetak, and Kwajalein); Psendojuloides atavai (Labridae; Fanning, Line Islands). (Labridae; southeast Oceania); Thalassoma undes- The total number of the three types of endem- cribed species (Heiser, 1981; Labridae; Ducie and ics (species only; subspecies excluded) mentioned Pitcairn); Zebrasoma rostratus (Acanthuridae; Tua- or alluded to in the previous discussion is equal motu, Marquesas, Line, and Rapa islands). Cres- to about 20 percent of the estimated nonmarginal sey and Randall (1978) list three additional spe- Pacific Plate shorefish fauna. I predict that total cies that are confined to the islands from Rapa or endemism of shorefishes on the Plate will reach Pitcairn to Easter: Pseudolabnis Juentesi (Labridae), 22 to 25 percent. The information offered here is Gymnothorax nasuta (Muraenidae), Goniistius in direct contrast to generally held opinions that (= Cheilodactylus) species (Cheilodactylidae). "the homogeneity of the Indo-Pacific [fish] fauna The most common type of endemism on the is at once apparent from the fact that great Plate consists of Type 3 endemics, and most of numbers of shore species exist from one end of it these endemics occur at high islands. For instance, to the other" (Myers, 1940:203; Cohen, 29 percent of the 442 species of Hawaiian shore- 1973:453). Great numbers, perhaps, but consid- fishes, about 10 percent of the approximately 350 ering the great numbers of families, genera, and species of Marquesas Islands shorefishes, and species that reach, but do not transgress the west- about 27 percent of the 99 species of Easter Island ern margin of the Pacific Plate, and the large shorefishes are Type 3 endemics (Randall, 1976, percentage of species that are restricted to the 1978a). The number of endemics present in these Plate, homogeneity is hardly descriptive at the three island groups totals about 190, or about 14 species level. I would agree wholeheartedly, how- percent of the estimated total number of shorefish ever, with recognition of the Indo-Pacific as a species on the Plate. Although Easter Island is biogeographic unit, with the Pacific Plate distin- not on the Pacific Plate, it is close to the Plate's guished as a major subunit. eastern margin, and I consider its fish fauna, The importance of knowing the sister-group which is most closely related to that of the rest of relationships of the endemic species on the Pacific the Indo-Pacific (Randall, 1976), to be part of Plate, and the distributions of the sister groups, the Plate fauna. Rehder (1980) similarly related was mentioned in the discussion of the Reality the molluscan fauna of Easter Island most closely Hypothesis, and the few Type 1 Pacific Plate to that of the Indo-Pacific. No recent compilation endemics for which sister species could be sur- of Marquesas Islands fishes has been published. mised reasonably, were mentioned. If more infor- Randall's (1978a) assessment of endemism at mation on sister-species relationships was gener- these islands was based on his extensive knowl- ally available it would serve to direct our search edge of Indo-Pacific fishes, including his collect- for the mechanisms (vicariant events) that di- ing expeditions to the Marquesas. Based on my vided ancestral populations into separate subpop- investigations of blenniid fishes at the Marquesas, ulations and allowed these subpopulations to di- I can concur completely with Randall. verge. For Indo-Pacific fishes, there is scant infor- Examples of low-island Type 3 endemics are mation on sister-group relationships. This deficit few, and most are known from one or a very few exists because cladistic analyses have begun to collections; probably they will be found to occur appear only recently in the literature, and be- more widely. Some apparent low-island Type 3 cause meaningful cladistic analyses of sister-group endemics are: Medusablennius chani (Blenniidae; taxa are often exceedingly difficult to produce. Raroia, Tuamotus; Figure 60), Amphiprion tricinc- Sister-species distributions and relationships for tus (Pomacentridae; Marshall Islands; Figure 32), Types 2 and 3 endemic Plate species are also Plectroglyphidodon flaviventris (Pomacentridae; Tua- lacking, but information bearing on the origin of motus), Acanthurus chronixis (Acanthuridae; Ka- these types of endemics is known or can be hy- pingamarangi), Cirricaecula johnsoni (Ophichthi- pothesized. In the sections that follow, I first treat dae; Enewetak, Figure 60); Bodianus prognathus the three main areas (Hawaiian, Easter, and Mar- 130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY quesas islands) of Type 3 endemism on the Pacific island groups: Marquesas. Society Is., S. Tuamotus and Plate, and follow these with a discussion of pos- Rapa but not with the approximately equidistant E. Caro- lines and Marianas or Revilla Gigedo and Galapagos Is. sible mechanisms in the formation of Types 1 and 2 endemics. Van Balgooy concluded (p. 146), "The (archipe- HAWAIIAN ISLANDS AND NORTH-SOUTH DISJUNCT lagic) connections of Hawaii towards Malesia DISTRIBUTIONS ON THE PACIFIC PLATE.—More is appear not to have been direct through Micro- known about the geology and biology of the nesia, but by way of SE. Polynesia." The impor- Hawaiian Islands than any other island area in tant point that van Balgooy makes is basically the Indo-Pacific. A current geological view of the the same as Zimmerman made, that the closest Hawaiian Islands is contained in Jackson et al. biotic similarity of the Hawaiian Islands is with (1980), and some interrelated aspects of the geo- the islands of Oceania, particularly southeast tectonics and biology of the Islands were discussed Oceania. Whether this similarity is strong or weak by Schlanger and Gillett (1976) and Rotondo et is secondary. Van Balgooy refrained from trying al. (1981). The latter paper will be treated in to explain the distribution patterns he found, but more detail in this section. noted significantly that the usual methods of Knowledge of the geology and terrestrial biota, dispersal did not seem to account for them (p. primarily the insects, of the Hawaiian Islands 143): through 1948 was eloquently presented by Zim- In the present state of our knowledge I cannot find a clear merman (1948), and his treatise still merits atten- correlation between distribution and dispersal. The impres- tion. Like all previous and subsequent students of sion is that the island plants "got there" despite lack of the Hawaiian biota, Zimmerman noted the high efficient dispersal ability rather than on account of superior degree of endemism in the islands, which he dispersal capacity. attributed to their great geographical isolation. Kay (1967) indicated that the Hawaiian Mol- He gave a long and classical discussion of disper- lusca showed a closer relationship with the wide- sal, which he emphasized as the primary mecha- ranging Indo-West Pacific Mollusca than with nism by which the Hawaiian Islands obtained its Pacific Basin restricted Mollusca. Kay (in litt.) biota. Interestingly, he accounted for adaptive now informs me that a small proportion of the radiations within the islands primarily by the Hawaiian mollusks have their closest relation- mechanism of vicariant events. Zimmerman con- ships (not cladistically determined) with mollusks cluded that the geographical affinities of the on the Pacific Plate. In a fine essay on Pacific greatest proportion of the terrestrial endemic Basin biogeography, Kay (1980) presented a gen- biota was with the biota of the Polynesian region, eralized explanation for the distribution patterns which includes only islands on the Pacific Plate in that area. She drew upon recent geotectonic (see his fig. 26). literature and the classical island biogeographic Van Balgooy (1971) published a comprehen- and ecological literature in proposing that a for- sive summary of plant biogeography in the Pa- mer, widely distributed, Mesozoic, or early Cen- cific. He stated (p. 116) that the "floristic alliance ozoic, biota became isolated in the Pacific Basin, [of the Hawaiian Islands] is most pronounced was variously extinguished because of changes in with SE. Polynesia," but rephrased this statement island morphology, positions, and climatically (P- 128), induced sea-level changes,5 and is today repre- Hawaii is another island group without any strong relations with other groups. The relatively high Kroeber coefficient ° Comment on interpreting the effects of paleoclimates on |a measure of biotic similarity] with Clipperton I. is easily sea level is offered here because these effects have generally explained by the fact that nearly all the widespread Clip- been invoked uncritically by biogeographers. For instance, perton genera are also found on Hawaii. The same is true Briggs (1974:427) stated that glacioeustatic sea-level changes for Easter I. Leaving these out of consideration, Hawaii would have affected the north Atlantic and north Pacific shows distinct but weak relations with some SE. Polynesian about equally. Perhaps, but geotectonic activity may have NUMBER 367 131
sented by relict elements in isolated islands or the five oceanic species, that occurs in these four island groups. Two of the studies Kay cited island groups. (Springer, 1981a; Rotondo, Springer, Scott, and The Hawaiian Islands-southeast Oceania type Schlanger, 1981, cited by Kay as Rotondo and distributions are, perhaps, part of a general pat- Scott, 1979) were unpublished at the time, and tern of north-south disjunct distributions that has this fact is noted here to stress the original con- most often involved organisms restricted to con- cepts independently arrived at by her and me, tinental seas. This pattern has been referred to and to be further developed in this section. variously as: bipolar, biboreal, bitemperate, an- Rehder (1980) reported that 10 percent of the titropical, or antiequatorial, depending on how well-studied mollusk species (99) that occur at distant the disjunct populations are from the Easter Island, which is on the Nasca Plate but equator (see Ekman's, 1953, discussion of bipo- close to the East Pacific Rise, are known otherwise larity). To explain the north-south disjunctions, only from the Hawaiian Islands. He reported that some authors, such as Rehder (1980), have as- another 9 percent of the Easter Island species, sumed that the low-latitude portions of the dis- consisting only of endemics, had their closest tributions of formerly widely distributed forms relationships (not cladistically determined) with were eliminated. Rehder explained these elimi- Hawaiian species. Rehder's finding of a mollus- nations by hypothesizing a combination of com- can relationship between the Hawaiian Islands petitive exclusion (a difficult, if not impossible, and Easter Island contrasts with van Balgooy's hypothesis to corroborate and which has often for plants. Van Balgooy's findings, however, were been invoked) and drowning of centrally located based on the generic taxonomic level, and a islands. Island drowning, or changes in island cladistic analysis of the species might provide a morphology as proposed by Kay (1980), would different conclusion. In any event, Rehder's find- have been enough. The fact that high islands on ings provide added support for an the Pacific Plate are today essentially restricted Hawaiian-southeast Oceania relationship. Fur- to southern Oceania and the Hawaiian Islands ther support is found in the distribution of the (excluding islands adjacent to the North Ameri- marine water-striders (Insecta). Halobates ha- can coast; Figure 2) and thus correlate with north- waiiensis is known only from the Hawaiian Islands south disjunct distributions on the Plate corrob- and the southeast Polynesian islands: Marquesas, orates the Kay-Rehder proposal (see section un- Society, and Tuamotus (Cheng, 1973). Halobates der "Extinction Hypothesis" for a geotectonic hawaiiensis is not among the five species of Halo- explanation of the distribution of high and low bates that have invaded the open ocean (Cheng, islands). 1973; Cheng and Shulenberger, 1980), and it is Other authors (most recently, Randall, in press the only species of Halobates, other than some of b, for fishes) have proposed that higher latitude forms were able to cross the tropics during times resulted in a different net effect in each area, or even in when temperatures were cooler and the extent of different parts of each area. Frakes (1979) summarized avail- the tropics considerably decreased. Briggs (1974) able information on the effects of past climates on sea level. and others have commented, however, that the He noted that it is possible presently only to recognize relative changes in sea level through time (p. 208): "It would climatic history of the Indo-Pacific has been sta- be desireable to estimate the extent to which the sea has ble. Recent information bearing on the paleosea- risen or fallen against a fixed bench mark but this is not surface temperatures is worth noting. A plot possible because the landmasses themselves may have moved (CLIMAP Project Members, 1976, fig. 1) of sea- vertically in unison . . . vertical tectonic activity continuously surface isotherms for the most recent glacial pe- affects the configuration of the sea floor itself. Glacioeustatic riod, 18,000 years ago, for example, shows little changes in sea level are recognized in the geologic record but tectonic movements of one sort or another would appear to be responsible shift toward the equator for the 20° C isotherms for most major changes in relative sea levef' (italics mine). when compared with the 20° isotherms of today 132 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
(Sverdrup et al., 1942, charts II and III), but deeper, cooler waters in equatorial regions and there may have been greater temperature shifts ascend to shallower waters once having traversed in earlier Cenozoic times.6 Frakes (1979) provided those regions. This mechanism may provide an some general estimates of the equatorial surface explanation for the distribution of some eury- temperatures during the Tertiary: 16-17° C in bathic, strictly biboreal or bitemperate taxa (C.L. the Pacific at some time during the mid-Miocene Hubbs, 1952), but it is doubtful that littorally (p. 214); 13° C in the eastern Pacific and 22° in restricted taxa, such as the blenniid fish Entoma- the western Pacific at some time during the Oli- crodus niuafoouensis (Figure 7) or the surgeonfish gocene (p. 194: fig. 7-3); 21° C in the eastern Acanthurus leucopareius (Figure 5), of the warmer Pacific and 29° C in the western Pacific at some antiequatorial regions on the Pacific Plate, are time during the Eocene (p. 193: fig. 7-2); 17-19° capable of isothermic submergence either as C for the global ocean during late Paleocene- adults or larvae. early Eocene (p. 214). If the coolest of these Rotondo et al. (1981) presented geological ev- temperatures existed in the equatorial regions of idence to show that the Hawaiian Islands have the Indo-Pacific, it would seem that the coral incorporated volcanic edifices that originated to reefs would have been eliminated. It is known, the southeast of the Hawaiian Islands. Using the however, that coral reefs have been continuously geological data, they hypothesized a vicariant present at Enewetak, Marshall Islands, since at mechanism, island integration, to explain the least early Miocene, and possibly since Eocene origin of some of the endemism in the Hawaiian (Schlanger, 1963; Ladd, 1966). Reefs were present Islands. They offered island integration primarily during early Eocene at Enewetak, but the Oli- as an alternative to founder principle (Mayr, gocene is represented by a hiatus. Even though 1942), a dispersalistic mechanism that is invoked Frakes stressed that all paleotemperature data most often to explain island endemism (for a should be used with caution because of problems dispersalistic view of endemism among Hawaiian in determination, and also noted that it may not fishes see Gosline, 1957, 1971). Island integration be proper to assume that temperature require- also may explain some north-south disjunct dis- ments of organisms of the distant past can be tributions on the Pacific Plate, particularly dis- inferred from the requirements of similar orga- tributions involving the Hawaiian Islands and nisms today, I am skeptical, given the evidence, islands in southeast Oceania. Rotondo et al. out- that north-south disjunct distributions on the Pa- lined island integration as follows. cific Plate resulted from a depression of sea-sur- face temperature in the low latitudes on the Plate. Islands of the Hawaiian-Emperor chain have been forming intermittently during the past 70+ million years over a For a recent, strong opinion to the contrary, but relatively fixed and isolated melting anomaly in the aesthen- with no more direct data from the tropics than osphere below the rigid Pacific lithospheric plate. As the presented here, see Fleming (1979). Pacific plate moved in a northwesterly direction over this Another scheme that has been used often to anomaly, the islands and seamounts so formed were carried with the plate. Gradually, the islands subsided, eroded, explain north-south distributional disjunctions of formed atolls while in warmer waters, were then inundated, marine forms is that of equatorial or isothermic and those that reached the continental margin were sub- submergence: cool-dwelling forms descend to ducted. Concurrent with the formation of some of the Ha- waiian-Emperor chain islands, other island groups formed well to the SE on the Pacific plate close the East Pacific Rise. It seems unnecessary to push the origins of present-day These latter islands shared a biota that was not present at/ distributions back beyond the Tertiary, as most modern on the emergent Hawaiian Islands of the time. As the Pacific family-level taxa of fishes appear to have originated no plate moved, many of these islands became deeply sub- earlier than Lower Tertiary (Andrews et al., 1967). Most merged, but a few may have remained emergent as volcanic modern genera and species probably date little, if any, earlier islands or atolls, thus becoming relatively isolated. As a result than Miocene, where there is a preponderance of first ap- of this isolation, components of their biota began to diverge pearances of modern, widely distributed taxa. (speciate). Ultimately, these emergent islands integrated NUMBER 367 133
