CalCOFI Atlas 33

Transcript

1 THE EARLY STAGES IN OF THE FISHES CALIFORNIA CURRENT REGION CALIFORNIA FISHERIES COOPERATIVE OCEANIC INVESTIGATIONS ATLAS NO. 33 BY THE SPONSORED STATES OF COMMERCE DEPARTMENT UNITED OCEANIC AND ATMOSPHERIC ADMINISTRA nON NATIONAL MARINE FISHERIES SERVICE NATIONAL CENTER FISHERIES SCIENCE SOUTHWEST SUPPORTED BY A CONTRACT WITH THE IN PART STATES DEPARTMENT OF THE INTERIOR UNITED MANAGEMENT SERVICE MINERALS CONTINENTAL SHELF REGION OUTER PACIFIC Editor Geoffrey Moser H. National Marine Fisheries Service Southwest Fisheries Science Center La Jona, California

2 THE CALCOFI SERIES ATLAS is the in a series of atlases containing data on the hydrography and plankton from the This thirty-third the Fisheries Current. The field work was carried out by the California Cooperative Oceanic of California region a program sponsored Investigations, agencies: following by the Fish and Game of California Department National and Administration, National Marine Fisheries Service Atmospheric Oceanic California, Scripps Institution of Oceanography University of Atlases are issued as individual units as they become physical, They provide processed available. CalCOFI and planktonic the California Current region of life history information on and biological chemical, measurements more the Each number may contain one or of contributions. A general description of the region. organisms program CalCOFI its objectives appears in the preface of Atlas No.2. with CalCOFI Committee: Doyle A. Hanan Michael M. Mullin Michael F. Tillman CalCOFI atlases related to the subject of Atlas No. 33: Recent No. 31. Moser, H. G., R. L. Charter, P. E. D. A. Ambrose, S. R. Charter, C. A. Myer, E. M. Sandknop, Smith, the in fish larvae and eggs of California Current region: Taxa with Distributional 1993. atlas W. and Watson. or more total larvae, 1951 through 1984. 1000 pp. 233 No. Moser, H. G., R. L. Charter, 32. Charter, E. P. A. Ambrose, S. R. Smith, C. A. Myer, E. M.Sandknop, D. and W. Watson. 1994. Distributional atlas of fish larvae in the California Current region: Taxa with less than 1000 total 1951 through 1984. 181 pp. larvae, ATLAS 33 CALCOFI NO. ISBN 0-935868-82-8 of Congress Catalogue Card Number Library 67-4238 George T. Hemingway, CalCOFI Coordinator Marine Life Research Program Scripps Institution of Oceanography La Jolla, 92093-0227 California 1996 April Printed in United States of America by the Kansas Press, Inc., Lawrence, Allen Paper). Printed on paper meeting the @ of ANSIINISO Z39.48-1992 (Permanence of requirements

3 PREFACE The of this guide are with the scientific investigations on the collapse of the Pacific sardine origins of the California Cooperative Sardine Research Program, later in the founding the 1940s and population late the California Oceanic Fisheries Investigations. Among the diverse assemblage CaICOFI, renamed Cooperative Ahlstrom, the its environment was Dr. Elbert H. "Ahlie" and who quickly took of scientists investigating sardine saw he and taxonomy was One of Ahlie's scientific talents a great history studies. early in role the lead life larvae about many other fish learning whose eggs and the were collected in plankton tows for potential species of the sardine. He began a vigorous research program to increase the and of identifiable taxa number those with and species. papers describing the eggs of larvae of a variety of progress this with documented a series of about 100 early an identification guide to the Ahlie life history stages and work on 1978, In I began most common species encountered in the CalCOFI ichthyoplankton collections. Early progress on this ended of the of M. Daniel Richards, J. (William colleagues his untimely five thereafter, shortly and, 1979 in death with his Michael P. Fahay, Arthur W. Kendall, Jr., and Sally L. Cohen, joined me in organizing an Richardson) international to honor Ahlie and his scientific achievements. Ahlie's notes and unpublished manuscripts symposium of California, San Diego made the symposium was held at the University available the contributors and were to was "Ontogeny and Systematics of Fishes," published by The American The volume entitled, 1983. in symposium Ichthyologists and Herpetologists in 1984. Society of I sought the from 1988, Minerals Management Service, In funding of Interior, to Department the U.S. illustrations required to publish begin comprehensive version the a more the guide originally envisioned by of and me. The proposal was successful, largely Ahlie the efforts of Dr. Gary D. Brewer of MMS. In through September Barbara Sumida McCall began the illustrations and my research group, William Watson, David 1989, of the guide. work Elaine M. Sandknop, began to R. on the various chapters Ambrose, Charter, and A. Sharon obvious became it soon but of about 500 pages covering about 250 taxa a book Initially, planned to we produce became this an underestimate. Work that each chapter was a significant research project, with exciting new on discoveries. What was to be a three-year project has taken six years to complete. Consequently, the book tripled of taxa and illustrations. A total of 1171 original in includes more than twice the planned number size and 181 unpublished illustrations on hand before the project was six years. With the illustrations the completed over time increased 1352 original illustrations. The extra of spent on the book has its scope the book a total began, has and to usefulness coverage scientists studying ichthyoplankton in all oceans. and its this our group have labored prodigiously on of project, showing great patience and research Members the of duties other the research with of the book along sections their to asked being while forbearance complete more William sole author or in collaboration, wrote as than division. Watson, the chapters, produced many half the illustrations, was responsible for the reduction of original illustrations and the paste-up of the plates, and of all assisted us with his writing skills during the development of our chapters. His central role in the realization of Ambrose this will be obvious to anyone using it. David of made major advances in our knowledge of the book ontogeny of Alepocephaloidei, Alepisauroidei, Gadiformes, Cottoidei, and Scombroidei. He was responsible for role of captions and headers and took the lead figure in preparing the indexes for the book. Sharon the paste-up and greatly the Elopomorpha chapters of improved our knowledge of eel for most was Charter responsible she was responsible for the sections on the Lampridiformes, Mirapinnidae, Pleuronectidae, and leptocephali. Also, She shared Cynoglossidae, contributions to chapters on the Carangidae and Bothidae. and important made indexes. of the Elaine Sandknop was responsible for chapters production responsibility Ambrose David for with on the Melamphaidae, Apogonidae, Mugilidae, Polynemidae, Howellidae, and Trichiuridae, and worked closely other comprehensive the Clupeiformes, Scopelarchidae, and on chapters in the book. Her with William Watson us of and the stages was helpful to each of egg as we developed our chapters. The editing of yolk-sac knowledge manuscript for this book was a long and arduous process and could not have been accomplished without the deeply enormous member of our research team. I am each indebted to them for the of effort they assistance their extended this process and for during dedication throughout this project. 111

4 Former La Laboratory scientists were important contributors and authors on chapters of the guide. Jolla H. Ahlstrom's on Bathylagidae and Myctophidae was important to the completion of those sections. work Elbert Elizabeth G. after work on the book began but made important contributions to sections on retired Stevens soon series Barbara MacCall worked out the larval Sumida for the numerous Synodontidae. and Bregmacerotidae sanddab species CalCOFI this and the was crucial to the completion of the chapter on in collections L. research who currently leads John on age and growth Paralichthyidae. Butler, in the La Jolla of fishes much contributed the chapter on Microstomatidae. Laboratory, to following scientists from other laboratories contributed chapters in special areas of their expertise: The of School of Fisheries, University Brogan, Washington, Seattle, Washington-Lutjanidae; Mr. W. Dr. Michael Bruce C. Marine Fisheries Service, Southwest Fisheries Science Center, Honolulu Laboratory, Mundy, National Dr. Fisheries 1. Richards, National Marine Hawaii-Bramidae; Service, Southeast Fisheries Honolulu, William Michael Florida- Triglidae; and Mr. A. Shane, Hubbs-Sea World Miami, Center, Miami Science Laboratory, Institute, San Diego, Califomia-Polyprionidae. Research illustrations the heart The are the guide. George Mattson produced many illustrations during his career of our throughout and his images, some previously unpublished, are found in the chapters. He used a split- laboratory pen for all his illustrations and the depth point is unique field with this technique of in line drawings of achieved the early life stages of fishes. The two years given to the project by Barbara Sumida McCall set the standard for the other Her images, exquisite illustrators. their detail follow from her experience as an in and accuracy, true from great and talent. The same is her for William Watson's illustrations. accomplished biologist artistic Nancy Robert C. Walker worked on the project for more than two years and Mary T. Vona was with Arthur: and in the illustration of fish eggs and larvae before joining for a year. All are fine artists us had no experience and the They were trained by William Watson and guided by each project. the as they worked on their of authors Esther M. Perez did all chapters. word processing and formatting of the guide. Her talents in design and the editing and her dedication became increasingly more important as the book progressed. This book could not have been completed her. We are deeply grateful for her efforts. without suggestion publish this book The to the CalCOFI Atlas series came from Dr. John R. Hunter, Chief in of the Fisheries Resources Division of the La Coastal This idea was enthusiastically accepted by Jolla Laboratory. the CalCOFI Committee and the collaboration with George T. Hemingway, the CalCOFI Coordinator, was an enjoyable experience. and inadequacies are part Mistakes any of of this scope and complexity and undertaking ultimately come to rest with the editor. I offer my apologies to all for any shortcomings that come to light. H. Geoffrey Moser Southwest Fisheries Science Center La Jolla Laboratory iv

5 CONTENTS PREFACE. . HI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... v CONTENTS ACKNOWLEDGEMENTS x ... G. H .. 1 INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Moser By Background 1 ... .. Geographic Coverage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Coverage Taxonomic List of Taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Terminology ... Principles and 27 Stage 28 ... Egg Larva Yolk-sac ... 30 Stage Larval Stage ... 31 ... 36 Transformation Stage Juvenile ... 36 Stage ... 37 Abbreviations ... 38 Glossary of Format and Methods ... Explanation 44 Orders ... . . . . . . . . . . . . . . . . . . . . . 44 Introductions to Families 45 ... Introductions to ... Descriptions 45 Species Identify Early Stages of Fishes How this Guide ... 50 to with ... 73 DESCRIPTIONS By S. R. Charter and H Elopomorpha. Moser ... 74 G. Elopiformes By Elopidae. Charter and H G. R. ... 75 S. Moser Albuliformes Albulidae. S. R. Charter and H G. Moser . ... 79 By Notacanthiformes. By H G. Moser and S. R. Charter . ... 82 Notacanthidae. By G. Moser and S. R. Charter ... 82 H By S. Charter and H G. Moser . ... 86 AnguHliformes. R. Muraenoidei G. R. Charter and H S. Moser ... 88 By Muraenidae. Congroidei 93 S. R. Charter ... By Ophichthidae. By S. Charter ... 100 Congridae. R. By R. Charter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119 Derichthyidae. S. Nemichthyidae. S. R. Charter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122 By Serrivomeridae. By S. R. Charter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131 Nettastomatidae. By S. R. Charter . ... 135 Incertae sedis. S. R. Charter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 By By R. Charter ... 144 Saccopharyngiformes. S. 145 S. R. Charter ... By Cyematidae. Saccopharyngidae. By S. R. Charter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151 Eurypharyngidae. By S. R. Charter . ... 155 Clupeiformes. By Watson ... 158 W By W. Clupeidae. and E. M Sandknop ... 159 Watson Engraulidae. By W Watson and E. M Sandknop ... 173 Salmoniformes. By H G. Moser ... 184 v

6 Argentinoidei By Argentinidae. Moser . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185 H G. • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. E. H Ahlstrom and 188 H Moser Bathylagidae. By G. Moser and J. Microstomatidae. Butler . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208 By H L. H G. Moser ... 216 By Opisthoproctidae. Alepocephaloidei D. A. Ambrose 224 ... By Alepocephalidae. By Platytroctidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234 A. D. Ambrose Salmonoidei G. Moser ... 240 H By Osmeridae. H G. Moser Stomiiformes. W. Watson ... 244 By and By Gonostomatidae. ... 247 W. Watson Stemoptychidae. By W. Watson ... 268 284 W. Watson . ... By Phosichthyidae. By Stomioidea. H ... 295 G. . Moser By Chauliodontidae. G. Moser ... 297 H H G. Moser . 301 ... By Stomiidae. H Moser . ... 305 Astronesthidae. G. By H G. Moser ... 308 Melanostomiidae. By By H G. Moser ... 321 Malacosteidae. By Idiacanthidae. Moser . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325 H G. By Aulopiformes. W. . ... 328 Watson Aulopoidei Aulopidae. By A. Ambrose ... 329 D. Scopelarchidae. By Watson E. M Sandknop ... 332 W. and 344 Watson and E. M Sandknop ... W. Notosudidae. By Alepisauroidei Synodontidae. By G. Stevens H G. Moser . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 348 E. and By Paralepididae. D. Ambrose ... 352 A. Anotopteridae. A. Ambrose ... By D. 369 D. A. Ambrose . ... 373 By Evermannellidae. By D. A. Ambrose ... Alepisauridae. 379 Myctophiformes. By G. Moser 382 H ... By Neoscopelidae. H Moser . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 G. ... H G. Moser 387 E. H Ahlstrom . and By Myctophidae. By Gadiformes. D. A. ... 476 Ambrose Bregmacerotidae. By E. G. Stevens and H G. Moser. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477 A. Ambrose ... 483 D. Macrouridae. By By A. Ambrose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 500 Moridae. D. By D. A. Ambrose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 508 Merlucciidae. Ophidiiformes. By A. Ambrose ... 512 D. Ophidiidae. By D. A. Ambrose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 513 Carapidae. By D. A. Ambrose . ... 532 Bythitidae. By A. ... 538 D. Ambrose Batrachoidiformes W. Watson ... 546 Batrachoididae. By Lophiiformes. By W. . ... 550 Watson Lophioidei Lophiidae. By W. Watson ... 553 Antennarioidei Vi

7 Antennariidae. By Watson . ... 559 W. Ogcocephaloidei Ogcocephalidae. By ... Watson 563 W. Ceratioidei W. 567 Watson ... Caulophrynidae. By ... 571 W. Watson By Melanocetidae. Watson Oneirodidae. ... 575 By W. . W. Watson ... 582 By Ceratiidae. Gigantactinidae. By ... W. Watson 589 Watson ... 592 W. Linophrynidae. By By Gobiesociformes. W. ... 596 Watson Gobiesocidae. By ... 597 W. Watson W. . ... 608 Watson Atheriniformes. By W. Watson ... 608 Atherinidae. By By W. Watson ... 620 Cyprinodontiformes. By Cyprinodontidae. Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 620 W. By Beloniforrnes. . ... , ... , ... 625 W. Watson W. 627 ... By Scomberesocidae. Watson By Belonidae. W. 630 ... Watson By Watson ... 634 Hemiramphidae. W. ... W. Watson . 643 By Exocoetidae. G. By Charter and H R. Moser . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 659 Lampridiformes. S. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. R. Charter and 661 G. Moser. H By Lophotidae. H By R. Charter Radiicephalidae. S. G. Moser . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 665 and Trachipteridae. By S. R. Charter and H G. Moser ... 669 . ... 678 W. Watson Beryciformes. By Berycoidei . ... 679 W. Watson Diretmidae. By By Anoplogastridae. Watson ... 683 W. W. Watson . Holocentridae. 686 By ... Stephanoberycoidei E. M Sandknop and W. Watson ... 692 By Melamphaidae. Mirapinnidae. By R. Charter and H G. Moser '" ... 713 S. Sandknop By and E. M Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 717 W. Syngnathiformes. Aulostomoidei By W. Watson and E. M Sandknop ... 718 Fistulariidae. W. Watson and E. M Sandknop ... 725 Centriscidae. By Syngnathoidei Syngnathidae. By Watson E. M Sandknop ... 728 W. and ... H G. 732 Moser By Scorpaeniformes. Scorpaenoidei Scorpaenidae. By G. Moser ... 733 H Triglidae. By W. J. Richards ... 797 Anoplopomatoidei H G. Moser . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 807 Anoplopomatidae. By Hexagrammoidei By Hexagrammidae. D. A. Ambrose ... 811 Cottoidei Cottidae. D. A. Ambrose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 821 By Psychrolutidae. By D. A. Ambrose ... 841 VB

8 Agonidae. By A. Ambrose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 844 D. By D. Ambrose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 860 Cyclopteridae. A. W. Perciformes. ... 872 Watson By . Percoidei M Shane, W. Watson, and H G. Moser ... 873 A. By Polyprionidae. By . ... 876 W. Serranidae. Watson Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 900 W. Priacanthidae. By E. M Sandknop and Apogonidae. Watson ... 904 By W. By Watson ... 912 Echeneidae. W. By Carangidae. W. S. R. Charter, H G. Moser, Watson, Ambrose and E. D. Sandknop ... 914 A. M By Nematistiidae. H G. Moser . ... 955 By D. A. Coryphaenidae. ... 959 Ambrose Bramidae. By G. Moser B. C. Mundy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 964 H and H Moser . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 973 G. By Caristiidae. W. Watson and M W. Brogan . ... Lutjanidae. By 977 G. Moser . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 990 H Malacanthidae. By W. Watson ... 995 Lobotidae. By Gerreidae. W. Watson . ... 998 By Haemulidae. By Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1002 W. ... By W. Watson and E. M Sandknop ... 1013 Sparidae. Sciaenidae. By G. Moser . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1017 H Mullidae. W. Watson ... , ... 1034 By Kyphosidae. By W. Watson ... 1038 Ephippidae. By A. Ambrose . ... 1047 D. By W. . ... 1050 Chaetodontidae. Watson W. ... 1054 By Watson Pomacentridae. W. Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cirrhitidae. 1064 By .. By W. Watson ... 1068 Opistognathidae. By E. M Sandknop and W. Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1072 Howellidae. Mugiloidei E. M Sandknop and W. Watson ... 1078 Mugilidae. By Polynemoidei Polynemidae. E. M Sandknop and W. Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1082 By By . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Watson Labroidei. .. 1088 W. By Watson Labridae. ... 1088 W. Scaridae. W. Watson ... 1104 By Zoarcoidei. By W. Watson ... 111 0 Bathymasteridae. By Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1112 W. By Watson ... 1116 Stichaeidae. W. By W. Watson ... 1120 Pholidae. Anarhichadidae. By Watson ... 1127 W. Trachinoidei. W. Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1130 By Chiasmodontidae. By W. Watson and E. M Sandknop . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1131 AmmodYtidae. By Watson ... 1138 W. By ... Watson Uranoscopidae. 1144 W. Blennioidei. W. Watson . ... 1148 By Tripterygiidae. By W. Watson ... 1148 Labrisomidae. W. Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1151 By Clinidae. By W. Watson ... 1164 viii

9 Chaenopsidae. By Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1170 W. By W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1177 Dactyloscopidae. Watson Blenniidae. By ... 1182 Watson W. . Icosteoidei By Icosteidae. W. ... 1201 Watson Callionymoidei By Watson ... 1205 Callionymidae. W. W. Watson . Gobioidei. 1208 By ... By Watson ... 1209 Eleotridae. W. W. ... 1214 By Gobiidae. Watson W. Watson . ... 1246 By Microdesmidae. Sphyraenoidei Sphyraenidae. W. Watson and E. M By ... 1250 Sandknop Scombroidei. D. A. Ambrose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1257 By By Gempylidae. Ambrose ... 1258 D. A. By Scombridae. A. Ambrose ... 1270 D. By E. M Sandknop and W. Watson ... 1287 Trichiuridae. Stromateiodei. By Watson ... 1295 W. By Watson . ... 1297 Centrolophidae. W. Nomeidae. W. Watson By 1300 ... Tetragonuridae. By W. Watson . ... 1312 Stromateidae. By W. Watson ... 1318 Pleuronectiformes. By G. Moser 1323 H ... Pleuronectoidei By Paralichthyidae. Moser and B. Y. Sumida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1325 G. H Bothidae. Charter Moser and S. R G. ... . . . . . . . . . . . . . . . .. 1357 By H Pleuronectidae. S. By Charter and H G. Moser ... . . . . . . . . . . . . . . . .. 1369 R. Soleoidei Achiridae. By G. Moser ... 1404 H H By R. Charter and Cynoglossidae. G. Moser ... 1408 S. Tetraodontiformes. By W. Watson ... 1415 Balistoidei Balistidae. By Watson ... W. 1417 By Monacanthidae. W. . ... 1422 Watson Watson W. 1425 . ... Ostraciidae. By Tetraodontoidei Tetraodontidae. By W. Watson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1428 Diodontidae. By W. Watson . ... 1435 Molidae. By Watson ... 1439 W. CITED LITERATURE ... 1442 INDEX OF SCIENTIFIC NAMES ... 1492 INDEX OF COMMON NAMES ... 1503 IX

10 ACKNOWLEDGEMENTS project the Southwest a joint is guide This of Benjamin thank and Remington Rasmussen Sherrie for Fisheries the (SWFSC), Center Science Minerals MMS the of their assistance in the administration of the (MMS) of Department the Service Management Roy Allen provided helpful grant. Ken Raymond and Oceanic Interior, Cooperative California and the advice on graphics. Fisheries enthusiastic (CaICOFI). Investigations The We deeply to the many people are indebted who of MMS was D. Gary support afforded Dr. by Brewer this produced guide: original used illustrations in J. Frank to obtaining the initial funding. Dr. critical Arthur, C. Robert Nancy MacCall, Sumida Barbara of the MMS provided support throughout Manago T. Walker, Mary Vona, M. Henry Mattson, George course the Hunter, R. John Dr. of project. Chief, C. Orr, Raymond, Mundy, Bruce Kenneth S. Richard Fisheries Resources Division, gave us Coastal the Kubo, Eloisa Gary Jordan, Atsuhiro A. Feeney, F. the out guide the research for carry to and opportunity Beverly and Matus-Nivon, M. Attributions for Vinter. throughout the project. Dr. Izadore us encouraged illustration species the in given are each descriptions. Southwest Fisheries Science Director of the Barrett, the Michael F. of project, Dr. the Center at start Marine Vertebrates Collection The the of Scripps and Dr. Director, Science Richard A. Tillman, current Institution of Oceanography was a major resource Neal, Director Deputy SWFSC heartily have of the and the was as project, this throughout assis- advice funds Additional work. the supported and encouraged curator of H. Richard Dr. by given tance Rosenblatt, needed enlarge the coverage and scope to of the guide H. by and collection, the Cynthia and Jr. Walker, J. and by Drs. Hunter, Tillman, Neal were made available principal our was collection The Klepadlo. source of Manago. Dr. Dr. of efforts the through MMS, and by data gathering meristic specimens to supplement for the and Neal generously Esther permitted Smith E. Susan transforming of an literature and source important was the do to duties many her from time take to Perez M. identification for needed specimens juvenile and the of entire manu- word processing and formatting for the larval species and for descriptions. Computer series to central were contributions Esther's script. the summaries capture of the in specimens for localities of this book. completion collection geographic ranges useful were in establishing provided. This always of species and graciously were helped with various SWFSC the at people Many guide the without completed been have not could an this on based was study The guide. of aspects Collection Vertebrates of the SIO Marine availability over collection ichthyoplankton a period accumulated the encouragement and assistance and Rosenblatt of Dr. scientif- debt to all the years of and we 45 owe a great and staff. his ic and ships' those who carried out We cruises. crews sorters many the to indebted are identifiers who and source Another major adult of larval, juvenile, and skillfully separated eggs, larvae, and juveniles the fish of Angeles Los Museum History Natural the was fishes from Richard plankton. L. Charter, CalCOFI the course the Throughout County. Dr. this of project, our is manager, data all ichthyoplankton partner in numerous Robert J. Lavenberg kindly loaned speci- us ichthyoplankton computer his and work CalCOFI to needed were mens that series ontogenetic complete this of support beginning the from essential been has and We are deeply grateful to Bob and meristic lists. Mark project. student and Frostad, Eric Yip, several and Seigel Richard Dr. A. Jeffrey curators, assistant his Upward College Mudd Harvey the from interns Bound needed much their for support. Feeney F. Dr. Daniel and specimens larval for search in aided Program the Division. Sciences Life Curator, Chief Cohen, M. helped Smith, and Mark Yip, the student interns Leo throughout support and advice helpful offered LACM, produce radiogJ;aphs and meristic data. Debra Losey, this project. SWFSC librarian, made numerous searches literature the given assistance The project. entire during us for thank Drs. We Steven B. Reilly and Timothy Barbara by con- on Watkins Barbara and Engstrand of the Marine SWFSC, Mammal Division, Gerrodette of to the completion tracts and supplies was essential for marine obser- mammal on tows neuston including the us assisted Hostler Virginia project. the many with Robert Pacific tropical and eastern the to vation cruises this with We actions personnel project. associated tows. L. Pittman and the other observers who made the x

11 The Inter-American the of collections plankton Tropi- sen. The kindness and generosity extended by Panama Laboratory cal Tuna Commission at Achotines, Dr. during Dr. Bertelsen and that Jargen Nielsen source a rich were of material for tropical We species. visit research and numerous the subsequent loan of thank Director Joseph, Dr. James IA for of the TIC are specimens appreciated. deeply allowing to use these collections and us Dr. Daniel journal of all authors acknowledge We gratefully Robert Margulies, Dr. Scholey, Vernon Olson, J. P. papers manuscripts were illustrations which from or collections the making for Wexler B. Jeanne and obtained. publications were major The following Many available. species edge western the from of the illustrations used in sources guide and for this we water mass central the from obtained were area study publishers, authors, and of those thank the editors Other tropical studies came of Valerie J. Loeb. material those illustra- publications reproduce to permission for the collections from of Marine the Institute Hawaii of Systematics of Fishes (1984), and Ontogeny tions: Biology, University thank Dr. of Hawaii and we ofIchthyologists Herpetologists; American Society and A. for The Clarke the use of that material. Thomas Oceanic Ichthyoplankton in the Western Studies the on the of the Honolulu Laboratory of collections plankton Press, Kyushu North Pacific, University ed. Ozawa, T. this for specimens of important an were SWFSC source (1986g); An Atlas of Early Stage Fishes in Japan, the C. available material made this Mundy Bruce guide. to Press, Okiyama, (1988a); ed. M. Tokai University species on information much provided also us and Laboratory Guide Early the to History Stages of Life distributions. advice his for grateful are We and Northeast A. Matarese, C. A. Fishes, Pacific W. assistance throughout generously this given project. Kendall, Jr., Blood, M. (1989). Vinter M. B. D. and collections Numerous were other and associated data the made available to us for gener- this through study All chapters this scientif- rigorous received guide of osity people. many of H. Sharon of collections The reviews group experts on each ic from of fishes. The contained critical specimens Kramer nearshore many of drafts of and lengthy were sections these of many all from specimens us loaned Matsui Tetsuo species. his much detailed contained descriptions that required in midwater collections the Southern Bight California effort and patience. to of each grateful deeply are We provided unpublished data. and allowed Preston Porter valuable reviewers these from time taking for their of flatfish from species several us use specimens to research to help us with comments, Their project. this provided Richard Dr. experiments. rearing Harbison improved and criticisms, vastly suggestions quality the him the in divers other and by captured specimens and usefulness Reviewers guide. of the sections the and open specimens sea. Fernando Nostratpour supplied they reviewed are (Cyprino- as K. follows: W. Able Stephen Birch Aquarium-Museum, the at reared Allen (Gobiesociformes), C. C. L. G. dontiformes), Scripps Institution Behrstock of Oceanography. Robert Barham (Aulopiformes, Serranidae), E. G. Baldwin the provided material from his collections in Gulf of (Paralichthyidae), M. S. W. M. Brogan (Blennioidei), We thank California. Life (Marine Tapper S. Guy Busby (Agonidae, Cyclopteridae), D. M. Cohen Group, Research SIO) B. advice on technical for Scom- (Beloniformes, Collette graphics. (Salmoniformes), B. SIO) Center, Smith M. Stuart (Geological sup- Data broidei), J. G. Ditty (Echeneidae, Coryphaenidae, with us plied locality for data SIO We expeditions. W. Stromateidae), Ephippidae, Sciaenidae, Lobotidae, Mark thank D. Annie W. Ohman and Townsend classification), and names Eschmeyer N. (taxonomic Research Group, Life (Marine P. M. Fahay (Gadiformes, Ophidiiformes), R. F. SIO) for providing station Feeney Fritzsche R. Psychrolutidae), (Cottidae, A. data from the SIO Invertebrate Collection Data (Syngnathiformes), Base (http://gdcmp (Trichiuridae), D. A. helpful F. J. Gago 1. uscd.eduJplankton) and for Hensley the manuscript. Ernesto Calix and comments (Pleuronectiformes), M. H. Horn on (Stromateoi- (Clupeiformes, Houde D. E. Bregmacerotidae, dei), made Analytical (MEC Isham William Systems) the D. G. Johnson Serrani- (Polyprionidae, of Callionymidae), helpful comments on the descriptive sections dae, Howel- manuscript. Caristiidae, Malacanthidae, Much Bramidae, background information of the for lidae), D. (HGM) of us L. (Mycto- Kawaguchi K. Jones (Labroidei), myctophids larval one by obtained was phiformes), during A. a Dana Collections at the Anoplo- (Scorpaenidae, Jr. Kendall, W. 9-month study of the Icos- Psychrolutidae, Hexagrammidae, pomatidae, Zoological Museum, University That of Copenhagen. by study was Foundation's made possible Carlsberg the teidae), Myctophiformes, Klepadlo (Stomiiformes, C. Triglidae, Batrachoidiformes, Mullidae, Carangidae, Erik Bertel- Johannes Schmidt Stipendium through Dr. Xl

12 Kyphosidae, Sphyraenidae, Mugilidae, Gobioidei, grateful to George T. Hemingway, We are CalCOFI Tetraodontiformes), T. E. Laidig Pleuronectiformes, and (Doyle Coordinator, A. the CalCOFI Committee J. R. (Salmoniformes, (Scorpaenidae), Lavenberg Michael and Mullin, M. Michael Hanan, Tillman) F. (priacanth- Leis M. J. Exocoetidae), Atheriniformes, project this and their willingness to of for support their idae, Chaeto- Lutjanidae, Nematistiidae, Apogonidae, publish in guide this the Atlas CalCOFI series. George Labridae, dontidae, Tetrao- Pomacentridae, Cirrhitidae, Hemingway was instrumental the in many of aspects Haemuli- K. C. Lindeman (Lutjanidae, dontiformes), of his Without guide. this and development publication Markle (Alepo- V. J. Loeb (Scorpaenidae), D. F. dae), never book it would the for funding gaining in efforts Gadiformes), cephaloidei, (Hexagram- Matarese C. A. support His published. been have and unwavering Trachinoidei, Zoarcoidei, midae, Psychrolutidae, during is truly project this us of encouragement to all McCosker E. J. Pleuronectidae), B. (Anguilliformes), maps appreciated. We thank Roy for drafting the Allen (Stomiiformes, C. Mundy Aulopiformes, Myctophi- in section in and for his help the introductory produc- the and Beryciformes, Caristiidae, formes, introductory brochure Valerie book. the for ing the advertisement Oda Okiyama M. (Pleuronectiformes), sections), D. provided Philbrick much needed computer assistance in E. J. (Aulopiformes), R. J. (Lampridiformes), Olney Kohichi preparing for lists mailing brochure. the (Aulopiformes, Pietsch Paxton W. T. Myctophiformes), Jeffrey Kawaguchi, Leis, William J. Richards, M. Mycto- (Aulopiformes, Richards J. W. (Lophiiformes), and Losey Debra Quero, C. J. J. H. Cynthia Klepadlo, Lutjanidae, phiformes, Serranidae, Priacanthidae, pre- the in instrumental making were Jr. Walker, Haemulidae, Labroidei, Scombroidei), R. H. Rosenblatt publication scientific brochure wide a available to Opistognathidae), Sparidae, (Elopomorpha, Gerreidae, audience; their help is greatly appreciated. We thank (Elopo- Smith G. D. (Scorpaenidae), Sakuma M. K. Foster Diane for assistance. secretarial V. Springer (Blennioidei), B. M. Vinter G. morpha), Pleuronectidae), Psychrolutidae, (Hexagrammidae, H. Ahlstrom H. a great Elbert to debt owe we Finally, J. Walker, Jr., (Batrachoididae, Triglidae, Carangidae, of leading publication the to us path the on started who Mugilidae, Kyphosidae, Polynemidae, Mullidae, way. the along much taught us and book this Pleuronectiformes, Sphyraenidae, Gobioidei, Tetrao- Williams J. T. (Blennioidei). dontiformes), and xii

13 INTRODUCTION H. MOSER G. Background Sars 1865, In the Norwegian biologist G. O. discov- higher each catego- taxonomic character among varies cod that had ered revealing thus eggs, free-floating the ries (Fahay 1983). in At present, most larvae collected knowledge for potential gaining both of stocks parent identified be can at to least family; plankton samples relative- of marine fishes by sampling their and recruits of species whose larvae are however, the proportion planktonic stages. Sars' easily-captured discovery ly in >80% of from regionally varies known a maximum investigations directed intensive stimulated towards the in 10% ca. Indo-Pacific to Atlantic the northeast the describing early life history (ELH) and rearing of ontogenetic Matarese (Kendall and 1994). The use of European Russell (see seas the of fishes stages research characters in systematic has provided new and Ahlstrom 1979; Hempel 1981). Moser 1976; phylogenetic into insights teleost among relationships development Hensen's (1895) gear of the tech- and "Ahlstrom the fishes. Contributors to Symposium" sampling quantitatively for niques ichthyoplankton early the on life information available summarized the provided larvae) and eggs (fish for impetus plank- stages systematic integrated and teleosts it of with ton extent and timing the at determining aimed surveys 1984b); al. this information on adults (Moser et fish important estimating of spawning and stocks of history life of charac- early evaluation comprehensive plankton off surveys of results biomass. The their greatly has ters enhanced our ability to identify fish Johannes by conducted Norway Hjort first the in and eggs regional published recently Likewise, larvae. century led this of fluctua- that postulate to him decade ichthyoplank- identification guides aid greatly in ELH interannual to tions in year-class strength were due Kendall Matarese (see 1994). and ton investigations variation in survival stage of the first-feeding larval National Marine Fisheries Service the of Scientists that proposed Hjort 1914). (Hjort the was starvation Northwest the for guides such produced have Atlantic and mortality early in factor causal year- that principal Pacific Northeast the et al. (Matarese and 1983) (Fahay prey of organisms populations in fluctuations to-year 1989). and their to first-feeding fish larvae ulti- availability guide This to the stages early ofthe fishes teleost of mately determined the size the year class. future of the California Current region, the third in of series oceanography hypothesis by in Hjort his incorporated regional guides, is the result of five decades of NMFS by currents ocean that suggesting could reduce survival sponsored the California by research ichthyoplankton transporting and from away larvae favorable eggs Oceanic Fisheries Investigations Cooperative feeding areas. debate has Much followed Hjort's (CaICOFI) and its predecessor, the California Coopera- has focus the and hypothesis seminal shifted to stage- Sardine Research Program. CalCOFI evolved from tive as and size-specific predation specific growth rates to the for reasons the understand need practical the principal factors determining early survival and year- of the scien- founding the and sardine Pacific demise (Anderson on research however, 1988); strength class complex problem a such would realized that tists of marine fishes continues the early life history stages that a require program comprehensive integrated size the assessing to provide practical means for and and ecology, marine analyses, population investigations the central is and populations spawning of to extent the of dynamics physical California the Current. of fluctuations factors for these of regulating search of discovery the are of this effort results the Among populations. fish to techniques plankton-based and stocks new new measure biomass of important of taxonomic competency development in stocks, the importance The overestimated. be cannot research ichthyoplankton series, of time biological and fund physical and a vast evolved have larvae and eggs fish Marine biology the on information an enormous published of and oceanog- and specializations, morphological forms, of array of California As region. the Current a conse- raphy for characters provide that patterns pigmentation these of a become has CalCOFI advances, quence use Effective them. identifying these for characters of for development model the biological/oceano- other of knowledge requires identification and their of intra- country and interna- graphic fisheries pro grams in this of how interspecific variation and an understanding tionally. 1

14 Geographic Coverage region covered in this guide is defined by The core et Kramer in described are a series in and (1972), aL a pattern survey which extends over the CalCOFI 1 > data reports of 24 that list ichthyoplankton associ- and 2 minion border Oregon-California the from area to km station survey CalCOFI each for conducted data ated southern tip the to offshore and Sur California Baja of al. 1951 from (Ambrose et 1987) to 1984 (Stevens et ca. 400 n.mi. (Figures Introduction 1-3). The sampling interannual, a1. 1990). Summaries of the geographic, coastal three grid Current, California the overlays seasonal and distribution on collected larvae fish all of San Oregonian, (the zoogeographic provinces Diegan, presented are to 1951 from surveys CalCOFI in 1984 zone, coastal and three a upwelling and Panamic), 32 Atlases CalCOFI 31 and that precede this guide oceanic Central, (Subarctic-Transitional, masses water (Moser et al. 1993, 1994a). Since larvae collected in 4). Introduction (Figure Pacific) Tropical Eastern and nets recently fairly and small are generally plankton oblique tows plankton 38,735 made CalCOFI (33,876 in reflects abundance relative their spawned, plankton tows, to 1951 from The 1994. tows) neuston 4,859 the spawning activity Thus, of the parent population. eggs oblique fish million 28.6 ca. yielded tows and in provided the summaries CalCOFI 32 and 31 Atlases million distributed larvae 13.6 fish in approximately guide to provide are used extensively in informa- this families (Moser et 152 aL 1993, 1994a) on and seasonality geography tion of spawning of the species described. Spatial and temporal coverage was most complete during the early years (1951-1960) surveys of the ELH In addition to the specimens obtained from occupied cruises multi-vessel when most survey of the regular CalCOFI survey cruises, other cruises, expedi- at reduced pattern Subsequently, intervals. monthly sources important research programs and tions, were of limitations ship availability and funding resulted in a early life history materia! (Table Introduction 1). Some reduction areal their in and cruises of number the of 7205 and of these cruises CalCOFI NORPAC, (e.g., coverage. were Surveys made the CalCOFI cruises to annually from 1951 to California, of Gulf the 7210, EASTROP during 1995 to 1985 from and 1969 and AC I and II) were special extended cruises triennially been has pattern survey the 1985, Since 1969-1984. the with associated expeditions or CalCOFI program. to quarterly coverage limited the Southern within cruises Survey the off and coasts California Oregon Division Resources Fisheries Coastal the by conducted 1993). California Bight (Moser et a1. were SWFSC a valuable material. of source of the Cruises the Marine Mammal Observation SWFSC of are coastline, the to perpendicular lines, survey The placed 40 40- at are stations principal and apart n.mi. SIO and Program Tuna-Oceanography Program the 1). Introduction (Figure lines the on intervals n.mi. south from waters the specimens many provided of Usually are occupied at 20-n.mi. intervals on stations Likewise pattern. CalCOFI of program sampling the inner the a Panama was pattern of half at closer intervals Laboratory near TTC IA the and the Achotines, at features. these accommodate to islands and source rich coast the speci- of materiaL Important sources for At the were mass water central the from mens cruises each station a double oblique plankton net is made of tow a standard (to to depth SWFSC during Dr. of cruises the Laboratory, Honolulu the 140 and 1951-1968 m to Thomas were m thereafter). Neuston (Manta net) 210 Biology, A. Clarke (Hawaii Institute of Marine tows taken 1995. 1978 from station to each on University expeditions Hawaii), many Details and of to of SIO that region. laboratory and methods sampling CalCOFI procedures 2

15 '00 '00 40· ,O? " . \'l-~ />.0 SAN fRANCISCO' ,,0 POINT ,/>.O ONCEPTION 35" ,'00 • '~ ... <>A • . DIEGO! ..:.. • : • ... ,'b? ( /. ... . . 'l-0O ... " ~,.~ ' • '00 -- - ,. 1° 30" '00 q,0· ,0° ,,0 25" '. 0· 20· ,'0 110· 105· 115· 130· 125" 135" 120° Figure Introduction 1. The CalCOFI survey pattern extending from the California-Oregon border (line 40) to south of Cabo San Lucas, onshore-offshore of lines indicate major survey lines; major station numbers Mexico (line 157). Numbers at the ends Baja Sur, California has survey the 1985, 40. Stations on extended lines 60 through 110 were occupied infrequently during 1951-1984. Since line shown are on 1993). al. et Moser (from 120 station to 77 to 93, with a maximum seaward extent lines Bight, California Southern the to limited been 3

16 • • • '00 • ~O • • • • • 40° • • • • • • • • • • • ~O • • • • • • • • • • • • • • • • • • • • • • • • • • q,O • • • • • • • • • • • • • • • • • • • • 35° • • '00 • • • • • • • • • • • COO • • ,/7 • • • San Miguel Island • • • • • \O~ • • • • • • • j' • • • • • • San Island Nicolas • • • • • • • • • • • • • • • • • • • • • • • • • • • • San Clemente Is. of Figure 2. Part of the northern section Introduction the CalCOFI survey pattern showing geographic features. 4

17 33° • • • • • • • • • • • 30° • • • • • • Guadalupe Isla \ de • • • • • \\() • •• • • • • Cedros-~ Isla • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • Alijos Rocas • • • 25° . / · • • · .. • • • Magdalena Babia • • • • • • . ~(). . • • • Almejas Bahia • • • • \ro() • • • • • • • 1() • • • • • • • • • • • • • • 9J() • • • • • • ell • • • • • • • • • • ",,() • • • • • • • • • • • • , \b<() • • • • • • • • Marias • • Tres Islas • • • • • • • • • • • • • • • • • • • • • • • • \<:'() 20° • • \<:,1 de Islas Revillagigedo Clarion • Figure Introduction 3. Part of the southern section of the CalCOFI survey pattern, including the Gulf of California, showing geographic features. 5

18 SUBARCTIC TRANSITIONAL CENTRAL I. "-,. • Introduction 4. Water masses and zoogeographic provinces Figure the northeast Pacific in relation to the CalCOFI survey area. of Smith of water masses are from Brinton (1962) and Moser et aI. (1987); zoogeographic provinces are modified from Allen and Boundaries is outlined. (1988). The CalCOFI survey area 6

19 Table Introduction expeditions, and research programs that were major sources of early life history specimens 1. Cruises, the this are approximate and include stations occupied enroute to Coordinates survey area. Specimens from guide. in used cruises, programs, listed below are archived in the or Reference Collection (CFRD Ref. ColI.); the expeditions CFRD contains from sources other than those listed in specimens table. The S10 Marine Vertebrates Collection and collection this Natural History Museum of Los Angeles were major sources for specimens used in this guide but are not included in this the table; their SIO and LACM, respectively. acronyms are Number Program, of Cruise, Acronym Locality Years or Expedition Cruises zone MEC Nearshore coastal 1977-1986 Commission California Coastal 126 and Onofre San between at Onofre San study Nuclear Margarita River, Santa conducted Station, Generating California; 33°N, 117°W by MEC Analytical Systems, Inc. California Current region; CalCOFI 1949-present 258 Oceanic California Cooperative 20°-42°N, llOo-123°W Fisheries Investigations CalCOFI west Current 7205, California to 1972 2 Cruise CalCOFI Extended Mass; Water Central 7210 on numbers (station 1972 o N, 121O-145°W 20o-31 extended lines are "degrees latitude. degrees longitude") CalCOFI 5602, Gulf of California; 1956-1959 8 Gulf CalCOFI of California 5612, 5702, 23°-32°N, 107°-115°W 5604, from (distinguished Cruises 5708, 5706, 5704, a regular cruises CalCOFI by 5902 line after in "G" number station designation) 1955 Pacific; NORPAC (August- Northeast CalCOFI Expedition NORPAC 20°-48°N, 111°-154°W September) 5508) (CaICOFI HMS-30 Pacific; 1955 (July- Northeast CalCOFI Expedition NORPAC 157°W~180oW 29°-48°N, August) M. Hugh (R/V Smith; NMFS Laboratory) Honolulu 1987-1992 8803, CFRD 8701, off Eastern Pacific Coastal Fisheries Resources 7 9104, 9001, 8903, and Oregon; California (SWFSC) Division 12F-129°W 34°-46°N; 9107,9203 Worldwide 1928-1930 Dana Danish RlV Dana Expedition 1967 EASTROPAC I Eastern Tropical Pacific; (February- 4 I EASTROPAC 76°-126°W 20oN-20oS, March) 1967 Pacific; Eastern Tropical (August- 3 II EASTROP AC II EASTROPAC o N-15°S, 20 77°-119°W September) coast Clarke-Hawaii Leeward (west) 1970-1972 of 6 Marine of Institute Hawaii Oahu, Hawaii; 21oN, cruises Oahu Biology 158°W by Thomas A. conducted Clarke 1985-present Pacific; Eastern Tropical numerous IATTC Achotines IATTC Laboratory o 7°N,80 W cruises Collections 7

20 Table Introduction Continued. 1. Cruise, Program, of Number Acronym Locality Years Cruises Expedition or IATTC MazatJ{m Eastern Tropical Pacific; 1966-1967 Project Mazatlan IATTC 8 210-23°N, 106°-1100W Project Eastern Tropical Pacific; MOPS 1986-1990 Division Mammal Marine 5 0 31 N-12°S, 78°-154°W (SWFSC) MOPS Cruises Eastern Tropical Pacific; 1992 (July- 92 PODS Mammal Division Marine 2°-27°N, 77°-115°W December) Cruise 1992 PODS (SWFSC) 1993 Coastal PODS 93 Coastal (July- California, Division Marine Mammal Mexico, & Gulf of November) PODS Cruise (SWFSC) 1993 18°-42°N, California; 105°-BloW 1986-1988 zone SK Nearshore coastal 24 Sharon Kramer H. collections between Mission Bay and Onofre, California; San 117°W 32°-33°N, central North Pacific gyre; 1971 (September- IX Aries Aries sm IX 124°-155°W 27°-32°N, October) Pacific (June-July) 1972 gyre; central North I Cato Cato I SIO 25°-3 ION, 133°-156°W South Pacific 1972 (July) and North Cato II Cato II SIO 10N-26°S, gyres; central 155°W North Pacific central gyre; (September) 1968 Climax I I Climax sm 155°-158°W 27°-29°N, gyre; (August- North Pacific central 1968 Climax II II Climax SIO 0 28°-29°N, 151 -155°W September) (October- 1955 Pacific; Tropical Eastern Eastropic 2 SIO Eastropic 0 30 December) N-7°S, 78°-122°W 1985 Front; (July) Ensenada 85 FRONTS 85 FRONTS SIO 0 -33°N, 119°-122°W 31 (August- 1951 Pacific; Northeast Holiday Northern Holiday Northern sm 0 28°-56°N, W September) 127°-160 Eastern Tropical Pacific; 1956 (November- Scope SIO Scope Expedition 4°-27°N, 79°-114°W December) Pacific; 1958 (April-June) Eastern Tropical TO 58-1 SIO Expedition SCOT 78°-119°W ON, 4°-28 58-1) (TO (May- 1952 Pacific; Tropical Eastern Shellback Expedition Shellback S10 28°-15°N, 78°-125°W August) 8

21 Table Introduction Continued. 1. Cruise, Program, Number of Locality Expedition or Acronym Years Cruises central Pacific gyre; North Southtow XIII XIII 1973 (J anuary- SID Expedition Southtow 28°-32°N, l24°-155°W February) gyre; central Pacific North Tasaday I Tasaday 1973 (June) SIO I Expedition o N, 155°W 28°-30 1973 gyre; central Pacific (July) II Tasaday North SIO II Expedition Tasaday 25°-28°N, 155°W gyre central Pacific XI Tasaday North 1974 (March) Expedition Tasaday XI SIO Eastern Tropical Pacific; 58-2 TO SIO Tuna-Oceanography (November- 1958 Gulf primarily of December) 58-2 Cruise Tehauntepec; 12°-26°N, 91°-114°W 1959 TO 59-1 Eastern Tropical Pacific; (January- Tuna-Oceanography SIO primarily Gulf of February) Cruise 59-1 8°-29°N, Tehauntepec; 87°-116°W Pacific; Eastern Tropical 1959 (August- TO 59-2 SID Tuna-Oceanography primarily Gulf of September) Cruise 59-2 14°-27°W, Tehauntepec; 94°-117°W Seamount, TC 8405, SE Hancock (July) 1984 SWFSC, Honolulu Laboratory, 3 TC 8501, Northern Ridge, 29 1985 (February) Hawaii Townsend Cromwell RN o °N-30oN, 179°-180 WI (August) TC 8605 1986 Cruises Seamount Hancock Johnston Atoll, 16°-17°N, TC 8406 1984 (November) Honolulu Laboratory, SWFSC, 169°-170OW RN Townsend Cromwell Johnston Atoll Cruise (September- 1985 of coasts East and west 8504 TC Laboratory, Honolulu SWFSC, 4 and December) Oahu, Hawaii; 22°N, 8505 TC Oahu RN Townsend Cromwell 158°W and 21°N, 1986 (April-June) 158°W TC 8602 Ichthyoplankton Vertical TC 8604 Cruises Distribution 9

22 Taxonomic Coverage The guide includes full descriptions of 467 species from taxa and area survey CalCOFI the within lie aries orders. 25 and families, 141 genera, 340 contained in provinces water masses and coastal zoogeographic that descriptions representative illustrations of and Partial of the from region CalCOFI the core the on impinge introductions the in species included are orders to 119 fluctu- south, and west. Faunal boundaries may north, to genera and families; these add 17 families and 78 to interannual, seasonal, dynamic the response in ate represented with the in the guide. Of the species total shifts and climatic that characterize the California New science. to new are (27%) 127 descriptions, full Current is reflected in the faunal system and this illustrations are undescribed previously of stages CalCOFI of the the composition various subregions in new information and (62%) species 364 for presented 1987, et Moser 1983; (Loeb area survey 1993, al. the guide. for all nearly 586 species in is included we of this, 1994a; Moser and Smith 1993). Because guide With this in included species the exceptions, few attempted to have than in exclusive rather inclusive be planktonic history those are or pelagic early life with described. selecting the species In the southern part of found that may be stages in coastal, oceanic, or estua- or the were included if their ELH pattern, stages taxa rine waters the regions. of CalCOFI area and adjacent coast the from of been have adults recorded Pacific stages Early freshwater of and a are species anadromous If area. Lucas San Cabo the or Sur California Baja nor not are the viviparous surf perches described, the from specimens criteria, these of met either taxon whose young new-born advanced (Embiotocidae) Gulf were or of more southern localities California that used to augment the for basis the formed material surf the in remain includ- fishes Most zone. subtidal or the criteria description. Analogous applied were spawn guide the in ed progress that eggs free-floating in a distinct planktonic larval stage before through in the western of regions northern and taxa of selection transforming into a pelagic or demersal juvenile. Many CalCOFI the These some- modified were criteria area. included shorefish groups in this guide (e.g., cottids, in northern part of what the the CalCOFI region labrisomids, gobiids) produce pomacentrids, blenniids, of the of because a comprehensive identifi- availability planktonic larvae from demersal eggs. When known, stages cation guide to ELH of northeast the of fishes the demersal eggs The species are groups these of some described. in al. 1989). Taxa et Pacific (Matarese rockfishes Sebastes some gadids, (e.g., to birth give but viviparous are ELH whose species) (Sebastes) or planktonic larvae that are at a stage treated rare are development stages adult in the CalCOFI area and of not were (1989), aL described in funy in Matarese et first-feeding the to equivalent larvae of typical ovipa- included guide this was them on information although are many rous species with planktonic eggs. There and tables text in in meristic the of chapter introduc- (ca. species rockfish 60) in CalCOFI region and the tions. an of the important component their larvae are fishes are included Also ichthyoplankton. CalCOFI whose species the contains list following The early such planktonic have which alepocephalids, as eggs life that and guide this in described are stages history larvae and are juveniles) relatively pelagic (or deep the where found. page number the description may be column water the stan- in in collected rarely are and Eschmeyer Classification and nomenclature follow plankton dard CalCOFI tows. (1990; pers. comm., December 1995 - 1996); February family. each within alphabetically listed are species bound- The guide includes taxa whose distributional 10

23 List of Taxa Elopomorpha Elopiformes Elopidae Elops 1909 ... 76 affinis Regan Albuliformes Albulidae Albula sp. ... 80 Notacanthiformes N otacanthidae ... 84 Leptocephalus giganteus 1959 Castle Anguilliformes Muraenoidei Muraenidae Gymnothorax mordax (Ayres 1859) 90 ... Congroidei Ophichthidae Jordan and Myrophis 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 vafer Gilbert triserialis (Kaup 1856) Ophichthus ... 96 zophochir Ophichthus Gilbert 1882) (Jordan and 98 ... Congridae gilberti (Ogilby 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 Ariosoma 104 ... 1891) macrurus Bathycongrus (Gilbert obtusus (Garman 1899) ... 106 Chiloconger cinctus 1899) Gnathophis (Garman 108 ... Heteroconger canabus and Rosenblatt 1974) ... 110 (Cowan Heteroconger digueti (pellegrin 1923) ... 112 ... 114 Paraconger californiensis Kanazawa 1961 (Jordan Bollman 1889) nitens and Rhynchoconger ... 116 Derichthyidae Gill Derichthys serpentinus 1884 120 ... Nemichthyidae Avocettina bowersii (Garman 1899) 124 ... ... 126 (GUnther 1878) infans Avocettina scolopaceus 1848 Nemichthys Richardson ... 128 Serrivomeridae sector Garman 1899 Serrivomer ... 132 Nettastomatidae ... 136 (Garman 1899) Facciolella gilbertii ... 138 Hoplunnis sicarius 1899) (Garman sedis Incertae 142 ... Castle Raju 1975 Thalassenchelys coheni and Saccopharyngiformes Cyematidae Cyema atrum GUnther 1878 ... 146 Leptocephalus Schmidt 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148 holti Saccopharyngidae Sacco pharynx lavenbergi Nielsen and Bertelsen 1985 ... 152 Eurypharyngidae 11

24 Eurypharynx pelecanoides 1882 ... 156 Vaillant Monognathidae Monognathus 1974 ... 144 ahlstromi Raju Bertelsen and 1987 ... 144 Monognathus rosenblatti Nielsen Clupeiformes Clupeidae Clupea Valenciennes 1847 ... 162 pallasii (DeKay 1842) ... 164 Etrumeus teres (Jordan thrissina Gilbert 1882) ... 166 Harengula and (GUnther ... libertate 168 Opisthonema 1867) sagax 1842) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170 Sardinops (Jenyns Engraulidae compressa (Girard 1858) ... 176 Anchoa delicatissima Anchoa 1854) ... 178 (Girard mysticetus (GUnther ... 180 Cetengraulis 1867) Girard ... 182 mordax Engraulis 1854 Salmoniformes Argentinoidei Argentinidae Argentina Giibert 1890 ... 186 sialis Bathylagidae bericoides (Borodin Bathylagus ... 190 1929) Bathylagus Maul 1948 ... 192 longirostris milleri Gilbert and Bathylagus 1898 ... 194 Jordan nigrigenys Parr 1931 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196 Bathylagus Bathylagus ochotensis Schmidt 1938 ... 198 Bathylagus pacificus 1890 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200 Gilbert wesethi Bolin ... 202 Bathylagus 1938 Rass 204 ... schmidti Leuroglossus 1955 stilbius 1890 ... 206 Leuroglossus Gilbert Microstomatidae sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210 Microstoma candida Cohen 1958 ... 212 Nansenia crassa 1965 ... 214 Nansenia Lavenberg Opisthoproctidae exilis Cohen Bathylychnops ... 218 1958 Dolichopteryx longipes (Vaillant 1888) ... 220 Macropinna microstoma Chapman 1939 ... 222 Alepocephalidae Alepocephalus Gilbert 1892 ... 226 tenebrosus burragei 1925 and Nichols Bajacalifornia ... 228 Townsend Goode and Bean 1896 ... 230 Bathylaco nigricans Talismania bifurcata (parr 1951) ... 232 P latytroctidae Holtbyrnia latifrons Sazonov 1976 ... 236 Sagamichthys abei Parr 1953 ... 238 Salmonoidei Osmeridae Hypomesus pretiosus (Girard 1854) ... 242 Stomiiformes 12

25 Gonostomatidae Cyclothone Garman 1899 ... 0 •••••••••••••••••••••••••••••• 250 acclinidens atraria Gilbert ... 252 Cyclothone 1905 Mukhacheva Cyclothone 254 1964 pseudopal/ida ... Garman ... 256 signata Cyclothone 1899 Parr 1931 ... 0 • • • • • • • • • • • Diplophos • • • • • • • • • • • • • • • • • • 258 proximus • taenia Gunther 1873 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 260 Diplophos atlanticum Norman 1930 Gonostoma 262 ... Gonostoma Grey 1960 ... 0 •••••••••• 264 ebelingi elongatum Gunther ... 266 Gonostoma 1878 Stemoptychidae affinis 1899 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argyropelecus 270 Garman hemigymnus 1829 ... 0 •••••••••••• 272 Argyropelecus Cocco lychnus Garman 1899 ... 274 Argyropelecus Argyropelecus sladeni 1908 ... 0 •••••••• 276 Regan oculatus 1899) ... 278 Danaphos (Garman spp. ... 280 Sternoptyx tripunctulatus (Esmark 1871) ... 0 •••••••••••••••••••••••••• 282 Valenciennellus Phosichthyidae Ichthyococcus irregularis Rechnitzer and Bohlke 1958 ... 286 Vinciguerria lucetia 1899) ... 288 (Garman poweriae (Cocco ... 290 Vinciguerria 1838) (Beebe ... 292 nonsuchae 1932) Woodsia Stomioidea ChauHodontidae Bean 1890 Chauliodus 298 macouni ... Stomiidae Stomias atriventer Garman 1899 ... 302 Astronesthidae sp. . ... 305 Astronesthes panarnensis and Trewavas 1929 ... 305 Borostomias Regan capensis (Gilchrist and von Bonde Neonesthes ... 306 1924) Melanostomiidae Bathophilus brevis Regan and Trewavas 1930 ... 309 Bathophilus filifer (Garman 1899) ... 314 Bathophilusjlemingi Aron McCrery 1958 ... 0 •••• 316 and sp. ... 0 •••••••••••••••••••••••••••••••••••••••••••• 309 Eustomias . boureei (Zugmayer 1913) ... Flagellostomias •••••••••••••••••••••••••••••• 309 0 Leptostomias sp. ... 311 Melanostomias sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310 Opostomias mitsuii Imai 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310 Photonectes sp 310 ... macro pus Tactostoma 1939 ... 318 Bolin Malacosteidae Aristostomias scintillans (Gilbert 1915) ... 322 Idiacanthidae ... antrostomus Gilbert 1890 ldiacanthus 326 Aulopiformes Aulopoidei Aulopidae 13

26 Aulopus bajacali and Kotlyar 1984 ... 330 Parin Scopelarchidae Benthalbella 1939) ... 334 dentata (Chapman (Rofen 1966) 336 Rosenblattichthys volucris ... 1929) 338 (parr ... nicholsi Scopelarchoides Scopelarchus ... 340 analis (Brauer 1902) Alcock 1896 ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 342 Scopelarchus guentheri Notosudidae brevis Kreffi: and Marsha111976 ... 344 Ahliesaurus Bertelsen, (Mead ... harryi 346 Scopelosaurus 1953) Alepisauroidei Synodontidae Synodus (Ayres 1855) ... 350 lucioceps Paralepididae risso 1840) Arctozenus ... 354 (Bonaparte neles (Harry ... 356 Lestidiops 1953) (parr ... 358 pacificum 1931) Lestidiops (Jordan Lestidiops Gilbert 1880) ... 360 ringens and atlantica (Kr0)'er 1868) ... 362 Magnisudis macrura (Ege 1933) ... 364 Stemonosudis atrox 1963 ... 366 Sudis Rofen Anotopteridae pharao Zugmayer 1911 ... Anotopterus 370 Evermannellidae Evermannella ahlstromi Johnson and Glodek 1975 ... 374 Evermannella indica Brauer 1906 ... 376 Alepisauridae Alepisaurusferox 1833 ... 380 Lowe Myctophiformes Neoscopelidae Alcock 1890 ... 384 Scopelengys tristis Myctophidae Lampanyctinae longipes (Brauer 1906) ... 394 Bolinichthys townsendi (Eigenmann and Eigenmann 1889) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396 Ceratoscopelus pacificus Parr 1931 ... 398 Diaphus theta and Eigenmann 1890 ... 400 Diaphus Eigenmann luminosa (Garman Lampadena ... 392 1899) Lampadena urophaos Paxton 1963 ... 402 Lampanyctus acanthurus Wisner 1974 ... 404 Lampanyctus nobilus 1928 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 406 Taning parvicauda 1931 ... 408 Lampanyctus Parr steinbecki Bolin 1939 Lampanyctus 410 ... Lampanyctus tenuiformis (Brauer 1906) ... 412 Lobianchia gemellarii (Cocco 1838) ... 414 Nannobrachium (Parr 1931) ... 416 idostigma Nannobrachium "niger" ... 418 Nannobrachium "no pectorals" ... 420 Nannobrachium (Gilbert 1892) ... 422 regalis Nannobrachium ritteri (Gilbert 1915) ... 424 14

27 Notolychnus valdiviae 1904) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 426 (Brauer 0 428 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 resplendens (Richardson Notoscopelus 1845) Hubbs and 1964 ingens Parvilux Wisner 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 430 0 0 0 0 and Eigenmann 1890) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Stenobrachius 0 0 0 0 0 0 0 0 0 0 432 leucopsarus (Eigenmann 0 minimus Taaningichthys 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1928) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 434 (Taning 0 0 0 1890) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 436 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (Gilbert Triphoturus mexicanus (Brauer Triphoturus nigrescens 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 438 1904) 0 0 Myctophinae (Gilbert and Cramer 1897) Benthosemafibulatum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 392 0 0 panamense Benthosema 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (Taning 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 440 1932) 0 suborbitale (Gilbert 1913) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 442 Benthosema nigroocellatus (Gunther Centrobranchus 1873) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 444 0 0 atlanticus Diogenichthys 1928) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 446 (Taning laternatus Diogenichthys (Garman 1899) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 448 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 450 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1829) risso Electrona (Cocco 0 (Garman 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 tenuiculus 0 1899) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 452 Gonichthys 0 454 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1899) (Garman atratum Hygophum Becker 1965 Hygophum proximum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0000000000000 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 456 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 458 0 1892) Hygophum reinhardtii (Lutken rara 1892) Loweina (Lutken 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 460 0 asperum 0 1845) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Myctophum 0 0 0 0 393 (Richardson aurolaternatum 1899 Myctophum Garman 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 462 0 0 brachygnathum 1856) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (Bleeker 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 393 Myctophum Myctophum lychnobium Bolin 1946 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 392 0 nitidulum Garman Myctophum 1899 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 464 0 0 0 obtusirostrum Myctophum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Taning 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 393 1928 0 0 Taning 1928 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Myctophum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 393 selenops 0 spinosum 1867) Myctophum (Steindachner 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 392 0 0 0 (Bolin 1939) crockeri Protomyctophum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 466 0 0 thompsoni (Chapman 1944) Protomyctophum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 468 0 Symbolophorus (Eigenmann and Eigenmann 1889) californiensis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 470 0 0 (Gilbert 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 evermanni 0 0 0 0 0 0 0 0 0 0 0 0 472 1905) Symbolophorus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1880) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 474 and (Jordan Gilbert crenularis Tarletonbeania Gadiforrnes Bregrnacerotidae 0 0 0 0 1890 0 0 0 0 0 0 0 0 0 0 0 0 0 0 478 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Bollman Jordan bathymaster Bregmaceros and Bregmaceros 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 spo 0 0 0 0 0 480 0 Macrouridae . 0 0 486 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (Gilbert Albatrossia 1892) pectoralis 488 0 0 1890) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (Gilbert scaphopsis Caelorinchus 0 acrolepis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1884) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 490 Coryphaenoides (Bean filifer (Gilbert 1896) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Coryphaenoides 492 494 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Giinther 1877 Coryphaenoides leptolepis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 496 and Iwamoto Hubbs 1977 Mesobius berryi Nezumia sppo 0 0 0 0 0 0 0 0 0 0 0 0 • 0 0 •• 0 • 0 0 ••• 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 498 Moridae Microlepidium (Jordan and Cramer 1897) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 verecundum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 502 0 Physiculus nematopus Gilbert 1890 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 504 0 Physiculus Gilbert 1890 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 rastrelliger 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 506 Merlucciidae 15

28 Merluccius productus 1855) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 510 (Ayres Ophidiiformes Ophidiidae Hubbs 1944 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 516 Brotula clarkae 518 1890) ... (Gilbert emmelas Cherub/emma (Girard taylori Chilara 1958) ... 520 pinna Hildebrand and Barton 1949 Lepophidium 522 negro ... 1890 ... 524 stigmatistium Lepophidium Gilbert ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . 526 scrippsae 1916) Ophidion (Hubbs sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 528 Ophidion ... 530 grandis (GUnther 1877) Spectrunculus Carapidae Echiodon exsilium Rosenblatt 534 1961 ... ... (Putnam 1874) 536 ius Encheliophis dub . Bythitidae ... 540 1854) Brosmophycis marginata (Ayres rubrirostris Cataetyx 1890 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 542 Gilbert diagrammus and Snodgrass Oligopus (Heller ... 544 1903) Batrachoidiformes Batrachoididae Porichthys notatus Girard 1854 ... 548 Lophiiformes Lophioidei Lophiidae Lophiodes 1899) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 554 caulinaris (Garman ... 556 spilurus 1899) Lophiodes (Garman Antennarioidei Antennariidae 560 ... 1907 and Jordan is avalon Antennarius Starks Chaunacioidei Chaunacidae Chaunax . sp. ... 552 Ogcocephalioidei Ogcocephalidae elater (Jordan and Zalieutes 1882) Gilbert ... 564 Ceratioidei Caulophrynidae Caulophryne spp.. ... 568 Melanocetidae Melanocetus johnsoni 1864 572 ... GUnther Oneirodidae 577 ... 1932 draco Chaenophryne Beebe 577 ... 1925 Regan longiceps Chaenophryne ... 578 Dolopichthys spp. Microlophichthys microlophus (Regan 1925) ... 577 ... 580 Oneirodes spp. . Phyllorhinichthys micractis Pietsch 1969 ... ; ... 577 Thaumatichthyidae Thaumatichthys sp ... 552 Centrophrynidae 16

29 Centrophryne spinulosa and Trewavas 1932 ... 552 Regan Ceratiidae Ceratias 1845 ... 584 holboelli Kreyer Gill 1883 586 Cryptopsaras couesii ...•... Gigantactinidae ... spp. 590 Gigantactis Linophrynidae Borophryne ... 594 apogon Regan 1925 "Hyaloceratias" larvae ... 593 Gobiesociformes Gobiesocidae Briggs ... 600 eugrammus 1955 Gobiesox (Girard 1858) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gobiesox maeandricus 599 rhessodon Smith 1881 ... 602 Gobiesox eigenmanni (Gilbert 1890) Rimicola 604 ... Rimicola ... 606 spp. Atheriniformes Atherinidae Jordan and Gilbert 1881 ... 610 Atherinella eriarcha (Myers and Wade 1942) ... 612 Atherinella nepenthe affinis (Ayres 1860) ... 614 Atherinops californiensis Atherinopsis 1854 ... 616 Girard tenuis 1860) ... 618 Leuresthes (Ayres Cyprinodontiformes Cyprinodontidae Fundulus Girard 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 622 parvipinnis Beloniformes Scomberesocidae Cololabis saira (Brevoort 1856) ... 628 Belonidae Ablennes 01 alenciennes 1846) ... 631 hians exilis (Girard ... 632 Strongylura 1854) Hemiramphidae viridus Euleptorhamphus Hasselt 1823) ... 635 (van saltator Gilbert and Starks 1904 ... 636 Hemiramphus Hyporhamphus rosae and Gilbert 1880) ... 638 (Jordan micropterus 1847) 01alenciennes Oxyporhamphus ... 640 micropterus Exocoetidae heterurus hubbsi Parin 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 646 Cheilopogon Cheilopogon pinnatibarbatus californicus (Cooper 1863) ... 648 Exocoetus monocirrhus 1846 ... 645 Richardson volitans 1758 ... 650 Exocoetus Linnaeus acutus rostratus (Gunther 1866) Fodiator 652 ... Hirundichthys marginatus (Nichols and Breder 1928) ... 654 Hirundichthys rondeletii 01 alenciennes 1846) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 654 Hirundichthys 01 alenciennes 1846) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 654 speculiger Hirundichthys spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 654 Prognichthys tringa Breder 1928 ... 656 Lampridiformes Trachipteroidei 17