with emergent Hawaiian Islands, thus combining non-Ha- Island (on the Nasca Plate near the East Pacific waiian biotas with the Hawaiian. Continued isolation of the Hawaiian Islands reinforced the developing endemism, Rise), and E. cymatobiotus, widely distributed, but which we recognize today. restricted to the islands of the Pacific Plate, ex- cluding the Hawaiian and Easter islands (Figure If this scheme is correct, and it is geologically 8). The halfbeak Hemiramphus depauperatus is reasonable, the relationships of the endemics so known only from the Hawaiian, Line, and Mar- produced should exhibit their closest affinities quesas islands (Parin et al., 1980). The trunkfish with different elements of the biota on the Pacific Ostracion whitleyi is known only from the Ha- Plate, particularly southeast Oceania. Undiffer- waiian, Johnston, Marquesas, Tuamotu, and So- entiated elements of the integrated biota might ciety islands (Randall, 1972b). The gobiid Eviota also be shared only with southeast Oceania. Cla- epiphanes is known only from the Hawaiian, John- distic analyses of the Hawaiian biota are essen- ston, and Line islands (Lachner and Karnella, tially nonexistent, but one can note, with much 1980). The gobiid Kelloggella oligolepis is known interest, the congruence of Zimmerman's and van only from the Hawaiian and Easter islands (Fig- Balgooy's generalized conclusions, and Rehder's ure 20). distributional data, with an island integration Gomon (1979) recognized three subspecies for hypothesis. the labrid Bodianus bilunulatus: one endemic to the With regard to the littoral marine fishes, there Hawaiian and Johnston islands, one restricted to are several patterns of relationships: (1) species the Marquesas, Henderson, and Ducie islands in that support direct relationships between Ha- southeast Oceania, and one ranging from the east waiian Islands' endemics and non-Hawaiian Pa- coast of Africa to Japan, Philippines, Moluccas, cific Plate endemics; (2) apparently undifferen- New Guinea, and New Caledonia (hence, re- tiated species that the Hawaiian Islands share stricted to the Indo-West Pacific). If a cladistic only with islands to the south and east on the analysis of these three subspecies should indicate Pacific Plate, or near the Plate's southeastern that the Hawaiian-Johnston subspecies is the sis- margin; and (3) species groups whose individual ter group of the subspecies in southeast Oceania, species distributions have good potential for sup- or the sister subspecies to a sister subspecies group porting Hawaiian Islands-southeast Oceania formed by the subspecies in southeast Oceania relationships, pending cladistic studies of the and the Indo-West Pacific, the relationships groups. Examples follow. A few taxa of shorefishes would be congruent with the island integration are known only from the Hawaiian Islands and hypothesis. A sister-subspecies relationship be- relatively nearby Johnston Island. For the pur- tween the Hawaiian subspecies and the Indo- poses of my study I consider these species to be West Pacific subspecies would not be congruent Hawaiian endemics. Gosline (1955) considered with the island integration hypothesis. Johnston Island to be an outlier of the Hawaiian Under the family account of the Blenniidae, I fish fauna. have asserted the monophyly of three of the five The littoral epipelagic halfbeak Hyporhamphus species of Enchelyurus: E. brunneolus, endemic to the acutus (Figure 22) and the benthic, sand-dwelling Hawaiian Islands, E. ater, ranging from Lord creediid Crystallodytes cookei (Figure 60) each com- Howe Island east to the Tuamotus, and E. kraussi, prises two subspecies, and each is represented by ranging from the Red Sea and western Indian an endemic subspecies in the Hawaiian and John- Ocean east to the Marianas, New Guinea, and ston islands and an endemic subspecies that is Great Barrier Reef (Figure 10). I am unable to widely distributed among, and limited to, the decide on the sister-species relationships within other islands of the Pacific Plate. this group. If a future analysis supports a sister- Springer (1967a) considered the Hawaiian en- species relationship between E. brunneolus and E. demic blenniid Entomacrodus strasburgi to be most ater, or between E. ater and E. kraussi, the relation- closely related to E. chapmani, endemic to Easter ships would be congruent with the island integra- 134 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY tion hypothesis. A sister-species relationship in- TIME PERIOD volving E. kraussi and E. brunneolus would be in- compatible with the island integration hypothe- PSJ[£HJIP|SJ|S>!|- sis. Although the island integration hypothesis is V straightforward, cladistic relationships that indi- cate congruence with the hypothesis could be misleading, because the same relationships may also be congruent with an extinction (or other) hypothesis, particularly where the area clado- P S HI | P| S H| | P1 S |H|J gram includes two antiequatorial (disjunct) areas on the Pacific Plate. To illustrate the problems, the following discussion7 is presented and dia- v v grammed (Figure 61). HYPOTHESIS IPSHI IPSTHI fPislHM The island integration hypothesis (Figure 61: II Hypothesis I) presupposes that at Time Period 1 v v there was a uniform biota widely distributed on the Pacific Plate (Figure 61: P), or more widely, IPHSI IPW1 fPiHlSl including southeast Oceania (Figure 61: S), but absent from the Hawaiian Islands (Figure 61: H). IT v At Time Period 2, two possible vicariant events FIGURE 61.—Area cladograms for two hypotheses involving occurred, one of which is island integration (Fig- relationships of some Hawaiian Islands endemics (brackets on the left denote two hypotheses and embrace cladogram ure 61: Hypothesis la) and one of which is not possibilities; pointed brackets on right indicate homoclado- (Figure 61: Hypothesis Ib); in the latter case, grams; P = Pacific Plate islands less Hawaiian Islands and southeast Oceania is isolated from the remainder islands of southeast Oceania; S = islands of southeast of the Pacific Plate. In Time Period 3, according Oceania; S* = islands of southeast Oceania that have inte- to Hypothesis la, a vicariant event isolated south- grated with Hawaiian Islands; H = Hawaiian Islands; see discussion under "Hawaiian Islands and North-South Dis- east Oceania from the remainder of the Pacific junct Distributions on the Pacific Plate"). Plate, and according to Hypothesis Ib, island integration occurred. The area cladograms given waiian Islands. A third area cladogram (Figure at each time stage reflect the possible cladistic 61: lie) can be developed from this same initial relationships of the endemic taxa formed as a hypothesis that would result in a three taxa state- result of the vicariant events. ment that is not compatible with the island inte- It is possible to arrive at two three-taxa clado- gration hypothesis. In summary, there are five grams that are identical with those resulting from possible three-area cladograms that can be devel- island integration, but that are the result of a oped from the two initial hypotheses, and from different set of vicariant events (Figure 61: He these one can arrive at three three-taxa clado- and Hd). To arrive at these "homocladograms" grams, of which only two are congruent with the it is necessary to hypothesize that there was a island integration hypothesis. On what basis can uniform biota that was widely distributed on the one determine which of the cladograms of each Pacific Plate (or more widely), including the Ha- homocladogram pair is to be preferred (i.e., la or Hd; Ib or lie), or in other words, how can one decide whether the antecedents of (some) Ha- 71 am indebted to R.P. Vari for the details of this analysis. A general discussion of the use of area cladograms in bio- waiian Islands' endemics were part of an undif- geography is provided by Platnick and Nelson (1978) and ferentiated biota that included the Hawaiian Is- Nelson and Platnick (1981). lands and other Pacific Plate islands, or only NUMBER 367 135 other Pacific Plate islands (a continental area rica, Australia, New Zealand, Rapa, Pitcairn, could be added to, or replace Pacific Plate islands and Easter (thus antiequatorially distributed; at time period one)? Unfortunately, there is no W.F. Smith-Vaniz, pers. comm.), antiequatorial- unequivocal answer to this question. ity on the Pacific Plate may only be a secondary There is one other aspect of the distribution of reflection of antiequatoriality that developed ini- Hawaiian Islands' fishes that may also have bear- tially in continental areas (see also family account ing on north-south disjunct distributions: the of Isonidae for another example). "Hawaiian exception." I have used this term With the exception of the young Eastern Car- frequently in the family accounts to denote Indo- oline Island chain and Nauru and Ocean islands, Pacific taxa that occur nonmarginally on the high islands on the Pacific Plate, as noted earlier, Pacific Plate only at the Hawaiian Islands. These are distributed antiequatorially (Figure 2). For taxa usually occur also in Japan and/or the ecological reasons other than temperature, then, Ruykyu Islands, and many are antitropically dis- one might expect antiequatorial distributions of tributed in the Indo-West Pacific; some are re- organisms on the Plate. The distribution of the stricted to north of the tropics and some are extant members of the land snail family Endo- moderately deep dwelling. In the following ex- dontidae (Figure 48) duplicates closely the distri- amples an asterisk denotes species discussed by bution of high islands on the Plate. The presence Randall (in press b), but not in the family dis- of fossil endodontids of Miocene to Pleistocene cussions herein: Microcanthus strigatus (Scorpidi- age at Bikini and Midway atolls, where there are dae), Gonorynchus (Gonorynchidae), Sarda orientalisno extant endodontids, is evidence that these two (Scombridae), Kentacapros aculeatus (Aracanidae), islands were high during the times of endodontid Cirrhitops fasciatus (Cirrhitidae), Oplegnathus fascia- extancy (Solem, 1977). Schlanger (1963) pre- tus and 0. punctatus (Oplegnathidae), Histiopterus sented data in support of a statement written me acutirostris (Pentacerotidae), Malthopsis jordani, M. (Schlanger, in litt.) that "entire atoll groups in mitrigera, and Dibranchus stellulatus (Ogcocephali- the Pacific were high limestone island groups in dae), Myripristis chryseres* and Ostichthys archiepis- Miocene and perhaps Eocene time as well as in copus* (Holocentridae), Seuzichthys species (Labri- Pleistocene time." The land snail data are con- dae). These species probably dispersed from the gruent, therefore, with an hypothesis of extinction Japanese area to the Hawaiian Islands via eddies as an explanation for north-south disjunct distri- from the Kuroshio extension, or in the case of the butions on the Plate.8 some of the deeper dwelling forms, possibly by Before concluding discussion of north-south dispersal northward from Japan along continen- disjunct distributions I wish to call attention to tal coasts to the Emperor Seamounts and down the special case of antiequatorially distributed the Emperor Seamounts to the Hawaiian Islands. high-seas organisms. Ralston (1981) noted that the common southern Cheng (1973) and Cheng and Shulenberger Japanese angelfish, Centropyge interruptus, is other- (1980) detailed the distribution of the five high- wise known only from Kure, northern and west- seas species of the surface-dwelling insect genus ernmost of the Hawaiian Islands. He made ref- Halobates (comprising 42 species). One of the five erence to unpublished studies of E.S. Hobson and species, H. sericeus, which is confined to the Pa- L.R. Taylor that indicated that the fish fauna of the northern Hawaiian Islands showed closer re- 8 One can wonder at the biogeographical significance of lationships to the Japanese fauna than to that of the fact that the two fossil endodontid genera that occurred the southern Hawaiian Islands. In a case like at the geographically somewhat intermediate Bikini Atoll Pseudocaranx dentex (Carangidae), which in the were not contemporaneous, and that each of the genera is extant in a different island group: the lower Miocene Cooke- Indo-Pacific portion of its distribution is known concha is extant only in the Hawaiian Islands; the Pliocene- only from Japan, Hawaiian Islands, southern Af- Pleistocene Minidonla is extant only in southern Oceania. 136 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY cific, is antiequatorially distributed (Cheng and explanation for the endemism has been proposed, Shulenberger, 1980, fig. 2c). In the Northern although the extreme geographic isolation of the Hemisphere, H. sericeus ranges along a broad island is usually mentioned as an implied expla- swath from Taiwan and Japan to Mexico, and in nation. In the following discussion I will propose the Southern Hemisphere, it ranges along a broad a geotectonic hypothesis for Easter Island ende- swath from Australia to off Chile. Cheng and mism. Shulenburger noted that the distributions of H. The geology and tectonics of Easter Island and sericeus and three of the other high-seas species of nearby Sala-y-Gomez were most recently dis- Halobates conform with the dominant oceanic sur- cussed by Clark and Dymond (1977) and Bonatti face current patterns, and in the case of//, sericeus, et al. (1977). Much of the following discussion is with two subtropical anticyclonic gyres. Numer- based on geological information contained in ous species of midwater zooplankton (Reid et al., these two studies. Most of the edifice ages I cite 1978) and fishes (R.H. Gibbs, Jr., pers. comm.) were published prior to 1977, and were compiled have similar distribution patterns to that of H. and assessed by Jarrard and Clague (1977). sericeus (and the other three oceanic species of Easter and Sala-y-Gomez rest on the Nasca Halobates). Restriction of the zooplankton and Plate, which is forming at the East Pacific Rise fishes to these anticyclonic gyres is believed to be and moving eastward to a subduction zone along dependent on the physicochemical characteristics the west coast of South America (Figure 1). Easter of the water. Whether the distribution of H. and Sala-y-Gomez are the westernmost edifices of sericeus is similarly limited is not known. No mech- an east-west oriented chain of edifices (Easter anisms have been proposed to explain the distri- volcanic chain) that terminates 2400 km east with bution of H. sericeus, but those that have been San Felix and San Ambrosio islands off the coast proposed to explain the distributions of the other of Chile. There are no islands between these two organisms are different from those so far discussed pairs of islands, but there is a fairly dense band by me for the shorefishes (and implied by me for of seamounts between them ^many of these sea- Halobates hawaiiensis; see early part of discussion mounts appear on Figure J, but more complete in this section). Brinton (1962) suggested that a representation can be found in Mammerickx et 2.5° C ocean-wide cooling (at 200 m depth) al., 1975, fig. 1—a bathymetric chart of the east- would permit southern gyre euphausids to trans- central Pacific). Easter lies about 250 km east of gress the equatorial regions in the eastern Pacific the East Pacific Rise in an area where the spread- (only) and become established in the northern ing half-rate of the ocean floor is the fastest known gyre. Subsequent warming would isolate certain (about 10 cm/yr; Herron, 1972). The ocean bot- organisms in the two gyres, resulting in antiequa- tom on which Easter rests is in the region between torial distributions. Reid et al. (1978) believed magnetic anomalies 2 and 3, and would be about that the southern gyre mixes with the equatorial 3 m.y. old (Heirtzler et al., 1968, fig. 3). Easter current system in the eastern Pacific (only) and Island cannot, therefore, be very old, and the age permits southern gyre organisms to enter the of its rocks confirm this. Subaerial volcanism at northern gyre. Because these organisms are Easter occurred during three periods: 2.5, 0.9, adapted to the physiocochemical parameters of and 0.4 m.y. ago. The spreading rate taken to- the gyres, it can be inferred that they cannot gether with the earliest date for subaerial Easter maintain populations in the convergence areas. I Island volcanism suggest that Easter Island was do not believe that either of these two mecha- formed at or adjacent to the East Pacific Rise (0.1 nisms can be used to explain shorefish distribu- m/yr X 2.5 m.y. = 250 km, the distance between tions. Easter Island and the crest of the East Pacific 9 EASTER ISLAND.—The first general acknowledg- Rise today). ment that high endemism might be characteristic of Easter Island is that of Briggs (1974). No Jackson et al. (1972) summarized available data on time NUMBER 367 137
Sala-y-Gomez rests on ocean bottom in the Island endemism, I hypothesize that islands have region of magnetic anomaly 5, which has been been appearing on the Nasca Plate near the East dated at about 9 m.y. Only a few rocks from Sala- Pacfic Rise at the latitude of Easter Island for y-Gomez have been dated, and these provide ages several million years. As erosion and subsidence of 1.3 and 1.9 m.y. While the latter age is consid- drowned these islands, elements of their biota ered a minimum for the island, and is based on were able to disperse to a nearby, newly formed a dredged sample, the former is based on sub- island. aerial samples and believed to indicate an average The edifice chain including Ducie Atoll rests age for subaerial volcanism at Sala-y-Gomez. The on the Pacific Plate and appears to commence spreading half-rate at anomaly 5 is only 5 cm/yr; about 300 km east of Ducie (or about 1200 km thus, Sala-y-Gomez was probably formed close to west of the East Pacific Rise), where a pair of its present location. seamounts is emplaced (Mammerickx et al., 1975, There are no dated rocks for San Ambrosio fig. 1) in the vicinity of magnetic anomaly 6 (20 and San Felix, but volcanic activity at San Am- m.y.). Between these seamounts and the East brosio last occurred in 1922, and sheer cliffs at Pacific Rise the seafloor appears to be devoid of San Felix are believed to indicate that it is at volcanic edifices (Figure 3), indicating that a most only a few million years old. These two volcanically quiet period has existed in this region islands are, thus, near or at their location of of the Pacific Plate for about 20 m.y. There is a origin. Only three of the over 50 seamounts be- paucity of data for this region, however, and a tween Sala-y-Gomez and San Felix have been remote possibility exists that other seamounts are dated: 8.0 m.y. (located presently about 1400 km present (Epp, 1978). east of the crest of the East Pacific Rise on ocean Ducie and Henderson atolls, the next edifices bottom with an age of 20 m.y.); 1.7 m.y. (1500 west of the paired seamounts, and the paired km east of the Rise, also on ocean bottom about seamounts, have not been dated. Ducie and Hen- 20 m.y); and 30 m.y. (about 2600 km east of the derson are emplaced on ocean floor about 20-27 Rise on ocean bottom about 30 m.y.; therefore, m.y. in age, and must be much older than Pit- this last seamount quite possibly originated near cairn Island, the nearest aged edifice, which is the crest of the East Pacific Rise). little, if any, older than 0.95 m.y. (Duncan et al., It is apparent from these data that edifices in 1974) and rests on ocean bottom about 30 m.y. in the Easter volcanic chain were not propagated age. Few Pacific atolls have been dated but those from a single fixed mantle plume (hot spot), as that have are usually more than 10 m.y. old, and were the Hawaiian Islands (however, see Rotondo often more than 37 m.y. old (Jarrard and Clague, et al., 1981, for exceptions). Two or more mantle 1977). Mururoa Atoll, in the northern Tuamotus, plumes have been active, and edifices have been appears to be an exception, and has been aged at propagated intermittently for at least 30 m.y. 7 m.y. Schlanger and Gillett (1976) have shown Some, if not many, of the edifices (including some that it takes about 15 m.y. for a high island in at each end of the chain) were islands that origi- the Hawaiian chain to become an atoll, and this nated near the crest of the East Pacific Rise. time period is probably reasonable for such con- For the purposes of this discussion of Easter versions in other island chains. High islands are, expectedly, relatively young, with ages for those required to form Hawaiian volcanic shields: "The available dated ranging generally from less than 1 m.y. to evidence strongly suggests that the tholeiitic phase of volcan- ism on individual shields in the Hawaiian-Emperor chain about 12 m.y. Exceptionally, Tubai, a high island may have a total lifetime of only 0.5 to 1.5 m.y., and the in the Austral chain, has been dated at about oldest K-Ar ages on subaerial tholeiitic flows . . . will be only 10.9±l to 24.9±10 m.y., but there is some ques- slightly younger than the date of birth of these volcanoes." tion about the data, and Mangaia, a high island I have assumed that the rate of Easter volcanism is probably in the Cook Islands, has been well dated at about commensurate with that of the Hawaiian volcanoes; the age- distance data are reasonably congruent with this assumption. 