30 Lophotidae 662 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1809 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Gioma Lophotus pede lace Radiicepha1idae Radiicephalus 1917 elongatus Osorio 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0·0 0 0 0 0 0 0 0 0 0 666 0 0 0 Trachipteridae 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 670 0 1977 Rosenblatt Butler lorum Desmodema and 0 1858 0 Kner 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 672 altivelis Trachipterus 0 0 0 0 0 0 0 0 0 0 0 0 0 674 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1964 Trachipterusjukuzakii Fitch (Bonelli 1819) Zu cristatus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 676 0 0 0 Beryciformes Berycoidei Diretmidae Johnson argenteus Diretmus 1864 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 680 0 0 Anop1ogastridae Anoplogaster corn uta (Valenciennes 1833) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 684 0 Holocentridae 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 688 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Myripristis Valenciennes 1855 leiognathos suborbitalis (Gill Sargocentron 1864) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 690 0 0 0 Stephanoberycoidei Melamphaidae 0 1891 0 0 696 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Gilbert lugubris Melamphaes Melamphaes parvus Ebeling 1962 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 698 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 700 1962 Melamphaes simus Ebeling 0 spo 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Melamphaes 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 702 0 Poromitra (GUnther 1878) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 crassiceps 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 704 0 Poromitra megalops (Liitken 1877) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 706 Poromitra oscitans 1975 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 695 Ebeling Poromitra spo 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 695 0 0 708 (GUnther 1887) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 •• 0 0 0 0 0 0 0 0.0 0 0 0 0 0 robustus Scopeloberyx (Gilbert 1915) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 710 Scopelogadus bispinosus Mirapinnidae Eutaeniophorusfestivus and Marshall 1956) (Bertelsen 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 714 0 Syngnathiformes Aulostornoidei F istu1ariidae 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 720 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Riippell1838 commersonii Fistularia Fistularia corneta Gilbert and Starks 1904 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 722 Centriscidae 726 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1839) Macroramphosus (Lowe gracilis Syngnathoidei Syngnathidae arctus (Jenkins and Evermann 1889) Cosmocampus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 731 0 0 excisus Kaup 1856 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Doryrhamphus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 731 excisus Hippocampus ingens Girard 1858 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 730 0 Syngnathus 1845 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Storer 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 731 californiensis Syngnathus leptorhynchus Girard 1854 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 731 Scorpaeniformes Scorpaenoidei Scorpaenidae Sebastinae 18

31 Sebastes atrovirens and Gilbert 1880) ... 742 (Jordan auriculatus Girard ... 744 Sebastes 1854 (Gilbert 1890) 746 Sebastes aurora ... and 748 1880) ... (Jordan Sebastes carnatus Gilbert 1844 ... 750 caurinus Sebastes Richardson (Jordan and Gilbert 1880) ... 741 Sebastes chlorostictus constellatus (Jordan Gilbert 1880) ... 752 Sebastes and (Eigenmann Beeson 1894) ... 754 dallii and Sebastes (Gilbert 1890) Sebastes 756 diploproa ... ensifer Chen 1971 ... 741 Sebastes eos (Eigenmann and Eigenmann 1890) ... 741 Sebastes Sebastes goodei and Eigenmann 1890) ... 758 (Eigenmann jordani (Gilbert ... 760 Sebastes 1896) (Eigenmann Eigenmann 1889) ... 762 levis and Sebastes (Eigenmann and Beeson Sebastes ... 764 macdonaldi 1893) melanops Girard 1856 ... 766 Sebastes melanostomus (Eigenmann and Sebastes 1890) ... 768 Eigenmann Sebastes (Jordan and Gilbert 1881) ... 770 mystinus ovalis 772 1863) ... Sebastes (Ayres paucispinis ... 1954 Sebastes 774 Ayres Sebastes (Gill 1864) ... 776 pinniger Sebastes rastrelliger (Jordan and Gilbert 1880) ... 778 Sebastes rosaceus 1854 ... 741 Girard rosenblatti Chen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 741 Sebastes 1971 (Eigenmann 780 Eigenmann 1890) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rufus Sebastes and serrano (Eigenmann and Eigenmann 1890) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 782 Sebastes ides umbrosus (Jordan and Gilbert 1882) ... 741 Sebastes Scorpaeninae Pontinus (Gilbert 1890) ... 784 sierra sp. 786 Pontinus ... Scorpaena guttata Girard 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 788 xyris (Jordan Scorpaenodes Gilbert 1882) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 790 and Sebastolobinae Sebastolobus alascanus Bean 1890 ... 792 Sebastolobus altivelis Gilbert 1896 ... 794 Triglidae Bellator (Jordan 1897) ... 800 loxias ruscarius and Starks 1904 ... 802 Prionotus Gilbert stephanophrys Lockington 1881 Prionotus 804 ... Anoplopomatoidei Anoplopomatidae Anoplopomajimbria (Pallas 1814) ... 808 Hexagrammoidei Hexagrammidae Hexagrammos decagrammus (Pallas 1810) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 813 Hexagrammos lagocephalus 1810) ... 813 (Pallas Ophiodon Girard 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 813 elongatus Oxylebius pictus Gill 1862 ... 814 Pleurogrammus (Panas 1810) ... 813 monopterygius Zaniolepis frenata Eigenmann and Eigenmann 1889 ... 816 19

32 r Zaniolepis Girard 1858 ... 818 latipinnis Cottoidei Cottidae (Hubbs 1926) ... 828 Artedius corallinus (Hubbs 1926) 834 Artedius creaseri ... and 828 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jordan Artedius Jenestralis Gilbert 1896) ... 828 harringtoni Artedius (Starks (Girard 1854) ... 830 Artedius lateralis (Jordan and Starks 1895) ... 834 Artedius meanyi Girard ... 828 notospilotus 1856 Artedius Jordan and Gilbert 1880 Ascelichthys 834 rhodorus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cirrhosus (Pallas 1814) ... 826 Blepsias pugetensis (Steindachner 1876) ... 836 Chitonotus Clinocottus acuticeps 1896) ... 830 (Gilbert analis (Girard ... 830 Clinocottus 1858) (Jordan Starks 1895) ... 830 embryum and Clinocottus (Girard 1858) Clinocottus 832 globiceps ... recalvus (Greeley 1899) ... 832 Clinocottus asper Richardson 1836 ... 826 Cottus bison Enophrys 1854) ... 836 (Girard hemilepidotus 1811) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 824 Hemilepidotus (Tilesius spinosus (Ayres 1854) ... 824 Hemilepidotus Icelinus quadriseriatis 1880) ... 836 (Lockington Leptocottus Girard 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 826 armatus Nautichthys oculoJasciatus (Girard 1858) ... 826 Oligocottus maculosus 1856 ... 832 Girard snyderi Greeley ... 832 Oligocottus 1898 Starks Mann 1911 ... 836 triacus and Orthonopias Eigenmann and Eigenmann Paricelinus ... 838 hopliticus 1889 asprellus Gilbert 1890 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 838 Radulinus boleoides Gilbert 1898 Radulinus 838 ... Rhamphocottus Gilnther 1874 ... 824 richardsoni marmoratus 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 824 Scorpaenichthys Girard gilli Bean 1890 ... 838 Synchirus Psychrolutidae Psychrolutes phrictus Stein and Bond 1978 ... 842 Agonidae Agonomalus mozinoi Wilimovsky and Wilson 1979 ... 847 Agonopsis sterletus 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 848 (Gilbert pentacanthus 1980) ... 850 Bathyagonus (Gilbert swanii (Steindachner 1876) ... 852 Bothragonus Occella verrucosa 1880) ... 847 (Lockington Odontopyxis Lockington 1880 ... 854 trispinosa Stellerina xyosterna (Jordan and Gilbert 1880) ... 847 Xeneretmus latifrons 1890) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 856 (Gilbert Xeneretmus Gilbert 1915 ... 858 leiops Cyclopteridae Careproctus melanurus Gilbert 1892 ... 864 Liparus Gilbert 1896 ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 866 fucensis Liparis mucosus Ayres 1855 ... 868 20

33 Liparis puehellus 1855 ... 870 Ayres pelagieus Gilbert Burke 1912 ... 863 Neetoliparis and Gilbert Paraliparis 863 1892 eephalus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 863 ... 1912 Burke Rhinoliparis attenuatus 1896 Rhinoliparus 863 barbulifer Gilbert . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Percifonnes Percoidei Polyprionidae Ayres 1859 Stereolepis gigas ... ... " 874 Serranidae Serraninae Dipleetrum sp. ... 882 ... 884 1854) clathratus Paralabrax (Girard ... 886 Paralabrax maeulatofasciatus (Steindachner 1868) nebulifer (Girard 1854) Paralabrax 888 ... sp. Serranus ... 890 Anthiinae signifer 1899) Hemanthias (Garman ... 892 Gill 1863 Pronotogrammus multifasciatus ... 894 Epinephelinae ... 881 Epinephelus analogus Gill 1863 eolonus (Valenciennes 1846) Paranthias 896 ... Grammistinae thaumasium (Gilbert Pseudogramma 898 1890) ... ... 881 Ryptieus nigripinnis 1861 Gill Priacanthidae ... 901 (Cuvier Cookeolusjaponieus 1829) Heteropriaeanthus (Lacepede 1801) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 901 eruentatus Pristigenys serrula (Gilbert 1891) ... 902 Apogonidae Apogon Jordan and McGregor 1898 atrieaudus 906 ... (Osburn and Nichols 1916) ... 908 Apogon guadalupensis (Gill 1863) ... 910 Apogon retrosella Echeneidae Eeheneis Linnaeus 1758 nauerates ... 913 Carangidae ... 920 eiliaris 1787) Aleetis (Bloch eaballus 1868 ... 922 GUnther Caranx ... 924 Quoy and 1825 sexfaseiatus Caranx Gaimard orqueta Chloroseombrus and Gilbert 1883 Jordan ... 926 Deeapterus spp. ... 928 (Quoy Gaimard 1825) bipinnulata and Elagatis ... 930 Gnathanodon 1775) ... 932 speciosus (Forsskal ... 934 1758) Nauerates duetor (Linnaeus ... 936 saurus inornatus Gill 1863 Oligoplites ... 938 (Bloch 1793) Selar erumenophthalmus ... 940 Selene 1863) (Gill brevoortii peruviana (Guichenot 1866) ... 942 Selene ... 944 Valenciennes 1833 Seriola lalandi ... 946 Traehinotus kennedyi Steindachner 1875 21

34 Trachinotus paitensis 1832 ... 948 Cuvier rhodopus Gill ... 950 Trachinotus 1863 (Ayres 1855) 952 Trachurus symmetricus ... N ematistiidae Gill pectoralis ... 956 Nematistius 1862 Coryphaenidae Linnaeus 1758 ... 960 Coryphaena equiselis hippurus Linneaus '" ... 962 Caryphaena 1758 Bramidae dussumieri 1831 Brama 966 ... Cuvier Hilgendorf 1878 ... 968 Brama japonica aesticola (Jordan and Snyder 1901) ... 970 Pteraclis Caristiidae Caristius Maul 1949 ... 974 maderensis Lutjanidae Hoplopagrus 1862 ... 980 guentherii Gill (Peters 982 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . argentiventris Lutjanus 1869) guttatus 1869) .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 984 Lutjanus (Steindachner novemfasciatus Gill 1862 ... Lutjanus 986 Lutjanus (Nichols and Murphy 1922) ... 988 peru Malacanthidae Caulalatilus (Jenyns 1842) ... 992 princeps Lobotidae surinamensis (Bloch 1790) ... 996 Lobotes Gerreidae Eucinostomus sp(p). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1000 Haemulidae Anisotremus (Steindachner 1875) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1004 davidsonii serrifer Jordan Gilbert 1882 ... 1006 Conodon and reddingi and Richardson 1895 ... 1008 Orthopristis Jordan californiensis (Steindachner 1875) ... 10lO Xenistius Sparidae Calamus (Lockington 1880) ... 1014 brachysomus Sciaenidae Atractoscion (Ayres 1860) ... 1020 nobilis saturnum (Girard Cheilotrema ... 1022 1858) Genyonemus lineatus (Ayres 1855) ... 1024 Menticirrhus undulatus (Girard 1854) ... 1026 Roncador stearnsii 1875) ... 1028 (Steindachner poUtus ... 1860 ... " Seriphus 1030 Ayres Umbrina Jordan and Gilbert 1882 ... 1032 roncador Mullidae Mulloidichthys dentatus (Gill 1863) or Pseudupeneus (Gin 1863) ... 1036 grandisquamis Kyphosidae GireUinae Girella nigricans (Ayers 1860) ... 1040 Kyphosinae Evermann azurea Jenkins and Hermosilla 1889 ... 1042 Scorpidinae 22

35 Medialuna californiensis 1875) ... 1044 (Steindachner Ephippidae Chaetodipterus 1858) ... 1048 zonatus (Girard Chaetodontidae humeralis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1052 Gunther Chaetodon 1860 Pomacentridae 1862) ... 1056 Abudefduftroschelii (Gill (Cooper 1863) ... 1058 Chromis punctipinnis (Girard rubicundus ... 1060 Hypsypops 1854) (Gill 1062 ... rectifraenum Stegastes 1862) Cirrhitidae Cirrhitichthys (Bleeker 1855) ... 1066 oxycephalus Opistognathidae sp. ... 1070 Opistognathus Howellidae Howella and Snodgrass 1903) ... 1074 pammelas (HeUer . 1076 sp. Howella ... Mugiloidei Mugilidae Mugil Linnaeus 1758 ... 1080 cephalus Polynemoidei Polynemidae Polydactylus (Lay and Bennett 1839) ... 1084 approximans opercu/aris 1863) Polydactylus ... 1086 (Gill Labroidei Labridae dispilus (Gunther 1864) ... 1091 Halichoeres Halichoeres semicinctus (Ayres 1859) ... 1092 Oxyjulis californica 1861) ... 1094 (Gunther pulcher (Ayres ... 1096 Semicossyphus 1854) ... .. sp(p). 1098 Thalassoma . . . . . . . . . . . . . . . . . 1962 ... 1100 Xyrichtys mundiceps Gill pavo (Valenciennes Xyrichtys ... 1102 1840) Scaridae denticulata 1106 and Radcliffe 1917) ... Nicholsina (Evermann sp. . ... 1108 Scarus Zoarcoidei Bathymasteridae Rathbunella alleni Gilbert 1904. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1114 Ronquilus jordani 1889) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1113 (Gilbert Zoarcidae Bothrocara ... 1111 sp. Stichaeidae Plectobranchus evides 1890 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1118 Gilbert Cryptacanthodidae Cryptacanthoides aleutensis (Gilbert 1896) ... 1111 Pholidae sanctaerosae Gilbert and Starks 1897 ... 1122 Ulvicola Xererpesfucorum (Jordan and Gilbert 1880) ... 1124 Anarhichadidae 23

36 Anarrhichthys ocellatus 1855 . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1128 Ayres Zaproridae Zaprora 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1111 silenus Jordan Trachinoidei Chiasmodontidae 1864 ... 1134 niger Chiasmodon Johnson 1909 ... 1133 indica Kali Lloyd (Parr 1931) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 113 6 Kali normani ... Pseudoscopelus sp. 1133 Ammodytidae hexapterus 1814 ... 1140 Ammodytes Panas Ammodytoides 1895) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1142 gilli (Bean Uranoscopidae zephryeus Gilbert and Starks 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astroscopus 1145 .. Kathetostoma Jordan and Bollman 1890 ... 1146 averruncus Blennioidei Tripterygiidae (Jordan and Gilbert carminalis ... 1150 Axoclinus 1882) Labrisomidae holderi (Jordan and Starks 1907) ... 1154 Alloclinus corallinum ... 1890 Cryptotrema 1156 Gilbert multiporosus 1953 ... 1158 Labrisomus Hubbs xanti (Gill 1860) Labrisomus 1160 ... Paraclinus integripinnis (Smith 1880) ... 1162 Clinidae Gibbonsia elegans (Cooper 1864) ... 1166 Heterostichus rostratus 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1168 Girard Chaenopsidae Neoclinus 1858 ... 1172 blanchardi Girard Hubbs 1174 ... stephensae Neoclinus 1953 Dactyloscopidae byersi Dactyloscopus Dawson 1975 ... 1179 heraldi lunaticus Gilbert 1890 Dactyloscopus 1179 ... Gillellus Gilbert 1890 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1180 semicinctus opercu/aris ... 1861 Myxodagnus 1179 Gill Blenniidae Entomacrodus (Jordan and Gilbert 1882) ... 1184 chiostictus Hypsob/ennius brevipinnis (GUnther 1861) ... 1186 Hypsoblennius gentilis 1854) ... 1188 (Girard gilberti 1882) ... 1190 Hypsoblennius (Jordan jenkinsi (Jordan and Evermann 1896) Hypsoblennius .. 1192 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypsoblennius proteus (Krejsa 1960) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1194 Ophioblennius steindachneri Jordan and Evermann 1898 ... 1196 Plagiotremus (Jordan and Bollman 1890) ... 1198 azalea Icosteoidei Icosteidae Icosteus aenigmaticus Lockington 1880 ... 1202 Callionymoidei Callionymidae Synchiropus atrilabiatus (Garman 1899) ... 1206 24

37 Gobioidei Eleotridae latifrons (Richardson ... 1212 Dormitator 1844) (Jordan Erotelis 1211 1895) armiger ... Gobiidae (Temminck Schlegel 1845) ... jlavimanus 1218 Acanthogobius and and Gilbert 1882) ... 1220 ios Clevelandia (Jordan (Bean Coryphopterus ... 1222 nicholsii 1881) newberryi 1856) ... 1224 Eucyclogobius (Girard Cooper 1226 ... mirabilis Gillichthys 1864 crescentalis 1892) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1228 Gobulus (Gilbert gilberti (Eigenmann and Eigenmann 1889) ... 1230 Rypnus lepidus Lepidogobius 1858) ... 1232 (Girard connectens Hubbs ... 1234 Lethops 1926 (Gilbert 1236 ... dalli Lythrypnus 1890) zebra 1890) ... 1238 Lythrypnus (Gilbert y-cauda (Jenkins and Evermann 1889) ... 1240 Quietula trigonocephalus (Gill 1859) ... 1242 Tridentiger californiensis 1879 ... 1244 Typhlogobius Steindachner Microdesmidae bilineatus (Clark 1936) ... Clarkichthys 1248 Sphyraenoidei Sphyraenidae Sphyraena argentea Girard 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1252 Sphyraena ensis Jordan and Gilbert 1882 ... 1254 Scombroidei Gempylidae multistriatus Maul ... 1260 Diplospinus 1948 Cuvier ... 1262 serpens 1829 Gempylus 1843) ... Lepidocybiumjlavobrunneum (Smith 1264 tripes Johnson 1865 ... 1266 Nealotus pretiosus Cocco 1829 Ruvettus 1268 ... Scombridae spp. 1272 Auxis ... Euthynnus lineatus Kishinouye 1920 ... 1274 pelamis Katsuwonus 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1276 (Linnaeus Sarda chiliensis (Cuvier 1832) ... 1278 Scomber japonicus Houttuyn 1782 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1280 Scomberomorus sierra and Starks 1895 ... 1282 Jordan albacares 1788) ... 1284 Thunnus (Bonnaterre Trichiuridae arigato 1995 Aphanopus ... 1289 Parin anzac (Alexander 1917)... 1289 Assurger Benthodesmus pacificus Parin and Becker 1970 ... 1289 Lepidopusfitchi and Wilson 1987 ... 1290 Rosenblatt Trichiurus nitens Garman 1899 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1292 Stromateoidei Centrolophidae Icichthys lockingtoni Jordan and Gilbert 1880 ... 1298 Nomeidae 25

38 Cubiceps baxteri 1923 ... 1302 McCulloch pauciradiatus Gunther ... 1304 Cubiceps 1872 (Gmelin Nomeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1306 1788) gronovii ... 1880 1308 Lutken Psenes pellucidus ... Haedrick ... 1310 sio Psenes 1970 Tetragonuridae 1839 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1314 Tetragonurus atlanticus Lowe Risso Tetragonurus ... 1316 cuvieri 1810 Stromateidae simillimus 1860) ... 1320 Peprilus (Ayres Pleuronectiformes Pleuronectoidei Paralichthyidae Group Cyclopsetta Citharichthysfragilis Gilbert 1890 ... 1330 gordae Beebe and Tee-Van 1938 ... 1332 Citharichthys platophrys Gilbert 1891 ... 1334 Citharichthys sordidus (Girard ... 1336 Citharichthys 1854) Jordan Gilbert 1882 ... 1338 stigmaeus and Citharichthys Gilbert 1890 Citharichthys 1340 xanthostigma ... panamensis (Steindachner 1876) ... 1342 Cyclopsetta querna (Jordan and Bollman Cyclopsetta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1344 1890) Etropus Jordan and Gilbert 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1346 crossotus ovale ... 1864) Syacium 1348 (Giinther Group Paralichthys Hippoglossina Eigenmann and Eigenmann 1890 ... 1350 stomata Paralichthys cali/ornicus (Ayres 1859) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1352 Xystreurys liolepis and Gilbert 1880 ... 1354 Jordan Bothidae Bothus 1862) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1360 leopardinus (Gunther Jordan 1890 Bollman sanctilaurentia ... 1362 Engyophrys and Clark 1936 ... 1364 Monolene asaedai taeniopterus (Gilbert 1890) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1366 Perissias Pleuronectidae Embassichthys (Gilbert 1890) ... " 1374 bathybius (Lockington ... 1376 Eopsettajordani 1879) zachirus Lockington 1879 ... 1378 Glyptocephalus Hypsopsetta guttulata 1856) ... 1380 (Girard /sopsetta (Lockington 1880) ... 1382 isolepis Lepidopsetta bilineata (Ayres 1855) ... 1384 Lyopsetta exilis and Gilbert 1880) ... 1386 (Jordan pacificus 1879) ... 1388 Microstomus (Lockington vetulus Girard 1854 ... 1390 Parophrys Platichthys stellatus 1787) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1392 (pallas Pleuronichthys Girard 1854 ... 1394 coenosis Pleuronichthys decurrens Jordan and Gilbert 1881 ... 1396 Pleuronichthys Starks and Morris 1907 ... 1398 ritteri Pleuronichthys verticalis Jordan and Gilbert 1880 ... 1400 Psettichthys melanostictus Girard 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1402 Soleoidei 26

39 Achiridae Achirus 1880) ... 1405 klunzingeri (Steindachner (Steindachner Achirus 1404 1869) mazatlanus ... Cynoglossidae (Jordan 1880) ... 1410 Symphurus and atricaudus Gilbert ... 1868) 1412 Symphurus elongatus (Giinther Tetraodontiformes Balistoidei Balistidae ... Balistes polylepis Steindachner 1876 1420 1786) ... 1418 (Bloch maculatus Canthidermis and Starks 1904) ... 1419 Sufflamen verres (Gilbert Monacanthidae (Osbeck ... 1423 scriptus Aluterus 1765) Ostraciidae (Bloch Schneider 1801) ... 1426 diaphana and Lactoria and Nodder 1796 ... 1427 Ostracion meleagris Shaw Tetraodontoidei Tetraodontidae 1842) ... , ... (Jenyns ... 1430 Sphoeroides annulatus , , (Steindachner 1870) ... 1432 Sphoeroides lobatus Diodontidae holocanthus 1758 ... 1436 Diodon Linnaeus Linnaeus Diodon .. 1437 1758 hystrix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Molidae (Linnaeus 1440 Mola 1758) mola ... ... 1776) (Pennant laevis Ranzania 1441 Principles and Terminology A marine fishes are oviparous large proportion are extruded in a gelatinous mass that floats to the of buoyant eggs that develop in and produce surface, separate, where development to the yolk-sac or first- the feeding occurs prior to hatching. Many shore- stage a rela- hatch and column water the of in part upper state with unpigmented eyes, tively undifferentiated the of waters (those fishes the species inhabiting a prominent Develop- yolk-sac. unformed mouth, and slope) upper and shelf continental eggs demersal have these with rapidly, progresses larvae yolk-sac of ment individually, either that are deposited on the substrate, nutrition larvae yolk, that the producing by supplied tended or more one by are that nests in masses, or in are capable parents. the small planktonic feeding organisms. on of often development species these In of proceeds the yolk-sac stage before hatching. beyond column Larvae reside in the upper water where they to the hatching, first-feeding larvae Upon travel the to continue develop and trans- eventually and grow, the of larval continue develop to they where column water upper The duration form fishes. into juvenile stage varies greatly among species, from several days the eventually and transform and settle back to bottom either as or In some species with juveniles. planktonic of the larval stage, bottom end At to many months. the demersal eggs, development within the egg progresses living juvenile the to settle must species (demersal) in the remain species pelagic of habitat whereas larvae the beyond newly first-feeding stage, and hatched water Eggs column. atherini- and beloniform many of morpho- and individuals other have well formed fins In larval advanced and features, pigmentation. logical attach they which with filaments have fishes form their are and retained in the fertilized eggs fishes, viviparous some of Eggs flotsam. mats algal floating to eggs or In Sebastes, scorpinines) fishes (e.g., most demersal lophiiforms, where development takes place. ovaries 27

40 young as first-feeding larvae and they are released Among species whose eggs have been nomic character. their larval development in the upper water continue a single oil 60% have glob- approximately described, lack and globules, 25% multiple have 15% ule, oil in typical oviparous species. The young just as column Embiotocidae, of the as born are surf family perches, The number, size, 1980). Moser and (Ahlstrom them or mature as even (or juveniles transforming larvae taxonomic are globules oil of important and position, characters. the do not become part and of some in males species) plankton. Teleosts telolecithal eggs, with the yolk mass have of egg Stage Egg type separate from developing embryo. This the cleavage, discoidal or meroblastic undergoes where plankton Most marine fish eggs encountered in yolk divi- cell is not incorporated in the cells during in transparent, ca. 1 mm and samples are spherical, contrasts sion. which eggs invertebrate most with This in the with mm 0.5-5.5 ca. is overall size the range diameter; (yolk cleavage have holoblastic incorporated 70% distinguishing the species ranging between 0.7 and of of a means and provides cells) dividing ca. in some mm (Ahlstrom and Moser 1980). egg of eggs. Development of the teleost Species kinds two 1.6 the initial begins that produces a mound cell with division eggs planktonic produce groups are that ellipsoidal egg. With scarids) some ophidiids, engraulids, g, (e. whereas of celis, the blastodisc, at one pole of the (blastomeres) cleavage the individual continued cells some demersal spherical be may eggs gadoids (e.g., A distinguish. to difficult blenni- and smaller become more flattened some and blennioids), somewhat (e.g., gobi- some (e.g., of urn-shaped or spindle-shaped oids), syncytial zone, the base the at present is periblast, the blastodisc continuous mass; yolk the around is and Eggs enclosed are oids). a thin in chorion or that shell nutrients transfer to be may functions its from of one smooth microscope but appears under the dissecting of this embryo. the At to developing the end yolk the pores has fine-scale surface features and when analyzed of base at a region development of the the early stage The distinctly be may shell microscopy. electron by to blastodisc begins proliferate cells, forming a thick- some (e.g., filaments or spines with ornamented thickened cell germ ring. A peripheral zone, the ened and atheriniforms, beioniforms, aulopiforms), hexago- forms at one mass, of the embryonic the shield, part various of (e.g., some sizes polygonal networks nal or the of axis the marking ring, germ longitudinal future clupeids, stomiiforms, aulopiforms, macrourids, and embryo. Cell proliferation continues in the blastodisc of the on the inner surface pustules pleuronectiforms), chorion (argentinoids), or a single protuberance or the germ and by a mass yolk enclosing gradually ring, (e.g., swelling engraulids and pleuronectiforms) some process known as epiboly. As this process continues, primordi- a rod-like into develops shield embryonic the of (e.g., fishes 5). (Figure groups a few In Introduction of epiboly (marked at completion embryo al the and, anguiHiforms and some stomiiforms) there is a second, by brain optic vesicles, "blastopore"), the of closure internal the chorion. egg membrane free from typically are otic lobes, myomeres some and capsules, tail continued With recognizable. organogenesis, the yolk chorion the the between space and The mass and the yolk the sac and begins to elongate off lifts is usually small, but in some fishes space) (perivitelline and heart, other organs become apparent. gut, Initial clupeoids, and stomiiforms, anguilliforms, (e.g., usually stage when this melano- forms at pigmentation is smaller considerably pleuronectiforms) yolk mass the region phores develop the from neural crest located shell and the relative width of perivitelline the than the in the dorsolaterally along the Later axis. embryonic yolk taxonomic is The character. useful a is space embryonic fin pectoral and finfolds median the period, in anguilliforms, dupeiforms, salmoniforms, segmented form and and other buds the eyes, brain, myomeres aulopiforms, most stomiiforms, myctophiforms, and differentiate. structures continue to Melanophores in homogeneous is most The teleosts. higher whereas it along the dorsum proliferate remain in may and rows of the yolk segments can be a useful taxonomic size the body, onto the yolk sac and or migrate ventrad on Trachurus, sphy- character. In a few perciforms (e.g., species In median finfolds. the and to globule, oil some and pleuronectiforms (e.g., some soleids) the raenids), on found pattern the assume to begin melanophores the initially yolk is segmented becomes but homogeneous In are useful and stages post-hatching identification. in in absence presence The stages. developmental later or species neustonic particularly others, or living at (those the taxo- oil globules within of yolk mass is a useful 28

41 Segmented A Pectoral fin bud B Yolk globule Oil Shell (Chorion) Migrating pigment Oil globule sac Yolk Dorsal fmfold finfold Ventral Anus Cleithrum c pigment series Ventral Foregut section gut -~----Trunk---~-----Tail--------I Head length Body D ~----- Snout-anus distance (Preanal lenl!Ttlll------~ length Head Snout length pigment Internal fin blade Pectoral Figure Introduction 5. Anatomical and morphometric features ofthe early stages of fishes. (A) Composite illustration of a fish egg modified larva from et ai. (1989). (B) Early and late yolk-sac larvae modified from Kramer (1960). (C) Preflexion Matarese modified from Fahay an (1983). Late preflexion larva modified from (D) original illustration by William Watson. 29

42 near surface), pigmentation is dense and covers sea the in diameter and, 3.5 at mm yolk-sac the hatching, hatching. before embryo the of develop- the approximately at are larvae stage same ca. mm 1.0 of eggs from hatching those as ment we For develop- egg divide purposes comparative 1980). (Ahlstrom and Moser diameter stage, early from 1) follows: as into ment stages formation the extrusion to blastodisc, advanced an of sac size The size of the yolk is related to initial egg of middle 2) end the to beginning the from stage, yolk used before hatching. of amount the to and the epiboly of to from end stage, late 3) epiboly, and is off-round sac yolk Typically, to oval at hatching the and development of appear- the degree hatching. The species with an may be elongate and but tear-shaped in structures, pigmentation in middle- organs, of and ance elongate stomiiforms). and clupeoids (e.g., gut If oil an and late-stage eggs provides a useful suite taxonom- of anteriorly present globule is it may be located or identifying the on information Additional ic characters. position in posteriorly or an intermediate in the yolk found be may eggs fish marine of in characteristics sac. Multiple oil or distributed evenly be may globules (1974), Leis and Watson and (1976), Russell Ahlstrom in globule(s) oil the of clumped in patches. Location Matarese (1980), and and Sandknop (1984). Moser identification. in character an important sac is the yolk Larva Stage Yolk-sac characterized is yolk-sac The stage the migra- by cells. of pigment rearrangement and coalescence, tion, black Melanophores pigment brown or containing Kendall this guide we follow the stage criteria of In with a yolk-sac et. al. (1984) and refer to individuals cells used for the the (melanin) are pigment principal identification of most fish eggs and that Other kinds those larvae, yolk-sac as yolk of remnants or have larvae. (e.g., present be may also cells pigment of erythro- yolk as larvae, and those that are in their all utilized of as juveniles to larvae from changing phores with red pigment, xanthophores with yellow process the transforming The specimens. stage transformation or pigment); in remains pigment melanistic only however, greatly varies at hatching development of state size and specimens formalin and preserved in fixed formalin in egg to related marine among is generally but fishes or guanophores alcohol. Silvery pigment produced in or, more precisely, diameter diameter. Typically, yolk remains in in specimens fixed in alcohol. juveniles of diameter the times 2.5-3.0 migrating body at hatching is length Melanophores capable are amoeboid and the of yolk (or shell in species with a narrow perivitel- from their in the neural crest to various sites on source be larva. migration the begin or may This com- may line space). Generally, yolk-sac larvae hatching from of eggs <1.5 mm have an unformed mouth, unpigmented majority late-stage in pleted embryos but, in the species, stage. yolk-sac the during occurs it The and pectoral eyes, buds, whereas yolk-sac larvae fin sites along principal migration is ventrad to the gut and well hatching from larger eggs are comparatively body may melanophores but margin also ventral formed, eyes mouth pigmented, the with developed, the of to and finfolds the to migrate regions various developed. pectoral cases, both In fins larval and first mela- head. engraulids), In (e.g., fishes some the is aided by a prominent median finfold that locomotion to on yolk-sac larvae is above nistic pigment appear caudal of around head the the the from extends top gut and on the ventral margin the of of the yolk appar- body; the posterior the to forward and region margin ently, demersal sac migra- ventral their complete melanophores the with (Figure Introduction 5). Most species of the go through the yolk-sac stage before the By eggs essentially synthesis melanin before tion end begins. are and coalesced yolk-sac stage, melanophores usually of hatching is development at hatching. The state first- of arranged in specific patterns characteristic a few families of of eggs the in advanced remarkably feeding larvae. have beloniforms typically example, For fishes. pelagic is fins median developed well notochord flexion and of the oil and yolk the stage yolk-sac the end At or either underway complete before hatching. In organ globule(s) the and up used been and have major pelvic and lampridiforms, the extremely elongate dorsal prey capture to required systems sensory functional. are visible clearly are and in form to beginning are rays the and mouth The formed, have at open is anus gut eggs. eggs, with large advanced species In some newly the margin undeveloped; hatched yolk-sac larvae are comparatively are pigment- of the ventral finfold, the eyes present. are fins pectoral primordial the and ed, 3.0- of Argentina sUus are for example, the eggs 30

43 Larval Stage larvae marine of diversity remarkable The teleost process an evolutionary that has produced an reflects et Following et al. Ahlstrom (1976) Kendall and al. immense array of specializations in form, pigment we larval stage into three sub-stages (1984), divide the and behavior and pattern, in is not if scope, equal and postflexion), (preflexion, based on flexion, the of Moser 1953b; (Orton adults the 1981; that scale, to degree of of flexion noto- the of section terminal the length Body 1984). al. beginning of the at Kendall et caudal fin. This the of system formation during chord times the larval stage is approximately 4 the 5 to convenient is by bias morphometric caused reduces and of the egg, therefore most first-feeding fish diameter slower in the body length relative to increase other larvae are 3-6 size in length. The ultimate ca. mm during the flexion structures of the notochord. varies attained greatly among species, with some (e.g., others and mm <5 (e.g., transforming achirids) at some reaching pleuronectiforms) before mm 100 > of the caudal fin begins with the Development maximum transformation. Most reach larval species initial formation ventral supports on fin of the the Shape range. varies greatly, 10-30 mm size within the Introduc- of the notochord near its tip (Figure surface elongate slender and elongate (e.g., clupeoids), to from plates flattened These 5). and parhypural (hypural tion (e.g., to short and elopomorphs), compressed and and subsequently first elements) are formed in cartilage (e.g., compressed to globular (e.g., acanthurids), become caudal fin rays are attached to The ossified. ). tetraodontiforms when forma- ray time the about at and, elements these tip the begins, tion begins to flex of the notochord differentiate systems organ The during the basic is developing When the flexion completed, upward. of these systems structure however, stage; larval the rays are in a longitudinal position and the posterior between and varies greatly within and orders a large preflex- the hypural plate(s) is vertical. The of margin unique stage. larval the to is specializations of array the of yolk-sac termination the at begins stage ion of the of One series obvious features of larvae is the and ends at initial notochord flexion. The flexion stage muscle body (Figure segments the or myomeres along stage ends stage begins) when the noto- (postflexion complement adult the species, most In 5). Introduction 0 the is its final position at approx. 45 in from chord tip myomeres of myo- since and, hatching by present is of posterior margin the stage, this At axis. notochord mere number of approximately number the is to equal 0 the from plate (90 vertical is the upper hypural provided with an form, is one will that vertebrae of lower the however, axis); margin notochord the important linking and characteristic adult larval early plate be not may or vertical completely be not may stages. the of upper aligned fully the posterior margin with plate Introduction 6). The postflexion stage (Figure Another the feature obvious in preflexion larvae, the to larval the from transformation ends when in basic trends gut, some exhibits major among form stage transformation on sections (see juvenile begins 6; and refer teleost Introduction (Figures groups 5 and criteria). defining for stages juvenile and 7 Introduction Figure to the end of introductory the at in shape gut of comparison for section inter-family and is elongate is straight The process described above produces what it Generally, larvae). mid-stage in teleosts (e.g., elopomorphs, clupeiforms, lower caudal a homocercal type basic the fin, of termed aulopiforms). some stomiiforms, some salmoniforms, teleosts. this is found in in to caudal exception An fin have it may anguilliforms some In with undulations the noto- of section terminal the where gadoids, the rays fin the and straight remains chord form symmetri- salmoniforms, clupeiforms, structures in and nodular differenti- is cally above and below it. Some other groups have primitive groups it stomiiforms, and other a hindgut into ated mucosal prominent a foregut, with lack anguilliforms) (e.g., fms caudal reduced highly or It from freely trails where In macrourids). (e.g., caudal fins cases entirely section. terminal and folds, a short in body aulopiforms some In stomiiforms. many it the not are notochord on based substages larval flexion saccular is and scopelarchids) (e.g., voluminous in and initiation the on are usually substages applicable, based development. In anguilliforms, of dorsal and and fin initially short is anal paralepidids) it some (e.g., others development. larval during larval substages are based on tooth morphology and In lengthens markedly from but to trailing, long, myctophids it varies to short, characters. several other 31

44 Supraoccipital Sphenotic Pterotic Posrtemporal Suprac1eithral Cleithral Rostral Opercular Posterior preopercular Subopercular Infraorbitals Interopercular Articular Anterior pre opercular jaw of lower Angle r-------------- length----------~ Standard Dorsal fin ---~ r---- Second First dorsal r dorsal1 caudal Principal " Soft rays fin rays Forebrain fin Caudal Dentary caudal Procurrent ray fin bud fin Pelvic I Cleithral symphysis ray Branchiostegal fin Anal Figure Introduction 6. Postflexion stage of a fish larva showing anatomical and morphometric features; the enlargement (above) shows of an spines. Modified from head original illustration by William Watson. examples 32

45 late usually moderate in length, slightly sigmoid, and has stage yolk-sac or embryonic is the in typically the functional, is fin Although 5). in gadoids Introduction is coiled prominent mucosal folds. The gut (Figure differentiation ray evolutionary higher shift. and teleosts, marking a major in is delayed until transformation in varies a gut The mass with rounded, form from teleosts lower many and (e.g., fishes higher some in fishes, pectoral fin-ray of section (e.g., gadoids, terminal short and coil compact majority flatfishes). In the to stage preflexion the early late begins formation in beloniforms, relatively to elongate (e.g., lophiiforms), the complement fin-ray pectoral adult and postflexion ophidiiforms, zoarcoids) but, in the majority some of present is by mid- or late postflexion. In some fishes, extends triangular roughly is teleosts, higher and to fins the which in those usually become highly pectoral The gut midbody. mass may about extend beyond well the profile ventral simultaneously or before developed, rays form early, cynoglossids) of (e.g., body or the some (e.g., fishes some In with caudal fin rays. the develop (e.g., elaborate trailing appendage some an fins the cynoglossids), and anguilliforms are pectoral ). bythitids and juveniles in but larvae in present adults. absent the jaws, head, and The relative size and shape of in the is great diversity of size and shape the There a large provide eyes suite characteristics identifying of in macrourids, (e.g., groups some fin and pectoral larvae (Figure Introduction in may be 5). The head and some myctophids) the fin base may be enlarged small snout a pointed with (e.g., and small mouth Such shape. a distinctive have are often characters a small clupeoids), small and rounded with mouth or useful in identifying species genera. globose and large hexagrammoids), (e.g., with a small large mouth large a with (e.g., tetraodontiforms), fin The appear. to caudal next the is usually The jaws toothed and mouth heavily scombroids), (e.g, or that are finfold caudal the those form to rays first in between forms intermediate of these in variety a of the the primary supporting elements with articulate size, in Larval extremes. shape, teeth are highly varied parhypural) (Figure Introduction 5 fin and (hypurals many and pattern and characteristics. identifying afford (superior) upper form rays principal These 6). in and of diverse. remarkably are eyes larvae Yolk-sac The with the superior groups (inferior) lower and associated elliptical, have teleosts most compressed dorsoventrally complement and inferior hypural elements. The full of eyes that become rounded, off-round, oval, or elliptical the rays completion usually of principal is present by and compressed early in the preflexion anteroventrally postflexion in the early or flexion notochord The stage. In stage. clupeiforms, lower many teleosts (e.g., be of principal within consistent rays to tends numbers aulopiforms) salmoniforms, stomiiforms, eyes the are clue orders or ordinal clusters, thus providing another relatively small and oval to strongly elliptical. In some linking early larvae with adults. Many lower teleosts tissue choroid of a mass groups, these within species salmoniforms, stomiiforms, aulopiforms, (clupeiforms, eye the of lies within scleral the at the base envelope have myctophiforms) 10 19 superi- rays principal with of and, in other species, the eyes are on stalks borne as 10+9). The 10+9 or and 9 inferior rays (indicated and shapes, sizes, of range wide A lengths. various formula in but beryciforms found is also formula the is found in larvae of the Myctophidae; specializations excep- the with teleosts, widely higher other in varies or of the subfamily Myctophinae have oval larvae usually that percoids have tion (9+8) 17 principal and some tissue, with some eyes, elliptical choroid of the at begins rays principal caudal rays. Formation generally Lampanyctinae of whereas larvae stalked, center of the fin and progresses (usually symmetrically) have round choroid no or little with eyes off-round or dorsally and ventrally. Additional caudal fin rays, the tissue. Usually, in higher teleosts, the eyes are round to the formed are rays, procurrent to anterior principal eyes later in round becoming oval with oval, slightly rays The by rays. are inferior procurrent supported larval stages. are superior procurrent rays the and spines haemal from derived elements or spines neural supported by fin development may occur Initial the as early as of procurrent rays general- numbers The spines. neural embryonic the or yolk-sac stage but, in most teleosts, consistent in major phylogenetic groups but not ly are in develop to begin fins median larvae. The preflexion of are specifically distinct. The unique structure often sequence a can fins individual of formation of be identifying charac- an provides fin caudal gadiform the character or higher useful taxonomic levels. at specific group that for teristic unique do III as arrangements a peduncular of The pectoral fin consisting primordial taxa. other is usually appear, and a thin blade base the first fin to 33

46 The and form simultaneous- dorsal anal fms usually in preflexion and are enlarged and highly developed and ly in the approximate in the fmfolds ventral dorsal the the in trends Phylogenetic stage. larval during positions they will along occupy in adults and this, are fins pelvic the of apparent and position composition and type the numbers with soft fin of rays (only rays providing when they begin forming in larvae, thus of charac- lower set teleosts), in most furnishes another set another of identifying characteristics. For example, help link larvae is that appear ters to First adults. and the pelvic fins are abdominal in position in lower a series of finfold; the of base the these at thickenings area teleosts and generally are thoracic (chest below the primordia are of elements the pterygiophores, the the pectoral fins) region) higher in (throat jugular or constitute the (Figure 5 and Introduction fin that base number The fishes. pelvic of greatly varies rays fin at the in form bases the argentinoids, In 6). finfold major among within trends some but groups fish and to some distance from the body and the are connected apparent: number the usually, are ranges rays of hyaline strands. In and with spinous body by fishes between 7 and lower in 12 but teleosts in most higher soft-rayed and anal fins, primordial pterygio- dorsal spinous one scorpaeniforms), perciforms, groups (e.g., are larger and usually soft-rayed the in phores part rays soft fewer) (or five and ray present. are part, pronounced more and than in the typical- spinous Elongation elaboration and of occurs in fin rays the in begin appear rays The earlier. forming finfold ly larvae throughout major teleost groups. the the Often thickenings as contiguous (bundles of actinotrichia) becomes fin entire many cases, one enlarged in but, to with the developing pterygiophores. Typically, differ- A and elongate become ornamented. rays several entiation the of dorsal and fins begins somewhat anal site the anterior region is common of the dorsal fin, anterior the middle to the of the soft-rayed portion of where a single spine (e.g., elongate be may ray or throughout fin and continues the posteriad and anteriad bothids) or a group melamphaids, ray(s) or of spines stage, with the adult complement larval becoming forming sometimes elongate, (e.g., lampridi- are a crest apparent by late larvae In postflexion. gadoids some of common site paralichthyids, forms, achirids). Another with and two dorsal anal posterior the fins, multiple the fin pelvic is the elongation for entire fm is where and dorsal separate within differentiate fins anal often enlarged (e.g., gadiforms, lophiiforms, beryci- more centers, whereas simultaneously, less or differen- or forms) several rays may become elongate and to one tiation Segmenta- the anterior dorsal of fin is delayed. paralichthyids). ornamented (e.g., lampridiforms, of usually apparent rays soft the in postflexion is tion some fins pectoral the Likewise are enlarged species in pronounced and becomes more larvae by the end the of myctophiforms, of groups many (e.g., aulopiforms, larval stage. In larvae species, of posterior- the some lophiiforms, or, perciforms) scorpaeniforms, rarely, one most spine the dorsal and anal fins of becomes also to several rays may become elongate (e.g., myctophi- rays. segmented its tip is fimbriated as in soft and forms, peristidiines). Elongation of rays fin is rare; anal Transformation a spinous into ray soft apparent this of provides Alectis fin anal anterior the in ray elongate the at occurs ray end in the larval stage or of early few the of examples. one and obscured segments the when juveniles, the become tip pointed (Mansueti 1958; Berry and becomes occurs, or elongation of rays fin Where enlargement Matarese Richards 1973; species many In al. et 1989). formation ray stage, early an at initiated usually is dorsal fins anal and is the of ray soft posteriormost the or even in the egg in early often preflexion yolk-sac or In base. fin the with articulation at its divided some stage. Elongation is usually associated with the devel- anal and dorsal may the fins fishes (e.g., clupeids) and opment of structural pigment special elaborations their migrate forward from initial position during the theme in these specializations is recurrent A patterns. there larval stage some and in others (e.g., myctophids) mimicry apparent their coelenterate structures, as of by allometric shift is an apparent in position caused exemplified by dorsal fin the of lampridi- carapids, region of the posterior of the tail. growth (Liopropoma forms, some epinephaline serranids and pelvic typically are the last fins to begin The fins some and in rays pectoral elongate the Diploprion), and, to forming begin not do teleosts, lower many in myctophids (Loweina and Tarletonbeania), and the of period, larval the just until the end differentiate of and bothid some pelvic rays and dorsal elongate before In 6). Introduction (Figure transformation some flatfishes. paralichthyid The recurrence of such similar aulopiforms, myctophiforms, (e.g., some groups structures in unrelated groups invites speculation on lampridiforms, form beryciforms), early the pelvic fins their possible function. siphono- If they are mimicking 34

47 are phores other poisonous coelenterates they could are larvae in present or that spines many and larvae adults. in lacking as Moser function in discouraging potential predators and al. (1984) (1984) al. Kendall et (1981), Govoni et highly is Pigmentation and fishes larval in evolved have suggested to serve may they or food, since attract of useful identifying characteristics. provides a wealth have siphonophores many parasitic or commensal Since observations are usually our preserved limited to for size fish of appropriate prey hyperiid amphipods remains, the in which only material, pigment melanistic 1991). Also, these complex al. larvae et (Baldwin of the overall pigment pattern. we see only a part involved larval sensory with structures or may be of Melanophore size, pattern, and sequence shape, have may they or functions, hydrodynamic multiple (Figure features taxonomic important formation are functions. be Individual 5 and Introduction melanophores may 6). round, dashes), linear (streaks steliate, or dendritic or become may finfold median The highly specialized open on shapes other or in based circles, the rectilinear, it usually larvae. higher teleosts, fish In in some distribution of the and melanophore the of shape becomes reduced as is the median fins form and within melanin it. Melanophores may be located postflexion relatively however, larvae; insignificant in beneath superficially the dermis, in epidermis, the or the it may remain a prominent feature throughout cavity, internally in muscle tissue, the cranial or on (e.g., teleosts lower argentinoids, in stage larval bladder. perivisceral the of and gas cavity membranes myctophids). The stomiiforms, some preanal finfo.ld melanin and be amoeboid melanophores are can Since perSIst the may (portion of anus) finfold anterior to the redistributed within them, there is for potential great and stage postflexion the into weB developed is often cells change in the appearance of individual pigment median may lower teleosts. in entire The finfold their aggregate pattern. These changes can occur in and become appearance inflated an have in and voluminous as such periods, a melanophore over short time when myctophids the Tar- (e.g., fishes some and Loweina from form apparent its changing contracts, to stellate letonbeania). pigment or round, through patterns develop when proliferation, migration, and coalescence. This is first In addition to the specialization in spines of fin initial the when stage yolk-sac the during apparent . larvae of many higher teieosts, there has been a is pattern Pigment is formed. pattern pigment larval evolution remarkable larval of body and head spination unique genetically determined; many species have in (particularly Scor- in spination rivals that adults patterns, and often there are pigment arrangements that and paeniformes Perciformes). of One the ~ost com- higher typify Apart from order. to genus from taxa mon sites for larval spine development the IS on unique patterns are many convergent pigment such lod and anterior angulate where bone, post~ri­ preopercular independently have that arrangements and in evolved or series develop with an enlarged, often serrated spme apparently clades, unrelated in response to common the Introduction of (Figure posterior series the at angle me!ano- of rows Examples are the selective pressures. common eye, where one the is above 6). Another site the in phores above intes- elongate an with gut larvae a bony or project- crest in present be may spines more that gut overlies mass in shield the melanistic the tine, bone. frontal scorpaeniforms Many the from ing the intestines, the overlying shield coiled with larvae a pair develop dorsolateral parietal crests (usually of gas ventral the melanophore series on the bladder, in whereas many spines, in serrated) terminating margin of tail, melanophores that cover the brain the crest serrated a single perciforms terminat~n? in a s~ine melanophores along the dorsa! of lobes, paired rows midline the regIOn. supraoccIpItal the develops of in many in species, and, of margin body, the neustonic on other cranial bones (e.g., often Spines develop sheath the of pigment covering the entire larva solid on the infraorbitals, pterotic, posttemporal), opercular, The for the caudal functions region). (except of these bones of the shoulder (supracleithrum and girdle to patterns specific highly and open are convergent the of cleithrum), or on specialized bones or scales but their 1984) al. et Govoni 1981; (Moser speculation body malacanthids, agonids, in (e.g., chiasmodontids, usefulness of characteristics larvae as fish identifying transient are spines achirids). In most fishes these at unquestionable. is levels taxonomic an and juveniles in reduced or lost are and features larval of larval the There are numerous other features spiny- a group that scorpaenids, in Even adults. has summarized those to addition in stage, are that above, spines headed adults, the in prominent more are head 35

48 in changes by accompanied is Transformation of recognizing useful species and higher taxa. One in I may species Epipelagic habitat. in often and behavior is the as these early presence of photophores in larvae the move or habitat same form in remain and schools apparent most are These stage. preflexion as the in leave to inshore nursery areas. Demersal species must patterns myctophid larvae where there are specific of and to common patterns photophores early-forming to the surface waters and settle the juvenile habitat; the critical this accomplish they which by mechanisms life genera 1970, Ahlstrom and (Moser genera of groups or larval history stage unknown. largely are The process 1972, 1974). larval Other specializations are the is protracted larvae where species the demersal some in balloon-like outer skin of lophiiforms and the special- may remain pelagic for many months often and attain ized body form of leptocephali. Surveys of elopiform a large size. Numerous examples this of may be found larval their and fishes of specializations many the in families several of flatfishes (paralichthyids, bothids, use potential found be may identification in in Orton pleuronectids); and achirid flatfishes provide an exam- Moser (1981), (1953b), Berry and Richards (1973), stage the is brief other extreme where of larval the ple and Kendall (1984), al. et et Govoni al. (1984). at and settlement occurs mesope- mm. of Larvae <5 juvenile descend to their lagic and bathypelagic fishes Transformation Stage from that habitat, which often is different the of adult, transformation. to complete stage The larval is followed by a transformation Stage Juvenile form and in general stage characterized by changes of adult detail structural that involve the acquisition produces Successful which ajuvenile transformation change characters. The larval of characters and loss a pre-reproductive as generally is individual defined and from larva to juvenile may be abrupt or gradual is that morphologically similar to the adult with involve or the In changes. large small relatively may Juve- complete fin-ray complements and squamation. great majority from is different shape larval of fishes, habitat niles may adult the occupy or often live sepa- that and of requiring juvenile, the substantial some- rately size. adult Some fishes near until they grow to growth allometric times rapid bony muscle of and polyprionids) (e.g., lampridiforms, oreosomatids, have of from different is larval the Usually tissue. gut that juveniles pigmentation that differ markedly in form and juvenile and transformation involves the formation the in and many families (e.g., adults from shorefish development and other of further the of a stomach serranids, pomacentrids, sciaenids, lutjanids, carangids, digestive transpar- Since most larvae have thin organs. adult and juvenile the labrids) color are patterns ent skin, transformation the development requires of markedly (e.g., shorefishes Many different. mugilids, pigmented the much thicker, solidly juvenile skin. pelagic a special have cottids) hexagrammids, stage appears in guanophores, in produced pigment, Silvery and a dark with is dorsum that sides silvery generally many at this time. Scale formation usually fishes herring-like in appearance and behavior (Kendall et al. begins during transformation in species that have 1984). Other pelagic and demersal fishes have highly defining is most the useful scales, and often this for modified remain may that stages juvenile pelagic characteristic of develop- Fin stage. transforming the in common are stages pelagic Such periods. extended ray where point the to progressed has ment adult described often were families reef tropical many and complements or are nearly complete. LarvaI present mistakenly as unique species (Hubbs 1958; Leis and fin structures such as and rays enlarged elongate fins, 1983; et Kendall Rennis the California aI. 1984). In free-trailing barbels, exten- eyes, stalked appendages, Sebas- Current the region scorpaenid species in genera sions the gut, specialized larval teeth, and volumi- of may that juveniles pelagic have Sebastolobus and tes resorbed fmfolds nous are or lost through allometric settlement. before months for offshore remain small growth. Larvae with relatively eyes may experi- with in and, size eye in increase a rapid ence those summary above the in used terms are of the Many eyes, the shape elliptical or oval narrow changes abbreviated in the descriptive accounts guide; of this eye the round achieve rapidly to of adults and juveniles are these and other terms used in these descriptions that In some clupeoids, the round dorsal, have eyes. defined in the following list abbreviations and in of the fins luminous shift position. In may anal, and pelvic institutions, glossary following it. Acronyms for other and photophores fishes, tissue develop luminous given and research programs are expeditions, in rapidly. Table Introduction 1. 36

49 Abbreviations L lD lower first dorsal fin line lateral LL 2D dorsal second fin 11 LVL length anal fin (e.g., A Bathophilus) of liver number vertebra fin anal of anterior to rays the 31st blood vessel vertical last LVBV A30 (carapids) MDR mid-dorsal bony (agonids) plates number rays anal of to vertebra 201st the anterior lower ML from of the anterior jaw, measured the length margin of the (belonids) eye (nemichthyids) MVR mid-ventral (agonids) plates bony fin adipose Ad anal initial development fin the (e.g., stage after MxL margin posterior from snout to distance of AF maxilla macrourids) NL notochord length number fin origin of vertebrae anterior to OG anal oil globule AFO length organ anterior margin light on barbel 00 (platytroctids) eye of of mandibular BbL base fin pectoral at depth body occipital ORL BD of dorsal fin-ray (bregmacerotids) length body depth at anus, including gut protrusion if pectoral fin BDA PI flatfishes) (e.g., base present (macrourids) fin of pectoral length PIBL exclusive body length depth (length inflated greatest ray; of of epidermis fin pectoral of longest BDi PIL formation) of blade before ray juvenile length benthic BJuv beak (e.g., hemiramphids) of length length elongate lobe of pectoral fin (e.g., Ichthyo- of BkL coccus) body length BL depth pectoral fin base of rays branchiostegal BrR fin caudal pelvic fin C caruncle diameter longitudinal measured pelvic of Ca the to disc, outer edges disk (cyclopterids) of rays principal fin caudal C I length pelvic fin P L caudal procurrent rays fin C 2 2 choroid eye of below tissue length SpL P CL spine fin pelvic length (gempylids) 2 length of fin from measured peduncle, anal caudal preanal length; length Sn-A as same the CPL PAL length spine of parietal PaSL apogonids) (e.g., insertion PdL origin fin of snout to dorsal tip distance from vertebrae caudal juvenile pelagic PJuv fin dorsal (carap- 31st dorsal of fin rays to the vertebra number PoF postflexion stage PrAF macrourids) (e.g., fin development anal initial to prior ids) number fin dorsal of vertebra 201st the to rays vertebrae precaudaJ PrCV (nemichthyids) PrF preflexion stage of anterior vertebrae to D origin number preopercular longest of length spine DFO PrSL PrVL to (carapids) base vexillum snout from distance bony plates (agonids) dorsolateral DLR rostrum length length RL dorsal spine longest of DSL disk (maximum width width ogcocephal- of head in light subopercular organ (platytroctids) SBO DW standard length SL ids) distance snout-to-anus Sn-A eye) round (in diameter eye ED eye length (long axis anal to snout from origin fin distance Sn-AO oval or elliptical eye) of EL distance from snout to tip loop gut of Sn-GL life history early ELH SnL snout length length stalk eye ESL oval of eye) of supraoccipital crest or spine elliptical or SoCL length (short eye width axis EW species ( singular) sp. stage flexion F be or (may more species) species one sp(p). greatest depth of body GD (plural) species gular photophore (platytroctids) spp. G0 2 length gut trailing section of of TGL gill rakers GR total length TL length head HL stage transformation Tr head width HW illicium length (Iophiiforms) upper U IL distance between internasal width; the nasal capsules V vertebrae InW (carapids) origin fin dorsal to vertebrae of number VDO (tetraodontids) between photophore bases of pelvic fins VLR (agonids) ventrolateral bony plates IVO origin VVO (carapids) vexillum of vertebrae to (platytroctids) number y-S stage) yolk-sac for yolk-sac (as used eye lower to of margin (ventral depth jaw angle of JD (referring itself) sac yolk to sac yolk YS lampridiforms) jaw; in Juv stage juvenile 37

50 Glossary (As Jones et al. 1978, Fahay 1983, Leis in Trnski 1989, and in this study) used and Threadlike fibers in the finfolds of fish larvae that Actinotrichia. the Bathypelagic. Zone below 1,000 ocean. open m in depth into eventually rays. fin develop the in or on Living Benthic. (substrate). bottom to Adhesive. to or substrate the other stick eggs adhesive Sticking; eggs. Blastodisc. Early embryo of consisting of a disc- or cap- teleosts yolk. mass of cells on the like the dorsal fin. Adipose fin. Fleshy fin posterior to the forming cells Individual Blastomere. ofteleosts. embryo early to usually Aliform. shape the Wing-like; referring pectoral the of fin. Blastopore. Circular area on the yolk has of teleost fish eggs that been during not covered by the advancing germ ring epiboly; not fishes Referring to rivers Anadromous. to ascend that spawn. the blastopore homologous with of amphibians other and some invagination is there where vertebrates the to surface of cells (usually Fin fin. the on gadiforms) in some double but Anal single interior through 1964). Ballard blastopore the (see embryo the of margin ventral of tail. the of the dorsal the from margin vertical depth. Body The distance gape jaw; region of of jaw. Bony prominence posterior to Angle margin ventral the to body the the of base at the measured body of where fin pectoral it attaches to the body; fins fin bases are not or bones. quadrate and articular, of the angular, of the junction measurement. included in the the Angular. The bone at angle the jaw or that region; used in of guide this angular in reference to pigmentation at, or near, the of a larva; length. term used to indicate the size General Body bone. snout the of (distance equivalent to notochord the from length tip tip to the the of preflexion and flexion stage larvae in notochord) or anatomical structure; plural of an organ Anlage. Primordial form the (distance snout tip length standard to from of the and equivalent postflexion larvae, plate) hypural the in of margin posterior the to is anlagen. juveniles, and adults. front; the Towards Anterior. cephalad. Branchiostegal rays. Ray-like bony elements attached to the hyoid a spine. of forward; to direction referring usually Angled Antrorse. and a arch, extending under the connected gill openings by membrane. surrounding Anus. Orifice and terminus tissue at the gut. of appears bump undifferentiated The Bud. at the that protuberance or initial formation fins. paired the of Aorta. chord that supplies blood Main vessel lying below the spinal heart. the from blood in ceratiids. rays fin dorsal modified Caruncles. Fleshy outgrowths; the of and the dentary between jaw lower Articular. A bone fish. the of fin end Caudal fin. Median posterior at the angular (some bones it part consider this the of in used angular); angle anterior guide in reference to pigment to the the of upper part Caudal peduncle. Narrow between ofthe tail end posterior the jaw). base the and fin the or anal caudal of fin; caudal the of dorsal peduncle length measured from the insertion posteriormost the of that of pigment contrasts with the adjacent back- Band. strip A ray the the anal or dorsal of (whichever is most posteriad) to fin or ground pigment unpigmented area; it may be on any part of the the anteriormost caudal fin-ray. of insertion may body, or fins and head, this position; any in oriented be in laterally a vertical band guide, body or occurring is the on head of vertebrae extending Caudal group posterior The vertebrae. is a stripe. as referred to as a bar band a horizontal to referred and the from urostyle. the to spine haemal first the bearing centrum A vertical band Bar. pigment of of on the head the surface lateral or The surrounding membrane Chorion. shell egg. an of body; often ocurring in a series. or a vertebra. The Centrum. body of barbel sensory Barbel. Slender projection on the lips or chin; of to the tip the on head attachment point the from measured length eye Choroid fissure. Indentation at the ventral margin of the the barbel. of marking invaginated borders the of fish. optic cup in larval the that supports the pelvic Basipterygium. Bone fin. or process Choroid tissue. Mass various of primordial vascular tissue of 38

51 shapes the eye; usually associated with narrow eyes; below lying jaws equal in length, no and nasal capsule, an lower undifferentiat- axis longitudinal its measured length its pigmented; often along finfold, ed and head the and straight, tip notochord the hypurals, no of the to eye the tip of portion with pigmented interface from the of large. relatively body the preanal region choroid the mass. The part Epaxial. myomeres of the above the lateral midline (horizontal septum). ventral of ends the or junction. Where the Cleithral symphysis bones meet. cleithral sea the with interface the at Epibenthic. to bottom; Zone refers Cleithrum. Elongate vertical bone in pectoral girdle at the junction sea with contact in living organisms the bottom. one body form to bones first the of in fish; the of and head the of body. and head the of junction the demarcates body; the Epiboly. Movement ofthe the embryonic of surface cell mass over ring boundary the yolk; the germ sheet marks the advancing the of gut where it is twisted or convoluted. cells. of the of Condition Coiled. depth m open the in 200 tb the from Compressed. Laterally surface Epipelagic. Zone flattened. ocean. CroWD. Top of the head. pigment Red Erythrophores. cells. An elongate or streak-like melanophore. Dash (pigment). lure Esca. The at the tip fishes; lophiiform most in illicium of species. luminous commonly deepsea in used stages the ocean bottom; At in reference to life Demersal. to adults. from eggs Euryodontic stage Advanced stage. larvae anguilliform of branched; used in reference to a usually Highly Dendritic. engyodontic smaller (following a relatively stage) characterized by of relatively teeth, short, broad melanophore. series region, head three preanal and capsules, lower shortened a relatively jaw, the formation of nasal the of notochord. teeth. bearing usually jaw, lower of fins, and hypurals, and the flexion Dentary. Major bone larvae rays Disk. diameter. Eye with round eyes, Flat the diameter In cup-like structure formed from modified pelvic fin the of (also in some ofthe eye, usually measured horizontal the through rays pectoral fin the onto holding for used fishes) part pigmented substrate cyclop- gobiesocids, (e.g., oval the on structures or substrate with larvae in midline; elliptical horizontal the eyes, or first, dimension. vertical dimension is given the followed by is measured (PPL) longitudinally terids, its diameter gobioids); is opposite one margin to the from margin; in echeneids, the disk Eye axis longer The length. spines; or measured eye elliptical oval an of on the top of the head and formed from modified dorsal fin ogcocephalids of one side of the where disk of from the pigmented margin through midline head disk-shaped the to refers also the the eye side. other to the disk. of transverse width dimension greatest is the (DW) width. The axis shorter Eye opposite or ellipical oval to proximal. an attachment; of measured eye of from Remote Distal. the point of one side of the the margin pigmented the through midline from Dorsal.. Upper part of ventral. of opposite body; to side. other the eye of the bearing length of the body. varying eyes; Dorsal dorsal on fin. Fin margin or fins peduncles stalks. Eye Movable stalk the where point attaches eye the from measured length stalk part Uppermost Dorsum. opposite eye. margin; body dorsal body; of the of to attachment at to point the to head the ventrum. (pterygio- Fin supporting and rays~ spines, Fin elements. bones Early life history. The early phase in life spanning the develop- phores). mental from egg to juvenile. stages the which within of skin surrounding body Finfold. fold Median ventral the of the anal, fins caudal and dorsal, the part develop; Indented; used to describe caudal fin shape. Emarginate. develops fin anal the which in finfold anal the as to referred is of differentiation and growth; Embryo. Organism at an early stage finfold. guide, in this fertiliza- between development of stage the to refers to anal and dorsal the posterior structures fin-like Small Finlet. hatching. tion and fins. Embryonic shield. Thickened area representing of the germ ring of axis longitudinal the future the embryo. of their larvae. most or all used have that Larvae First-feeding of prey. capturing capable are and yolk (preceding of anguilliform larvae Engyodontic Early stage stage. teeth, euryodontic stage) characterized upper the few needle-like by dorsally Flexion. Stage (or the process) when the urostyle bends 39

52 concurrently the development of principal rays and hypural with horizontal the (in tissue connective of A sheet septum. Horizontal caudal bones in the fin; also, the process notochord flexion; of plane) the masses; muscle hypaxial and epaxial the separates that to bending initial the to preflexion refers prior the stage of the pigmentation. midline area in reference to lateral stage the comple- after notochord tip and postflexion refers to the tion Refer and Principles the to Terminol- flexion. notochord the of Hyomandibular. cheek Bone or cartilage (usually elongate) in the section ogy for additional explanation. that region functions in suspension. jaw Anterior region Forebrain. developing the of includes that brain of midline the below myomeres the lateral part The Hypaxial. septum). (horizontal lobes. olfactory the Anterior part Foregut. alimentary primitive from of the canal Hypurals. A series of of bones derived from the haemal spines the esophagus which the and stomach develop. caudal last support which vertebrae rays. fin principal (fused Large bones. Frontal in that some species) bones paired consisting the of zooplankton the of part That Ichthyoplankton. form top the egg larval stages and fishes. of of over cranium anteriorly; they often form a ridge the each spines. more or one bear may that crest) (supraocular orbit Tentacle-like spine Illicium. of snout fin the the on located dorsal attract to a lure as used fishes, lophiiform most genus of chiasmodontid illicium prey; Gargaropteron stage. Pelagic stage of the greatly Kali, fin pelvic and pectoral elongate rays. by characterized length measured from the point the of insertion of illicium on the to the tip of the structure. head lying Gas-filled bladder. Gas in the column spinal the beneath sac Inferior. Spatial anatomical term meaning lower in position; referred to bladder. region; abdominal also as air bladder or swim to superior. opposite Thickened margin ofthe Germ that over advances ring. blastodisc point (usually) Posterior Insertion. attachment a fin. of of yolk the epiboly. during bone InteropercIe. Lower the of the gill cover lying below crystals Guanophores. Silvery pigment cells containing iridescent preopercle. guanine. of Interorbital. Region on top the between head the of orbits the of Gular. Ventral region the the isthmus head and anterior to of eyes. jaw. below the lower narrow, of the head below often gills, the Ventral region Isthmus. Alimentary Gut. tube and organs. associated gular the connecting regions. cleithral and found Gut Loop, fold, or curve loop. along axis of the gut. the indicate the region; usually throat to the Referring Jugular. used to position fishes. some of the pelvic fin in spine A median on the ventral surface Haemal spine. a vertebral of an arch form centrum, attached to the centrum by two bones that is Stage after transformation from the larval stage that Juvenile. arch). (haemal there a full is active; reproductively yet not but adult the like formed rays, that species (in have are scales fin of complement length. Head Horizontal distance from the tip the of the snout to and form the adult. them), the to similar is fundamentally margin posterior the of cleithrum. anteriormost The Lachrymal. the of infraorbital (circumorbital) width. Transverse Head distance between the lateral margins ofthe with species. some in spines and enlarged bones; orbit. the of at head the posterior margin measured the is unlike that Stage adult or hatching following juvenile Larva. developing the brain that of includes region Hindbrain. Posterior and metamorphose in form or pigmentation and must transform medulla. the juvenile/adult assuming before characteristics. the the includes that of canal alimentary Hindgut. part Posterior vessel. Last vessel blood large vertical blood posteriormost The and intestine rectum. dorsal extending from the as aorta to the kidney (nephros); used eel and point leptocephali. reference in character taxonomic upper of caudal fin in teleosts where the and Type Homocercal. lower lobes are symmetrical; internally, the major structural elements fin ofthe formed by of are an the upturned urostyle (several fused body verte- line. Lateral A line along the lateral surface parhypural) and (hypurals flattened with articulates that brae) a series bones are sensory usually that of modified pores associated with scales. rays. fin that support the principal opposite to segmented in Homogeneous. Uniform in composition; the of body between epaxial and Lateral midline. The region referring yolk. egg to hypaxial myomeres; the region septum. of the horizontal 40

53 Lepidotrichia. the of soft form that structures Scale-like segments tissue between Myosepta. Connective myomeres. adjacent fishes. bony in rays region Nape. Dorsal the of body immediately behind the head. Transparent, large of Leptocephalus. ribbon-like, often larvae with fishes an elopiform internal cavity filled with acellular openings of the nasal organs Nares. rosettes. The or and head a small have usually leptocephali material; mucinous prominent teeth. spherical Paired, more-or-less Nasal structures on the capsule. organs. the snout that contain olfactory reproduction the eggs develop within where of Type Live-bearing. or juveniles; as larvae vivipa- the and are born the young ovaries in open than (rather Nekton. Motile, marine organisms living water rous. the sea floor) and capable to small currents; against swimming of moderate-sized organisms (e.g., midwater fishes, juvenile nektonic as that separates of sheet tissue micronekton. A septum. Longitudinal connective referred are cephalopods) some fishes, to and the on the right masses left sides; the dorsallongitudi- muscle nal septum separates the epaxial muscle masses and the ventral Kidney; used to refer to larval kidney. often Nephros. septum longitudinal separates the hypaxial masses. to seaward of Pelagic zone coastal margin the extending Neritic. shelf. the continental shaped. Lunate. Crescent the Maxillae. located jaw; Longest upper the of paired bones above spine surface the dorsal A median of spine. Neural a vertebral on centrum, to the centrum by two bones that form attached arch an premaxillae. spinal the which through arch) (neural cord passes. Median. Referring to the midline plane that divides a bilaterally usually synonymous symmetrical animal into right and left halves; the of ridge developing neural the of Region crest. Neural embryo that differentiates into many kinds including of tissue and cells, with mesial. melanophores. or black ameboid melanin; containing A cell Melanophore. brown plankton ocean; sizes the of surface derived the Inhabiting Neustonic. of neural various shapes from the and in living cells pigment this zone as neuston. are referred to of the embryo. crest region Notochord. that rod cartilaginous the axis Longitudinal supports in series occurring characters. (e.g., Countable structures of Meristic body. the myomeres, vertebrae, fin rays). Notochord the tip of the to snout length (NL). The distance from open the in depth m 1000 to m 200 from Mesopelagic. Zone notochord. the of tip posterior the ocean. end at the N Metamorphosis. A marked change in form or structure nchal. nape; the of region the to Referring behind immediately loss and characters adult of acquisition involving stage larval ofthe the head, dorsally. transformation. with synonymous characters; larval of head. crest. Occipital ridge located posteriorly on top of the Bony of developing brain that includes the optic the Region Midbrain. and cerebellar lobes. the of seaward zone shelf or Oceanic. sea Open slope. continental Midline. The median plane or line the of the body either on dorsal fatty Oil of yolk the Spheres within material globule. or oil of used sometimes surface; ventral or the as part. a body of middle fish some eggs. Mucosal folds. Folds lining tissue of intestine; the of cavity the Ontogenetic develop- with associated characters Those characters. rugose some in appear- fish larvae giving a striated or pronounced mental stages. ance. the of Upper Opercle. bone largest usually and posterior cover gill series; in occurring body the of Myomeres. Muscle segments of used as synonym of operCUlum. a fish; often approximately equal to number of vertebrae in adults; in this guide, myomeres the preanal anterior myomeres all as defined are to a gill Operculum. Bony plate of the cover. anus of the postanal the and vertical from posterior margin the are those myomeres posterior to a vertical from the myomeres salariin some of stage Pelagic stage. Ophiohlennius blenniids preanal of the anus; in many fishes the number of margin posterior enlarged, by enlarged teeth hooked pectoral and fins characterized of precaudal number and postanal myomeres approximate the in or jaw both jaws. the in lower anteriorly (abdominal) and caudal vertebrae, respectively. larval or embryonic The Opisthonephros. kidney. Myomereformula. Preanal myomeres+ postanalmyomeres = total myomeres. 41

54 Orbit. of eye. socket bony The the membrane lining tissue connective associated and The Peritoneum. the gut cavity. period advanced relatively The Organogenesis. of embryonic the systems. organ of by development characterized formation space. Fluid-filled Perivitelline between the embryo and space shell or chorion of an egg. Origin. (usually) point Anterior of attachment of a fin. Photophores. Luminous organs. Ossification. formation calcification involving of Process of bone tissue. cartilage or connective Referring to the Phylogenetic. evolutionary lineage an organism. of Region Otic. the hearing ofthe head organs. containing auditory or Small, free-living, weakly swimming passively or Plankton. organisms or fresh the with drift that water floating marine develop Oviparous. Producing eggs that the maternal body. outside currents. Referring Paedomorphic. retention phylogenetic to the of juvenile Posterior Postanal. to the anus. stage. characters larval) (or in the adult Posterior. Towards the back caudal region; opposite to anterior. or of the roof of the the margin outer on lying Bones Palatine. often bearing teeth. mouth; or eye socket. Postorbital. Behind the eye Papilla. or protuberance. fleshy A projection posttemporal the from emerges that A spine spine. Posttemporal located posterolateral region upper the on bone skull. of the bone of Parhypural. Lowermost the principal supporting caudal fin rays. anterior Located Preanal. to the anus; preanal length (synonymous the from measured distance) snout-anus with tip of snout to the the Pectoral fin. behind fins ventrolateral) (sometimes lateral Paired anus. the of posterior margin pectoral prior to ray formation, head; fin length is measured the from the base distal ofthe blade of margin greatest the to (finfold) of includes vertebrae that Precaudal vertebrae. The anterior group is fin length blade; from the after ray formation, pectoral measured anterior to the centrum with the all first centra heamal spine. point the of of insertion of the ray. tip the to ray longest the specialized As Precocious. formation early guide, this in used of Pectoral fin Supporting structure base. the pectoral fin; of in a structure or fin elements) compared to typical develop- (e.g., fins fin; the operate that muscles the contains and it is peduncular larvae most abnormality. infer not does fishes; mental in sequences longitudinal from point the its length is measured along the axis of blade or point the to body the on insertion of attachment of the fin Premaxillae. of Paired the upper jaw anterior to the maxillae bones is axis widest its at rays; transverse the on measured its depth and bearing teeth; often protrusile, and extending ventrad usually (usually point. distal) most the in advanced teleosts. of most of maxillae small, A stalk. short Pedicel. Preopercle. the of bone anterior Upper bearing often cover, gill serial spines; spines the margin at as to referred are bone the of posterior and those on a bony ridge forward preopercular spines the to a structure connects that stalk or part narrow A Peduncle. of caudal fin to body). connecting peduncle caudal (e.g., body are the margin referred to as anterior preopercuiar spines. away sea usually the from Pelagic. sea the in Free-living bottom, caudal Principal on rays fin-rays. originating Caudal-fin the and parhypural elements. hypural beyond the continental shelf. Pelvic fins. Paired fins usually located ventrally on the body; the Procurrent fin-rays. Small dorsal and ventral rays of caudal Ougular), the in various head position to below the beneath from fin caudal located supported not and rays principal the to anterior by elements. hypural/parhypural region gut high on pectoral region (thoracic), to the (abdominal); fin pelvic formation, stomioids; to prior some in sides the ray length measured from the base is the of (finfold) to the blade point the Near Proximal. attachment of or to opposite origin; distal margin greatest distal. of the blade; after ray formation pelvic fin of insertion of the longest ray to is measured from point the length Pterotic spines. bone from the the pterotic Spines ray. the of tip emerging located of region temporal the in orbit the of region upper the to posterior the skull. surrounding Periblast. Thin lying below the embryo and membrane surrounding the yolk is invaded the yolk in teleosts; the space by of a syncytial nuclei, forming region unknown function. fin the form bony elements that or Cartilaginous Pterygiophores. of a fish. and support the fin rays base associated body or the of Region Peritoneal. the gut the with synonymous peritoneum; perivisceral. the with of often membrane melanophore. Punctate melanophore. Round or dot-like 42

55 Querimana Silvery pelagic juvenile stage of mugilids. stage. to snout the of tip the from distance The (SL). length Standard of plate. hypural the the posterior vertical margin fin and Rays. paired bilaterally usually are that supports Segmented melanophore. Stellate the melanophore; or dendritic A star-shaped branched. often condition can be temporary and stellate melanophore can the punctate after become contraction. Net-like Reticulated. or web-like in appearance. in Striations. Surface features form the of lines or bands. backward; Angled Retrorse. referring direction to usually of a spine. surface lateral the on the of band A horizontal Stripe. pigment of or a series. head sometimes body; occurring in of stage juvenile holocentrids, Pelagic Rhynchichthys stage. of opercular, preopercular, rostral, presence the by characterized skin. beneath Occurring the Subcutaneous. silvery and spines supraoccipital and color. below lying cover gill of bone Posterior Subopercle. the the in a spine ending Rostrum. A prolongation of the snout, sometimes opercle. of edge anterior from measured length rostrum spine); (rostral the to tip (holocentrids). spine or rostrum the of the jaw the upper back Subterminal mouth. Underneath or set from the tip of the an to referred sometimes snout, inferior mouth. as bone describe to used often appearance; a wrinkled Having Rugose. with the describe to used sometimes surface; textured a highly upper Superior. Spatial anatomical position; term meaning in numerous by folds. mucosal caused of the gut, appearance inferior. to opposite straddles of that pigment and dorsum the Saddle. A broad band the A spine from spine. Supracieithral originating supracleithrum posterior margin near located the upper body. the of head. the of extends ventrad on each side with shell egg an to Referring Sculptured. or ornamentation the Supraoccipital of the of midline back on crest or Spine spine. textures. of various shapes and surface features the the head originating from supraoccipital bone. Scutatus characterized antennariids, of stage juvenile Pelagic stage. or Supraorbital spine. One eye the above a crest on spines more from posteriorly extending cranium. the plates by bony in some fishes. opposite divided; or Particulate Syncytium. Segmented. multinucleate mass A in of protoplasm. homogeneous of referring to egg yolk. Tail. Portion the of the postanal region. anus; the posterior body to Shell. The membrane that encloses egg; generally, equivalent to an Type eye. of elongate, cylindrical eye that protrudes Telescopic chorion. of skin. or upward within an envelope forward Snout. Forward part of the head anterior to the eye; snout length in the axis from the anteriormost pigmented longitudinal measured Specialized stage. stage following developmental benthic Tenuis the snout. of the to eye the of tip larval a small the by head, thin characterized carapids; in stage surface a vexillum. of lack and body, from Distance to the snout of tip (Sn-A). Snout-anus the the at of the longitudinal axis; the anus, measured margin posterior typical that opens anteriorly; of of Type Terminal mouth mouth. most fish larvae. (PAL). equivalent to preanal length Spatulate. spoon-shaped. or Flattened stage. Tholichthys of stage juvenile chaetodontids, Pelagic characterized by broad, enlarged preopercular spines, supracleithral the Bone upper part of at the skull, often forming part Sphenotic. and posttemporal pigmentation. silvery often and plates, orbit. the of of the chest the Referring area; usually the region Thoracic. to pectoral Spines. of fin below a fish the unsegmented, are that fins the in elements Supporting base. body sharp and unbranched, unpaired, (sometimes and stiff usually referred to refers also, projections pointed rays); spinous as to the the of tip anterior fish from Measurement (TL). length Total arising from various bones, usually on the head. the finfold fin caudal or caudal the including part, posterior most to rays. scales Spinous spines (not the with larval Specialized scale. scale serrated ctenoid fishes). adult of Transformation. with metamorphosis) (synonymous process The in stage, characterized by a marked larval change the of end at the a stalk peduncle. Stalked eye. Eye borne on or form or structure and involving acquisition adult of juvenile or 43

56 to characters characters; also refers larval the stage of loss and Ventrum. Lowermost part ventral opposite margin; body body; of to dorsum. "transitional" term sometimes is used process occurs; the this where undergo (e.g., that larvae transformation scom- for in a gradual brids). blood ofthe vessel. One or blood Vertical vertically oriented more in vessels leptocephali. connecting kidney or gut the with aorta the of between the head and Trunk. Portion the the anus. body VexiIlifer stage Larval stage. carapids of the by characterized tentacle-like dorsal fin ray. Tubercle. protuberance. knobby Small elongate modified Highly VexiIlum. in larval dorsal-finray anterior in region which A median bone the the Urohyal. throat to carapids. of arch. hyoid the attach; sternohyoid muscles not part Viviparous. Type where develop embryos of reproduction the structure Urostyle. Complex bony (usually upturned) at the termi- nutrition. maternal receive and ovary the within fusion from of several ver- formed column vertebral the of the nus caudal the attached to it. of fin are supporting tebrae; elements the Vomer. Elongate palate. that forms the roof of the bone of a fish where ventral the Opening Vent. the alimentary on surface cells. pigment Xanthophores. Yellow anus the to equivalent essentially canal urinary and larval in open; fin. a sac-like mass (yolk sac) egg or in of the Nutritive material Yolk. the below region abdominal larva. of a newly hatched to opposite body; of dorsal. part Lower Ventral. Yolk-sac larva. Early larval stage yolk present in a sac-like with fins. fins. Ventral Pelvic of gut. the region of Format Explanation Methods and Introductions to Orders Atlantic (see Bigelow 1963), Smith and western North Masuda (1984), al. et (1986), Heemstra Moser et al. this The generally guide descriptive accounts in Whitehead et (1984), al. (1984,1986), Paxton and of stages used follow formats guides in for ELH the (1994), et al. (1995), and individual Fischer Eschmeyer Northwest of fishes Atlantic the (Fahay the 1983) and is followed this Usually, revisions. systematic brief by Orders and 1989). Northeast Pacific (Matarese et al. in the kinds of fishes found the of characterizations subordinal sequentially presented are subfamily to taxa and, their in some cases, order reproductive modes and Eschmeyer of the classification as they appear in ELH stages. Finally, the representatives order of in the Eschmeyer's Likewise, nomenclature follows (1990). of the a list with along are area CalCOFI the indicated classification (1990) and generic listings. Within in the guide. treated families families or subfamilies, the genera and species are listed alphabetically. Occasionally, the classification or order the for Where is scanty information ELH guide from differ this in sequence phylogenetic and known descriptions brief its families, of of certain for (1990) recent by warranted when are section, this in given sometimes stages larval that in Eschmeyer 1993; Johnson (e.g., 1994). Nelson studies For stages. larval single of illustrations by accompanied some orders (e.g., Lophiiformes) containing numerous and an most orders For is there suborders, some families with few representatives in the CalCOFI area, section briefly summarizes introductory its system- that all the is area in taxa for meristics of a summary table status, atic salient defining characteristics, taxonomic Tables of containing this section. at included the end literature. components, and lists pertinent These of information needed to interpret descriptive accounts were most recent available the on based summaries the section. this in included be may order in families the literature, primarily Nelson (1994), papers in photophores of diagrams For example, definitions and on phylogeny a symposium of percomorph proceedings Stomiiformes. to introduction in the listed are a few In the of Fishes in volumes 1993), Johnson (see the family cases, where an order is represented by a single 44

57 in Batrachoidifonnes), (e.g., CalCOFlarea the general shape, (e.g., planktonic or demersal, size, features order the on infonnation is in the family included of of the eggs the color, identifying characteristics) Figures introduction. tables section, and' in and this Rosen family. Breder (1966), Moser et and (1984b), al. throughout the sequentially guide, within are numbered al. sources were (1989) for major et Matarese and each Table section Figure 1 or Introduction (e.g., and eggs. Next, on infonnation reproduction general 2). Anguillifonnes are summarized, such features larval at hatch- size as size, maximum ing, general morphology (shape of head length relative shape and body, and and of gut, shape Introductions to Families pigment myomere counts, fins), of pattern, location This section provides basic infonnation on the stage-specific unique development (if warranted), of of the families each 152 ELH stages and adults some in and, and ecology cases, specializations, intended is to infonnation included This guide. the in similar fami- to those in other If larvae are behavior. orient help users interpreta- of the guide and aid in the be may comparisons brief lies, to aid in identifi- given tion species. accounts descriptive the of individual of cation. Comparisons species within the family in the of complete or be These not intended to are introductions CalCOFI area are provided where larvae to are likely reviews exhaustive family, the of serve rather they as limited important to are be confused; these usually the a starting and in some point for users of guide since larvae, similar very with species diagnostic The cases to suggest future research on that family. If in presented are descriptions. species the features on infonnation brief some with begins introduction the diag- necessary, more comprehensive comparisons of of and taxonomic defmition, composition, distribution or more for are one features nostic presented in stages section. family. this This in infonnation is based largely on Nelson the fonn tabular cited sources other the and the preceding in (1994) adult and ELH the of and specimens overview an is Next section. A final paragraph documents the species the occurring in the CalCOFI area. of stages descriptions are species the which on sources literature significant For in the families with representation size with of measured, specimens The numbers based. meristic area, CalCOFI characters adults of these of are given for each species for ranges each stage, species the at table accompanying an in listed end are are in families where two or more species described; Other is given in a table sources for species lists and introduction. the usually infonnation this described, section. the of at are adults the on infonnation in the region end the the following meristic table detailed this table of specimens and in ranges size Numbers are these primarily, cited; and Lavenberg (1968, Fitch represent and the in study measured specimens only al. (1972), Miller and Lea 1975), Hubbs 1971, et those not sources in used from the al. et Eschmeyer (1979), al. et Thomson (1979), literature. Literature descriptions (1983), al. (1991), Allen and Robertson the et are Robins in foot- or paragraph the in cited al. (1995). Comprehensive et and (1994), Fischer and in the Tables table. the (including figures notes species are numbered sequentially illustration plates) interannual and seasonal the on distribution infonnation within 1 or Muraenidae Table (e.g., family each Figure is CalCOFI of larvae in the time series and abundance 2). Ophichthidae in CalCOFI Atlases 31 available al. et (Moser 32 and are atlases These 1994a). 1993, of much for basis the Species Descriptions the given on larval distribution throughout infonnation but, to avoid cited. are not A the guide repetition, proportion large A (ca. 23%) of early life the second paragraph briefly describes the adult morpholo- general new to infonna- species described in this guide of giving family, the of are stages and gy ecology tion size, shape, identification The science. accom- color, was habitat, stages of these fm arrangement, scales, based is series plished primarily by establishing ontogenetic commercial This and summary importance. a series was Once sources cited above. on the literature morphology on pigment. based and it was through several means. established, identified repro- of fins, vertebrae, or (myomeres structures Countable Next overview an paragraphs are giving of of the family. Where the ELH stages duction, and counts scales, gill rakers, etc.) were matched with infonnation on spawning, mode available, general transfonnation of possible species. For some species, along with reproduction, and fecundity are given series exhibiting both larval and juvenile characters 45

58 (meristics, and pigment) were available. morphology gath- was tables these for Information available). (if Sometimes a juvenile specimen with remnants a of to introduction in the listed sources literature from ered provided the identifica- larval pigment pattern unique The from the family and original observations. litera- was method for Another identifying ELH series tion. on made counts by primarily augmented ture was experimen- SWFSC to in the series developmental rear specimens the from Vertebrates Marine SIO Collection. facilities. Developmental aquarium at other and tal 10 more species each of specimens or were Usually field-collected series eggs from artificially spawned and if available. Counts of vertebrae and median examined radiographs from taken were fins prepared in our to identifiable were grown juveniles. em- Methods struc- other fins paired of and and counts laboratory ployed been have described studies aquarium these in Basally specimens. the from taken were tures directly al. (Moser elsewhere and Butler 1981, 1987; Butler et rays anal or posteriormost divided counted were dorsal 1983; 1982; Stevens and Moser 1982; Ambrose et al. range and highest the represents The spine. a single as Moser al. 1983; Hunter 1984; Ambrose and Moser et lowest counts obtained from all sources. Generally, the 1988; al. et Stevens 1989). modal our counts represent original observations since given they are not often in the literature. For wide- Descriptions on presented are species each two for worldwide with those or species ranging distributions the pages facing with written description on the left stages of developmental on counts the for ranges usually represent the extreme the page illustrations the and the right page. representative and the modal counts are species family page, left the On the of name a CalCOFI the from specimens If nominal area. located and comer left upper the in is font) case (upper the CalCOFI in region worldwide counts has species comer. For in right the upper is name species the the of species from in other the localities, counts are disjunct that the only simplicity, binomial portion from the only represent modes and ranges population is given name with describ- name species full the here; cells table the indicate the in CalCOFI Blank region. at the of taxa in the list date be found may er and right the On guide. the to introduction the of beginning counts that information was lacking the literature or in page, the species name is in the upper right comer and to information insufficient was there If made. not were comer. the common name is in the upper left Nomen- given. count, a modal one was not determine Counts were stages ELH of to critical identification the for clature genera and families follows Eschmeyer indicates table the "0" in obtained A cases. all in nearly (1990) with consistent is nomenclature species and the meristic is, that category; that for zero of a count catalogue of prepared world the of species fish being elements in the species for structure absent are consid- for N. W. Nomenclature comm.). Eschmeyer (pers. by ered. common names follows and Lea (1972), Hubbs Miller Robins (1983), aL and et et al. (1979), Eschmeyer et categories Three to vertebral common sources of Other supply (1991). aL counts used given: are Robert- and Allen Precaudal aL (1979), were Thomson et names vertebrae. total and caudal, precaudal, the (abdominal) vertebrae include all centra anterior to Where different (1995). et Fischer and (1994), son al. spine; verte- haemal first the with centrum caudal the were in given species same the for names common haemal centrum the with include brae the first spine frequently most we selected these publications, the urostyle it, including the posterior to those all and name this guide. for used vertebra). The as of precaudal sum a single (counted not may the total counts vertebrae caudal and equal of two for The format the description page consists a larger counts total represent usually count, since ELH life history, meristics, on sections columns; caudal precaudal for samples and than sample verte- original illustrations are of literature, pattern, and a list in and given caudal rarely are counts Precaudal brae. the the left and in description column early of life counts were these made of most literature the and the history stages and a morphometric table are in right of 10 this usually guide, or from preparation the during explana- and sections these of The contents column. fewer for In specimens. radiographed counts fins, the below. discussed are them to pertaining tions rays are given as roman numerals and soft rays spinous both Where numerals. arabic indicated are spinous by Meristics. counts and ranges list table This modal rays and fin, same the in occur rays soft are they fin (usual count, not statistical mode) for vertebrae, are fins Multiple a comma. by separated by separated rays (if available), and branchiostegal rays, gill rakers 46

59 a appropriate, Where "+". principal and procurrent ray activity popula- the in parent spawning of reflective given for the are counts caudal superior fin; and tions. This information was obtained primarily from "+". principal a by inferior separated are counts ray 1994a); 1993, (Moseretal. 32 CaiCOFIAtlases31 and (inferior) Counts for upper (superior) and lower was obtained from Matarese supplemental information rays some procurrent In separately. listed cases are et Doyle (1992). and al. et Walker (1987), al. (1987), given. Gill is rakers (e.g., count total the only gadoids) pattern, mode gives ELH section, of this part last The limb the given upper and lower are for first (e.g., reproduction of oviparous), gill the egg habitat (e.g., of the the count for the first habitat the and demersal), arch; total planktonic, arch given is when of the or larvae and lower limbs. The from came information This not was juvenile data special for available upper stage. above cited literature from sum original observations. and upper the lower limbs of the gill rakers on and number the given may not the total reason for equal fishes a gill raker In counts. vertebral above with for are here Listed Literature. publications contain that at the between the upper and the lower angle limb, the descriptions and/or illustrations ELH of this of stages the is lower included in the count for at the raker angle aspects other species. Literature pertaining to this of is the to introduction cited in the limb. usually species family. on range, geographic Life information Basic History. Original previously Illustrations. Original, unpublished ELH habitat, is given in and season, spawning pattern are illustrations with mm, in size and stage by listed literature on this section. Geographic ranges are based of speci- illustrator parentheses. in acknowledged the on original data from and localities capture mens Verte- Marine SIO the (primarily museums in History Life section gives Early This Description. north-south the Usually Collection). brates extreme of wide- the aid to intended information detailed identifica- in ranges are given distributions for shorefishes; to generalized be may species ranging zoogeographic tion under listed egg of the ELH stages of the species, or masses. temperate) (e.g., provinces water tropical, stages. These and accounts are and larval stage juvenile in Figures Major geographic are shown landmarks on previous descriptions (see literature based citations geographic Spellings of 3. localities Introduction 2 and in and on original observations. left column) south the listed those follow border U.S. the of in information Sailing known, If the and Mexico of Coasts West the for eggs of the species are Directions diameter, yolk Defense is Map- U.S. given in a format that includes shell Central by the published America and diameter characteristics, number of oil globules, ping Agency (DMA 1991). shell surface, (chorion) and the of pigmentation, nature habitat The includes usually description brief are features. diagnostic given mm <1.0 Measurements mm >1.0 those and mm 0.01 mm. 0.1 to given are zone (e.g., epipelagic, mesopelag- depth on information to homogenous) yolk segmented, (e.g., of the nature The and coastal life zones (e.g., species oceanic for ic) on with the yolk given, diameter when available. is along type demersal continental shelf, slope), bottom (for a relatively In species with the space, perivitelline wide species), recorded maximum depths for and shore- is information it can so listed is mass yolk the of litera- the usually largely based that on width fishes. This (1972), Lea and Miller primarily width to give ture, be compared with et shell Eschmeyer al. of estimate an perivitelline the and other sources cited in the introduction to width the of (1983), diame- of A range space. each ters season spawning on Information family. oil with given along globules a multiple for of some is their arrangement, when available. from CalCOFI of area is available sources species description the in Lavenberg (1968, 1971, and Fitch and as inner the of ornamentations and features Surface 1975), such et al. (1989), Love if a with visible al. (1974), Matarese shell et described, are surface Feder outer from al. (1992) and dissecting (1991), Doyle (1992), and Leet and sequence microscope. The et arrangement embryo, yolk, fishes, such as rockfishes of the on melanophores of development of studies on groups specific Love et briefly are globule oil and (e.g., on Information Phillips described. Wyllie Echeverria 1987; 1964; of of the chorion, yolk, or oil globule(s) al. is larvae of occurrence of Seasonality 1990). color the proportion included since a large of refers preservative. in color the to on captured larvae eggs pelagic The and are ichthyoplankton surveys are recently spawned egg, the of provides diagnostic features section final 47

60 may that one or a combination of the charac- include conversely, melanophores sometimes and, descriptions given above ters features. unique other or described are that were not present specimen on the illustrated, illustration. or could not be portrayed in the larval and juvenile stages includes on The section are The descriptions and illustrations intended to be milestones developmental at ranges size of estimates are melanistic Major supplementary. mutually features (hatching, flexion, transformation), the sequence fin of when appear in a specific stage and they described formation, pigment patterns, and a summary of diag- stages only generally are in described when subsequent nostic features these Smallest and stages. largest of their arrangement changes or they are significantly sizes at ranges the represent milestones developmental The augmented or depleted. description of pigment stages these at sizes of or literature the from obtained because emphasized is pattern in of importance its in overlap be may There from original observations. larvae at all stages and at identifying taxonomic all size ranges for any milestones; these of developmental (Russell levels 1976). the descriptions, these In words larva be for example, the largest known preflexion may "melanophore" since "pigment" usually are left or out larger largest and larva flexion smallest the the than the are they that it is understood subject the of majority smallest postflex- be than larger the flexion larva may of descriptive phrases and their constant repetition larva. ion be would inefficient. Words indicating specific types ranges of In and initiation for such size cases, completion for pigment, of for Criteria given. are flexion melanistic such as "series" (equivalent to the determining beginning and end melanophores), "row") or "patches" (of individual of each stage are explained introductory section on general "blotches", in the "dashes" are appropriate. or when indicated defined and size principles in the glossary. Specimen descriptions the space, conserve To in presented are throughout in this as the and section given is guide abbreviated, with verbs and style semi-telegraphic "body length (BL); whether the indicated body length" articles included only when needed for clarity. Num- is "notochord (NL) length" the bers descriptions refer to in number the length" melano- (SL) "standard or of be by referring to size-at-stage data ascertained can phores in the or area in described. the being series this section. given in for conventions Other used to conserve space, are example, abbreviation "ID" (the for the first dorsal fm) to portion spinous the indicate may used be is based first the on of formation fin the of sequence The though the spinous and soft-rayed even fin, appearance dorsal of support elements or fin rays for each fin portions determined most accurately from examination fin, completely or continuous are fin the of of not separate. cleared series. Observations developmental and stained original either on cleared and stained material, made or character- presents section features diagnostic The when the from literature, were used available; other- transformation larvae, separate help will that istics material wise, observations on unstained formed the or specimens, juveniles early for basis sequence estimating all of a species from of fm development. The in others given family same the sequence is in a formula with the abbreviations or Usually, a combi- genus. nation of but is needed to accomplish this characters separated fms appearing successively for by a comma fins and simultaneously appearing united Often, sometimes a single is diagnostic. character total amper- an by myomere myomere counts and/or preanal and postanal sand cases In ("&"). where fin development appeared given because are counts to be simultaneous impor- of sizes their were paramount appropriate of larvae but in tance identification. confirm fms to were this, unavailable shown as the simultaneously forming. Comprehensive Morphometries. infor- morphometric and higher is important in separating species mation Descriptions for of melanophore patterns are given morphometrics Stage-specific taxa. for transforming and stage and larval each early are of key features intention is to The when stages juvenile appropriate. a quantita- is in given The table. a fixed-format table description a broad-brush give of components main the tive portrayal of basic form; this is especially important describe every and no attempt the is made to pattern of in shown usually not are that structures demonstrating the patterns melanophore; however, general of melanis- morpho- in illustrations (e.g., head width). Also, the given. illustra- are always tic In some pigment cases, metric tables in this guide may prove useful in compar- studies ative hydrodynamic and ecological tions include melanophores that are not in these of a broad 48

61 array dis- snout-anus of Proportions taxa. teleost of necessarily proportionally) with specimen size to head length, and pectoral length, tance, depth, body fin is not permit more illustrations to be included. Detail length presented fin are pelvic percentage body of as of the earlier the illustration reducing by lost size diameter length, snout and length and eye head width, usually struc- in complex less stages because are they avail- as length. head of percentage When are given with ture pigment pattern compared and stages. later for mean are and given values each the range able, of flexion preflexion, these and yolk-sac, for features of a with produced were illustrations the Most and larvae postflexion specimens transformation for a dissecting lucida attached to microscope, thus camera and pelagic as specified (sometimes juveniles or proportions accurate assuring placement. pigment and of proportions Where juveniles). benthic appropriate, pencil Initial and then finished drawings were made in anus, given body depth at (e.g., other features key are and The line-and-stipple technique. using india ink stages the where In base). fin pectoral of cases depth during this was specimen process. constantly checked (e.g., applicable not are notochord on based flexion in style, with body are Illustrations semi-diagrammatic for calculated were means and ranges eels, macrourids) shown outline and major surface features with solid or For stage categories or for size intervals. other some pigment stippling by indicated and lines dashed line. or the taken table for information species, this from was is lightly stippled. No attempt Internal pigmentation from either literature, directly or after calculation was stippling to show depth perspective with made measurements; however, for most species, published with because interferes portrayal the that pigment of original represents information observations. the appropriate, Where pattern. enlarged sections, or dorsal to nearest made the 1.0 mm > were Measurements to of specimens are illustrated in aid views or ventral mm and those <1.0 0.1 were made to the nearest mm identification. sources Literature mm). 0.01 to (estimated mm 0.02 for used specimens and are original measurements specimens and stage by The list captions figure the in family, the to introduction the in given either of followed size, localities the by parentheses in a table. descriptions morphometric Published in or text for capture illustrations or original by the literature all include features the of and on data always not do illustrations. citations Where for published previously is same the this guide. Generally, used included only stages in we locality literature citation or collection the or the illustrations, sequential more or two for locality the did and descriptions these from available data not fill to specimens additional measure out missing given the series. The cruise of end the is at citation the table of in the (0%) A zero table. indicates sections number station are given for specimens from and other and of the feature and a blank indicates the CalCOFI CalCOFI For cruises. survey absence the measured. not was feature maps of specimens, be found can on the these localities Figures Introduction in survey CalCOFI the pattern are given for 1-3. Where no cruise and station number Illustrations. The of illustrations containing plate CalCOFI specimens (i.e., only the attribution selected developmental stages the species is on the of and "CaICOFI" cruise given), is not are data station of each species description, facing the right hand page available. other sources Information on (research an of available, If illustrations egg descriptive account. Table in summarized is and yolk-sac at are one) than more (sometimes larva cruises) expeditions, programs, 1. of the plate with illustrations of preflexion, for given are numbers Catalogue Introduction the top Marine Vertebrates Collection, specimens the from and stages juvenile transformation, postflexion, flexion, (SIO), of Oceanography of Section Institution Scripps has a label indicat- in sequence below. Each illustration Los of Museum History Natural Fishes, Angeles of (diameter body eggs, for the specimen size the ing museum collections other and (LACM), County fish juve- length for larvae, transforming specimens, and et Leviton to according (acronyms 1985). al. niles). (not Usually, the size of the illustration increases 49

62 How Guide this with Fishes of Stages Early Identify to of fishes is of Identification the stages fin rays are important. With this information one early should by many reasons. First, their size presents order to specimen the identify for to able challenging be with working in experienced not those for a problem the matching the characteristics of the with specimen dissecting solves small good microscope specimens. A 2. characters in Introduction Table ordinal summarized of part this problem but it may take time considerable determine the family, one compare the total should To skills to develop the manipulative and observational count myomere of range the total with specimen the of for the California in present families counts vertebral necessary Another work with material of to this size. problem Table has larva the If 3). diversity (Introduction region Current is the the large variety of species and and counts be can fins anal and dorsal for ray the fins, of sizes, shapes, pigmentation one encounter may the in region families sample. Workers familiar with in ichthyoplankton for ranges an the with compared 4 and Tables (Introduction overall the Next, shape 5). identify organisms are to dichotomous using keys in compared be should structures major of and the shape to applicable frustrated these are not generally find with identifying ontogenetic stages. This is true because the those on the outline illustrations of typical mid- of each family in the CalCOFI area larvae stage any nature individual character of usually over changes (Figure This procedure should identify Introduction 7). unique have species few and development of the course possible of the specimen, or at least several the family consistently that are characters throughout recognizable further The identification can then carried families. be early are keys Also, inappropri- history. entire the life by consulting the information in the introductory ate portions large because of the ichthyoplankton of a accounts order the is guide this in informa- Our of this If family(ies). and intention region may be unknown. tion is consistent with the identification, preliminary summarize to gained over years knowledge studying of of meristics may the narrow for family the table the the stages ELH of fishes California the Current in one species possible before identification consults the as novices by identification their to aid an as region individual accounts for illustrations descriptive and weB experts. as This species. identification should procedure permit of of history life early general the A knowledge for the CalCOFI region, of most larvae descrip- which patterns interpreting of for essential marine fishes is exist. tions guide this and gaining competency in identification of eggs, juveniles. A summary of basic fish larvae, and eggs presents a of more difficult Identification this information is presented in the on Princi- section far fewer are they since for challenge species known and ples In the Explanation Terminology. Format of examination initial the During larvae. with compared of each and structure the Methods and components characteristics of a specimen, the following structural section of the along described are guide the with noted: be should nature shape; size; egg of shell the to obtain and present the informa- methodology used inner surface; color of the shell; presence or of absence contained tion them. in with Familiarity will this aid nature the space; perivitelline the of width membranes; the of interpretation and use the in guide. greatly segmenta- of the yolk and, segmented, the type if of the and arrangement and location, size, tion; number, one examining a from gains impression first The be of that oil globules. characters the Among should sometimes and important is larva permit may fish at eggs note in late-stage are: state of development immediate identification. Its overall shape (also size, the of development eyes, of state closure; blastopore eyes, gut), of head, shape body, and develop- of stage mouth, fins, finfold, gut, myomeres and late-stage of pigment pattern, and myomere count are basic ment, of sequence the Also, pigment embryos. appearance of noted. features that should be present, If fins are their on important yolk, embryo and globule, oil the are and number the and location, structure, and shape features identification. be can characters These for them within rays soft and spinous of are arrangement egg of those compared with in the ordinal summary characters. of The presence important early-forming brief and 2) Table (Introduction the with characters identifying important is an fins ("precocious") charac- and orders to introductions the in families. summaries head such features prominent other Also, teristic. as be can identification family, the to down narrowed If or elongate markedly and barbels, spines, ornamented 50

63 Miller a of the species accounts may reveal its browsing in descriptions and keys the using by identified Martin and Lea (1972) and Eschmeyer et al. (1983). identity. series), the in volumes other and (1978; Drewry and Other sources may be helpful in confirming egg Ozawa al. (l984b), Leis et Moser (1983), Rennis and with the aid larval and identifications made Leis (l988a), (1989) Tmski and and Okiyama (l986g), of this much et Matarese guide. identification identification on information present of (1989) includes life aL early the they that CalCOFI species the of of species stages, although most from the northern boundary history area region. can juveniles Often, guide. this in treated not be CalCOFI the in occur not do cover 51

64 Table Introduction useful for the identification of early stages of fishes, listed by order or suborder. Information 2. Characters al. and Fahay (1983), Leis and Tmski (1989), Matarese et (1976), (1989), and from authors of chapters Ahlstrom from Moser guide. in this Elopiformes/ Character Albuliformes Notacanthiformes Anguilliformes Saccopharyngiformes Clupeiformes Undescribed Eggs Undescribed Undescribed P or ~Ianktonic P; rarely D Qemersal Round Shape Round oval to Shell Smooth surface Smooth Segmented Yolk Segmented Wide Perivitelline Narrow to wide space globules 0,1, or more Oil 1 to 0, many Larvae Leptocephalus Body Leptocephalus shape Leptocephalus; Leptocephalus slender Elongate, tail forked 40-95 50-80 Elopiformes, 75-80; >95 length Preanal 48-95 BL) (% AIbuliformes, 90-95 more Straight Straight; looped in some Straight; some with Straight, with I or type Gut Straight loops; rarely trailing loops Round to oval Round to oval Round to oval; Round to oval Eyes Round slightly to choroid tissue in oval some some; telescopic No No No Head spines No No 225-330 97-400+ (most 68-250 39-76 51-82; Elopiformes, Vertebrae 65-92 100-250) AIbuliformes, No No No No Early fins forming No shrinkage & Marked; shrinkage & Marked; shrinkage Marked; shrinkage & growth Marked; D, A, & Marked; Transformation growth & growth growth P migrate 2 No No No No pelagic No Special juveniles Rays Rays rays & Spines Rays Rays fm of elements Type Late Late Late Late Late formation fm Pectoral Late Absent Absent Late Pelvic fin formation Late Abdominal Absent Absent Abdominal Pelvic fin position Abdominal Absent Absent with 7-11; some Var., 10-16; Elopiformes, usually Var., 7-10 formula Pelvic fin AIbuliformes, 9-11 spines 1 fin 1 fin 1 fin 1 fm 1 fin fin Dorsal 1 fin I fin fill 1 1 fin Anal fill 1 fin No No No Adipose fin No No some in absent 0-4; 5-11; Reduced or absent Usually (10+9) 19 19 (10+9) Principal caudal fin absent in some rays 52

65 Table Introduction Continued. 2. Argentinoidei Stomiiformes Salmonoidei Aulopiformes Myctophiformes Character I I Eggs II ~; P P P D P ~Ianktonic or !2emersal Round Round oval Round to Round Round Shape Smooth; pustulate Smooth (most); some Smooth; attachment Smooth (most); disc Smooth Shell surface chorion in many in membrane; some double pattern hexagonal some in synodontids with facets Segmented Homogeneous Segmented or Segmented Segmented Yolk segmented Narrow Narrow Wide to narrow Narrow Narrow Perivitelline space I to many or o I or o I many to o globules Oil Larvae Elongate Var., slender; to & elongate Elongate; some slender Elongate, Var., to elongate shape Body compressed moderately stout moderately stout some stout 40-70 20-80 70-95 30-95 60-80 (% length Preanal BL) saccular in Straight; trailing & Straight; or Straight; folded Straight; var. shapes; Straight Gut type ornamented in some some in some trailing in I species saccular narrow; Round narrow; very Round to very to Round oval, to Round to very narrow; oval Round to Eyes choroid tissue in some stalked in sometimes Idiacanthus choroid tissue in some; stalked in stalked some Usually none None None Usually none None spines Head 45-65) (most 40-85 30-100+ 45-120 28-45 42-74 Vertebrae lchthyococcus in PI D & in some PI PI & P in some No No fms forming 2 Early marked, Marked Marked; photophore Var.; Marked osmerids; in Marked Transformation formation prolonged in prolonged, or delayed gradual in alepocephalids some No Present in some No No No juveniles pelagic Special Rays Rays Rays Rays Rays of fm elements Type • elongate & early Var.; elongate & Var.; most; in early elongate in Late elongate Late; in Late osmerids in fm formation Pectoral in some lchthyococcus some in some Var.; early elongate & late; elongate Late Usually elongate & early Var.; Late in osmerids formation Pelvic fin in some in some some in Abdominal Abdominal Abdominal Abdominal Abdominal fin Pelvic position usually Var., usually 8-10 Var., usually 5-8; up to Var., 7-11 Var., usually 8-12 5-11 formula fin Pelvic in Bathophilus 26 rays in I fin; elongate I fin I fm I fm I fin fm Dorsal some I fin I fm I fm I fin I fin Anal fin present Usually present present Usually Usually present Often osmerids in Present fin Adipose (10+9) 19 19 (10+9) 19 19 (10+9) (10+9) 19 (10+9) rays fm caudal Principal 53

66 Table Introduction Continued. 2. Gadiformes Ophidiiformes Lophiiformes Character Batl'achoidifol'mes Gobiesociformes Eggs P; rarely D P; some live-bearers rafts nests in some P; often ~lanktonic or Qemersal D; in D to slightly oval Round Round to slightly oval Round Round Shape var. shapes Round or Smooth smooth; Usually Smooth Smooth Shell surface Smooth pattern in hexagonal some macrurids Homogeneous Homogeneous Homogeneous Yolk rarely Homogeneous; Homogeneous segmented space Perivitelline Narrow Narrow Narrow Narrow Narrow globules 0, 1, or mOTe o or I O-several Oil or J o J many to Larvae Var., Elongate elongate to Stout Globular shape Body to Moderate stout moderately stout 33-55 Usually <50 BL) length 38-44 30-90; usually >75 Preanal (% 50-85 Usually coiled Coiled Coiled Gut Coiled, voluminous type Initially later straight, coiled Round Round Round Round Eyes Round to oval Usually none Opercular Opercnlar spines in some spines spines Head Rarely o J opercular spine or 44-66 18-31 most; 80-116 40-150 30-58 in Vertebrae 21-54 in macrourids some; yolk-sac in fonn PI in Fins vexillum in P2 some in Early forming fins D, Often No PI' P, stage carapids Gradual Gradual Marked Gradual Transformation Marked to gradual No No Present in some Present in some juveniles pelagic Special No Rays Rays rays & rays Spines & Spines elements of fin Type Rays Early yolk-sac In some in stage some in Late fonnation fin Pectoral Late in most Late stage yolk-sac In Early in some Late fonnation Pelvic Often absent; early to fin Late late Thoracic Jugular Thoracic or jugular Jugular position Pelvic fin Thoracic VaL, 1,2-3 3-5 2-8 0-2 I, o or fin Pelvic I, 4 or 1, 5 fonnula an fin I to 3 fins I fin 2 fins 2 fins; anterior fin Dorsal I fin iIlicium 1 fin I or 2 fins 1 fin fin 1 Anal fin I fin No No No No fin Adipose No 8-10 12-14 0-14 8-70; Total rays rays fin caudal Principal 8-14 lacking in some 54

67 Table 2. Introduction Continued. Cyprinodontiformes Atheriniformes Beloniformes Character Lampridiformes Beryciformes Eggs D D; many live-bearers Var., P or D P ~lanktonic or 12emersai P Round Round oval Round to Round Round Shape or ornamented; many Usually with Smooth with Smooth or Usually smooth; some surface Shell Smooth with filaments or spioes filaments spinules with filaments or other ornamentation Homogeneous Homogeneous Homogeneous Homogeneous Homogeneous Yolk Narrow Narrow Narrow Narrow space Narrow Perivitelline Many some 0; Usually with many to many o Usually globules Oil minute many; often Larvae to elongate Moderately stout Moderately elongate Moderately Usually elongate & Slender to stout shape Body elongate compressed 30-{50 65-80 47-59 ca. iocreases with 45-90 20-50; Preanal length BL) (% development Straight Coiled Coiled Coiled Coiled type Gut Round Round Round Round Round Eyes No No No No to heavy None spioes Head 24-54 36-97 21-55 33-200 Usually 23-33 Vertebrae C (often before hatchiog) C D anterior more or & often other fms I No formiog fins Early D in P2' some Often P & rays hatchiog) (before 2 Gradual Gradual Gradual Gradual gradual; Usually marked Transformation io some some Present No No io No Present in some pelagic Special juveniles Rays Spioes rays Rays Rays & rays & Spioes of elements fin Type some io Var.; early Late Late Late Intermediate Pectoral fm formation Late io many Usually early; I or Early Late Late formation Pelvic fm more elongate, usually heavily ornamented Abdomioal Abdominal thoracic Abdominal thoracic, Abdomioal to thoracic Var., to fin position Pelvic abdominal, or jugular 4-7, usually 6 6 0-17 Var.; usually I, 6 or 1,5 Pelvic formula fm 1,5 1,7; not more 1 or I fin I fm; 1 fin or 2 fins 1 2 fins Dorsal fin anterior rays elongate & highly ornamented Oorlfin 1 fin I fm spines fm,O-1 1 fin, I usually 1-4 spines Aual fm No No No No No Adipose fm 3-32 11-18 15 19 (10+9) caudal fin rays Principal 17 55

68 Table Introduction 2. Continued. Scorpaeniformes Perciformes Syngnathiformes Pleuronectiformes Character Tetraodontiformes Eggs D P; some in rafts; rarely P; D or some brooded D; D or P or D or ~lanktonic Qemersal P live-bearers some or slightly oval Var., round in most Round some round Var., Round Round Shape Usually smooth smooth Usually Smooth Smooth surface Smooth or sculptured Shell Homogeneous Homogeneous; rarely Homogeneous; rarely Homogeneous Homogeneous Yolk segmented; secondarily secondarily segmented granular in blennioids Narrow Narrow Usually narrow; rarely Narrow to moderate Narrow Perivitelline space wide to 1, 0, o to many many 1, or more 0, many o to Many Oil globules Larvae moderately Var.; elongate Var.; Var.; elongate often Var.; stout markedly usually Stout to globular shape Body compressed to stout 35-60 20-80 Usually <40 40-90 45-90 BL) (% length Preanal Coiled; distended in Usually Coiled Var.; coiled in most straight Coiled; voluminous type Gut some Usually round; Round some Round Round Round Eyes with narrow; some tissue choroid heavy Usually present; heavy Var.; none to to heavy; often on None Present heavy in often; Rarely present Head spines in some some bony plates 20-100+ 24-28 ; many 25-65 25-65 16-30 19-87 Vertebrae Drays, Often; anterior 2 I or in some; 1st D more None; None large PI can be fins forming Usually none; D spine, PI Early P, & or 3 P, rays spine, spine rays some rays in eye Usually gradual Marked; migrates Gradual Gradual Gradual Transformation larval prolonged Often; e.g., scorpaenids Present in some Rarely; No Rarely pelagic Special juveniles stage in many Spines family rays Spines & rays Rays; spines in i & rays & or rays Spines rays Spines & elements of fin Type only Late early late; not Var.; in late Var.; not in Early some Late formation fin Pectoral some some in Early late Often absent; Intermediate Var., early in some late absent; Often fin Pelvic formation Thoracic jugular to Usually thoracic; some Thoracic Thoracic Abdominal Pelvic fin position or jugular abdominal 5/5, 6/6, e.g., Var., 0-1,5 0/4 fewer I, 5 or 0-6 fewer or 1,5 fin formula Pelvic I fin I or 2 fins 2 fins I or 2 fins; 3 in I or 2 fins I or Dorsal fin tripterygiids usually I fin, fin I spines 1-3 0-3 spines I fin I fin, I fin, spine 0-1 Anal fin No No No No No fin Adipose total 9-12 10-23 Var., usually Usually 17 (9+8); fewer <17 Var., 0-15 Principal caudal fin rays in some rays 56

69 Table Introduction of total vertebral counts for fish families in the CalCOFI region. Only species found in the 3. Ranges included Carapidae the ranges. Vertebral counts for Macrouridae and are are given rarely in the literature and are region in in this listed Counts not table. Counts rings given instead of body for Syngnathidae. are furnished by authors of of vertebrae in this guide. chapters Ostraciidae 24-29 Labridae 18 Molidae 24-36 Gobiesocidae 17-19 25 Tetraodontidae Kuhliidae 17-19 Balistidae 25 Scaridae 18-19 Ogcocephalidae 25-26 Polyprionidae 18-19 Chaunacidae 25-27 Kyphosidae 19 Lophiidae 25-27 Triglidae 19 Antennariidae 25-28 Eleotridae 19 Linophrynidae 25-29 Anoplogastridae 19-20 Oneirodidae 25-30 ranoscopidae U 19-21 25-48 Melanocetidae Cottidae 20 26 Ceratiidae Cirrhitidae 20 26 Haemulidae Caulophrynidae 20 26 Diodontidae Howellidae 20-23 Callionymidae 26 Pomacentridae 21 26 Centrophrynidae Xiphiidae 21 26-27 Monacanthidae Holocentridae 21 26-27 Malacanthidae Thaumatichthyidae 21-22 Gigantactinidae 26-30 Achiridae 21-23 26-38 Gobiidae Acanthuridae 22-23 26-41 Luvaridae Echeneidae 22-23 27-29 Diretmidae Priacanthidae 23 Cyprinodontidae 27-37 Centriscidae 23-24 27-46 Carangidae Myctophidae 23-26 28-30 Melamphaidae Opistognathidae 23-31 29-31 Giganturidae Scorpaenidae 23-31 29-32 Neoscopelidae Apogonidae 24 29-37 Stromateidae Chaetodontidae 24 29-41 Stemoptychidae Ephippidae 24 29-42 Zeidae Gerreidae 24 Blenniidae 29-50 Istiophoridae 24 29-52 Agonidae Lobotidae 24 Gonostomatidae 29-99 Lutjanidae 24 30-34 Coryphaenidae Mugilidae 24 Embiotocidae 30-41 Mullidae 24 30-42 Nomeidae Polynemidae 24 Scombridae 30--66 Pomacanthidae 24 31--61 Gempylidae Nematistiidae 24 Paralichthyidae 32-40 Sparidae 24 32-47 Labrisomidae Sphyraenidae 24 Psychrolutidae 33-35 Pentacerotidae 24-25 33-43 Sciaenidae Tripterygiidae 24-26 33-46 Serranidae Chiasmodontidae 24-26 Rondeletiidae 33-83 Melanostomiidae 24-27 57

70 Table Introduction Continued. 3. Pleuronectidae 34-66 48-54 Bregmacerotidae 49-51 34-83 Cyclopteridae Cetomimidae Opisthoproctidae 49-65 Bythitidae 34-84 Caristiidae 35-40 Stylephoridae 50 Microstomatidae 52-100 35-50 Paralepididae 53-72 35-60 Chaenopsidae Osmeridae Syngnathidae 54-58 35-72 Malacosteidae Hexagrammidae 55-91 36-63 Ophidiidae Bramidae Chauliodontidae 56-62 37-46 Atherinidae 57-74 Ammodytidae 37-54 Alepocephalidae 58-61 Centrolophidae 37-74 Bothidae 61-62 38-41 Scytalinidae Phosichthyidae 61-62 38-47 Zaproridae Batrachoididae Scomberesocidae 62-69 38-48 Belonidae Oreosomatidae 62-99 39-40 Engraulidae 62-111 Trachipteridae 39-47 Exocoetidae Icosteidae 65-70 39-51 Dactyloscopidae Albulidae 68-74 39-52 Bathylagidae 69-74 Stomiidae 39-55 70-80 Cyematidae Barbourisiidae 40-44 71-85 Clupeidae Cryptacanthodidae 40-58 Fistulariidae Platytroctidae 75-86 42-52 Notosudidae Zoarcidae 75-127 42-61 Stichaeidae Elopidae 78-82 43-83 44-47 Trichodontidae 79-80 Anotopteridae 80-108 Pholidae Scopelarchidae 44-55 Moridae 81-85 Idiacanthidae 44-58 Tetragonuridae Trichiuridae 84-158 44-58 Microdesmidae 97-125 Eurypharyngidae 45-56 101-175 Anoplopomatidae Muraenidae 45-66 Lamprididae 105-120 Monognathidae 46 Astronesthidae 114-121 Radiicephalidae 46-58 Cynoglossidae Congridae 116-199 46-61 124-153 Argentinidae Lophotidae 47-49 Alepisauridae Derichthyidae 126-136 47-52 Evermannellidae 143-151 Regalecidae 47-52 144-162 Ophichthidae Leptochilichthyidae 47-52 151-153 Merlucciidae Serrivomeridae 47-55 Saccopharyngidae Mirapinnidae 175-220 42-55 177-203 N emichthyidae Clinidae 47-58 221-251 Synodontidae Anarhichadidae 47-63 Nettastomatidae 223-258 Hemiramphidae 47-75 225-255 N otacanthidae Aulopidae 48 Bathymasteridae 48-51 58

71 Table 4. of total dorsal-fin ray counts (counts for spinous and soft rays combined) for fish families in Introduction Ranges region. given formulas (spinous rays shown in bold type) are CalCOFI after each family, except for families with the Fin included fms rays. Only species found in the region are spinous in the ranges. Macrourids have 2 spinous rays lacking single 8-16 soft rays in the first dorsal fin; counts for the second dorsal fin are not available since they are given rarely in the and include for total rays anterior to the is vertebra. Counts for Echeneidae do not the 31st for range Carapidae literature. The of disc. Counts for fmlets (Scomberesocidae, Carangidae, the Scombridae) are in parentheses. Counts Gempylidae, laminae guide. this of chapters in authors by furnished 12-25 Anotopteridae Hemiramphidae 0 Linophrynidae 12-37 Alepocephalidae 3 13 Luvaridae 3-5 Ceratiidae 13 13-14 Ogcocephalidae Callionymidae 4-6 4+9-10 Gigantactinidae 13-15 Lophiidae 4-7 3+3-4,7-8 13-16 0-11+4-9 Agonidae 4-20 Rondeletiidae 13-17 Chauliodontidae 5-7 Melanocetidae 13-18 Oneirodidae 5-8 Diodontidae 13-28 Gobiesocidae Gobiidae 5-16 2-9+0-3,8-19 Centrophrynidae Chaunacidae 6-7 14 3+11 Thaumatichthyidae 14-15 Aulopidae 6-7 Scopelarchidae 14-16 Sphyraenidae 6-10 5+1,8-10 Stemoptychidae 14-18 Eleotridae 6-11 5-8+1,8-9 Bathylagidae 14-20 F istulariidae 6-14 14-21 Paralepididae Centriscidae 4-8+10-13 7-13 14-22 Tetraodontidae Caulophrynidae 7-13 Microstomatidae 14-34 8-11 Cottidae 5-18+8-30 15-17 8-13 Antennariidae Osmeridae 3+12-14 15-17 Astronesthidae Mullidae 8-21 6-8+1,8 15-18 Ostraciidae Uranoscopidae 0-5,13-16 9 15-20 N otosudidae Albulidae 9-12 9-12+(4-6) 15-20 Platytroctidae Scomberesocidae 9-12 16-17 Cyprinodontidae Apogonidae 9-15 6+1,9-10 Exocoetidae 16-19 Giganturidae 9-15 Mirapinnidae Opisthoproctidae 16-20 9-16 16-20 Stomiidae Gonostomatidae 9-18 Myctophidae Molidae 17-19 9-24 17-20 Argentinidae Anoplogastridae 10-13 17-20 Gerreidae Evermannellidae 10-13 9,8-11 17-40 Synodontidae Echeneidae 10-17 17-48 Syngnathidae Melanostomiidae 10-30 18-19 N eoscopelidae Howellidae 11-13 8+1,9-10 18-43 11-15 Scombridae Leptochilichthyidae 8-27+0-3, 9-10+(4-10) 19 Phosichthyidae Kuhliidae 10,9 11-16 19 Clupeidae Scaridae 11-22 9,10 19-22 Belonidae Barbourisiidae 11-27 9-37+0-2 19-43 Carapidae 11-37 Notacanthidae 20-23 4+1,7-9 Mugilidae 12-14 Labridae 0-2+7-12,9-14 Polynemidae 20-23 12-19 Engraulidae 8+1,11-14 2-3,9-18 20-24 12-21 Melamphaidae Priacanthidae 10,10-14 2-9+1,8-14 20-27 12-24 Atherinidae Elopidae 59

72 Table Introduction Continued. 4. Scorpaenidae 12-17,8-17 20-30 Oreosomatidae 36-43 6-7,30-36 9-12+10-12 Triglidae 20-30 Clinidae 36-51 31-38,5-13 20-32 36-51 1-11,11-27 Serranidae Monacanthidae 2,34-49 39-46 Embiotocidae 20-39 6-18,9-28 Centrolophidae 3,36--43 21-23 Cirrhitidae 10,11-13 41-51 Scytalinidae 11+1-2,9-10 21-23 Polyprionidae 42-47 Bathymasteridae 1-6,37-45 42-50 Xiphiidae 21-25 Malacosteidae 38-45,4-5 Pentacerotidae 21-25 13-15,8-10 Stromateidae 43-52 2-4,41-49 Lutjanidae 10--12,11-15 43-59 Istiophoridae 21-26 36-53,6-7 21-37 Kyphosidae Moridae 9-14,11-27 43-78 4-13+39--67 Opistognathidae 10-11,12-17 22-28 Bregmacerotidae 44-53 1+43-52 Pomacentridae 45-60 Microdesmidae 22-30 11-13,10-17 15--19,28-43 Tripterygiidae 3-4+12-19+7-14 46-57 Merlucciidae 22-37 0--1,10-13+36--44 46--63 22-45 Blenniidae 7-13,11-36 Ammodytidae Stichaeidae 22-78+0-43 Lamprididae 22-78 48-52 11,12-15 Icosteidae Holocentridae 50-56 23-26 Sparidae 12-13,11-13 52-117 23-26 Pleuronectidae Haemulidae Coryphaenidae 10--14,12-18 52-66 23-31 Tetragonuridae 14-21,10-17 Achiridae 53-63 24-38 Carangidae 3-8+ 1,17-39+(0-1) Zaproridae 54-57 24-48 54-57 8-36+0--2, Idiacanthidae 54--66 Gempylidae 24-77 +(0-7) 60-77 15-41 Cryptacanthodidae 73-77 Paralichthyidae Lobotidae 11-13,15-16 60-102 26-29 26-29 7-9+19-20 70-128 Zoarcidae Psychrolutidae Diretmidae 1,25-29 Pholidae 26-30 74-98 74-98 Caristiidae Bothidae 75-102 26-35 3,23-34 79-150 Trichiuridae Balistidae 26-37 3-46,54-128 Labrisomidae 0-4+17-33,0-13 80-114 Cynoglossidae 26--41 Nomeidae 0--12+1-13,15-32 82-116 Monognathidae 26-46 Sciaenidae 7-15+1-2,18-41 26-51 Ophidiidae 85-229 8+1,18-23 Cyematidae Ephippidae 27-32 85-93 Chiasmodontidae 7-14,18-29 Bythitidae 95-115 27-42 Anoplopomatidae 12-30+ 15-21 115-122 Stylephoridae 27-51 Hexagrammidae 15--28,11-30 120-197 Trachipteridae 28-54 Serrivomeridae Chaenopsidae 18-46,0-38 145-148 28-59 152-160 Radiicephalidae Cyclopteridae 28-78 Malacanthidae Eurypharyngidae 7-10,22-27 155-196 29-35 162-432 Chaetodontidae 11-13,18-25 Nemichthyidae 29-37 Congridae 202-647 Cetomimidae 30-31 206-263 Lophotidae Alepisauridae 30-45 Anarhichadidae Pomacanthidae 11-14,17-25 218-250 218-250 31-36 Saccopharyngidae Trichodontidae 14-16+18-20 224-285 32-36 7-9,22-43 226-262 Derichthyidae Acanthuridae 32-50 Bramidae 258-344 Ophichthidae 32-55 Zeidae 8-9,25-27 260-412 Regalecidae 33-36 Batrachoididae 2+31-39 33-41 367 Muraenidae Dactyloscopidae 7-22,18-36 370-382 Nettastomatidae 33-50 Nematistiidae 7-8+ 1,26-28 34-37 60

73 Table Introduction Ranges of total anal-fin ray counts (counts for spinous and soft rays combined) for fish families in the 5. region. Fin (spinous rays shown CalCOFI formulas for type) after each family, except given families with single bold are in spinous rays. Only fms found in the region are included in the ranges. Counts for Macrouridae are given rarely lacking species the literature and are not listed here. Counts for Carapidae are for total rays anterior to the 31 st vertebrae. Counts for fmlets in (Scomberesocidae, Carangidae, Scombridae) are Gempylidae, parentheses. Counts by authors of chapters in this in furnished guide. 9-23 Regalecidae Scombridae 9-23+4-10 0 9-27 Trachipteridae Clupeidae 0 Syngnathidae 2-5 9-46 Melanostomiidae 10 Linophrynidae Apogonidae 3 2,8 3-5 Melanocetidae Howellidae 10 3,7 10-12 Mugilidae Ceratiidae 2-3,7-10 4 10-14 Lutjanidae Ogcocephalidae 3,7-11 4 Thaumatichthyidae 10-17 Haemulidae 2-3,10-14 4 10-17 Oneirodidae Tetragonuridae 4-6 1-2,9-15 10-28 Gigantactinidae Bathylagidae 4-7 11-12 Agonidae Polyprionidae 4-14 3,8-9 11-13 Leptochilichthyidae Centrophrynidae 5-6 11-14 Chaunacidae 5-6 Giganturidae 11-15 Argentinidae Albulidae 5-10 11-18 Diodontidae Gobiesocidae 5-15 11-24 Hemiramphidae Lophotidae 5-20 11-26 Sternoptychidae Lophiidae 6-7 11-27 Myctophidae Radiicephalidae 6-7 11-37 Gempylidae Tetraodontidae 6-12 1-3+10-35+(0-7) 11-51 3,3-11 Scorpaenidae Alepocephalidae 6-14 12 Scaridae Cottidae 3,9 6-25 1-2,6-7 12-13 Aulopidae Mullidae 7-9 12-14 Anoplogastridae Neoscopelidae 7-10 12-14 Microstomatidae Psychrolutidae 7-11 12-15 Exocoetidae Scomberesocidae 12-15+(4-7) 7-14 12-16 0-3,7-18 Labridae Gobiidae 3,9-13 7-19 12-17 Holocentridae Antennariidae 4,8-13 8-9 12-17 Phosichthyidae Callionymidae 8-9 12-17 Pomacentridae 1,7-11 2,10-15 Melamphaidae 8-12 Elopidae 12-18 8-13 Cyprinodontidae 12-19 Caulophrynidae Opisthoproctidae 8-14 12-23 Osmeridae Synodontidae 8-15 12-24 1-2,6-23 Kyphosidae Sciaenidae 3,9-21 8-25 12-28 3,6-7 Astronesthidae Cirrhitidae 9-10 1,8-9 Kuhliidae Eleotridae 13 3,10 9-10 13-14 Sparidae Ostraciidae 3,10-11 9-10 13-14 Uranoscopidae 3,6-8 9-11 Gerreidae 1-2,8-10 13-15 Lobotidae 3,10-12 Sphyraenidae 9-12 1,8-11 13-16 Rondeletiidae Triglidae 9-12 13-18 Alepisauridae Chauliodontidae 9-14 3-5,6-9 13-18 Priacanthidae Pentacerotidae 9-14 3,10-15 Luvaridae Serranidae 0-3,6-19 14 9-22 14 61

74 Table Introduction 5. Continued. Anotopteridae 27 14-17 Cetomimidae Platytroctidae 14-17 Zeidae 27-29 F istulariidae 14-19 Centrolophidae 27-32 3,24-29 Molidae 14-19 27-32 Evermannellidae 2,12-30 14-32 Blenniidae 27-38 Batrachoididae Barbourisiidae 15-18 28-34 Zaproridae 4,24-30 Mirapinnidae 28-38 Microdesmidae 15-18 3,12-15 Polynemidae 28-43 Dactyloscopidae 15-18 2,26-41 Opistognathidae 2-3,13-17 15-20 28-43 Idiacanthidae 12-16,3-4 15-20 Xiphiidae 28-52 Monacanthidae 28-56 Stichaeidae Caristiidae 15-23 1-3,26-55 Hexagrammidae 0-4,12-28 29-30 Trichodontidae 15-28 1,28-29 Belonidae 30-36 Oreosomatidae 15-29 30-55 Stylephoridae 16-17 Pholidae 1-2,29-53 Notosudidae 16-21 33-35 Bathymasteridae 1-2,31-33 33-42 Stomiidae Lamprididae 16-22 Anoplopomatidae 2-3,14-23 34-44 Icosteidae 16-26 Nomeidae 1-3,14-31 36-41 Scytalinidae 16-34 36-44 Merlucciidae Echeneidae 16-38 Gonostomatidae 36-54 Carapidae 16-72 Nematistiidae 2-3,15-17 36-76 Moridae 17-20 17-76+0-29 Chaetodontidae 3,14-19 37-47 Stromateidae 17-22 2-3,35-44 Scopelarchidae Pleuronectidae 38-102 17-29 2,15-28 41-47 Tripterygiidae Achiridae 17-30 Engraulidae 43-106 Trichiuridae 17-33 1-2,41-105 Carangidae 1,14-33+(0-1) 45-51 Cryptacanthodidae 1-2+ 17-34 2-3,43-49 45-60 Bregmacerotidae Centriscidae 18-19 Ephippidae 2-3+16-20 46-81 Paralichthyidae 18-23 Labrisomidae 1-2,16-27 55-88 Monognathidae 18-29 0-1,17-29 58-85 Bothidae Chiasmodontidae 18-30 Diretmidae 62-114 Zoarcidae 1,18-24 19-25 Istiophoridae 13-20,6-7 62-181 Ophidiidae 19-27 Balistidae Cynoglossidae 63-98 20-30 Atherinidae 1,19-31 72-86 Cyematidae 20-32 Bythitidae Paralepididae 72-91 20-38 2,18-39 118-147 Eurypharyngidae Chaenopsidae 20-41 Pomacanthidae 118-201 N otacanthidae 3,18-22 14-59+ 104-142 21-25 Malacanthidae 1-2,20-26 139-460 Congridae 21-28 3,18-35 143-152 Serrivomeridae Embiotocidae 21-38 Cyclopteridae 155-180 Derichthyidae 21-73 Ammodytidae 170-372 Nemichthyidae 22-32 Clinidae 180-234 Anarhichadidae 1-3,21-35 22-37 0-1,180-233 Acanthuridae 3,19-35 190-256 Ophichthidae 22-38 226 Coryphaenidae Muraenidae 23-31 Saccopharyngidae Malacosteidae 235-284 25-29 Bramidae 283-317 Nettastomatidae 25-43 62

75 present Figure Outline illustrations of representative larvae of families of fishes 7. in the CalCOFI survey area. Introduction Flexion stage larvae were illustrated if they were available. Original Illustrations prepared by W. Watson and R. C. Walker. Derichthyidae Elopidae Saccopharyngidae Nemichthyidae Eurypharyngidae Albulidae N ettastomatidae N otacanthidae Clupeidae Serrivomeridae Muraenidae Engraulidae Monognathidae Ophichthidae Argentinidae Congridae Cyematidae Bathylagidae 63

76 Figure 7. Introduction Continued. ... \ ) ~,~ Stemoptychidae Malacosteidae Microstomatidae Phosichthyidae Idiacanthidae Opisthoproctidae ~= ~ ... Chauliodontidae Aulopidae Alepocephalidae Stomiidae Scopelarchidae Platytroctidae Astronesthidae N otosudidae Osmeridae ~ ... ~:. . ... - .. _ ... - .. _ .. _._.-_.- .. -.. ~:.~~ ...•.. .. ~ Melanostomiidae Synodontidae Gonostomatidae 64

77 Figure Continued. 7. Introduction Paralepididae Myctophidae Ophidiidae Bregmacerotidae Carapidae Anotopteridae Macrouridae Bythitidae Evermannellidae Batrachoididae Moridae Alepisauridae Lophiidae Merlucciidae N eoscopelidae 65

78 7. Continued. Introduction Figure Oneirodidae Linophrynidae Antennariidae /" ,/ Thaumatichthyidae Gobiesocidae Chaunacidae Atherinidae Ogcocephalidae Centrophrynidae , <),~=r >., Cyprinodontidae Ceratiidae Caulophrynidae Gigantactinidae Scomberesocidae Melanocetidae 66

79 Continued. Figure Introduction 7. Trachipteridae Mirapinnidae Belonidae ] '" e Fistulariidae Diretmidae Hemiraphidae Anoplogastridae Centriscidae Exocoetidae Syngnathidae Holocentridae Lophotidae ? ~'~ Melamphaidae Scorpaenidae Radiicephalidae 67

80 Figure 7. Continued. Introduction I ... . ~.; d···· Agonidae Triglidae Apogonidae - .. .----------_ Cyclopteridae Echeneidae Anoplopomatidae Polyprionidae Carangidae Hexagrammidae Nematistiidae Serranidae Cottidae Priacanthidae Psychrolutidae Coryphaenidae 68

81 Figure Continued. 7. Introduction Gerreidae Bramidae Kyphosidae Haemulidae Ephippidae Caristiidae Chaetodontidae Sparidae Lutjanidae Pomacentridae Sciaenidae Malacanthidae Cirrhitidae Mullidae Lobotidae 69

82 Figure Introduction Continued. 7. Scaridae Opistognathidae Pholidae Bathymasteridae Anarhichadidae Howellidae Mugilidae Zoarcidae Zaproridae Polynemidae Stichaeidae Chiasmodontidae ~ ___ n ___ ~ Cryptacanthodidae Labridae Ammodytidae 70

83 Figure 7. Continued. Introduction Blenniidae Sphyraenidae Uranoscopidae Icosteidae Gempylidae T ripterygiidae Scombridae Callionymidae Labrisomidae Trichiuridae Eleotridae Clinidae Centrolophidae Gobiidae Chaenopsidae ...:::.:.:.:.:.:.:.:.:.;,;,;:.:.::.;; Nomeidae Microdesmidae Dactyloscopidae 71