18 m.y. Mangaia has been greatly uplifted, and 138 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY the normal erosional processes and subsidence formation on the Pacific Plate adjacent to the that would have converted it into an atoll or same vicinity ceased. Westward movement of the seamount have been offset (although it probably Pacific Plate gradually increased the distance existed as an atoll in the past). between Pacific Plate islands and islands of the The aged edifice nearest to Pitcairn is in the Easter volcanic chain until interchange between high Gambier Islands group (4.7-7.1 m.y.) of the these two groups of islands was no longer possible. southern Tuamotu Archipelago. This archipel- The resulting isolation of the more western islands ago, which otherwise consists almost entirely of of the Easter volcanic chain allowed many species atolls formed on a rise, arches northwest from in these islands to diverge, accounting for the Pitcairn Island and is continuous with the Line high degree of endemism that we recognize today. island chain (together called the Tuamotu-Line Within the Easter volcanic chain, the islands Rise by Morgan, 1972), apparently following an that appeared in the more central parts of the earlier pole of rotation (direction of movement) chain permitted dispersal of biotic elements to of the Pacific Plate. Morgan (1972) believed that islands in the eastern portion of the chain. Sub- the Tuamotu-Line Rise and the Sala-y-Gomez- sequent drowning of these more central islands Nasca Rise originated contemporaneously from resulted in isolation of the easternmost islands, the same hot spot on the crest of a spreading rise permitting elements of their biota to diverge (see (East Pacific Rise). Morgan's hypothesized origin following discussion). Because the present islands of the Tuamotu-Line and Sala-y-Gomez-Nasca at the eastern end of the chain are young, they rises has not survived (Bonatti et al., 1977; Clark must have derived their endemic elements from and Dymond, 1977), but the contemporaneity now drowned islands. There are numerous sea- and juxtaposed origins of the Tuamotu and mounts in the vicinity of San Felix and San Nasca portions of the rises on opposite sides of the Ambosio (Mammerickx et al., 1975, fig. 1) that East Pacific Rise at about the latitude of Easter possibly served as biotic suppliers to these young Island still seems probable (H.W. Menard, pers. islands. One particularly shallow seamount comm., April 1981), indicating that littoral con- (hence, possibly drowned relatively recently) is ditions existed on the rise contemporaneously shown in Figure 2). with littoral conditions in the Tuamotus, and The fishes and mollusks of Easter Island were ocean bottom ages in the two areas cover about discussed by Randall (1976) and Rehder (1980). the same time span. These two groups of organisms exhibit 27 and 42 Based on the previous discussion, I propose the percent endemism, respectively. Although there following scenario to account for Easter Island are no cladistic studies of these organisms avail- endemism (a similar, but generalized scenario for able, the general relationships of both groups are Pacific island plants was proposed by Axelrod, clearly with the fishes and mollusks of the central 1972). and western Pacific (see discussion of the Ha- Until about 20 m.y. ago, there was a common waiian biota in the previous section). biota inhabiting the islands of southeast Oceania The mollusks and fishes of Ducie Atoll were on both sides of the East Pacific Rise at about reported by Rehder and Randall (1975). These the latitude of Easter Island. The uniformity of two groups are represented overwhelmingly by this biota was maintained in the face of diverging species that also occur farther west. A few of the oceanic lithospheric plates (seafloor spreading) species are restricted to southeast Oceania, and and the subsidence and drowning of islands on some of these range to Easter Island. each side of the East Pacific Rise by the periodic Nothing is known about the fishes of Sala-y- formation of new islands within biotic dispersal Gomez, and only three mollusk species have been distance on each side of the Rise. Island formation reported from there, all common at Easter Island on the Nasca Plate in the vicinity of Easter Island (Rehder, 1980). On the basis of nearness and has been continous, but about 20 m.y. ago, island small size of the island (maximum length 700 m, NUMBER 367 139 maximum width 400 m), one can expect that the Easter Island species occurs otherwise only in biota of Sala-y-Gomez will be closely allied to, southeast Oceania, including Pitcairn Island). and a subsample of, that of Easter Island. Russell (pers. comm.) beieves that a Rapa Island The closest islands to the east of Sala-y-Gomez endemic species of Pseudolabrus is the sister species are San Felix and San Ambrosio, which are 2400 of the Easter Island endemic. The distribution km distant. I know of no faunal reports devoted and supposed relationships of the Pseudolabrus spe- to the mollusks or fishes of these two islands, cies, thus, seem to support the scenario developed although various revisions include mention of above to account for Easter Island endemism, selected San Felix fishes. The most comprehensive and endemism at San Felix (and, perhaps, Juan collection of San Felix fishes (includes some types) Fernandez) Island. resides at Scripps Institution of Oceanography. In closing this discussion of Easter Island ende- R.H. Rosenblatt provided me with a listing of mism, I wish to reiterate the fact that nine percent the 31 species (not all identified to species) in the of the Easter Island mollusks and several species San Felix collection. As best I can determine, well of the island's fishes are found otherwise only in over half of these species appear to be either the Hawaiian Islands. I explained this distribu- endemic to San Felix or to San Felix and the tion pattern by invoking island integration (Ro- Juan Fernandez Islands,10 situated to the south tondo et al., 1981; see discussion of Hawaiian of San Felix (Figure 2). About six of the species Islands). Island integation and the scenario pro- also occur on the mainland coast of the Western posed here to account for the high degree of Hemisphere, and a few others range widely in the endemism at Easter Island are compatible.Taken southern Pacific. The overall relationships of the together, they serve to demonstrate the complex- fishes seem to lie more closely with western Pacific ity of geotectonic events in the Pacific Basin and genera and species than with eastern Pacific gen- how these events can complicate and contribute era and species. Greenfield and Woods (1980: to the resolution of biotic distributions involving 633-634) noted the same general geographical a few remote and well separated islands. direction of the relationships for San Felix-Juan MARQUESAS ISLANDS.—The Marquesas Islands Fernandez fishes, based on the few available pub- are a short (about 350 km long), relatively young, lished references to these fishes and their own high-island chain increasing gradually in age study of Chromis meridiana (Pomacentridae). Quite from about 1.3 m.y. for Fatu Hiva at its south- possibly, the strong, cold, northward flowing Peru eastern end (Duncan and MacDougall, 1974) to Current along the west coast of South America about 5.1-8.7 m.y. for Eiao at its northwestern prevents the eastward dispersal of San Felix-Juan end (Brousse and Bellon, 1974; because of prob- Fernandez fishes to the South American coast. lems with argon enrichment, these authors pre- The labrid fish genus Pseudolabrus is known ferred an average date of 6.3 m.y., but other from the coasts of Japan and China in the North- authors citing their work—e.g., Jarrard and Cla- ern Hemisphere, and from southeast Australia gue, 1977, Epp, 1978—do not). The progression east to San Felix and Juan Fernandez islands in rate of the chain is about 10 cm/yr, so a presumed the Southern Hemisphere (B.R. Russell, pers. hot-spot origin of the chain would probably be comm.). The single Pseudolabrus species at San not more than 300 km southeast of Fatu Hiva Felix and Juan Fernandez islands is endemic (Brousse et al., 1978, fig. 6), and perhaps as little (Russell and Randall, 1980), as is one of the two as 12 km southeast (Duncan and MacDougall, Pseudolabrus species at Easter Island (the other 1974). Although there is no question about the youth of the Marquesas, the alignment of the chain, relative to alignments of the Hawaiian- 10 In the Juan Fernandez Islands, Mas-a-tierra has been Emperor chains, would indicate that the Mar- dated at 2.0-3.9 m.y. and 6±3 m.y., and Mas-afuera at 0.87- quesas were initiated more than 68 m.y. ago 1.3 m.y., with two, apparently radiogenic Ar enriched, dates of 13 and 19.6 m.y. (Ferrara et al., 1969). (Epp, 1978). For this reason, Epp conjectures that 140 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY the Marquesas may be a recent group that took to exhibit the highest endemism for fishes and its alignment from a propagating fracture system mollusks of any island or island group in Oceania. that formed more than 68 m.y. ago, rather than This high endemism is unexpected in view of the from a hot spot. The Marquesas alignment, as fact that the Marquesas are less isolated geo- would be expected in such a case, is perpendicular graphically than several other Pacific Plate is- to the Marquesas Fracture Zone and the south- lands that exhibit little or no endemism, for in- eastern end of the islands almost abuts the Zone stance, Wake Island, Johnston Island (which, (Mammerickx et al 1975, fig. I11), which tends to however, shares some endemics with the Ha- support Duncan and MacDougall's (1974) plac- waiian Islands, but has none of its own), Rapa ing the origin of the chain only 12 km southeast. and Marotiri (together), Maiden and Starbuck, Such an origin would make Brousse and Bellon's and Howland and Baker; Rapa and Marotiri, 5.1-8.7 m.y. age for Eiao about 1.6-5.2 m.y. older among these, are the only high islands. than expected (350 km length + 10 cm/yr prop- There are other unexpected conditions existing agation rate = 3.5 m.y., probable age for Eiao). in the Marquesas. In spite of the fact that the The Marquesas are separated from their near- islands are situated between 07°S and 12°S lati- est island neighbors, in the Tuamotus, by about tude, well within the tropics, no marine groups 500 km. There are no seamounts that could have are particularly well represented there. Coral been littoral or subaerial and served as dispersal reefs, which surround the other tropical islands of stepping stones between the Marquesas and any the Indo-Pacific, are reduced and present only in other islands. Even if the Marquesas were 300 km protected bays in the Marquesas (Chevalier, southeast of their present position at some time 1978). The lack of reefs undoubtedly limits the between 5.1 and 8.7 m.y. ago, and the Tuamotus variety of the reef-associated biota. Brousse et al. are rotated (by eye) back up to 300 km along the (1978) discussed possible causes for this poor reef Tuamotu alignment, the Marquesas do not ap- development, including those proposed by earlier pear to approach the Tuamotus more closely than authors. They were able to discount the impor- they do today. The geographical isolation of the tance of all proposed explanations, which in- Marquesas, thus, appears to have remained the cluded the following: rapid subsidence; absence same during their entire existence. of a shallow, pre-existing platform (necessary for Concerning the marine biota, the only list of colonization); cold water temperatures (surface Marquesas fishes is that of Plessis and Mauge waters range from 25.2° to 29.4°C, depending on (1978), which is uncritical, incomplete for the time of year); hydrodynamic forces (heavy wave period covered, and contains numerous errors. action); terrigenous deposition; and geographic Randall (1978a), based on his own collecting, isolation. They concluded that the reasons for estimated that 10 percent of the Marquesas fishes poor reef development were unknown and pro- are endemics, and my own studies of the blenniid posed that the solution to the problem probably genera Entomacrodus and Alticus (Springer, 1967a; concerned physiochemical properties of the local Smith-Vaniz and Springer, 1971) also indicate a seawater, for which they lacked information. high degree of endemism. There are no compila- The general relationships of Marquesas fishes tions of Marquesas mollusks, but Rehder (1968) can be assigned to the Indo-Pacific, to which estimated that 20 percent of the Marquesas mol- region many of the Marquesas genera and species lusks are endemics. Next to the Hawaiian Islands are restricted. Formal cladistic relationships of and Easter Island, the Marquesas, thus, appear the Marquesas endemics have not been proposed, but these relationships are clearly with other spe- This figure indicates that the fracture zone is separated cies on the Pacific Plate. For instance, according into eastern and western segments with a broad discontinuity to Randall (1978a), "the fiery red angelfish Cen- between the segments. The Marquesas abut the region of tropyge loriculus [a widely distributed Pacific Plate discontinuity. endemic] has lost its dark bars [which character- NUMBER 367 141
ize the species at other Pacific Plate localities]," For an oceanographical scenario, I propose the and Springer (1967a) considered Entomacrodus ma- following: past local ocean currents (or eddies) crospilus, a Marquesas endemic, to be most closely that permitted biotic dispersal from the Tuamotu related to E. thalassinus, which is widely distrib- region to the Marquesas were gradually modified uted in the Indo-Pacific, but absent from the (deflected) and eliminated as the Marquesas in- Marquesas (these two species share two speciali- creased in linear extent. It might be possible to zations, synapomorphies, in Entomacrodus: absence test such a proposal by preparing a scaled model of lip crenulae, smallest maximum size) .Two of the Pacific in the Tuamotu-Line islands region other Marquesas endemic species of Entomacrodus without the Marquesas, applying a fan-diven (E. randalli, E. comeliae) appear to be most closely wind to simulate present-day wind circulation in related to other Indo-Pacific species on the Pacific the region, and then gradually adding the Mar- Plate that are absent from the Marquesas. With- quesas Island lineation to see what effect it would out pursuing this line of evidence further, I would have on the water movement. Surface currents propose that the ancestors of the Marquesas en- around the Marquesas (Sverdrup et al., 1942, demics were widely distributed and included in chart 7) indicate that present-day transport is their range both the Marquesas and other Pacific generally from the Marquesas to the Tuamotus, Plate islands. A vicariant event occurred that thus isolating the former against "infection" from isolated the Marquesas from the other islands on the latter, but, seemingly not the latter from the the Pacific Plate and allowed divergence of (at former. The coarse, general rendition of surface least) the Marquesas populations. currents on charts can be misleading. For in- It is difficult to decide the nature of the vicar- stance, localized, but persistent, eddies (Royce, iant event that gave rise to Marquesas endemism. 1978) can provide countercurrent transport, as Unlike the Hawaiian Islands and Easter Island, can repeated, seasonal changes in current pat- where geological information allows plausible vi- terns. It would be desirable, therefore, to have cariance scenarios, the Marquesas do not appear detailed information on water transport in the to have been isolated by a geological event, at Marquesas region. least not in the same way as were those two other The alternative to a vicariant event as an island groups. While there are probably several explanation for Marquesas endemism is founder ways that could be proposed to isolate the Mar- principle (Mayr, 1942; critically reviewed by quesas, I will present only two here, one geologi- MacArthur amd Wilson, 1967, who combined it cal, the other oceanographical; the first is, per- with founder effect). In a founder principle scen- haps, less likely. ario, numerous different species would have in- The existence of a former intra-Pacific Plate dependently dispersed to the Marquesas, each as spreading ridge (Tamaki et al., 1979) and an a small propagule. Selection on descendants of inactive intra-Plate fracture system (Farrar and each of these propagules would have had to op- Dixon, 1981) have been proposed (the fracture erate rapidly in order that subsequent dispersal system presumably split the Pacific Plate into from the parent population of each species would eastern and western halves that underwent differ- not dilute the gene pool of the founding propa- ential offsets). The ages of these two features are gule's descendants. Additionally, the newly too old, more than 60 m.y., and their positions evolved taxa would have to competitively exclude too removed to have affected the Marquesas Is- subsequent dispersals from the parental popula- lands. If, however, younger, similar features tion, otherwise both members of each pair of should be found to have existed between the sister taxa would occur together, at least occasion- Marquesas and the Tuamotus, either type feature ally. There is no evidence that the sister species of could have resulted in an increase in distance any Marquesas Island endemic occurs in the between the two island groups and effected their Marquesas. For that reason and because of the isolation. repeated complex, multiple conditions necessary 142 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
-190 MY -135 MY
»/ (1 // If (1 /I II
i Ikj /) FARALLON = —"1 KULA \j'' M 1 1. PHOENIX "*
hr* "•-,,'•
Fig. 3. (a) Plate, ridge, transform fault, and subduction zone locations in the Western Fig. 4. Plate, ridge, transform fault and subduction zone locations at —135 m.y. Pacific at —190 m.y. (b) Pacific plate origin at about —185 m.y. Features shown in this figure and Figs. 4—7, except for Asia and Antarctica, are schematically located on a Mer- -65 MY cator projection in their paleopositions as determined from paleomagnetic data. Because of the complex and less certain paleomagnetic determination for Asia and because Asia and Antarctica have apparently been subject to relatively little movement for most of the period of our reconstruction (—190 m.y. to present) when compared to the other plates involved, we have assumed they have remained fixed in their present positions.
-100 MY
Fig. 6. Plate, ridge, transform fault and subduction zone locations at —65 m.y. The —135 m.y. and —100 m.y. magnetic lineation and fracture zone pattern of the Pacific plate is shown by short dashed lines.