84 Figure 7. Continued. Introduction Achiridae Tetragonuridae Tetraodontidae Cynoglossidae Diodontidae Stromateidae Balistidae Paralichthyidae Molidae Monacanthidae Bothidae Ostraciidae Pleuronectidae 72

85 DESCRIPTIONS 73

86 ELOPOMORPHA S. H. G. MOSER R. CHARTER AND related closely al. Elopi- and notacanthiforms et grouped the the to (1966) orders Greenwood anguilliforms Anguilliformes, than they are and formes, together Notacanthiformes follow to and megalopids. We elopids on based Elopomorpha superorder the in cranial Eschmeyer's (1990) the which in classification elopo- presence osteology and the orders, the Elopiformes, morphs into five are divided of a leptocephalus larva. In Albuliformes, and Anguilliformes, Notacanthiformes, includes Elopiformes the classification, their two Saccopharyngiformes. consists Elopiformes of the the suborders: Albuloidei, consisting of the Albulidae with and and Megalo- one Elopidae, species, six genus (bonefishes), and the Elopoidei composed of the and (ladyfishes) Elopidae (tarpons). Megalopidae the whereas and two species, genus one with pidae, Albuli- Albulidae, with of family, a single the consists formes et al. (1966), much debate has five least at and genera two species. Since of Characteristics Greenwood Elopiformes are summarized in the and Albuliformes elopomorph with relationships, some surrounded Albulidae. Elopidae for sections introductory and (Nelson albulids the to status ordinal workers assigning Anguilliformes, Notacanthiformes, Saccopharyngi- and 1973a; Greenwood 1977; Eschmeyer and others 1990) separate have formes sections. introductory and 1977). them as a suborder (Patterson listing Rosen agree workers Recent generally are albulids that more Orders included: Elopiformes Albuliformes Notacanthiformes Anguilliformes Saccopharyngiformes 74

87 ELOPIDAE: Tenpounders R. AND H. G. MOSER S. CHARTER The only in the California Current region, Elops elopid are leptocephali Elopid described. been not have eggs affinis, Peru ranges from Ventura, California, to to (BD length depth in and moderate in moderate small Whitehead Schultz 1909a; 1962; Fitch and (Regan and the 5-12% BL). The head is depressed snout is are adults The 1983). al. et Eschmeyer 1978; uncom- The relatively short. origin is under anal fin the of the mon north Baja Magdalena, Sur, of California Bahia dorsal pigment it. behind just or is base fm Midlateral collected in larvae regular CalCOFI and have not been postanal melano- (four larvae yolk-sac from persistent samples. ichthyoplankton later- and larvae Yolk-sac between phores) through postflexion (vertical dashes samples stage the larvae occur in ichthyoplankton from myomere segments). a Elops affinis leptocephali have count lower myomere (78-82) than anyeelleptocepha- of California (pers. obs.) and south to Panama Gulf except li more myomeres (72-120), and 1963a). (Hildebrand Cyematidae also (70-72). from albulids larval differ than They The consists Elopidae family of to subtropical six leptocephali positions in albulid the relative of the and presence fms, anal dorsal species shallow-water that may enter estuaries. tropical lateral pigmentation, of by light-tackle anglers. prized They are and bladder. a vertical in having Elops affinis larvae Elopid gas mm) transform at a smaller size (40-50 albulids than 91 cm reach (Miller and Lea 1972). may 4.5 kg and be may Elopids recognized by a bony plate in the gular larvae Megalops larval resemble Elops mm). (60-70 with region, a terminal mouth a large gape extending Megalops but have more larvae myomeres than eye, branchiostegals 23-35 and the beyond (Hildebrand not Elops larvae but (51-68). The head is depressed in 1963a). The pectoral fins are low on the body and the Elops in fm anal than dorsal Megalops. more have rays All pelvic fins are abdominal. slender are species Elops while fm anal more have Megalops dorsal rays than and sides, silvery dorsum, bluish a with bodied, (Richards 1984; 1989m). Smith slightly yellowish ventral anal and dorsal The regions. lack filaments last fins (Hildebrand rays the to attached (1963a) captures reported Hildebrand affinis of E. 1963a). have species other and affinis a small Elops the in and August, autumn leptocephali in February, at spot lower dark the 1979). jaw symphysis (Nybelin along the Pacific coast of Panama. Our larvae were collected raker count February, Gulf the from June, E. affinis has (16-20) a high gill in and August, Also, (Regan 1909a; Hildebrand 1943; Whitehead 1962; yolk-sac of California. Detailed examination of two Whitehead and mm), (5.4, 5.8 one flexion (12.3 mm), and four Rodriguez-Sanchez 1995b). postflexion (16.9-33.6 mm) larvae provided the meris- and description. tic this for data morphometric their Elopids are oviparous (Hildebrand 1963a); 75

88 ELOPIDAE Elops a/finis MERISTICS DESCRIPTION EARLY LIFE HISTORY Range Mode EGGS Vertebrae: diam.: Shell Yolk: 78-82 Total 80 No.ofOG: Diam.ofOG: Precaudal 52-54 54 Shell surface: Caudal 25-29 27 Pigment: Fins: features: Diagnostic Dorsal spines 0 Dorsal rays 20-27 LARVAE Anal spines Hatching <5.4 mm length: 0 12-18 rays Anal Flexion mm <12.3 length: 12-16 Pelvic mm length: Transformation >33.4 17 17 Pectoral sequence: PI' C development Fin D, A, C & P & l 2 2 Caudal: pigment Pigmentation: Yolk-sac-preflexion-Eye 5.8 not complete at 10+9 10+8-9 Principal mm; around 4 midlaterally starting at anus; scattered over anus; Procurrent: of tail. Flexion-l dorsally on gut; on margin ventral notochord tip; Upper 8 over heart; at midlaterally beginning ca. 23% BL, ending over & eye Lower 8 Crays on developing & less notochord; over over anus; anus; Gill rakers: on ventral margin hypurals; along dorsally on vertically of caudal gut; Upper 10-12 form to eye over Poslflexion-Increasing a crescentic peduncle. 12-20 Lower dashes heart; over enlarging vertical becoming midlaterally patch; 23-35 Branchiostegals of gut from throat, surface myomere between dorsal segments; on becoming dashes. horizontal HISTORY LIFE Leptocephalus; Diagnostic features: total myomeres ca. 79 (77-81); D overlap depressed pigment; forked & A head; midlateral so; nearly or becoming thickens, & long shrinks body transformation, at gut; tail; rare generally Peru; to California, Ventura, from Pacific Eastern Range: A. & D of movement forward apparent opaque; north of California Sur Baja %) in mean & (range MORPHOMETRICS areas; inshore Shallow Habitat: in schools & August season: Spawning June in February, collected Larvae Y-S PrF F PoF Tr Juv & eggs Oviparous; planktonic leptocephali pattern: ELH Sn-AlBL 81-88 93-98 85 97 96 LITERATURE 11-12 BDIBL 5-7 12 7 6 1943, Hildebrand 1963a 12-16 HLlBL 8-12 1984 Richards 14 13 10 1989rn Smith 64-67 HWIHL 51-55 54 66 52 ORIGINAL ILLUSTRATIONS (Illustrator) SnUHL 31-37 34-34 34 34 34 Yolk-sac Arthur) 5.8 mm (N. larva, EDIHL 14-18x mm (N. Arthur) Flexion larva, 12.3 17-22 Sumida mm (B. MacCall) 16.9 larva, Postflexion 20x22 28* 16x20 mm 16.9 of MacCall) postflexion larva (B. Sumida Head 3-3 2-4 3 3 3 One yolk-sac larva with becomes slightly oval; near-round eye intact; eye * 0-0 0-0 given is axis vertical second. first, horizontal axis o o 0 76

89 Machete Elops a/finis mm 12.3 16.9 mm ':) ')) \~--' "'.. / / Figure Elopidae 1. Yolk-sac larva, 5.8 mm (CalCOFI 5607, station 145G.40); flexion larva, 12.3 mm (CalCOFI 5708, station 103G.70); and postflexion 16.9 mm, whole body larva, head (CalCOFI 5708, station 131G.94). 77

90

91 ALBULIDAE: Bonefishes G. s. H. MOSER AND CHARTER R. CalCOFI single the in occurs species albulid A survey Pfeiler 1984; et California of Gulf the in al. (pfeiler guides Regional identification (Fitch and Laven- area. CalCOFI in the Gulf of Califor- single A 1988). cruise 1972; Lea and berg Miller 1971; 1978; Castro-Aguirre 574 nia larvae (3.2-48.2 mm) 1957 August in yielded Alhula as species this list 1983) al. Eschmeyer et from (pers. Gulf the of regions all obs.). vulpes; of and however, Hawaiian the discovery two Indo-West species Pacific electro- through Alhula of and have presumably Bonefish are oviparous Tamaru 1981; and (Shaklee analysis phoretic Shaklee have they although eggs, planktonic de- been not et the brought 1982) identity of the question into al. scribed. Larvae unpigmented as 3.2 mm, with small as eastern eyes, nonfunctional mouths and large yolk reserves the species and Pacific that Alhula concept have been collected in the Gulf vulpes (1981) circumtropical Pfeiler Albulids California. of species. is a single that and Smith (1989m) suggested the bonefish in our have be leptocephalus distinguished can that larvae region, here referred to as albulids count; myomere by eels most of those from Alhula sp., is an undescribed total while eel leptocephali have myomeres <100 have species; this hypothesis is supported by electrophoretic 1984). Total > 100 (Smith 1979, 1989d; Richards evidence (pfeiler 1996). in the family saccopharyngiform myomere count are is <100 but their leptocephali extreme- Cyematidae tropical through- Bonefish inhabit nearshore waters The deep-bodied ly characteristics. other in unique and fish recreational a valuable are and world the out and fin caudal forked and anal separate large distinctly in Cal- the Bonefish ability. fighting their for prized leptocephali albulid late-stage and of mid- dorsal fins 46 and COFI (Miller and cm 0.8 ca. reach may area kg anguilliform from and notacanthiform distinguish them mud with water shallow prefer They 1972). or Lea of Albulid larvae from differ leptocephali leptocephali. They snout sand bottoms. recognized by a conical are of elopids in gut length, and fin, anal the position almost an lower the beyond projects that inferior jaw, extends almost lateral pigmentation. In albulids the gut the before ending mouth horizontal pectoral low eye, body origin the of the and anal the of length entire the is grey- fins, abdominal pelvic and fins. Coloration the gut is In is well behind the dorsal fin. elopids fin brown above lateral line and silvery below. the fin anal the below the and shorter relatively is origin dorsal albulid larvae lack lateral melano- Also, fin. collections from CalCOFI Larval albulids are rare in elopids. in present phores the Baja California. Juveniles Pacific coast of and Francisco, San as north far as California occur adults as larval series (1961) Alexander by described The 15 sp. There 1950). Alhula of occurrences are (Fitch 26 mm A. vulpes is similar to a partial series (12 NL to larvae mm) in CalCOFI samples in the (5.7-53.5 Pfeiler described sp. Alhula of SL) mm by al. et nearshore region southern between California of Baja that the (1988) Gulf of California. We the from assume Lucas. Punta have larvae No Abreojos San Cabo and is conspecific herein described complete larval series off or northern Baja Califor- California been collected (1988). al. et Pfeiler of use We the series partial with although nia and transformation specimens demersal 30 Pacific coast specimens mm) (3.8-117.2 from the mm have been collected juveniles as as 18.6 small of description. following in the California the and Gulf obs.). California (pers. bays In southern in from contrast, sp. Alhula widespread common are and larvae 79

92 Alhula ALBULIDAE Sp. EARLY HISTORY DESCRIPTION LIFE MERISTICS EGGS Range Mode diam.: Yolk: Shell Vertebrae: Diam.ofOG: No.ofOG: 68-74 Total surface: Shell 52-65 Precaudal Pigment: 5-16 Caudal Diagnostic features: Fins: 0 Dorsal spines LARVAE 3-20 16 Dorsal rays length: <3.2 Hatching mm 0 Anal spines Flexion length: ca. mm 15.5 9 4-10 rays Anal 71.0 ca. length: Transformation mm 9-11 Pelvic , D, , C PI' sequence: development Fin C P A, 15-18 Pectoral I 2 2 on edge of alOiJg 1 ventral yolk sac; Pigmentation: Yolk-sac-4 gut; Caudal: radiating finfold caudal on notochord. from 11+11 Pre flexion-Paired rows 8-11+9-12 Principal along anus; of dashes on dorsal edge of gut from throat to caudal Procurrent: over Posiflexion-A eye crescentic patch by mm. 16 ca. by fmfold 4-9 Upper blotches dusky initially becoming 29 ca. yellow; with mixed mm; 3-8 Lower crossbands 12 Juvenile---ca. to lateral extending back on silver. dusky Gill rakers: above on line; eye; snout & by grey-brown ca 82 mm, on D & C; 15-17 Upper on below, D & C yellowish-brown above lateral line, silvery white Lower margins. 10-16 Branchiostegals Diagnostic ca. total myomeres Leptocephalus; features: gut (70-72); 71 elopiforms; overlap no in than longer lateral A; & D of pigment; no LIFE HISTORY occurs at thickens; & shrinks body transformation, at tail; forked size in than D larger of movement forward apparent eIopids; & A. the Range: Pacific from Eastern San Francisco, California, throughout to Gulf of California, south Panama (range in mean & %) MORPHOMETRICS Shallow Habitat: fresh enters occasionally waters, subtropical or tropical PrF Y-S Tr PoF F Juv sand flat & mud soft on bottom water; or brackish Sn~AlBL 87-98 98-99 94-97 7-87 68-73 76-90 season: Spawning 93 83 84 96 71 99 pattern: leptocephali, planktonic Oviparous; in tide ELH found shallow 7-14 6-9 BDIBL 4-5 4-4 10-30 18-20 bay shallow in specimens transformation offshore; miles 60 pools to 16 4 5 7 11 19 around areas, juveniles piers, settling to sand bottom; in shallow tide 8-10 9-12 7-9 HLlBL 6-9 16-32 26-32 sand hard over pools shell & bottom 8 10 9 8 21 31 38-54 40-47 51-95 36-52 HWIHL 35-43 44-47 LITERATURE 44 46 46 39 44 80 27-30 29-36 25-33 17-29 25-32 SnLIHL 34-40 1961 Alexander 28 22 30 29 33 37 1950 Fitch 24-26 25-34 23-26 21-26 21-27 49-58 EDIHL 1984 Pfeiler 29 25 53 23 24 24 et 1988 Pfeiler al. Richards 1984 2-4 2-3 2-3 0-1 6-16 15-32 PILlBL 3 3 2 0.3 20 9 1989m Smith 1979, 1989d, 0-1 0-0 0-0 0-0 5-14 L1BL P 14-28 2 ORIGINAL ILLUSTRATIONS (Illustrator) 0 0 17 8 0 0.1 (B. Sumida MacCall) Yolk-sac mm larva, 5.2 MacCall) Sumida mm (B. 10.6 Preflexion larva, Flexion larva, 23.6 mm (N. Arthur) Postflexion larva, 33.4 mm (B. Sumida MacCall) Arthur) (N. mm 34.2 specimen, Transformation 80

93 Bonefish Albula sp. mm 5.2 10.6 mm 23.6 mm 34.2 mm FigureAlbulidae 1. Yolk-sac larva, 5.2 mm (CaICOFI 5708, station 105G. 70); preflexion larva, 10.6 mm (CaICOFI 5708, station 129G.89); (SIO 68-168). 5708, mm; postflexion larva, 33.4 mm (CaICOFI larva, station 131G.94); transformation specimen, 34.2 mm flexion 23.6 81

94 NOTACANTHIFORMES G. AND S. R. CHARTER H. MOSER eel-like by the recognized fishes Notacanthiforms are Halo- the pers. Smith, of Representative comm.). directed spine on the upper a backwardly of presence sauridae the from reported been not have California posterior margin maxilla, a projecting of snout, the a result Current region although be may this of limited abdominal a pelvic fins connected by membrane, sampling in their deep continental slope abyssal to plain habitat. and tapering tail with pectoral fms high on the body, caudal a reduced at posteriorly fin anal a long ending leptocephalus larvae that can have Notacanthiforms is fin some species). The eye absent in fin (caudal and m) (1-2 size large numer- a remarkably reach have that is continous covered by a transparent membrane including features, distinctive a long straight gut ous head. with the skin (1973) McDowell covering the extends length the entire that body, the a short-based of 6 genera, 3 families, recognized and the reviewed order late-developing dorsal V-shaped fins, pelvic fin, other with elopo- 22 species. Their relationships and and (Smith filament caudal a delicate myomeres, 1970, are Nelson (1973a) proposed unsettled. morphs that 19891; The caudal filament is broken off Castle 1,984). a similar on based albulids arrange- to closest they are of presence literature. in the described larvae in all The canal of the mandibular sensory in the two ment -shaped pelvic fins, a compact dorsal myo- fin, and V whereas are they suggested (1962) Marshall groups, meres leptocephali notacanthiform separate clearly closer anguilliforms similarities on based to in the from those types Three of of notacan- Anguilliformes. swim bladder. recognized leptocephali thiform are primarily by eye The order Notacanthiformes represented is in the relative shape has none however, and length; snout 1970, family the in by Current California two region species been to assigned (Smith definitively a family inhabit deep-shelf, conti- Notacanthidae. Notacanthids 19891; Leptocephalus Castle 1984). One larval type, world. the throughout abyssal and slope, nental regions giganteus, a worldwide distribution has supposedly their deep captures of these few distribution, Owing to although it that apparent is different from larvae North the for documented been have fishes eastern We a describe in pigment. regions vary and structure monotypic The Pacific. Lipogenyidae from known is specimens of fairly complete developmental series Japan Zealand, the North and New Atlantic, G. (D. mm) from CalCOFI region. (18-560 the Spinyeels NOTACANTHIDAE: H. MOSER AND S. R. CHARTER G. notacanthids are represented in the CalCOFI short, isolated dorsal spines and a long anal fin with Adult that to region by two species, Notacanthus chemnitzi and posteriorly grade anteriorly of spines consisting covered rays. The body and compressed are head and N challengeri. Polyacanthonotus of Six specimens is small and small, with mouth The scales. cycloid California central from recorded been have chemnitzi comb- teeth inferior; are small and arranged a single in of 1100-1500 m (peden British to Columbia at depths The row. like pelvic fins are abdominal and have one Lea four and 1987) Rosenblatt and 1976; P. challen- The caudal spines. several to fin minute is (McDowell geri from Vancouver to Island have been reported Sulak 1986). 1983; al. 1973; et Eschmeyer 1968; (Peden m 2100-2450 of depths at Oregon Stein additional and N We examined three Butler 1971). described The notacanthiform leptocephalus series and chemnitzi include counts and challengeri P. one represent type, a possibly a single to appears herein measurements (Table literature the from those with and to similar those single species, whose characters are of Notacanthidae 1). 1967, (e.g., giganteus Leptocephalus Castle 1959, One Tabeta 1970). of 1970; 1973; Nielsen and Larsen Adult (up notacanthids medium-sized to small are our filament, caudal intact a relatively has specimens but fishes ca. 125 cm eel-like most species <50 cm) to 82

95 demonstrating morphology and pigmen- the remarkable pelvic fin anlagen in relation to myomeres dorsal and tation the mm specimens 18-22 and structure, that of the in larval form are similar to the positions of these the first larvae are early of reported. to giganteus L. be in the adult. relation fins to The number in vertebrae of All :::::108 our specimens of have a series of paired mm giganteus L. in pterygiophores fin dorsal developing is gut the to similar be to appear which pouches lateral similar in count ray dorsal the to N The chemnitzi. a (1970) on by observed tubercles" "fleshy Tabeta only number the in is difference major of larval giganteus They Peru. off captured L. 75 cm develop and myomeres adult vertebrae. When countable, total entire the along hindgut, initially as slight protrusions ranged larvae our in myomeres from to 330 350. The gut lateral the along specimen largest our In wall. range is literature the in (Smith 19891). The 300-486 of the (560 they appear hollow outpouchings mm) as range of chemnitzi in from the number vertebral N wall that ca. gut and body project the 0.5 mm beyond northeast Pacific and Rosenblatt (Lea 225-239 is 1987; nutritional lateral body contour. They may have a be explained could difference This study). this if function absorptive surface of the by the increasing in vertebrae do not form association with the myo- hindgut. in meres the posterior region Myomeres of the larva. difficult and minute The region this in to count are larvae in giganteus L. of early stage presence The last of only about 10% 100 the body myomeres occupy the CalCOFI samples indicates local origin. No nota- length and their coalescence trans- degeneration at or canthiforms are known except to occur in the region N much count a vertebral formation produce could lower chemnitzi and P. challengeri. The southernmost record than the loss However, fishes. other in found ratio 1 1: of is ca. at Oregon, Newport, off challengeri P. 45°28' seems of large numbers of myomeres at transformation N N while specimens of chemnitzi have been captured unlikely, and this of we know documentation of no in occasions on several California. The occur- central off for few counts notacanthi- The fishes. other vertebral rence giganteus L. of stages CalCOFl early in of larval forms the literature are <250 while myomere in counts from ranging localities at samples Conception, Pt. types other for leptocephali notacanthiform putative of California, to us California, Eugenia, leads Punta Baja (e.g., 331 Tiluropsis) (Smith range from 229 19891). to that suspect to form this be the of giganteus may L. It that have the is possible L. adult forms of giganteus larva chemnitzi. N chemnitzi of P. N than Other and not been collected, although this seems given unlikely the notacanthiform reported only challengeri, from the worldwide the distribution the of larvae. A comprehen- area (identifi- specimen a mutilated is survey CalCOFI sive review existing in counts vertebral of notacanthi- cation uncertain) of the halosaur Halosaurusattenuatus specimens, form including the frequency and of degree Baja California off central (Townsend Nichols and to caudal regeneration, would provide valuable clues 1925). the that possible is It are leptocephali CalCOFI the identification larvae. of their the larvae been not have of halosaurs whose adults based giganteus is L. description following of The collected; is this however, and larvae since unlikely on detailed examination the of nine larvae of following specimens transforming of 1965; (Mead halosaurs lengths: 18.4,20.3,22.0, and 191,301,311,314, 108, Harrisson 1966; have characteristics 19891) Smith 560 were from several obtained Meristic mm. data different than from those notacanthids of (Merrett sources 1971; (Peden 1968, 1975; Stein and Butler Lea 1981; Smith 19891). and Rosenblatt 1987) and from counts made during A number of morphological features of L. giganteus this study. of chemnitzi. N adult of those match the positions The Table N 1. Meristic characters for the notacanthid species recorded from the California Current region. Vertebral otacanthidae are counts given precaudal + caudal = total. The origins of as dorsal, anal, and pelvic fins are indicated by the sequential the origin. of at the point vertebral number rays Fin D A P 2 Vertebrae Species D P A C PI 2 origin origin origin IH-IV,6-7 14-16 04-133 IX-X,2 XIV-XVII, 1 54-58+171-183=225-239 42-55 5 41--48 38--47 Notacanthus chemnitzi 1+8-10 11-15 XXXIII- XXXIX-LIX, 55-58+187-197=242-255 5 12 55 45 Polyacanthonotus XXXVII 126-142 challengeri 83

96 NOTACANTIDDAE Leptocephalus giganteus EARLY LIFE HISTORY DESCRIPTION MERISTICS EGGS Range Mode diam.: Vertebrae: Shell Yolk: Total No.ofOG: Diam.ofOG: surface: Shell Precaudal Pigment: Caudal features: Diagnostic Fins: Dorsal spines LARVAE Dorsal rays Hatching length: Anal spines length: Flexion Reduced C Anal rays Transformation length: Pelvic , D, PI> sequence: P Fin A, C development Pectoral 2 Paired Pigmentation: number in increase that blotches ventral Caudal: throughout larval period, in at 9-11 108 18-22 mm larvae, ca. 35 Principal mm, 560 at 340 as 301-314 at 104-122 mm, 191 at 75 mm, mm; Procurrent: ventral across fuse they enlarge blotches midline extend dorsad to & Upper new with embedded blotches added between larger ones; trunk, Lower pair median gut between in large an & ventricle of heart; larvae Gill rakers: heavily leaflike, heart; of develops embedded pair above atrium Upper pigmented, structures on caudal filament. Lower features: Leptocephalus body gut straight form Diagnostic with Branchiostegals of body; >300 caudal myomeres; length entire extending almost filament (usually broken) has series of pigmented leaflike structures; HISTORY LIFE ca. D with short appears pterygiophores at ca. 100 mm; P 2 appears 10 at ca. mm; eye nearly 300 hindgut with lateral outpouchings. round; from California eastern Pacific in worldwide; Apparently Range: central Peru to in mean MORPHOMETRICS (range & %) demersal are deep-shelf to abyssal from notacanthids Adult Habitat: mm 18-22 108-314 mm depths 560mm 99-99 September- Sn-A/BL* collected mm) Spawning season: Smallest larvae (18-22 87 99 99 December Current California in BDIBL 5-5 2-3 Oviparous; pattern: larvae ELH planktonic 5 3 2 HLlBL 8-9 3-4 LITERATURE 3 9 2 30-44 HWIHL 23-42 1967, Castle 1973, 1984 35 29 21 Nielsen & 1970 Larsen 26--28 SnLIHL 26-43 19891 Smith 1970, 27 36 46 1970 Tabeta 12-19x x 20-25 EDlHLt 23-30 13-22 ILLUSTRATIONS ORIGINAL (Illustrator) 23x27 IIx12 15x17 Vona) T. (M. 22.0 larva, Early mm 0-0 0-0.03 P1LlBL T. (M. Vona) mm 22.0 of 0.Q2 larva early Head 0 0.02 Mid-stage larva, 314.0 mm MacCall) Sumida (B. P 0-0 L1BL 0-0 bud 2 Head body Sumida (B. larva MacCall) mm 314.0 of section & mid-stage 0 0 since mm range given for 18-22 * No gut was intact only in 22-mm larva. t Eye is is slightly larvae <314 mm; first, given in axis horizontal elliptical vertical axis second. 84

97 Giant leptocephalus giganteus Leptocephalus mm 22.0 314.0 mm ~-- ------~.---- .. ./ _-----::::::::::-==:::::::::=::::::=:--:::--------- --.-- ... - Figure Notacanthidae 1. Early larva, 22.0 mm, whole body and enlargement of head (CaICOFI 6612, station 120.55); mid-stage larva, body 314.0 whole body and enlargements of head and mm, section (hindgut region) (CaICOFI 8410, station 110.45). 85

98 ANGUILLIFORMES CHARTER G. MOSER s. R. H. AND and order of diverse Anguilliform eels comprise a large of AnguiHiform array a diverse have leptocephali families the inhabit They species. 600 15 ca. and allowed morphological and characters that pigment midwater regions and benthic enter and oceans an of workers early to describe their even species though brackish Seven freshwater. and families of a total with resulted This unknown. were stages adult a dual in species not presently assigned one 20 species, including taxonomy This of larvae and adults (Smith 1989d). a California to the in identified been have family, by problem has been resolved largely progress made Current region. in decades several past the during linking and larval stages adult almost all larvae can of identified eels; be order (1989) Robins summarized the and listed a many family genus to to and species (Castle 1969, or total 1984; 1984, Smith Raju 1985). 1989d; of 42 anguiHiform characters, including: very gills extending elongate body and head with the wen Developmental stages based on notochord flexion neurocranium, behind pelvic no fins, vertical confluent appropriate are not for eelleptocephaH because fins girdle, or fins high on the pectoral absent, body or of their larva, reduced caudal fin. Leiby (1979, 1989) markedly leptocephalus gill membranes united to the stages. two into development pretransformation divided and widely nostrils reduced, openings isthmus gill Engyodontic a few have larvae scales one separated but all re- or absent in family, the teeth, needle-like duced embedded, and lack lower or of no jaw, upper the and caeca, pyloric jaw equal to than longer characters. numerous no hypurals, Most nasal capsule, undifferentiated finfold, osteological of these preanal and lengths head and notochord, unflexed reductions or losses represent may be and characters to length. total respect with long relatively the In convergent; however, Robins (1989) argued that the and the stage euryodontic teeth are needle-like shed found such number large characters, not of in many three replaced series by elopomorphs, other support monophyly. anguiHiform shorter, broader relatively of Also, teeth. lower the the than shorter becomes jaw Apparently, a nasal hyp- upper, differentiate, fins forms, capsule are anguiHiforms an oviparous, al- preanal and head and flexes, notochord the form, urals though for known is a few species. only spawning become relatively smaller. Most lengths are Eggs planktonic, relatively large CalCOFI (3-4 mm in some have species), a wide perivitelline space and a seg- anguilliform larvae are euryodontic; however, when mented yolk (Eldred 1969). Like other elopomorphs engyodontic were identified, the morphometric larvae a planktonic a large have with larva leptocephalus they both tables, tables these In stages. ontogenetic include mucinous acellular transparent body filled with an values engyodontic for larvae are the under listed loops a small or head, a tubular gut with material, smaller ranges. values size two of were Morphometric thickenings in some species, and a pectoral fin that and transforming larvae when calculated yolk-sac for identified. specimens were leptoceph- be may lost AnguiHiform at transformation. a rounded or pointed have ali myomeres, W-shaped Methods caudal and fins, anal and dorsal the with fin confluent myomeres follow counting Leiby of pelvic fins. Larvae lack myomere first The (1989). have other elopomorphs reach of the not may mayor the midline, is but counted. Nephric still myomeres are and (1) characters: following elopiforms albuliforms- counted the from a vertical to midline the along caudad developed caudal W -shaped myomeres, a well forked terminus as determined easily of most nephros, the the and dorsal fin that is separate from short-based anal blood (L vessel point where the last vertical VBV) (2) and fins, fins present; pelvic saccopharyngiforms- myomeres are enters the kidney. Preanal and predorsal confluent -shaped V pointed tail a myomeres, with the terminus counted to a vertical from and anus the of and dorsal and anal fins, no and (3) pelvic fins; the origin these of AU respectively. fin, dorsal of the short myomeres, -shaped V notacanthiforms- -based are and total but identification larval in helpful counts elongate an fins, anal long-based and dorsal caudal are counts they since useful most the nephric myomere filament, 1989d). (Smith present fins pelvic 86

99 approximate measured precaudal vertebral counts in was (PdL) length predorsal when the dorsal total and muscle formed, attachment to fin prior were bundles to usually was (GD) depth body greatest The adults. length or at ca. body vent. The 50% the pterygiophores developing 1989). (Leiby measured at Muraenoidei Suborders and families included: Muraenidae Congroidei Ophichthidae Congridae Derichthyidae N emichthyidae Serrivomeridae N ettastomatidae 87

100 MURAENIDAE: Morays G. MOSER CHARTER R. S. AND H. live and subtropical all tropical in oceans Muraenids perivitelline 1969). Muraenid lepto- (Eldred space seas; and a few species and inhabit temperate waters length in moderate are cephali compressed strongly and (Bohlke adults and juveniles freshwater enter some as a moderate with BL), 5-15% (BD preanal long to region CalCOFI the in occur species Four 1989). al. et species). length (59-87% depending on The BL, the 1): Echidna nebulosa, Califor- (Table Muraenidae nostril liver is undivid- posterior is above The mid-eye. verrilli, Gymnothorax mordax, G. and nia moray the ed gut is a simple and tube. Morays are the straight lentiginosa. have been recorded Muraena species Five eel fins to completely resorb the pectoral larvae only Gulf in the of California (Anarchias galapagensis, region broadly transformation. is caudal The during zebra, octavianus, Enchelycore Gymnomuraena rounded. The fin may be dorsal 20-98% of body Gymnothorax castaneus, and panamensis) six G. and may length the anal genus, on depending and, fin Gulf the from range species south California of along at or immediately originate anus. the behind a distance Gymno- nocturna, (Echidna American the Central coast morphologically, similar are leptocephali Muraenid argus, G. equatorialis, Muraena M thorax dovii, separated be may species and genera but a combina- by clepsydra, and Uropterygius macrocephalus). Muraenid of pigmentation, myomere count at characteristics: tion CalCOFI ichthyoplankton leptocephali are rare in of nephros terminus, total count, myomere and position of the common relatively are but collections south mordax Gymnothorax anus. and origin fin dorsal is region. CalCOFI identified total myomere by the nephros terminus and respectively), 146-154, and (77-84 counts fin dorsal (ca. m) 1.2-3 large to medium common are Adults 19-53, 24), and pigmentation. (myomere origin usually residents benthic rock or coral reefs, occupying of leptocephali occur in the distinct other muraenid Two to zone subtidal the from caves and crevices shallow of of the CalCOFI area; they California and south Gulf ca. 500 m (usually shallower than 20 m). Morays ca. identified transformation without critical be cannot are with slightly confluent compressed, elongate and identified (1983) Lopez specimens. a larva Gymno- of and dorsal, lack both lateral line caudal, anal fins. They thorax verrilli from Rica. Costa pectoral and and round, is small, opening gill The fins. The may low on the be mouth is large; some body. muraenid Pacific is sparse for eastern Literature posterior have strong, sharp species teeth. The canine Castle Australasian described (1965b) leptocephali. thick, The eye. the above located is nostril scaleless north western described 1989g) (1979, Smith species, is light skin with color in greenish to brown dark or (1984) 15 genera with Smith Castle (1984) and and species Atlantic bars, spots, or mottling. Presently, of (Nelson recognized are species 200 approximately general examination Detailed summaries. provided Eschmeyer 1990). 1994; 46 mm) G. of (36.5-108.0 specimens larval mordax the this morphometric and meristic provided data for study. Muraenid eggs are large a wide (3-4 mm) with 88

101 Table Muraenidae Ranges (above) and means (below) of vertebral counts for muraenid species in the California Current 1. based on study and McCleneghan (1976). Geographical ranges are based on Miller and Lea (1972), McCosker and region this SIO Marine McCleneghan et al. (1984), Bohlke (1975), al. (1989), and specimens in the et McCosker Rosenblatt (1976), fin Abbreviations: number of vertebrae anterior to dorsal DFO, origin; AFO, number of vertebrae Vertebrates Collection. to anal rm origin; Gulf, Gulf of California; Galapagos, Galapagos Islands; CA, California; BCS, Baja California Sur. anterior Vertebrae DFO AFO PrCv Number Total Species ranges Geographical 102 II 101-109 56 103 & Galapagos Anarchias Gulf 105 galapagensis 5-6 14 56-58 120--128 Echidna 64-66 nebulosa Lucas to Cabo Rica, San Costa 57 65 123 6 Clipperton Is. & Eniwetok II 56-59 6-9 8-122 II 65-67 E. Gulf, Clipperton Is., nocturna Galapagos, 120 66 58 7 Panama Cocos Is., 67-70 141-146 8-9 56-59 Gulf Galapagos 8 & Enchelycore 8 68 143 57 octavianus ca. 84-85 95-97 14-14 128-137 7 zebra Gymnomuraena Gulf & Galapagos 14 85 ca. 96 131 57-59 138-146 II 6-8 & Galapagos Gymnothorax Gulf 66-68 67 7 58 142 castaneus 5-7 57-59 68-71 140--147 10 to Panama, Galapagos Gulf dovii G. 70 143 6 58 53-54 7-7 ca. 61-62 141-147 Panama to 8 Gulf equatorialis G. ca. 61 144 53 7 5-17 77-81 145-152 64-66 21 Bahia to Conception, CA Pt. mordax G. 148 65 Magdalena, 79 9 Is., BCS, Catalina Santa Guadalupe, Gulf, Galapagos Isla de 51-52 123-131 Galapagos 13-14 58-60 Gulf & 20 G. panamensis 51 126 13 59 173-175 76-77 Gulf, ca. 89-92 BCS, Bahia Magdalena, 4 6-6 verrilli G. 174 91 Panama ca 77 6 49 4 124-124 2 57 Peru, Rocas Alijos, BCS to Gulf, argus Muraena Galapagos 124 124-131 50--52 3-5 Cabo 10 San Lucas, BCS, Gulf, 59-59 clepsydra M 4 51 59 128 Panama, Galapagos 46-49 53-56 II4-119 3-5 Magdalena, Bahia BCS, Gulf, Costa 15 lentiginosa M II7 55 Galapagos Rica, Panama, 47 4 Revillagigedo, 52-56 lO6-IIl Gulf, Islas de 88-91 15 86-88 Uropterygius Clipperton 87 90 54 108 Colombia, Is., Panama, macrocephalus Galapagos 89

102 MURAENIDAE Gymnothorax mordax EARLY LIFE HISTORY DESCRIPTION MERISTICS Range Mode EGGS Yolk: diam.: Shell Vertebrae: No.ofOG: Diam.ofOG: 150 146-154 Total Shell surface: 74 Precaudal 84 Pigment: Caudal features: Diagnostic Fins: 0 Dorsal spines LARVAE Dorsal rays 367 length: Hatching 0 spines Anal Flexion length: 226 rays Anal length: mm*

103 mordax Gymnothorax California moray mm 44.1 --~---------- ----~ ~----~- 97.7 mm 71.1 mm Figure Muraenidae 1. Larvae, 44.1 mm, whole body and sections showing head, nephros, and caudal regions; 97.7 mm, whole body and sections head, detail showing of dorsal fin base pigment, and nephros regions (SIO 73-263); caudal region of a 71.1 mm larva (lower right) 5708, station 116G.37). (CaICOFI 91

104

105 OPHICHTHIDAE: eels worm and eels Snake S. R. CHARTER two family, eel largest the is sub- Ophichthidae with . 1989) (Leiby ichthinae) species . families, 250 and genera, 55 tribes, six Six 1989). al. et (McCosker occur species in the Ophichthids unknown. are Eggs oviparous. are Myrichthys CalCOFI study area: Ethadophis merenda, Leptocephali a moderate length (maximum ca. attain Myrophis triserialis, Ophichthus vafer, xysturus, O. three liver lobes and at mm), least two with 150 two or specimen A single miurus. Scytalichthys and zophochir, 0-15 gut swellings loops. The nephros terminates or merenda E. of was taken from the stomach of a white (Leiby 1989). Pigmenta- the anus anterior myomeres to California Baja Thurioe, de Bahia in caught seabass tion frequently found is on the myosepta midlaterally M Bahia from Sur. xysturus Baja Magdalena, ranges postanally and in most notochord the to ventral just to Sur Galapagos Islands, including the the California species (Leiby 1989). Larval ophichthids may be Gulf California. zophochir O. range of and M vafer characters: and number of separated a combination by Bay, Pedro San to Peru California, Humboldt and from gut loops, myomere lobes liver and of the morphology Gulf the including respectively, California. O. of and nephros the at counts and origins fin anal dorsal counts, myomere total and preanal terminus, and from ranges to triserialis Humboldt Bay, California patterns late In 1989). (Leiby transfor- pigmentation Peru, the Gahipa- the Gulf of California and including de rays, branchiostegal of gos the specimens mation Islands. Scytalichthys Isla miurus ranges from number Guadalupe to tip the and number placement of Apparently, all and of Baja California. pores, sensory cephalic gill osteology may separate species arch caudal and rare relatively are Baja of north although California, 1989). (Leiby liver have leptocephali vafer M three habits make them inaccessible to most their fossorial nephros at lobes with three gut swellings, the typically collecting methods (Rosenblatt and McCosker 1970). myomere the adult 60 (55-63), closely matching At least 20 ophichthid species occur in the additional count vertebral precaudal of a total myo- 58-62, and in collected been have Leptocephali California. of Gulf count mere (152-164) that overlaps the adult vertebral at the ichthyoplankton CalCOFI coastal samples count (146-155). triserialis O. and o. zophochir may southern end of triserialis and O. the study area. O. vs. 8, loops (9 of gut number the by separated be zophochir San Punta Juanico, from leptocephali range triserialis O. identifications leptocephali respectively). Califor- of Sur into south and Baja California Gulf the were juvenile (99.9 mm an based on early nia. Larval vafer range from Punta Marques, Baja M EASTROPAC I, station 14.069), which retained larval California. E. of Sur California Gulf the into and south and the rays, branchiostegal 22 had: pigmentation larvae miurus and xysturus, M merenda, S. are un- and count myomere a total nephros 63, myomere at of known. 150. by identified were leptocephali zophochir O. matching counts (56-62) and adult nephric myomere of habitats in ophichthids occupy a variety Adult total myomere and (57-58), counts vertebral precaudal temperate the including waters, subtropical to intertidai (146-163 and 144-162, and adult vertebral counts zone, coral the and substrates, soft shallow reefs, mid- ophichthid of kinds distinctive other Five respectively). are to sman (McCosker realm water They 1982). CalCOFI ichthyo- from examined were leptocephali in scaleless, almost and moderate-sized, cross round plankton not adults. with linked be could but samples, nostril is usually within or section. posterior The Meristic data were taken from morphological and 31 piercing 1994). Branchiostegal the (Nelson lip upper leptocephali, M and leptocephali triserialis O. 10 vafer numerous are rays mid- the at and (15--49) overlap and 1 juvenile, 21 Addi- leptocephali. zophochir O. ventral strengthens that apparatus a branchial in line information tional was taken from Fitch and ecological prey struggling hold gill presumably basket, the to (1970), McCosker Lavenberg and Rosenblatt (1968), be 1989). (McCosker et. al. The caudal fin may et. et. McCosker (1989), Leiby (1983), al. Eschmeyer or confluent with the dorsal and anal fins (Myrophinae) (1994). a!. (1989), and Nelson (Oph- a hard to reduced rays embedded containing tip 93

106 OPIDCHTHIDAE Myrophis valer MERISTICS EARLY LIFE HISTORY DESCRIPTION EGGS Mode Range Vertebrae: Shell diam.: Yolk: Total 146-155 No.ofOG: Diam.ofOG: Precaudal 58--61 60 Shell surface: Caudal Pigment: features: Diagnostic Fins: Dorsal spines 0 344 rays Dorsal LARVAE length: spines Hatching Anal 0 length: Flexion rays Anal 256 length: Transformation Pelvic 0 Pectoral Fin C, development sequence: A, 0, PI Caudal: Pigmentation: Few on upper jaw; anterior & posterior to heart; Principal 3+3 on irregular notochord; & often just below notochord on irregular Procurrent: body wall; laterally ventral lobes; few on dorsolaterally on gut; liver patchy on pterygiophores; on C pterygiophores. 0 Upper A 60 Lower (55-63); total 0 features: myomere ca. at Nephros Diagnostic myomeres (152-164); gut swellings (not 3 3 liver rakers: Gill ca 156 lobes; length <50% length; body of gut 0 usually Upper midlateral looped); 0 Lower myosepta. pigmentation not limited to 49 Branchiostegals %) in mean & (range MORPHOMETRICS HISTORY LIFE mm 38.6-98.0 of California Gulf Range: San Pedro, the to Peru, including California Sn-AlBL 41-53 45 pools Habitat: Temperate to subtropical waters; tide & shallow water to II m; benthic but not burrowing BDIBL 3-4 3 Spawning year season: caught throughout the Leptocephali 7-11 GDIBL 9 pattern: Oviparous; leptocephali planktonic ELH HLlBL 4-7 5 LITERATURE 21-34 HWIHL 27 SnLIHL 25-44 35 ORIGINAL ILLUSTRATIONS (Illustrator) 14-19x EDIHL* 16-22 mm 80.5 Larva, (N. Arthur) 17xl9 body, Head, mm sections & Arthur) (N. larva caudal 80.5 of 0.4-1 PILlBL 0.8 PdLIBL 22-33 horizontal axis Eye is slightly oval; * axis is given first, vertical second. 26 94

107 Myrophis valer Pacific eel worm - ~ ... 80.5 mm Figure Ophichthidae 1. Larva, 80.5 mm, whole body and sections showing head, nephros, and caudal regions (CaICOFI 5612, station 140G.20). 95

108 OPHICHTHIDAE Ophichthus triserialis DESCRIPTION EARLY LIFE HISTORY MERISTICS EGGS Range Mode Shell diam.: Yolk: Vertebrae: 150 No.ofOG: Diam.ofOG: 144-160 Total Shell surface: 67-68 68 Precaudal Pigment: Caudal Diagnostic features: Fins: 0 spines Dorsal LARVAE 296 Dorsal rays length: Hatching <8.1 mm 0 spines Anal length: Flexion 197 Anal rays ~9.9 to shrinking mm ~123.0 length: mm Transformation 0 Pelvic sequence: development Fin PI D, A, C, Pectoral dorsally of lower jaw; on esophagus tip near PI; 9 On Pigmentation: Caudal: gut; on ventrally patches 4-5 on ventrally gut; patches on dorsally 3+3 Principal by 10.6 2 mm, & ventrally on tail tip; dorsally postanal myomeres; Procurrent: postanal by patches; 5 13.3 mm, 4 postanal patches; by ca. 20 mm, 0 Upper ca. hypaxial myomeres; postanal subcutaneous mm, by in 27 patches 0 Lower on A pterygiophores, in myomeres, hypaxial patches 6 subcutaneous Gill rakers: myoseptum. Transformation-Series on each on & A pterygiophores. 0 Upper patch heart; 9th gut Juvenile-Internally anterior & posterior to 0 Lower 22 patches barely 6 subcutaneous nephros; at anus to posterior visible. Branchiostegals ca. at Nephros features: Diagnostic myomere (61-68); total 66 myomeres ca. 9 149 2 liver (146-158); lobes; gut loops, straightening LIFE HISTORY juvenile at pigment on loops into stage; pigmentation transformation; transformation, myoseptum; each on midlaterally C rays at enclosed including Range: of Gulf the Peru, to California, Bay, Humboldt skin, in spike-like. appear Baja California Islands; Galapagos & California north rare of %) in mean & (range MORPHOMETRICS waters; to burrows m; 23 to water shallow Habitat: subtropical Temperate in soft bottoms Juv Tr mm mm 10.6-21.3 28.4-123.0 mm 113.6 99.9 mm April, Spawning season: Leptocephali caught in June, September & Sn-AlBL 73-75 49-68 pattern: Oviparous; planktonic ELH leptocephali 74 58 50 41 LITERATURE 6-10 BDIBL 3-6 4 8 3 4 7-10 10-14 GDIBL 12 7 8 8 5-9 11-15 HLlBL ORIGINAL ILLUSTRATIONS (Illustrator) 6 6 13 9 25-32 22-27 HWIHL Walker) 99.5 mm (R. C. Larva, 28 25 26 27 body, Head, & C. caudal of Walker) sections 99.5 mm larva (R. 32-35 27-37 SnLIHL 32 34 21 16 15-17x 16-21x EDIHL* slightly oval, becoming round in juvenile stage; is horizontal axis is * Eye 18-22 16-19 second. given firSt, vertical axis 16x17 19x20 13x14 t Predorsal measured larva. on one length 11 1-3 0.4-2 PILlBL 1 0.5 3 36-56 PdLlBLt 45 51 21 36 96

109 Ophichthus triserialis Spotted eel snake 99.5 mm Figure Ophichthidae 2. Larva, 99.5 mm, whole body and sections showing head, nephros, and caudal regions (CaICOFI 5704, station 141G.29). 97

110 OPHICHTHIDAE Ophichthus zophochir MERISTICS EARLY LIFE mSTORY DESCRIPTION Range Mode EGGS Vertebrae: diam.: Shell Yolk: 154 Total 144-162 No.ofOG: Diam.ofOG: 57-58 Precaudal Shell surface: Caudal Pigment: Diagnostic features: Fins: Dorsal spines 0 Dorsal rays 258 LARVAE Anal spines 0 length: <7.5 Hatching mm 190 Anal length: Flexion rays Pelvic 0 mm ~137.2 to shrinking mm, :::147.2 length: Transformation Pectoral Fin development sequence: C, A, D, PI Caudal: heart; jaw; posterolateral & anterior lower on Few Pigmentation: to 3+3 Principal myosepta; hypaxial 1-4 dorsolaterally on liver lobes & gut loops; on Procurrent: 6 subcutaneous patches ventral to notochord; postanal A most on 0 Upper patchy pterygiophores, transformation; at notocord to ventral & dorsal tip. Lower 0 rakers: Gill Diagnostic Nephros features: at total (56-62); 58 myomere ca ca. myomeres (146-163); 157 in loops 9 loops; 0 Upper gut 8 lobes; liver 2 0 leptocephali midlaterally pigmentation transformation; before larger Lower on 27 1-4 myosepta. Branchiostegals in %) mean (range & MORPHOMETRICS mSTORY LIFE California Peru, including the Gulf Humboldt Range: Bay, of to Tr 7.5-30.6 mm north uncommon California; of Baja California mm 58.6-147.2 137.2 mm Habitat: Temperate to subtropical m; rocky & sandy areas to 64 waters; Sn-AlBL 46-52 66-77 49 72 46 burrows in soft bottoms BDIBL 3-3 5-12 season: Spawning 7 3 3 GDIBL 8-14 7-8 ELH leptocephali planktonic Oviparous; pattern: 11 7 9 8-17 HLlBL 4-6 LITERATURE 11 4 5 25-36 HWIHL 20--29 30 24 24 SnLIHL 36-40 26-37 38 30 18 ILLUSTRATIONS ORIGINAL (Illustrator) EDIHL* 15-23x 14-17x 16-27 15-19 Larva, 82.2 mm (R. C. Walker) Walker) (R. larva mm 82.2 of C. sections caudal & body, Head, 19x22 16x17 14x15 2-2 0.3-1 PILlBL 2 0.7 0.5 oval; slightly is Eye * second. vertical axis first, is given axis horizontal PdLIBL 0--0 41-49 0 43 41 98

111 Ophichthus zophochir YeHow eel snake 82.2 mm Figure Ophichthidae 3. Larva, 82.2 mm, whole body and sections showing head, nephros, and caudal regions (CaICOFI 5612, station 170.20). 99

112 CONGRIDAE: Conger eels R. CHARTER s. Congridae eel family, with three subfami- is a speciose They waters. temperate into range species few oceans; lies, ca. 1989a). 17 genera and 103 species (Smith lack vary snout The shape. in considerably and scales Smith follows 1) Nomenclature Congridae (Table The and varies from long slender to short and stout. and (in Smith (1995), and Smith Lavenberg (1989a), genera most in lip and lower genera some in lip upper fin origin dorsal The flanges. into turned are is up the California in Current occur prep.). Five species Columbia, (British atrarius Xenomystax Canada to closer pectoral the fin than to the vent; preanal region: The 20-50% Rosa (Santa cinctus Gnathophis is BL. dorsal, caudal and anal fins to Chile), Valparaiso, length dorsal and anal fins with commonly are confluent, the Island, the including Colombia, to California Gulf of 1989a). (Smith black in The caudal region edged and gilberti Ariosoma California), punctata Gorgasia almost and stiff to from ranges filamentous. stout to Panama), and Chilo conger (Baja California Sur of interest to sport are eels Garden (Heterocongrinae) Three Colombia). (Baja California Sur to obtusus, divers. other species (Bathycongrus macrurus, B. varidens, and the canabus) Heteroconger have been recorded from are unknown. are Eggs oviparous. Lepto- Congrids Gulf of California, and Heteroconger digueti, in moderate are cephali (maximum elongate to length californiensis Paraconger Rhynchoconger and nitens 300 1989f). with a rounded or pointed tail (Smith mm) California of Gulf the from range mainland Mexico, to some pigment have species oval; to round is The eye Geographic Peru and to Colombia, respectively. to above the eye. Midlateral pigment varies and below determine the for or ranges are difficult to benthic oblique row along single row, or an a to absent from fossorial adults. The dorsal fin be long or short the may myosepta. straight is a simple tube, BL). The gut (22-99% of CalCOFI the in occur species seven Leptocephali 71-99% BL. area; study A. gilberti is the most abundant and G. abundant. Larval A. gilberti, R. nitens, cinctus least the and (Table Congridae 2) morphological Meristic and as south far occur as cinctus G. and Ecuador range specimens from taken study data this in examined were northward the to Gulf of California and the outer coast 3). mor- (Table Congridae Additional ecological and gilberti and California. Northern Baja limits for of A. (1985) phological information was taken from Raju Sur, California Baja Abreojos, Punta are nitens R. and (1989f). Raju from lengths Specimen (1985) Smith and Santos, Baja California, respectively. Bahia Todos Chiloconger illustrations. his from calculated were have of G. cinctus been captured as far north as Larvae obtusus identified the from leptocephali were examina- San B. and californiensis P. Larval California. Pedro, of G. D. by identified specimen, transforming one tion California Baja Lucas, San Cabo from range macrurus larval Smith (SIO 70-362). This specimen retained range Sur, from obtusus C. of Larvae Colombia. to myomere pigmentation, and the nephros terminus count Larval to H. California, Loma, Colombia. Point of 49 matched precaudal counts of 46 and 49 from Cabo, Baja California from range San Jose canabus del of two adult specimens. Heteroconger radiographs to Gulf mainland Mexico, including the Califor- of Sur canabus by were and H digueti leptocephali identified nia; of Califor- is known only from the H digueti Gulf (72-79 matching the nephros terminus myomere counts mao vertebral precaudal respectively) and and 68-72, adult respectively). 66-69, and (70-76 counts and tropical in live grids con Adult subtropical 100

113 and Table this study (following Smith 1989a, 1995; Smith in Lavenberg, in used 1. Congridae names taxonomic Congrid in the literature. prep.) and synonyms sometimes used Other names study This (Garman 1899) Chiloconger obtusus Atopichthys dentatus Garman 1899 A. Garman 1899 obtusus Chiloconger labiatus Myers & Wade 1941 Paraconger dentatus (Garman 1899) 1899) Gnathophis 1946) (Garman Gnathophis catalinensis (Wade cinctus Rosenblatt 1974) Heteroconger canabus 1974 Rosenblatt & Cowan canabus Taenioconger & (Cowan digueti T. 1923 Pellegrin H 1923) (Pellegrin digueti Rhynchoconger 1890) (Jordan nitens Hildebrandia nitens (Jordan & Bollman 1890) Bollman & CalCOFI 2. Congridae vertebral counts for adult congrids in the Ranges study area and Gulf of California based on Table of 1972, Raju 1985, Seigel 1987, and original counts. Peden Vertebrae Species PreV Postanal Preanal CV Total 47-50 77-89 124-139 gilberti Ariosoma 98-107 30-36 128-143 macrurus Bathycongrus 109-115 38-41 147-156 varidens B. 46-49 70-71 116-120 obtusus Chiloconger 32-36 84-85 127-130 cinctus Gnathophis 140-155 punctata Gorgasia 111-123 70-76 185-199 canabus Heteroconger 115-126 66-69 184-193 digueti H 137-148 californiensis Paraconger 138-150 168-182 30-32 nitens Rhynchoconger 48-57 107-123 159-177 atrarius Xenomystax and 3. of specimens, size ranges, Congridae morphometric, Number meristic characters (range above, Table pigmentation, data). original and 1985 (Raju California of below) study Gulf CalCOFI the in leptocephali congrid for and mean area Shape Myomeres Pigmentation Size range Nephros No. of Eye Midlateral Eye Tail (rnm) specimens terminus Total Species rounded none oblique round 16.5-227.2 66-71 126-138 23 Ariosoma rows trans.) 68 (3 134 gilberti row 135-148 crescentic patch single pointed oval 36.0-146.7 20 46-50 Bathycongrus 141 over and under 48 macrurus none single row round rounded 121-124 41.4-128.3 50-57 14 Chiloconger 52 (1 123 trans.) obtusus oval 20.1-122.5 crescentic patch none II 42-46 126-137 rounded Gnathophis over and under 44 130 cinctus off- row rounded single 34.1-122.6 72-79 189-201 none 21 Heteroconger round (2 trans.) 75 196 canabus off- 68-72 rounded none single row 37.3-101.5 189-196 7 digueti H round 69 193 137-148 57-61 46.3-133.3 none single row round rounded 18 Paraconger californiensis 59 143 row single 44.8-126.3 46-49 patch 20 crescentic oval pointed 169-187 Rhynchoconger below nitens 48 177 101

114 CONGRIDAE Ariosoma gilberti DESCRIPTION HISTORY LIFE MERISTICS EARLY Range Mode EGGS Yolk: Vertebrae: diam.: Shell 124-139 No.ofOG: Total Diam.ofOG: surface: Shell 47-50 Preanal Pigment: 77-89 Postanal Diagnostic features: Fins: Dorsal spines 0 LARVAE rays Dorsal length: Hatching spines Anal 0 length: Flexion Anal rays Pelvic 0 Transformation length: mm shrinking to ~139.4 mm '

115 Ariosoma gilberti Gilbert's conger ca. 78.0 mm ca. 70.0 mm Figure Congridae 1. Larva, ca. 78.0 mm, whole body and sections showing head and caudal regions; late transformation specimen, ca. 129.0 whole body and head; juvenile, ca. mm, 70.0 mm, whole body, head and body section (Raju 1985). 103

116 CONGRIDAE Bathycongl'US macl'UI'US MERISTICS DESCRIPTION EARLY LIFE HISTORY EGGS Mode Range Vertebrae: Shell diam.: Yolk: No.orOG: 128-143 Total Diam.ofOG: surface: Shell 30-36 Preanal Pigment: 98-107 Postanal features: Diagnostic Fins: o Dorsal spines LARVAE Dorsal rays Hatching length: o Anal spines Flexion length: rays Anal Transformation iength: o Pelvic Fin PI development C, D, A & sequence: Pectoral Caudal: eye; melanophores 1-2 below & over patches Crescentic Pigmentation: midlaterally; 2 over heart; series Principal paired series ventrally operculum; on along gut; on bases. D, C, A Procllrrent: myomere features: (46--50); 48 total ca. at Nephros Diagnostic Upper (135-148); 141 elongate body; crescentic pigment ca. myomeres Lower patches over eye; & pointed under tail. Gill rakers: Upper o MORPHOMETRICS %) in (range & mean o Lower Bnmchiostegais 36.0-146.7 mm LIFE HISTORY 90--95 Sn-AlBL 92 of recorded Range: Cabo Gulf San California; larvae Baja from Lucas, Sur California 4--5 BD/BL 5 Habitat: Nearshore tropical waters GDIBL 10-13 12 season: Larvae collected in February, May, June, August, & Spawning HLIBL 5-8 July; October (maximum in June); transformation individuals collected in 6 juveniles in collected January 29-38 HWIHL 35 planktonic leptocephali ELH pattern: Oviparous; SnLIHL 28-36 31 LITERATURE 19-24x ED/HL* 22-27 1985 Raju 22x25 1-2 Eye * second. axis vertical first, given is axis horizontal oval; is I PdLIBL 66-83 71 104

117 Shorthead conger macrurus Batnycongrus &::::: ···=··t,;,i ... ... ~~ ... mm ca. 70.0 \ ... ...•••••...•.•..•••.••..•.•...•••.••... ... ~.:'~ .. ;~ ... :.;-:. 0 • ~ u =.~ ... ... · ... u. \i~ ~.= 143.0 mm ca. i}i2 ------~ mm 68.0 ca. specimen, Congridae 2. Larva, ca. 70.0 mm, whole body and sections showing head, midbody and caudal regions; transformation Figure 143.0 68.0 whole body and head; juvenile, ca. mm, mm, whole body and head (Raju 1985). ca. 105

118 CONGRIDAE Chiloconger obtusus MERISTICS EARLY LIFE HISTORY DESCRIPTION Range Mode EGGS Vertebrae: Shell diam.: Yolk: Total 116-120 No.ofOG: Diam.ofOG: Precaudal 46-49 Shell surface: 70-71 Caudal Pigment: Fins: features: Diagnostic 0 Dorsal spines rays Dorsal LARVAE Anal spines 0 Hatching length: Anal rays length: Flexion Pelvic 0 length: Transformation mm <110.0 to shrinking mm >128.3 Pectoral Fin development sequence: C, D, A & PI Caudal: ventral Pigmentation: Evenly spaced series on lateral midline; dorsal & 4-5+3 Principal A 3-4 bases. over heart; 2 on lower jaw; on D, C & on gut; series Procurrent: features: Nephros Diagnostic total myomere (50-57); at ca. 52 Upper myomeres absence teeth; compressed laterally (121-124); 123 ca. of Lower Paraconger- pigment crescentic under eye; compared to patch Gill rakers: midlateral and more count, myomere lower regular pigment, relatively Upper mm). «130.0 relatively smaller size Lower o o Branchiostegals in mean & (range MORPHOMETRICS %) LIFE HISTORY Tr 41.4-128.3 mm mm 110.0 Sur to Colombia Range: Tosco, Punta California Baja Sn-AlBL 88-91 90 52 Habitat: subtropical to tropical Shallow waters BDIBL 5-7 Spawning season: 6 6 GDIBL 10-15 pattern: Oviparous; ELH leptocephali planktonic 12 12 HLlBL 6-8 LITERATURE 7 8 26-33 HWIHL 1899 Garman 28 32 1985 Raju SnLIHL 27-35 32 16 (Illustrator) ILLUSTRATIONS ORIGINAL EDIHL* 16-19 17 15x19 mm Arthur) Larva, 94.1 (N. larva mm Arthur) (N. 94.1 of Head, sections body, & caudal 1-3 2 2 PdLlBL 59-74 transformation oval becomes in specimens; horizontal axis is given * Eye 64 24 second. first, vertical axis 106

119 obtusus Chiloconger Thicklip conger O' ... . ... 94.1 mm DDDIJ Figure Congridae 3. Larva, 94.1 mm, whole body and sections showing head, nephros and caudal regions (CaICOFI 6001, station 143.55); mm 96.0 (Raju 1985 as Paraconger californiensis). (below head) from larva ca. teeth 107

120 CONGRIDAE Gnathophis cinetus EARLY LIFE HISTORY DESCRIPTION MERISTICS Mode Rallge EGGS Shell Vertebrae: diam.: Yolk: No.orOG: 128 Total 127-130 Diam.orOG: 32 surface: Precaudal 32-36 Shell Pigment: 84-85 84-85 Calida I featnres: Diagnostic Fins: 0 Dorsal spines LARVAE 205 Dorsal rays 202-207 Hatching length: Anal spines 0 length: Flexion Anal 148 139-162 rays to shrinking mm ~123.l 0 length.: Transformation mm ::;122.5 Pelvic Pectoral PI' & A D, C, Fin development seqllence: & above patch Crescentic Pigmentation: heart; over patch eye; below Calldal: 3+3 margin series on ventral bases. D posterior & A, C on gut; of Prillcipal ProCUITent: Nephros at ca. myomere features: (42--46); total Diagllostic 44 ca. myomeres 130 Upper blllnt absence pigment; midlateral of (126-137); & below above patch pigment eye; transformation crescentic tail; Lower a blunt, snout. develop individuals hook-like rakers: Gill o Upper o (range & meall in %) MORPHOMETRICS Lower Branch.iostegals Tr HISTORY LIFE mm 20.1-123.1 mm 122.5 Sn-A/BL Island, to California Catalina Range: of Gulf Colombia, including 90-94 92 75 & Galapagos Islands California BDtBL 5-7 to subtropical shallow Temperate water; benthic in sand Habitat: 6 5 GDIBL 12-16 season: Spawning l3 12 6-10 HLIBL planktonic Oviparous; pattem: ELM leptocephali 8 7 HwtHL 29-35 LITERATURE 32 34 35-42 SnLlHL 1985 Raju 38 30 16-19x EDIHL* 19-25 axis horizontal oval; slightly is Eye * axis second. vertical first, given is 18x21 19x22 0.8-2 I PdLIBL 73-79 76 56 108

121 Catalina conger cinctus Gnathophis 6.7 mm ca. ca. 106.0 mm ca. 142.0 mm Figure Congridae 4. Head of early larva, head length ca. 6.7 mm; larva, ca. 132.0 mm, whole body, head and caudal regions; transformation specimen, ca. 106.0 mm, whole body and head; juvenile, ca. 142.0 mm, whole body and head (Raju 1985 as Gnathophis catalinensis). 109

122 CONGRIDAE Heteroconger canabus EARLY LIFE HISTORY DESCRIPTION MERISTICS Range EGGS Mode Sheil diam.: Yolk: Vertebrae: 185-199 No.ofOG: 194 Total Diam.ofOG: 74 snrface: 70-76 Shell Precaudal Pigment: 111-123 Caudal Diagnostic features: Fins: o Dorsal spines LARVAE 644 Dorsal rays Hatching length: o Anal spines Flexion length: 460 rays Anal mm :::; o Transformation length: >123.0 mm shrinking to 86.0 Pelvic 9-18 Fin & A, 13 PI development seqllellce: C, D, Pectoral series Pigmentation: Several over heart; dorsal on lateral midline; on Caudal: 3+3 of D posterior & A C, on gut; along ventrally series gut; surface Principal bases. Procurrent: total myomere 75 (72-79); ca. features: Diagnostic Nephros at Upper small conical teeth; rounded tail; (189-201); 196 myomeres ca. Lower compared to of maxillary, lateral midline H on diguetf-lack pigment rakers: Gill at snout, relatively relatively begins pigment myomere longer 16-22, o Upper relatively larger eye, larger head. o Lower Branchiostegais in mean & (range MORPHOMETRICS %) HISTORY LIFE Tr 34.1-122.6 mm off mm of California; larvae Baja collected Gulf San Jose del Cabo, 86.3-95.7 Range: California Sur Sn-AlBL 71-85 42-46 78 44 Burrows in depth); sandy bottoms (generally <60 m shallow, Habitat: water tropical currents moderate in clear, BDIBL 5-5 4-6 5 5 February & October Larvae collected in Spawning season: GDIBL 10-14 9-10 11 10 leptocephali planktonic Oviparous; pattern: ELH HLlBL 5-10 7-7 7 7 LITERATURE HWIHL 20-36 38-41 29 40 1985 Raju SnLlHL 29-40 17-19 18 36 (Illustrator) ORIGINAL ILLUSTRATIONS 17-22x EDIHL* 20-23 18-25 22 CR. mm Walker} Larva, C. 102.3 Head, body, larva mm & caudal Walker) C. (R. sections 102.3 of 19x22 1-1 0.6-3 PILIBL 2 1 horizontal axis is * Eye is oval, becoming round in transformation stage; PdLlBL 24-54 13-14 given axis vertical first, second. 37 13 110

123 Cape eel Heteroconger canabus garden ca. 103.0 mm Figure Congridae 5. Larva, 102.3 mm, whole body and sections showing head, nephros, and caudal regions (SIO 73-249); early juvenile, ca. 103.0 mm, (Raju 1985 as Taenioconger canabus). 111

124 CONGRIDAE Heteroconger digueti DESCRIPTION HISTORY LIFE EARLY MERISTICS EGGS Mode Range Vertebrae: diam.: Shell Yolk: 184--193 Total No.ofOG: 186 Diam.ofOG: Shell 66-69 Precaudal surface: 68 Pigment: 115-126 Caudal Diagnostic features: Fins: Dorsal spines 0 647 LARVAE Dorsal rays Hatching length: 0 spines Anal Flexion length: 346 rays Anal Transformation length: 0 Pelvic D, C, sequence: P Fin development & A, Pectoral J Caudal: heart; over Several Pigmentation: lateral on series maxillary; 3 on 3+3 gut; A C, on gut; of Principal series ventrally along surface dorsal midline; on & posterior D bases. Procurrent: ca. features: Diagnostic myomere 69 (68-72); total at Nepbros Upper ca. myomeres conical teeth; (189-196); tail; 193 small rounded Lower canabm-pigment maxillary, midline H lateral on to compared rakers: Gill pigment snout, relatively shorter relatively 11-15, at myomere begins o Upper head. smaller relatively smaller eye, o Lower Branchiostegals & %) MORPHOMETRICS (range in mean HISTORY LIFE mm 37.3-101.5 California of Gulf Range: to de Banderas, Mexico Bahia 71-83 Sn-AlBL 77 in m); Burrows shallow, sandy bottoms (generally <60 clear, Habitat: water tropical in moderate currents BDIBL 4--6 5 Larvae Spawning season: collected in February October & GDIBL 9-15 11 ELH planktonic Oviparous; leptocephali pattern: 5-8 HLlBL 6 LITERATURE 26-30 HWIHL 28 Raju 1985 31-38 SnLIHL 34 ILLUSTRATIONS ORIGINAL (Illustrator) 17-20x EDIHL* 20-24 Walker) C. (R. mm 101.5 Larva, body, Head, larva (R. C. Walker) & caudal sections of 101.5 18x21 1-2 PJLlBL 2 horizontal oval; is Eye * axis second. vertical first, given is axis 22-35 PdLIBL 27 112

125 Heterocongel'digueti Cortez eel garden 101.5 mm ca. 102.0 mm Figure Congridae 6. Larva, 101.5 mm, whole body and sections showing head, nephros, and caudal regions (SIO 73-249); early juvenile, 102.0 head whole body and mm, (Raju 1985 as Taenioconger digueti). ca. 113

126 CONGRIDAE Paraconger californiensis LIFE DESCRIPTION HISTORY EARLY MERISTICS EGGS Mode Range Yolk: Shell diam.: Vertebrae: 137-148 139 Diam.ofOG: No.ofOG: Total surface: Shell Precaudal Pigment: Caudal features: Diagnostic Fins: o Dorsal spines LARVAE Dorsal rays length: Hatching o spines Anal Flexion length: Anal rays length: Transformation o Pelvic Fin sequence: development PI 15-18 D, & A C, 16 Pectoral lateral on Series Pigmentation: along ventrally series paired midline; Caudal: gut; on over 4+3 D, C & A bases; 3-4 heart. Principal Diagnostic Nephros at ca. features: myomere (57-61); total 59 Procurrent: myomeres ca. 143 compressed teeth; (137-148); crescentic of absence Upper eye; pigment. pigment patch under midlateral Lower Gill rakers: in mean MORPHOMETRICS %) (range & o Upper o Lower Branchiostegals 46.3-133.3 mm LIFE mSTORY Sn-AlBL 85-97 93 to Colombia, including San Baja California Cabo Lucas, Range: Sur 4-5 BDIBL southern Gulf of California 4 8-11 GDIBL tropical Habitat: Shallow depth 33-50 & subtropical waters; m 10 6-9 HUBL Larvae Spawning season: collected August in all months & except 7 May); transformation individuals in January & December (maximum in HWIHL 24-30 April; juveniles in March 27 leptocephali Oviparous; pattern: ELH planktonic SnLIHL 29-44 38 LITERATURE EDIHL 13-19 16 1985 Raju 1-2 PILlBL I PdLIBL 76-94 86 114