-25 MY
Fig. 5. Plate, ridge, transform fault and subduction zone locations at —100 m.y. The ASIAN PLATE —135 m.y. magnetic lineation and fracture zone pattern of the Pacific plate is shown by short dashed lines.
FIGURE 62.—Hypothesized orgin and development of the Pacific Plate (from Hilde et al., 1977, figs. 3-7). Apices of triangles indicate direction of subduction; double lines in- dicate spreading ridges, associated arrows indicate direction of spreading; straight lines indicate transform faults, associ- ated arrows indicate relative movement on either side of fault; open arrows denote direction of plate movements.
Fig. 7. Plate, ridge, transform fault, and subduction zone locations at —25 m.y. The —135 m-y-. —100 m.y., and —65 m.y. magnetic lineation and fracture zone pattern of the Pacific plate and other inactive features are shown by short dashed lines. NUMBER 367 143
to establish and evolve each of the numerous spreading rise of the Pacific Plate (Figure 62: fig. Marquesas endemics under a founder principle 5 shows this offset rise due east of Borneo) and is scenario, I believe a vicariant explanation for the most recognizable by a bend in the Hawaiian- high degree of Marquesas endemism is preferable Emperor volcanic chain. Similar bends, although to one based on founder principle. less obvious than the Hawaiian-Emperor bend, GEOLOGICAL HISTORY OF THE PACIFIC AND PHIL- presumably appear in island chains that were LIPPINE PLATES AND ADJACENT MARGINS AS RE- being actively constructed more than 43 m.y. ago LATED TO FORMATION OF TYPES 1 AND 2 ENDEM- and continued forming after that time (this model ICS.—The origin and general development of the has been modified by Farrar and Dixon, 1981, Pacific Plate, particularly as pertains to the tec- who proposed that the Pacific Plate was formerly tonically complex western portions, has been split into eastern and western portions, and that summarized by Hilde et al. (1977), and their rotation affected only the western portion and account distilled by Uyeda (1977). A schematic volcanic edifices, such as the Emperor chain, on presentation of their ideas is contained in Figure that portion). Volcanic edifices have also ap- 62, which reproduces figs. 3-7 in Hilde et al. peared irregularly as the result of magmatic (1977; the same figures, except for the first, are release along intraplate transform fault or frac- also reproduced in Uyeda, 1977). These authors ture zones and at the East Pacific Rise. There hypothesized that the formation of the Pacific was also an unexplained episode of massive Cre- Plate commenced about 190 m.y. ago, prior to taceous volcanism over an extensive area of the the breakup of Gondwanaland. The oldest extant Pacific Plate that produced numerous edifices, ocean bottom representing the Plate, however, is most of which are now drowned (for instance, the from the Upper Jurassic (Figure 4), and dates Mid-Pacific Mountains); some of the atolls of the from about 160-140 m.y. ago. Older, non-extant Marshall and Line islands probably orginated portions of the Pacific Plate are believed to have during this episode (Schlanger and Premoli-Silva, been subducted below the continental areas that 1981). border the Plate's western margin. The propelling As the Pacific Plate moves, it gradually sub- force that produces the northwestward movement sides into deeper water and subducts adjacent of the Plate (Figure 1) from its active area of plates at deep trenches along its western margin. formation (East Pacific Rise) is not known. As the subducting Plate approaches the mantle Movement of the Plate, volcanism, transform under adjacent plate margins, it remelts and faults, and subduction are major contributors to induces melting in the supra-adjacent mantle, the topography of the Plate and areas along its which gives rise to andesitic volcanism on the western margin (Figure 3). Some island chains on margins of the obducting plates (Arculus, 1981). the Pacific Plate appear to be ascribable to melt- The western margin of the Pacific Plate, hence, ing anomalies (hot spots, mantle plumes) in the delineates the Andesite Line of biogeographers aesthenosphere below the Plate. The hot spots are and geologists (for an historical discussion of the relatively fixed in position, and as the Plate Andesite Line see Appendix 1). Andesitic volcan- moves, intermittent, basaltic volcanism occurs ism is the source of island arcs on the eastern above the hot spots and gives rise to island chains margins of obducting plates. For unclear reasons, (the magma forming the Plate at the East Pacific an island arc tends to divide along its longitudinal Rise is also basaltic). A change in the direction of (volcanic) axis. The portion closest to the margin movement (from NW to WNW) of the Plate of the plate moves opposite to the direction of (which occurred about 43 m.y. ago), is associated subduction, leaving a sunken, inactive (fossil, with subduction along the margin of the Asian back arc) ridge behind the volcanically active Plate of a spreading rise that presumably existed frontal arc. Between the back arc and frontal arc as a western offset extending from the southern is a widening sea basin. Hence, part of an island 144 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY arc that formed adjacent to a continental plate Philippine Basin subducting) along the Philip- margin may, with time, become increasingly dis- pine Islands lineament. Sea-floor spreading about tant from any land mass on the plate. the Central Basin Ridge (Figure 63) to the west The account of the Philippine Plate (also of the Palau-Kyushu Ridge, occurred prior to known as the Philippine Sea Plate) that follows entrapment of the West Philippine Basin and is distilled mostly from information contained in continued for a short while after entrapment. The Hilde et al. (1977), Shih (1980), and Scott et al. West Philippine Basin is believed to have moved (1981). north about 15°-20° and rotated clockwise about Late in the Paleozoic, before the formation of 5O°-7O° during the past 35-40 m.y. (Shih, 1980). the West Philippine Basin (Figure 63), subduc- The West Philippine Basin and the basins and tion presumably occurred along the eastern side islands arcs to the east form the Philippine Plate of the Asian Plate giving rise to the Ryukyu Arc. (Figure 63; often called the Philippine Sea Plate). Except for some Paleocene to early Oligocene The formation of the eastern basins have a con- igneous activity in the Ryukyu Arc, no volcanism siderably different history from that of the West occurred along that portion of the Asian Plate Philippine Basin. After formation of the Palau- boundary until late Miocene, when spreading Kyushu Ridge in middle Oligocene, volcanism began in the Okinawa Trough behind (west of) continued on the active Ridge until early Mio- the Arc (the Arc and Trough are not labelled on cene, terminating about 29 m.y. ago, when the Figure 63). In the Mesozoic (Upper Cretaceous, Ridge sundered along its volcanic axis. The west- Hilde et al., 1977; see their figs. 5 and 6 on my ern segment of the sundered ridge became the Figure 62) subduction presumably continued and inactive Palau-Kyushu Ridge, and the eastern consumed the proto-Pacific Plate along the Asian segment (fore-arc) became the precursor of the continental margin. Early in the history of the West Mariana Ridge. Back-arc-basin spreading Pacific Plate it is believed that there were a series commenced between the two portions of the sun- of transform faults extending north-south just dered ridge driving the fore-arc eastward and west of the Pacific and Kula plates (Figure 62: opening the Parece Vela Basin. The Shikoku fig. 4). These transform faults intersected the Basin opened simultaneously, and it and the Pa- spreading rise that was continuous (with offsets) rece Vela Basin are considered to be sections of a with the spreading rise forming the southwestern single, large marginal basin (Karig, 1975). There margin of the Pacific Plate. The two easternmost is no evidence of volcanism in the area for the of these transform faults bordered a region of period 29-20 m.y. ago. About 20 m.y. ago, island- ocean bottom that was to become the West Phil- arc volcanism began again, immediately behind ippine Basin of the Philippine Plate (Figure 63). the fore-arc, and formed the West Mariana island Subduction along the margin of the Asian Plate arc (ridge). of the spreading rise intersected by the transform Formation of the Parece Vela Basin continued faults occurred about 43 m.y. ago (noted above) until sometime between 18 and 14 m.y. ago. and is associated with the change in direction of About 11 to 9 m.y. ago, arc volcanism on the movement of the Pacific Plate and spreading West Mariana Ridge subsided, and roughly 5 behind the Oki Daito Ridge (Figure 63). This m.y. ago sundering of the Ridge formed the change in movement appears to have converted Mariana Trough and eastwardly displaced the the easternmost of the two transform faults into new West Mariana fore-arc region, which even- a subduction zone (Pacific Plate subducting) that tually became the present Mariana Island Ridge. resulted in the formation of a rise (Palau-Kyushu Subsidence of the Parece Vela Basin and the Ridge, Figure 63), which entrapped the ocean West Mariana Ridge occurred at the same time bottom to the west (West Philippine Basin). The as the Mariana Trough was spreading, and con- westernmost of the two transform faults was sim- tinues to spread. In the Pliocene, volcanism began ilarly converted into a subduction zone (West along the Mariana Ridge and has continued to NUMBER 367 145
j Rldgx tall
FIGURE 63.—Topographic features in the Philippine Plate area (redrawn from Scott et al., 1981, fig. 1). the present. The Iwo Jima Ridge is a northern ago (Karig, 1975). The Iwo Jima Ridge is pres- extension of the West Mariana and Mariana ently in an early stage of sundering, ridges, and it takes its origin from the combined It can be surmised from the geologic history of Iwo Jima-Palau-Kyushu ridge of 25 to 18 m.y. the Marianas that these islands had their origin 146 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY on the Palau-Kyushu Ridge about 40 m.y. ago the eastern distributional limits, in this region, of and over 1000 km west of their present-day loca- some otherwise continental plate limited taxa (for tion. Sea-floor spreading that occurred after for- example, Terapontidae and Uranoscopidae, Fig- mation of the Palau-Kyushu Ridge, clockwise ures 43 and 46). rotation, and considerable northward movement The geotectonic history of the islands along the of the West Philippine Basin, probably indicate Pacific Plate margin extending from New Guinea that the Marianas originated south and even to the Tonga Islands is highly complex, and much further west, probably close to the Indonesian- remains to be explained. These islands have been Philippine arc systems (Karig, 1975). Species of termed the Melanesian Borderlands by Coleman continental origin would, consequently, have had and Packham (1976), who attempted a regional a shorter distance to disperse to colonize the geological synthesis. The discussion here is mainly Marianas. If other (now subducted) islands ex- limited to that portion of the margin extending isted on the Pacific Plate adjacent to the proto- from the New Hebrides to the Tonga islands. Marianas, these in turn could have been colo- Most of the Type 2 endemics (see discussion of nized and led to the colonization of other Pacific species endemism) I report from the Pacific Plate Plate island groups. Later distancing, and asso- are limited to the islands ranging east from the ciated rotation, of the Marianas created an is- Tonga Islands to Easter Island. If one presumes landless void between the Marianas and the con- that the Type 2 endemics in the Polynesian area tinental coasts to the west. The void could have of today have sister taxa whose distributions ex- isolated the Pacific Plate in this region, at least, tend westward from the Pacific Plate margin and contributed to subsequent divergence of the toward the Australian region, then one must also Plate biota (or the formation of Type 1 endemics). presume that the common ancestor of the two Myers and Shepard (1981) additionally attribute sister taxa had a distribution that included, at present-day isolation of the Marianas Islands least, contiguous portions of the ranges of both from the continental coast to the prevailing west- sister taxa. A logical search for the vicariant event ward flow of ocean currents in the region, making that divided the ancestral population would, the islands net exporters of fish larvae, perhaps then, be expected to have occurred in the general accounting for the western distributional limits in region of Australia (New Hebrides to Tonga is- this region of taxa otherwise restricted to the lands). The same vicariant event could enforce Pacific Plate. the endemism of the widely distributed Pacific It is because of the integral association and Plate endemics (Type 1). development of the Philippine and Pacific plates With the possible exception of its north coastal that I treat distributions of Pacific Plate species portion, which may be an accreted arc segment, having their western limits along the western New Guinea has always been part of the Indian- margin of the Philippine Plate as Pacific Plate Australian Plate, but the remaining islands of the distributions.12 In general, I believe such distri- Melanesian Borderlands formed variously during butions to have been attained by westward dis- the past 80-odd m.y., probably as island arcs. The persal from the western Pacific Plate margin. It New Hebrides, Fiji, and Tonga islands are em- appears that the species involved are uncommon placed on a ridge that originated during the on the western margin of the Philippine Plate, Eocene (at about the time Australia separated and one may question whether breeding popula- from Antarctica, about 50 m.y. ago) on the Aus- tions exist (some examples: Acanthurus achillies, A. tralian side of a west-dipping subduction zone leucopareius, Figure 5). The Marianas also serve as that marked the boundary between the Indian- Australian and Pacific plates. The present ar- 12 Moberly, 1972:43, indicated the possibility that the rangement of the islands on the ridge is believed northern part of the Philippine Plate was probably the old to be the result of fracturing of the ridge effected western part of the Pacific Plate. by complex tectonic processes. In some areas NUMBER 367 147 subduction zones shifted westward reversing arc near the subduction zone, are much closer today polarity, as occurred in the New Hebrides linea- than they were in the Miocene (if the Samoan ment. islands were in existence then; the Samoan Is- The oldest rocks of Fiji are upper Eocene, of lands have not been dated, but Hawkins and Tonga, pre-upper Eocene, of Lau, Miocene, and Natland, 1975, proposed an age of only 1 to 2 of the New Hebrides, Oligocene. Gill and Gor- m.y., or less). ton's (1973, fig. 5) proposed reconstruction of the The Cenozoic history given here seems to be of original ridge supporting these islands consisted little consequence for an explanation of today's of a northwest to southeast trending segment (the biotic isolation of the Pacific Plate in the Polyne- New Hebrides Ridge, on which the New Hebrides sian area. While there is possibly pertinent infor- and Fiji islands were positioned) bent rather mation available that I have overlooked that sharply at its eastermost limit and continuing as might afford such an explanation, I must leave a southerly extending segment (the Lau-Tonga that for the future. Although I am reluctant to Ridge, on which the Lau and Tonga islands were extend the search for an explanation to the pre- positioned). The Lau-Tonga Ridge was probably Cenozoic, it is possible that Cretaceous to Paleo- continuous with the Kermadec Ridge (but today cene events are responsible for some Pacific Plate there is a discontinuity between the ridges), which endemism. My reluctance stems from the fact supports the Kermadec Islands and continues to that endemism of fishes (but not necessarily other New Zealand. groups) on the Pacific Plate is almost entirely The Lau-Tonga Ridge sundered along its vol- restricted to the species level; whereas, the fossil canic (longitudinal) axis about 5.5 m.y. ago into record (limited to continental areas for fishes), in the present-day Lau-Coleville Ridge to the west general, gives little evidence for the existence of and Tongan Ridge to the east (these two ridges extant generic taxa earlier than Eocene (Romer, can be seen on Figure 3 extending north from 1966; I question whether many of the Eocene New Zealand). The Lau-Coleville Ridge (a rem- genera are correctly identified with the extant; nant arc) is currently inactive, but the volcani- my limited experience with fossil fishes would cally active Tonga Ridge continues its eastward indicate that few extant genera are present prior movement, carrying the Tonga Islands away from to the Miocene). The following discussion pre- Fiji and over the edge of the Pacific Plate. During sents a possible scenario for pre-Cenozoic vicari- early separating, the Lau-Coleville Ridge moved ance events that could lead to Pacific Plate ende- westward, and the northern end of the Ridge mism. together with Fiji underwent a counterclockwise Schlanger and Premoli-Silva (1981) and rotation. The separation of Fiji and the New Schlanger et al. (1981) have reasonably defended Hebrides is believed to have occurred earlier the existence of a Cretaceous episode 110 to 70 (Middle Miocene or Tertiary, 9-12 m.y. ago) m.y. ago of widespread volcanism on the Pacific than the Fiji-Tonga separation. Fiji today is much Plate. The volcanism resulted in a regional uplift- further from the New Hebrides than it was 9-10 ing of the Plate reminiscent of Menard's (1964) m.y. ago.13 It also appears that the eastward Darwin Rise. Subsidence has now drowned most, moving Tonga Islands and the westward moving if not all, the islands that were formed, and Samoan Islands, which are on the Pacific Plate increased the depth of the seamounts. Before subsidence the numerous volcanic edifices could 13 The movements that separated Fiji from the New He- have served as stepping stones (Schlanger et al., brides to the east and the Tonga Islands to the west may 1981, fig. 5) reaching from Australia across the have been the vicariant events that permitted the develop- Pacific Plate, and onto the now mostly subducted ment of endemic fish species in the area. Smith-Vaniz (1976) Farallon Plate (the Farallon and Pacific Plates recognized the following endemic blenniids: Meiacanthus bun- doon, M. atrodorsalis ovalauensis, Plagiotremus laudandus flavus (all shared a common spreading ridge, Figure 62, from Fiji), and Meiacanthus ptocne (from Tonga). and, thus, the age progression of the ocean bottom 148 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY on the two plates would form mirror images; the islands have a decided continental component in Farallon Plate split into the Cocos and Nasca their biota (Appendix 2: Table B, column A), plates during the Pliocene, Figures 1, 2, and 4). which is explainable by dispersal from New Schlanger and Premoli-Silva's (1981) evidence for Guinea along the closely spaced islands to Tonga, the action of stepping stones is based on the and even Samoa. Raven and Axelrod (1972), who presence of a