127 californiensis Paraconger California conger 96.0 ca. mm DDDl) 97.0 ca. mm U~"======~ ca. 85.0 mm Figure Congridae 7. Larva, ca. 96.0 mm, whole body and sections showing head, midbody, and caudal regions and enlargement of teeth nead, midbody, mm, whole body and sections showing 97.0 and caudal regions; juvenile, ca. 85.0 mm, whole ca. below; juvenile, early body (Raju 1985). 115

128 CONGRIDAE nitens Rhynchoconger DESCRIPTION mSTORY LIFE EARLY MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: 168-182 No.ofOG: Total Diam.ofOG: surface: Shell 30-32 Preanal 138-150 Pigment: Postanal Diagnostic features: Fins: 0 spines Dorsal LARVAE rays Dorsal length: Hatching 0 Anal spines Flexion length: Anal rays Transformation length: 0 Pelvic Fin PI development C, D, A & sequence: Pectoral Crescentic Pigmentation: 1-3 eye; below Caudal: patch on maxilla; 2-3 at jaw 4+4 ventral Principal paired articulation; 3-4 over heart; series below lateral midline; series A, vent; to isthmus from on bases. & C D, Procurrent: features: Diagnostic myomere Nephros at ca. total 48 (46-49); Upper 177 (169-187); crescentic pointed elongate body; tail; ca. myomeres Lower patch; eye on midlateral maxilla; dots pigment 1-3 pigment rakers: Gill o Upper & mean in %) MORPHOMETRICS (range o Lower Branchiostegals 44.8-126.3 mm mSTORY LIFE Sn-AlBL 83-93 89 Gulf of California to Gulf of Tehuantepec, Mexico; Northern Range: Pacific Todos Bahia to coast Santos, Baja northward range larvae along 4-6 BDIBL California 5 GDIBL 9-12 Shallow temperate tropical & Habitat: waters II 5-8 HLlBL season: December except months all in collected Larvae Spawning 6 (minimum October, maximum in May); transformation individuals in in HWIHL 30-41 January, May, June, October; juveniles in March, & July 35 pattern: Oviparous; planktonic leptocephali ELH 28-38 SnLIHL 34 LITERATURE 18-23x EDIHL* 22-28 Raju 1985 20x24 1-2 1 is oval; Eye * second. vertical first, is given axis horizontal axis PdLIBL 38-51 45 116

129 nitens Rhynchoconger Bignose conger ... = ... = ... = ... = ... ~ 6?:~:I«===rt]f==r=iJ = ... 123.0 mm ca. o ~ 120.0 mm~~~~~~~~===~~=========:::::::::::::::::::::::::::::::::::::::::==~~ ca. ca. mm 91.0 •••.• , ,ret".,I'&"'£' Figure Congridae 8. Larva, ca. 123.0 mm, whole body and sections showing head, nephros, and caudal regions; transformation specimen, ca. body mm, whole body and head; juvenile, ca. 91.0 mm, whole 120.0 and sections showing head and midbody (Raju 1985 as Hildebrandia nitens). 117

130

131 DERICHTHYIDAE: eels Longneck s. R. CHARTER the temperate live throughout regions of Derichthyids Derichthyids are oviparous, with planktonic eggs. mesopelagic the Indian in Atlantic, Oceans Pacific, and Derichthys serpentinus ovaries described Beebe (1935) m zones from 457 to 1829 and bathypelagic multi- indicating eggs of two which batches contained genera two Presently, al. and 1983). (Eschmeyer et. mm in average eggs Planktonic spawning. ple 2.33 Only three are recognized this family. in species a wide have and perivitelline diameter space, segment- Derichthys region, CalCOFI the in occurs serpentinus globule(s) and yolk, ed (Stibane 1983). oil and ranging from southern California to Chile (Fitch not abundant and 1968). The adults are Lavenberg Derichthyid are small to moderate in leptocephali regular in collected been not have larvae the CalCOFI moderately and mm) (60-90 deep-bodied length gut BL). The liver is undivided (13-24% the is a and samples. ichthyoplankton in The tube. straight simple dorsad turns generally tail dorsal fin origin preserved The specimens. scar-like striations by recognized easily are Adults is at mid- on The area. neck the in a constriction and head the during changing BL), (43-59% body transformation to cephalic sensory system has large pores that are the adult position in BL). (20-30% Pigmentation sometimes large, for nostrils, especially the mistaken Derichthys leptocephali is limited to minute serpentinus anteriorly directed (Castle snout the on one tubular melanophores on the midlateralline. just below the tail 1970). gill slits are nearly horizontal and posi- The larval from distinguished be may larvae serpentinus D. anteroventrally tioned The vertical fins. pectoral the to terminus, nephros the of position the by serrivomerids are fins contiguous, the dorsal and anal fins with respectively. 34-39, myomeres at versus 58-65 the fin dorsal The area. caudal abruptly reduced in information was taken and larval, Egg, ecological at 20-30% BL. Derichthys serpentinus is ca. is origin 1935; from the literature (e.g., Beebe Karmovskaya specimen fresh A scaleless. olive mouse to tawny was of meristics and Morphometrics 1989h). Smith 1986; preserved material is dark brown dorsally while gray, brown on the abdomen (Beebe 1935; Kar- and light serpentinus D. taken from detailed examination of were oval, opaque, an have serpentinus D. 1986). movskaya specimens (8.1-43.6 mm) 16 two transfor- larval and whitish patch in each web anal of the fins, dorsal and mation mm). 67.0 and (65.1 specimens is lost in preservation (Beebe possibly luminescent; this 1935). 119

132 Derichthys serpentinus DERICHTHYIDAE EARLY LIFE HISTORY DESCRIPTION MERISTICS EGGS Mode Range Shell diam.: 2.3 mm Segmented Yolk: Vertebrae: of Multiple OG: No. 133 Diam.ofOG: 126-136 Total surface: Shell 56 53-57 Precaudal Pigment: Caudal segmented space; perivitelline wide size; Large features: Diagnostic Fins: yolk. 0 spines Dorsal 244 226-262 Dorsal rays LARVAE 0 Anal spines length: mm >5.5 Hatching 159 155-180 Anal rays length: Flexion 0 Pelvic Transformation length: ca. 55.0 mm 13 11-18 Pectoral PI Fin development sequence: D, A, C, Caudal: through series postanal stages; all (minute), Similar Pigmentation: just 10 9-11 Principal below the lateral midline. Procurrent: ca. (5~5); total myomere 62 Nephros features: Diagnostic at 0 Upper ca. 134 leptocephali; moderate-sized (131-137); caudal myomeres 0 Lower region curved dorsad pigmentation. distinctive specimens; in preserved Gill rakers: 0 Upper (range & MORPHOMETRICS %) in mean 0 Lower 7 6-8 Branchiostegals Tr HISTORY mm LIFE 8.1-8.8 mm 10.6-43.6 65.1-67.0 mm 78-78 Sn-AlBL 74-83 50-58 Range: Worldwide in warm & temperate seas; eastern Pacific from 78 54 79 Chile to California southern 10-l2 BDIBL 6-12 6-8 m & 11 depth bathypelagic, 500-2000 Habitat: Meso- 9 7 18-19 GDIBL 15-16 16-24 Spawning season: 19 16 19 16-16 10-10 HUBL 10-15 planktonic leptocephali pattern: ELH & eggs Oviparous; 16 10 12 22-31 28-38 36-38 HWIHL LITERATURE 34 37 27 22-22 28-33 31-33 SnLIHL Beebe 1935 32 22 30 Castle 1970 1986 Karmovskaya 27-30x 14-14x 20-26x EDIHL* 31-34 21-33 12-16 Smith 1989h 1979, 1983 Stibane 28x33 23x28 14x14 3-3 1-3 3-3 3 2 3 is Eye * second. vertical first, axis given is axis horizontal oval; slightly 45-51 PdLIBL 43-59 21-29 48 52 25 120

133 Derichthys serpentinus Narrow neck eel ... , . ... ~ 7mm ~==--- 42mm I ••• ... 54mm ---c-- -~ €f_® 1 ~ ~L~=~=- '7 7 ~- 1 ___ m~m • o 0 o 0 Figure Derichthyidae 1. Early larva, 7 mm, head and whole body (Castle 1970); larva, 42 mm, whole body and sections showing head 71 mm, head and body; transformation specimen, 54 mm, head and whole caudal late transformation specimen, and region (Smith 1979); whole body (Castle 1970). 121

134 NEMICHTHYIDAE: Eels Snipe CHARTER R. S. nemichthyid Four CalCOFI the in occur species study 4-11 slender 400 mm) and with greatest depth % BL. area Nemichthyidae (Table 1): bowersii,A. Avocettina straight gut is a simple and is undivided liver the The and N. scolopaceus (Niel- infans, Nemichthys larseni, tube caudal The is attenuate or BL. 89-96% region melanopho- of most com- scolopaceus, N. the Smith 1978). sen and filiform. A characteristic internal series trawl collections, ranges from res mon species in midwater notochord. above lies the Canada, Columbia, British to the Gulf including Peru Current Nemichthyid leptocephali in the California A. infans, the second most common of California. from species, Canada, occurs British to Columbia, region separated into two groups (Table may be Nemichthys has a filiform tail and 2): Nemichthyidae central including the Gulf Mexico of California. The has while ca. attenu- myomeres of N. larseni is similar to that of A. infans an Avocettina 300-400 distribution ate tail and ca. 200 and myomeres. scolo- N. N. larseni Francisco, and A. bowersii ranges from San California, sympatric and Pacific eastern have the in are pace us to Peru. precaudal and (Nielsen overlapping counts vertebral Nemichthys leptocephali (11.0- Smith Ten 1978). recognized mesopelagic are nemichthyids Adult eels are to identical 184.8 mm) examined in this study lower scaleless, elongate bodies with upper and by bodies are jaws Their other. each from away curve that Castle by scolopaceus those N. as described (1965a) compressed and their anal are higher than their fins and are un- Smith (l989i). N. larseni leptocephali dorsal Mature males undergo a transformation, fins. have infans Avocettina A. and non- bowersii known. of their jaws, and at one time a shortening most visibly overlapping counts (59-66 vertebral versus precaudal in different conspecific from genera placed were bowersiileptocephali examined in this study A. 69-72). species Nemichthys (Nielsen 1978). Smith and females had nephros at terminus the ca. (60--64), 62 myomere filamentous tails, line have consisting a lateral of three 5-12 melano- of consisting patches pigment lateral and of pores, and may have >750 vertebrae. Avocet- whereas 80-84, rows at myomeres 46-51, and phores 24-25, tina line with tails, attenuate have species a a lateral nephros A. the had infans myomeres ca. at terminus 71 Although vertebrae. 200 ca. and pores, of row single 1-4 and of consisting patches pigment (69-74) lateral depths at caught usually scolopaceus N. m, >300 have at myomeres melanophores 97-100. and 28-31,59-61, California of Gulf the tows surface in caught been in to those similar are A. infans larvae in this study 1966). Fitch (Lavenberg and gut in Differences (l989i). Smith by described pig- in mentation noted are morphometry and the species unknown. are eggs planktonic oviparous; are Adu.lts descriptions. length (maximum long are Leptocephali bodied 122

135 Table Nemichthyidae Meristic characters for the nemichthyid species in the California Current vicinity (Nielsen and Smith 1. and The are often broken in adult Nemichthys; counts for the dorsal filaments anal fins are the numbers of rays 1978). caudal to the anterior vertebra (those counts referred to as D 1978) and A Smith, by 201st and Nielsen 200 200 Vertebrae Fin rays PrCv CV Total D Species A 59-66 124-129 177-195 233-333 208-297 bowersii Avocettina 116-131 185-203 69-72 279-432 A. 240-372 irifans 79-86 164-218 173-222 Nemichthys larseni 77-105 170-253 scolopaceus 186-273 N. Table Nemichthyidae 2. Morphology and pigmentation of nemichthyid leptocephali examined in this study. Relative positions of the end of the liver, the nephros terminus, and vent (preanal), are indicated by the corresponding myomere number posterior and head. counting posteriad from (range Also listed for each species are the number mean), of melanophores the lateral in patches of myomeres. to the respect with the patches position and Myomeres Pigmentation Total Number Myomeres Nephros Liver Preanal Species & 187-191 123-158 80-84 5-12 24-25,46-51, 60-64 Avocettina bowersii 141 189 26 62 97-100 & 69-74 59-61, 187-198 1-4 28-31, 163-167 A. irifans 165 194 30 71 1-5 275-354 134-251 & 81-88 103-123 69-81, 18-23,35-44, 39-47 Nemichthys scolopaceus 210 315 41 85 123

136 NEMICHTHYIDAE A vocettina bowersii DESCRIPTION EARLY LIFE HISTORY MERISTICS Mode EGGS Range lIiam.: Shell Vertebrae: Yolk: No.ofOG: 177-195 Total Diam.ofOG: surface: 59-66 Shell Precaudal 124-129 Pigment: Caudal features: Diagnostic Fins: 0 spines Dorsal 233-333 LARVAE rays Dorsal Hatching length: 0 spines Anal 208-297 length: Flexion Anal rays 0 Transformation length: Pelvic 15-19 C, & PI Fin development sequence: D & A Pectoral attenuate Dorsally three notochord; on each) Pigmentation: below (5-12 clusters Caudal: dorsally gut, on 46-51, 80-84; & midline lateral 24-25, at myomeres Principal past liver; heavier ventrally on gut, heavier before liver. Procurrellt: myomere at ca. nephros 26; myomere at ca. Liver features: Diagnostic Upper tail; myomeres ca. 189 attenuate (187-191); total (60-64); 62 Lower compared A. longer snout, to & infans--relatively smaller eye head rakers: Gill 0 at & position pigment lateral of less number (5-12) depressed, Upper 0 ventral & & pigment dorsal 24-25, 46-51, & 80-84, gut myomeres Lower anus. to heart from Branchiostegals %) MORPHOMETRICS (range & mean in HISTORY LIFE San Francisco, California to Peru; offshore Pacific mm 155 Eastern from Range: 88.7 mm* 0 150 W to Sn-A/BL 87 93 waters; collection Habitat: tropical & temperate warm Mesopelagic in m 92-641 depths BDIBL 8 3 GDIBL Spawning season: 15 7 HLIBL changes ELM pattern: Oviparous; degenerative in females & males 11 4 planktonic leptocephali suggests semelparity; HWIHL 31-33 34 32 LITERATURE SnLIHL 44-46 44 45 EDIHL 19-20 21 20 (Illustrator) ILLUSTRATIONS ORIGINAL PjLIBL 3 Arthur) (N. mm 88.7 Larva, PdLIBL body, Head, (N. Arthur) & caudal sections of 88.7 mm larva 0 99 specimens calculations. One of two available * ripped, not usable for BL 124

137 A vocettina bowersii ,';-",,, 88.7 mm Figure Nemichthyidae 1. Larva, 88.7 mm, whole body and sections showing head, gastric, and caudal regions (EASTROPAC I, station 12.51). 125

138 NEMICHTHYIDAE Avocettina in/ans EARLY LIFE HISTORY DESCRIPTION MERISTICS Mode EGGS Range Shell diam.: Yolk: Vertebrae: 193 No.ofOG: 185-203 Total Diam.ofOG: surface: Shell 71 69-72 Precaudal Pigment: 120 116-131 Caudal features: Diagnostic Fins: 0 spines Dorsal LARVAE 279-432 339 rays Dorsal length: Hatching 0 spines Anal length: Flexion 299 240-372 Anal rays length: Transformation Pelvic 0 14--18 16 & D sequence: development A C, Fin & PI Pectoral dots attenuate Pigmentation: Dorsally on notochord; three clusters (1-4 each) Caudal: at myomeres midline below lateral dorsally 97-100; & 28-31,59-61, Principal from gut along liver. to heart from gut along ventrally liver; Procurrent: 30; nephros at ca. myomere myomere Liver features: Diagnostic at ca. Upper total (69-74); 194 myomeres length moderate ca. 71 (187-198); Lower large attenuate relatively head; 150 mm); depressed tail; (maximum rakers: Gill eye 0 & short snout; at (1-4) & position of lateral pigment number Upper at & 97-100; dorsal gut pigment starting 0 59-61, 28-31, myomeres Lower at ending liver. ventral liver; pigment gut Branchiostegals in MORPHOMETRICS (range & mean %) HISTORY LIFE S; mm 46.1--62.9 oceans Range: All north of 20° in the eastern Pacific from Queen Canada Islands, Charlotte to central Mexico, including Columbia, British Sn-A/BL 94-96 Gulf the California of 95 m m depth; usually 1200 to 2000 Surface depth to 2000 Habitat: BDIBL 4--5 5 season: Spawning 8-10 GDIBL 9 females ELH pattern: Oviparous; degenerative changes in males & HLIBL 5-7 planktonic suggest leptocephali semelparity; 6 30-34 HWIHL LITERATURE 32 SnLIHL 40-41 Castle 1965a 41 Smith & 1978 Nielsen 1989i Smith EDIHL 23-24 24 1-1 PILIBL 1 PdLIBL 90-91 90 126

139 Avocettina infans Blackline eel snipe 20.0 mm 45.0 mm 111.0mm Figure Nemichthyidae 2. Larvae, 20.0 mm, 45.0 mm; larva, 111.0 mm, whole body and sections showing head, gastric, and caudal regions (Smith 1979). 127

140 NEMICHTHYIDAE Nemichthys scolopaceus HISTORY DESCRIPTION EARLY LIFE MERISTICS Mode EGGS Range diam.: Shell Yolk: Vertebrae: No.ofOG: Diam.ofOG: Total* 77-105 surface: Shell Precaudal Pigment: Caudal Diagnostic features: Fins: 0 Dorsal spines LARVAE 170-253 Dorsal (Dz rays )* •• length: Hatching 0 Anal spines 186-273 Flexion length: Anal rays (Azoo)* 0 Transformation length: Pelvic sequence: development Fin PI C, & A & D 10-14 Pectoral clusters each) dots (1-5 Dorsally four notochord; on Pigmentation: Caudal*: atmyomeres midline below lateral 103-123; 18-23,35-44,69-81, & reduced to filament Principal dorsally on gut; D & A fin bases; larvae >120.8 have ventral mm Procurrent: patch posterior to heart. Upper at Liver features: Diagnostic myomere ca. myomere 41; nephros at ca. Lower total 305 (230-354); (81-88); number (1-5) & 85 ca. myomeres Gill rakers: position & 69-81, 35-44, 18-23, at of pigment myomeres lateral 0 Upper body 260 ca. length maximum 103-123; mm. tail; filamentous 0 Lower 7-15 Branchiostegals MORPHOMETRICS in mean (range & %) LIFE HISTORY mm 11.0-32.4 mm 49.6-184.8 eastern Pacific Range: tropical & temperate seas; Cosmopolitan in the in Columbia, British from to Canada Peru, including the Gulf of California Sn-AlBL 86-94 79-91 92 86 storms; during eastern ashore Habitat: Mesopelagic; occasionally stranded BDIBL 5-7 2-4 depths collection Pacific 24-4337 m, usually <500 m 3 6 9-11 4-9 GDIBL Spawning season: 10 6 lllJBL 3-6 7-11 in males females & changes degenerative Oviparous; ELH pattern: 9 4 leptocephali suggest semelparity; planktonic 33-39 28-33 HWIHL 30 35 LITERATURE SnLIHL 36-43 44-50 41 47 Crane 1937 & Beebe 1965a Castle 23-29x 19-22 EDlHLt 22-25 20 Nielsen & Smith 1978 Smith 1989i 25x24 0.9-2 0.2-0.7 PILlBL 2 0.4 Table See 1 for explanation. Nemichthyidae * PdLIBL 0-0 84-93 first, is axis given oval have larvae t Small horizontal eyes; slightly 90 0 second. vertical axis 128

141 Slender snipe eel scolopaceus Nemichthys 15.0 mm 182.0 mm "'ill ~I'"'''''' Figure Nemichthyidae 3. Larvae, 15.0 mm, 72.0 mm; larva, 182.0 mm, whole body and sections showing head, gastric region, and caudal (Smith 1979). region 129

142

143 SERRIVOMERIDAE: eels Sawtooth S. R. CHARTER tropical Serrivomerids occur and temperate the in planktonic oviparous; is S. sector un- are eggs regions nearly all oceans. Tighe (1989a) recognized of known. size (>63.7 mm in moderate are Leptocephali Serrivomer family. species 10 the in two genera and deep-bodied before transformation) and relatively known species CalCOFI the only the in occur to sector, (16-24% is gut the a is liver The BL). undivided, central California from to Chile ranges area, study and simple straight tube, and the tail The dorsal is pointed. BL. (62-81%) 75% ca. 1000 m, although irregular An > depths at collected is usually fin is at origin specimens after the near collected been surface have just minute below the of melanophores is present series at beginning midline, lateral BL. 50% ca. Other Lavenberg Adults are uncom- sunset 1968). (Fitch and collections and larvae are rare trawl midwater in mon is pigmentation the dorsal posterior the on of portion samples. single A ichthyoplankton CalCOFI S. in sector S. eye. fin on and base, fin anal the on base, the California of Gulf the in collected specimen jesperseni serpentinus from distinguished Derichthys be may by (1989) aL reported is held by LACM, and Matarese et nephros anterior more its 34-39 myomeres (at terminus the CalCOFI study area. of north them a more posterior respectively), 58-65, dorsal fin vs. midlateral of preanal pigmen- origin, presence and the tation. Fresh specimens of are Serrivomer colored; bronze preserved specimens (Tighe in pigment this persist may 1989a). attenuate a with jaws moderately has sector S. meristics and Morphometries sector S. of were 50 of larval brown, slender, scaleless body. The dorsal and dark examination detailed from obtained collected the of west largely mm), (4.7-63.7 specimens with anal and fin caudal the confluent are fins anal the survey fin. S. CalCOFI adults on information Additional area. dorsal the than higher and is more fin anterior (1968). and Fitch from taken was Lavenberg large, is sector easily recognized compressed by the vomer. teeth on the 131

144 SERRIVOMERIDAE Serrivomer sector MERISnCS EARLY LIFE HISTORY DESCRIPTION EGGS Mode Range Shell diam.: Vertebrae: Yolk: 153 Total No.ofOG: 151-153 Diam.ofOG: 36 35-36 caudal Shell surface: Pre 117-118 Pigment: 117 Caudal Fins: Diagnostic features: 0 Dorsal spines 145-148 LARVAE Dorsal rays length: Hatching <4.7 mm Anal 0 spines 143-152 Flexion length: Anal rays 0 <63.7 Transformation mm length: Pelvic reduced Pectoral & A & D C, sequence: development Fin PI Caudal: irregular series just Pigmentation: below lateral midline Minute 3+3 of ca. 50% A base; on BL; Principal portion posterior on D base; at beginning surface dorsal on of eye. Procurrent: Upper 0 ca. at Nephros Diagnostic featnl"es: total (34-39); 37 myomere midline ca. 153 (145-161); minute pigment below lateral Lower 0 myomeres BL 50% at rakers: Derichthys to compared bases; A & D on & Gill beginning Upper 0 ca. ca. at ca. BL 33% at orgin D BL; 75% vs. serpentinus-nephros 0 Lower pigment 75% vs. anus. before 50% midlateral BL; BL Branchiostegais 7 in mean & (range MORPHOMETRICS %) LIFE HISTORY mm 4.7-9.8 mm 10.0-63.7 California Range: to Chile Central 65-95 Sn-AlBL 76-94 82 87 usually Habitat: bathypelagic; to Epi- >305 m depth BDIBL 6-14 6-16 summer & season: early Spawning Spring 11 9 17-22 GDIBL 16-24 planktonic pattern: leptocephali ELH Oviparous; 19 21 HLIBL 12-16 8-14 LITERATURE 15 11 HWIHL 36-61 25-42 1959 Bauchot 44 33 Tighe 1989b SnLIHL 28-41 33-41 37 37 (IlIlIstrator) ILLUSTRATIONS ORIGINAL EDIHL 26-40 x 17-30x 30-42 17-34 Larva, (M. T. 46.5 mm Vona) body, & caudal Vona) sections of 46.5 mm larva (M. T. Head, 32x34 25x27 1-4 PILlBL 1-4 3 2 axis horizontal oval; slightly is Eye axis * second. vertical first, given is PdLIBL 65-70 60-82 67 69 132

145 Sawtooth eel Serrivomer sector 46.5 mm Figure Serrivomeridae 1. Larva, 46.5 mm, whole body and sections showing head, nephros and caudal regions (TC 8406, station 210). 133

146

147 NETTASTOMATIDAE: eels Duckbill CHARTER s. R. family Nettastomatidae . 6 genera and 25 The contains granular patches to large and fang-like. from small, undescribed species more 10 possibly species, with and caudal have confluent dorsal, anal, Nettastomatids gilbertii and Only Facciolella (Smith 1982). and Castle regen- may be which tail, attenuate a slender with fins CalCOFI the in commonly occur Venefica tentaculata Hoplunnis, pectoral retains erated. Only one genus, F. 1). from area ranges gilbertii Nettastomatidae (Table are fins. Nettastomatids benthic but not burrowers. tows Adults collected in deep have midwater been Pt. Conception, California, to Panama, while V. feed on a from subtropical to tropical latitudes and to tentaculata ranges from San Francisco, California, 1989b). small of and (Smith invertebrates fishes variety Chile. along occur the Baja F. gilbertii leptocephali from Bahia coast, California Sebastian Vizcaino to are Nettastomatids oviparous, presumably with Cabo Lucas; one early San transformation specimen eggs. planktonic female One H. sicarius (470 mm) San Diego, a midwater collected was in trawl off free containing ovaries mature with eggs (ca. 1 mm) suggested California. Smith (1989b and pers. comm.) Mazatlan, Mexico (Lane and Stewart off was collected gilbertii may be synonymous with F. F. that larvae are slender (GD 3-14% 1968). Nettastomatid equatorialis, the of study a definitive however, prob- with HL), (34-49% a long-snout and BL) compressed, Larval lem not been done. has are tentaculata V. undivided and is gut is the liver and attenuate tail. The sicarius pers. Smith, (D. Hoplunnis G. unknown. low to moderate (21-57% BL) and may have short from sp. the Gulf of Saurenchelys occur comm.) and undulations two swellings distinct or (Facciolella) A Ecuador. to south of California specimen single H. pectoral (Hoplunnis). In larger Facciolella larvae, the ichthyoplankton a CalCOFI sicarius was collected in fins resorbed. be may Punta off 133.45), station 6509, (CaICOFI San sample Baja California Sur. Additional specimens Juanico, meristic and Morphometric this for information were collected south Ecuador. to sp. Saurenchelys detailed from came study examination 25 of larval Gulf the larvae California of collected been have from Hoplunnis 12 and mm) (7.8-204.1 gilbertii Facciolella and south Ecuador. to sicarius data ecological Additional mm). (17.4-155.2 Lavenberg came Fitch Smith and (1968) and from adults Nettastomatid are long-snouted; and elongate (1989b). some Vomerine proboscis. a fleshy have vary teeth in Nettastomatidae Meristic characters for the nettastomatid species Table the California Current vicinity and Gulf of 1. California based on Garman 1899 and this study. Vertebrae Fin rays Total D CV PrCv Species A 51-57 370 172-181 223-238 Facciolella 283 gilbertii 252-258 ca. 215 ca. 43 ca. 382 ca. Hoplunnis sicarius 317 a 135-172a 61-61 ca. 196-233 tentaculata Venefica 397 ca. 285 possibly due regenerated and broken to tails. counts of range a Large 135

148 NETTASTOMATIDAE Facciolella gi/bertii EARLY LIFE HISTORY DESCRIPTION MERISTICS EGGS Mode Range Shell diam.: Vertebrae: Yolk: No.ofOG: 223 223-238 Total Diam.orOG: surface: Shell 51 51-57 Precaudal 172 172-181 Pigment: Caudal Diagnostic features: Fins: 0 Dorsal spines LARVAE rays 370 Dorsal Hatching length: mm <7.8 0 spines Anal 283 Flexion length: rays Anal ca. length: Transformation m 204.1 0 Pelvic 0 Pectoral PI resorbed C, sequence: development Fin PI' A, & D Pigmentation: & posteriorly Caudal: & On snout laterally lower jaw; on (regenerated 5-7+5-7) 3-4+3-4 Principal PI series midlateral base; braincase; on heart; to posterior & anterior in around myosepta vertically arranged cluster) per (l-several Procurrent: (stellate) ventrolaterally along gut. series notochord; Upper 0 at Diagnostic ca. myomere 57 (52-59); total features: Nephros 0 Lower myomeres ca. gut snout; 220 elongate (213-229); BL <40% generally rakers: Gill 0 undulating Upper specimen, midlateral gut; series low BL); 57% mm 11.3 (1 0 of Lower melanophores around myosepta in vertically arranged notochord; PI base. pigmented 12 Branchiostegals %) mean in MORPHOMETRICS (range & HISTORY LIFE Y-S Range: Pacific from Pt. Conception, California to Panama Eastern mm 11.3-23.4 mm 7.8 mm 26.1-204.1 Habitat: Benthopelagic Sn-AlBL 49-57 30-46 34 51 37 Larvae season: December-Iuly Spawning collected BDIBL 5-6 2-5 leptocephali Oviparous; planktonic pattern: ELH 9 5 4 GDIBL 8-9 5-10 LITERATURE 9 9 7 llllBL 11-12 7-11 Castle 1978 4 11 9 Smith & Castle 1982 22-30 HWIHL 14-23 106 25 18 ILLUSTRATIONS ORIGINAL (Illustrator) SnLIHL 34-44 34-49 47 42 45 (M. 38.1 Larva, T. mm Vona) EDIHL* 16-24 body & section Head of T. Vona) (M. larva mm 38.1 9-14 59x68 18 11 1-2 PILlBL 0.7-3 by 11.3 mm; * Eye in yolk-sac larva is slightly oval, becomes round 0 2 1 given is axis horizontal second. vertical first, axis PdLIBL 0-0 11-21 0 0 15 136

149 Facciolella gilbertii Dogface eel witch ~~============?:==~ .. ~ 38.1 mm --r:; eo 0 6 is 0""1$~ 0 0 -0 0 r) 0 0 (') 0 ¢ 0 0 () 0 0 Q 0 0 0'" g 0.- sections Figure 1. Larva, 38.1 mm, whole body and Nettastomatidae showing head and nephros regions (CaICOFI 6501, station 130.40). 137

150 NETTASTOMATIDAE Hoplunnis sicarius MERISTICS LIFE HISTORY DESCRIPTION EARLY Mode Range EGGS Vertebrae: diam.: Shell Yolk: 252-258 Total No.ofOG: Diam.ofOG: 43 ca. surface: Shell Precaudal ca. 215 Pigment: Caudal features: Diagnostic Fins: o Dorsal spines ca. 382 LARVAE rays Dorsal length: Hatching Anal spines o 317 ca. rays length: Flexion Anal o Pelvic length: >155.2 mm Transformation 11 Pectoral PI & A, C, sequence: development Fin D regenerated commonly Caudal: snout Similar Pigmentation: through all stages; on tip jaw; lower & of 3-4+3-4 Principal & posteriorly on braincase; to & heart; posterior anterior laterally Procurrent: midline along lateral gut; along scattered row along postanally tail; to o Upper myomeres. of ventral edge o Lower features: Nephros Diagnostic total (45-48); at 47 myomere ca. rakers: Gill 254 ca. (245-261); generally gut myomeres <30% BL with two well Upper developed swellings (2 specimens mm, BL); pigment <54.0 <53% Lower of myomeres. postanally ventral edge along 8 Branchiostegals %) & mean in (range MORPHOMETRICS LIFE HISTORY mm 17.4 54.0-155.2 mm Range: Eastern from Mazatlan, to Mexico Panama Pacific Sn-AlBL 21-30 53 25 burrowing not Habitat: but Benthic BDIBL 3-4 Spawning season: 7 3 GDIBL 5-7 leptocephali Oviparous; ELH pattern: planktonic 14 6 HUBL 6-7 LITERATURE 11 6 HWIHL 20-25 1982 Castle & Smith 23 22 SnUHL ILLUSTRATIONS ORIGINAL 38-42 (Illustrator) 38 40 EDIHL C. Walker) mm Larva, 107.5 (R. 13-16 38 15 body, Head, of C. sections caudal & Walker) (R. larva mm 107.5 PILlBL 0.2-1 0.9 PdLlBL 7-12 0 9 138

151 Hoplunnis sicarius 107.5 mm Figure Nettastomatidae 2. Larva, 107.5 mm, whole body and sections showing head, nephros and second intestinal swelling, and caudal regions (Eastropic, station H 43). 139

152

153 INCERTAE SEDIS CHARTER S. R. coheni, from larval known only Thalassenchelys pointed BL). (4-7% small is the head and is snout the in Chlopsidae specimens, was originally placed Castle is Pigment lacking. and Raju (1975) described and Presently (Castle 1975). Raju T. be not can coheni specimens ova defined and well with transformation 1975; Lavenberg (Castle and Raju assigned to a family suggested species the has a short posttransformation eastern 1988). In Pacific, the from ranges coheni T life, to mature and reproduce. sufficient only Seattle, Washing- to California Baja Guadalupe, de Isla (Castle and west to the Hawaiian Islands ton, and Raju from meristic data were obtained Morphometric and west (1958) 1975). Aron collected two leptocephali of 11 specimens (137.2-299.2 mm) collected in midwater MOCNESS Vancouver Island, Canada. British in depth, Columbia, samplers at 0-100 m or trawls March and July. Aron (1958) collected leptocephali in depth Leptocephali are large (up to September in a midwater trawl (maximum ca. cm), have of 30 Cohen's with W-shaped a from and were specimens (1959) m), 30 (33-43% BL) bodies deep extremely m (27-30 June depth). trawl (47-54% BL). gut undulating midwater The taken in an and myomeres, 141

154 INCERTAE SEDIS coheni* Thalassenchelys HISTORY LIFE EARLY MERISTICS DESCRIPTION EGGS Range Mode Shell diam.: Vertebrae: Yolk: No.ofOG: Total Diam.ofOG: surface: Shell Precaudal Pigment: Caudal features: Diagnostic Fins: spines Dorsal LARVAE rays Dorsal length: Hatching spines Anal Flexion length: Anal rays Transformation length: Pelvic Pectoral C, Fin development sequence: D, A & PI None Pigmentation: Caudal: Principal (58-66); 63 total ca. features: Nepbros at myomere Diagnostic Procurrent: myomeres relatively moderate length, extremely (154-161); ca. 158 lacks leptocepbali; pigment deep-bodied Upper Lower MORPHOMETRICS (range %) in mean & rakers: Gill Upper Lower 137.2-299.2 mm Branchiostegals Sn-AlBL 47-54 LIFE HISTORY 50 BDIBL 5-7 known; California Range: Not well Baja larvae from Isla de Guadalupe, 6 Hawaii to Vancouver Island, British Columbia & GDIBL 33-43 37 Epipelagic Habitat: HLlBL 4-7 5 season: Spawning HWIHL 28-41 33 ELH pattern: 35-44 SnLIHL 39 LITERATURE 13-19x EDlHLt 15-22 Castle 1975 & Raju 1988 Lavenberg 16x18 PILlBL 0.4-1 0.9 Adult * unknown PdLlBL 45-51 horizontal axis Eye is slightly oval; t axis is given first, vertical second. 49 142

155 Thalassenchelys cohen; Figure Incertae sedis 1. Larva, 246 mm, whole body and head (Castle & Raju 1975). 143

156 SACCOPHARYNGIFORMES CHARTER R. S. with greatly reduced skeletons (e.g., pelagic fishes no Saccopharyngiformes order the of relationships The are and bones, opercular or rays branchiostegal a relatively unresolved. be to gulpers considered (1984) Smith families small Larvae are known for all neurocranium). eels, midwater (1989) Robins whereas modified highly damaged the Monognathidae. A single except derivative to them be an independent considered of a Monog- (1989) Robins stock. elopomorph separate listed 12 nathus represent a larval specimen, ca. 40 mm, may Saccopharyn- of specialized characters that distinguish and stage Representatives 1987). Nielsen (Bertelsen giformes Saccopharyngidae, Eurypharyn- (Cyematidae, ahlstromi (M area study CalCOFI the this family in most from Monognathidae) and gidae, Anguilliformes; and Saccopharyng- rosenblatti) are shown in Figure M mouth huge is the apparent elongate, the posterior- and if been has Monognathus undescribed An 1. ormes collected CalCOFI Rosen- (R. region Current the in ly projecting suspensorium. comm.). pers. blatt, slender abysso- to Saccopharyngiformes are meso- included: Families Cyematidae Eurypharyngidae Saccopharyngidae 48.5 mm 60.0 mm mm Figure 1. Representatives of the Monognathidae: Monognathus ahlstromi, 48.5 Saccopharyngiformes (Raju 1974); Monognathus rosenblatti,60 mm (Bertelsen & Nielsen 1987). 144

157 CYEMATIDAE: eels Bobtail S. R. CHARTER midwater Cyematids are highly eels, modified widely (Fitch and profile a dart-like create fin caudal reduced temperate in oceans (Fitch all tropical and distributed Smith Lavenberg 1968; Eschmeyer, et al. 1983; and Lavenberg 1968; Eschmeyer et. Smith 1983; al. 1989c). with Historically 1989c). nemichthyids in allied as Anguilliformes, presently they are recognized are are eggs planktonic oviparous; Cyematids Eschmeyer 1989c; (Smith Saccopharyngiformes 1990). are The distinctive cyematid leptocephali unknown. the eastern Pacific, Cyema atrum ranges from In 70 mm, 1989j) Smith moderate in size (maximum ca. to Panama, Oregon relatively rare, and usually is is (16-53% They have fewer deep-bodied and BL). midwater trawls at depths caught in m (Fitch >610 and Other leptocephali, any myomeres other eels. than Eschmeyer 1968; Lavenberg et. al. 1983). Larval C. also Albula and affinis Elops low myomere have sp., collected the CalCOFI study area atrum are throughout respectively), 70-72, body but (77-82 counts and northern from California Baja to California Sur, shape, gut depth, fin morphology, and pigmentation Mexico. ranges Leptocephalus holti, Another cyematid, and atrum haiti C. L. will easily separate the groups. CalCOFI northern from to southern in the California may number the by distinguished myomeres of be be Pacific. area to the L. holti may central the north respectively) pigmenta- lateral and 95-120, vs. (75-83 tion. larva erythrosoma relatively showed haiti which (Smith 1989j), Neocyema of large L. of specimens Our and myomere in terminus nephros total variation m 2000 from two specimens collected from is known and respectively) 95-120, pig- in and (47-62 counts in the south Atlantic (Castle 1977). mentation. elongate C. atrum have jaws, and small eyes Adult and meristic data Morphometric were obtained from scaleless The bodies. compressed stubby, short, dorsal detailed and anal fins originate at midbody and are confluent examination of and leptocephali atrum C. 28 reduced jaws with and caudal fin. The elongate the leptocephali. 9 L. haiti 145

158 CYEMATIDAE Cyema atrum HISTORY DESCRIPTION EARLY LIFE MERISTICS EGGS Range Mode Shell diam.: Yolk: Vertebrae: No.ofOG: Diam.ofOG: 70-80 Total surface: Shell 47 Precaudal Pigment: Caudal Diagnostic features: Fins: 0 spines Dorsal 85-93 LARVAE rays Dorsal Hatching length: 0 Anal spines 72-86 length: Flexion rays Anal length: Transformation 0 Pelvic Fin development sequence: 12-15 Pectoral on dorsum; Pigmentation: one laterally; scattered Individual blotches Caudal: irregular each gut loop peak; on ventrally on gut 4 Principal features: Diagnostic at Nephros ca. myomere 37 total (35-40); Procurrent: myomeres ca. Leptocephalus greater holti, 80 compared with (75-83); 0 Upper & HL), (13-26% eye larger relatively BL), (23-58% depth body 0 Lower laterally. melanophores scattered rakers: Gill 0 Upper mean & (range MORPHOMETRICS %) in 0 Lower 0 Branchiostegals 10.2-69.8 mm LIFE HISTORY 63-73 Sn-AlBL 69 Range: from Oregon the to Panama in eastern Pacific, Worldwide; 18-36 BDIBL Habitat: Midwater in all tropical usually oceans; & temperature >610 m 26 depth 23-58 GDIBL 46 June, season: Spawning except Leptocephali collected all in months 13-18 HLlBL & December September, 15 19-36 HWIHL Oviparous; ELH leptocephali pattern: planktonic 25 47-63 SnLIHL LITERATURE 54 EDIHL 13-26 Bertin 1937 17 Raju 1974 Smith 1979, 1989j 3-5 4 PdLIBL 60-68 64 146

159 Bobtail snipe-eel Cyema atrum 24.9 mm Figure Cyematidae 1. Larva, ca. 24.9 mm (Raju 1974); head of 42.0 mm larva (Smith 1979). 147

160 Leptocephalus holti* CYEMATIDAE EARLY DESCRIPTION HISTORY LIFE MERISTICS EGGS Mode Range diam.: Shell Yolk: Vertebrae: No.ofOG: Diam.ofOG: Total surface: Shell Precaudal Pigment: Caudal features: Diagnostic Fins: spines Dorsal LARVAE rays Dorsal length: Hatching Anal spines Flexion length: rays Anal Transformation length: Pelvic development sequence: Fin Pectoral Over liver; dorsally heart; laterally Pigmentation: dorsally on over & Caudal: on & two front heavy pigment on lower jaw tip most gut loops; have Principal to as heavy as on lower jaw; dorsal from on teeth; upper none jaw, Procurrent: body surface with 1 or 0 or 2 blotches; lateral body with 0, 5 Upper specimens), of or gut on on lower loop (33% laterally blotches; last Lower of specimens). all on lower (67% loops rakers: Gill Diagnostic features: myomere ca. at Nephros (47-62); total 55 Upper with compared (95-120); 108 Cyema atrum, relatively myomeres ca. Lower (8-15% eye smaller relatively BL), 16-30% HL), (body depth slender Branchiostegals melanophores, lateral 5) I or along (usually present, when large midline. LIFE HISTORY & in %) mean (range MORPHOMETRICS region to the Current the California Pacific, North the In Range: from central water mass mm 14.3 mm 15.7-48.2 Habitat: 70-80 Sn-AlBL 76 74 season: Spawning 11-14 BDIBL ELH 11 pattern: 13 GDIBL 23-30 LITERATURE 16 26 17-19 HLlBL 1984 Castle 22 18 1974 Raju HWIHL 10-26 1909 Schmidt 25 21 1989j Smith 43-55 SnLIHL 51 48 Leptocephalus species one than more possibly unknown; a Adults * form EDIHL 8-15 15 12 holti group (Castle 1984). P L1BL 0.7-4 1 3 2 53-76 PdLIBL 0 65 148

161 Leptocephalus holti 40.0 mm Figure Cyematidae 2. Larva, 40.0 mm, whole body and head (Raju 1974). 149

162

163 SACCOPHARYNGIDAE: WhiptaH gulpers CHARTER S. R. in study occurs area, the CalCOFI saccopharyngid One organ, lines" "white longitudinal and caudal slender (Fitch and ranging from northern California to Peru the to that start closer fin pectoral the to than snout is It Nielsen 1985). Lavenberg 1968; Bertelsen and (Nielsen Bertelsen and 1985). in midwater caught relatively rare; generally adults are <2400 collected was one although m, depths at trawls planktonic Saccopharyngids are oviparous; eggs are bathy- at to meso- are Leptocephali surface. the alive Bertelsen 1985). Their lepto- unknown and (Nielsen pelagic, <1525 m (Nielsen depths at collected usually an cephali suspensorium are distinguished by elongate 1985). Bertelsen and yolk-sac small as as larvae in seen be can which mm, 43 mm. 10.4 (maximum small are Leptocephali are large Saccopharyngids (total 1070 mm) length bodied 1909), (23-39% BL), with an deep Schmidt The acute tail. number greater myomeres of total meso- and highly fishes; most bathypelagic modified numerous postanal vs. more 122-136), (210-244 1985). Bertelsen species are cosmopolitan (Nielsen and by Adults a huge mouth, distensi- are easily identified slits (4 fewer gill myomeres (168-194 vs. 84-94), and abdomen, distin- and a caudal filament. ble They are saccopharyngids separate larval from will vs. 6) 1963). (Orton having guished from eurypharyngids by relatively eurypharyngids jaws, a larger abdomen, a more posterior dorsal shorter Examination openings closer to the provided leptocephali ten 20% the gill of BL), (ca. origin fin study. this for data to the Body large curved teeth, wen devel- snout than morphometric and meristic anus, slit and head length and dermal gill first filaments (Nelson 1984, the pectoral fins, depth was measured at oped the of the of top the to snout measured from the tip Bertelsen and et al. Bertelsen 1985; 1994; Nielsen lavenbergi has scaleless, brownish-grey skin first gill slit. S. 1989). simple, a relatively posteriorly, lighter becomes that 151

164 SACCOPHARYNIGIDAE Saccopharynx lavenbergi MERISTICS EARLY LIFE HISTORY DESCRIPTION Range Mode EGGS Vertebrae: diam.: Shell Yolk: ca. Total 175-220 No.ofOG: Diam.ofOG: Precaudal surface: Shell Caudal Pigment: Fins: features: Diagnostic spines Dorsal 0 rays 224-285 Dorsal LARVAE spines Anal Hatching mm 0 40.4 length: 33-36 Pectoral Fin development sequence: Caudal: gut. of Pigmentation: scattered over posterior portion Lightly 0 Principal total ca. Diagnostic 45 (42-52); Nephros features: at myomere Procurrent: 225 ca. myomeres suspensorium; elongate with a (210-244); gut Upper 0 swelling. pigmented single Lower 0 (range in & MORPHOMETRICS rakers: %) Gill mean 0 Upper Lower 0 Y-S Branchiostegals 0 lOA mm 10.5-15.7 mm mm 25.1-40.4 LIFE HISTORY Sn-AlBL 53-58 50-57 52 56 53 to Peru Northern California Range: BDIBL* 18-23 12-25 11 20 17 m & bathypelagic; Meso- depth Habitat: primarily 2000-3000 GDIBL 23-36 27-39 18 29 33 season: Spawning 12-13 HLlBLt 9-11 9 12 10 pattern: changes degenerative ELH Oviparous; females in males & HWIHL 47-55 37-47 suggest semelparity; planktonic leptocephali 59 50 41 LITERATURE SnLIHL 24-36 29-41 4 31 35 1966 BOhlke 37-42x EDlHLt 23-33x 38-46 25-38 Nielsen Bertelsen 1985 & OrtOn 1963 58x53 40x41 27x30 1974 Raju 2-4 P1LIBL 2-2 1989k Smith 2 3 2 PdLIBL§ 14-51 ORIGINAL (Illustrator) ILLUSTRATIONS 0 19 27 larva, Yolk-sac 10.4 mm (M. T. Vona) Vona) section of 10.4 mm yolk-sac larva (M. T. & body Head Larva, 25.1 mm (M. T. Vona) of25.1 Vona) Body mm larva (M. T. section fIrst at depth Body slit. gill * to slit. gill of snout of top length tip from t Head fIrst, given is axis oval; slightly is Eye t horizontal second. axis vertical § Predorsal larva. fm length measured on one small 152

165 Whip gulper tail Saccopharynx lavenbergi 10.4 mm 25.1 mm Figure Saccopharyngidae 1. Yolk-sac larva, 10.4 mm, whole body and sections showing head and nephros regions (CFRD 9104, station MOCNESS); section 25.1 mm, whole body and larva, of nephros region (CaICOFI 6301, station 127.65). 73.3.90, 153

166

167 EURYPHARYNGIDAE: gulpers Umbrellamouth R. S. CHARTER pelecanoides, known eury- only the Eurypharynx them adults Mature saccopharyngids. from separate species, bathy- occurs and meso- in pharyngid the undergo transformation. In males, the significant a (e.g., temperate and tropical of seas zones pelagic head the view dorsal in that so enlarges rosette nasal 1966; Fitch Bohlke Lavenberg the In 1968). and 1989). a1. appears (Gartner 1983; Nielsen et notched from ranges pelecanoides E. northern Pacific, eastern 1.0-1.3 mm eggs ovarian oviparous: Eurypharynxis Peru (Fitch and Lavenberg 1968; to California were Planktonic reported by Nielsen et aI. (1989). eggs are rare in the CaICOFI a1. 1989). Adults et Matarese 35-40 are unknown. Leptocephali are small (maximum larvae the in collected been not have area and study BL), (22-36% bodied an with Smith deep 1989k), mm, our of One speci- larval two regular samples. survey tail. acute fewer myomeres Eurypharynx than Pacific and gyre has the was collected from mens north of 210-244), a greater the saccopharyngids vs. (122-136 west slightly trawl midwater a one from heavier and 1963) Orton 4, vs. (6 slits gill of number CalCOFI sampling grid. gut. pigment the on E. pelecanoides adults are highly modified and, like descriptions from (1966), were taken Adult Bohlke a mouth, an have saccopharyngids, large enormous (1989), a1. and et Bertelsen 1994), (1984, Nelson The tai1. distensible tapering long abdomen, and a and (1989), a1. from taken were meristics et Nielsen smaller black, scaleless body, relatively longer jaw, al. et Raju from length Specimen (1989). Matarese to closer are which smaller gill abdomen, the openings calculated was illustration. (1974) his from fins, snout, anus than to the minute teeth and pectoral easily of the dorsal fin and more anterior origin 155

168 EURYPHARYNGIDAE Eurypharynx peiecanoides MERISTICS DESCRIPTION HISTORY LIFE EARLY Mode Range EGGS Vertebrae: diam.: Shell Yolk: Total 97-125 No.ofOG: Diam.ofOG: Precaudal Shell surface: Caudal Pigment: Fins: Diagnostic features: Dorsal spines o 155-196 rays Dorsal LARVAE Anal o spines Hatching length: Anal rays 118-147 length: Flexion o Pelvic Transformation length: ca. 30--40 mm Pectoral II sequence: Fin development Caudal: posterior Pigmentation: Heavily scattered over portion of gut. Principal o 40 (38-42); total Diagnostic features: Nepbros at ca. myomere Procurrent: myomeres with gut suspensorium; a (122-136); 129 ca. elongate Upper o single swelling. pigmented Lower o Gill rakers: mean %) MORPHOMETRICS & (range in o Upper Lower o mm 8.2 rom 22.7 Branchiostegals o Sn-AlBL LIFE HISTORY 55 56 BDIBL northern in to California Worldwide; from Pacific, eastern the Range: 19 25 Peru GDIBL 22 36 m depth 500--3000 Habitat: bathypelagic; & Meso- HLlBL 21 12 season: Spawning & March Leptocephali collected in May HWIHL 33 44 pattern: ELH Oviparous; degenerative changes in males & females leptocephali planktonic semelparity; suggests SnLIHL 14 37 LITERATURE EDIHL* 27x30 25 a1. 1989 Nielsen et P1LlBL Orton 1963 3 1974 Raju PdLIBL Smith 1979, 1989k 0 41 Van Utrectht 1988 becoming round * Eyes are slightly larvae; larger in oval larvae, small in second. is given fIrSt, vertical axis horizontal axis 156

169 Eurypharynx peiecanoides Umbrellamouth gulper 27.0 mm Figure Eurypharyngidae 1. Larva, ca. 27.0 mm (Raju 1974 as Leptocephaluspseudolatissimus). 157

170 CLUPEIFORMES WATSON W. a catch-all was Clupeomorpha oviparous. polyphyletic Clupeiform fishes are assemblage Most spawn planktonic eggs that range in shape from spherical to teleosts (e.g., Berg 1940) until Green- of "primitive" elliptical, strongly and (1966) defined superorder based on al. the in size from ca. 0.6-3.8 mm, et wood gas and oil without or with a segmented globules, the between connection a unique characters: three with Some yolk. bladder a unique demersal adhesive spawn species and connection), otophysic (the ear clupeid the preopercular eggs planktonic from between hatch that Larvae eggs. and infraorbital connection typically are 2-5 long, with unpigmented eyes, un- ca. mm an lateralis), recessus (the canals line lateral and or formed and large yolk mouth, formed partially sac. unusual, unique, caudal skeleton. Clupeiformes but not and Larvae of species with demersal eggs are larger further diagnosed as having the parietals has been a bery- as lacking developed at hatching. Clupeiform larvae are better and the supraoccipital, by separated by with elongate, characterized preanal length being in the ciform foramen Thus 1985). (Grande ceratohyal The length. body of 60-90% ca. hindgut from ranging Clupeiformes generally accepted been has defined, as marked with prominent annular striations, is and to thought is presently a monophyletic order. The order stage of postflexion the through at least the early part herrings, and sardines, of species 356 contain about is pattern fiber muscle cross-hatched a characteristic distributed among anchovies 83 genera in four families the on the myomeres. on There spines no are visible al. taxonomic The 1988). 1984; et (Nelson Whitehead in postflexion stage or the Late girdle. pectoral or head composition of of the clupeiform subgroups, and some typically a fin shifts dorsal the transformation during relationships are subgroups, the unsettled between cephalad. myomeres few a moves also few anus The and Grande 1984; Berry McGowan 1973b; (Nelson cephalad myomeres the gut as shortens during comple- 1988). al. Whitehead et 1985; Whitehead 1985; of fin development. anal Larval pigmentation tion size Most clupeiform fishes are small to medium to the gut and is and largely light restricted typically (ca. 10-30 cm) marine planktivores that in school at the early part least ventrum through of the post- deep- to moderately They waters. coastal are elongate flexion stage. usually with bodied, slightly to strongly compressed, in round herrings one (only abdomen the along scutes of species 18 occur engraulids and clupeids About in the and spines no New are There anchovies). World the of larvae these, the area. study in CalCOFI Among two or one first the although fins, dorsal unbranched anchovy northern the far by are mordax) (Engraulis and small are typically rays fin anal and spine-like. ichthyo- CalCOFI larvae abundant most in taken Clupeiform fishes typically are brown to green or blue plankton The collections. Pacific sardine (Sardinops dorsally silvery below. They are key components and rank larvae whose abundance, overall in eighth sagax), econom- most the among are and coastal ecosystems of of CalCOFI the establishment for was responsible of accounting for annually fishes, ically important collapse the in fishery of virtue by program its of the world's (e.g., catch fish of the one-quarter about decades early this of 1982). Radovich (e.g., century al. 1988). et Whitehead 1985; Whitehead included: Families Clupeidae Engraulidae 158

171 CLUPEIDAE: Herrings WATSON AND E. M. SANDKNOP W. family Clupeidae contains cally 180-190 species The near school that fishes planktivorous cm) 10-30 about waters. coastal warm in surface the (Nelson 1994; Whitehead 1985), nine species few A of occur which area study CalCOFI (Table 1). Clupeidae The in cool temperate waters and some species occur in the precipitous demise) ultimate (and decline to inhabit brackish or fresh water. Clupeids are slightly of the major with strongly compressed, preanal length about half to the these, Pacific of one for fishery commercial with of of a body and typically length, the three-quarters for establishment basis principal was sardine, the are fins Pectoral mouth. terminal small on low inserted program Cooperative original (the 1947 in CalCOFI sides each the fins, pelvic the and with 7-9 rays, are Sardine Research Program was renamed the California 2-4 Fisheries Investigations program anal and dorsal the in rays Oceanic abdominal. The first Cooperative are fins commonly unbranched, and the first 1-3 1953). in spine-like. Clupeids and small usually are (except herrings) round a row have scutes of belly. the along the is Clupeidae not family of The composition Pigmentation commonly is green to blue on the dorsum entirely and settled (McGowan Berry 1984). For of dorsolateral stripes and below, silvery with patterns round the example, herrings are in the Clup- placed among the most species. in spots some or Clupeids are as many by Dussumieriinae subfamily the authors eidae species economically important fishes, several with Nelson Whitehead others while 1985) (e.g., 1994; (e.g., large commercial fisheries Whitehead supporting them family (e.g., Eschmeyer et al. consider a separate fisheries Some important in sport species are as 1985). 1983). Within the Dussumieriinae, some authors well. to Etrumeus a cosmopolitan species, of consist consider (e.g, species E. teres, and an endemic South African life histories are known Early many for well of the Whitehead 1985; Whitehead and 1986) Wongratana and marine important commercially anadromous of the E. teres at some others consider while least (see 1984). Berry and McGowan review species by populations to be separate species (e.g., Hildebrand Clupeids spawn plank- spherical Most oviparous. are 1963b; 1983). et al. Eschmeyer follow We Whitehead's 0.6-3.8 mm (typically 1-2 from ranging tonic eggs in Clupeidae the of chapter. this arrangement (1985) with a smooth, often very thin, chorion and a mm), narrow to very wide perivitelline space, a segmented ichthyo- CalCOFI larvae taken in clupeid Most oil more or species) (most one usually and yolk, California plankton collections off and Baja California Clupea, Dorosoma, and Spratelloides species globules. are Sardinops rela- sagax. teres Larval Etrumeus are spawn adhesive demersal eggs (McGowan and Berry common in summer and fall tively Cali- off southern ca. 1984) in in 1-2 enclosed are that diameter mm larvae Opisthonema while California, Baja and fornia sheathes. the from hatch Larvae gelatinous thick off southern in occasionally have taken summer been with ca. unpigmented mm 3-5 eyes, at planktonic eggs the Gulf of in Baja California common are (they large from Larvae sac. unformed mouth, and yolk are Larval California). Clupea pallasii sometimes taken mm 4-7 ca. with at demersal eggs typically hatch from stations CalCOFI central at spring in winter and formed to fully eyes, pigmented partially partially or specimens few A northward. California of Harengula are mouth, and small yolk sac. Larvae functional and thrissina probably unidentified an Lile clupeid, usually cylindrical compressed, slightly to elongate, stolifera, dip collections from net in taken been have becoming moderately and compressed deeper-bodied Sur, CalCOFI during California Baja Magdalena, Bahia stage. transformation postflexion the during or gut The Dorosoma and sapidissima Alosa surveys. petenense long is (preanal length 75-90% ca. BL, shortening to species whose estuarine, introduced, primarily are during with transformation), ca. 70-80% postflexion or apparently larvae collected during been have not prominent annular striations on the hindgut. The CalCOFI surveys. cross-hatched of muscle clupeiform pattern typical above forms fin dorsal The visible. is clearly fibers the (typi- Most adult clupeids are small to medium size 159

172 fin and characters, pigment counts), ray fin origin, anal of the hindgut, and typically shifts middle posterior to to forward a position nearer midbody during transfor- location. occur- seasonal and size Larval geographic mation. Pigmentation late embryos and dorsal in is (e.g., some rence may be useful in cases Clupea versus can species all of be all Not Sardinops). distin- stages melanophores all or most but newly-hatched larvae, guished it example, for presently; and the during ventrum migrate ventrad to the gut how unknown the is King stage Orton 1953a; O'Toole and (e.g., yolk-sac of H. of thrissina can be distinguished from those eggs 1974; yolk Following 1989b). al. et Matus-Niv6n the Opisthonema species, or how larvae of the Opis- absorption, dorsal and lateral pigment commonly are thonema distinguished from species another. are one caudal the on except lacking, until area, postflexion the Fore (1973) give and Houde charaCters for distinguish- ing the eggs the of H. jaguana and 0. oglinum in Gulf stage. for Mexico; of characters might be useful similar the length, The elongate body with long preanal pig- eastern Pacific species. mentation largely limited to the gut and ventrum, pattern striated fiber hindgut, muscle cross-hatched and literature on based are descriptions following The clupeids of larval from all should separation allow detailed and observations on of of 4-46 specimens larvae other lightly pigmented in the Cal- elongate, each Alosa 2). Clupeidae (Table species sapidissima area COFI except engraulids. Clupeids commonly have refer and to Dorosoma petenense are not described; than length preanal a longer engraulids have (usually Jones et descriptions al. (1978) and Wang (1981) for BL versus usually <75%), >75% when the dorsal and references. additional and obtained were data Meristic the it inserts stage) flexion during (beginning fin forms Clothier from (1950), Peterson (1954), McHugh (or a few anal several myomeres anterior to the anus to Berry (1956), Berry and (1963), McGowan and Barrett origin) fin clupeids, in to myo- a few one to contrast in Matus-Niv6n and (1984), et (1989b). al. and Dorsal the behind or anus the to posterior meres anal (above which totals are small the counts include fin-ray anal fin origin) in engraulids. Identification clupeid the of Ecological rays. was information spine-like first requires species a combination characters of meristic obtained Whitehead al. (1983), et Eschmeyer from myomeres, total and postanal, preanal, number (e.g., of and (1985), and Whitehead Rodriguez-Sanchez anus or myomeres between dorsal fin insertion and (1995a). 160

173 the Table geographic ranges for the clupeid species in and CalCOFI study area. All have characters Clupeidae 1. Meristic fin rays. Northern range limits are followed by southern limits. Abbreviations: Gulf, Gulf of California; 10+9 principal caudal BCS, Baja California Sur. BC, Baja CA, California; California; rays Fin Vertebrae Gill CV Total D A PrCv P Species C Range PI 2 2 rakers 27-29 55-58 15-19 19-23 13-18 27-29 7-8+7 Alosa sapidissima 9 14-25+ Alaska-Ensenada BC 28-47 15-21 46-58 29-33 13-20 15-20 19-23 9+8-9 20-21+ 9 Clupea pallasii Sea-BC Bering 41-50 30-32 12-17 11-15 17-27 11-13 40-45 9+6-7 7-8 petenense 300-440 Dorosoma Bay-Long Humboldt total Beach CA 15-20 16-22 9-19 8-9 48-55 14-17 6-9+ 32-36 12-15+ & teres Etrumeus Monterey CA 6-8 28-35 Gulf-Chile 19-22 16-20 8-9 7-10+ 15-16 9-18+ 40-43 14-17 20-23 La Jolla CA & thrissina Harengula 24-34 5-7 Gulf-Peru 8-11+ 21-23 19-21 42-44 15-18 15-23 13-14 8 13-18+ Life stolifera Bahia Ballenas BCS 6-9 32-36 & Gulf-Peru 17-20 19-22 15-18 110-149+ 22-24 44-48 8-9 6-9+ Pedro CA San libertate & Opisthonema 6-8 161-224 Gulf-Peru 45-48 17-20 19-23 8-9 16-19 7-8+ 70-99+ Redondo Beach CA 0. medirastre 5-6 Gulf-Peru & 110-156 48-54 16-19 8-9 6-9+ 17-20 22-23 17-20 28-30 21-23+ Alaska-Gulf sagax Sardinops 5-8 44-45 of specimens (above) Table size ranges (in mm, below) used in the preparation of the clupeid Clupeidae 2. Number and species descriptions. An used in description. indicates ilL" literature the Flexion Postflexion Preflexion Yolk-sac Transformation Egg Juvenile Species La,b 5 7 6 3 3 3 Clupea pallasii 19.5-26.8 8.3-12.8 15.8-18.5 25.7-29.5 7.4-8.5 53.8-58.4 Lc,d,e 5 2 3 6 6 3 teres Etrumeus 5.2-9.8 10.7-13.7 10.0-12.1 32.5-35.1 3.8-4.8 63.8-77.4 Lf Lf 10 5 5 3 3 thrissina Harengula 3.7-6.7 13.0-22.0 19.3-25.2 7.6-9.3 29.2-39.4 Lg Lg Lg Lg Lg Lg 4 Opisthonema libertate 37.3-49.7 5 8 7 15 3 5 3 Sardinops sagax 4.5-10.8 11.3-14.7 14.0-23.9 31.7-38.3 3.7-5.6 36.2-50.4 1.5-1.7 al. a Matarese et 1989 1981 b Wang Leis and Watson C 1974 1974 d O'Toole and King 1973 e Houde Fore and Marinas, Postal La Paz, BCS, Mexico) f Matus-Niv6n et aI. (manuscript; Apdo. Ciencias de Interdisciplinario Centro 592, 1989b al. et g Matus-Niv6n 161

174 CLUPEIDAE Clupea pallasii MERISTICS EARLY LIFE HISTORY DESCRIPTION Mode Range EGGS Vertebrae: Shell mm 1.2-1.7 diam.: Yolk: Segmented 51-53 46-58 Total No.ofOG: 0 Diam.ofOG: 29-33 31 surface: Precaudal Smooth Shell 21-22 Pigment: yellowish Caudal 19-23 Yolk clusters, primarily on seaweed features: Attached in Diagnostic large Fins: 0 yolk segmented eelgrass; & 0 Dorsal spines 18-19 15-21 rays Dorsal 0 LARVAE Anal spines 0 13-20 16-17 Hatching Anal rays length: mm 5.6-7.5 Flexion mm length: ca. 16-19 9 9 Pelvic 15-20 17 length: 35 mm mm through Pectoral 25-27 Transformation P A, , D, , C sequence: development Fin C Caudal: & PI I 2 2 10+9 10+9 Yolk-sac-prejlexion-Dorsolateral along each side Pigmentation: row Principal gut; of row along ventral margin of Procurrent: anterior 40-50% of single scattered few mm; 10 ca. by isthmus on 1-2 gut; of 50-60% posterior Upper 9 9 8-9 Lower 8 tip. around notochord Flexion-I-2 dorsolaterally on hindbrain by 17 internal mm; over 18 0-2 midbrain over series mm, after increases; Gill rakers: 21 Upper 20-21 19 mm & spreading cephalad; cord ca. by posteriorly beginning spinal 41-50 45-46 of gut; near increasing lower on isthmus; 1-2 on surface dorsal series Lower 8-9 end internally cleithmm; proximally on C rays; of on lower hypurals Branchiostegals parhypural ca. by 18 mm. Posiflexion--On jaws by 21 mm; on & snout 25 opercle & by spreading c1eithrum; along increasing mm; LIFE HISTORY ventrad gut; on on Dbase & midlaterally on caudal 21 by peduncle spreading & posteriorly beginning dorsolaterally scattered mm; California Baja to Alaska in the eastern Northern Pacific, Range: by mm. 25 cephalad western Pacific in Korea to southward the Diagnostic 53-55)=40-44+10-14 (usually 50-56 Myomeres features: (usually before transformation; 5-8 myomeres 6-7) between D spawn over Schools Habitat: continental shelf; enters bays & estuaries to & insertion notochord prominent stage; postflexion through origin A pigment; at large size hatching tip/caudal & flexion. Spawning season: Late fall-spring, primarily December-March in Californian waters in %) MORPHOMETRICS & (range mean larvae ELH Oviparous; attached demersal eggs pattern: planktonic & Y-S PrF F PoF Juv • Tr LITERATURE 83-86 80-91 84-85 Sn-AlBL 80-87 75-77 75-77 84 84 84 83 76 75 1989 et al. Matarese McGowan & 1984 Berry 7-9 BDIBL 3-4 5-6 8-15 15-19 19-21 3 6 7 10 17 20 1988 Takita 1958b al. et Uchida 12-14 16-20 12-15 HUBL 19-23 26-29 26-28 1981 Wang 13 17 21 13 27 28 54-71 44-52 52-63 38-47 HWIHL 33-36 30-33 ORIGINAL ILLUSTRATIONS (Illustrator) 56 60 49 43 34 32 17-24 8-19 25-29 20-28 SnLIHL 27-29 26-31 mm (R. C. Walker) 7.6 Yolk-sac larva, 21 12 24 27 28 28 Transformation C. (R. mm 29.5 Walker) specimen, 7-23 I I x x EDIHL* 9-23 18-28 24-27 22-28 23-25 25-25 27-30 22x25 21x24 24 24 25 29 * horizontal Eye slightly initially oval, becoming round by flexion stage; second. first, given is axis axis vertical 2-3 2-3 3-4 5-9 0-3 14-15 I 3 3 3 15 7 0-0 0-0 0-0 1-6 8-9 9-10 0 0 0 3 10 8 162

175 pallasii Clupea Pacific herring ... ... ... ;; ... ;;;0;;;>;. 0 ..$.· ...• ~ ... ··, ~0E0t'" ~ 12¥ mm 7.6 1.4 mm ~.~/;<;> .. : ... ~ 0., ~=:>::; , ... ""0" ••• , ••••• , ." '.' •••• • ~ ... ~- ... 10.4 mm mm 19.0 Q "- - •• ' .' 23.8 mm 29.5 mm (CFRD Clupeidae Egg, 1.4 mm (Uchida et al. 1958b); yolk-sac larva, 7.6 Figure 1. Ref. Coil., OSU 247C, Buoy 15, Yaquina Bay, mm Oregon); preflexion larva, 10.4 mm; flexion larva, 19.0 mm; postflexion larva, 23.8 mm (Matarese et al. 1989); transformation specimen, 29.5 (CFRD Ref. mm Coil., San Francisco Bay, California, May 15, 1974). 163

176 Etrumeus teres CLUPEIDAE EARLY LIFE HISTORY DESCRIPTION MERISTICS EGGS Mode Range diam.: Shell mm 1.2-1.5 Yolk: 1.0--1.3 mm diam. Segmented; Vertebrae: No.ofOG: Diam.ofOG: 0 54 48-55 Total Shell Smooth surface: 35 32-36 Precaudal pale on dorsum melanophores yellow; to colorless Yolk Pigment: & 19 15-20 Caudal of late gut embryos stage of terminus near Fins: features: ofOG; perivitelline lack Diameter; narrow space; Diagnostic 0 0 Dorsal spines segmented yolk 19--21 16-22 Dorsal rays 0 0 spines Anal 11-12 LARVAE 9-19 rays Anal length: 3.8-4.8 mm Hatching 8 8-9 Pelvic ca. Flexion length: 10--10.5 mm through ca. 12-13 mm 16 14-17 Pectoral Transformation 32-35 ca. mm length: Caudal: sequence: C Fin D, A, C development P PI' , 10+9 10+9 2 b 2 Principal & snout, Pigmentation: Yolk-sac-Scattered dorsally over midbrain Procurrent: number in decreasing end by disappearing & initially series stage; of 7 6-9 Upper dorsum on from nape by to end to last myomere, migrating ventrum 6 6-8 Lower intestine, on stage; patch posteriorly preanal of may extend onto Gill rakers: finfold. or Prejlexion-pos(jlexion-None laterally; dorsum on 12-15 Upper each along side series ventrolateral of ventral margins gut; series along 28-35 Lower of few on proximal 25% tail; outer principal hindgut, isthmus, & of 14-16 Branchiostegals caudal & fm rays in postflexion Transformation-Dorsally stage. spreading dorsolateral, caudad; dorsum, on head; on laterally LIFE mSTORY & caudal peduncle. on C spreading caudad; increasing 40-45+11-15=51-58 (usually Diagnostic features: Myomeres Monterey Bay, California, to in Worldwide; eastern Pacific from Range: D long, origin; A & insertion between myomeres 3-4 41-42+13); Chile wedge-shaped snout patch of melanophores HL); (SnL=27-40% yolk-sac & early preflexion stage; no dorsal in hindgut on posteriorly bays enter shelf, may continental over Schools Habitat: lateral or trunk late through preflexion or melanophores tail from stage. postflexion Spawning season: Summer-fall in area CalCOFI MORPHOMETRICS (range in mean & %) ELH eggs planktonic Oviparous; pattern: larvae & Y-S PoF F PrF Tr Juv LITERATURE 78-82 84-87 84-87 76-84 Sn-AlBL 80--82 80--83 1980 Moser & Ahlstrom 80 80 81 85 81 85 1973 Fore Houde & 8-8 7-9 7-9 BDIBL 14-16 7-10 16-17 1978 al. Jones et 8 8 8 8 15 17 1979 al. et Miller 1961a Mito 19--23 17-19 18-20 20--21 26-29 28-30 HLlBL & 1974 O'Toole King 28 27 20 19 18 21 Takita 1988 45-59 46-59 45-50 50--56 30--32 HWIHL 35-43 Watson 1974 Leis & 53 52 53 49 39 31 27-33 7-9 35-40 31-33 33-40 28-29 SnLIHL (Illustrator) ORIGINAL ILLUSTRATIONS 8 31 29 36 37 32 17-22x 36-48x 18-20x 17-32x EDIHL* (R. Walker) C. mm 12.1 Flexion larva, 21-25 17-21 21-28 22-23 23-24 28-31 27.1 Walker) Transformation C. (R. mm specimen, 20x24 23x24 19x20 42x22 24 31 2-4 3-4 2-5 7-11 13-14 0--0 by round becoming oval, horizontal initially Eye * transformation; axis is 4 3 3 14 0 10 vertical given first, axis second. 0--0 0--0 0--0 8-10 7-8 0--0 0 0 0 8 0 8 164

177 Round herring teres Etrumeus ... ... "'''''''''./''' ,' ... ,r .' " " .. : : " ',." . .. ". . .. " * • 5.7mm 1.4 mm 6.6mm ',' : 12.1 mm 17.7 mm 27.1 mm Figure Clupeidae 2. Egg, 1.4 mm (Ahlstrom and Moser 1980); yolk-sac larva, 5.7 mm; preflexion larva, 6.6 mm (Miller et al. 1979); flexion 17.7 specimen, (CFRD n03AH, station 2213 No.7); postflexion larva, mm mm (CFRD Ref. Coll.); transformation 12.1 27.1 mm larva, the (SID scales developing posteriorly along 85-176; lateral midline are not shown). 165

178 CLUPEIDAE thrissina Harengula LIFE EARLY DESCRIPTION mSTORY MERISTICS EGGS Mode Range mm 1.7 diam.: Shell Yolk: Segmented Vertebrae: 41 I No.ofOG: Diam. of 40-43 mm 0.09 OG: Total surface: Shell Smooth 20-23 Precaudal Pigment: 19-22 Caudal features: Diagnostic perivitelline space; wide with egg Large Fins: small yolk; segmented OG 0 o spines Dorsal 18-19 16-20 Dorsal rays LARVAE o 0 Anal spines 16 3.7 mm ca. length: Hatching 14-17 rays Anal Flexion length: 6.6-8 mm through 9.5 mm 8 8--9 Pelvic Transformation ca. length: mm 19.5 15-16 15-16 Pectoral , D, C sequence: Fin development C , , PI P A, Caudal: 2 2 I Yolk-sac-On dorsum moving to gut by end of initially, 10+9 10+9 Pigmentation: Principal notochord Preflexion--Dorsally & around tip. remaining stage except Procurrent: at PI base; ventrally isthmus; 7 on notochord tip; on cleithrum on 7-10 Upper sides along dorsolaterally margin 6 ventral on bladder; gas over gut; of 5-7 Lower except on none 10-20%. of gut posteriorly from gas bladder last Gill rakers: caudad 9-18 Flexion-postflexion--At nape along 6.6 mm, extending by Upper by reaching & urostyle becoming deeply internal margin, dorsal 24-34 LoWer over 14 externally midbrain by 8 mm, mm; mandible & opercle by on Branchiostegals mm; pair under brain by 6.6 mm; in otic capsule by 9.5 mm; 14 over & cleithrum after 9.5 mm; 9 by foregut on isthmus increasing mSTORY LIFE midline lateral on posteriorly mm; by on extending cephalad; mm, 14 D ventral oftail by 14 mm; on margin base by 14 mm; proximally on Peru, including Gulf of California; Jolla, California, to Callao, La Range: internally on rays; C urostyle hypurals. & north rare California Baja of Sur Myomeres features: Diagnostic (33-35+7-8 through flexion, 40-43 to myomeres 4-6 stage); posttlexion late by 28-31+11-14 shifting waters coastal in Schools Habitat: moderate over pigment heavy & A origin; to D between insertion dorsally pigment notochord tip through early posttlexion stage; on April-October Spawning season: Larvae collected beginning early & posttlexion in stage. dorsal margin head planktonic Oviparous; ELH pattern: larvae & eggs %) MORPHOMETRICS (range & mean in LITERATURE PrF Y-S PoF F Tr Juv de Interdisciplinario Centro (manuscript; al. et Matus-Niv6n Ciencias 84-87 85-87 Sn-AIBL 75-84 73-75 74-76 Apdo. Marinas, Mexico) BCS, 592, Paz, Postal La 86 85 79 74 75 ORIGINAL ILLUSTRATIONS (Illustrator) BDIBL 7-10 8-9 18--23 8--16 23-24 9 8 13 20 24 Egg, 1.7 mm (E. Matus-Niv6n) 17-19 17-19 lllJBL 27-28 21-26 27-28 Yolk-sac larva, mm 3.7 Matus-Niv6n) 18 (E. 23 18 28 28 (E. Matus-Niv6n) mm Preflexion 6.6 larva, 43-48 46-53 34-42 HWIHL 32-37 36-36 (E. Matus-Niv6n) mm 9.0 larva, Flexion 47 51 34 37 36 larva, mm (E. Matus-Niv6n) Posttlexion 18.5 17-22 19-25 SnLIHL 25-29 24-32 27-30 C. 20.8 mm (R. Walker) Transformation specimen, 20 22 27 28 28 29-30 23-27 EDIHL 23-28 27-31 28-32 30 25 25 29 30 0-2 1-2 2-7 12-20 16-18 PILlBL I 2 4 17 17 P 0-0 L1BL 0-0 2-11 12-15 11-13 2 0 0 7 12 13 166

179 Flatiron herring thrissina Harengula 3.7mm 1.7 mm (" " : ... :": ,:";,' / .. :' ~. :" :'" ... \ . " .. • • !" ; :',' ::.;.:: 6.6mm 9.0mm 18.5 mm 20.8 mm Figure Clupeidae 3. Egg, 1.7 mm; yolk-sac larva, 3.7 mm; preflexion larva, 6.6 mm; flexion larva, 9.0 mm; postflexion larva, 18.5 mm are 70-139; mm (SIO scales 20.8 not shown; damaged dorsal, anal manuscript); transformation unpublished specimen, (Matus-Nivon et ai., and caudal fins on this specimen were modified in the illustration based on fins of a 19.3 mm specimen from the same lot). 167

180 CLUPEIDAE Opisthonema fibertate EARLY LIFE HISTORY DESCRIPTION MERISTICS Mode EGGS Range 1.3-1.9 mm diam.: Vertebrae: Shell diam. 0.~.9 mm Yolk: Segmented; 46 No.ofOG: 1 44-48 Total of mm 0.08-0.17 OG: Diam. Smooth surface: Shell 23 22-24 Precaudal 23 Melanophores on after of embryo dorsum Caudal Pigment: beginning soon closure blastopore Fins: thin, with egg Large features: Diagnostic wide chorion; fragile Dorsal spines 0 0 OG 19 yolk; segmented space; perivitelline small 17-20 Dorsal rays 0 Anal spines 0 20 LARVAE 19-22 Anal rays mm length: Hatching 2.3-2.6 8 Pelvic 8-9 6.5-8 mm length: Flexion 16-17 15-18 Pectoral mm Transformation length: 19.4-32.2 Caudal: A, C P , C & Fin PI 10+9 10+9 development sequence: D, , Principal I z z migrating rows Yolk-sac--Initially two along Pigmentation: dorsum, Procurrent: ventrad gut 7 to & Preflexion--On stage. ventral end by margin of Upper 6-9 6 Lower to side each along 6-8 dorsolateral cleithra; to adjacent isthmus row near middle posteriorly; of margin ventral along gut double gut, of row Gill rakers: 110-149 series dorsal anus; near along forward gut on dorsally spreading Upper 161-224 hindgut after 4 pair on Posiflexion-Internal dorsolateral mm. Lower & eye, hindbrain on opercle mm; 12 over mid- & hindbrain; below by Branchiostegals by capsule otic mm; 19 in by cleithrum of part lower on mm; 12 by of along 12 mm; on dorsal margin HISTORY caudal peduncle, on hypurals, & LIFE C rays mm; on mm; 19 by midline lateral along series internal 12 by D mm. 19 by bases A & Peru & Pacific Eastern Range: from San Pedro, California to throughout Myomeres features: Diagnostic (usually 32-40+5-13=45-48 Sur. north of Baja California coast outer on rare California; of Gulf between myomeres 5-9 <16 in larvae 37-40+5-10 (usually 6-8) mm); on & A origin; pair ·of dorsolateral melanophores D insertion Schools in coastal waters Habitat: melanophore(s) external & dorsal caudal of margin on hindbrain in postflexion stage. early peduncle July-September Spawning season: Spring-fall, principally (range MORPHOMETRICS mean & %) in larvae planktonic Oviparous; ELH eggs & pattern: PrF F Y-S PoF LITERATURE Tr Juv Sn-AlBL 79-89 86-96 84-87 79-84 74-85 & 1988 Esquivel-Herrera Funes-Rodriguez 88 85 86 82 87 78 Matus-Niv6n et al. 1989b McGowan & 1984 Berry BDIBL 7-10 6-10 7-9 9-14 25-27 16 8 9 8 12 26 15-18 15-21 HLlBL 15-19 19-25 26-31 16 17 16 16 21 28 HWIHL 31-39 35 13-18 16-22 21-27 SnLIHL 25-28 24-28 17 19 23 18 26 27 EDIHL 34-42 25-35 26-31 23-27 28-34 36 30 39 28 25 31 15-17 PILlBL 16 PzLIBL 10-10 10 168

181 Deepbody thread Opisthonema tibertate herring 4.2mm ---__ 1.7mm ',', ',: ',,',.: .;~ :. ~. :, :.: .;:: . • .. I' 5.1 mm 12.1 mm 19.2 mm 22.6 mm Figure Clupeidae 4. Egg, 1.7 mm; yolk-sac larva, 4.2 mm; preflexion larva, 5.1 mm; late flexion larva, 12.1 mm; postflexion larva, mm 19.2 transformation specimen, 22.6 mm; (Matus-Niv6n et al. 1989b). 169

182 Sardinops sagax CLUPEIDAE HISTORY EARLY LIFE DESCRIPTION MERISTICS EGGS Mode Range Yolk: Shell mm 1.3-2.1 diam.: mm 0.9-1.4 Segmented; diam. Vertebrae: of of No. mm 1 (occasionally 0.12-0.19 OG: OG: Diam. 51 48-54 Total o part northern in of range) 29 28-30 Precaudal Smooth surface: Shell 22 22-23 Caudal along Melanophores dorsum of embryo soon after form Pigment: Fins: blastopore closure 0 0 spines Dorsal features: thin, fragile chorion Diagnostic Large egg with wide & 19 17-20 Dorsal rays segmented space; small perivitelline with yolk OG 0 0 Anal spines 18-19 17-20 Anal rays LARVAE 8 8-9 Pelvic 3.5-3.8 Hatching mm length: 18-19 16-19 Pectoral through mm 9-10.5 length: Flexion mm 13-14.7 Caudal: length: 25 mm through 35-40 mm Transformation 10+9 10+9 Principal , C , D, A, P Fin & C development PI sequence: z I z Procurrent: Yolk-sac--Initially dorsum, migrating ventrad to on Pigmentation: gut 6-7 6-9 Upper end by margin ventral & none Preflexion-flexion-Usually stage. of 5-6 5-8 Lower on dorsum; PI I origin, above 1-2 on isthmus; 1 on c1eithrum just rakers: Gill ca. of below PI insertion after side 13 mm; dorsolateral row on each 21-23 Upper posteriorly gut, shifting to single irregular bladder; gas from dorsally 44-45 Lower row double gut posteriorly from ventral bladder; few gas margin of on 6-10 Branchiostegals hypurals on & on C rays, usually 1 prominent ventral margin of tail; Posiflexion-1-2 anteriorly hypurals. middle between distally spot HISTORY LIFE by over hindbrain midbrain mm, 13-14 mm; 16 by over posteriorly in capsule otic by 15 mm; on internal & jaws by end of stage; snout eastern Pacific coast to Cabo San Range: Alaska & Kamchatka along to along series spinal cord beginning anteriorly by ca. 15 & mm Sur; in Lucas, Baja Gulf of California to north California Guaymas, caudad. spreading Sonora, Mexico features: Diagnostic Myomeres myomeres 5-8 38-42+10-14=50-54; a often origin; A & insertion D between 6) (usually prominent over shore near often shelf, continental Schools Habitat: hypurals. at distal margin between middle melanophore in Spawniug season: Throughout the year with peaks spring & summer, %) MORPHOMETRICS (range & mean in fall in minimum California off winter & Y-S PrF Tr PoF F luv Oviparous; planktonic eggs larvae pattern: & ELM 82-86 82-86 Sn-A/BL 76-77 76-78 79-81 80-85 LITERATURE 84 84 80 83 77 76 14-1S 6-7 6-8 7-11 16-17 BD/BL 1960, 1966 Ahlstrom 4-6 1959, IS 17 5 7 7 8 1970 Kramer Sandknop & Matarese 1984 13-\S 24-25 13-15 15-21 13-15 26-27 HLIBL Matarese al. et 1989 18 15 14 14 24 27 & McGowan 1984 Berry 52-70 38-48 SI-72 28-31 47-57 28-31 HW/HL Scofield 1934 57 43 52 30 30 60 15-30 24-29 SnLIHL 26-30 30-30 28-31 8-14 29 23 27 11 30 30 21-25 21-28 24-28 23-33 31-39 ED/HL 25-27 28 23 23 26 26 36 13-15 1-3 1-3 8-12 3-3 0-0 PILIBL 2 3 3 14 0 10 0-0 0-4 0-0 0-0 9-10 9-10 PzLIBL 10 0 0 0 1 9 170

183 Pacific sardine sagax Sardinops 3.0mm 1.4-2.1 mm mm 6.1 mm 9.1 mm 19.7 31.3 mm Figure Clupeidae 5. Egg, 1.4-2.1 mm (Matarese and Sandknop 1984); yolk-sac larva, 3.0 mm; preflexion larva, 6.1 mm; flexion larva, specimen, 9.1 postflexion larva, 19.7 mm; transformation mm; 31.3 mm (modified from Kramer 1970). 171

184

185 ENGRAULIDAE: Anchovies WATSON AND E. M. SANDKNOP w. to eleven of the approximately 139 engraulid Nine for Early life histories of a number are known well in species the occur by review (see species CalCOFI study area (Table All 1984). Berry and McGowan Engraulidae widely are anchovies the Although 1). small large moderately to spawning oviparous, are that dimension 0.7-2.3 mm) planktonic eggs (largest and as a monophyletic group, the composition accepted with strongly to spherical from shape in vary elliptical, relationships the within family are not of the genera a knob species. genera Most some in pole one at lack Whitehead fully resolved (McGowan and Berry 1984; a segmented oil globules; all have yolk. Larvae ca. et polyphyletic al. 1988). The Engraulinae may be a partial- 2-3.5 mm long hatch with unpigmented eyes, undescribed there 1985) and (Grande both are anchovy mouth, formed ly no or little and sac, yolk a large (Whitehead split be may et that species and species al. preanal with elongate, are Larvae body. the on pigment Nelson 1988). Cetengraulis be that suggested (1984) ca. length hindgut length. body of 60-80% is The in primarily on a subgenus Engraulis based included as gas strongly marked with annular striations. The shared intestinal synapomorphy arch a gill and coiling bladder, between which first inflates mid-preflexion the with New World Engraulis species. However, flexion early and near typical The midbody. is stage, characters, for example the order larval initial fin of clupeoid cross-hatched muscle fiber pattern is visible and Anchoa in ray formation, which is the same fin dorsal The development. larval of during much eastern different but Cetengraulis the from both in typically inserts 1-4, overlapping at postanal myomere Pacific Engraulis (e.g., Orellana and Balbontin 1983; During the origin fm by one the to several anal rays. this Caddell study), suggest the possibility 1988; of the transformation, postflexion stage and of part latter affinities. other myomeres the dorsal fin and anus typically shift a few forward. largely and light is pigmentation Larval in collected larvae and Nearly all anchovy eggs gas of the bladder, gut, the dorsal to surface limited the CalCOFI ichthyoplankton samples are Engraulis the caudal the fin through the late post- ventrum, and mordax, abundant fish species collected the most stage. flexion during Anchoa larvae apparently are CalCOFI surveys. to bays, harbors, and estuaries. Larvae largely restricted area, study the In larval engraulids are CalCOFI as mysticetus Cetengraulis have identified tentatively be clupeids. The larval with confused to likely most Gulf taken been in the of California. engraulids commonly the than myomeres fewer have (overall clupeids with species most 38-47, range :::;;43 small are Anchovies (most ca. 2-37 range cm; <20 and (usually 40-58, vs. a shorter preanal length ~43) that cm), primarily planktivorous fishes school in inserts The dorsal BL). >75% usually vs. BL <75% fin coastal Adults water. fresh in live species Some waters. after the in myomere preanal last the and engraulids slightly to strongly compressed, with preanal length are last the before clupeids. the in myomere preanal about half a with to two-thirds of body length, and more engraulid larvae of is species the Identification elongate mouth, inferior lower large jaw. and snout, Presently, characters known problematic. allow no the although fms, one the in first spines no are There discrimination certain of one from species Anchoa the anal two or dorsal and spine- fin rays are small and from Anchoa, before to or another, Cetengraulis mid- brown Anchovies like. are to (occasionally blue) green late postflexion stage. Geographic location, together species have a dorsally and silvery below; some with Table (e.g., counts fin-ray and Engrauli- myomere Anchovies prominent midlateral silver stripe. are dae in identification species cases, allow some may 1) among the most economically important fishes, sup- of will commonly more number but the reduce only (White- of the world's largest fisheries porting some Larval potential species. three or two to identifications 1985; are They 1988). used al. et head Whitehead from distinguished easily relatively mordax are E. some fisheries, bait primarily in reduction and although basis Anchoa and Cetengraulis on the several of direct for species are prepared in human various ways E. mordax typically is more characters, for example consumption. pigmented elongate and slender, and is more heavily 173

186 than the Most useful is the dorsal other two genera. retained augmented perhaps and is apparently tation always is present which nearly after pigment, hindgut of the early the much specimens Panamanian in while always but mordax, E. is nearly in absorption yolk the postflexion of is lost part early the by pigment gut late before absent postflexion stage in the C. mysticetus stage). literature the from obtained were data Meristic The principal Anchoa. in transformation before and 1951; Howard Peterson 1956; Nelson (McHugh 1954; 1983, and Berry 1984; White- McGowan 1986; 1984, and caudal fin rays begin to form before the dorsal while true is E. mordax, opposite the anal fin in rays and 1995c) Rodriguez-Sanchez counts from head and and made this study. counts during anal fin ray Dorsal for Cetengraulis. Anchoa and and both the branched rays include the first 1-4 (usual- ly rays 2 (sometimes unbranched sepa- 3) or counted The descriptions are based on literature following rately and indicated with numerals, lower case roman of specimens 8-33 of examinations detailed on and in all, at counted not or literature). Ecological the Engraulidae and larval The 2). each species (Table (1956), Eschmeyer et al. is from information Peterson juvenile described Gulf from the C. mysticetus are here Rodriguez-Sanchez and and (1983), Whitehead Nicoya, of Gulf the from and Panama, of Rica. Costa limits the (1995c). Peterson (1956) gave northern range the from mysticetus C. tentatively as identified Larvae and curta Anchoa of Juanico, Bahia San as exigua A. are Gulf of California somewhat more heavily pig- Baja Rodriguez- and Whitehead while Sur, California mented the mid-preflexion through mid-postflexion in the to only ranging both show main- (1995c) Sanchez (in specimens Central the than stages Gulf American of California. of Gulf of the lower coast land pigmen- all specimens California stage early preflexion Table Engraulidae characters and geographic ranges for the engraulid species in the California Current vicinity. Meristic 1. 10+9 caudal fm rays. Abbreviations: BC, Baja principal BCS, Baja California Sur; CA, California; California; All species have Gulf, Gulf California. of Vertebrae Fin rays Gill Prev CV Total D A Species C P PI z z rakers Range 20-23 39-41 12-14 29-33 13-16 19-22+ 18-19 8-9+ Anchoa 7-8 Bay CA-Bahia Morro 6-8 24-27 compressa Todos BC Santos 14-16 19-22 19-22 39-42 21-27 8-10+ 13-14 6-8 16-20+ curta A. Bahia Juanico San 8-10 22-26 BCS-Ecuador 13-16 19-21 19-21 delicatissima 39-41 23-27 12-14 A. 18-21+ 8-9+ 6-8 CA-Bahia Long Beach 26-32 7-9 Magdalena BCS 14-15 18-23 12-15 22-24 43-46 20-23 9+9 7 17-23+ exigua Bahia San Juanico A. 22-28 BCS-Panama 14-15 22-24 41-45 17-22 16-17 6-9+8 19-21 6-8 ischana 14-19+ A. Magdalena Bahia BCS 17-23 & Gulf-Costa Rica 17-20 19-22 38-42 13-15 26-32 8-10+ 14-15 16-20+ 6-7 A.lucida Bahia & Magdalena 7-8 18-22 Gulf-Peru 8-10+ 14-16 19-21 19-22 39-42 14-16 22-28 6-7 17-24+ nasus Bahia San Juanico A. 8-10 20-28 BCS-Peru 18-20 16-24 40-42 13-14 30-35 13 7 9-11+ 17-22 walkeri Lucas San A. & Cabo lower 7-9 Gulf-Panama 19-21 19-22 40-41 28-32 14-16 Anchovia 13-14 6-7 8-10+ 70-135 Bahia Magdalena & 7-8 total Gulf-Peru macrolepidota 9-10+ 21-23 39-43 13-17 18-26 19-20 6-7 15-17 Cetengraulis 63-145 Los Angeles CA-Peru mysticetus 7-11 total 24-26 19-21 43-47 14-19 19-26 moniax Engraulis 13-20 6-8 7-10+ 28-41+ British Columbia--Gulf 7-10 37-45 174

187 Table Engraulidae Number of specimens (above) and size ranges (in mm, below) used in the preparation of the engraulid 2. descriptions. literature "L" indicates species used in the description. An Egg Postflexion Flexion Preflexion Transformation Yolk-sac Species Juvenile La La La La 5 0 compressa Anchoa 3 2.5-3.2 28.5-38.2 La La La 5 U 0 5 A. delicatissima 2.4--3.0 44.7-62.3 Lb Lb 9 3 7 5 3 Cetengraulis mysticetus 6.1-6.4 6.5-9.3 12.0-23.8 23.7-28.6 32.3-34.0 C 6 L 5 9 6 3 4 Engrauiis mordax 7.6-12.0 13.7-26.2 2.7-3.2 2.8-8.1 28.5-36.7 49.1-64.3 a Caddell 1988 b Simpson 1959 c Bolin 1936 175

188 Anchoa compressa ENGRAULIDAE HISTORY EARLY LIFE DESCRIPTION MERISTICS EGGS Mode Range diam.: Shell 0.63-0.90 mm Yolk: Segmented Vertebrae: Diam.ofOG: No.ofOG: 0 40 39-41 Total Shell Smooth surface: 18-19 19 Precaudal None Pigment: 21 20-23 Caudal yolk; segmented Round to features: Diagnostic no slightly elliptical; Fins: OG 0 0 Dorsal spines 13-14 12-14 Dorsal rays LARVAE 0 0 spines Anal mm Hatching 1.5-2.5 length: 31 29-33 Anal rays 10.0-10.5 mm 5.5-6.5 mm through Flexion length: 7-8 7 Pelvic mm, <28 mm >20 Transformation length: 14 13-16 Pectoral Fin sequence: development , , D, A, , C PI P C Caudal: 2 I 2 dorsolaterally along gut & on ventral Pigmentation: Yolk-sac-Little 10+9 10+9 Principal Preflexion-flexion--On isthmus; 1-6 dorsolaterally tail. of margin Procurrent: gas to anterior gut along ca. at beginning bladder gas on bladder; 8-9 8-9 Upper ventral 7 during few mm; number on in margin of hindgut, decreasing 6-7 6-8 Lower over flexion stage; rarely 1 or 2 anteriorly hindgut; rarely 1 or 2 over rakers: Gill end tip, of tail; under notochord on hindgut; of ventral margin 19-22 Upper on Posiflexion-Internally along developing Crays. becoming aligned 24-25 24-27 Lower brain after 12 along on urostyle 15 mm; cleithrum after 11 mm; internally 10-13 Branchiostegals hypurals; lower & posteriorly mm; 16.5 after colunm vertebral along increasing on C. LIFE HISTORY through features: Myomeres 38-41 (20-26 Diagnostic preanal myomere (postanal 5-7 2 above A ray D insertion stage); postflexion Bahia Range: California, Sur California Baja Magdalena, Bay, Morro to postflexion through 3) or pigment no or little count; A ray high stage; with compared hindgut; over none) (usually Engraulis mordax- Habitat: estuaries Schools in shallow coastal waters, commonly in bays & preanal body, deeper relatively given at size smaller length, shorter stage, developmental pigmented. lightly more & Summer season: Spawning %) MORPHOMETRICS (range & mean in & larvae eggs planktonic Oviparous; pattern: ELH Y-S Tr PoF* F* PrF* Juv LITERATURE 61-72 69-72 Sn-AlBL 52-54 70-76 70-75 1988 Caddell 68 73 72 71 53 1892 Eigenmann 7-15 5-10 4-5 17-19 BDIBL 6-7 4 10 19 7 6 length 1) Values * larval using Appendix (1988: Caddell from calculated HUBL 17-23 14-18 16-22 16-27 25-26 each represent exclusively 4.00-4.99; that classes stage: preflexion, 20 15 19 18 25 mm; 7.00-8.99 flexion, 11.00-19.99 mm. postflexion, 46-59 HWIHL 33-38 51 36 7-15 13-15 SnLIHL 12 14 21-29 23-30 21-27 25-27 EDIHL 33-43 26 24 26 38 26 14-16 0-0 PILlBL 15 0 9-10 P 0-0 L1BL 2 0 9 176

189 compressa Anchoa anchovy Deepbody . ., ... . ' ... -. ...•. ', . ... . . _\ ... ... ... ~... ..-" , .. , ... . , ... 2.2mm 0.76-0.84 mm mm 3.1 8.2mm mm 11.7 Figure Engraulidae 1. Egg, 0.76-0.84 mm; Anchoa sp. yolk-sac larva, 2.2 mm; Anchoa sp. preflexion larva, 3.1 mm; Anchoa sp.late flexion larva, A. compressa postflexion 11.7 mm (Caddell 1988). larva, 8.2 mm; 177

190 ENGRAULIDAE delicatissima Anchoa MERISTICS DESCRIPTION HISTORY LIFE EARLY Mode Range EGGS Vertebrae: mm Shell diam.: x 0.48-0.63 0.81-1.07 Yolk: Segmented 39--41 40 0 ofOG: Total No. Diam.ofOG: 19-21 20 Precaudal Smooth surface: Shell 19-21 None 20 Pigment: Caudal Fins: mm;segmented features: average 0.57 0.91x Elliptical, Diagnostic unpigmented yolk; o 0 Dorsal spines 13-16 15 Dorsal rays LARVAE Anal spines 0 o 25 23-27 rays Anal mm 2.0-2.5 length: Hatching mm Pelvic through mm 6-7 7 9.S-11 6-8 length: Flexion 12-14 13 Transformation length: Pectoral Fin development sequence: , , C C" A, D, P Caudal: P, 2 2 10+9 10+9 Principal gut; few on ventral Pigmentation: Yolk-sac-Few dorsolaterally along Procurrent: oftai!. margin Preflexion-flexion--2-4 dorsolaterally isthmus; few on 8-9 8-9 of Upper bladder; dorsally foregut; along gas on margin ventral on few 7-9 8 Lower flexion hindgut, decreasing during on stage; few ventral margin of tail; along Crays. developing & finfold caudal on ventrally little rakers: Gill 18-21 Upper 15 after hindbrain & mid- on Pos!'flexion--Internally on mm; 26-32 after mm; increasing on gas bladder, 11 margin of cleithrum ventral Lower 11-12 11 Branchiostegals none ventral margin tail, of & C; internally over caudal hindgut; on vertebrae after hypurals. lower urostyle on intema1ly mm; 16.5 & LIFE HISTORY 20-26+14-18=38-42 (usually Diagnostic features: Myomeres posttlexion stage); through 14-16 24-25+ D insertion at postanal early stage; posttlexion through 6-8 rays A above 2-3, myomere no or little Beach Range: Long Harbor, California, to Bahia Magdalena, Baja 23-27; A rays hindgut; on dorsally none) (usually pigment compared Sur California D with mordax-deeper-bodied, greater Engraulis between overlap D insertion A origin, ray C, before begins formation ray A & & & bays, in estuaries, harbors Habitat: Coastal primarily waters; schools formation, & more lightly pigmented. coast occasionally near shore along open MORPHOMETRICS (range & mean in %) season: Summer Spawning eggs ELH Oviparous; larvae & planktonic pattern: Y-S F* PrF* Tr PoF* Iuv Sn-AlBL 69-74 71-75 LITERATURE 66-76 59-72 53-55 72 74 71 67 54 1988 Caddell 3-6 5-7 5-10 7-15 BDIBL 18-20 4 6 7 II Eigenmann 1892 19 11-18 16-21 16-23 HUBL 17-23 23-26 19 14 19 20 25 larval from Caddell (1988: Values length * calculated Appendix using 2) 42-58 HWIHL 32-44 classes preflexion, stage: each that exclusively represent mm; 3.00-5.99 49 39 mm. posttlexion, mm; 7.00-8.99 flexion, 12.00-19.99 7-11 SnLIHL 18-21 8 19 32-38 23-36 21-30 19-32 EDIHL 30-31 34 30 26 24 30 0-3 P,LlBL 9-15 I 12 P L1BL 0-0 8-9 2 0 9 178

191 Anchoa delicatissima Slou.gh anchovy - ... .. " .. •... - , ... . ... ... · .••. \ ~~ ... \ .. ... ~0 /.,:~::·~· ~ ~._... .••• f ••••••••.•• _ \ ',- ',r.-: '.- 2.2 mm 0.81-1.07 x '. .. ", ... ',:"', ..•.. ". ". , 1\ -' . mm 0.48-0.63 ' ... '. : '. . ... '. .""'. :. ~ ... . "'''--'. ~ ... ...:... '" ,', ',_"i. '.'" ,," .. ,', -',,"" .. :-',' ... \ ," ._.,.'-'- ,.-_... """'" ... /' ,. "-:':-":".',-"----- - ..'" .. ... _-_.-_ ... . -"'", ... _-... , -' ... ~~~ mm 3.1 9.0mm mm 15.5 Figure Engrau/idae 2. Egg, 0.81-1.07 x 0.48-0.63 mm; Anchoa sp. yolk-sac larva, 2.2 mm; Anchoa sp, preflexion larva, 3.1 mm; A. delicatissima delicatissima flexion larva, 9.0 mm; A, late postflexion larva, 15.5 mm (Caddell 1988). 179

192 ENGRAULIDAE Cetengraulis mysticetus MERISTICS DESCRIPTION EARLY LIFE HISTORY Range Mode EGGS Vertebrae: Segmented diam.: mm Yolk: x 0.49-0.66 mm 0.99-1.4 Shell 39-43 41-42 Total OG: Diam. of of OG: 0 No. 21-23 surface: Shell Precaudal Smooth 19-20 Caudal Pigment: None Fins: 1.2 features: x Elliptical, Diagnostic average 0.56 segmented mm; 0 o Dorsal spines yolk; unpigmented 13-17 15 rays Dorsal Anal spines 0 o LARVAE 21-23 18-26 rays Hatching length: mm Anal ca. 2 6-7 Pelvic 6 Flexion length: mm 6.5-7.5 16 15-17 23-30 ca length: Transformation Pectoral mm Caudal: , , PI , C P C A, & D sequence: development Fin I 2 2 10+9 10+9 Principal on dorsolaterally Yolk-sac-Few Pigmentation: ventrally on foregut & Procurrent: hindgut isthmus; near stage. Preflexion-flexion-I-3 on of tail end & 9-10 Upper 9 on posterodorsally pair dorsolaterally 0-1 1-4 on foregut; cleithrum; 7-1I 9 Lower margins of hindgut on tail; little ventral 4-6 each & on gas bladder; Gill rakers: head at Posiflexion-External dorsally on on region. ventrally caudal 63-145 Total* ca. 21 by 21 mm; on opercle at 18 mm; internally on mm; brain I Upper of tail, gas cleithrum, isthmus, on increasing margin ventral bladder, 25-60 Lower after 18 mm; decreasing to 0-2 ventrally C gut by 14 mm; & on 7-9 8 Branchiostegals on gut after ca. 19 internally & mm; dorsolaterally dorsally increasing caudal in dorsal beginning margin on mm; 18 after peduncle HISTORY LIFE posteriorly after snout increasing on Transformation--On 21 mm. & mm; heavy on head at 23 gut; spreading cephalad on dorsal margin; laterally on trunk & tail. Peru, to California, Angeles, Los Range: California of Gulf including 40-43 (23-25+15-18 through early Diagnostic features: Myomeres to posttlexion stage, shifting 20-23+19-22 D transfonnation); by & Habitat: open along coast, estuaries, bays, in schools waters; Coastal A 5-8) through 2-4 rays (above myomere postanal at inserts well often offshore & becomes elongate & snout pointed coils hindgut stage; posttlexion mm. 18 before hindgut dorsal no transfonnation; during pigment Fall-winter (panama) with December peak Spawning season: mean %) MORPHOMETRICS (range & in & planktonic Oviparous; pattern: ELH larvae eggs y-S PrF LITERATURE F PoF Tr Juv Sn-A/BL 73-77 73-75 64-72 61-63 63-65 Simpson 1959 74 75 68 62 64 ORIGINAL ILLUSTRATIONS (Illustrator) 7-9 BDIBL 8-9 9-13 16-20 22-23 8 8 18 II 23 mm 6.1 larva, Pretlexion C. Walker) (R. HURL 16-18 18-20 20-24 27-31 33-35 (R. mm Walker) C. 7.8 Flexion larva, 19 17 22 34 30 mm Walker) (R. C. mm, 14.0 21.7 larvae, Posttlexion HWIHL 54-66 46-57 32-34 36-46 25-28 specimen, 28.6 Transfonnation Walker) mm (R. C. 52 59 40 33 26 SnLIHL 21-29 18-19 16-23 16-17 17-17 18 24 19 16 17 adult increasing of gill length. increases with rakers number * The 27-30x EDlHLt 22-27x 20-22x 19-23 x 18-21x or stage; posttlexion by so nearly becoming eye is oval, t The round 22-26 17-26 18-22 22-23 20-22 vertical axis second. given axis is first, horizontal 29x24 24x23 22x21 21x22 20x21 2-3 PILlBL 3-4 2-4 12-12 6-1I 2 3 3 8 12 P L1BL 0-0 0-0 0-8 8-9 9-9 2 0 0 5 9 9 180

193 Anchoveta mysticetus Cetengraulis 1.99 mm mm 6.1 x 0.99-1.36 mm 0.49-0.66 7.8 mm mm 14.0 mm 21.7 mm 28.6 Engraulidae 3. Egg, Figure x 0.49-0.66 mm; yolk-sac larva, 1.99 mm (redrawn from Simpson 1959); preflexion larva, 6.1 mm; 0.99-1.36 flexion larva, 7.8 (IATTC, mm of Panama 1721L); postflexion larvae, 14.0 mm Gulf Gulf of Panama 1687L), 21.7 mm (IATTC, (IATTC, Gulf of Panama 1721L); transformation specimen, 28.6 mm (IATTC, Gulf of Panama 1687L; developing scales are not shown). 181

194 ENGRAULIDAE Engraulis mordax MERISTICS EARLY LIFE HISTORY DESCRIPTION EGGS Range Mode Vertebrae: Shell diam.: mm 1.23-1.55 x 0.65-0.82 Yolk: Segmented 45-46 No.ofOG: Total 43-47 0 Diam.ofOG: Smooth 25 Shell surface: 24-26 Precaudal 20-21 None Pigment: Caudal 19-21 Fins: Ellipsoidal shape; size Diagnostic features: 0 0 spines Dorsal LARVAE 14-19 16-17 Dorsal rays Hatching length: mm 2.5-3 Anal spines 0 0 19-26 22-23 rays Anal length: Flexion mm through 10.5-13.5 6.5-10 mm 6-7 Pelvic 35-40 6-8 mm Transformation length: ca. 16 Pectoral 13-20 , C" P PI D C A, & , sequence: development Fin 2 2 Pigmentation: Yolk-sac-Few form gut on & on ventral Caudal: dorsolaterally 10+9 10+9 stage. through mid-way tail of on Prejlexion-jlexion-l-few Principal margin Procurrent: PI c1eithrum few near isthmus; origin by 11 mm; on dorsolaterally 9 7-10 Upper level of gas bladder, to dorsal over hindgut; on becoming gut along 9 7-10 Lower notochord under tail; of margin ventral on hindgut; of margin ventral becoming developing along proximal tip, Crays. Posiflexion- Gill rakers: 38-40 28-41 Upper mid- over on mm; 18 after opercle on Dorsally & hindbrain & 41-43 37-45 16 increasing on Lower hindbrain after isthmus, mm; cleithrum, ventral gut, 11-14 14 Branchiostegals margin C; & tail, of midlaterally mm; 16 by hypurals on internally on beginning posteriorly by ca. body 18 mm. Transformation-On HISTORY LIFE body posteriorly, extending length dorsum full of beginning by ca. 37 mm; mm; snout & mandible by 37 on increasing generally, especially on dorsum, midlaterally on & body, on gut Queen Range: Island, British Columbia, to Cabo San Lucas, Charlotte Diagnostic features: Myomeres 43-47 (26-30+13-18 to late Baja California Sur, California of Gulf lower & postanal 2-4 (at myomere above D insertion stage); ray A postflexion postflexion through 2) or I more hindgut; on pigment dorsal stage; coastal waters in Schools Habitat: stage, at larger elongate, given & Anchoa. more pigment than the year with late season: winter-spring Throughout Spawning peak, late %) MORPHOMETRICS (range & mean in minimum summer-fall larvae pattern: ELH eggs Oviparous; & planktonic Y-S PrF F PoF Tr Juv Sn-AlBL LITERATURE 60-71 70-77 74-79 64-67 65-76 64-66 68 74 76 72 65 66 1965, Ahlstrom 1966 BDIBL 8-11 7-9 8-9 8-10 12-14 16-18 8 8 9 9 1936 Bolin 13 17 Caddell 1988 15-19 HLlBL 14-17 16-18 18-22 22-26 30-32 Ahlstrom 1968 & Kramer 16 17 17 19 24 31 Matarese Sandknop 1984 & 49-67 40-69 HWIHL 45-56 34-44 33-36 24-29 al. 1989 et Matarese 59 59 49 40 35 27 Berry 1984 McGowan & SnL/HL 4-10 16-28 21-29 19-27 17-19 18-20 Moser & Ahlstrom 1985 8 21 25 23 17 19 Wang 1981 EDIHL 37-43 22-33 21-26 20-24 24-26 21-26 40 27 23 21 25 25 2-4 2-4 0-0 3-4 PILlBL 4-6 13-14 0 3 3 3 5 14 P L1BL 0-0 0-0 0-0 0-5 2 7-9 9-10 0 0 0 2 8 9 182

195 mordax Engraulis Northern anchovy .' 2.5mm x 1.23-1.55 7.5 mm mm 0.65-0.82 11.5 mm 18.4 mm mm 31.0 Engraulidae 4. Egg, 1.23-1.55 Figure 0.65-0.82 mm (Moser and Ahlstrom 1985); yolk-sac larva, 2.5 mm; preflexion larva, 7.5 mm; x flexion larva, 11.5 mm; postflexion larvae, 18.4 mm, 31.0 mm (Kramer and Ahlstrom 1968). Most stippling on the Kramer and Ahlstrom and (1968) is not pigment (especially along the dorsal and ventral margins illustrations on the head), but is shading which was intended to convey shape. 183

196 SALMONIFORMES H. G. MOSER Greenwood et al. a taxonomically (1966) hypothesized and these small characters (a added three others 8 37 and of suborders complex order consisting of the vomer, extension an anterior mouth, terminal subsequent methods by cladistic of Use work- families. of caudal the in supraneurallamellae the and presence ambiguities some resolved ers relation- concerning the al. distinct four defined (1984b) complex). et Ahlstrom ships the these fishes and brought question of into Bathylagidae, Microstomatidae, lineages (Argentinidae, monophyly (Rosen order the of Weitz- and Fink 1974; Argentinoidea, the within Opisthoproctidae) and based man Lauder 1982; and Leim 1983). Fink (1984) and and ontogenetic on adult characters. Begle (1991) suggested that Salmoniformes (sensu is longer since no useful, 1966) al. et Greenwood in these of systematics the Despite the great interest of salmoniform left has characters no evaluation of their fishes, phylogenetic relationships and those monophyletic Salmonidae. Begle group larger than primitive been not have general in fishes euteleostean for Osmerae taxon the proposed the non- 1992) (1991, we resolved. Eschmeyer's follow Hence, fully (1990) the Argentinoidei plus of order, members salmonid the includes that Salmoniformes sub- of classification several based Osmeroidei, on characters (e.g., fusion of and Argentinoidei, Lepidogalaxioidei, orders Esocoidei, a presence centra, ural to arches neural rudimentary of Salmonoidei. anterior cartilaginous vane on and basibranchials, the loss osmeroids, the of nuptual tubercles). Within the of the Salmonoidei and Early history stages life were defined by numerous characters Osmeridae California region. Argentinoidei are present in the of an anchor membrane on the the including presence Some osmerid of species smelts spawn intertidally, pattern the egg, caudal and skeleton, the in fusion of sand grains, while other to eggs coarse attaching their the palatine. of shape the reported larvae are species to spawn offshore. Osmerid are stations. taken occasionally in nearshore CalCOFI Monophyly two clades principal the of in superficially clupeoids resemble but lack They the the Argentinoidea superfamilies and Argentinoidei, layer differ transverse and clupeoids in muscle found of a is based on Alepocephaloidea, the presence and the gut. Eggs of pigment in the and structure crumenal a unique branchial outpocketing with organ, larvae of occur families argentinoid all CalCOFI in the structural specialized and Greenwood (Rosen elements and of the Alepocephalidae stages region. ELH Alepo- the of Begle Monophyly 1992). 1971; deep relatively Platytroctidae found water the in are opercle, reduced a dorsally 1) is based on: cephaloidea and, column are rare in CalCOFI consequently, sam- of the spine margin the dorsal on a specialized 2) individuals yolk-sac are eggs Their large, ples. and 3) opercle, specialization of 4) and basibranchial, the of larvae directly and Eggs juveniles. into develop al. rakers gill the on teeth (Begle et Ahlstrom 1992). are Microstomatidae and Argentinidae, Bathylagidae, Argen- four used synapomorphies to define (1984b) well represented collec- ichthyoplankton CalCOFI in 1) fins Bathylagidae are particularly abun- and of initial development of dorsal anal tinoidea: those tions and in ornamentation 2) fmfold, the of inner the of surface three these of stages ELH distinctive highly The dant. shell, 3) a unique rete mirabile in the swim the egg characters of array a large by identified be can families detail below. The spookfishes pala- and vomer in for tendency the 4) and bladder, described the tines to assume the function of the premaxillary and fishes (Opisthoproctidae) are relatively rare midwater samples. in CalCOFI rare are larvae whose of maxillary. (1991, 1992) fourth dismissed the Begle Alepocephaloidei and families included: Argentinoidei Suborders Argentinidae Alepocephalidae Bathylagidae Platytroctidae Microstomatidae Salmonoidei Opisthoproctidae Osmeridae 184

197 ARGENTINIDAE: Argentines MOSER G. H. Argentinidae family The includes genera and about two perivitel- large a moderately with mm), (1.3-1.85 large Atlantic, the regions coastal in distributed species 19 of globule, and line a large yolk, segmented space, oil Pacific, Indian oceans. The single argentine species and the surface pustules inner (visible in early stages) on of Argentina ca. CalCOFI area, hatch at sialis, ranges from larvae the Yolk-sac shell. with mm 3.5-5.0 the the in mouth that characteristic ventral on forms pigmentation to San the River south Cabo Columbia of elongate late-stage embryos. The larvae are with an Sur and Lucas, into the of Gulf California Baja Cali- of gut elongate series ventrolateral distinct a and distributional fornia. The range for larvae is from Pt. At blotches. end of the the preflexion stage a series of Arena, California, to Southern Baja with California, overlay ventrolateral blotches. Larvae California. Baja the off lateral bars begins to central highest abundance California. One also blotches median ventral postanal have two or of Gulf the in rare relatively are pigment bars. associated This pigmentation, along with (<22 and cm) medium-size the are argentines snout the in pigment embedded characteristic Adult silvery, unmistakable. the make opercular region, flat-sided fishes with a wedge-like head, larvae large eyes, and a small mouth 1964b). The jaws are tooth- (Cohen Argentina sialis is fm of tongue. less teeth present on the but The dorsal description following The are based examination detailed on 23 of (4 yolk-sac, larvae origin is just forward of the and at midbody pelvic fin 4 flexion, mm; 7.2-10.0 5 preflexion, 5.2--6.8 mm; fin adipose an with is far fin anal the above posteriad mm) 16.6-27.2 7 postfiexion, mm; 10.6-14.7 from are ventrolateral. it. The pectoral fins are Adults collections central to California southern off CalCOFI benthopelagic over the outer shelf and continental (37.0-47.9 mm) from California; 3 juveniles Baja also, There behavior. their of are known slope. Little is collections bottom-trawl (SIO) were examined in detail. off commercial fisheries for them in the Atlantic and the Meristic information was derived from literature Japan. and 1984b) al. from (Cohen 1964b; Ahlstrom et are of argentines relatively during this made study. original counts eggs The planktonic 185

198 Argentina sialis ARGENTINIDAE DESCRIPTION HISTORY LIFE EARLY MERISTICS EGGS Mode Range diam.: 1.3-1.7 mm Shell Yolk: Segmented Vertebrae: Diam. 0.27-0.46 1 No.ofOG: of OG: mm 48 47-49 Total Faint surface: surface inner on pustulation Shell 29 28-30 Precaudal Pigment: Nape; tip terminus; gut trunk; anterior tail of of embryo 19 18-21 Caudal Diagnostic Large shell featnres: inner pustulate OG diameter; faintly & Fins: embryo on pattern pigment characteristic surface; shell o 0 Dorsal spines 11 10-13 Dorsal rays LARVAE o 0 Anal spines Hatching length: mm 3.5-4.0 12 11-15 Anal rays length: Flexion 10.5-15.0 mm 11 10-12 Pelvic mm 35 ca. Transformation length: 15-19 16 Pectoral PI> C development seqnence: CI> D , P & Fin & A 2 2 Caudal: Yolk-sac-Blotch Pigmentation: 2 region; gut terminal above 10+9 10+9 Principal pairs juncture; ventrolateral above widely spaced along gut/trunk & Procnrrent: yolk above internally sac; below tail tip; ventral to anterior region of 12 10-13 Upper roof of series gut, blotches added above Preflexion--4 mouth. 12 10-12 Lower median becoming blotch blotch; embedded; postanal ventral & tail-tip rakers: Gill trunk bars beginning enlarged; embedded opercular streak; to form 7-9 Upper blotches. above Flexion-posiflexion--8 embedded gut blotches, 14-20 Lower bars, 3 postanal & 8 trunk of base; PI caudal; including maximum 5-5 5-5 Branchiostegals on opercle; ventrally along lower jaw; brain; & posteriorly anteriorly on P base 2 transformation. before & on D Juvenile-Bars just LIFE HISTORY become coalesce, saddles. of series straight; & features: Diagnostic elongate gut body; Elongate Columbia Baja Lucas, San Cabo of to Oregon, River, Mouth Range: later-developing with associated gut above blotches embedded trunk California Sur, into Gulf of & California bars; embedded pigment above roof opercular in of mouth region; & flexion & in protuberance fleshy slight blotch; caudal large snout on Habitat: bottom to m 274 On or near at depths larvae. postflexion stage Spawning season: Larvae present year-round with in CalCOFI samples & %) in mean (range MORPHOMETRICS in abundance peak January-March Y-S PrF F Tr PoF Juv Oviparous; planktonic eggs & pattern: larvae; demersal at ca. ELH 35 mm SL 75-79 78-82 77-78 72-73 Sn-AlBL 80-85 80 82 78 73 77 LITERATURE 8-10 9-10 10-12 15-16 BDIBL 9-18 12 10 11 15 9 a1. 1984b et Ahlstrom 15-19 16-19 20-28 20-22 HUBL 29-30 (Illustrator) ILLUSTRATIONS ORIGINAL 18 21 24 30 17 37-58 42-50 33-39 39-41 HWIHL 31-34 Egg, Arthur) MattsonIN. (G. mm 1.5 45 40 32 36 47 (G. Arthur) mm MattsonIN. larva, Yolk-sac 5.8 11-20 26-29 20-31 SnLIHL 32-35 26-30 15 34 29 27 24 25-32x 22-26x 23-27 x x 28-33 EDIHL* 20-27x * Eye trapezoidal oval, becoming round by postflexion stage; horizontal to 22-28 23-30 26-32 20-27 23-27 axis is first, vertical axis second. given 31x26 28x29 24x25 24x24 26x26 2-4 11-14 0-2 4-6 4-10 PILlBL 3 5 7 12 0.4 P L1BL 11-12 0-0 0-0 0-0 0-6 2 0 0 2 0 11 186

199 Pacific argentine sialis Argentina mm 5.8 : ... . 1.5 mm .. \ ... \~; ... ;--.', 'j :~, .. ::;;i..:~.~, .. ~:. :' ... ... '. . ... i .. 7.0mm 9.0mm 15.0 mm 21.0 mm Figure Argentinidae 1. Egg, 1.5 mm; yolk-sac larva, 5.8 mm (CaICOFI 5103, station 117.40); preflexion larvae, 7.0 mm, 9.0 mm; postflexion larvae, 15.0 mm, 21.0 mm (Ahlstrom et al. 1984b). 187

200 BATHYLAGIDAE: smoothtongues and Blacksmelts AHLSTROM' G. H. MOSER E. H. AND occur the California in bathylagids of species Ten pole of the egg (Ahlstrom I 969b). the vegetal In (Table regions Current adjacent and I). Bathylagidae Leuroglossus a migrate first globules the stilbius to Larvae of stilbius, Leuroglossus these, of three to coalesce blastodisc, the under position a smaller wesethi, Bathylagus the among ochotensis, B. and are divide number, to polar migrate and groups two into CalCOFI time series, most abundant species in the positions axis embryonic the (consid- to respect with ranking 5th, 12th, and in overall abundance. The 13th equatorial the in be to ered plane), then coalesce to of L. stilbius and B. wesethi are essentially distributions single globules, migrate embryo the below position to a Current, the California to confined while B. ochotensis before and, finally, coalesce further to a single globule zone in occurs subarctic-transitional to westward the (Ahlstrom 1969b). In hatching species Bathylagus of three of subarctic-transitional other Japan. Larvae embryogenesis whose cluster the documented, been has species, and of periphery B. pacificus, B. milleri, the initially migrates globules of to L. the schmidti, are CalCOFI samples. B. bericoides that abundant in much to migrate less groups two into blastodisc, then divides widely are apparently longirostris B. and in distributed of They there remain each during the remainder pole. the egg stage, undergoing some in coalescence partial are rare subtropical and tropical and seas their larvae species 1969b). (Ahlstrom in of CalCOFI the at the western margin samples survey larvae and Adults pattern. an of B. nigrigenys, occur species, the southern at Pacific eastern tropical have larvae are Bathylagid elongate and large CalCOFI the of 1971). (Ahlstrom pattern extreme finfolds in which the dorsal and anal fins develop. eastern Another urotranus, L. species, Pacific tropical the except is all of delayed caudal fins Development 1965). (Bussing region CalCOFI the in occur not does until the larval period; ray formation in in late the metamorphosis. The occurs fins pelvic and pectoral at (most medium-size are bathylagids Adult species relatively gut is straight with a valve separating the <20 cm) fishes that inhabit epi- bathypelagic zones to foregut long hindgut the valve another from and oceans. Some species have a deep body and a all of shorter terminal gut separating hindgut from the the some a narrow with robust are peduncle, caudal deep some section. diverse is development Larval with (Cohen peduncle, while others are slender caudal complex patterns eyes and species developing stalked small short snouts, and mouths, have They 1964b). pigmentation (Ahlstrom et Leuroglossus 1984b). of al. origin relatively large is at eyes. The about dorsal fin B. short stalks has ocho- eye whereas larvae early in of is at the origin fin the midbody, anal posterior third is tens stalks of has larval throughout length moderate the and the pectorals are ventrolateral. Some body, stalks Eye are longer in development. B. pacificus and are narrow gill openings and species colored dark with in long extremely B. bericoides B. longirostris. and paler with are others extending slits gill comparatively wesethi Bathylagus nigrigenys B. eyes small have and halfway up the body (Kobylyanskiy 1985). Most without stalks whereas moderately large B. milleri has Little is ha\'e of their species an adipose fm. known pigmentation varies eyes. Larval markedly. sessile behavior and there is no them. for fishery with Bathylagus a pigmented sparsely is bericoides melanophores the of inconspicuous along gut. series with oviparous are Bathylagids and eggs planktonic have milleri B. and pacificus, Leuroglossus,B. relative- of bathylagids stages Ontogenetic larvae, highly are ly isolated large the whereas blotches or melanophores evolved. Eggs of all known species have a pustulate trunk the region have serial gut and species in other multiple glob- yolk, segmented chorion, inner oil and pigmentation. head extensive and melanophores This glob- oil the Moreover, 1984b). et (Ahlstrom ules al. maximum reaches wesethi B. in pattern development of patterns remarkable undergo species some of ules nigrigenys, and which also are remarkable in having B. position initial coalescence from and migration at their with foregut a voluminous rugae. transverse Service, Laboratory, , Formerly, at La Jolla National Marine Fisheries California. Southwest Fisheries Science Center, La Jolla, 188

201 examination on and literature on based are descriptions Ahlstrom summarized knowledge of aL et (1984b) larvae 6-28 species each of juveniles and (Table of ontogeny and presented single illustrations bathylagid known eggs and of the summary and descriptions are to similar Eggs urotranus L. of 2). Bathylagidae larvae of the species discussed above. Matarese et at their of L. stilbius (Ahlstrom 196%) as are those larvae; since neither eggs nor larvae presented (1989) B. larvae urotranus of on information additional of L. B. ochotensis, B. pacificus, B. samples not is development their bericoides, CalCOFI B. mil/eri, in occur this described in guide. and schmidti, L. following The wesethi, stilbius. L. Bathylagidae 1. characters for the bathylagid species in the California Current vicinity. All species have 10+9 Table Meristic fin rays and 2 pairs of branchiostegal rays. principal caudal rays Vertebrae Fin Prey Total D A P, CV C, Species PI 8-12 9-11 17-22 8-10 12-15+13-17 52-54 22-24 29-31 bericoides Bathylagus 48-53 9-12 18-21 9-13 18-23 9-10 11-14+10-13 28-31 longirostris B. 9-16 49-55 18-21 6-9 20-28 6-9 14-18+13-17 28-34 milleri B. 9-11 8-10 11-15+11-14 14-18 10-14 41--45 21-26 17-21 B. nigrigenys 12-17 9-13 8-12 13-16 47-49 12-15+ 21-23 9-11 25-27 B. ochotensis 13-15+12-15 7-10 7-11 21-32 15-25 8-13 44-51 18-25 B. pacificus 10-14 13-17 9-12 8-1l 13-16+13-16 19-23 43--47 23-26 B. wesethi 8-9 8-10 10-17+11-15 10-11 11-14 47-52 25-27 22-29 schmidti Leuroglossus 8-11 11-15 9-12 39--42 8-10 11-16+9-15 18-21 20-21 stilbius L. 10-11 9 8 40--42 20-23 19-20 9 urotranus L. Bathylagidae 2. Number of specimens (above) and size Table (in mm, below) used in the preparation of the bathylagid ranges ilL" indicates literature used in the description. An species descriptions. Posttlexion Preflexion Flexion Transformation Juvenile Yolk-sac Egg Species 2 0 0 2 2 0 0 Bathylagus bericoides 14.2-15.2 5.9-7.5 16.4-23.0 6 3 3 4 3 0 B. longirostris 31.3-37.3 4.7-10.8 11.2-15.6 16.4-21.0 23.8-34.7 4.7 7 4 3 2 1 3 0 B. milleri 26.0-28.7 8.5-16.1 22.5-28.6 19.3-24.0 35.6-43.8 7.8 La 7 3 9 3 0 3 nigrigenys B. 12.0-21.2 9.4-11.3 22.5-28.0 5.0-9.1 3.3-3.6 La 6 2 2 3 7 2 B. ochotensis 12.7-16.2 16.4-26.6 26.0-27.6 5.0-11.3 24.9-26.4 4.8-5.0 2 7 3 3 3 0 20 B. pacificus 6.4-14.7 16.1-19.3 19.1-22.6 20.9-24.2 5.6-5.8 1.4-1.6 La 4 9 8 0 3 3 wesethi B. 4.5-9.6 10.4-12.8 14.0-23.7 22.4-28.2 3.7--4.2 La,b Lb Lb Lb Lb 0 0 schmidti Leuroglossus La 7 3 4 8 3 3 L. stilbius 27.2-30.3 15.4-27.4 10.5-14.7 6.2-10.0 28.8-36.4 4.2--4.3 a Ahlstrom et aI. 1984b b Dunn 1983 189

202 BATHYLAGIDAE Bathylagus bericoides EARLY HISTORY DESCRIPTION LIFE MERISTICS Mode Range EGGS Shell diam.: Yolk: Vertebrae: 53 No.ofOG: 52-54 Diam.ofOG: Total surface: Shell 29-31 30 Precaudal Pigment: 22-24 23 Caudal Diagnostic features: Fins: 0 0 Dorsal spines LARVAE 9 9-11 Dorsal rays Hatching length: 0 0 spines Anal 20 length: Flexion ca. 14 mm 17-22 rays Anal mm >23 length: Transformation 9 8-10 Pelvic Fin development sequence: & P PI D & A, CI> & C 8 8-12 Pectoral 2 2 Preflexion-posiflexion-Lateral on dorsally liver; to Pigmentation: Caudal: 10+9 of series terminal gut; section in surface; gut lateral along 11 to up of 10+9 Principal base, PI on larvae, some late postflexion opercle, lower jaw, isthmus, Procurrent: 14 12-15 of caudal peduncle. region & lateral Upper 15 13-17 Extremely long eye stalks, ca. Diagnostic features: in 50% HL Lower of 70% stage; ca. at sparse flexion preflexion maximum stage, Gill rakers: specimens; available in region gut to limited pigmentation, extremely Upper postflexion Atlantic long especially in larvae stage; differs gut, from Lower stalks & in brain 2 2 species the of in having relatively longer eye same Branchiostegals versus length, of optic lobe region than slightly greater shape (width equals width differs larvae); Atlantic in length its of 80% B. from HISTORY LIFE pigment & eye longer relatively /ongirostris in having less stalks. Widely distributed in tropical & subtropical Atlantic, Pacific, & Range: in %) (range MORPHOMETRICS & mean Indian Oceans Y-S F PrF Tr PoF bathypelagic Habitat: Epi- & Juv Larvae season: Spawning summer in collected & autumn 82-83 71-83 Sn-AlBL 90-91 83 77 90 larvae Oviparous; pattern: ELH planktonic eggs & 8-8 BDIBL 8-10 9-9 9 8 9 LITERATURE 22-23 25-25 HLlBL 24-27 25 22 26 a1. Ahlstrom 1984b et HWIHL 35-41 36-36 39-40 1989 a1. Matarese et 38 36 39 & 1991 Olivar Fortuno 11-13 SnLIHL 24-27 22-22 12 25 22 (Illustrator) ORIGINAL ILLUSTRATIONS 10-10x 12-l4x 18-19x EDIHL* 24-24 27-31 19-20 Head M. of 6.4 mm preflexion larva (H. Orr) IOx19 18x29 13x25 7.3 Orr) mm (H. larva, Preflexion M. 14.5 mm (H. M. Flexion Orr) larva, 3-3 2-5 PILlBL 4-6 4 3 5 P L1BL 0-0 0-0 2 first, axis given is axis horizontal elliptical; Eye * second. vertical 0 0 0 190

203 Bathylagus bel'icoides mm 6.4 ~ 7.3 mm 11.8 mm ... ," :' .: .f .. ,',:',"".f ... ... ' ... :, ... : ... : ... ... "",' ... ,',' ... ,' J ,/ 14.5 mm 17.7 mm Figure Bathylagidae 1. Head of 6.4 mm preflexion larva, dorsal view; preflexion larva, 7.3 mm (CaICOFI); head of 11.8 mm preflexion flexion larva, (Ahlstrom et al. 1984b); view larva, 14.5 mm (CaICOFI dorsal 7210, station 39.137); postflexion larva, 17.7 mm (from tropical eastern Atlantic, Ahlstrom et al. 1984b). 191

204 BATHYLAGIDAE Bathy/agus /ongirostris DESCRIPTION EARLY LIFE mSTORY MERISTICS Mode Range EGGS Sbell diam.: Vertebrae: Yolk: 48-53 51 No.ofOG: Total Diam.ofOG: Sbell 28-31 surface: 29 Precaudal 18-23 22 Pigment: Caudal features: Diagnostic Fins: 0 0 Dorsal spines LARVAE 9-12 10 rays Dorsal length: Hatching 0 0 spines Anal 18-21 Flexion mm length: 19 11-16 Anal rays 9-10 10 mm Transformation length: 24-35 Pelvic 11-12 , PI> Pectoral sequence: P & D Fin 9-13 C), C A, development 2 2 ca. gut; on laterally 12-14 of Yolk-sac-Series Pigmentation: in 7 Caudal: 10+9 10+9 postanal Pre flexion-Lateral gut becomes series Principal midline series. ventral Procurrent: subdivided (series terminal hindgut, up to section, on 8-12 on 4-5 of 11-14 11 Upper on patch foregut); on 12 ca. series liver; to 3 lateral of series opercle; 10-13 11 jaw from subocular, jaw; lower to isthmus; at 3 embedded of Lower on tip series opercle; to p) from myomeres hypaxial on C; below of Gill rakers: 8-9 8 foregut & Flexion-posiflexion-Hind- to Upper increase series notochord. 14 13-18 entire in body series 12-15; on hypaxial myomeres late along Lower 2 2 base. p) of Branchiostegals postflexion stage; on snout; on anterior surface epaxial myomere series. Traniformation-Several per in hypaxial & HISTORY LIFE HL in 53% average stalks, eye long features: Diagnostic Extremely preflexion & heavy stage; postflexion flexion in stages, decreasing pigment myomeres. & gut on series & subtropical Atlantic, Pacific, & Range: in tropical distributed Widely Indian Oceans MORPHOMETRICS (range & %) in mean Habitat: Epi- to mesopelagic Y-S PrF PoF F Tr Juv Spawning season: Larvae captured from April-October in CalCOFI Sn-A/BL 91-94 88-91 79-87 surveys 81-82 74-75 79 83 89 92 82 74 larvae & eggs planktonic Oviparous; ELH pattern: 7-9 BDIBL 8-10 9-12 11-14 15-16 6 9 9 12 10 15 LITERATURE HUBL 21-26 24-26 26-28 22-25 24-27 25 17 25 27 24 26 aI. 1984b et Ahlstrom HWIHL 30-42 36-48 40-43 42-52 44-51 40 41 37 41 47 48 ORIGINAL ILLUSTRATIONS (Dlustrator) 13-15 13-15 SnLIHL 16-22 28-33 27-29 14 14 8 19 28 30 mm (H. 9.1 M. Orr) Late larva, preflexion 12-22x 11-14x 13-14x EDIHL* 15-17x 21-26x 19-32 16-20 17-19 17-20 25-31 25x35 15x24 13x19 13x18 16x19 24x28 elliptical; * Eye second. vertical first, given is axis horizontal axis P)LlBL 2-4 2-4 7-11 4-6 0 3 3 3 5 9 P L1BL 0-0 0-0 0-0 2-4 5-7 2 0 0 0 0 6 3 192

205 Longsnout blacksmelt Bathy/agus /ongirostris 9.1 mm 12.4 mm 20.1 mm Figure Bathylagidae 2. Late preflexion larva, 9.1 mm (CaICOFI 7210, station 31.143); head of 12.4 mm flexion larva, dorsal view; 20.1 mm (Ahlstrom et al. 1984b). postflexion larva, 193

206 BATHYLAGIDAE Bathy/agus milleri EARLY LIFE HISTORY DESCRIPTION MERISTICS Mode Range EGGS Yolk: Shell diam.: Vertebrae: 52 Diam.oCOG: No.oCOG: 49-55 Total surCace: Shell 19 18-21 Precaudal Pigment: 31 28-34 Caudal Ceatures: Diagnostic Fins: 0 spines Dorsal 0 7 LARVAE rays Dorsal 6-9 mm Hatching length: 5 ca. 0 0 spines Anal 19-24 mm 23 Flexion length: 20-28 rays Anal 7-8 Transformation length: 26-29 mm 6-9 Pelvic sequence: & C 12 Fin development C , A , PI> D & P 9-16 Pectoral I 2 2 Prejlexion- tip. Pigmentation: Yolk-sac-B1otch anterior to notochord Caudal: 10+9 of juncture at pair Large section gut terminal where trunk & gut 10+9 Principal begins; below to liver, then moves anterior to liver); base PI (lateral Procurrent: I above embedded forebrain; hindbrain; on lower jaw; at midgut; 14 14-18 Upper 13-17 15 lateral on postorbital; opercle, surface of Flexion- base. PI Lower pos{/lexion--On of region lower head at jaw; upper nares; posterior rakers: Gill jaws; to Upper fin A I at dorsal margin & at ventral margin (at Ad & bases); 16-18 15-20 I on I at dorsal margin above anus; postanal ventral on margin; Lower 2 2 terminal gut of blotch over hypural region. section; ventral surface Branchiostegals Relatively large unstalked eyes, nearly Ceatures: Diagnostic short gut; isolated orientation; round, developing large slight dorsolateral HISTORY LIFE melanophores & clumps in distinct pattern; large size at hatching, notochord flexion, pigment larval transformation; & in visible Range: to Bering Sea & west to Japan Southern California to ca. 50 juveniles mm SL; low preanal up myomere count (ca. high postanal 24-25), myomere count (ca. & high total count 27-28), mesopelagic to Epi- Habitat: high (6-9); count ray D low (49-53); count ray A (20-28). in surveys CalCOFI in season: abundant most Larvae Spawning MORPHOMETRICS in & (range %) mean December-February Y-S ELH pattern: Oviparous; eggs planktonic PoF PrF & larvae F Tr Juv Sn-AIBL 58-60 54-63 60-62 LITERATURE 63-65 59--61 59 60 56 61 64 60 1984b al. et Ahlstrom 8-11 8-10 BDIBL ll-14 12-13 20-20 12 10 9 5 12 20 Matarese et 1989 al. 18-20 HUBL 18-20 20-25 23-25 24-27 (Illustrator) ORIGINAL ILLUSTRATIONS 14 18 19 22 24 25 52-54 53-59 49-59 HWIHL 61-65 49-50 mm larva (D. M. flexion Orr) 23.1 of Head 40 53 56 49 53 63 SnLIHL 27-38 9-28 27-28 27-31 16-20 15 21 32 28 29 18 vertical axis Eye oval; * horizontal axis is given first, second. 25-32x 24-25 x EDIHL* 20-24x 20-22x 33-38x t in juvenile specimens. broken rays Fin 29-35 26-30 22-24 24-28 36-38 35x29 29x33 24x28 23x26 21x23 36x37 1-2 1-3 0.4-1 PILlBL 3-4 t 0 1 1 2 4 P 0-0 L1BL 0-0 0-0.8 2 t 0 0 0 0.3 194