benthonic, reef-associated fauna of were concerned with terrestrial biogeography, de- large pseudorbitoid foramnifera known only from veloped a partially similar scenario, postulating the Upper Cretaceous of Central America, the that Fiji was formerly much closer to Australia, Line Islands, the Nauru Basin, and New Guinea. but derived its continental biota, in part, by this Later subsidence of edifices on the western por- continguity and, in part, by overwater dispersal tion of the Pacific Plate could have isolated the from both Malaysia and Polynesia. shallow-water and terrestrial biotas of the edifices Schlanger et al. (1981) noted similarities in the to the east and allowed them to diverge. More geology of the Nauru Basin (area labelled 8 just recent island building on the Pacific Plate would west of New Guinea on Figure 4), on the western have provided havens for the biota of subductng part of the Pacific Plate (Figure 1), and the edifices; however, this would not allow for the Caribbean Plate, which has been considered a extinction of the pseudorbitoid foramniferan detached relic of the Darwin Rise (Mattson, fauna in the area, which may have entailed wide- 1969), hence, possibly a former part of the Far- spread, deep subsidence of the entire occupied allon Plate (Figure 62). It is interesting to specu- area. late that the seemingly anomalous distribution of It is possible that the opening of the Tasman iguanine lizards (Fiji, Tonga, Galapagos, Carib- and Coral sea basins (Figure 2; sea-floor ages for bean, and South America; none of the genera are these basins are indicated on Figure 4) were as trans-Pacific; G. Zug, pers. comm.) may be ex- important as subsidence for isolating the Pacific plained by Cretaceous stepping stones, but Plate biota. The Tasman Sea Basin began open- whether the stepping stones have anything to do ing in the Upper Cretaceous, about 80 m.y. ago with Pacific Plate endemic fish species, is another (Hayes and Ringis, 1973) and the Coral Sea Basin matter. At the risk of undermining my own res- began opening in the Paleocene about 62 m.y. ervations, it will be recalled that I mentioned in ago (Weissel and Watts, 1979). Both basins ceased the discussion of the shark family Carcharhinidae, opening in the Paleocene, about 56 m.y. ago the difficulty of explaining how Triaenodon obesus, (Weissel and Watts, 1979). The opening of these a sluggish, shallow-dwelling, obligate coral-reef basins drove the margin of the Indian-Australian inhabitant, which is widely distributed in the Plate perhaps as much as 1500 km east of the Indo-Pacific and otherwise known only in the coast of Australia (personal inference based on eastern Pacific, came to be so distributed. Creta- current positions). If no islands that could have ceous stepping stones? served as stepping stones were present in these basins during spreading, increased distance of Epilogue Australia from islands on the western Pacific In my study I have attempted to mesh the Plate margin could have served as a barrier to relatively little that is known about the distribu- dispersal between the Indian-Australian and Pa- tions and relationships of the shorefish fauna of cific plates in this region. Islands along the New the Pacific Plate with the relatively little that is Hebrides to Tonga lineament would have been known about the geotectonics of the Plate. The further distanced from Australia by opening of information available in both subject areas is the South Fiji Basin (Figure 2) in Oligocene, greatly imperfect, but it is difficult for me not to about 29-35 m.y. ago (Watts and Weissel, 1977). draw certain conclusions. (1) A large proportion Despite their remoteness from Australia, these of the Indo-Pacific shorefish fauna is divided into NUMBER 367 149 two geologically associated types: continental restricted, or nearly restricted, to a continental lithospheric plate and oceanic lithospheric plate. plate area, often including the joint continental The distinction is not as striking as the distinction and oceanic plate margins. If formal cladistic between black and white, but there is only a studies (now lacking) corroroborate these rela- narrow gray line that separates great expanses of tionships, and I predict that they will, a causal the shades on either side. (2) If a Pacific Plate relationship between present distributions and endemic taxon is widely distributed, there are past geotectonic events along the joint continental often hints that its closest relative is to be found and oceanic plate margins will be unavoidable.
Addendum
While this study was in press I had the opportunity to collect fishes in the Fiji Islands. I obtained the first records for Fiji of four shorefish families (specimens to be deposited at USNM): Acanthoclinidae {Acanthoclinus hiatti), Gobioididae (Taenioides sp.), Opistognathidae {Opistognathus sp.), and Schind- leriidae (Schindleria praematura). The first two families were not indicated as expected to occur in Fiji (or New Caledonia), but the Opistognathidae and Schindleriidae were listed as expected (present or expected in New Caledonia) in Appendix 2, Table A. These new records increase by two the number of families present or expected in Fiji (Appendix 2, Table B; Figure 58). I now expect that the Acanthoclinidae and Gobioididae will also be found to occur in New Caledonia, thus increasing by two the number of families expected at that locality. A.D. Lewis, Principal Fisheries Officer in Fiji, informed me that he had records, but no specimens, of Rastrelliger (Scombridae) from the Marshall Islands. J.E. Randall (pers. comm.), who has spent much time collecting in the Marshall Islands and is preparing a checklist of Marshall Islands fishes, has expressed doubt that Rastrelliger occurs in the Marshalls. Verification by specimens is desirable. I am informed by J.I. Ford (in litt., June 1982) that he recently obtained the following genera of eleotridids and gobiids in the freshwaters of Ponape, Eastern Caroline Islands (specimens to be deposited at the Bishop Museum): Ophiocara, Bunaka, Oxytleotris (all Eleotrididae), Stigmatogobius (= Pseudogobius?), Pandaka, Sicyopus (all Gobiidae). These genera represent first records for the nonmarginal area of the Pacific Plate, and give futher evidence for operation of the Caroline Islands conduit. Appendix 1 The Andesite Line
Gutenberg and Richter (1939) stated that Suess Linie, which he illustrated on a map of the western (1888) was among the first to notice that there Pacific (reproduced here as Figure 64). Born's was a line of demarcation separating structures intent was to stress the sharp difference in the of two different types in the western Pacific, but composition of the rocks of the islands of the it was Born (1932) who coined the term Andesite- Pacific Basin from the rocks of the land masses to
an haII „ • •••• •• Vo r w i e g e n
FIGURE 64.—The Andesite Line (from Born, 1932, fig. 306). 150 NUMBER 367 151 the west of the Basin. Born's term was incorpo- included the Indian Ocean side of the Indonesian rated into the geological literature, where it is still island arc in the zone. used, although Born's name is rarely associated Gutenberg and Richter (1941) stressed the with the term. Phytogeographers frequently men- strong association of Pacific Basin seismicity with tion the Andesite Line in passing, but seemingly the Andesite Line. The distribution of earthquake have not found the feature meaningful. In con- epicenters caused them to favor Born's delinea- trast, the zoogeographical literature, and partic- tion of the Andesite Line over Chubb's definition ularly the ichthyological, often invokes the An- of the limits of the andesite zone, where the two desite Line as an important faunistic boundary, conflicted. describing the easternmost limits of the distribu- MacDonald (1949) incorporated the Andesite tion of Indo-Pacific species. Line (without mentioning the term), with a mod- Chubb (1934), in a literature compilation of ification in the eastern Pacific, to define a hypo- the limits of the "andesite zone" of the Pacific, thetical boundary of the Pacific Basin (Figure 65 did not refer to Born (1932). Chubb's Pacific reproduces MacDonald's figure 11). It is interest- Basin (1934, fig. 1), margined by andesite, rep- ing, but not surprising, to note how closely resented, in effect, an extension of the Andesite MacDonald's Pacific Basin compares with pres- Line around the northern Pacific and down the ent concepts of the Pacific Plate (Figure 1). coast of the Western Hemisphere. Chubb's an- After MacDonald, the importance of andesite desite zone in the western Pacific deviated from and the integrally associated patterns of seismic- Born's by improperly including many of the East- ity were rapidly incorporated into the developing ern Caroline Islands. Chubb, however, correctly plate tectonic paradigm. The Andesite Line is
NORTH PACIFIC OCEAN
HAWAIIAN "• • .' ISLANDS
^MARQUESAS ISLANDS
. • • ' SALA-V^OOMEZ SAN FEUX^N
JUAH FERNANDE V T it PACIFIC JOCEAN ISLANDS
FIGURE 65.—The Andesite Line (from MacDonald, 1949, fig., 11). 152 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY still referred to frequently in the geological liter- may occur in other oceanic rocks of hot spot ature as a descriptive convenience. The current origin and are unlike the andesite of continental geological significance of the Line has been de- margins, hence they are not the result of stranding scribed by Holmes (1978:664). (Bandy, 1937; Bonatti et al., 1977). For geological purposes, and probably biologi- The boundary between the basalts of the oceanic crust and cal also, the western margin of the Pacific Basin islands, and the andesite-dacite-rhyolite volcanic rocks of the can be described best by the andesitic line of circum Pacific belt, is called the andesite line, and it turns out to be essentially the boundary between the continental greatest seismicity, with a secondary margin crust with sial and the oceanic crust without sial, and to along the boundary between the Philippine and follow the ocean trenches. The only exceptions to this gen- Pacific plates. Bom's Line must also be shifted to eralization are due to the occurrence of a few isolated shreds the east of the subduction zone in the region of and patches of sial left stranded in the Pacific floor, eg the Solomon to New Hebrides islands, where arc Macquarie Island and possibly Easter Island. The andesite polarity has been reversed. line around the Pacific and its continuation around the Indonesian Arc is ... "one of the fundamental geological The Andesite Line has also been referred to boundaries" of the earth. It has turned out to be the line infrequently as the Sial Line or Marshall Line, where oceanic lithosphere is subducting along Benioff zones. but I have been unable to determine the origins of these designations, which are apparently more The term "stranded" is open to question. Easter recent. I have not seen them used without indi- Island andesitic basalts are typical of those that cation that they are synonyms of the older term. Appendix 2 Tables
TABLE A.—Distribution of the Indo-Pacific families (and one order and two suborders) of warm-water, marine shorefishes (A = absent from Pacific Plate except possibly marginally; B = present on Pacific Plate nonmarginally only in Hawaiian Islands; C = present on the Pacific Plate nonmarginally only in the Eastern Caroline Islands; D = present on the Pacific Plate nonmarginally other than only in the Hawaiian or Eastern Caroline Islands; AE = Africa, southeast coast; IM = Indo-Malayan region; PI = Philippine Islands; NG = New Guinea and Bismarck Archipelago; QD = Queensland; GU = Guam; PO = Ponape; MI = Marshall Islands; HI = Hawaiian Islands; NC = New Caledonia and New Hebrides; FI = Fiji Islands; SA = Samoan Islands; SI = Society Islands; RI = Rapa Island; DI = Ducie Island; El = Easter Island; + = applicable distribution pattern; X = present; 0 = expected; - = absent; * = questionably characterized as shorefishes; ** = questionably characterized as warm-water inhabitants)
Family, Suborder, Order A B C D AE IM PI NGQDGUPO MI HINC FI SA SI RI DI El Acanthoclinidae + X X X 0 X _ 0 X ______Acanthuridae + X X X X X X X X X X X X X X X X Albulidae + X X X X X 0 0 - X X 0 X X X - - Ambassidae + X X X X X - X - - X X X - - _ - Ammodytidae* * + X - - X X - - - X ------Anguillidae + X X X X X X X - - X X X X X - - Anomalopidae + 0 X X X - X 0 X - X X X - - - - Antennariidae + X X X X X X X X X X X X X - - X Aploactinidae + X X X X X - - - - 0 X - - - - - Apogonidae + X X X X X X X X X X X X X X X X Aracanidae* +" X - - - X - - - X ------A "" A • y\.x x x x /\rnoae x + X X X X X X X X X X X X 0 X _ _ AulostomidaAtherinidae e + X X X X X X X X X X X X X X X X Balistidae + X X X X X X X X X X X X X X X X Batrachoididae + X X X X X Belonidae + X X X X X X X X X X X X X X X X Blenniidae + X X X X X X X X X X X X X X X X Bothidae + X X X X X 0 X X X X X X X X X X Brachaeluridae + - - - - X Branchiostegidae + X X X 0 X X - - - X 0 - - - - - Bregmacerot idae * + X X X X X X 0 X X 0 X 0 - - - - Bythitidae + X X X X X X X X X X 0 X - - X - Caesionidae + X X X X X X X X - X X X X X - - Callionymidae + X X X X X X X X X X X X X - - - Caracanthidae + X X X X X X X X X X X X X - - - Carangidae + X X X X X X X X X X X X 'X X X X Carapidae + X X X X X X 0 X X X X X X - - - Carcharhinidae + X X X X X X X X X X X X X X X X Centriscidae + X X X X X - X - - X ------Centropomidae +b - X X X X Cepolidae + X X X X X - - - - X ------Chaetodontidae + X X X X X X X X X X X X X X X X Chanidae + X X X X X X X X X X X X X - - - Cheilodactylidae + X - - - X - - - X X - - - X - X Chirocentridae +b X X X X X - - - - X X - - - - -
153 154 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE A.—Continued
Family, Suborder, Order A B C D AE IM PINGQDGU PO MI HI NC FI SA SI RI DI El Cirrhitidae 4- X X X X X X X X X X X X X X X X Citharidae* + X 0 X 0 Clinidae + X X X - X - - - - X ------Clupeidae 4- X X X X X X X X X X X X X - - - Congridae + X X X X X X X X X X X X X X - X Congrogadidae + X X X X X Coracinidae* * + X Coryphaenidae + X X X X X X 0 X X X X X 0 X 0 0 Creediidae + X X 0 0 X 0 0 X X 0 0 X X 0 X X Cynoglossidae X X X X X - - - X ------Dactylopteridae + X X X X X X 0 - X X X X X X - - Dasyatididae + X X X X X X 0 X X X X X X - - - Diodontidae + X X X X X X 0 X X X X X X X X X Drepanidae +b X X X X X - - - - 0 0 X - - - - Echeneididae + X X X X X X 0 X X X X X X X 0 X Eleotrididae + X X X X X X X X X X X X X X - X Elopidae + X X X X X X 0 0 X X X X X - - - Emmelichthyidae* +" X - X - - X 0 - X ------X Engraulididae + X X X X X X 0 0 X X X X X - - X Enoplosidae* * + - - - - X Ephippididae + X X X X X X X X - X X X X - - - Exocoetidae* + X X X X X X X X X X X X X X 0 X Fistulariidae + X X X X X X X X X X X X X X X X Fnrminn iH UP 4-c J. Ul IlllUllIlittv x x x x x Gerreidae X X X X X X X X _ X X X _ _ _ _ Ginglymostomatidae + X X X X X X 0 X - X X X X - - - Girellidae* * + - - - - X _ ------X Glaucnsomatidae 4. x x Gobiesocidae X X X X X X 0 X _ X X X _ _ _ _ Gobiidae + X X X X X X X X X X X X X X X X Gobioididae 4- X X X X X X X ______Gonorynchidae* * + X - _ _ X _ _ _ X _ _ _ _ _ Gramma t idae +d X X X _ _ _ _ _ X X ______Gymnuridae x x x x Haemulidae 4- X X X X X X X X _ X X X X X _ _ Harpadontidae 4- x x x x Hemiramphidae X X X X X X X X X X X X X X 0 X Hemiscylliidae 4- x x x x x Heterodontidae 4- x x x Holocentridae X X X X X X X X X X X X X X X X Isonidae + X ______X 0 _ X _ _ _ _ Istiophoridae* + X X X X X X 0 0 X X 0 X X X 0 X Kraemeriidae + X 0 X X X X 0 X X 0 0 X X _ _ _ Kuhliidae + X X X X X X X X X 0 X X X X X X Kurtidae 4. x x x Kyphosidae 4- X X X X X X X X X X X X X X X X Labracoglossidae* * + - _ X ______0 X _ _ X _ X Labridae + X X X X X X X X X X X X X X X X Lactariidae + _ X X X X _ _ _ _ 0 X _ _ _ _ _ Lam n idae* 4- X X X 0 X 0 0 X X X X X X X 0 X Leiognathidae + X X X "X X X X _ _ X X X _ _ Leptobramidae 4. x x Lethrinidae 4. X X X X X X X X X X X X X X X NUMBER 367 155
TABLE A.—Continued
Family, Suborder, Order A B C D AE IM PI NGQDGU PO MI HINC FI SA SI RI DI El e Lobotidae + X X X X X _ X _ X 0 X _ _ _ _ _ Lophiidae* +" X X X - X - - - X ------Lutjanidae + X X X X X X X X X X X X X X X X Malacanthidae + X X X X X X X X X X X X X - - - Menidae +c X X X X X - - - - 0 X - - - - - Microdesmidae + X X X X X X X X X 0 X X X - - - Mobulidae + X X X X X 0 0 X X X 0 0 X X - - Monacanthidae + X X X X X X X X X X X X X X X X 4. 1V1(JI1LH_CI1LI lUIUdC x x x x Monodactylidae X X X X X X X _ _ X X X _ - - _ Moridae* ** +f X X X X 0 - - - X 0 X - - X - - Moringuidae + X X X X X X X X X 0 X X X X X X Mugilidae + X X X X X X X X X X X X X X X - Mugiloididae + X X X X X X X X X X X X X - - - Mullidae + X X X X X X X X X X X X X X X X Muraenesocidae + X X X X X - - - - 0 X - - - - - Muraenidae + X X X X X X X X X X X X X X X X Myliobatididae + X X X X X X 0 X X X X X X - - - Nemipteridae + X X X X X X X X - X X X - - - - Neostethidae + - X X Nomeidae* + X X X 0 X X 0 X X 0 0 0 - - - - Notograptidae + - - - X X Odontaspididae +a X 0 - - X - - - X ------Ogcocephalidae* +a X X X X X - - - X ------Ophichthidae 4- X X X X X X X X X X X X X X X X Ophidiidae + X X X X X X X X X X X X X - - X Opistognathidae + X X X X X - - - - X 0 X - - - - Oplegnathidae** + X ------X ------Orectolobidae + X X X X X Ostraciidae + X X X X X X 0 X X X X X X X 0 X Pataecidae" — — ~ ~ •^ Pegasidae +g X X X X X X - - X X ------Pempherididae + X X X X X X X X - X X X X X X - Pentacerotidae* ** + X - X X - - - - X ------Pholidichthyidae + - X X X - - - - - X ------Platycephalidae + X X X X X X X X X X X X X - - - Plesiopiodae + X X X X X X X X - X X X - - - - Pleuronectidae + X X X X X 0 0 X X 0 X X X - - - Plotosidae X X X X X - - - - X X X - - - - Polynemidae + X X X X X X 0 X X X X X X X - - Pomacanthidae + X X X X X X X X X X X X X X X X Pomacentridae + X X X X X X X X X X X X X X X X Porrtatomidae + X - - - X ------Priacanthidae + X X X X X X X X X X X X X X X X Pristidae + X X X X X ------v V x x Psettodidae* + A A A A Pseudochromidae + X X X X X X X X - X X X - - Pseudotrichonotidae T — ~ ~ ~ Rachycentridae +b X X X X X ------v v Rajoidei (suborder) + A A A Rhinobatoidei (suborder) + X X X X X 0 - X X X + Rhinoprenidae x x x x x Rhinopteridae yv 156 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE A.—Continued