207 Robu.st blacks melt milleri Bathylagus \')"""------- --.<'/_---- -- ... ~ 9.5 mill mm 10.6 mm 23.1 .. ~ •...•. _- ____ ----"!~'7' 27.5 mm Figure Bathylagidae 3. Head of9.5 mm preflexion larva, dorsal view; preflexion larva, 10.6 mm (Moser 1981); head of23.1 mm flexion larva, larva, view (CaICOFI); postflexion dorsal 27.5 mm (Moser 1981). 195

208 Bathylagus nigrigenys BATHYLAGIDAE EARLY LIFE HISTORY DESCRIPTION MERISTICS EGGS Mode Range mm diam.: 0.83-1.09 Shell Segmented Yolk: Vertebrae: No. OG: of 12-20 Various OG: Diam. of 43 41--45 Total Pustulate inner surface Shell surface: 23 21-26 Precaudal migrates dorsum of Pigment: in ventrad On nape; on heavy embryo; 19 17-21 Caudal some sac; yolk adjacent to light embryos, late-stage on finfold Fins: globules clumped at oil multiple shell; Pustulate Diagnostic features: 0 0 spines Dorsal pigment some to poles; dorsal pigment migrates yolk sac; heavy nape 12 10-14 rays Dorsal 0 0 spines Anal LARVAE 15-16 14-18 Anal rays Hatching 3 mm ca. length: 9 8-10 Pelvic 9-12 mm Flexion length: ca. 10 9-11 Pectoral Transformation 22 mm length: ca. Caudal: sequence: A, C" & D, PI & P, 10+9 Fin C development 10+9 lo Principal Pigmentation: Yolk-sac-Coalescing ventrally pairs 8-10 into Procurrent: below & above to gut, from dorsolateral P I base to anus; cluster above 12-13 11-15 Upper on gut; to dorsolateral pairs preflexion-8-11 Early notochord tip. 12-13 11-14 Lower ventral surface optic each at 1 at lobe; ventral notochord tip; of rakers: Gill on symphysis; on c1eithral membrane; postorbital; branchiostegal 7-9 6--10 Upper late jaws; on snout. Mid- & lower preflexion-Dn upper & 15-16 14-19 Lower margin ventrolateral of hypural over patch notochord ventral head; 2 2 Branchiostegals of C rays; irregular & series begins; peppering region base hypaxial finfold & adjacent area; PI of surface lateral & inner on on ventral HISTORY LIFE on I or more ventral postanai base; midline; patch at c1eithral 1 or gut; on dorsolaterally more symphysis. Flexion-Up to 12 above Pacific tropical Eastern Range: dorsal on brain; laterally series; epaxial Poslflexion-Irregular finfold. on hindbrain; dorsally on peduncle. Habitat: mesopelagic to Epi- complex DiaglJostic features: Early migration of dorsal pigment; & & rugose foregut; pairs more unique pigment pattern; voluminous of Spawning year-round Possibly season: gut pigment ventral ventrolateral (6--8); wesethi B. in than (8-10) dorsal in tip pigment versus notochord & ventral B. wesethi; optic ELH planktonic pattern: larvae & eggs Oviparous; sparser series trunk pigment; lobe & more irregular than in B. wesethi; than vs. fewer total vertebrae B. wesethi (41--45 43--47). LITERATURE (range & mean in %) MORPHOMETRICS et Ahlstrom 1984b aI. Castillo 1979 Y-S PoF F PrF If Juv 1973 Rass & Pertseva-Ostroumova 91-96 75-77 86--91 Sn-AIBL 79-88 70-77 ORIGINAL (Illustrator) ILLUSTRATIONS 94 89 82 76 74 21-23 15-18 14-16 11-15 BDIBL 6-8 Mattson) Egg, 0.96 mm (G. 15 17 13 22 7 Yolk-sac Arthur) 2.8 mm (G. MattsonfN. larva, mm MattsonfN. (G. Arthur) Preflexion larva, 4.5 27-29 19-26 29-31 24-29 14-16 HLIBL 27 28 15 23 30 Head of 21.8 mm postflexion larva (G. MattsonfN. Arthur) 27.0 MattsonfN. Arthur) (G. mm Juvenile, 52-60 57-68 55-60 44-58 42--49 HWIHL 45 64 57 51 55 19-28 26-30 20-24 7-16 26--32 SnLlHL slightly second. oval; horizontal axis is given first, vertical Eye * axis 26 28 22 28 12 40--48 23-27x 17-24x 14-16x 11-17x x ED/HL* 17-18 12-19 24-31 17-26 32-35 15x18 43x33 20x21 15x16 25x27 1-5 2-6 2--4 8-10 0-0 P,LIBL 9 3 3 4 0 0-0 0-0 0-0 0-0.3 P,LIBL 0.04 0 7 0 0 196

209 Blackchin blacksmelt nigrigenys Bathylagus ~~iE,;;~_·e/_~ 2.8mm -0·················'··'················, ."'. .co-1'·····~···:;:: ~ ~ ., .".. .. ~ 0.96 mm 4.5mm 8.7mm .. .. .. 21.8 mm 21.8 mm 27.0 mm Figure Bathylagidae 4, Egg, 0.96 mm (Shellback, station 19); yolk-sac larva, 2.8 mm (Shellback, station 28); preflexion larva, 4.5 mm (Shellback, 86); head station of 8.7 mm preflexion larva, dorsal view; postflexion larva, 21.8 mm (Ahlstrom et al. 1984b); head of 21.8 juvenile, mm larva, dorsal view (Shellback, station 92); postflexion 27.0 mm (CaICOFI 5106, station 157.10). 197

210 BATHYLAGIDAE Bathy/agus ochotensis MERISTICS EARLY LIFE mSTORY DESCRIPTION Mode Range EGGS Vertebrae: mm Yolk: Segmented 0.92-1.1 Shell diam.: 48 47-49 Total OG: of Diam. 2 ca. to coalesce >10, OG: of No. Various Precaudal 25-27 26 inner surface Pustulate surface: Shell Caudal 21-23 22 Pigment: Fins: shell; lack of pigment; OG Diagnostic features: Size; pustulate Dorsal spines 0 0 coalescence 11-12 Dorsal 9-13 rays Anal spines 0 LARVAE 12-17 14-15 Anal rays 3 mm ca. length: Hatching 10 9-11 Pelvic Flexion length: ca. 12-16 mm 8-12 length: 10 Pectoral mm 26 ca. Transformation Caudal: , , P C sequence: development Fin & PI C A, D, 2 I 2 10+9 10+9 Principal base PI notochord Pigmentation: tip. Yolk-sac-On Pre flexion-Below Procurrent: gut to series of ca. 5 on lower trunk, liver; on terminal section; lateral 12-15 14 Upper increasing terminal gut pair stage; of end at 12 ca. to to increases on 13-16 14 Lower series extending forward isthmus; entire hindgut; internal series on to Gill rakers: series expaxial on myomeres above hindgut. Flexion-posiflexion- 8-9 8 Upper ca. 1 per myomere to from PI base to C; on Hypaxial increases series 13-20 Lower 16 & lower jaws; on laterally brain; on upper hypural; on opercle; 2 2 Branchiostegals postorbital regions. & otic Diagnostic features: ca. Stalked eyes with length increasing to stalk mSTORY LIFE ca. to decreasing mm, 9 ca. at 20% HL to & stage flexion at 12% 2-4% postflexion stage; notochord tip pigment in early preflexion in series hypaxial pigment on series epaxial body, entire along stage; Sea, & Baja Range: California north Central to Gulf of Alaska, Bering posterior 33% body; pigment series along of hindgut. Japan to westward MORPHOMETRICS (range mean & %) in Epi- Habitat: mesopelagic; depth ca & 50-900 m y-S Spawning winter-spring in highest abundance larval season: CalCOFI PrF F PoF Tr Juv peak with February-March in 71-76 78-84 Sn-AlBL 87-92 84-91 78-78 76-78 81 73 88 90 78 77 pattern: ELH Oviparous; eggs planktonic larvae & 7-9 7-8 BDIBL 9-11 11-14 15-15 16-16 LITERATURE 8 8 10 12 15 16 12-17 21-22 HLlBL 18-22 21-23 25-25 25-27 Ahlstrom 1972a 22 20 14 21 25 26 1984b a1. et Ahlstrom 37-53 38-53 HWIHL 47-50 46-48 41-45 42-45 et Matarese a1. 1989 49 48 45 47 43 44 13-13 11-22 SnLIHL 18-25 24-36 26-26 24-27 ORIGINAL ILLUSTRATIONS (Illustrator) 19 13 22 29 26 26 13-15x 13-19x EDIHL* 38-43 x 14-20x 21-25x 27-30x Egg, mm Mattson) 1.1 (G. 28-33 22-33 21-23 20-22 31-31 24-26 postflexion mm of21.5 Head larva (G. Arthur) MattsonIN. 40x30 14x22 16x28 16x20 23x25 29x31 0-0 1-5 3-5 PILlBL 4-5 8-10 7-8 Eye * vertical axis second. given is axis horizontal oval; to fIrst, elliptical 4 0 5 5 7 9 P L1BL 0-0 0-0 0-0 0-3 5-6 8-9 2 0 0 0 0.8 8 6 198

211 ochotensis Bathylagus Popeye blacksmelt mm 7.4 1.1mm 8.5 mm mm 13.0 mm 21.5 21.5 mm 25.5 mm Figure Bathylagidae 5. Egg, 1.1 mm (CaICOFI 5002, station 60.90); preflexion larva, 7.4 mm (Ahlstrom 1972a); head of 8.5 mm preflexion larva, view (Moser 1981); flexion larva, dorsal 13.0 mm (Ahlstrom 1972a); postflexion larva, 21.5 mm (Ahlstrom et al. 1984b); head of (CaICOFI); 21.5 postflexion larva, dorsal view mm juvenile, 25.5 mm (Ahlstrom 1972a). 199

212 BATHYLAGIDAE Bathylagus pacificus LIFE MERISTICS EARLY HISTORY DESCRIPTION Mode Range EGGS 1.4-1.6 Segmented Vertebrae: mm Shell diam.: Yolk: 45 44-51 to Total condensing (>50), Many OG: of No. 10-15 ca. 18 18-25 Precaudal Various, Diam. 0.02-0.2 of OG: mm 21-32 27 bumps Outer Caudal contiguous small Shell surface: with surface covered Pigment: Fins: features: shell 0 0 Diagnostic surface; numerous diameter; large Pebbled Dorsal spines 8-13 8-10 oil globules rays & Dorsal coalescing pole, vegetal at clumped size varied of forming 2 polar in late-stage eggs 0 0 groups spines Allal 15-25 20 Allal rays 7-10 Pelvic 8 LARVAE 10 3.5-4.0 Hatching length: mm 7-11 Pectoral mm 16-19 Flexion length: Caudal: 10+9 Transformation 10+9 mm 20-24 length: Principal PI & D A, C sequence: development Fin C & P & , Procurrellt: I z z 14 13-15 late larvae Upper 2 yolk-sac Pigmentation: Yolk-sac-At notochord tip; in 12-15 trunk, 13 other the & midbody at Lower (large) & gut of at juncture laterally 1 region at posterior liver, to flexion-Lateral hindgut. of Pre rakers: Gm to added 1 Upper on original 2; on terminal between section; gut ca. gut midway 17-19 13-20 jaw. Late preflexion-posiflexion-In some on lower opercle; Lower 2 2 specimens 1 original to 2 anterior or 1 & ones 2 between Btrallcltiostegais or original jaw; one at midbody; on upper embedded, lateral to hindbrain. with averaging Diagnostic 28% length stalk features: eyes Stalked HISTORY HL LIFE preflexion in % 7-11 in to decreasing stage, flexion in 33% & stage postfiexion late & transformation stages; 2 large blotch-like lateral Bering & Alaska of Gulf to Range: northern Sea Baja California; with melanophores smaller D low between; gut on ones count 1-2 ray westward between 54°-66° N to Japan high & count relatively A ray (15-25). (8-13) mesopeJagic at ca. 149-1,000 m depth Habitat: Epi- & %) MORPHOMETRICS & (range ill mean most winter- in abundant larvae surveys, In CalCOFI Spawning season: PrF Y-S F spring with PoF February-March Tr in peak Juv & larvae 81-84 ELH planktonic pattern: Oviparous; eggs Sn-AlBL 72-78 75-82 77-82 83-85 74 78 83 84 80 LITERATURE 9-11 7-9 8-10 BDIBL 11-13 13-13 8 9 10 12 13 et Ahlstrom 1984b al. 19-25 22-24 16-19 HLIBL 24-25 26-26 Matarese al. et 1989 22 17 23 24 26 42-47 36-50 40-46 HWIHL 48-50 46-50 ORIGINAL (Illustrator) ILLUSTRA nONS 42 44 42 48 49 11-19 20-28 SnLIHL 26-30 27-32 24--32 mm Arthur) 5.6 larva, Yolk-sac (N. 14 24 29 29 28 (G. mm MattsonIN. Arthur) 7.1 larva, Preflexion 1 15-18x x 7-23 36-37x 18-18x 15-16x EDIHL Arthur) (G. mm 16.0 MattsonIN. larva, Flexion 22-24 23-35 26-33 20-23 21-21 36x28 19x30 17x23 15x22 18x21 horizontal to oval, becoming axis round by transformation; Eye elliptical * 1-5 2-2 0-0 3-3 3-4 first, given axis is second. vertical 0 2 3 4 3 0-0 0-0 0-0 0-0 PzLIBL 0.05-2 0 0 0 0 1 200

213 Pacific blacksmelt pacijicus Bathy/agus 5.6mm '.! '/! =;; > .~.; •... ;L=z -->':>::'.-// /'/ '-, ... ... . ... ".' ~' G-rr,/yq ' ... ". ... ... ... ...,!: " ... ". '. ", '. . . . . ... ~;r'~""~ 7.1 mm 16.0 mm 21.4 mm Figure Bathylagidae 6. Yolk-sac larva, 5.6 mm (CalCOFI 8803, station 74.9.54.6); preflexion larva, 7.1 mm (CalCOFI 5204, station 70.55); postflexion flexion 16.0 mm (CalCOFI 5204, station 87.50); larva, larva, 21.4 mm (Ahlstrom et al. 1984b). 201

214 Bathy/agus wesethi BATHYLAGIDAE EARLY LIFE IDSTORY DESCRIPTION MERISTICS EGGS Mode Range diam.: 0.90-1.1 mm Yolk: Segmented Shell Vertebrae: of Various OG: of Diam. mm 12-20 OG: No. 44 43-47 Total Shell surface: Pustulate inner surface 24 23-26 Precaudal dense late of embryo, on head; on ventral less On dorsum Pigment: 21 19-23 Caudal fmfold Fins: Diagnostic features: Pustulate shell; differs from lack in B. nigrigenys 0 0 Dorsal spines patch pigment pigment of to migration no having nape in & of heavy 11-12 10-14 rays Dorsal yolk sac 0 0 Anal spines 16 13-17 rays Anal LARVAE 9 8-11 Pelvic Hatching 3 ca. length: mm 10 9-12 Pectoral 10-14 mm length: Flexion Caudal: mm Transformation length: 22-24 10+9 10+9 Principal P C Fin development D, PI sequence: , & A, & C I 2 2 Procurrent: migrating ventrad Pigmentation: Yolk-sac--Peppering over entire body, 14 13-16 Upper & coalescing. Pre dorsolateral pairs c1eithraljunction; flexion-At 6-8 14 13-16 Lower from gut to above gut anterior on region; surface terminal to base PI Gill rakers: lower & PI of margin tip; notochord to base; opposing pair anterior 9 8-10 Upper & lower jaws; opercular region; on upper postorbital; on lower 16-17 14-19 Lower embedded membrane; branchiostegal lateral cerebellum; to 2 2 Branchiostegals on head ventrolaterally series to jaws from postanal opercle; along midline; along ventral ventral peppering lateral gut surface & & LIFEIDSTORY hypaxial f1nfold. Flexion-8-IO dorsolateral gut pairs; associated Posiflexion-transformation- series from head to hindgut; on Crays. California Range: California Baja southern to Oregon from Current along 2 or then entire series epaxial & Hypaxial body, per more myomere. Habitat: mesopelagic to Epi- features: Eyes not stalked; complex Diagnostic pigment & unique gut ventrolateral of foregut; fewer pairs rugose voluminous pattern; year-round; some Spring-summer spawning season: Spawning with in than (6-8) melanophores nigrigenys B. ventral & dorsal (8-10); highest larval abundance in CaiCOFI surveys May-August in at notochord pigment tip nigrigenys; B. in ventrally only vs. trunk more & heavier series total more nigrigenys; B. in than regular ELH eggs planktonic Oviparous; pattern: larvae & vs. vertebrae than B. nigrigenys (43-47 41-45). LITERATURE (range mean MORPHOMETRICS & %) in 1972a 1969b, 1965, Ahlstrom Y-S PoF F PrF Juv Tr 1984b aI. et Ahlstrom et Matarese 1989 aI. 77-86 86-91 92-98 77-78 76-76 Sn-AIBL 82 77 94 88 76 (Illustrator) ILLUSTRATIONS ORIGINAL 7-9 13-14 8-14 17-18 13-18 BDIBL 17 15 14 11 8 larva Arthur) MattsonIN. of (G. postfiexion mm 20.5 Head Arthur) MattsonIN. (G. mm 27.0 Juvenile, 27-29 16-19 16-24 24-26 23-27 HLlBL 20 25 29 17 25 45-49 49-55 55-57 55--65 46-50 HWIHL * axis axis horizontal oval; first, slightly Eye vertical given is second. 46 56 61 52 49 29-34 25-30 20-25 20-31 16-18 SnLIHL 25 32 23 27 17 14-15x 11-14x 28-33x 31-34x 16-25x EDIHL* 30-35 19-32 16-18 13-16 23-27 13x14 33x25 30x32 20x25 15x17 8-10 4-7 5--6 4--6 0-0 PILlBL 5 6 6 9 0 0.6-1 6-10 0-0 0-0 0-0 L1BL P2 8 0 0 0.9 0 202

215 Snubnose blacksmelt wesethi Buthylagus S.7mm 1.0 mm mm 8.5 11.3 mm 20.5 mm 27.0 mm Figure Bathylagidae 7. Egg, 1.0 mm (Ahlstrom 1969); preflexion larva, 5.7 mm (Ahlstrom 1972a); head of 8.5 mm preflexion larva, dorsal view mm postflexion larva, dorsal (CaICOFI); postflexion larva, 1972a); head of20.5 view 1981); flexion larva, (Moser 11.3 mm (Ahlstrom mm 24.5 (Ahlstrom 1972a); juvenile, 27.0 mm (CaICOFI 4903, station 92.117). 203

216 Leurog/ossus schmidti BATHYLAGIDAE LIFE HISTORY DESCRIPTION EARLY MERISTICS Mode EGGS Range diam.: 1.65-1.90 Shell mm Yolk: Segmented Vertebrae: I OG: 1-9, coalescing to of No. Diam. ofOG: 0.35-0.40 mm 49 47-52 Total 24 22-29 before fusing Precaudal 25 surface: Pustulate inner surface Shell 25-27 Caudal On tip & OG notochord Pigment: Fins: shell Diagnostic coalescence & features: Pustulate inner surface; o 0 Dorsal spines larger diameter shell globules; oil of migration stilbius. L. in 10 than 10-11 Dorsal rays o 0 Anal spines 12-13 11-14 LARVAE Anal rays 4-5 mm 9-10 length: Hatching 8-10 Pelvic Flexion length: 13-18 mm 8-9 9 Pectoral Transformation length: 31-35 mm Caudal: 10+9 , CI> A, D, C development Fin PI' P sequence: 10+9 Principal 2 2 Pre on tip; notochord Yolk-sac-Around Pigmentation: OG. flexion- Procurrent: 12-13 midgut midgut, extends to trunk when expanded; Blotch above on 10-17 Upper 12 trunk dorsad to hindgut, when expanded. on extends Flexion- 11-15 blotch Lower (large) of2-8 posiflexion--Series hindgut; on of3-6 series trunk; on Gill rakers: 8-9 1-4 base on peduncle; caudal on 1-2 jaw; lower of at tip opercle; on Upper C rays of postfiexion. late in 17-19 Lower 2 Slightly early eyes stalked stage; in features: preflexion Diagnostic 2 Branchiostegals pigment body more lateral than in L. count vertebral & higher total (47-52 gut melanophores blotch-like large stilbius & vs. 39-42); on HISTORY LIFE body. & to the Bering & eggs Okhotsk Seas; Columbia Southern British Range: MORPHOMETRICS (range & mean in %)* south larvae eastern in N Pacific 46° to Y-S PrF F PoF Epi- Habitat: Tr mesopelagic & Juv Sn-AlBL 64-80 67-81 74-81 Summer-winter 71-77 season: Spawning 76 72 78 74 pattern: ELH larvae Oviparous; planktonic eggs & 4-9 7-9 12-17 BDIBL 8-13 10 14 8 7 LITERATURE 7-20 17-23 17-28 HLlBL 28-30 23 16 19 29 Ahlstrom 1969b HWIHL 1984b al. Ahlstrom et Dunn 1983 24-46 11-39 15-32 26-30 SnLIHL 1989 et. al. Matarese 26 24 32 27 15-24x x 23--63 11-33x 26-32x EDlHLt ILLUSTRATIONS (Illustrator) ORIGINAL 26-78 22-32 18-31 24-31 20x26 20x25 30x39 29x28 Preflexion larva, 8.4 mm Arthur) (G. MattsonIN. Flexion larva, 20.5 mm (G. Arthur) MattsonIN. PILlBL P L1BL 2 . Data from Dunn (1983). * is given fust, vertical axis second. slightly oval; horizontal axis t Eye 204

217 schmidti Leuroglossus Northern smoothtongue •.• .• "'."""'~ ~,0ri."""""" '."' 6.3 mm 1.8mm 8.4 mm 20.5 mm mm 31.5 33.1 mm Figure Bathylagidae 8. Egg, 1.8 mm (Ahlstrom 1969); yolk-sac larva, 6.3 mm (Dunn 1983); preflexion larva, 8.4 mm (Northern Holiday, larva, 31.5 mm (Northern Holiday, station 29); postflexion 20.5 mm (Ahlstrom et aI. 1 984b); juvenile 33.1 mm larva, station flexion 47); (Dunn 1983). 205

218 Leuroglossus stilbius BATHYLAGIDAE HISTORY EARLY LIFE DESCRIPTION MERISTICS EGGS Mode Range 1.0-1.2 mm Shell diam.: Segmented Yolk: Vertebrae: 15-25 No.orOG: initially, Diam. OG: 40 inner Pustulate of 39-42 Total coalescing & 1 fmally 2, to 21 surface 20-21 Precaudal surface: Shell 20 18-21 Caudal Pigment: Around notochord tip, on OG Fius: surface of inner Pustulate shell; coalescence & features: Diagnostic o 0 Dorsal spines pigmentation globules; oil of migration 10 9-12 Dorsal rays o 0 Anal spines LARVAE 12 11-15 rays Aual 3 length: ca. Hatching mm 8 8-10 Pelvic mm 3.5-15 length: Flexion 8 8-11 Pectoral mm 24-29 length: Transformation Caudal: 10+9 , C], C sequence: Fin D & A, PI' development P 10+9 Principal 2 2 on Pigmentation: tip; notochord Yolk-sao-Around OG. Procurrent: globule Preflexion-Oil right either to dorsally migrate melanophores 11-16 13 Upper side or left notochord & of trunk; diminishes from by disappears 13 9-15 Lower a series; form to on hindgut, flexion others appear 1 (large) stage; then Gill rakers: pair below specimens in half the base PI (ca. 10% have I on one side 7-9 Upper & none); on lower & upper jaws. Flexion-posiflexion- have 40% 17-20 Lower on opercular 2 patch; to isthmus; over Hindgut series 5-6; increases 2 Branchiostegals girdle. & nape; on pectoral brain Diagnostic eyes stalked Slightly features: in early preflexion stage; LIFE HISTORY sparse pigmentation; distinctive hindgut pigment series; 0-1 2-5 L. ventrolateral melanophores caudal schmidti; on in trunk vs. California of Oregon to Gulf Range: total lower count than in pigment; lacks peduncle vertebral schmidti L. (39-42 vs. 47-52). Epi- Habitat: & mesopelagic %) (range & mean in MORPHOMETRICS January- season: Spawning Winter-spring; highest larval abundance in April surveys in CalCOFl Y-S PrF PoF Tr F Juv ELH pattern: Oviparous; planktonic eggs & larvae Sn-NaL 75-78 72-76 78-82 73-77 67-70 76-76 74 76 79 74 76 69 LITERATURE 7-10 15-15 15-16 11-14 10-11 8-15 BDIBL 10 12 15 16 11 9 1969b, Ahlstrom 1965, 1972a a1. 1984b Ahlstrom et 20-22 23-26 28-30 17-22 14-15 30-31 HLlBL Matarese et 1989 a1. 15 20 22 24 29 31 52-67 52-54 43-50 36-41 38-42 HWIHL 33-60 (Illustrator) ILLUSTRATIONS ORIGINAL 53 40 47 58 49 39 17-26 13-17 26-33 26-31 SnLIHL 25-30 24-27 1.1 Mattson) (G. mm Egg, 22 29 28 29 15 26 mm Yolk-sac larva, 3.9 MattsonIN. Arthur) (G. 18-24x 18-19x 15-21x 21-25x 36-50x 24-29x EDIHL* of Head MattsonIN. Arthur) (G. larva preflexion mm 5.4 27-33 26-39 24-27 22-26 20-23 27-29 larva, Preflexion mm 9.0 MattsonIN. Arthur) (G. 20x30 19x25 17x21 43x31 27x28 23x24 11-11 3-6 2-5 7-10 0-0 5-6 PILlBL vertical axis slightly second. Eye * oval; horizontal axis is given first, 0 6 4 8 11 3 0-0 5-9 P 0-0 L1BL 0-0 0-1 9-9 2 0 0 0.4 0 7 9 206

219 California smoothtongue stilbius Leuroglossus 3.9mm 1.Imm 5.4 mm 8.5 mm mm 5.4 9.0mm 15.7 mm -~ ···i!/o; 23.2 mm 28.5 mm Bathylagidae Figure Egg, 1.1 mm (CaICOFI); yolk-sac larva, 3.9 mm (CalCOFI 5103, station 112.65); preflexion larva, 5.4 mm 9. dorsal larva, preflexion of 5.4 mm preflexion larva, dorsal view (CalCOFI 5101, station 93.40); head of 8.5 mm head (Ahlstrom 1972a); view (Ahlstrom et al. 1984b); preflexion larva, 9.0 mm (CalCOFI 5104, station 70.90); postflexion larvae, 15.7 mm (Ahlstrom 1972a), juvenile, 23.2 (Ahlstrom et al. 1984b); mm 28.5 mm (Ahlstrom 1972a). 207

220 MICROSTOMATIDAE: and Argentines pencilfishes 1 G. 1. AND H. L. BUTLER MOSER genera The family and Microstomatidae includes three Microstomatids are oviparous, with relatively large 17 species and is found from equato- worldwide about inner eggs characterized planktonic by a pustulate the of in occur genera three boreal to rial waters. Two of segmented chorion, a single and yolk, the surface the of species five least at are there region: CalCOFI al. (Ahlstrom large oil et globule Larvae 1984b). are is Nansenia whereas Microstoma a by represented unusual an develop and slender in fold" the foregut that undescribed (Table single Microstomatidae species 1). is rounded has often pigmentation. The head associated are distributions Species by mass delimited water profile; head the species some in anterior a blunt with boundaries. candida California occurs in Nansenia the from is bent downward the longitudinal axis. The eye of to Alaska northern Baja Gulf Current from the is elliptical becoming at the round species, most in end California and N. crassa has a subtropical distribution the first to the larval period. of Pectoral fin rays are off Baja California and overlaps that extends northward at least or is heavy Pigmentation most in form species. in characteristic is the feature A common species. most that California of N. candida off southern (Kawaguchi of series a longitudinal of two or of these melanophores presence abundances Larval 1984). Butler and onto the the in offshore peak blotches coast the near low are species posteriad above the gut, extending and of Nansenia ahZstromi, N. the California Current. core tail into anteriad species, some in the head region. and, peZagica, the N. Zongicauda occur in primarily and Of of larvae these, only the N. et al. Ahlstrom summarized information on (1984b) Pacific. eastern tropical eggs and larvae of candida, N. crassa, N. and Micro- been identified in offshore CalCOFI have peZagica flexion sp. eggs single and with of illustrations stoma adult et samples. Ahlstrom (1984b) listed and al. of series developmental Complete species. these larvae that characters ontogenetic eastern the distinguish of primari- are here presented species three these based of microstoma M from Microstoma of species Pacific 29 N. of lyon detailed 23 candida, N. examination in occur sp. Microstoma of Larvae Atlantic. north the crassa, and 19 Microstoma sp. (Table Microstomatidae CalCOFI from central California to southern samples 2). distributional and meristic adult for sources Primary California peak abundance seaward a more with Baja information Kawaguchi (1964b), and were Cohen N. larvae. crassa N. candida and compared with of (1984), Butler Larvae (1984b). al. et Ahlstrom and Nansenia in samples are rare and in peZagica <20 CalCOFI (usually medium-size are microstomatids Adult species condition not are and in the poor included midwater fishes. They occur slender, cm), silvery, by distinguished descriptions. They are easily a deep primarily and in epi- some but waters mesopelagic in and BL 22% averages depth (body body postflexion) species may be benthopelagic where they encoun- also blotches diagnostic pigmentation. Heavy pigment ter the continental slope. The lateral line well is develop fin adipose and caudal, pelvic, dorsal, the on with distinct scales that extend posteriad developed larvae bases the gular region. In in late postflexion and the of the caudal fins onto sides the on are fin. Pectoral bases and pelvic extend dorsal the on blotches the fin is either fin dorsal at midbody and body the (Nansenia) of rays. the onto regions basal the well posterior (Microstoma). The snout or is short and blunt (Cohen 1964b; and Butler 1984). Kawaguchi National Laboratory, Jolla 1 La Southwest Service, Fisheries Marine Fisheries California. Science Center, La Jolla, 208

221 Table Microstomatidae characters for the microstomatid species in the California Current vicinity. All species have 1. Meristic caudal 10+9 rays. principal Vertebrae Fin rays Total D CV A PrCV P, BrR Species C, PI 49-50 9-11 7-8 15-17 11 32-35 10-12+ 9 4 sp. Microstoma 10-11 8-11 7-8 35-36 12 9-11 3 ahlstromi Nansenia 44-47 9-10 8-9 9-11 27-29 16-19 9-10+ 9-11 3 N. candida 8-9 8-9 9-10 43-46 17 11-13 10-11 29 4 10+10 crassa N. 11-14 47-50 10-11 10-11 9-11 4 longicauda N. 9-11 38-39 9-10 8-9 10-11 4 pe/agica N. the 2. of specimens (above) and size ranges (in mm, below) Table in Number preparation of the Microstomatidae used ilL" indicates literature used in the description. microstomatid species descriptions. An Pre flexion Flexion Postflexion Yolk-sac Transformation Egg Juvenile Species La 3 4 3 3 3 2 sp. Microstoma 7.8-9.0 9.5-12.7 13.8-16.0 18.6-21.2 25.6-28.9 6.7-7.9 La 6 8 6 4 2 3 Nansenia candida 17.2-23.3 5.2-7.0 7.1-9.8 10.5-15.0 24.3-25.9 4.4-5.3 La 2 8 4 6 0 3 N. crassa 18.4-20.4 10.0-14.2 7.5-9.5 4.9-7.1 3.9-4.8 a Ahlstrom et aI. 1984b 209

222 MICROSTOMATIDAE Microstoma sp. MERISTICS EARLY LIFE HISTORY DESCRIPTION Mode Range EGGS diam.: mm 2.1-2.4 Vertebrae: Shell Segmented Yolk: No.ofOG: I Total 50 49-50 mm Diam. OG: 0.49-0.82 of 34 Precaudal 32-35 surface: Shell Pustulate inner surface 16 15-17 Caudal extending ventral surface of trunk Pigment: Along onto gut, above Fins: head on sac yolk & tail region; spines Dorsal o 0 & inner pustulate Diagnostic features: Large shell diameter; oil globule II 9-11 rays Dorsal pattern shell surface; pigment on embryo Anal 0 spines o 7-8 LARVAE 7 Anal rays 9 Hatching mm 5-6 length: Pelvic 9 11 11 Pectoral Flexion length: 9-13 mm 18-25 Transformation mm Caudal: length: 10+9 10+9 Principal C" P" sequence: development Fin C P2' A, & D 2 Procun-cnt: ventral Pigmentation: Yolk-sac-Paired posteriad trunk series extends 10-12 Upper on snout; tail to hypural region & anteriad through branchial region to 10-11 Lower isthmus from midline ventral on concentrated sac; yolk anus. on to Gill rakers: gill fold; gut on flexion-Heavy Pre of at Flexion- arches. angle 9 Upper notochord ray Paired embedded series above C between D & C; on 15 Lower in & on finfolds caudal surface lateral Posiflexion-On region. bases 4 4 Bnmchiostegals C. Transformation-Over entire surface, heaviest D & between to posterior D. HISTORY LIFE with fold appearing in yolk-sac late Diagnostic features: Long gut snout from heavy paired ventral pigment continuous stage; to C; rays form early in on back far P A, D, PI; & embedded Ad; no body; Range: Pacific, south to central Baja eastern Subarctic-transitional 2 between guanine above & notochord pigment layer silvery C; D California count (49-50) than developed at transformation; total vertebral higher candida Nansenia (43-46). crassa N. or (44-47) in Habitat: Mesopelagic ill MORPHOMETRICS (range & mean %) with present year-round in Larvae season: Spawning samples CalCOFI abundance August in peak y-s PrF F PoF Tr luv eggs & planktonic pattern: Oviparous; larvae ELH Sn-AlBL 73-75 74-78 72-79 79-81 81-82 79-81 74 76 76 80 80 81 LITERATURE BDIBL 10-13 8-10 9-10 9-10 10-11 10-11 11 1984b 9 9 et Ahlstrom al. 10 11 II HLIBL 16-18 13-15 17-20 18-19 18-19 17-18 ORIGINAL ILLUSTRATIONS (Illustrator) 14 17 19 18 17 18 HWIHL 54-62 56-63 50-56 39-50 39-42 37-39 Artbur) larva, 4.3 Yolk-sac mm (G. MattsonIN. 57 59 53 46 41 38 larva, (G. mm 8.6 Arthur) Preflexion MattsonIN. SnLIHL 8-\8 17-27 21-25 22-25 19-19 18-22 mm (G. Arthur) MattsonlN. Juvenile, 28.3 12 23 23 24 20 19 33-42x EDIHL* 28-30x 26-32x 25-30x 29-30x 26-26x 33-40 33-40 33-39 29-33 30-32 26-30 oval, slightly from Eye * horizontal vertical; inclined typically axis long 39x36 28x37 29x35 28x30 30x31 26x28 second. axis vertical first, given is axis P,LIBL 0-3 2-6 3-5 6-9 10-10 14-14 3 4 4 8 10 14 P LIBL 0-0 0-0 0-0 2-3 2 4-7 7-12 0 0 0 2 6 9 210

223 Dusky pencilsmelt sp. Microstoma 4.3 mm 2.2mm 8.6mm 12.0 mm 28.3 mm Figure Microstomatidae 1. Egg, 2.2 mm (Ahlstrom et al. 1984b); yolk-sac larva, 4.3 mm (CaICOFI 5104, station 90.120); preflexion larva, mm (Ahlstrom 5104, station 90.120); flexion larva, 12.0 mm (CaICOFI et al. 1984b); juvenile, 28.3 mm (CaICOFI 5101, station 97.40). 8.6 211

224 Nansenia candida MICROSTOMATIDAE LIFE DESCRIPTION HISTORY EARLY MERISTICS EGGS Mode Range mm 1.4-1.6 Segmented diam.: Yolk: Shell Vertebrae: 1 No.ofOG: Diam.ofOG: mm 0.41-0.49 46 44-47 Total Shell surface: Pustulate inner surface 27-29 27-29 Precaudal trunk Between & gut; Pigment: on yolk sac 17-19 16-19 Caudal shell Diagnostic features: Moderately large diameter; pustulate OG & Fins: surface; embryonic inner shell pigment o 0 Dorsal spines 10 9-10 Dorsal rays LARVAE o 0 spines Anal mm Hatching length: 3.5-4.0 8-9 9 Anal rays mm 7.0-10.0 Flexion length: 9-11 Pelvic 17 mm Transformation length: ca. 9-11 Pectoral , , D PI' A, P C C & sequence: development Fin I Caudal: 2 2 PI from gut & trunk between series Yolk-sae-Paired Pigmentation: 10+9 10+9 Principal anterior on gut; of region of base to midgut; above terminal section Procurrent: below yolk sac. Pre flexion-Ventral midline patch liver. 9-10 Upper ventral Flexion-Paired hindgut; above forming midline patch series 8-9 Lower Pos!flexion-Hypural extends hypural region. on to hindgut; posteriad rakers: Gill base; on body surface, A on expanded; blotch heaviest above 8-11 Upper Transjormation-Covering heaviest at body entire myosepta. surface, 18-21 Lower region. hypural 3 3 Branchiostegals features: Diagnostic Moderately developing fold lateral with gut long gut at melanophore paired stage; preflexion fold unpigmented; series LIFE HISTORY trunk & gut; slightly origins hypural P D & pigment; 2 between layer developing guanine PI; posterior early in rays midbody; to form Range: Subarctic-transitional eastern 30° N 55° to from Pacific in late postfiexion stage. Habitat: Mesopelagic MORPHOMETRICS mean (range & in %) with samples CaiCOFl year-round Spawning in present Larvae season: y-S PoF PrF Tr Juv F February abundance peak in 72-80 82-84 74-80 78-83 Sn-AlBL 80-85 68-76 larvae & planktonic Oviparous; pattern: ELH eggs 80 76 75 72 83 82 LITERATURE 13-14 15-17 13-15 13-15 10-14 BDIBL 14-15 14 15 12 14 14 13 al. et 1984b Ahlstrom 24-26 24-27 19-24 7-9 24-26 HUBL 24-26 1989 al. et Matarese 22 25 25 25 25 9 49-57 50-59 42-48 HWIRL 96-118 38-47 41-45 (Illustrator) ILLUSTRATIONS ORIGINAL 54 45 53 43 105 43 21-34 23-27 21-25 19-20 20-23 29-56 SnLIHL Egg, (G. Mattson) rum 1.4 27 44 22 25 19 21 larva, Preflexion 7.0 MattsonIN. (G. rum Arthur) 72-86x 25-29x 23-27x 25-29x 25-29x 27-29x EDIRL* Arthur) Postfiexion larva, 14.0 mm (G. MattsonIN. 32-38 27-33 26-30 29-31 56-76 28-33 MattsonIN. Arthur) 20.0 mm (G. Juvenile, 28x30 28x31 27x28 26x29 27x35 80x67 6-12 9-15 0-0 2-6 15-16 12-15 long oval, slightly Eye vertical; horizontal * from inclined typically axis 11 4 9 14 0 15 axis is given first, vertical axis second. 0-0 0-1 1-7 8-8 9-9 0-0 0 0 0.3 4 9 8 212

225 Bluethroat argentine candida Nansenia 7.0mm 1.4 mm 8.4 mm 14.0 mm 20.0 mm Figure Microstomatidae 2. Egg, 1.4 mm (CaICOFI); preflexion larva, 7.0 mm (CaICOFI 5003, station 40.100); flexion larva, 8.4 mm (Ahlstrom al. 1984b); postflexion larva, et 14.0 mm (CaICOFI 5004, station 50.100); juvenile, 20.0 mm (CaICOFI 5005, station 40.110). 213

226 MICROSTOMATIDAE lVansenia crassa EARLY MERISTICS DESCRIPTION HISTORY LIFE Mode Range EGGS Shell diam.: 1.1-1.3 mm Vertebrae: Segmented Yolk: No.ofOG: I 43-46 Total niam. OG: 0.30-0.35 mm of inner surface Shell snrface: Pustulate 29 Precaudal 17 Paired series between trunk & gut; Pigment: on yolk sac Cauda! Diagnostic features: Large & shell diameter OG Fins: 0 o Dorsa! spines LARVAE 9-10 Dorsal rays Hatching ca. 3 mm o length: 0 spines Anal Flexion 8-9 length: mm 7.5-9.5 rays Anal length: Transformation mm >18 10-11 Pelvic Fin PI sequence: development Pectoral 11-13 C P2, A, D, & C & I 2 trunk Caudal: Yolk-sac-early Pigmentation: between prejlexion-Paired series 10+9 10+9 midgut elongate isthmus & Principal & hindgut; patch on ventral midline from Procllrrent: dorsum on blotches jlexion-6-7 prejlexion--early Late liver. to 10 fold. posterior surface of on anus; Late jlexion-early to gut anterior Upper 10 postjlexion--On blotches above & below region; Lower hypural median rakers: Gill jaw; lower postjlexion--On Late snout. peduncle. caudal 12-14 Upper covered anterior to A origin; heavy on surface Transformation-Body 23-24 P2 anus. & above Ad, D, Lower & bases 4 4 Branclliostegals fold Diagnostic forms Gut on pigmented stage, preflexion in features: posterior blotches along dorsum; heavy embedded heavy surface; LIFE HISTORY below isthmus & midline liver; heavy hypural & pigment ventral early-fonning rays. with PI caudal large blotches; peduncle Current south region Range: N Subtropical; southern of35° of California (range & mean in %) MORPHOMETRICS adults benthopelagic Mesopelagic; in continental slope regions Habitat: PrF Y-S F PoF Tr Juv Larvae present Spawning year-round in CalCOFl samples with season: January-March peak in abundance Sn-A/BL 56-71 74-79 75-83 71-77 80-80 74 65 79 76 80 eggs planktonic Oviparous; larvae ELH & pattem: 9-12 10-11 BDIBL 11-13 14-17 14-15 10 10 13 15 14 LITERATURE 13-17 20-25 HLlBL 24-27 26-29 27-30 22 16 25 28 27 1984b al. Ahlstrom et 53-60 54-68 48-56 HWIHL 42-46 38-43 55 44 52 60 40 ILLUSTRATIONS ORIGINAL (Illustrator) 12-25 16-23 22-26 SnLIHL 21-25 20-21 24 21 20 23 21 3.6 Yolk-sac Arthur) larva, mm (G. MattsonIN. 25-29x 43-50x EDIHL* 19-26x x 23-25 21-25x Preflexion larva, 6.0 mm (G. MattsonIN. Arthur) 27-33 31-40 31-35 24-29 25-27 (G. larva, 14.5 Arthur) MattsonIN. mm Postflexion 24x30 45x33 27x36 22x27 23x26 mm 18.8 Transformation specimen, (G. Arthur) MattsonlN. 0-3 4-6 5-7 6-20 15-16 0.8 5 6 14 16 from vertical; horizontal axis * Eye slightly oval, is long axis inclined 0-0 0-0 0-0 1-8 vertical first, axis given second. 0 0 0 5 10 214

227 Stout argentine crassa JVansenia 3.6mm 1.5 mm 6.0mm mm 8.5 14.5 mm 18.8 mm Figure Microstomatidae3. Egg, 1.5 mm (Ahlstrom et al. 1984b); yolk-sac larva, 3.6 mm (CaICOFI 5103, station 133.40); preflexion larva, (Ahlstromet 14.5 5102, station 137.35); flexion larva, 8.5 mm (CaICOFI al. 1984b); postflexionlarva, mm mm (CaICOFI 5103, station 6.0 120.45); transformation specimen, 18.8 mm (CaICOFI 5106, station 127.40). 215

228 OPISTHOPROCTIDAE: Spookfishes G. MOSER H. Opisthoproctidae is a small family of relatively The 1964b). (Cohen differ Dolichopteryx of Species fin in rare The midwaters. oceanic in worldwide found fishes position and ray length (Table In Opisthoproctidae 1). and about family includes six genera the species; 10 dorsal, near D. binocularis the anal, and pelvic are fins to due unsettled somewhat is of the group taxonomy caudal the fin the and rays fin pectoral beyond extend existence the their rarity and to numerous of the In base. caudal fin anal, the longipes D. dorsal, and Six undescribed species (Cohen 1964b; Badcock 1988). pelvic fins are more anterior and the pelvic fin rays are in occur genera three in species opisthoproctid the Dolichopteryx sp., the than longer pectoral In rays. California Opisthoproctidae (Table Current 1). Larvae is relatively anterior origin fin pelvic than more D. in of longipes, Dolichopteryx species, three Bathylychnops (Table relatively is head the and larger longipes CalCOFI in occur microstoma, Macropinna and exilis, Opisthoproctidae The for delayed tendency general 1). has a D. longipes apparently sampies. ichthyoplankton paedomorphosis metamorphosis and in the group B. while worldwide distribution temperate to tropical culminates in a gelatinous has which Dolichopteryx, in exilis eastern occur microstoma and M Pacific appearance. larvoid sheath and microstoma M of transitional waters. A few specimens been of opisthoproctids have not Planktonic eggs D. Pacific. central tropical the in collected been have usual below reside may and depths. identified sampling Beebe (l933a) reported large ovarian eggs in an B. and 85 brachyrhynchus, primarily known binocularis from the Atlantic, and an undescribed Dolichopteryx Stein and Bond (1985) reported mm D. longipes and species are known from a few juvenile or adult speci- a 464 ovarian eggs (2.2-2.6 mm diameter) in mm ripe mens in the CalCOFI sampling region. collected exilis. Large eggs B. and large size at hatching may be typical opisthoproctids. This of part, in explain, may are special- highly most the among Opisthoproctids absence the of in ichthyoplank- small larvae preflexion stage) ton The smallest known larvae (flexion samples. teleosts. ized dorsally-directed Bathylychnops has round are microstoma M and are that eyes tubular have genera other longipes aU recog- easily D. of eyes and (Dolichopteryx, Macropinna, directed dorsad either prominent nized by their weB-developed tubular eyes, abdominal Rhynchohyalus) Opisthoproctus, rostrad pelvic or (Winteria). flexion gut; long relatively and fins, stage B. the extend that of arrangement alternating a unique have exilis diverticulae AU retinal species have blotches. pigment dorsal and ventral field of vision and, in Bathylychnops, the diverticulum a is secondary eye with separate lens and cornea (Pearcy et 1977; Lockett 1966; Munk 1965; aL Stein descriptions following The exilis, B. longipes, D. of 1985). Bond and morphs the body Two occur in microstoma are based M on detailed examinations and (Dolichopteryx and Bathyly- an form elongate family: Opisthoproctidae Counts of 19 specimens (Table 2). compressed form and a foreshortened, laterally and information were obtained morphometric from chnops) (1960, 1964b), Cohen al. (1984b), Ahlstrom et Stein Macropinna). Opisthoproctidshave smaHmouths, (e.g., Bond and from counts (1988) and Badcock and (1985), numerous modifications of the bones associated head taken measurements and study. this in eye and with jaw specializations, and enlarged pectoral pelvic some species fins that are high on the body in or 216

229 Table 1. characters and fm ray and vertebral counts of Opisthoproctidae species that occur Opisthoproctidae Morphometric principal California region based on the or adult specimens. All species have 10+9 Current caudal fm rays. in juvenile characters Morphometric Fin rays Total Sn-D/SL Sn-A/SL D HUSL Sn-PiSL A P PI 2 vertebrae Species a 82 64 25 Bathylychnops hrachyrhynchus II 13 73 10 8 58 13-16 71-73 70 82-85 16-21 10-13 7-8 10-14 exilis B. 78-84 b 88 82 86 II 25 13 hinocularis 14 Dolichopteryx 7 57 77-79 31-34 69-70 8-11 74-77 10-11 8-10 11-13 D.longipes 41 14-15 sp.c 84 37 9 8 63 80 Dolichopteryx 45 II 11-12 70-76 59-61 61--66 45-46 9-10 10-14 17-19 34-37 microstoma Macropinna SID 106.5 mm a 72-58, b 75-51, SID mm 121.7 c SID 93-246, 73.4 mm Table Opisthoproctidae 2. Number of specimens (above) and size ranges (in mm, below) used in the preparation of the opisthoproctid species descriptions. Flexion Postflexion Transformation Juvenile Species 2 2 0 exilis Bathylychnops 14.2-16.7 25.4-60.3 99.9 2 2 0 2 Dolichopteryx longipes 13.0-13.4 70.7-101.9 20.8-29.7 2 2 4 microstoma Macropinna 12.2-20.2 21.5-23.7 11.8 26.1-29.5 217

230 exilis Bathylychnops OPISTHOPROCTIDAE mSTORY LIFE EARLY DESCRIPTION MERISTICS EGGS Mode Range Yolk: Shell diam.: Vertebrae: No.ofOG: 81 Diam.ofOG: 78-84 Total surface: Shell Precaudal Pigment: Caudal features: Diagnostic Fins: 0 0 Dorsal spines LARVAE 14 13-16 Dorsal rays Hatching length: 0 0 spines Anal Flexion mm length: 14-16 12 10-14 Anal rays Up length: Transformation 120 to mm ca. 7 7-8 Pelvic , A, & P PI> D, C development Fin C sequence: 12 10-13 2 Pectoral I 2 Flexion-posiflexion--On & jaws; Pigmentation: lower internal upper Caudal: 10+9 ventrally snout; in patches elongate filaments on brain; on laterally & 10+9 Principal gill to of lower limbs of paired arches; internally posteroventral eye; Procurrent: in gular 12 at & basibranchial region; at c1eithrum; & base PI internally 12 Upper symphysis; c1eithral at 6-7 paired blotches along gut trunk/gut 12 12 Lower hypural on postanal bar; heavy juncture extending ventrad over gut; Gill rakers: blotches with dorsal median 5 region; ventrolateral alternating Upper blotches; on but PI & P also bases. Transformation--Same as above Lower 2 2 & A bases. on D 2 Branchiostegals Elongate features: eyes round, dorsolateral orientation; Diagnostic gut; ; early protuberance; snout fleshy forming PI & P unique head mSTORY LIFE 2 stomach & ventrolateral blotches; alternating median dorso- pigment; longitudinal dorsal side; right early on forms total high open; septum Range: Central California to Columbia British vertebral count (78-84). Habitat: rnesopelagic & Epi- in MORPHOMETRICS & (range mean %) Spawning season: F Y-S PrF PoF Tr Juv larvae Oviparous; pattern: planktonic ELH Sn-AlBL 79-80 83-83 79 83 85 LITERATURE 7-9 BDIBL 6-8 7 8 6 Ahlstrom et al. 1984b Badcock 1988 21-22 18-21 HUBL 1960, 1964b Cohen 19 22 21 Matarese 1989 al. et 31-37 40-44 HWIHL Stein & Bond 1985 42 34 33 26-34 SnLIHL 37-43 (Illustrator) ORIGINAL ILLUSTRATIONS 30 40 47 EDIHL 17-19 Postflexion larva, 25.4 mm (N. Arthur) 21x22 18 20 4-10 PILlBL 7 5 8 in P transforming * broken specimen. rays 2 P L1BL 0-7 • 2 4 7 218

231 exilis Bathylychnops Javelin spookfish mm 15.6 mm 15.6 mm 25.4 A. M'CRER)", PETER 112.5 mm Figure Opisthoproctidae 1. Flexion larva, 15.6 mm, lateral and ventral views (Ahlstrom et al. 1984b); postflexion larva, 25.4 mm (CFRD AO 71, station 43); juvenile, 112.5 mm (Cohen 1960). 219

232 OPISTHOPROCTIDAE Dolichopteryx longipes MERISTICS LIFE DESCRIPTION HISTORY EARLY Range Mode EGGS Shell diam.: Vertebrae: Yolk: 41 Total No.ofOG: Diam.ofOG: surface: Shell Precaudal Caudal Pigment: Fins: Diagnostic features: 0 0 Dorsal spines 10-11 11 Dorsal rays LARVAE Anal spines 0 0 Hatching length: 8-10 Anal 10 rays 13 ca. length: mm Flexion 8-11 11 mrn Transformation Pelvic length: 30 ca. 11-13 13 Pectoral D , C P & PI & A & Fin sequence: C & development I 2 2 Caudal: gill arches & Pigmentation: mostly at angle; Flexion--On filaments, 10+9 10+9 Principal gut; on of terminal to base P I from trunk & gut between series section Procurrent: Pos(flexion--On on lateral series P jaws; base. lower upper & 2 10-11 10-11 Upper lateral of angle posterior extending series from isthmus; jaw; lower 11-12 11-12 Lower to brain; elongate patch & below vomer; lateral on arches; gill on rakers: Gill series on both surfaces gut; above from ventrad extending blotches of Upper on bases; ofP D Juvenile-adult- C. of at base patch median base; 2 Lower Similar on to hypural on base; PI above sheath; in enclosed gut except 2 2 Branchiostegals region. Dorsally features: Diagnostic early gut; elongate eyes; tubular directed HISTORY LIFE & on pigment bases; pedunculate gill vomer P & PI with forming 2 & juveniles larvoid arches; sheath; gelatinous in enclosed adults forms septum longitudinal dorsal side; left on early open. stomach in Range: Apparently worldwide temperate & tropical waters %) (range in mean & MORPHOMETRICS Habitat: Mesopelagic Y-S PrF Spawning season: PoF F Tr Juv paedomorphic larvae planktonic; Oviparous; pattern: ELH Sn-AlBL 74-75 70-72 77-79 71 75 78 LITERATURE BDIBL 6-8 7-8 12-16 7 8 14 Ahlstrom a1. 1984b et HLlBL 21-25 24-27 31-34 Cohen 1964b 23 25 32 HWIHL 41-43 28-35 30-40 ORIGINAL ILLUSTRATIONS (Illustrator) 42 32 35 SnLIHL 31-36 37-43 42-44 Arthur) (N. mm larva, Flexion 13.4 34 40 43 mrn, lateral & ventral larva, (N. Arthur) Postflexion 27.8 views EDIHL* 11-16 16-16 14-21 16 14 17 tubular of PILlBL eye. * Maximum diameter 9-12 t 8 11 t specimen. in broken rays PI flexion specimen. t in broken rays flexion P 2 P L1BL 20-25 2 t 8 23 220

233 longipes Dolichopteryx Brownsnout spookfish mm 13.4 mm 27.8 ./ -~ - ~,~:--- ::--.------ '. '_., '~ .. - .. , 27.8 mm Figure Opisthoproctidae 2. Flexion larva, 13.4 mm (CaICOFl 7203, station 97.90); postflexion larva, 27.8 mm, lateral and ventral views (CFRD 8701, station 62.0.54.7). 221

234 OPISTHOPROCTIDAE Macropinna microstoma MERISTICS LIFE DESCRIPTION EARLY HISTORY Mode Range EGGS Vertebrae: diam.: Shell Yolk: 34-37 Total No.ofOG: 35 Diam.ofOG: 15-18 16 Precaudal Shell surface: 17-20 19 Caudal Pigment: Fins: Diagnostic features: spines Dorsal 0 0 11-12 12 LARVAE rays Dorsal 0 Anal spines 0 length: Hatching Anal rays 10-14 13 Flexion length: <12 mm 9-10 Pelvic mm 21-24 length: Transformation Pectoral 17-19 Fin development sequence: C PI P C & & A, D & , I 2 2 Caudal: of posteriad from angle extending Flexion-Streak Pigmentation: 10+9 10+9 Principal at liver; jaw; dorsal to lower along cleithra; midventrally internally Procurrent: P in blotch heavy liver; & myomeres over dorsad extending base, 2 10-11 10 Upper over blotches; hypural lower ventrad two blotch; hypural median gut; 10-11 Lower hypaxial myomeres; diagonal on streaks dorsally of on terminal series rakers: Gill myomeres epaxial on Posiflexion-Series gut. of increases section Upper Transformation-Gut enclosed posteriad, at transformation. complete Lower rays. PI & P except on Juvenile-Solid pigmentation in black sheath; 2 3 3 Branchiostegals for snout. occipital features: hump Diagnostic with head Large & dorsally mSTORY LIFE origin anus eyes; tubular directed P midbody; to posterior slightly 2 on series elongate; rays 2 anterior P fins paired midbody; to large, epaxial on later & blotch; hypural myomeres; hypaxial myomeres Pacific Eastern Guadalupe, de Range: to southeast Sea Bering from ofIsla compared (34-37) count total vertebral lower in genera other to Baja California region. Habitat: Epi- & mesopelagic MORPHOMETRICS & (range in %) mean Spawning season: Y-S PrF PoF F Tr Juv Oviparous; pattern: ELH larvae planktonic Sn-A/BL 58-65 60-61 61-66 64 61 60 64 LITERATURE BDIBL 12-25 30-31 34-36 14 Ahlstrom et 1984b al. 20 31 35 et 1989 al. Matarese HUBL 26-37 41-43 45-46 27 42 33 46 ORIGINAL ILLUSTRATIONS (Illustrator) HWIHL 49-55 51-52 53-54 47 53 52 54 (N. mm Arthur) Posttlexion 21.5 larva, SnLIHL 40-52 45-48 42-46 38 46 46 44 EDIHL* 18-21 24-25 28-33 diameter Maximum * eye. tubular of 19 20 24 31 PILlBL 12-19 25-26 9 15 25 24 P L1BL 20-25 47-50 2 16 22 48 47 222

235 Barreleye microstoma Macropinna 12.2 mm 21.5 mm Figure Opisthoproctidae 3. Flexion larva, 12.2 mm, lateral and ventral views (labelled 11.7 mm in Ahlstrom et at. 1984b); postflexion larva, mm (CFRD 9203, station 56.7.80.3). 21.5 223

236 Slickheads ALEPOCEPHALIDAE: A. D. AMBROSE 2.7 mm (Nielsen and Larsen 1968) nigricans are and more with genera 24 about includes Alepocephalidae those mm. >3 are tenebrosus A. of than 90 species (Sazonov and Markle 1990; Nelson smallest The available specimens one and species Ten 1994). related leptochili- closely (T. bifurcata and B. nigricans, ca. chthyid Current California the in occur may species mm) 15 mm; A. tenebrosus and B. burragei, 18 ca. Alepocephalidae vicinity (Table general have the shape and pigmentation 1); however, early young of caudal pelagic ray below reside usually stages juveniles. The dorsal, anal and primary fin 200 m and are by complements are complete ca. collected rarely on mm 30 surveys. plankton CalCOFI SL, although Only those fin are pectoral fin of the pro current caudal and Alepocephalus tenebrosus, Bajacalifornia burragei, than completed greater not until SL. 60 mm the For and Talismania bifurcata occur nigricans, Bathylaco four common species most in our study area, the California regularly in deep collections from the proportions relative large the of ca. head (HL 33-42% Current region. preanal the SL) and length (Sn-AL ca. 59-74% SL) early are stage; however, juvenile the through little change Slickheads (ca. medium-sized to small 10-100 (BD ca. increases depth body the SL). 11-22% Eyes and meso- cm) demersal, (45-5500 bathypelagic m, are horizontally elongate, becoming rounder in early aU oceans. The lepto- of residents m) 1000 usually> juveniles. Black tissue lines the mouth, branchial chilichthyid species, a Leptochilichthys agassizii, is peritoneum cavity, and smallest the in specimens bathypelagic poorly rare, denizen known (2000- examined. top The skull of is translucent the through branchiostegal 3000 m) with a higher ray count than stage. early juvenile the from this area (13 vs. the alepocephalids The 4-10). that 1994) (Nelson fishes are slickheads osmeriform may be confused with Young alepocephalids deep somewhat have compressed, moderately to and early platytroctids bathylagids. juvenile Alepo- fins bodies, elongate on the posterior vertical of half the is shoulder organ that present cephalids lack even and lack fins, pelvic abdominal body, the shoulder 1991) they in yolk-sac platytroctid larvae (Matsui and organs, fin, bladder, premaxillary tusks, adipose gas also present in early which is fin, lack an adipose and scales. head fins, when present, are Pectoral low counts meristic (Table The bathylagids. juvenile on are most pigmented body. Slickheads uniformly the position the and 1) Alepocephalidae anal the fin of closely related to platytroctids, which possess a unique dorsal the to relative origin differenti- to help origin fin shoulder structure, tube-shaped postcleithral the organ ate slickheads young the region. fin in this anal The and (Matsui Rosenblatt 1987). Begle (1992) suggested fin anterior is origin to the dorsal well origin in that the platytroctids and Leptochilichthys be included and Leptoderma lubricum Conocara salmoneum; though alepocephaloids Alepocephalidae. in Even the posterior Bajacalifornia origin in fin dorsal the to argentinoids epibranchial a similar possess and organ and Bathylaco nigricans, Narcetes stomias, burragei, (crumenal organ), early suggest characters life history agassizii; Leptochilichthys dorsal and subequal the to have they not that closely related. are Alepocephalids do undergo direct argentinoids, the larger eggs than in origin fin Asquamiceps Alepocephalus tenebrosus, in platytroctids) the (like to contrast development Asquamiceps velaris, pacificus, Photostylus juvenile form larval to metamorphosis from a distinct Features useful and Talismania biforcata. pycnopterus, the in no specialized they and share ids, argentino species are in separating young of the more common with argentino ids (Ahlstrom et ontogenetic characters prominent chin projection the Bajacalifornia of of HL) the relatively large mouth (>60% burragei, al. 1984b). nigricans, Bathylaco rays fin pectoral produced two the group this about known is Little its including ca. the and SL), Talismania of (by bifurcata 26 mm reproduction. Beebe to found alepocephalids (l933a) total myomere (50-54) count of Alepocephalus large eggs from hatch (3-4 direct have to and mm) tenebrosus. eggs intraovarian Late-stage development. of B. 224

237 The following are based on detailed descriptions literature (Beebe the from obtained were data Meristic examinations mm) 3 postflexion of larvae (18.5-24.2 Nielsen 1976, Markle 1968; Larsen and 1933a; 1980; and Markle 1985; Krefft and Markle 1984; of Quero Alepocephalus tenebrosus; 2 postflexion larvae 1988; 1989; al. et Matarese Shiogaki and Machida specimens (32.0-56.0), (18.3-26.0 mm), 4 transitional 1 scaleless Baja- from unpublished counts and 1990), early juvenile (66.6 mm) of Markle Sazonov and from T. this during made counts and Matsui, by made california burragei; 5 postflexion larvae (14.8-28.8 study. Ecological information was obtained from Parr 1 scale- and mm), (31.6 specimen 1 transitional mm), (1954), Larsen and Nielsen (1966), Perkins and Berry nigricans; (76.0 mm) juvenile less of Bathylaco early Quero and Markle (1983), aL (1968), et Eschmeyer mm), 3 transition and 5 postflexion larvae (14.3-26.6 and Machida and (1985), Krefft and Markle (1984), specimens scaleless 2 and mm), (32.5-48.4 early (1988). Shiogaki juveniles (58.5-68.2 mm) Talismania of bifurcata. Table Alepocephalidae 1. Meristic characters for the alepocephalid and leptochilichthyid species in the California Current vicinity. All 10+9 principal caudal rays. have Fin rays Vertebrae CV A Total D PrCV C GR BrR Taxon P2 PI 2 Alepocephalidae 15-18 10-12 51-55 24-26 15-18 26-29 6-7 14-16+14-15 6-7+17-18 6-7 tenebrosus Alepocephalus 9+9 18-21 21-24 42 17-18 16-17 15-17 6 9+16-17 6 Asquamiceps pacificus 2{}--21 10-12+9 37 16-17 16-17 13-15 5 16-17 5-6+9-12 5-6 Asquamiceps velaris 46--48 14-17 12-14 19-22 26-28 7-8 19-20+14-17 7 7-10+24-29 15-17 Bajacalifornia burragei 12-14+11-14 6-9 6--11 16-17 41--42 17-22 11-12 25 3-5+8-13 9-10 nigricans Bathylaco 51-53 7-12 16-23 25-28 5-7 31-33 20-21 6 salmoneum Conocara 4-5 42-51 48-59 63-74 27-37 7 4-6 15-17 Leptoderma lubricum 7-10 8-12 14-17 17-21 47-50 25-28 21-23 8-9 stomias Narcetes 17-20 44 16-19 12-15 6 18 6-7 26 Photostylus pycnopterus 26-31 6-8 43-50 17-24 17-24 8-13 14-18+12-16 5-8+15-19 7-8 16-18 Talismania bifurcata Leptochilichthyidae 11-15 12+9 8-11 8-11 18-24 47-52 11-13 6-11+16-21 27-30 13 Leptochilichthys agassizii 225

238 ALEPOCEPHALIDAE Alepocephalus tenebrosus MERISTICS HISTORY LIFE EARLY DESCRIPTION Range Mode EGGS Vertebrae: Shell diam.: Yolk: Total 53 51-55 No.ofOG: Diam.ofOG: 24-26 Shell surface: Precaudal 25 Pigment: 29 Caudal 26-29 Fins: Diagnostic features: 0 spines Dorsal 0 15-18 LARVAE Dorsal rays 17 Hatching length: 0 0 spines Anal length: Flexion Anal rays 15-18 17 length: Transformation 6-7 7 Pelvic 10-12 10 Pectoral D PI P A, & , C" sequence: development Fin , C z z Caudal: Pigmentation: tissue Poslflexion--Black lining digestive tract from 10+9 10+9 Principal body chamber & margin branchial at concentrated anus; to & mouth Procurrent: skull of top entire body; entire over spreading translucent. 14-16 Upper 15 15-18; A A origin rays 51-55; vertebrae Diagnostic features: Total 15 14-15 Lower mm. 26 >ca. PI rays not elongate on specimens subequal to origin; D rakers: Gill in mean & %) Upper 7 6-7 MORPHOMETRICS (range 17-18 18 Lower 6-7 6 Branchiostegals Y-S PrF F PoF Tr Juv HISTORY LIFE Sn-AlBL 60-66 63 Range: Bering Sea to at least Isla de Guadalupe, Mexico; also reported BDIBL 13-15 from Chile 14 HlJBL 36-38 Habitat: Demersal, depth m 46-1524 37 HWIHL 40-44 season: Spawning 41 SnLIHL 30-31 meso- pattern: ELH bathypelagic & larvae Oviparous; 30 LITERATURE EDIHL* 21-24x 15-19 22x17 PILlBL 1-2 2 ILLUSTRATIONS (Illustrator) ORIGINAL 5-6 PzLIBL 6 (R. mm 24.2 larva, Postflexion Walker) C. MxLIHL 32-37 34 horizontal axis vertical frrst, given is second. oval; is Eye * 226

239 Alepocephalus tenebrosus California slickhead 24.2 mm Figure Alepocephalidae 1. Postflexion larva, 24.2 mm (SIO 67-101). 227

240 ALEPOCEPHALIDAE Bajacalifornia burragei EARLY MERISTICS LIFE HISTORY DESCRIPTION Range EGGS Mode diam.: Shell Vertebrae: Yolk: 48 No.ofOG: 46-48 Total Diam.ofOG: 26 Shell surface: Precaudal 26-28 22 Pigment: Caudal 19-22 Diagnostic features: Fins: o Dorsal spines 0 15 14-17 LARVAE rays Dorsal o 0 spines Anal Hatching length: 12-14 13 rays Anal Flexion length: 8 7-8 length: Transformation all fins in forming rays mm, 32 ca. By Pelvic 17 15-17 Fin development sequence: & Pectoral C PI , D C A, b 2 Caudal: digestive Pigmentation: Posiflexion--juvenile-Black tissue lining tract 10+9 10+9 Principal from mouth concentrated anus; to chamber & branchial body at entire & of top body; skull over spreading translucent. Procurrent: margins vertebrae Total features: Diagnostic 46-48; origin A 12-14; 19-20 Upper rays A well to projection. chin prominent origin; D posterior 14-17 Lower rakers: Gill %) & 8 (range in 7-10 mean MORPHOMETRICS Upper 28 24-29 Lower 7 7 Branchiostegals Y-S F PrF PoF Tr Juv HISTORY LIFE Sn-NBL 67-72 66-70 70 68 65 0 the of California, Gulf & N), Chile 34 (ca. California Southern Range: BD/BL 13-17 13-15 0 (ca. 21 S) 15 14 16 HLlBL 39-40 34-39 associated also probably the with but Habitat: Mesopelagic, bottom 39 36 33 HWIHL 40-45 40-41 Spawning season: 42 40 40 SnLIHL 27-38 32-38 Oviparous; ELH deeper larvae mesopelagic pattern: or 32 28 34 LITERATURE EDIHL* 23-27x 22-29x 17-20 18-23 25xl9 25x20 28x21 3-4 PILlBL 4-5 4 4 4 ORIGINAL ILLUSTRATIONS (Illustrator) P L1BL 5-9 5-7 2 7 7 6 Walker) C. (R. Postflexion mm larva, 18.3 MxLIHL 41-46 35-44 43 39 46 axis second. is oval; horizontal * Eye is given first, vertical 228