Family, Suborder, Order A B C D AE IM PINGQDGU PO MI HI NC FI SA SI RI DI El Scaridae + X X X X X X X X X X X X X X X X Scatophagidae + X X X X X - X - - X X - - - - - Schindleriidae + X 0 0 X 0 0 0 X X 0 0 X X - - - c • • J -4-b X ociaemaae yxv x yxv yvx 4- X X X X X X X X X X X X X Scombridae X X X Scorpaenidae + X X X X X X X X X X X X X X X X Scorpididae + X - - - X - - - X X ------Scyliorhinidae +a X X X - 0 - - - X ------Serranidae + X X X X X X X X X X X X X X X X Siganidae 4- X X X X X X X X - X X X X X - - Sillaginidae 4- X X X X X - - - - X ------Soleidae 4- X X X X X X X X X X X X X - - X Solenostomidae 4-h X X X X X X 0 X - 0 X - - - - - Sparidae 4- X X X X X - - - - X ------Sphyraenidae 4- X X X X X X X X X X X X X X - - Sphyrnidae 4- X X X X X X 0 X X X X X X X - - Squatinidae 4- X - 0 X Stegostomat idae 4- X X X 0 X - - - - 0 X X - - - - Stromateidae 4- X x x Syngnathidae 4- X X X X X X X X X X X X X X _ X Synodontidae 4- X X X X X X X X X X X X X X X X Terapontidae 4- X X X X X _ - - - X X X _ _ _ _ Tet raodont idae 4- X X X X X X X X X X X X X X X 0 Torpediniformes (order) 4-* X X X - X - _ - X ______Toxotidae x x x x Tnacanthidae + x x x x Triakidae 4- X x x x Trichiuridaeb>' 4- X X X X X X X j Trichonotidae 4- X X X X X X 0 X _ X X _ _ _ _ _ Triglidae 4- X x x x x Triodontidae* 4- X X X 0 X _ _ _ X X X Tripterygiidae 4- X X X X X X X X X X X X X X X _ Trypauchenidae 4- X X X X X _ _ _ _ _ X _ _ _ _ _ Uranoscopidae 4- X X X X X X _ _ _ _ X X _ _ _ _ Urolophidae 4-' X X - X X _ _ _ X ______Xenocongridae 4- X X X X X X X X X 0 X X X _ _ _ Zanclidae 4- X X X X X X X X X X X X X X X - Totals 58 15 6 100 Grand Total 179 * Known nonmarginally on Pacific Plate only from deep f A shorefish species of Lotella occurs at Rapa; otherwise, water in Hawaiian Islands. family is known on Pacific Plate only from deep water. b Nonmarginal Pacific Plate distribution is indicated for g Known nonmarginally on Pacific Plate only from Ha- some species on generalized distribution maps in Fischer and waiian Islands and from stomach of tuna caught off Whitehead, 1974; I have been unable to verify such distri- Tahiti. butions, and in many instances these nonmarginal represen- Known nonmarginally on Pacific Plate only from a tations are known to be erroneous (for example, Scomberomonis single, juvenile specimen taken in a plankton tow off Bikini commerson, fide B.B. Collette, pers. comm.). Atoll (Schultz, 1953c) and from a photograph of an adult c Indicated for Hawaiian Islands by Matsubara (1955), taken by a resident at Kwajalein (both localities in Marshall probably erroneously. Islands). d Family composition undecided; Pacific Plate species is Here considered to include only Trichiurus and Leptura- deep dwelling in Hawaiian Islands. canthus. ' Apparently occurs nonmarginally on Pacific Plate only J Known nonmarginally on Pacific Plate only from Ene- as a straggler. wetak, Marshall Islands. NUMBER 367 157
TABLE B.—Summary of data on numbers of families of shorefishes in Table A (for a geographic plot see Figure 58)
Locality A B C D Known + Expected = Total Africa, SE coast (AE) 42 14 6 99 160 1 161 Indo-Malayan Region (IM) 47 8 6 % 153 4 157 Philippine Islands (PI) 46 9 6 97 155 3 158 New Guinea-Bismarck (NG) 42 4 6 97 141 8 149 Archipelago Queensland (QD) 50 10 6 97 160 3 163 Guam (GU) 2 2 5 93 94 8 102 Ponape (PO) 0 1 6 95 73 29 102 Marshall Islands (MI) 0 0 0 91 88 3 91 Hawaiian Islands (HI) 0 15 0 83 98 0 98 New Caledonia-New (NC) 19 3 5 98 104 21 125 Hebrides Fiji Islands (FI) 15 0 4 97 106 10 116 Samoan Islands (SA) 8 0 2 92 99 3 102 Society Islands (SI) 0 0 0 83 82 1 83 Rapa Island (RI) 0 0 0 63 62 1 63 Ducie Island (DI) 0 0 0 47 40 7 47* Easter Island (El) lb 1 0 52 50 2 52
* Not included here as present or expected at Ducie are 20 families obtained by John E. Randall (in litt.) from Ducie's nearest neighboring islands (nearest about 600 km west): Henderson, Pitcairn, and Oeno. It is possible that some of these families reach east to Ducie. Of note here is that Randall spent only 2.5 days collecting at Ducie, an atoll, but a month at Pitcairn, a high island. The families are Ammodytidae, Antennariidae, Caesionidae, Calliony- rnidae, Caracanthidae, Carapidae, Congridae, Dactylopteridae, Gobiesocidae, Isonidae, Mu- giloididae, Ophidiidae, Platycephalidae, Pleuronectidae, Polynemidae, Pseudochromidae, Si- ganidae, Soleidae, Sphyraenidae, Xenocongridae. b Eastern Pacific representation; not present on Pacific Plate nonmarginally.
TABLE C.—List of Indo-Pacific families and genera of warm-water shorefishes that occur other than marginally on the Pacific Plate. A few of the families and genera are questionably warm- water and/or shorefishes. Some families and genera not included in this list have been indicated as occurring on the Pacific Plate nonmarginally in Fischer and Whitehead, 1974 (see footnote b to Table A). Numerous genera have been omitted on the strength of unpublished synonymies from several sources. Numbers following family names indicate known or estimated total numbers of warm-water shorefish species that occur nonmarginally on Pacific Plate. Numbers following generic names indicate known total numbers of warm-water shorefish species that occur nonmarginally on Pacific Plate allowing for numerous unpublished records and unde- scribed species.
ACANTHOCLINIDAE 1 ALBULIDAE 2 ANOMALOPIDAE 2 Acanthoplesiops 1 Albula 2 Anomalops 1 ACANTHURIDAE 39 AMBASSIDAE 1 Photoblepharon 1 Acanthurus 20 Ambassis 1 Ctenochaetus 5 AMMODYTIDAE 1 ANTENNARIIDAE 10 Naso 9 Bleekeria 1 Abantennahus Paracanthurus 1 ANGUILLIDAE 3 Antennarius Zebrasoma 4 Anguilla 3 Phrynelox 158 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE C.—Continued
APOGONIDAE 50 Omox Galeocerdo 1 Apogon Parenckelyurus Negaprion 1 Apogonichthys Petroscirtes 5I Triaenodon 1 Archamia Parablennius CENTRISCIDAE 1 Cheilodipterus Plagiotremus I Aeoliscus 1 Foa Praealticus CHAETODONTIDAE 40 Fowleria Rhabdoblennius Chaetodon 30 Gymnapogon Salarias Forcipiger 2 Neamia Stimulus Hemitaurichthys 3 Pseudamia Xiphasia I Heniochus 5 Pseudamiops BOTHIDAE 5 CHANIDAE 1 Rhabdamia Arnoglossus Chanos 1 Siphamia Bothus CHEILODACTYLIDAE 2 Sphaeramia Engyprosopon Cheilodactylus 2 New genus* BREGMACEROTIDAE 3 ClRRHITIDAE 17 ATHERINIDAE 6 Bregmaceros Amblycirrhitus 1 Atherinomorus BYTHITIDAE 10 Cirrhitichthys (includes Pranesus) Brosmophyciops Cirrhitops 2 Atherion Dinematichthys Cirrhitus 1 Hypoatherina Microbrotula Isocirrhitus 1 Stenatherina Oligopus Neocirrhites 1 AtTLOSTOMIDAE 1 CAESIONIDAE 10 Oxycirrhites 1 Aulostomus 1 Caesio Paracirrhites 6 BALISTIDAE 19 Pterocaesio New genus 1 Balistapus 1 CALLIONYMIDAE 10 CLUPEIDAE 6 Balistoides Callionymus 2 Dussumiena 1 Canthidermis Diplogrammus 2 1 Etrumeus 1 Melichthys 2 Pogonymus [ Herklotsichthys 1 Odonus Synchiropus I Sardinclla 2 Pseudobalistes CARACANTHIDAE 2 3 Spratelloides • Rhineacanthus J Caracanthus ' i CONGRIDAE 8 Sufflamen ? CARANGIDAE 29 Ariosoma Xanthichthys J Alectis I 2 BELONIDAE 4 Atule Conger 2 Gorgasia 1 Abletmes 1 Carangoides l \ Heteroconger 2 Platybelone I Caranx ! Poeciloconger 1 Strongylura I Decapterus l \ Tylosurus I Elagatis 1 CORYPHAENIDAE 2 BLENNIIDAE 62 Gnathanodon Coryphaena 2 Alticus Naucrates CREEDIIDAE 4 r Aspidontus > Pseudocaranx Chalixodytes 1 Atrosalarias Scomberoides Crystallodytes 1 CirripecUs Selar Limnichthys 2 Ecsenius ' ! Seriola 4 DACTYLOPTERIDAE 1 Enchelyunts ) Trachinotus ',> Dactyloptena 1 Entomacrodus 13 Uraspis \) DASYATIDIDAE 5 Exallias CARAPIDAE 6 Dasyatis Glyptoparus Carapus Himantura Istiblemius Encheliophis Taeniura Medusablenmus Onuxodon Urogymnus Meiacanthus Snyderidia DlODONTIDAE 5 Nannosalarias CARCHARHINIDAE 12 Chilomycterus 1 Omobranchus Carcharhinus \) Diodon 4 NUMBER 367 159
TABLE C.—Continued
ECHENEIDIDAE 6 Feia 1 Myripristis 12 Echeneis Fusigobius New genusb 1 Phtheirichthys Gladygobius Ostichthys 2 Remora Glossogobius Plectrypops 1 ELEOTRIDIDAE 15 Gnatholepis ISONIDAE 2 Allomicrodesmus 1 Gobiodon Iso 2 Bos tryc thus Gobiopsis ISTIOPHORIDAE 2 Butis 1 Istigobius Istiophorus 1 Calumia 1 Kelloggella 4 Makaira 1 Eleotris Lentipes 1 KRAMERIIDAE 2 Hypseleotris Macrodontogobius Kraemeria Xenisthmus Mugilogobius KUHLIIDAE 3 ELOPIDAE 3 Nemateleotris 2 Kuhlia Elops 2 Oplopomops KYPHOSIDAE 4 Megalops 1 Oplopomus Kyphosus ENGRAULIDIOAE 4 Opua Sectator 1 Encrasicholina 1 Oxyurichthys LABRACOGLOSSIDAE 1 Stolephorus 3 Paragobiodon 1 Bathystethus 1 EPHIPPIDIDAE 2 Parioglossus 2 LABRIDAE 100 Platax 2 Periophthalmus Anampses 6 EXOCOET1DAE 15 Pleurosicya Bodianus 7 Cypselurus Priolepis Cheilinus 9 PExocoetus Psilogobius Cheilio 1 Fodiator Ptereleotris 2 Cirrhilabrus PParexocoetus Redigobius Coris 8 Prognichthys Sicydium Cymolutes FlSTULARIIDAE 2 Sicyopterus Epibulus 1 Fistularia 2 Stenogobius Gomphosus 1 GERREIDAE 3 Stiphodon Halichoeres 12 Genes Stonogobiops Hemigymnus 2 GlNGLYMOSTOMATIDAE 1 Trimma Hologymnosus 2 Nebrius 1 Valenciennea Labrichthys 1 GOBIESOCIDAE 3 Vanderhorstia Labroides 5 Lepadichthys 1 Vitraria 1 Labropsis 4 Liobranchia 1 Waitea Macropharyngodon 3 Pherallodus 1 Yongeichthys Novaculichthys 2 0 GOBIIDAE 150 Undescribed genus Polylepion 1 Acentrogobius GOBIOIDIDAE 2 Pseudocheilinus 5 Amblyeleotris Brachyamblyopus 1 Pseudocoris 2 Amblygobius Taenioides 1 Pseudodax 1 Asterropteryx 2 GONORYNCHIDAE 1 Pseudojuloides 3 Awaous Gonorynchus 1 Pseudolabrus 3 Bathygobius HAEMULIDAE 5 Pteragogus 1 Cabillus Plectorhynchus Stethojulis 4 Callogobius HEMIRAMPHIDAE 9 Suezichthys 1 Cryptocentroides 1 Euleptoramphus 1 Thalassoma 9 Cryptocentrus Hemiramphus Wetmorella 1 Ctenogobiops Hyporhamphus Xyrichtys Discordipinna 1 Oxyporamphus 1 LAMNIDAE 3 Drombus Zenarchopterus 1 Carcharodon 1 Eilatia HOLOCENTRIDAE 29 Isurus 2 LEIOGNATHIDAE Eviota Adioryx 13 2 Exyrias Flammeo 4 Gazza 1 160 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE C.—Continued
Leiognathus 1 Enchdynassa PLESIOPIDAE 3 LETHRINIDAE 7 Gymnomuraena Calloplesiops 1 Gnathodentex 1 Gymnothorax Plesiops 2 Lethrinus 5 ?Muraena PLEURONECTIDAE 2 Monotaxis 1 Rhinomuraena 1 Samariscus LOBOTIDAE 1 Uropterygius POLYNEMIDAE 3 Lobotes 1 MYLIOBATIDIDAE 2 Polydactylus L.UTJANIDAE 15 Aetobatis 1 POM ACANTHI DAE 25 Aprion 1 Pteromylaeus 1 Apolemichthys 3 Lutjanus NEMIPTERIDAE 5 Centropyge 15 Macolor 1 Pentapodus Euxiphipops 1 MALACANTHIDAE 5 Scolopsis Genicanthus 4 Hoplolatilus 3 NOMEIDAE 5 Pomacanthus 1 Malacanthus 2 Cubiceps Pygoplites 1 MlCRODESMIDAE 4 Nomeus 1 POMACENTRIDAE 72 Gunnellichthys 3 Psenes Abndefduf 6 Amblyglyphidodon Paragunnellich thys 1 OPHICHTHIDAE 40 3 MOBULIDAE 2 Achirophichthys 1 Amphiprion 5 Mania Apttrichtus Chromts 24 Mobula Brachysomoph is Chrysiptera 6 MONACANTHIDAE 17 Callechelys Dascyllus 7 Aluterus 2 Cirrhimuraena Lepidozygus 1 Amanses 1 Cirricaecula 1 Plectroglyphidodon 8 Brachaluteres Ichthyapus Pomacentrus 4 Canthrrhines 6 Lamnostoma Pomachromis 3 Oxymonacanthus 1 Leiuranus 1 Stegastes 5 Paraluteres 1 Muraenichthys PRIACANTHIDAE 3 Pervagor 2 Myrichthys Cookeolus 1 Thamnaconus 1 Afytophis Priacanthus MONODACTYLIDAE 1 Ophichthus PSEUDOCHROMIDAE 5 Monodactylus 1 Phaenomonas 1 Pseudochromis MORIDAE 1 Phyllophichthus 1 Pseudoplesiops Lotella 1 Schismorhynchus 1 SCARIDAE 31 MORINGUIDAE 3 Schultzidia 1 Bolbometopon 1 Moringua OPHIDIIDAE 2 Calotomus 2 MUGILIDAE 10 Brotula 2 (includes Scaridea) Chelon Cetoscarus 1 OPLEGNATHIDAE 2 Crenimugil Hipposcarus 1 Oplegnathus 2 Liza Leptoscarus 1 OSTRACIIDAE 6 Mugil Scants 25 Neomyxus Lactoria 3 SCATOPHAGIDAE 1 Plicomugil Ostracion 3 Scatophagus 1 Valamugil PEGASIOAE 1 SCHINDLERIIDAE 2 MUGILOIDIDAE 7 Eurypegasus 1 Schindleria 2 Parapercis 7 PEMPHERIDIDAE 3 SCOMBRIDAE 7 MULLIDAE 20 Parapriacanthus 1 Euthynnus 2 Mulloides 4 Pemphtris Grammatorcynus 1 Parupeneus PENTACEROTIDAE 1 Gymnosarda 1 Upeneus Histiopterus 1 Rastrelliger 1 MURAENIDAE 35 PLATYCEPHALIDAE 5 Sarda 1 Anarchias Onigocia Scomber 1 Echidna Thysanophrys SCORPAENIDAE 35 Enchelycore Wakiyus Dendrochirus NUMBER 367 161
TABLE C.—Continued
Iracundus 1 Plectranthias 5 Hippocampus Pterois 3 Plectropomus 4 Micrognathus Rhinopias 1 Pogonoperca 1 Oostethus 1 Scorpaena Pseudogramma 4 Penetopteryx 1 (includes Sebastapistes) (includes Aporops) Phoxocampus 1 Scorpaenodes Suttonia 1 Syngnathoides 1 Scorpaenopsis Variola 1 SYNODONTIDAE 12 Synanceia 1 SlGANIDAE 5 Saurida Taenianotus 1 Siganus Synodus 8 SCORPIDIDAE 1 (includes Lo) Trachinocephalus 1 Microcanthus 1 SOLEIDAE 4 TETRAODONTIDAE 20 SERRANIDAE 82 Aseraggodes Arothron Anthias 16 Soleichthys Canthigaster 13 (includes Miro- SOLENOSTOMIDAE 1 Lagoctphalus labrichthys and SoUnostofnus 1 Sphoeroides Pseudanthias) SPHYRAENIDAE 5 TRICHONOTIDAE 1 Anyperodon 1 Sphyraena Trichonotus 1 Aulacocephalus 1 Belonoperca 1 SPHYRNIDAE 2 TRIPTERYGIIDAE 10 Sphyma 2 Cephalopholis 8 Enneapterygius Epinephelus 19 SYNGNATHIDAE 25 Helcogramma Gracila 1 Choeroichthys 2 Norfolkia Grammistes 1 Corythoichthys 4 XENOCONGRIDAE 5 Grammistops 1 Cosmocampus Chilorhinus Holanthias 3 Doryrhamphus 1 Kaupichthys Liopropoma 9 Dunkerocampus 2 Powellichthys Luzonichthys 2 Festuealex 1 ZANCLIDAE 1 Odonlanthias 3 Hippichthys 1 Zanclus 1 1 E.A. Lachner, pers. comm. ' J.E. Randall, pers. comm. and in litt. D.F. Hoese, pers. comm. Literature Cited
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of the National Science Museum, Tokyo, 12 (4): 783-794, Lake, J.S. plates 1-7. 1971. Freshwater Fishes & Rivers of Australia. 61 pages, 16 Kotthaus, A. plates. 1975. Fische des Indischen Ozeans ... A. Systematischer La Monte, F. Teil, XVI. "Meteor" Forschungs-Ergebnisse, series D, 1952. Marine Game Fishes of the World. 190 pages. 21:30-53. Larson, H.K., and D.F. Hoese 1976. Fische des Indischen Ozeans ... A. Systematischer 1980. The Species of the Indo-West Pacific Genus Calu- Teil, XVII. "Meteor" Forschungs-Ergebnisse, series D, mia (Pisces: Eleotridae). Proceedings of the Linnean 23:45-61. Society of New South Wales, 104(1-2): 17-22. 1979. Fische des Indischen Ozeans ... A. Systematischer Lavenberg, R.J., and L.A. Yanex A. Teil, XXI. "Meteor" Forschungs-Ergebnisse, series D, 1972. A New Species of Cirrhitus from Easter Island 28:6-54. (Pisces, Cirrhitidae). Gayana, Zoologia, 21:1-11. Koumans, F.P. Le Danois, Y. 1933. On a New Genus and Species of Apogonidae. 1964. Etude anatomique et systematique des antennaires Zoologische Mededeelingen, 16(1—2):78, plate 1: figure de l'ordre Pediculates. Memoires du Museum National 2. d'Histoire Naturelle, Nouvelle Serie, series A (Zoolo- Kuramochi, T. gie), 31(1):1-162. 1980. A New Record of Kelloggella cardinalis (Family Lee, S.-C, and J.-T. Chang Gobiidae) from Taiwan. Japanese Journal of Ich- 1981. Comparative Studies of the Fishes of the Genus thyology, 27(3):256-258. Girella from Taiwan. Bulletin of the Institute of Zoology Kuthalingam, M.D.K. Academia Sinica, 20(l):9-16. 1967. Some Observations on the Fishery and Biology of Leis.J.M. Kurtus indicus (Bloch) of the Bay of Bengal. Indian 1978. Systematics and Zoogeography of the Porcupine- Journal of Fisheries, 10(A): 159-166. fishes (Diodon, Diodontidae, Tetraodontiformes) Kyushin, K., K. Amaoka, K. Nakaya, and H. Ida. with Comments on Egg and Larval Development. 1977. Fishes of the Indian Ocean. 392 pages. Fishery Bulletin, 76(3):535-567. Lachner, E.A. Liem, A.H., and W.B. Scott 1978. Echeneidae. In W. Fischer, editor, FAO Species 1966. Fishes of the Atlantic Coast of Canada. Fisheries Identification Sheets for Fishery Purposes, Western Cen- Research Board of Canada, Bulletin, 155:1-485. tral Atlantic, 2: 2 unnumbered pages. Lindberg, G.U., and Z.V. Krasyukova Lachner, E.A., and S.J. Karaella 1971. Teleostomi: XXIX. Perciformes ... CXLIV. 1980. Fishes of the Indo-Pacific Genus Eviota with De- Champsodontidae. In Fishes of the Sea of Japan and scriptions of Eight New Species (Teleostei: Gobi- Adjacent Areas of the Sea of Okhotsk and the Yellow Sea, idae) . Smithsonian Contributions to Zoology, 315:1-127. 3: v + 498 pages. [Translated from the 1969 Lachner, E.A., and J.F. McKinney Russian original.] 1978. A Revision of the Indo-Pacific Fish Genus Gobiopsis Lobel, P.S. with Descriptions of Four New Species (Pisces: 1981a. Invasion by the Mozambique Tilapia (Sarotherodon Gobiidae). Smithsonian Contributions to Zoology, mossambicus; Pisces; Cichlidae) of a Pacific Atoll 262:1-52. Marine Ecosystem. Micronesica, 16(2):349-355. 1979. Two New Gobiid Fishes of the Genus Gobiopsis and 1981b. Bodianusprognathus (Labridae, Pisces), a New Long- a Redescription of Feia nympha Smith. Smithsonian nose Hogfish from the Central Pacific. Pacific Sci- Contributions to Zoology, 299:1-18. ence, 35(1): 45-50. Ladd, H.S. Lourie, A., and A. Ben-Tuvia 1966. Chitons and Gastropods (Haliotidae through 1970. Two Red Sea Fishes, Pelates quadrilineatus (Bloch) Adeorbidae) from the Western Pacific Islands. and Crenidens crenidens (Forskal) in the Eastern United States Geological Survey Professional Paper, Mediterranean. Israel Journal of Zoology, 19:203- 531:1-98, plates 1-16. 207. 1972. Cenozoic Fossil Mollusks from Western Pacific Lubbock, R. Islands: Gastropods (Turritellidae through Strom- 1975. Fishes of the Family Pseudochromidae (Perci- bidae). United States Geological Survey Professional formes) in the Northwest Indian Ocean and Red Papa, 532:1-79, plates 1-20. Sea. Journal of the Zoological Society of London, 1977. Cenozoic Fossil Mollusks from Western Pacific 176:115-157. Islands: Gastropods (Eratoidae through Harpi- 1976. Fishes of the Family Pseudochromidae (Perci- dae). United States Geological Survey Professional Paper, formes) in the Central Indian Ocean. Journal of 533:1-84, plates 1-23. Natural History, 10:167-177. NUMBER 367 173
1977. Fishes of the Family Pseudochromidae (Perci- the Influence of Hydrography on Resident and formes) in the Western Indian Ocean. J.L.B. Smith Nonresident Scotian Shelf Ichthyofauna. Canadian Institute of Ichthyology, Ichthyological Bulletin, 35:1- Journal of Fisheries and Aquatic Sciences, 37(l):49-65. 21, 24-28. Marshall, N.B. 1978. A New Hawkfish of the Genus Amblycinhitus Gill 1950. Fishes from the Cocos-Keeling Islands. Bulletin of 1862, from Ascension Island, South Pacific. Senck- the Raffles Museum, 22:166-205, 2 plates. enbergiana Biologica, 58(5/6):261-265. 1952. The "Manihine" Expedition to the Gulf of Aqaba, 1980. Five New Basslets of the Genus Pseudochromis (Te- IX: Fishes. Bulletin of the British Museum (Natural leostei: Pseudochromidae) from the Indo-Austra- History), Zoology, 1(8):221-252 lian Archipelago. Revue Suisse de Zoologie, Masuda, H., C. Araga, and T. Yoshino 87(3):821-834. 1975. Coastal Fishes of Southern Japan. 379 pages. Lubbock, R., and G.R. Allen Matsubara, K. 1979. Canthigaster leoparda a New Sharpnose Pufferfish 1955. Fish Morphology and Hierarchy. Volume 1, xi + 789 (Teleostei: Tetraodontidae) from the Central pages. [In Japanese.] Indo-Pacific. Revue Franqaise de Aquariologie, Matsui, T. 6(3):87-90. 1967. Review of the Mackerel Genera Scomber and Ras- Lubbock, R., and A.J. Edwards trelliger with Description of a New Species of Ras- 1980. A New Butterflyfish (Teleostei: Chaetodontidae) trelliger. Copeia, 1967(1): 71-83. of the Genus Chaetodon from Saint Paul's Rocks. Matsuura, K. Revue Franqaise d'Aquariologie, 7(1): 13-16. 1980. A Revision of Japanese Balistoid Fishes, I: Family Lubet, P. Balistidae. Bulletin of the National Science Museum, 1973. Family Sepiidae. In W. Fischer, editor, FAO Species Tokyo, series A (Zoology), 6(l):27-69. Identification Sheets for Fishery Purposes, Mediterranean 1981. Xenobalistes tumidipectoris, a New Genus and Species and Black Sea (Fishing Area 37), 2: 2 unnumbered of Triggerfish (Tetraodontiformes, Balistidae) pages. from the Marianas Islands. Bulletin of the National Lund, W.A., Jr., and G.C. Maltezos Science Museum, Tokyo, series A (Zoology), 7(4): 191- 1970. Movements and Migrations of the Bluefish, Pom- 200. otomus saltatrix, Tagged in Waters of New York Mattson, P.H. and Southern New England. Transactions of the 1969. The Caribbean: A Detached Relic of the Darwin American Fisheries Society, 99(4): 719-731. Rise. Transactions of the American Geophysical Union MacArthur, R.H., and E.O. Wilson (EOS), 5O(3):317. 1967. The Theory of Island Biogeography. xi + 203 pages. Mauge, A.L. MacDonald, G.A. 1980. La Denture entopterygoidienne chez les suba- 1949. Hawaiian Petrographic Province. Bulletin of the dultes de Holanthias borbonius (Valenciennes, 1828) Geological Society of America, 60:1541-1596. (Pisces: Serranidae, Anthiinae). Cybium, series 3, Macfarlane, J.M. 1980(11):23-28. 1923. The Evolution and Distribution of Fishes. 564 pages. Maciolek, J.A. Mayr, E. 1942. Systematics and the Origin of Species, xiv + 334 pages. 1978. Taxonomic Status, Biology, and Distribution of Hawaiian Lentipes, a Diadromus Goby. Pacific Sci- McCosker, J.E. ence, 31(4): 355-362. 1971. A New Species of Parapercis (Pisces: Mugiloididae) Mammerickx, J., R.N. Anderson, H.W. Menard, and S.M. from the Juan Fernandez Islands. Copeia, Smith 1971(4):682-686, 1975. Morphology and Tectonic Evolution of the East- 1977. The Osteology, Classification, and Relationships Central Pacific. Geological Society of America, Bulletin, of the Eel Family Ophichthidae. Proceedings of the 86(1): 111-117, foldout figure. California Academy of Sciences, series 4, 41 (1): 1 —123. Mann, G. 1978. Synonymy and Distribution of Calloplesiops 1954. La Vida de los peces en aquas Chilenas. 2nd edition, (Pisces: Plesiopidae). Copeia, 1978(4):707-710. 339 pages. 1979. The Snake Eels (Pisces, Ophichthidae) of the Ha- Manning, R.B. waiian Islands, with Description of Two New 1969. A Review of the Genus Harpiosquilla (Crustacea, Species. Proceedings of the California Academy of Sci- Stomatopoda), with Descriptions of Three New ences, 42(2): 57-67. Species. Smithsonian Contributions to Zoology, 36:1-41. McCulloch, A.R. Markle, D.F., W.B. Scott, and AC. Kohler 1929. Check-List of the Fishes Recorded from Australia. 1980. New and Rare Records of Canadian Fishes and Australian Museum, Memoir, 5(3):329-436. 174 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
McGowan, J.A. logical Society of America, Memoir, 132:7-22. 1958. The Trochus Fishery of the Trust Territory of the Mouneimne, N. Pacific Islands. 46 mimeographed pages. [A pho- 1977. Liste des poissons de la cote du Liban (Mediter- tocopy of this paper is available in the reprint files ranee orientale). Cybium, series 3, 1:37-66. of the Division of Mollusks, National Museum of Munro, I.S.R. Natural History.] 1945. Postlarval Stages of Australian Fishes, No. 1. Mem- McKenna, M.C. oirs of the Queensland Museum, 12(3): 136-153. 1972. Sweepstakes, Filters, Corridors, Noah's Arks, and 1949. Revision of Australian Silver Breams Mylio and Beached Viking Funeral Ships in Paleogeography. Rhabdosargus. Memoirs of the Queensland Museum, In D.H. Tarling and S.K. Runkorn, editors, Impli- 12(4): 182-223, plates 16-22. cations of Continental Drift to the Earth Sciences, 1:295- 1958. The Fishes of the New Guinea Region. Papua New 308. Guinea Agricultural Journal, 10(4):97-369. [Also is- McKenzie, KG. sued as Territory of Papua and New Guinea, Fisheries 1967. The Distribution of Caenozoic Marine Ostracoda Bulletin, 1.] from the Gulf of Mexico to Australia. In Aspects 1967. The Fishes of New Guinea, xxxvii + 651 pages, 84 of Tethyan Biogeography. The Systematics Associa- plates. tion, Publication, 7:219-238. Myers, G.S. McKinney, J.F., and .V.G. Springer 1936. A New Polynemid Fish Collected in the Sadong 1976. Four New Species of the Fish Genus Ecsenius with River, Sarawak, by Dr. William T. Hornaday, Notes on Other Species of the Genus (Blenniidae: with Notes on the Genera of Polynemidae. Journal Salariini). Smithsonian Contributions to Zoology, 236:1- of the Washington Academy of Sciences, 26(9):376-382. 27. 1940. The Fish Fauna of the Pacific Ocean, with Espe- McKnutt, M., and H.W. Menard cial Reference to Zoogeographical Regions and 1978. Lithospheric Flexure and Uplifted Atolls. Journal Distribution as They Effect the International As- of Geophysical Research, 83:1206-1212. pects of Fisheries. In Proceedings of the Sixth Pacific Mead, G.M. Science Association, 3:201-210. 1970. A History of South Pacific Fishes. In Scientific Myers, R.F. Exploration of the South Pacific, pages 235-251. 1981. Chaetodon flavocoronatus, a New Species of Butterfly- Washington: National Academy Sciences. fish (Chaetodontidae) from Guam. Micronesica, Menard, H.W. 16(2):297-303. 1964. Marine Geology of the Pacific. 271 pages. Myers, R.F., and J.W. Shepard 1973. Depth Anomalies and the Bobbing Motion of 1981. New Records of Fishes from Guam, with Notes on Drifting Islands. Journal of Geophysical Research, the Ichthyofauna of the Southern Marianas. Mi- 78:5128-5137. cronesica, 16(2):305-347. Menon, A.G.K. Nair, R.V., and R. Soundararajan 1977. A Systematic Monograph of the Tongue Soles of 1973. On the Occurrence of the Deep Sea Sting Ray, the Genus Cynoglossus Hamilton-Buchanan (Pisces: Urotrygon daviesi Wallace in Indian Waters. Indian Cynoglossidae). Smithsonian Contributions to Zoology, Journal of Fisheries, 20(l):245-249. 238:1-129. Nelson, G., and N. Platnick 1980. A Revision of the Fringed-Lip Tongue Soles of the 1981. Systematics and Biogeography. xi + 567 pages. Genus Paraplagusia Bleeker, 1865 (Family Cyno- Nelson, G., and N. Rothman glossidae). Matsya, 5:11-22. 1973. The Species of Gizzard Shads (Dorosomatinae) Miller, C.N., Jr. with Particular Reference to the Indo-Pacific Re- 1978. Mezozoic Conifers. The Botanical Review, gion. Bulletin of the American Museum of Natural 43(2) :217-280. History, 150(2): 131-206. Mizuno, S., and Y., Tominaga Nelson, J.S. 1980. First Record of the Scorpaenoid Fish Caracantkus 1974. Fishes of the World, xiii + 416 pages. unipinna from Japan, with Comments on the Char- 1978. Limnichthys polyactis, a New Species of Blennioid acters of the Genus. Japanese Journal of Ichthyology, Fish from New Zealand, with Notes on the Tax- 26(4):369-372. onomy and Distribution of Other Creediidae (In- Moberly, R. cluding Limnichthyidae). New Zealand Journal of 1972. Origin of Lithosphere behind Island Arcs, with Zoology, 5:351-364. Reference to the Western Pacific. Geological Society 1979. Some Osteological Differences between the Blen- of America, Memoir, 132:35-55. nioid Fishes Limnichthys polyactis and L. rendahli, Morgan, W.J. with Comments on Other Species of Creediidae. 1972. Plate Motions and Deep Mantle Convection. Geo- New Zealand Journal of Zoology, 6:273-277. NUMBER 367 175
Norcross, J.J., S.L. Richardson, W.H. Massman, and E.B. de I'Atlantique Sud (1948-1949), 4(1): 12-154, plates Joseph 1-13. 1974. Development of Young Bluefish (Pomatomus salta- Poss, S.G., and W.N. Eschmeyer trix) and Distribution of Eggs and Young in Vir- 1975. The Indo-West Pacific Scorpionfish Genus Ocosia ginian Coastal Waters. Transactions of the American Jordan and Starks (Scorpaenidae, Tetraroginae), Fisheries Society, 103(3) :477-497. with Description of Three New Species. Matsya, Norman, J.R. 1:1-18. 1966. A Draft Synopsis of the Orders, Families and Genera of 1978. Two New Australian Velvetfishes, Genus Para- Recent Fishes and Fish-Like Vertebrates. 649 pages. ploactis (Scorpaeniformes: Aploactinidae), with a Ogasawara, Y., S. Akatsu, and Y. Taki Revision of the Genus and Comments on the 1978. Juvenile Stages and Effect of Salinity on the Sur- Genera and Species of the Aploactinidae. Proceed- vival of Larvae and Juveniles in the Striped Fin- ings of the California Academy of Sciences, 41 (18): 401- gerfish, Monodactylus sebae. Japanese Journal of Ich- 426. thyology, 24 (4): 246-250. 1979. Prosoproctus pataceus, a New Genus and Species of Parin, N.V., B.B. Collette, Yu.N. Shcherbachev Velvetfish from the South China Sea (Aploactin- 1980. Preliminary Review of the Marine Halfbeaks idae: Scorpaeniformes). Japanese Journal of Ichthyol- (Hemiramphidae, Beloniformes) of the Tropical ogy, 26(1): 11-14. Indo-West Pacific. Transactions of the P. P. Shirshov 1980. Xenaploactis, a New Genus for Prosopodasys asperrimus Institute of Oceanology. 97:7-173, foldout figure. [In Giinther (Pisces: Aploactinidae), with Descrip- Russian with English summary.] tions of Two New Species. Proceedings of the Califor- Pearson, J.C. nia Academy of Sciences, 42(8):287-293. 1941. The Young of Some Marine Fishes Taken in Ralston, C. Lower Chesapeake Bay, Virginia, with Special 1981. A New Record of the Pomacanthid Fish Centropyge Reference to the Gray Sea Trout Cynoscion regalis interruptus from the Hawaiian Islands. Japanese (Bloch). Fishery Bulletin of the Fish and Wildlife Journal of Ichthyology, 27(4):327-329. Service, 50:79-102. Randall, H.A., and G.R. Allen Penrith, M.J. 1977. A Revision of the Damselfish Genus Dascyllus (Po- 1967. The Fishes of Tristan da Cunha, Gough Island macentridae) with the Description of a New Spe- and the Vema Seamount. Annals of the South African cies. Records of the Australian Museum, 31(9): 349- Museum, 48(22): 523-548, plate 48. 385. 1976. Distribution of Shallow Water Marine Fishes Randall, J.E. Around Southern Africa. Cimbebasia, series A, 1955a. An Analysis of the Genera of Surgeonfishes (Fam- 4(7): 137-154. ily Acanthuridae). Pacific Science, 9(3): 359-367. 1978. An Annotated Checklist of the Inshore Fishes of 1955b. A Revision of the Surgeon Fish Genera Zebrasoma Southern Angola. Cimbebasia, series A, 4(11): 179- and Paracanthurus. Pacific Science, 9(4):396-412. 190. 1955c. A Revision of the Surgeon Fish Genus Ctenochaetus, Pietschmann, V. Family Acanthuridae, with Descriptions of Five 1930. Remarks on Pacific Fishes. Bemice P. Bishop Mu- New Species. Zoologica, 4(4)-.149-166, plates 1-2. seum, Bulletin, 73:1-24, plates 1-4. 1955d. Fishes of the Gilbert Islands. Atoll Research Bulletin, 1938. Hawaiian Shore Fishes. Bemice P. Bishop Museum, 47:i-xiv, 1-243. Bulletin, 156:1-55, plates 1-18. 1956. A Revision of the Surgeon Fish Genus Acanthurus. Pinkas, L. Pacific Science, 10(2): 159-235. 1961. Descriptions of Postlarval and Juvenile Bonito 1958. A Review of the Labrid Fish Genus Labroides, with from the Eastern Pacific Ocean. California Fish and Descriptions of Two New Species and Notes on Game, 47(2): 175-188. Ecology. Pacific Science, 12(2):327-347. Platnick, N.T., and G. Nelson 1960. A New Species of Acanthurus from the Caroline 1978. A Method of Analysis for Historical Biogeography. Islands, with Notes on the Systematics of Other Systematic Zoology, 27(1): 1-16. Indc-Pacific Surgeonfishes. Pacific Science, Playfair, R.L., and A.C.L.G. Giinther 14(3):267-279. 1866. The Fishes of Zanzibar, xiv + 153 pages, 21 plates. 1961. A Record of the Kyphosid Fish Sectator ocyurus (= Plessis, Y.B., and L.A. Mauge azureus) from the Society Islands. Copeia, 1978. Ichthyologie des iles Marquises. Cahiers du Pacifique, 1961(3):357-358. 21:215-235. 1963. Review of the Hawkfishes (Family Cirrhitidae). Poll, M. Proceedings of the United States National Museum, 1951. Poissons, II: Selaciens et chimeres. In Expedition 114(3472): 389-451, plates 1-16. Oce'anographique Beige dans les Eaux Cotieres Africaines 1964. A Revision of the Filefish Genera Amanses and 176 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Cantherhines. Copeia, 1964(2):331-361. Significance in the Classification of the Grammis- 1968. Caribbean Reef Fishes. 318 pages. tidae. Publications of the Seto Marine Biological Labo- 1972a. A Revision of the Labrid Fish Genus Anampses. ratory, 29(2-3): 157'-190. Micronesica, 8(1-2): 151-195. Randall, J.E., and G.R. Allen 1972b. The Hawaiian Trunkfishes of the Genus Ostracion. 1973. A Revision of the Gobiid Fish Genus Nemateleotris, Copeia, 1972(4):756-768. with Descriptions of Two New Species. Quarterly 1973. Tahitian Fish Names and Preliminary Checklist Journal of the Taiwan Museum, 26(3-4) :347-367. of the Fishes of the Society Islands. Occasional Randall, J.E., and J.E. Bohlke Papers of the Bemice P. Bishop Museum, 24(11):167- 1981. The Status of the Cardinalfish Species Apogon 214. evermanni and A. anisolepis (Perciformes: Apogoni- 1974. Notes and Color Illustrations of Labrid Fishes of dae) with Description of a Related New Species the Genus Anampses. UO, 21:10-16, plates 1-2. from the Red Sea. Proceedings of the Academy of 1975a. Notes on the Hawaiian Filefish Pseudomonacanthus Natural Sciences of Philadelphia, 133:129-140. garretti. Japanese Journal of Ichthyology, 21(4): 223- Randall, J.E., and A. Cea Egana 226. In press. Native Names of Easter Island Fishes. Occasional 1975b. A Revision of the Indo-Pacific Angelfish Genus Papers of the Bemice P. Bishop Museum. Geniacanthus, with Descriptions of Three New Spe- Randall, J.E., and J.R. Chess cies. Bulletin of Marine Science, 25 (3): 393-421. 1980. A New Species of Garden Eel (Congridae: Het- 1975c [1976]. Ostracion trachys, a New Species of Trunkfish erocongrinae) of the Genus Gorgasia from Hawaii. from Mauritius (Ostraciontidae). Matsya, 1:59-62. Pacific Science, 33( 1): 17-23. 1976. The Endemic Shore Fishes of the Hawaiian Is- Randall, J.E., and J.H. Choat lands, Lord Howe Island and Easter Island. Office 1980. Two New Parrot fishes of the Genus Scarus from de la Recherche Scientifique et Technique Outre-Mer, the Central and South Pacific, with Further Ex- Trauvaux et Documents, 47(1973):49-73. amples of Sexual Dichromatism. Zoological Journal 1977. Contribution to the Biology of the Whitetip Reef of the Linnean Society, 70(4): 383-419, 7 plates. Shark (Triaenodon obesus). Pacific Science, 31 (2): 143- Randall, J.E., and C.J. Ferraris, Jr. 164. 1981. A Revision of the Indo-Pacific Labrid Fish Genus 1978a. Marine Biological and Archeological Expedition Leptojulis with Descriptions of Two New Species. to Southeast Oceania. In National Geographic Society Revue Franqaise d'Aquariologie, 8(3):89-96. Research Reports, 1969 Projects, pages 473-495. 1978b. A Revision of the Indo-Pacific Labrid Fish Genus Randall, J.E., and P. Gueze Macropharyngodon, with Descriptions of Five New 1980. The Goatfish Mulloidichthys mimicus n. sp. (Pisces, Species. Bulletin of Marine Science, 28(4): 742-770. Mullidae) from Oceania, a Mimic of the Snapper 1980a. Westward to the Marquesas. Freshwater and Marine Lutjanus kasmira (Pisces, Lutjanidae). Bulletin du Aquarium, 3(3): 7-10, 84-92. Museum National d'Histoire Naturelle, Paris, series 4, 1980b. Revision of the Fish Genus Plectranthias (Serrani- 2:603-609. dae: Anthiinae) with Descriptions of 13 New Spe- 1981. The Holocentrid Fishes of the Genus Myripirslis of cies. Micronesica, 16(1): 101-187. the Red Sea, with Clarification of the Murdjan and 1981a. New Records of Fishes from the Hawaiian Islands. hexagonus Complexes. Natural History Museum of Los Pacific Science, 34(3):211-232. Angeles County, Contributions in Science, 334:1-16. 1981b. A Review of the Indo-Pacific Sand Tilefish Genus Randall, J.E., and R.K. Kanayama Hoplolatilus (Perciformes: Malacanthidae). Fresh- 1972. Hawaiian Fish Immigrants. Sea Frontiers, 18(3): water and Marine Aquarium, 4(12):39-46. 144-153. In press a. Revision of the Labrid Fish Genus Labropsis Randall, J.E., and J.C. Kay with Descriptions of Five New Species. Micronesica, 1974. Stethojulis axillaris, a Junior Synonym of the Ha- 17(1):125-155, 5 plates. waiian Labrid Fish Stethojulis balteata, with a Key In press b. Examples of Antitropical and Antiequatorial to the Species of the Genus. Pacific Science, Distribution of Indo-West-Pacific Fishes. Pacific 28(2): 101-107. Science, 35(3): 197-209. Randall, J.E., and W. Klausewitz In press c. A Review of the Fishes of the genus Cheilodac- 1973. A Review of the Triggerfish Genus Melichthys, with tylus, Subgenus Goniistius, with Description of a Description of New Species from the Indian New Species from Easter Island and Rapa. Occa- Ocean. Senckenbergiana Biologica, 54(l/3):57-69. sional Papers of the Bemice P. Bishop Museum. Randall, J.E., and A. Kotthaus Randall, J.E., K. Aida, T. Hibiya, N. Mitsuura, H. Kamiya, 1977. Suezichthys tripunctatus, a New Deep-Dwelling Indo- and Y. Hashimoto Pacific Labrid Fish. In "Meteor" Forschungs-Ergeb- 1971. Grammistin, the Skin Toxin of Soapfishes, and Its nisse, series D, 24:33-36. NUMBER 367 177
Randall, J.E., and R. Lubbock Akademie van Wetenschappen, Proceedings, C84(l): 1981a. A Revision of the Serranid Fishes of the Subgenus 131-141. Mirolabrichthys (Anthiinae: Anthias), with Descrip- Rau, N., and A. Rau tions of Five New Species. Natural History Museum 1980. Commercial Marine Fishes of the Central Phillipines of Los Angeles County, Contributions in Science, 33:1- {Bony Fishes). 623 pages. 27. Raven, P.H., and D.I. Axelrod 1981b. Labrid Fishes of the Genus Paracheilinus, with De- 1972. Plate Tectonics and Australasian Paleobiogeogra- scriptions of Three New Species from the Philip- phy. Science, 176:1379-1386. pines. Japanese Journal of Ichthyology, 28(10): 19-30. Rehder, H.A. Randall, J.E., K. Matsuura, and A. Zama 1968. The Marine Molluscan Fauna of the Marquesas 1978. A Revision of the Triggerfish Genus Xanthichthys, Islands. Bulletin of the American Malacalogical Union, with Description of a New Species. Bulletin of Inc., Annual Reports, 1968:29-32. Marine Science, 28(4) :688-706. 1980. The Marine Mollusks of Easter Island (Isla de Randall, J.E., and J.E. McCosker Pascua) and Sala y Gomez. Smithsonian Contributions In press. Two New Serranid Fishes of the Genus Anthias to Zoology, 289:1-167. from the Central Pacific. Journal of Aquariculture, Rehder, H.A., and J.E. Randall 2(3):59-69. 1975. Ducie Atoll: Its History, Physiography and Biota. Randall, J.E., and H.A. Randall Atoll Research Bulletin, 175-185(183): 1-40, figures 1981. A Revision of the Labrid Fish Genus Pseudojuloides, 1-29. with Descriptions of Five New Species. Pacific Reid, J.L., E. Brinton, A. Fleminger, E.L. Venrick, and J.A. Science, 35(l):51-74, 3 plates. McGowan Randall, J.E., T. Shimuzu, and T. Yamakawa 1978. Ocean Circulation and Marine Life. In H. Char- In press. A Revision of the Holocentrid Fish Genus Ost- nock and G. Deacon, editors, Advances in Oceanog- ichthys, with Descriptions of Four New Species and raphy, pages 65-130. a Related New Genus. Japanese Journal of Ichthyol- Roeleveld, M.A. ogy- 1972. A Review of the Sepiidae (Cephalopoda) of South- Randall, J.E., and Y.H. Sinoto ern Africa. Annals of the South African Museum, 1978. Rapan Fish Names. Occasional Papers of the Bemice 59(10): 193-313, plates 35-45. P. Bishop Museum, 24(15):291-306. Romer, A.S. Randall, J.E., M.M. Smith, and K. Aida 1945. Vertebrate Paleontology. Second edition, x + 687 1980. Notes on the Classification and Distribution of the pages. Indo-Pacific Soapfish, Belonoperca chabanaudi (Per- 1966. Vertebrate Paleontology. Third edition, x + 468 pages. ciformes: Grammistidae). J.L.B. Smith Institute of Rosenblatt, R.H., J.E. McCosker, and I. Rubinoff Ichthyology, Special Publication, 21:1-8. 1972. Indo-West Pacific Fishes from the Gulf of Chiri- Randall, J.E., and V.G. Springer qui, Panama. Contributions in Science, Natural History 1973. The Monotypic Indo-Pacific Labrid Fish Genera Museum, Los Angeles County, 234:1-18. Labrichthys and Diproctacanthus with Description of Rosewater, J. a New Related Genus, Larabicus. Proceedings of the 1961. The Family Pinnidae in the Indo-Pacific. Indo- Biological Society of Washington, 86(23): 2 79-298. Pacific Mollusca, 1 (4): 175-226. 1975. Labroides pectoralis, a New Species of Labrid Fish 1970. The Family Littorinidae in the Indo-Pacific, Part from the Tropical Western Pacific. UO, 25:4-11, I: Littorininae. Indo-Pacific Mollusca, 2(11): 22, plate 1. 417-506. Randall, J.E., and P. Struhsaker 1972. The Family Littorinidae in the Indo-Pacific, Part 1981. Naso maculatus, a New Species of Acanthurid Fish II: The Subfamilies Tectariinae and Echinininae. from the Hawaiian Islands and Japan. Copeia, Indo-Pacific Mollusca, 2(12): 507-529. 1981(3): 553-558. Rotondo, G.M., V.G. Springer, G.A.J. Scott, and S.O. Randall, J.E., and S.N. Swerdloff Schlanger 1973. A Review of the Damselfish Genus Chromis from 1981. Plate Movement and Island Integration—a Pos- the Hawaiian Islands, with Descriptions of Three sible Mechanism in the Formation of Endemic New Species. Pacific Science, 27(4):327-349. Biotas, with Special Reference to the Hawaiian Rao, DM., and K.S. Rao Islands. Systematic Zoology, 30(1): 12-21. 1981. A Revision of the Genus Scolopsis Cuvier (Pisces: Royce, T.C. Nemipteridae) with Descriptions of Two New Spe- 1978. Ocean Eddies Generated by Sea Mounts in the cies from Indian Waters. Koninklijke Nederlandse North Pacific. Science, 199(4333): 1063-1064. 178 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Russell, B.C., and J.E. Randall Schultz, L.P. 1980 [1981]. The Labrid Fish Genus Pseudolabrus from 1943. Fishes of the Phoenix and Samoan Islands Col- Islands of the Southeastern Pacific with Descrip- lected in 1939 during the Expedition of the U.S.S. tion of a New Species from Rapa. Pacific Science, "Bushnell." United States National Museum, Bulletin, 34(4): 433-440. 180:i-x, 1-316, plates 1-9. Ryan, P.A. 1953a. Family Congridae. United States National Museum, 1980. A Checklist of the Brackish and Freshwater Fish Bulletin, 202(1) :83-85. of Fiji. South Pacific Journal of Natural Science, 1:58- 1953b. Family Isuridae. United States National Museum, Bul- 73. letin, 202(1) :4. Sakai, H., and M. Nakamura 1953c. Family Solenostomidae. United States National Mu- 1979. Two New Species of Freshwater Gobies (Gobiidae: seum, Bulletin, 202(l):229-230. Sicydiaphiinae) from Ishigaki Island, Japan. Jap- 1956. A New Pinecone Fish, Monocentris reedi, from Chile, anese Journal of Ichthyology, 29(l):43-54. a New Record for the Eastern Pacific. Pro- Sale, P.F. ceedings of the United States National Museum, 1980. The Ecology of Fishes on Coral Reefs. Oceanography 106(3365):237-239, plate 1. and Marine Biology, Annual Review, 18:367-421. 1957. The Frogfishes of the Family Antennariidae. Pro- Sardou, J. ceedings of the United States National Museum, 1974. Contribution a la connaissance de la faune ich- 107(3383) :47-105, 14 plates. thyologique Malgache: decourverte de poissons de 1966a. Order Percomorphida. United States National Mu- la famille des Schindleriidae dans le canal de seum, Bulletin, 202(3): 1-13. Mozambique, a Nosy-Be et etude d'une collection 1966b. Order Gobiesocida. United States National Museum, de Schindleria. Cahiers O.R.S.T.O.M., Oce'anographie, Bulletin, 202(3): 137-138. 1967. A Review of the Fish Genus Labracinus Schlegel, Sato, T. Family Pseudochromidae, with Notes on and Il- 1978. A Synopsis of the Sparoid Fish Genus Lethrinus, lustrations of Some Related Serranoid Fishes. with the Description of a New Species. University Ichthyologica, The Aquarium Journal, 39(1): 19-40. of Tokyo, University Museum, Bulletin, 15:i-v, 1-70, Schultz, L.P., and A.D. Welander 12 plates. 1953. Family Dussumieridae: Round Herrings. United Saunders, W.B. States National Museum, Bulletin, 202(1):23-29. 1981. The Species of Living Nautilus and Their Distri- Scott, E.O.G. bution. The Veliger, 24(1) :8-17, 2 plates. 1973. A Tasmanian Record of Sphyma (Sphyma) zygaena Schlanger, S.O. (Linne, 1758) (Sphyrnidae), with a Consideration 1963. Subsurface Geology of Eniwetok Atoll. United of the Species of Hammerhead Sharks in Austra- States Geological Survey, Professional Paper, 260-BB: lian Waters. Records of the Queen Victoria Museum, 991-1066, plates 282-288, chart. 48:1-22. Schlanger, S.O., and G.W. Gillett Scott, J.S. 1976. A Geological Perspective of the Upland Biota of 1959. An Introduction to the Sea Fishes of Malaya, xii + 180 Laysan Atoll (Hawaiian Islands). Biological Journal pages. of the Unman Society, 8(3): 205-216. Scott, R.B., L. Kroenke, G. Zakariadze, and A. Sharaskin Schlanger, S.O., H.C. Jenkyns, and I. Premoli-Silva 1981. Evolution of the South Philippine Sea: Deep Sea 1981. Volcanism and Vertical Tectonics in the Pacific Drilling Project Leg 59 Results. In Initial Reports of Basin Related to Global Cretaceous Transgres- the Deep Sea Drilling Project, 59:803-815. sions. Earth and Planetary Science Letters, 52(2): Scrutton, R.A. 435-449. 1976. Fragments of the Earth's Continental Lithosphere. Schlanger, S.O., and I. Premoli-Silva Endeavour, 35(September):99-103. 1981. Tectonic, Volcanic and Paleogeographic Implica- Seale, A. tions of Redeposited Reef Faunas of Late Creta- 1906. Fishes of the South Pacific. Occasional Papers of the ceous and Tertiary Age from the Nauru Basin and Bernice Pauahi Bishop Museum of Polynesian Ethnology the Line Islands. In Initial Reports of the Deep Sea and Natural History, 4(1): 1-89. Drilling Project, 61:817-827. 1940. Report on Fishes from Allan Hancock Expeditions Schmidt, J. in the California Academy of Sciences. Allan Han- 1925. On the Distribution of the Fresh-Water Eels (An- cock Pacific Expeditions, 9(1): 1-46. guilla) throughout the World, II: Indo-Pacific Re- Shaboneev, I.E. gion. Memoires de I'Academic Royale des Sciences et des 1980. Systematics, Morphoecological Characteristics Lettres de Danemark, Copenhague, Section des Sciences, and Origin of the Genus Trachurus, Horsemacker- series 8, 10(4):329-382, plates 1-2. els. Journal of Ichthyology, USSR, 20(6), 125:787- NUMBER 367 179
799. [In Russian.] versity, Ichthyological Bulletin, 29:567-578, plates Shaklee, J.B., and C.S. Tamaru 87-91. 1981. Biochemical and Morphological Evolution of Ha- 1966a. Fishes of the Sub-Family Nasinae with a Synopsis waiian Bonefishes (Albula). Systematic Zoology, of the Prionurinae. Rhodes University, Ichthyological 30(2): 125-146. Bulletin, 32:633-682, plate 104. Shen, S.-C, and P.-C. Lim 1966b. An Interesting New Eel of the Family Xenocon- 1973. Ecological and Morphological Study on Fish- gridae from Cook Island, Pacific. Annals and Mag- Fauna from the Waters Around Taiwan and Its azine of Natural History, series 13, 8:297-301, plate Adjacent Islands, 5: Study of the Surgeon Fishes 10. (Family Acanthuridae) and Its Distribution with Smith, M.M. Descriptions of Three New Species. Report of the 1979. Rhabdosargus thorpei, a New Sparid Fish from South Institute of Fishery Biology of Ministry of Economic Africa, with a Key to the Species of Rhabdosargus. Affairs and National Taiwan University, 3(1):76-157, Copeia, 1979(4): 702-709. foldout table. Smith-Vaniz, W.F. Shepard, J.W., and K.A. Meyer 1975. Supplemental Description of Rare Blenniid Fish 1978. A New Species of the Labrid Fish Genus Macro- Phenablennius heyligeri Bleeker. Proceedings of the Acad- pharyngodon from Southern Japan. Japanese Journal emy of Natural Sciences of Philadelphia, 127 (6): 53-55. of Ichthyology, 25(3): 159-164. 1976. The Saber-Toothed Blennies, Tribe Nemophini Shepard, J.W., and J.E. Randall (Pisces: Blenniidae). Academy of Natural Sciences of 1976. Notes on the Labrid Fish Stethojulis maculata Philadelphia, Monograph, 19:1-1%. from Japan. Japanese Journal of Ichthyology, Smith-Vaniz, W.F., and V.G. Springer 23(3): 165-170. 1971. Synopsis of the Tribe Salariini, with Descriptions Shih, Tai-Chang of Five New Genera and Three New Species 1980. Marine Magnetic Anomalies from the Western (Pisces: Blenniidae). Smithsonian Contributions to Philippine Sea: Implications for the Evolution of Zoology, 73:1-72. Marginal Basins. In D.E. Hayes, editor, The Tec- Smith-Vaniz, W.F., and J.C. Staiger tonic and Geologic Evolution of Southeast Asian 1973. Comparative Revision of Scomberoides, Oligoplites, Seas and Islands. American Geophysical Union, Mon- Parona, and Hypacanthus with Comments on the ograph, 23:49-75. Phylogenetic Position of Campogramma. Proceedings Shindo, S., and U. Yamada of the California Academy of Sciences, fourth series, 1972. Descriptions of Three New Species of the Lizard- 39(13): 185-253. fish Genus Saurida, with a Key to Its Indo-Pacific Solem, A. Species. UO, 11:1-13; 12:1-14, 2 foldout tables. 1976. Endodontoid Land Snails from Pacific Islands (Mollusca: Sirgel, W.F. Pulmonata: Sigmurethra), Part I: Family Endodontidae. 1980. Two New Species of the Genus Trachycyslis from xii + 508 pages. South Africa (Mollusca, Gastropoda, Pulmo- 1977. Fossil Endodontid Land Snails from Midway nata, Endodontidae). Zoologische Mededelingen, Atoll. Journal of Paleontology, 51 (5) :902-911. 55(9) :97-113, 2 plates. 1981. Land-Snail Biogeography: A True Snail's Pace of Smith, D.G., and P.H.J. Castle Change. In G. Nelson and D.E. Rosen, editors, 1972. The Eel Genus Neoconger Girard: Systematics, Os- Vicariance Biogeography: A Critique, pages 197-221. teology, and Life History. Bulletin of Marine Science, Soljan, T. 22(1): 196-249. 1963. Fishes of the Adriatic. 432 pages. [Translated from Smith, J.L.B. the 1948 Serbo-Croatian original, Fauna et Flora 1953. The Sea Fishes of Southern Africa. Revised enlarged Adriatica, I: Pisces). edition, xv + 564 pages, 107 plates. Springer, S. 1958. Fishes of the Families Tetrarogidae, Caracanthi- 1979. A Revision of the Catsharks, Family Scyliorhini- dae and Synanciidae from the Western Indian dae. Department of Commerce, National Oceanic and Ocean with Further Notes on Scorpaenid Fishes. Atmospheric Administration, National Marine Fisheries Rhodes University, Ichthyological Bulletin, 12:167-181, Service, Circular, 422:i-v +1-152. plates 7-8. Springer, V.G. 1959. Gobioid Fishes of the Families Gobiidae, Per- 1964. A Revision of the Carcharhinid Shark Genera iophthalmidae, Trypauchenidae, Taenioididae, Scoliodon, Loxodon, and Rhizoprionodon. Proceedings of and Kraemeriidae of the Western Indian Ocean. the United Stales National Museum, 115(3493): Rhodes University, Ichthyological Bulletin, 13:185-225, 559-632. plates 9-13. 1967a. Revision of the Circumtropical Shoreflsh Genus 1964. Fishes of the Family Pentacerotidae. Rhodes Uni- Entomacrodus (Blenniidae: Salariinae). Proceedings of 180 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
the United States National Museum, 122(3582): 1 -150, ablennius) of Blenniid Fishes Based on Petroscirtes 30 plates. heyligeri Bleeker. Copeia, 1972(1):64-71. 1967b. The Pacific South American Blenniid Fish, Hyp- Springer, V.G., and A.E. Spreitzer soblennius sordidus. Copeia, 1967(2):461-465. 1978. Five New Species and a New Genus of Indian 1968. The Indo-Pacific Blenniid Fish Genus Stanulus, Ocean Blenniid Fishes, Tribe Salariini, with a with Description of a New Species from the Great Key to Genera of the Tribe. Smithsonian Contribu- Barrier Reef (Blenniidae; Salariini). Proceedings of tions to Zoology, 268:1-20. the Biological Society of Washington, 81 (15): 111 -121. Springer, V.G., and K.D. Woodburn 1971. Revision of the Fish Genus Ecsenius (Blenniidae, 1960. An Ecological Study of the Fishes of the Tampa Blenniinae, Salariini). Smithsonian Contributions to Bay Area. Florida State Board of Conservation Marine Zoology, 72:1-74. Laboratory, Professional Papers Series, 1:1-104. 1972a. Additions to Revisions of the Blenniid Fish Genera Stauch, A., and J. Cadenat Ecsenius and Entomacrodus, with Descriptions of 1965. Revision du genre Psettodes Bennett 1831 (Pisces, Three New Species of Ecsenius. Smithsonian Contri- Teleostei, Heterosomata). Cahiers O.R.S.T.O.M., butions to Zoology, 134:1 -13. Oce'anographie, 3(4): 19-30. 1972b. Synopsis of the Tribe Omobranchini with Descrip- Steene, R.C. tions of Three New Genera and Two New Species 1978. Butterfly and Angelfishes of the World. Volume 1, 144 (Pisces: Blenniidae). Smithsonian Contributions to pages. [A German language edition appeared in Zoology, 130:1-31. 1977.] 1976. Cirrisalarias bunares, New Genus and Species of Steindachner, F. Blenniid Fish from the Indian Ocean. Pro- 1869. Ichthyologische Notizen (viii). Sitzungsberichte der ceedings of the Biological Society of Washington, Akademieder Wissenschqften, Wien, 60(1): 120-139, 7 89(13): 199-203. plates. 1978. Synonymization of the Family Oxudercidae, with 1893. Ichthyologische Beit rage (xvi). Sitzungsberichte der Comments on the Identity of Apocryptes cantoris Kaiserlichen Akademie der Wissenschaften in Wien, Day (Pisces: Gobiidae). Smithsonian Contributions to 102(l):215-243, 3 plates. Zoology, 270:1-14. Stephenson, A.B., and D.A. Robertson 1981a. Comments on Solem's Land-Snail Biogeography, 1977. The New Zealand Species of Trachurus (Pisces: with an Hypothesized Explanation of the Distri- Carangidae). Journal of the Royal Society of New bution of the Endodontidae. In G. Nelson and Zealand, 7(2): 243-253. D.E. Rosen, editors, Vicariance Biogeography: A Cri- Sterba, G. tique, pages 225-230. 1962. Freshwater Fishes of the World. Translated and re- 1981b. Notes on Blenniid Fishes of the Tribe Omobran- vised by D.W. Tucker, 878 pages, 192 plates. chini, with Descriptions of Two New Species. Steyskal, G.C. Proceedings of the Biological Society of Washington, 1980. The Grammar of Family-Group Names as Exem- 94(3) :699-707. plified by Those of Fishes. Proceedings of the Biolog- Springer, V.G., and W.C. Freihofer ical Society of Washington, 93(1): 168-177. 1976. Study of the Monotypic Fish Family Pholid- Strasburg, D.W. ichthyidae (Perciformes). Smithsonian Contributions 1956. Notes on the Blennioid Fishes of Hawaii with to Zoology, 216:1-43. Descriptions of Two New Species. Pacific Science, Springer, V.G., and M.F. Gomon 10(3) :214-267. 1975. Revision of the Blenniid Fish Genus Omobranchus 1958. Distribution, Abundance, and Habits of Pelagic with Descriptions of Three New Species and Notes Sharks in the Central Pacific Ocean. U.S. Fish and on Other Species of the Tribe Omobranchini. Wildlife Service, Fishery Bulletin, 58(138):iv + Smithsonian Contributions to Zoology, 177:1-135. 335-361. Springer, V.G., C.L. Smith, and T.H. Fraser Suess, E. 1977. Anisochromis straussi, New Species of Protogynous 1888. Das Antlitz der Erde. Volume 2, iv + 703 pages. Hermaphroditic Fish, and Synonymy of Aniso- Sulak, K. chromidae, Pseudoplesiopidae, and Pseudochrom- 1977. The Systematics and Biology of Bathypterois (Pisces, idae. Smithsonian Contributions to Zoology, 252:1-15. Chloropthalmidae) with a Revised Classification Springer, V.G., and W.F. Smith-Vaniz of Benthic Myctophiform Fishes. Galathea Report, 1968. Systematics and Distribution of the Monotypic 14:49-108, plates 4-7. Indo-Pacific Blenniid Fish Genus Atrosalarias. Pro- Sverdrup, H.U., M.W. Johnson, and R.H. Fleming ceedings of the United States National Museum, 1942. The Oceans. 1087 pages, 7 charts. 124(3643): 1-12, plate 1. Tabata, R.S., editor 1972. A New Tribe (Phenablenniini) and Genus (Phen- 1981. Transcripts of "Taape: What Needs To Be Done?" NUMBER 367 181
Workshop. University of Hawaii, Sea Grant College ilabrus natalensis Fowler. Bulletin of the British Mu- Program, Working Paper, 46:v + 1-31. seum (Natural History), Zoology, 25(l):27-37. Takita, T. 1977. The Sciaenid Fishes (Croakers and Drums) of the 1975. Spawning and Embryonic Development of Syne- Indo-West Pacific. Transactions of the Zoological So- chogobius hasta in an Aquarium, with Description ciety of London, 33(4):253-541. of Larvae and Juveniles. Japanese Journal of Ich- Tsuda, R.T. thyology, 22(l):31-39. 1976. Occurrence of the Genus Sargassum (Phaeophyta) Talbot, F.H. on Two Pacific Atolls. Micronesica, 12(2):279-282. 1970. The South East Asian Area as a Centre of Marine Tucker, D.W. Speciation: An Ecological Analysis of Causes. In 1956. Studies on the Trichiuroid Fishes, 3: A Prelimi- Symposium on Marine Sciences. Reports of the Aus- nary Revision of the Family Trichiuridae. Bulletin tralian Academy of Sciences, 12:43-50. of the British Museum (Natural History), Zoology, Tamaki, K., M. Joshima, and R.L. Larson 4(3): 73-130, plate 10. 1979. Remnant Early Cretaceous Spreading Center in Tyler, J.C. the Central Pacific Basin. Journal of Geophysical 1967. A Redescription of Triodon macropterus Lesson, a Research, 84(B9) :4501 -4510. Phyletically Important Plectognath Fish. Konink- Taniuchi, T. lijke Nederlandse Akademie van Wetenschappen, Proceed- 1975. Reef Whitetip Shark, Triaenodon obesus, from Ja- ings, series C, 70(l):84-96, plates 1-2. pan. Japanese Journal of Ichthyology, 22(3): 167-170. 1968. A Monograph on Plectognath Fishes of the Super- Thomas, P.A. family Triacanthoidea. Academy of Natural Sciences 1969. Goat Fishes (Family Mullidae) of the Indian Seas. of Philadelphia, Monograph, 16:1-364. Marine Biological Association of India, Memoir, 3:vi + 1970. New Records of Triacanthoid Plectognath Fishes. 1-174, plates 1-8. Notulae Naturae, 435:1-7. Thome, R.F. Tyler, J.C, and C.L. Smith 1963. Biotic Distribution Patterns in the Tropical Pa- 1970. A New Species of Blennioid Fish of the Family cific. In J.L. Gressitt, editor, Pacific Basin Biogeog- Notograptidae from Eastern Australia. Notulae Na- raphy, pages 311-350. turae, 431:1-12. Thorson, T.B. Uyeda, S. 1978. Observations on the Reproduction of the Sawfish, 1977. Some Basic Problems in the Trench-Arc-Back Pristis perotteti, in Lake Nicaragua, with Recom- Arc System. In M. Talwani and W.C. Pittman III, mendations for Its Conservation. In T.B. Thorson, editors, Island Arcs, Deep Sea Trenches, and Back-Arc editor, Investigations of the Ichthyofauna of Nicaraguan Basins, pages 1-14. Lakes, pages 641-650. Vaillant, L.L. Tinker, S.W. 1887. Materiaux pour servir a 1'histoire ichthyologique 1978. Fishes of Hawaii. Pages i-xxx + 1-532 + i-xxxvi. des Archipels de la Societe et des Pomotous. Bul- Toksoz, M.N. letin de la Societe Philomalhique de Paris, series 7, 1975. The Subduction of the Lithosphere. Scientific Amer- ican, 233(5) :88-98. van Balgooy, M.M.J. Tominaga, Y. 1971. Plant-Geography of the Pacific. Blumea, Supplement 1968. Internal Morphology, Mutual Relationships and Volume, 6:1-222. Systematic Position of the Fishes Belonging to the van Balgooy, M.M.J., editor Family Pempheridae. Japanese Journal of Ichthyology, 1975. Pacific Plant Areas. Volume 3, viii + 386 pages, 15(2):43-95. van Steenis, C.G.G.J. Tong, L.J., and CM. Vooren 1963. Pacific Plant Areas, Volume I. Monographs of the 1972. The Biology of the New Zealand Tarakihi, Chei- National Institute of Science and Technology, Republic of lodactylus macropterus (Bloch and Schneider). New the Philippines, 8(1): 1-297. Zealand Ministry of Agriculture and Fisheries, Fisheries van Steenis, G.C.G.J., and M.M.J. van Balgooy, editors Research Bulletin, 6:1-60. 1966. Pacific Plant Areas, 2. Blumea, Supplement Volume, Toriyama, R., T. Sato, T. Hamada, and P. Komalarjum 5:1-312. 1965. Nautilus Drifts on the West Coast of Thailand. Vari, R.P. Japanese Journal of Geology and Geography, 36(1): 1978. The Terapon Perches (Percoidei, Teraponidae): A 149-161, plate 3. Cladistic Analysis and Taxonomic Revision. Bul- Trewavas, E. letin of the American Museum of Natural History, 1973. A New Species of Cichlid Fishes of Rivers Quanza 159(5): 175-340. and Bengo, Angola, with a List of the Known Vari, R.P., and J.B. Hutchins Cichlidae of the Rivers and a Note on Pseudocren- 1978. New Species of Terapon Perches (Percoidei, Ter- 182 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
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If illustrations are intended to be printed separately on coated stock following the text, they should be termed Plates and any components should be lettered as in figures: "Plate 9b_." Keys to any symbols within an illustration should appear on the art and not in the legend. A few points of style: (1) Do not use periods after such abbreviations as "mm, ft, yds, USNM, NNE, AM, BC." (2) Use hyphens in spelled-out fractions: "two-thirds." (3) Spell out numbers "one" through "nine" in expository text, but use numerals in all other cases if possible. (4) Use the metric system of measurement, where possible, instead of the English system. (5) Use the decimal system, where possible, in place of fractions. (6) Use day/month/year sequence for dates: "9 April 1976." (7) For months in tabular list- ings or data sections, use three-letter abbreviations with no periods: "Jan, Mar, Jun," etc. Arrange and paginate sequentially EVERY sheet of manuscript—including ALL front matter and ALL legends, etc., at the back of the text—in the following order: (1) title page, (2) abstract, (3) table of contents, (4) foreword and/or preface, (5) text, (6) appendixes, (7) notes, (8) glossary, (9) bibliography, (10) index, (11) legends. i