241 Sharp chin slickhead burragei Bajacalifornia I t mm 18.3 2. Postflexion larva, Figure mm (SIO 70-19). AlepocephaJidae 18.3 229

242 ALEPOCEPHALIDAE Bathylaeo nigrieans MERISTICS EARLY LIFE mSTORY DESCRIPTION Mode Range EGGS Vertebrae: diam.: Largest ovarian eggs Shell Yolk: 41-42 Total 41-42 ca. (Nielsen & Larsen 1968) 2.7 mm Precaudal 25 25 No.ofOG: Diam.ofOG: 16-17 16-17 Caudal Shell surface: Fins: Pigment: 0 0 spines Dorsal Diagnostic features: Dorsal rays 17-22 18-19 0 spines Anal 0 LARVAE Anal rays 12 11-12 length: Hatching Pelvic Flexion length: 7 6-9 7 Pectoral 6-11 Transformation fins all in forming rays mm, 36 ca. By length: Caudal: PI , C , & A, P C , D Fin development sequence: I 2 2 10+9 10+9 Principal Posiflexion-juvenile-Black tract digestive lining tissue Pigmentation: Procurrent: mouth from chamber to & concentrated at body branchial anus; 14 Upper 12-14 entire body; tip of skull translucent. spreading over & margins Lower 11-14 11-13 41-42; A rays 11-12; maxilla Total vertebrae Diagnostic features: ca. rakers: Gill extending low >50% ofHL, body; on well small, PI orbit; to posterior Upper 3-5 4 to D origin. posterior well A origin Lower 10 8-13 10 Branchiostegals mean & (range in %) MORPHOMETRICS 9-10 mSTORY LIFE Y-S F PrF PoF Tr Juv Sn-AlBL & S Range: Circumglobal between 40° N 40° 59-74 69 71 72 bathypelagic Meso- Habitat: & BDIBL 11-15 14 15 21 season: Spawning HLlBL 33-37 35 37 33 Oviparous; pattern: ELH or mesopelagic larvae deeper HWIHL 38-47 43 43 33 LITERATURE SnLIHL 19-22 21 21 23 EDIHL* 26-30x 19-23 ORIGINAL (Illustrator) ILLUSTRATIONS 28x20 22x19 23x23 PILlBL 0.5-2 16.3 larva, Posttlexion C. (R. mm Walker) 1 0.8 3 P L1BL 6-9 2 7 7 9 oval; is Eye * second. first, given is axis horizontal vertical MxUHL 59-66 62 58 70 230

243 Bathylaco nigricans Boney throat 16.3 mm Figure Alepocephalidae 3. Postflexion larva, 16.3 mm (CFRD 9107NH, station 64.5.85.6). 231

244 ALEPOCEPHALIDAE Talismania bifurcata MERISTICS EARLY LIFE HISTORY DESCRIPTION Range EGGS Mode Shell diam.: Vertebrae: Yolk: No.ofOG: Total 48 43-50 Diam.ofOG: Precaudal 16-18 Shell surface: 17-18 26-31 30-31 Caudal Pigment: Fins: features: Diagnostic 0 spines 0 Dorsal Dorsal 17-24 21 rays LARVAE Hatching length: 0 spines Anal 0 17-24 21-22 Flexion length: rays Anal 6 6-8 Pelvic ca. By length: Transformation fins all in forming rays mm, 32 8-13 Pectoral 12 , C PI A, P2' development sequence: C , D & Fin I 2 Caudal: digestive tissue tract lining Posiflexion-juvenile-Black Pigmentation: 10+9 10+9 Principal body at concentrated al1us; to & mouth from chamber branchial Procun-ent: margins spreading over entire body; & of top skull translucent. 14-18 17 Upper Total features: Diagnostic vertebrae A 17-24; A 43-50; rays origin 15 12-16 Lower ca. by evident rays PI second & first origin; to elongated D subequal Gill rakers: 8L; 26 ca. by present D & A P2; on spot dark 81. 32 mm mm 5-8 Upper 7 in %) 15-19 18 MORPHOMETRICS (range & mean Lower 7-8 7 Brallciliostegals PrF Y-S F PoF Tr Juv LIFE HISTORY Sn-AlBL 61-66 60-62 63-64 63 61 63 to Peru Canada Columbia, British Range: BDIBL 12-18 18-19 21-22 Habitat: depth m 300-2000 between usually Demersal, 15 19 22 HLIBL 33-42 41-42 39-42 Spawning season: 38 42 41 HWIHL 40-49 38-42 41-42 ELH or pattern: Oviparous; eggs & larvae mesope\agic deeper 45 41 40 8nLIHL 26-30 24-28 27-28 LITERATURE 28 26 28 EDIHL* x 24-26x 23-29 x 22-23 10-24 21-24 19-20 25x19 25x22 23x20 ORIGINAL (Illustrator) ILLUSTRATIONS P,LIBL 2-4 4-26 60-85 3 12 72 C. Walker) Postflexion larva, 14.3 mm (R. P,LIBL 3-7 7-10 5-7 5 6 9 MxLIHL 38-42 35-43 44-48 second. vertical first, given is axis horizontal oval; is Eye * 40 40 46 232

245 Threadfin Talismania bifurcata slickhead 14.3 mm Figure Alepocephalidae 4. Postflexion larva, 14.3 mm (SIO 66-41). 233

246 PLATYTROCTIDAE: Tubeshoulders AMBROSE D. A. 13 includes and 37 species; 5 Platytroctidae genera anus, and anlagen. Pigment fin caudal and anal, dorsal, California Current region (Table occur in species the forming is shoulder the of margins the at eyes, the in Platytroctidae and the yolk, dorsally laterally on the body organ, on Matsui and Rosenblatt 1987). Begle 1; the placed the tubeshoulders in (1992) Alepocephali- shoulder the organ. In later yolk-sac larvae, behind and lines the mouth, branchial cavity, yolk, pigment collected in been dae. The young stages have not to and dorsal in forming are rays peritoneum anus, the and CalCOFI plankton surveys. Holtbyrnia latifrons fins, anal and the complement full of principal caudal range abei Sagamichthys and area study the throughout rays increases head The present. is length (10+9) from from midwater tows in occur commonly rn 300-800 ca. 15-18% the least developed yolk-sac speci- BL in deep. and eubranchus taningi Mirorictus Pellisolus mens to ca. is yolk by the time absorbed, 30% BL the 900 below reside generally and depth rn less are the though even body (10- little changes length common, with ranging eubranchus P. central from elongate, horizontally is eye The mm). becoming 16 17° N) to N) central Mexico (ca. California (ca. 35° more round in juveniles. yolk Once the sac is ab- is agripalla Maulisia S). (12° Chile to taningi M and development sorbed, is direct. The top the skull of collected uncommonly in the off Pacific from central early remains translucent through the stage. juvenile California, California, Baja usually at depths and Chile, below 900 m. alepocephalids, which Young platytroctids resemble similar undergo development, and transitional larval cm) 12-30 Platytroctids are medium-sized (ca. stages juvenile and bathylagids, some of both although meso- and 200-2000 m bathypelagic residents (usually of the posteriorly directed shoulder these taxa lack depth) continents, near waters eutrophic of islands, tubule yolk-sac early in even present is that larval ridges, and ocean in seamounts an oceans. Tube- platytroctids. a Matsui to the (1991) presented key their papilla derive shoulders from a tubular name platytroctids young presence a of The California. off located line and posterior the lateral the to below just gular photophore separates most com- the (G0 two ) shoulder girdle through which blue-green a luminous 2 and Sagamichthys species (Holtbyrnia latifrons mon fluid discharged. Platytroctids are can be moderately abei) area. abei our in species other all from S. study dorsal elongate and somewhat compressed, with and a light has the margin dorsal of anterior the on organ anal fins the rear to the of with small pectoral body, eye (00); H this latifrons. photophore is on lacking abdominal with body, the low fins on and fins, pelvic Maulisia argipalla the between photophore a small has tail. a forked Light organs when present are directed horizontaHy in the young and ventrally bases pelvic eubranchus Pellisolus (IVO). fin the of in adults. sac whereas, maxilla; the bordering nearly nasal its has on Lacking are head scales, radii the scales, an adipose Body fin, and fin spines. pigmentation is uniform on the sac nasal the Mirorictus is taningi at midlength neither snout; has photophores. the of species ventrally. and dorsally Little is known of reproduction characteristics the to In is considered stage last yolk-sac the study this (ca. completely absorbed is sac yolk the until 1960). 13- only pub- (Parr fishes oviparous these of The the mm), 15 of descriptions lished until stage of those are most platytroctids larval persists postflexion and Matsui and (l933a) Beebe Murray (1912), Hjort fin (ca. formed have rays pectoral and mm), 30 the are (1991). They elongate ca. (BD slender fairly and the stage until persists transitional complement full of 20% ca. increasing to yolk-sac in BL 12% BL larvae scale of beginning the until or complete is rays fin formation, (ca. ca. in juveniles). The preanal length is fonowing The 64-70% mm). 50 of are descriptions examinations based on detailed throughout devel- least The development. body length yolk-sac 4 of larvae available oped platytroctid larva (10 SIO 78-160) mm, (12.2-13.5 mm), 6 postflexion larvae (13.0-26.2 mm), flexion an is specimen with a large yolk sac early 3 transitional specimens (29.0-39.4 mm), and 3 juve- to extending the from the cleithrum to about halfway niles (49.1-65.6 mm) 5 of Holtbyrnia latifrons, and 234

247 yolk-sac larvae mm), 6 postflexion larvae (14.8-16.0 study our in species platytroctid common less the of area 3 (30.2- specimens transitional (16.0-27.0 mm), Meristic are presented in Table Platytroctidae 2. from primarily obtained data were ecological and 38.0 juveniles (54.1-81.0 mm) of 3 and mm), Sagam- and Rosenblatt (1987) and Matsui (1991). Matsui morphometry comparative The abei. young ichthys of have 1. charactersfor the platytroctid Platytroctidae in the California Current vicinity. All species Meristic 10+9 Table species caudal rays and 12-15+ 11-13 procurrent caudal rays. principal Fin rays Vertebrae Total Dorsal Anal P Precaudal Caudal PI 2 BrR Gill rakers Species 17-20 14-16 16-20 26-29 20-21 8-9 46-50 7-9+18-21 8-9 Holtbyrnia latifrons 15-17 17-20 46-47 7-8 7-8+16-18 19-22 18-19 8-9 25-27 agripalla Maulisia 14-17 16-20 19-22 43-46 15-20 4-6+12-15 7-9 6-7 22-26 Mirorictus taningi 18-21 15-16 20--22 42-44 20--23 6-8 5-6+17-18 17-19 6-8 Pellisolus eubranchus 14-16 14-18 19-21 50--52 16-18 30--31 7-8+16-18 9-10 8 abei Sagamichthys less Platytroctidae Table morphometry of young of the 2. common platytrocids in the California Current region. Comparative Proportions are in percent with the range above the mean. The number of listed in parentheses. examined specimens are EDIHL EDIHL stage Developmental MaxLIHL HUBL (vertical) (horizontal) SnLIHL BDIBL Sn-A/BL HWIHL (mm) SL Species 40--50 19-19 25-27 43-46 24-27 35-36 15-15 postflexion (2) 67-69 20.0-28.0 Maulisia 26 26 15 36 44 45 19 68 24.0 agripalla 20 24 43 26 17 52 transitional (I) 36 66 38.0 19 27 27 42 58 juvenile (I) 23 43 64 51.2 23"':28 27-31 19-22 40-49 48-50 (3) postflexion 16-17 33-39 64-65 20.0-28.5 Mirorictus 44 21 29 25 36 49 17 64 24.8 taningi 22-22 28-28 45-48 21-21 35-38 33-35 transitional (2) 62-64 18-19 40.0-43.0 28 37 21 34 22 46 19 41.5 63 21 31 24 51 (1) 21 36 40 juvenile 63 52.0 42-48 24-25 26-32 21-26 (2) 48-51 transitional 37-39 16-20 67-69 29.2-46.0 Pelliso/us 45 23 24 29 50 18 38 68 37.6 eubranchus 26-27 41-43 19-19 32-35 45-48 21-23 30--33 (2) juvenile 61-63 58.4-71.0 42 26 22 31 46 19 34 62 64.7 235

248 PLATYTROCTIDAE Holtbyrnia latifrons HISTORY EARLY DESCRIPTION LIFE MERISTICS Mode EGGS Range Shell diam.: Vertebrae: Yolk: 46-50 No.ofOG: Total Diam.ofOG: 26-29 surface: Shell Precaudal 20--21 Pigment: Caudal features: Diagnostic Fins: 0 o Dorsal spines LARVAE 17-20 rays Dorsal

249 Streaklight tubeshoulder latifrons Holtbyrnia 15 mm 14mm YOLK IVa 14 mm Figure Platytroctidae 1. Yolk-sac larvae, 15 mm, 14 mm; postflexion larva, 14 mm (Matsui 1991). G0 = = posterior gular organ; IVO 2 intraventral organ. 237

250 abei Sagamichthys PLATYTROCTIDAE LIFE EARLY mSTORY DESCRIPTION MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: Diam.ofOG: No.ofOG: 51 50-52 Total Shell surface: 30 30-31 Precandal Pigment: 21 19-21 Caudal features: Diagnostic Fins: o 0 spines Dorsal LARVAE 17 16-18 rays Dorsal 12 mm Hatching length: D & A, P development PI C sequence: , , IS 14-18 z Pectoral z Pigmentation: Yolk-sac-On shoulder organ, subopercular eye, Caudal: on intraventral photophore, gular dorsally photophore, yolk sac, organ, 10+9 10+9 Principal on internally increasing chamber; gill & mouth in & & peritoneum Procurrent: & lower gut Postflexion--On nostrils along line jaw; dashed lateral 12-15 Upper D midline between & A, & D between internally laterally; A, spreads 11-13 Lower & later posteriorly. Transition-juvenile-Patch anteriorly spreading Gill rakers: on body whitish blue. hypurals & base; A 7 7-8 Upper Diagnostic features: 16 to Yolk-sac melanophore juvenile-Photophore or 16-18 Lower (GOJ; & silvery photophore region gular in photophore of in shape 8 8 Branchiostegals reflector on anterior dorsal margin (00) & of eye on subopercle opercle present; (IVO) photophore intraventral (SBO); opening mSTORY LIFE whitish coloration body mid-eye; about to dorsally extending blue. in Range: Canada Columbia, British from coastally Pacific eastern Japan; MORPHOMETRICS %) in mean & (range to N) 52° N) to Baja California (ca. 27° & Peru (ca. 16° S) Chile (ca. (ca. S) 39° F Y-S PrF Juv Tr PoF Habitat: m Mesopelagic, 300-900 depth 66-69 65-68 66-70 67-71 Sn-AlBL 68 67 68 69 season: Spawning Year-round 13-16 BDIBL 18-19 13-18 15-17 Oviparous; pattern: ELH mesopelagic larvae large 15 18 16 14 27-34 24-19 HUBL 34-37 33-37 LITERATURE 27 32 36 35 46-52 38-48 HWIHL 42-43 39-41 Matsui 1991 44 41 42 49 20-25 21-24 SnL/HL 11-23 17-21 22 23 22 19 horizontal second. (horizontally); elongate Eye * given vertical axis first, 26-34x 26-31 x EDIHL* 26-31x 25-26x 21-26 12-25 22-25 24-25 28x24 28x24 30x24 26x25 2-5 1-4 4-4 4-6 PILlBL 2 4 3 5 1-3 6-9 2-7 7-9 PzLIBL 2 5 7 8 34-46 36-43 44-50 48-54 MaxLIHL 51 39 41 47 238

251 Shining tubeshoulder abei Sagamichthys SBO I YOLK 16mm 14 mm 16 mm Figure Platytroctidae 2. Yolk-sac larvae, 16 mm, 14 mm; postflexion larva, 16 mm (Matsui 1991). 00 = orbital organ; G0 posterior = 2 organ; gular SBO = suborpercular organ; IVO = intraventral organ. 239

252 OSMERIDAE: Smelts MOSER G. H. and temper- Osmerid in shallow subarctic smelts occur (O'Connell the to contrast in 1981), directions opposite waters ate Northern Hemisphere, the of mostly in near-horizontal most other teleosts, alignment in anadromous habitats, although some species marine are osmerids. The clupeoid a much has including hindgut fresh landlocked or in water; a total 7 species in of 13 conspicuous has and foregut the than diameter larger genera are recognized (Eschmeyer Nelson al. 1983; et has hindgut transverse folds, while in osmerids the Six species in four reported 1994). genera are from foregut and the than diameter larger a slightly only three 1972); Lea and (Miller California however, of transverse clupeoid In inconspicuous larvae, the folds. ventral melanophore series is limited to the midline transpacificus, these Thaleichthys paci- (Hypomesus Spirinchus ficus, thaleichthys) elongate of are it is often and dashes, anadromous hindgut and consists region consists irregularly series osmerids, unlikely that their early larvae would occur in plankton In paired. the of samples from the often are located in the melanophores large that CalCOFI single sampling region. The below the foregut and hindgut. finfold pretiosus, Hypomesus Spirinchus species, marine starksi, and range elongatus, as far south as Allosmerus only been have There 13 of osmerid occurrences southern south spawn not do but California central of larvae in 1981 from surveys ichthyoplankton CalCOFI California. were all 1984; to between stations at nearshore one but Bay, larvae The California. Monterey and Reyes Pt. silvery cm), «40 fishes are Adults medium-size species. mm) appear to represent a single (5.2-15.2 They have school that usually in shallow a short water. of the specimen largest The total myomere count (71) and fin, adipose an midbody, at located fin dorsal indicates the that series pretiosus, is Hypomesus since pelvic fins. abdominal highly are species Some regard- are lower in the vertebral other counts species two ed recreational commercial provide and and fishes food distance might early larvae occur at this from whose fisheries during spawning runs. shore (Table Osmeridae 1). melanophore However, the Hypomesus pretiosus, the surf smelt, the in spawns differs somewhat from that pattern pretiosus H. of Eggs the day. intertidal attached upper zone during are by His Hearne larvae have larvae (1983). described to sand grains and the species coarse to select appears and preanal 13-21 4-8 postanal ventral midline mel- spawning on the basis beaches of sand grain size while anophores have 1-4, and 8-11 larvae CalCOFI (Yapchiongo 1983). Hearne 1949; starksi, Spirinchus of the CalCOFI one-third about Also, respectively. nocturnally and surf, the in spawns smelt, night the or two postanal one dorsal midline specimens have composed selects beaches apparently of coarse sand that while Hearne (1983, 1984) stated melanophores, (Fitch and Lavenberg 1971; Hearne 1983). Allosmerus the lack melanophores a diagnostic feature of these is elongatus, be an to believed is whitebait the smelt larvae. it is Without osmerid a transformation of series spawner. ocean eggs Osmerid in mm 0.8-1.1 are possible are differences these whether determine not to adhesive, double a characteristic have and diameter, to incorrect identifica- due to individual variation or chorion numerous oil globules (Hearne 1983, and tion. 1984; al. 1989). Matarese et following The description series larval a partial of larvae are slender, have an elongate gut, a Osmerid through notochord flexion stages), tentatively (yolk-sac melanophores of depressed a single and head, series identified as pretiosus, is based detailed on Hypomesus 1983, (Hearne gut the of along ventral midline the examination 5.2 mm; (1 larvae 12 of 8 yolk-sac, 1984). those resemble superficially larvae Their of mm) 13.0-15.2 preflexion, 3 flexion, mm; 6.8-11.7 of clupeoids but are easily distinguished on the basis the from CalCOFI obtained were data Meristic surveys. musculature, and their ventral morphology, gut body literature the from 1983, Hearne 1972, Lea and (Miller pigmentation. The myotomes of clupeoid have larvae Matarese this 1989) and et al. from counts made during a characteristic appearance caused by cross-hatched study. in obliquely aligned are fibers of that layers two 240

253 Table Osmeridae Distributional ranges and meristic characters of the marine osmerid species that may be encountered in 1. principal samples Central California. All species have 10+9 off caudal fin rays. CalCOFI Vertebrae Fin rays A D PrCv CV Total Range P Species 2 PI 14--17 9--11 65--67 23-27 40-44 12-14 Calif. Brit. Col.-S. elongatus Allosmerus 8 8-12 62-70 42-44 22-24 12-17 Bering Sea-So Calif. 14--17 8 pretiosus Hypomesus 10--11 15-21 33-36 25-29 60--65 8--11 Alaska-S. Calif. SE 8 starksi Spirinchus 241

254 OSMERIDAE Hypomesus pretiosus LIFE MERISTICS EARLY HISTORY DESCRIPTION Range Mode EGGS Shell 1.1 diam.: mm Vertebrae: Yolk: Segmented 66 OG: of Numerous No. 62-70 Total Diam.ofOG: surface: Smooth Shell 43 42-44 Precandal 23 22-24 on melanophores Numerous Pigment: region of yolk Caudal ventral Diagnostic multiple chorion; double disc; oil features: Attachment Fins: globules 0 Dorsal spines 0 10 8-12 Dorsal rays LARVAE 0 spines Anal 0 Hatching length: mm 3-4 14 12-17 rays Anal length: 8 mrn 13-15 Flexion 8-9 Pelvic 14-17 Transformation length: ca. 40 14 mrn Pectoral & P, C" & A Caudal: sequence: development Fin P D, 2 10+9 10+9 Principal posterior surface of yolk sac; 9 on Pigmentation: Yolk-sac--Cluster on Procurrent: some gut, below ventral midline ventral postanal 4 on finfold; entering 12 12-13 base. PI Upper tip; lateral pair midline; below I below notochord 10-11 11 Lower Preflexion-flexion-8 to lion on 2-4 midline; ventral preanal notochord of one-third ca. tip; rakers: Gill 1-4 below midline; postanal ventral 10-13 Upper have pairs between larvae gut 1-2 on postanal dorsal midline; lateral stage; 20-25 tlexion of end 15-16 to increasing musculature trunk & at Lower 7-8 terminal on pair lateral section gut. of Branchiostegals features: High total myomere count Diagnostic number (65-68); & HISTORY LIFE in arrangement of preanal ventral midline melanophores (lacking in present larvae); absence of diagonal muscle layers that are clupeoid clupeoids. Prince Range: to Alaska, Long California Beach, William Sound, %) MORPHOMETRICS (range & mean in Habitat: Marine, sometimes nearshore; entering freshwater larvae CaiCOFI in January-October; season: Spawning samples occur PrF Y-S PoF F Tr Juv January from to May Sn-A/BL 69-81 78-81 75 80 77 demersal ELH pattern: Oviparous with eggs attached to coarse sand grains are intertidal zone; larvae planktonic in upper 5-{i BDIBL 6-6 6 8 6 LITERATURE HLlBL 12-15 13-13 15 13 13 Hearne 1983, 1984 53-63 HWIHL 55-59 et a1. 1989 Matarese 63 60 57 & Okiyama 1988 Saruwatari SnLIHL 14-23 20-22 1949 Yapchiongo 20 18 21 17-23x EDIHL* 18-20x ILLUSTRATIONS (Illustrator) ORIGINAL 24-29 23-34 20x27 28x30 19x27 5.2 mm larva, Yolk-sac Arthur) (N. larvae, Pretlexion 8.3 11.2 mm (N. Arthur) mm, P,LlBL 2-4 2-3 (N. 15.2 mm Arthur) Flexion larva, 2 3 2 L1BL 0-0 P 0-0 2 0 0 0 * Eye axis is slightly oval; horizontal second. is given first, vertical axis 242

255 pretiosus Hypomesus Surf smelt mm 5.2 8.3 mm 11.2 mm 15.2 mm Figure Osmeridae l. Yolk-sac larva, 5.2 mm (CaICOFI 6304, station 60.52); preflexion larvae, 8.3 mm, 11.2 mm; flexion larva, 15.2 mm (CaICOFI 7203, station 60.52). 243

256 STOMIIFORMES MOSER WATSON G. H. AND W. medium-size Stomiiforms are small that fishes to be stable classification has yet to the order. A within all of oceans. zones bathypelagic the epi- inhabit to Weitzman and Fink 1974; 1967a, achieved (Weitzman diverse are They (ca. 50 >300 species), abun- genera, Fink 1984, 1985; Nelson et al. Ahlstrom 1982; 1984c; of oceanic the and constitute part major a dant, (1974) 1994). reduced the number Weitzman gono- of ichthyofauna. Species of the genera Cylothone and by the to genera stomatid in some genera placing six Vinciguerria may be the most numerous vertebrates. the and Stemoptychidae family, a separate in some phylogenetic and morphological their of Because Fink monophyly based 1985) (1984, Phosichthyidae. stomioids of including characters of pres- the numerous on they one are importance ecological and richness most extensively studied ence of of lack of gill rakers in infraorbital a single Mono- groups. teleost all bone, of unique the associated with barbel a mental photophore on based is order the of phyly presence adults, mandibulae of of the adductor oftooth type hyoid specialization attachment, a portion morphology, apparatus, inserting muscle on the postorbital photophore and a the of location a posterior muscle, mandibulae adductor liga- the mirabila on the swim bladder, a unique rete geniohyoideus divided not could he Since muscle. ment and premaxilla, the and ethmoid the connecting for monophyly compo- traditional demonstrate the six nent a single families combined them into he family, several osteological specializations of the branchial (1990) retained Eschmeyer 1985). (Fink the Stomiidae Weitzman 1982). and (Fink region traditional the arrangement and families stomioid six of we classification follow in this guide. his stomiiforms were thought to consist Traditionally, lightfishes of the (Gonostomatidae), the hatchetfishes of diagnostic (Stemoptychidae), fishes, the stomioid and a valuable are arrangements Photophore a group viperfishes, (dragonfishes, predators midwater larger of The identification. stomiiform in tool nomenclature traditional the differs photophores between somewhat segregated snaggletooths, and loose jaws) into six stomioid groups. These and are lightfish, hatchetfish, families. Subsequent phylogenetic altered have studies in 1, and defined Table shown Stomiiformes Figure in have stomiiform of relationships and views traditional 1. Stomiiformes at identified systematic problems levels all taxonomic included: Gonostomatidae Families Stemoptychidae Phosichthyidae Chauliodontidae Stomiidae Astronesthidae Melanostomiidae Malacosteidae Idiacanthidae 244

257 et al. Table of symbols used for designating photophore groups in stomiiform fishes (Ahlstrom Definitions Stomiiformes 1. 1984c). Deep bodied stemoptychids stomiiforms Other Definition Code Code Defmition SO to §ubQPercie photophore which is equivalent SO tip at located (Qrgans) photophores §ymphyseal of posteriormost in series photophore opercular lower jaw. of gonostomatids. Photophore located (Qre) to Qrbit. anterior Orb PO located the Photophores associated with anterior eye and posterior orbit. of PTO Op Qrbit, I!osterior located Photophore !O may be series, QI!crcle of Photophores coded three generally follows: 1/(1+1). as series opercular of upper to equivalent photophore gonostomatids. of (BRP) Br PRO photophore. f!:eQjJercular Photophores l2!:anchiostegal the on located membranes. definition. gonostomatid as Same Is Br i§.thmus. the Photophores located on (I) definition. Same as gonostomatid the from the ventral IP IPhotophores Is of series found the of fin. Qectoral base to isthmus the abdominally Photophores of ventral series located PV the series ventral the of from found Photophores AB base pelvic fin pectoral and fin between and base pelvic the to fin base fin Qectoral base. (yentral) equivalent to PV in gonostomatids, plus a few posterior photophores the of lIP series. VAV ~a1 yentral the of the and fin from to (Qre) anterior found found Photophores series Photophores PAN (yentral) the to base fm may be equivalent to ,!!nai pelvic V A V or VA in fin base. gonostomatids. found ~a1 above found Photophores Photophores of the ventral series AC from the fin. AN ,!!I1a1 fin base. base to £audal fin IC series Photophores lower ~ub) £audal peduncle. from on ventral the of photophores of found Summary SC Together with be equivalent to AC may group AN the isthmus to (IP+PV+VAV+AC). £audal fin base in gonostomatids. Photophores ~upra) to the abdominal above located IV Summary of from series ventral the of photophores SAB isthmus pelvic (yentral) fin (IP+PV). base series and may to to equivalent be VALin gonostomatids (SAB lacking in Sternoptyx). Photophores QPercie from series lateral the OV the of to fin I!ectoral the ~upra) above located Photophores SP (yentral) equivalent to OV in gonostomatids. base. fin be pelvic and may (found pelvic the from PAN VAL (VALA) above laterally located Photophores Photophores of the series lateral L Polyipnus). in (yentral) fin base only the ,!!llal fin base. to Summary to OA opercle the from ores of lateral photo ph anal fin base (OV+VA). (OAA,OAB) lateral to Entire ventral series on body just dorsal OAC (OC) just or series and extending from QPercular border, fin fin it, over ,!!I1a1 to to £audal medial base. ODM Photophores (Qrgans) found .Qorsal to the lateral gracile). midline (found only in Gonostoma 245

258 ~~+=====~O~A : ________ ORB , ,------0 ' ~------,' -:~--~'------------"~ -.-, VAV IV SO '- BR AC --, SUBORBITAL ORGAN ~ORB FIN ADIPOSE DORSAL POSTORBITAL ~ ==- o I GAN I ~-------OA----------~ AC --BULB VENTRAL ADIPOSE FIN STEM BARBEL Figure Stomiiformes 1. Generalized photophore patterns and their terminologies for Iightfishes (top; Fahay 1983), hatchetfishes (middle; The 964a). and stomioids (bottom; Morrow I 1980), photophore groups are defined in Table modified Badcock from and Baird 1. Stomiiformes 246

259 GONOSTOMATIDAE: Bristlemouths WATSON W. trum, swimbladder, Diplophos and in except gut, and in the species gonostomatid occur Eleven CalCOFI prominent display which Manducus, melano- dorsal Gonostomatidae four only but (Table area (Cyclo- 1), series or part during phore all of larval development taenia, Diplophos signata, C. thone acclinidens, Ahlstrom (e.g., 1984c; al. et Ozawa and Oda 1986b; Gonostoma atlanticum) with in regularity any occur 1991). al. et Smith Photophores during develop trans- others CalCOFI ichthyoplankton samples. Four These formation. originate as photophores," "white (Cyclothone pseudopallida, proximus, Diplophos most with ventral or the photophores within all each of Gonostoma ebelingi, G. principally occur elongatum) simultaneously. forming series In BR the Cyclothone offshore their southern, at distribu- stations; CalCOFI Gonostoma in while first, form V VA and and series area. the CalCOFI tions apparently are largely outside first Gono- (Table is photophore OP the Diplophos Larval C. Cyclothone alba, C. atraria and have pallida 3 2). stomatidae been recognized in CalCOFI ichthyoplankton not small samples, juveniles although the last of two Although a good deal of literature exists concerning occasionally are taken. larval stages of the gonostomatids, especiaHy Cyclo- thane and Diplophos (e.g., Gor- 1964; Mukhacheva cm), :::;;20 (usually Adult gonostomatids are small 1984c; 1982; bunova Ahlstrom et al. Oda and Ozawa fishes bioluminescent elongate, moderately compressed, not Cyclo- are identities known. well larval 1986a,b), bathypelagic that occur in the epi-, zones. meso-, and taxonomic They closely resemble phosichthyids; the thone have been confused in the species, especially, status literature. following species accounts are based on these two The been entirely yet not has families of literature the on detailed (Cyclothone atraria) and resolved. C. C. 10 acclinidens, of specimens 40 of examinations are oviparous (some are protandrous Gonostomatids 24 atraria, 36 C. pseudo pallid signata, C. 36 a, hennaphrodites, Nemoto Miya 1983; and Fisher e.g., proximus, Diplophos Gonostoma 37 taenia, D. 35 planktonic 1985), with eggs and larvae; eggs have been 24 elongatum G. 30 and ebelingi, G. atlanticum, described for (l986a) Oda and Ozawa 3). Gonostomatidae (Table Gonostoma denudatum (Sanzo 1931a), planktonic and larvae are six known for of the seven which described and illustrated as identified they larvae 1984c; et Ahlstrom (e.g., genera al. Oda Ozawa and and C. pallida; C. descriptions are not alba those eyes, unpigmented with hatches Diplophos 1986b). an data were from obtained reproduced here. Meristic yolk mouth; a functional without elongate and sac, Kobayashi (1984a), et Schaefer aL (1973), Fujii gonostomatid other the presumably a hatch species at et Ozawa and (1990), counts (1994), Miya al. (1986), similar stage of development. Larvae of aU species are given counts Myomere study. under this during made to a moderate preanal elongate and slender, with long features" "Diagnostic in the following refer descriptions (ca. length on depending species). 40-70% The BL, on made counts this in examined specimens to the trails slightly, gut only all. or more at not commonly, and the reflect not study may for range fun each slightly strongly oval, depending on Eyes are to a species. Ecological information was obtained from species, become round, but or nearly so, transfor- by variety of literature sources including Grey (1964), Larval pigment is absent or mation in most species. DeWitt (1972), Kobayashi (1973), Clarke (1974), Miya ven- light; when present it occurs the on principally and Nemoto (1985, 1991), and (1990). Ozawa et aL 247

260 Table Gonostomatidae Selected meristic characters for the gonostomatid species that occur in the CalCOFI region. 1. All species principal caudal fin rays. have 10+9 rays Fin Vertebrae Total P A CV PreV D C z PI z Species GR 13-14 17-19 30-32 13-15 18-20 8-10 6-7 6-7+5-6 acclinidens Cyclothone 6-9+12-17 18-19 31-32 12-15 17-20 C. 9-10 6-7 6+5-7 alba 13 4+10-11 9-11 17-20 31-33 12-15 17-20 12-14 6-7 5-7+5-8 C. atraria 7-10+13-18 9-11 16-19 17-21 31-34 12-15 13-15 6-7+6-7 6-7 7-10+13-18 pallida C. 12-15 17-21 9-10 6-7 6-7+6-7 17-21 29-34 12-14 4-7+11-14 pseudopallida C. 30-32 13-14 18-20 17-19 8-10 6 6-8+5-7 13 3-5+9-11 signata C. 3+9 9-10 85-93 9-11 54-62 48-55 3-5+3-5 36-39 proximus 8 Diplophos 10-11 89-99 37-41 59-72 8-10 3-6+3-4 52-59 3+7-9 7-8 D. taenia 16-18 26-31 7-8+6-7 38-39 9-11 6-8 20 6-7+11-12 18 Gonostoma atlanticum 12-14 26-29 9-13 8 10-12+7-10 19-20 23-27 8+11-12 43-46 ebelingi G. 10-13+8-10 39-41 11-15 27-32 10-13 24-26 7-9+11-12 7-8 15-16 elongatum G. of 2. of photophore group development (first Gonostomatidae Order photophores in each group) in some Table appearance Photophore groups are shown in Figure Stomiiformes 1 and defined in Table Stomiiformes 1; "&" indicates gonostomatids. development or specimens to determine sequence. insufficient simultaneous Photophore group development sequence Species , OP OA & VAV & AC, ORB & IV, OP & BR acclinidens Cyclothone 3 I IV, & OP & OA AC, & OP & ORB BR & VAV signata C. 3 z & ORB OP SO & AC, BR & IV, OA, & OP & & VA V, OP others proximus Diplophos z I 3 , & OP SO & OP & VAV IV, BR ORB & AC, OP OA, , Gonostoma atlanticum z I 3 ORB, OP OA, & AC IV, BR, VAV & , OP ebelingi G. I 3 OP SO OA & BR, YAY, AC, ORB & OPI' IV, , G. elongatum 3 248

261 mm, Number specimens (above) and size ranges (in of below) used in the preparation of the Table Gonostomatidae 3. species descriptions. gonostomatid An available literature in the description. No indicates were used for any of the ilL" eggs species. Yolk-sac Preflexion Flexion Postflexion Transformation Juvenile Species 10 6 10 9 0 5 acclinidens eyclathane 4.3-5.8 5.7-l3.7 23.4-32.8 5.0-5.9 12.8-21.7 La 4 0 0 6 0 C. atraria 11.6-14.2 l3.6-28.0 11 5 5 3 0 0 pseudapallida C. 6.0-16.7 16.4-20.2 22.2-36.8 5.3-5.7 4 11 5 10 6 0 signata C. 12.0-14.0 4.8-12.2 12.8-17.8 2.4-5.0 4.7-6.1 9 2 5 5 10 5 praximus Diplaphas 6.5-18.6 18.4-21.6 25.3-42.9 45.1-56.6 37.6-39.1 3.3-5.7 7 4 10 6 3 5 D. taenia 30.6-41.2 5.9-19.4 19.2-27.3 28.4-43.0 44.6-61.8 3.4-6.3 12 5 10 5 5 0 Ganastama atlanticum 21.9-30.1 4.7-14.6 17.6-20.7 4.5-6.0 3.5-5.4 10 2 4 7 0 G. ebelingi 16.1-32.7 15.4-16.9 5.2-16.0 3.4-4.6 7.0 10 5 5 5 5 0 G. elangatum 6.1-11.0 11.9-18.9 23.2-31.1 4.7-6.4 3.9-5.3 Ozawa and Oda 1986a a 249

262 GONOSTOMATIDAE Cyc/othone acclinidens MERISTICS DESCRIPTION mSTORY LIFE EARLY Range Mode EGGS Vertebrae: Shell diam.: Yolk: Total 31 30-32 No.ofOG: Diam.ofOG: 13-14 14 Precaudal Shell snrface: 17 17-19 Caudal Pigment: Fins: features: Diagnostic 0 spines 0 Dorsal 13-15 rays 14-15 Dorsal LARVAE Anal 0 0 spines length: Hatching rays 18-20 19 Anal ca. length: Flexion 5-6 mm 6-7 Pelvic 6 length: 22 to mm 13-14 ca. ca. Transformation mm 8-10 Pectoral 9-10 , C" sequence: development Fin , P, C P & A, D z z Caudal: Pigmentation: gut; of sides along Preflexion-posiflexion-Pairs over 10+9 10+9 Principal bladder; gas (usually gut ca. 4-9 9-13 after on over myosepta 4 mm; Procnrrent 10-11) (usually 1-3 myomere postanal from tail of margin ventral on 6-7 6-7 Upper gradually internal; row double 2), becoming on A base; 1 under 5-6 6 Lower of 0-1 at center hypural margin; 0-1 parhypural; tip/along notochord Gill rakers: over at myomere over beginning series tip; notochord notochord Upper 6-9 7 extending to myomere 12-15 6.5-7 ca. 10 mm. mm, 28-30 by at 12-17 12-14 Lower on & neural Transformation-Between on spines; hypurals; haemal 13-15 14 Branchiostegals membranes; ofbranchiostegal margins on hindbrain; bases; A & D on beginning midlaterally external 15 ca. at mm. HISTORY LIFE pigment Diagnostic features: Myoseptal above gut; pigment over spreading posteriorly forward to notochord beginning mm, after 6.5 by transformation; pigment ventrally melanophores 2-3 past D origin & temperate CalCOFl region to tropical in Throughout Range: worldwide (usually 1); 8 0-2 on myomere postanal at beginning tail waters rakers gill ceratobranchial ca. (by mm); 12-16 preanal myomeres II (usually 31-33 15-16 before, 12-13 during transformation), (usually maximum Habitat: m with 50-1900 ca. at & mesopelagic, Epi- myomeres. total 31-32) abundance ca. 400-800 depth m in MORPHOMETRICS & mean (range %) Probably summer-fall, based on occurrence of small Spawning season: larvae Y-S PrF PoF F Tr Juv eggs & ELH planktonic Oviparous; pattern: larvae Sn-AlBL 59-64 56-63 56-64 49-55 48-52 61 60 61 51 53 LITERATURE 9-11 BDIBL 8-11 14-17 9-11 14-15 1982 Gorbunova 10 10 10 16 15 al. et 1989 Matarese HLlBL 16-19 17-19 16-19 23-29 24-26 1991 and Olivar Fortufio 17 18 17 27 25 1986a Oda & Ozawa HWIHL 53-60 49-57 43-57 27-44 29-42 55 53 47 35 36 ORIGINAL ILLUSTRATIONS (D1ustrator) 18-23 SnLIHL 20-24 17-23 14-20 14-19 21 21 22 17 16 mm (B. Sumida MacCall) 5.8 larva, Preflexion 26-31x EDIHL* 24-30x 18-25x 7-9x 6-7x Flexion larva, 5.8 Sumida (B. mm MacCall) 28-32 26-32 7-10 18-26 7-8 mm, Postflexion larvae, mm (B. Sumida MacCall) 10.0 13.7 (W. Watson) mm Juvenile, 21.1 29x30 26x29 21x22 8x8 7x7 P,LlBL 3-5 3-4 2-4 12-18 9-13 4 4 3 15 12 axis horizontal oval; axis slightly Eye * second. vertical first, given is 0-0 PzLIBL 0-0 0-2 10-19 8-11 0 0 0.4 10 13 250

263 acclinidens Cyclothone Benttooth bristlemouth mm 5.8 5.8 mm mm 10.0 13.7 mm 21.1 mm Figure Gonostomatidae 1. Late preflexion larva, 5.8 mm (CaICOFI 6210, station 103.80); flexion larva, 5.8 mm (CalCOFI 6610, station (CalCOFI 123.50); larva, 10.0 mm (CaICOFI 6601, station 120.55); late postflexion larva, 13.7 mm postflexion 6210, station 103.80); juvenile, 21.1 mm (CFRD 8701, station 78.3.54.5, deep bongo). 251

264 GONOSTOMATIDAE Cyclothone atraria EARLY LIFE HISTORY DESCRIPTION MERISnCS EGGS Mode Range Shell diam.: 0.46-0.58 mm* Yolk: Vertebrae: No. of OG: OG: of Diam. 31-33 32 Total Shell surface: 14 12-14 Precaudal Pigment: 17-20 18 Caudal features: Diagnostic Fins: 0 0 spines Dorsal 13-14 LARVAE 12-15 Dorsal rays length: Hatching 0 0 Anal spines length: Flexion 18-19 17-20 Anal rays Transformation 11.5-14 ca. mm length: 6 6-7 Pelvic sequence: development Fin 9 9-11 Pectoral sides along pair symphysis; cleithral Posiflexiorr-At of Pigmentation: Caudal: 10+9 gas on gut anteriorly; over 10 bladder; lover end of hindgut; 10+9 Principal internal, II ventrally along tail; above myosepta gut anteriorly; Procurrent: center A on bases; external ray at parhypural; on hypural of 5-7 6-7 Upper extending anteriorly to 6 melanophores margin; over notochord, 5-8 6 Lower dorsal over Transformatiorr-Scattered surface of origin. past D rakers: Gill upper of half head 14 7 7-10 & by body mm; along margins of Upper branchiostegal membranes ca. 11.5 by membranes covering mm, 13 13-18 Lower by 12-14 mm. 14 12 Juvenile-Black. Branchiostegals above gut; Diagnostic over pigment Myoseptal features: pigment notochord (Ozawa & Oda 1986a); melanophores predorsal 6-7 & LIFE HISTORY by 8-9 (usually 9) ceratobranchial gill least at rakers, present with 11.5 mm, probably sooner; transformation specimens 13-14 Ocean Range: Pacific North 14) preanal (usually myomeres, 31-32 (usually 32) total myomeres. at Habitat: Meso- and bathypelagic, depth ca. m ~400 400 to m (range & mean in %) MORPHOMETRICS Spawning season: PrF Y-S F Iuv PoF Tr oviparous with planktonic pattern: Protandrous hermaphroditism; ELH & larvae eggs Sn-AJBL 54-57 50-53 55 51 LITERATURE 15-17 15-16 BDIBL 16 15 Mukhacheva 1964 Miya 1991 Nemoto & 24-27 HLlBL 24-30 1986a & Ozawa Oda 27 26 29-32 32-37 HWIHL nONS ORIGINAL ILLUSTRA (Illustrator) 34 31 14-17 SnLIHL 14-18 mm 14.2 Watson) (W. specimen, Transformation 16 16 8-lOx 7-8x EDlHLt 8-10 7-8 1991. Miya ova: * Hydrated & Nemoto 9xlO 8x8 vertical axis second. oval; axis slightly t Eye is horizontal given first, 15-16 13-15 P1LIBL 16 14 P L1BL 10-12 8-13 2 10 11 252

265 bristlemouth atraria eye/othone Black mm 7.5 I [./' l i 1\ I 1 14.2 mm Figure Gonostomatidae 2. Postflexion larva, 7.5 mm (Ozawa and Oda 1986a); late transformation specimen, 14.2 mm (CFRD 9104, station 70.85, MOCNESS). 253

266 GONOSTOMATIDAE Cyclothone pseudopallida MERISTICS EARLY DESCRIPTION HISTORY LIFE Range Mode EGGS Vertebrae: Shell 0.48-0.58* diam.: Yolk: 29-34 33 Total No.orOG: Diam.ofOG: 12-14 surface: Shell Precaudal 13 Caudal 17-21 Pigment: 20 features: Fins: Diagnostic spines Dorsal 0 0 12-15 13-14 rays Dorsal LARVAE Hatching length: 0 0 spines Anal Anal rays 19-20 17-21 mm 5-6 ca. length: Flexion 6-7 Pelvic 6 mm 16.5-22 ca. length: Transformation 9-10 9 Pectoral , , C & A & D C" PI P Fin sequence: development 2 2 Calldal: pairs along Pigmentation: of gut Flexion-transformation-I-2 sides 10+9 10+9 Principal gas over symphysis; cleithral at pair anteriorly; end over bladder; of Procurrent: 7-12 increasing anteriorly, to hindgut; gut on myosepta above 10-12; 6-7 6-7 Upper A on 9-11 internally, ventrally along tail; on bases; ray externally 6-7 Lower 7 C rays; hypural margin; on lower of on upper parhypural; 1 at center rakers: Gill 12 mm; over notochord tip in flexion larvae; over ca. after C rays 4-7 4 notochord posteriorly, 8-11 to anteriorly extending beginning Upper 11-14 11 mm; 16.5 ca. by head D past melanophores on origin Lower in specimens. transformation 14 14 Branchiostegais over pigment gut; above pigment Myoseptal features: Diagnostic LIFE HISTORY after notochord predorsal 8-11 to melanophores up mm, 5.5 by pigment transformation; at beginning tail on ventrally postanai gill & (usually I) on lower Crays; 7 ceratobranchial 0-1 myomere Range: Worldwide north of 30° S (usuaiiy 10-11 before, (by rakers mm); preanal myomeres 12-18 16 myomeres. total 33) (usually 32-34 transformation), during 13 depth m 300-1400 ca. at bathypelagic, and Meso- Habitat: (range & mean in %) MORPHOMETRIes Spawning Postflexion season: larvae collected in most months with May-October occurring smallest specimens Y-S PrF PoF F Tr Juv pattern: Oviparous; planktonic eggs & larvae ELM Sn-AJBL 59-72 62-62 48-58 47-52 62 63 51 49 LITERATURE BDIBL 9-9 7-10 9-17 12-13 9 1982 Gorbunova 9 13 12 Nemoto & Miya 1991 HLIBL 15-20 18-19 16-27 21-23 1991 Olivar and Fortufio 18 18 23 22 Ozawa & Oda 1986a HW/HL 42-54 37-51 26-30 27-34 49 44 27 29 ILLUSTRATIONS ORIGINAL (Illustrator) SnL/tiL 16-24 18-25 13-18 15-20 21 23 17 15 Watson) (W. 5.5 larva, Flexion mm x 24-25 EDlHLt 15-25x 7-ox 6-7 x Postt1exion (W. Watson) mm 10.8 larva, 27-31 16-26 7-8 8-9 (W. Watson) Transformation specimen, 14.4 mm 20x21 25x29 7x8 7x9 Watson) Juvenile, 21.1 mm (W. P,UBL 4-4 2-3 3-14 11-12 4 3 11 11 Miya ova: Nemoto Hydrated * 1991. & LIBL 0-0 0-1 2-14 10-12 P2 oval; slightly Eye t second. vertical first, given is axis horizontal axis 0 0.3 10 11 254

267 Slender bristlemouth pseudopallida Cyclotlwne mm 5.5 mm 10.8 14.4 mm 21.1 mm Gonostomatidae 3. Flexion larva, 5.5 Figure (CaICOFI 7210, station 20.90); postflexion larva, 10.8 mm (CaICOFI 7210, station mm part early transformation specimen, 14.4 mm (FRONTS 85, station 15, tow 1); juvenile, composite of 21.0 mm (head, anterior 39.141); (CFRD gut, some OA and IV photophores) and 21.1 mm (remainder of fish) specimens of 9104, MOCNESS; 21.0 mm specimen from 70.80). station 66.7.90, 21.1 mm specimen from station 255

268 GONOSTOMATIDAE Cyclothone signata MERISTICS HISTORY DESCRIPTION LIFE EARLY Range EGGS Mode Vertebrae: Shell diam.: Yolk: Total No.ofOG: 30-32 31 Diam.ofOG: surface: Shell Precaudal 13 13 Pigment: 18 17-19 Caudal features: Diagnostic Fius: Dorsal spines 0 0 LARVAE 13-14 13-14 Dorsal rays Hatching length: <2.4 mm 0 0 Anal spines Flexion length: 19-20 mm 5-6 ca. 18-20 rays Anal mm 12-14 ca. length: Transformation 6 6 Pelvic 8-10 , , D, C sequence: development Fin PI P C A, , Pectoral 9 2 I 2 along gut; over Pigmentation: Preflexion-flexion-2-4 pairs gas Caudal: 10+9 10+9 (>4.5 bladder margin ventral on 6) (usually 5-10 mm), tail of Principal at postanal beginning a gap myomere 3-5, usually of in middle series Procurrent: present (gap I under 75%); in situated along becomes tip, notochord Upper 6 6-8 5-7 6 Lower 6-8; increase to Posiflexion-transformation--Gut pairs parhypural. Gill rakers: 7-12 to 2-4 myomere postanal at beginning tail series increases 3-4 3-5 Upper 50%; in present gap 3), (usually of A base middle along row double 9-II (>ca. hindbrain (>10.5 mm); over under column, 8 mm); vertebral 9 Lower 13 12-14 Branchiostegals posteriorly beginning on stomach (~12.5 mm). mm); (> 1I.5 above gut; no pigment Diagnostic features: No myoseptal pigment HISTORY LIFE until notochord over end first of stage; postflexion 2) (usually 1-3 in postanal myomeres unpigmented; usually a gap middle of ventral preanal tail eyes usually distinctly oval; 13-15 (usually 14-15) series; throughout CalCOFI Eastern region Range: & central Pacific; myomeres, 30-32 (usually 31) total myomeres. Epi- Habitat: usually & mesopelagic, in upper 800 with depth m (range in %) MORPHOMETRICS & mean abundance maxima upper in 400-500 & m 100 m season: Possibly year-round with sununer-fall Spawning summer-fall, or Y-S PrF F PoF Tr Juv maximum Sn-AlBL 53-61 55-59 50-61 48-51 44-48 56 57 58 46 50 larvae & ELH Oviparous; planktonic eggs pattern: BDIBL 8-10 9-II 10-12 12-14 15-16 LITERATURE 9 10 II IS 13 HUBL 15-18 14-17 14-19 23-27 22-23 Ahlstrom 1974 17 16 18 22 26 a1. 1984c Ahlstrom et 50-68 54-59 HWIHL 40-64 34-41 32-42 et Matarese a1. 1989 58 57 50 35 36 18-27 SnLIHL 14-26 20-26 17-18 16-18 ORIGINAL ILLUSTRATIONS (Illustrator) 21 20 23 17 17 EDIHL* 27-37x 27-29x 100IIx 16-30x 8x Sumida MacCall) 5.0 mm (B. larva, Preflexion 29-42 32-37 11-12 19-34 9-10 mm 5.2 larva, Flexion MacCall) Sumida (B. 28x34 30x36 24x27 8x9 IIxII larva, mm 9.2 Postflexion (H. M. Orr) M. mm 13.4 Orr) (H. Transformation specimen, 4-5 4-5 2-5 PILlBL 7-II 14-16 mm 13.3 Juvenile, MacCall) Sumida (B. 5 3 4 9 IS P L1BL 0-0 0-0 0-2 8-9 10-13 2 0 0 I 8 12 is second. Eye oval; horizontal axis * given first, vertical axis 256

269 Showy bristlemouth signata Cyciothone 5.0mm 5.2mm ORK 9.2mm 13.4 mm 13.3 mm Figure Gonostomatidae 4. Preflexion larva, 5.0 mm (CalCOFI 6609, station 97.55); flexion larva, 5.2 mm (CalCOFI 6609, station 110.50); postflexion mm (CalCOFI 7210, station 100.90); late mm larva, 13.4 (CalCOFI 6608, station l23.40);juvenile, 13.3 larva, 9.2 postflexion mm (CFRD 8701, station 78.2.54.6, MOCNESS). 257

270 Dip/ophos proximus GONOSTOMATIDAE DESCRIPTION EARLY LIFE HISTORY MERISTICS EGGS Range Mode diam.: Shell Yolk: Vertebrae: No.ofOG: Diam.ofOG: 86--88 85-93 Total surface: Shell 36--38 36--39 Precaudal 49-52 Pigment: 48-55 Caudal features: Diagnostic Fins: 0 0 spines Dorsal LARVAE 10 9-11 Dorsal rays mm Hatching <4.9 length: 0 0 Anal spines length: ca. Flexion 18-19 mm 22 ca. through mm 57-59 54-62 rays Anal Transformation length: 38-43 ca. mm* 8 8 Pelvic , , A, PI C sequence: development Fin P C D, 9-10 9 Pectoral b 2 2 flnfold, median to dorsal in Yolk-sao-Many Pigmentation: migrating Caudal: patches. into aggregating & margins body ventral & flexion-- Pre 10+9 10+9 Principal single postflexion-Patches condense to melanophores: 14-23 Procurrent: tail, 4 numbers 3-5 on dorsally, 8-15 10-17 size; with increase ventrally Upper 3-5 isthmus; on 2-3 gut; over on mm; 18-19 C by on hindbrain; 1 under 4 Lower surface inner mm; base by ca. 22 mm; on mandible by ca. of 35 PI Gill rakers: over mm. becomes half Tran.y{ormation-Upper 39 ca. by hindbrain 3 3 Upper 9 9 pigmented. Lower Diagnostic features: Very elongate & slender; at size large relatively Branchiostegals & transformation stages; dorsal & large of series ventral flexion 11-12 dorsally, 14-20 or patches, melanophores usually pigment LIFE HISTORY ventrally (on tail), during most development; of larval 40-48 (usually total 87-89) (usually 85-91 myomeres, preanal 42-43) myomeres. southern to Range: California; uncommon north Pacific tropical Eastern southern California of Baja MORPHOMETRICS mean %) & in (range Epi- Habitat: & mesopelagic PoF Y-S PrF F Juv Tr from Spawning season: Larvae October-February principally collected 61-72 60-63 66-68 Sn-AlBL 49-53 48-49 56--61 67 67 62 59 51 48 larvae & eggs planktonic Oviparous; pattern: ELH 5-8 BDIBL 8-8 7-9 4-6 4-5 4-6 LITERATURE 4 6 5 4 8 8 11-14 12-17 11-17 HUBL 15-15 10-13 14-15 15 15 11 13 13 14 31-35 26--30 HWIHL 32-43 36--53 31-34 26--39 44 32 37 28 32 32 ILLUSTRATIONS (Illustrator) ORIGINAL 4-18 31-35 19-33 32-37 SnLIHL 34-36 30-34 26 33 9 32 35 35 (W. Yolk-sac Watson) 5.1 larva, mm 11-15x 11-17x 15-26x 33-45x 13-13x 15-16x EDlHLt (W. mm 7.1 Watson) larva, Preflexion 13-17 11-14 i6--26 13-14 22-38 15-16 larva, 19.2 mm Flexion (W. Watson) 21x20 15x15 41x30 16x15 12x13 13x13 Watson) larva, (W. Posttlexion mm 48.1 1-3 2-2 2-3 0-3 5-9 2 2 1 3 7 lOt * Larvae shrink during transformation. 0-0 0-0 0-3 5-8 5-6 0-0 round by transformation; becoming oval, initially Eye t nearly "horizontal" 1 0 0 0 5 7 axis of "vertical" second (long first, given eye typically inclined axis slightly from true vertical). specimen. t Pectoral fin rays broken in all but one juvenile 258

271 Diplophos proximus esi<~:;;:~;:2:~=;!;~~~<:~.~~~C.~/i:-~) 5.1 mm 7.1 mm 19.2 mm 48.1 mm Gonostomatidae 5. Yolk-sac larva, Figure mm (CalCOFI 6001, station 147.50); preflexion larva, 7.1 mm (IATTC Mazathin Project 5.1 4-8-0); flexion larva, 19.2 mm (EASTROPAC II, station 45.379); postflexion larva, 48.1 mm (EASTROPAC I, station MZ 8.9). The fine not transverse along the hindgut are folds shown in these illustrations. 259

272 GONOSTOMATIDAE Diplophos taenia MERISTICS mSTORY DESCRIPTION LIFE EARLY Mode EGGS Range Shell diam.: Yolk: Vertebrae: No.ofOG: 93-98 Diam.ofOG: Total 89-99 Shell surface: 38-40 37-41 Precaudal Pigment: 52-59 55-57 Caudal features: Diagnostic Fins: 0 0 spines Dorsal LARVAE 11 10-11 Dorsal rays Hatching length: ca. 0 mm 3.5-5 0 Anal spines ca. mm Flexion 27.5-28 ca. through mm 19-19.5 length: 63-66 59-72 rays Anal 7-8 mm* 8 through 30 ca. length: Transformation 43 Pelvic 9 8-10 Fin PI , , P C D, A, Cl> sequence: development Pectoral z z to limited Yolk-sac-Apparently of Pigmentation: end notochord. Caudal: 10+9 10+9 double continuous Prejlexion-pos!flexion-Initially on nearly rows Principal Procurrent: dorsal tail, & ventral margius of dorsal condensing into ca. 20-40 4 Upper 3-6 11-20 patches; pigment & 9-22 ventral (tail) melanophores or 3-4 3-4 Lower 1-4 pairs over gut; on isthmus after ca. mm; ventrolaterally on 8 by rakers: Gill 17-20 mm; decreasing mrn; 39 ca. on mandible at ca. hindbrain 3 3 ca. on notochord tip after 19 Upper on mm. Trans/ormation-Increasing 7-9 lateral entire midline. along 9 surface & dorsal Lower 12-14 features: Diagnostic Branchiostegals elongate & slender; large size at flexion & 13 Very or melanophores of series & dorsal stages; transformation ventral usually 25-35 dorsal, 15-17 tail), (on ventral pigment patches, HISTORY LIFE most during (usually 43-50 development; larval of preanal 47) myomeres. total 91-95) myomeres, 89-98 (usually Range: 30° to N 40° ca. Cosmopolitan, in eastern S except not from Pacific tropical MORPHOMETRICS (range %) & mean in at m 100 upper in mesopelagic, & ca. 450- & at night Habitat: Epi- PrF Y-S PoF F 610 m depth during the day Tr Juv highest on based year-round occurrence; season: Spawning Perhaps larval Sn-AlBL 62-69 63-70 57-61 49-52 60-67 47-51 67 66 63 59 51 49 October-November catches 5-8 4-7 BDIBL 4-4 7-9 4-5 7-9 ELH pattern: Oviparous; planktonic eggs & larvae 4 6 5 4 8 8 11-18 HUBL 14-17 11-13 10-13 16-17 14-16 LITERATURE 15 14 12 17 11 15 31-42 HWIHL 29-44 28-32 25-30 22-27 22-23 1984c al. et Ahlstrom 39 33 30 27 25 23 Grey 1964 2-12 SnLIHL 20-28 30-38 32-35 33-37 30-33 Jespersen 1934 24 34 35 5 33 32 Jespersen 1919 & TAning 27-48x 16-30x 1I-14x 14-17x 11-13 x EDlHLt 12-16x Oda & 1986b Ozawa 21-31 18-27 12-14 16-18 9-15 11-16 Rudometkina 1981 29x24 38x26 15x16 12x12 12x13 14x14 (Illustrator) ILLUSTRATIONS ORIGINAL 2-2 2-4 1-2 0-2 PILlBL 5-6 ~ 2 1 2 2 6 Watson) 5.1 mm (W. Yolk-sac larva, 0-0 0-0 5-5 PzLIBL 0-0 0-1 5-5 mm (W. Watson) Preflexion larva, 17.2 0 0 0 0.3 5 5 Flexion larva, 21.5 (W. mm Watson) (W. Watson) 43.0 mm larva, Postflexion transformation. shrink Larvae * during becoming t Eye initially oval, nearly round in postflexion; "horizontal" of inclined eye second typically (long axis firSt, "vertical" given axis vertical). true from slightly in ~ fin rays broken Pectoral all juvenile specimens examined. 260

273 Diplophos taenia mm 17.2 21.5 mm ~ .. ~:: = .. mm 43.0 Gonostomatidae 6. Yolk-sac larva, 5.1 mm (CaICOFI 5908, station 100.85); preflexion larva, 17.2 mm (CaiCOFI 5910, station Figure mm 6010, 120.80); postflexion larva, 43.0 station (CaICOFI 7205, station 20.141). The fine flexion larva, 21.5 mm (CaICOFI 107.70); not transverse along the hindgut are folds shown in these illustrations. 261

274 GONOSTOMATIDAE Gonostoma atlanticum DESCRIPTION EARLY LIFE HISTORY MERISTICS Range Mode EGGS Shell diam.: Yolk: Vertebrae: 38 Total Diam.ofOG: No.ofOG: 38-39 18 Shell surface: 18 Precaudal 20 20 Pigment: Caudal features: Diagnostic Fins: 0 0 spines Dorsal 16-18 17 rays Dorsal LARVAE mm length: Hatching <3.5 0 0 spines Anal 28-30 Flexion length: ca. 4.5-6 mm 26-31 Anal rays mm mm, >14.8 length: Transformation to ca. 21 6-8 mm <17.6 7 Pelvic 9-11 development sequence: Fin , P 10 C & A PI C & D, Pectoral b 2 z Preflexion-flexion-Series of 5-13 Pigmentation: ventral along Caudal: 10+9 10+9 sparse tail; of bladder gas over fmfold; caudal on ventrally margin Principal after ca. behind just gut of sides along pair Procurrent: or PI; 4 mm; present 7-8 7-8 in Postflexion-transformation-17-18 loop. hindgut on absent Upper 7 6-7 Lower ventral tail series; double row forms along A base; each along row Gill increasing from PI to hindgut; gut of entire gut area over rakers: side 6-7 Upper peduncle on few transformation; during caudal edges; hypural near 11-12 II Lower ca. by hindbrain 6-7 mm; on anteriorly ca. by midbrain pair on ll-12 on head. Branchiostegals 1l.5 mm; increases 37-40 myomeres (usually 17-18), preanal 17-19 features: Diagnostic in loop below hindgut characteristic HISTORY 38); (usually myomeres total LIFE oval BL; 55-60% bladder; usually length preanal gas prominent tail, of bladder, gas on dorsally & margin ventral on pigment eye; well area, Worldwide; CaiCOFI in from Range: offshore Oregon, pigment. laterally on gut; no internal parhypural & California California, Baja %) in mean & (range MORPHOMETRICS m at Habitat: Meso- & bathypelagic, ca. 100-2500 primarily depth; 490-560 during at night & m m 150-300 the day PrF F v-S PoF Tr Juv smallest with months most in collected larvae Larvae season: Spawning May-October occurring primarily Sn-AlBL 55-61 56-63 54-60 55-58 53-56 58 59 56 56 55 eggs & larvae Oviparous; pattern: planktonic ELH 8-11 BDIBL 10-12 ll-14 17-20 14-16 II 12 10 15 19 LITERATURE HUBL 17-22 19-23 19-25 23-26 21-23 20 21 21 22 25 1974 Ahlstrom 47-54 HWIHL 46-58 37-50 32-37 32-43 1984c a1. Ahlstrom et 51 50 43 35 35 1988a 1986a, Ozawa SnLIHL 20-27 18-31 17-29 24-27 18-22 1980 Rudometkina 23 22 24 21 26 28-33x 25-32x 30-33x EDIHL* 22-26x 18-20x (Illustrator) ILLUSTRATIONS ORIGINAL 31-43 34-37 29-37 23-28 18-20 31x37 29x33 32x36 25x25 19x19 Preflexion larva, 5.4 mm (8. Sumida MacCall) 6.0 mm (8. Sumida MacCall) Flexion larva, 3-6 4-5 4-5 PILlBL 9-12 3-4 specimen, Transformation 19.8 (W. Watson) mm 4 5 4 5 II 0 0-2 0 5-7 4-5 L1BL P2 0 0 0.2 4 7 horizontal Eye initially oval, becoming round during transformation; * second. axis vertical first, given is axis (narrow) 262

275 atlanticum Gonostoma Atlantic fangjaw mm 5.4 6.0mm : : . ';' . . 12.0 mm 19.8 mm Figure Gonostomatidae 7. Preflexion larva:, 5.4 mm (CaICOFI 7205, station 24. 137); flexion larva, 6.0 mm (CaICOFI 6912, station 1 13.80); postflexion larva, 12.0 mm (Ahlstrom 1974); transformation specimen, 19.8 mm (SIO 73-160). 263

276 GONOSTOMATIDAE Gonostoma ebelingi MERISTICS LIFE HISTORY DESCRIPTION EARLY Range Mode EGGS Vertebrae: Shell diam.: Yolk: 44-45 43-46 No.ofOG: Total Diam.ofOG: 19-20 Shell surface: 19-20 Precaudal 23-27 Pigment: Caudal 24-26 Diagnostic Fins: features: 0 0 spines Dorsal 12-14 rays 13 Dorsal LARVAE 0 spines mm Anal <3.4 length: Hatching 27-28 mm Anal mm, >4.6 length: Flexion 26-29 rays ~5.2-7 8 8 mm 15-16 ca. length: Transformation Pelvic 9-13 Pectoral , & P, D C A, & P C" sequence: development Fin 2 2 flexion-None. Pos!flexion-Initially Caudal: gas Pigmentation: over Pre 10+9 10+9 hindbrain on only; bladder ventrolaterally mm. 6.5 ca. at Principal Procurrent: by mm; pigmented Transformation-Stomach 15.4 completely 10-12 II Upper area, gut over anteriorly bladder. around farther extends gas 7-10 8 Lower 42-45 Diagnostic 15-19 preanal myomeres (usually 16), features: Gill rakers: between total 6-7 myomeres (usually 44-45); usually myomeres Upper 8 8 preanal origin; A anus & ca. usually length BL; 47-50% 12 II-12 largely pigment before eye narrow moderately transformation; Lower Branchiostegals transformation. & hindbrain before restricted to gas bladder HISTORY mean in %) (range & MORPHOMETRICS LIFE water; in in CalCOFI area, well Pacific Y-S warm Ocean offshore PoF F PrF Tr Range: Juv California California & Baja from Sn-NBL 39-53 44-56 46-56 49-54 48 51 48 51 52 depth night m 125-300 ca. at primarily at Mesopelagic, & Habitat: day the during m 520-700 10-14 BDIBL 8-12 13-16 15-18 12 II 10 14 16 Spawning primarily collected Larvae season: April-October HUBL 21-25 17-24 22-24 19-22 23 20 20 21 23 ELH pattern: planktonic eggs larvae & Oviparous; HWIHL 46-53 38-48 35-42 29-46 49 42 42 38 39 LITERATURE 18-29 SnLIHL 19-26 23-26 19-28 25 24 25 22 21 Ahlstrom 1974 22-27x 22-28 x EDIHL* 14-19x 19-20x Ahlstrom et al. 1984c 38-41 28-40 15-24 26-27 25x40 23x31 (Illustrator) ORIGINAL ILLUSTRATIONS 23x33 17x20 19x26 P,LlBL 4-7 7-21 ~ 5-6 MacCall) Sumida (B. mm larva, 3.5 Preflexion 5 5 5 6 II Flexion larva, Watson) mm (W. 7.0 P 0-0 L1BL 2-6 0-3 7-14 2 (R. C. mm Walker) 21.0 Juvenile, 0 I 4 0 9 axis narrow narrow; second. Eye * moderately axis long first, is given 264

277 Ebeling's fangjaw ehelingi Gonostoma 3.5 mm 7.0mm 15.0 mm 21.0 mm Figure Gonostomatidae 8. Preflexion larva, 3.5 mm (TC 8406, station 191, Tucker trawl); late flexion larva, 7.0 mm (CaICOFI 7205, juvenile, station early transformation specimen, 15.0 mm (Ahlstrom 1974); 27.145); 21.0 mm (TC 8406, station 192, Tucker trawl). 265

278 GONOSTOMATIDAE Gonostoma elongatum MERISTICS LIFE DESCRIPTION mSTORY Range Mode EGGS Vertebrae: Shell diam.: Yolk: 40-41 39-41 Total No.ofOG: Diam.ofOG: 16 15-16 Precaudal Shell surface: 24-26 Caudal 25 Pigment: Fins: Diagnostic features: 0 0 spines Dorsal 11-15 12 Dorsal rays LARVAE Anal spines 0 0 Hatching length: <3.9 mm 27-32 28 rays Anal mm 4.7-5.3 ca. mm Flexion 6.1-6.4 ca. through length: 7-8 Pelvic mm 8 ca. length: Transformation 12-19 12 Pectoral 10-13 sequence: development Fin , C P & D A, PI C & h 2 2 Caudal: bladder gas over initially; Preflexion-flexion-None Pigmentation: by 10+9 10+9 Principal Posiflexion- hindbrain on at 5.2 mm. ca. 4.7 mm; ventrolaterally ca. Procurrent: stomach, mm; Dorsally surrounding most of gut by ca. 10.2 on 10-13 11 Upper mm; hindbrain, ca. beginning on dorsolaterally 11 beginning at 8-10 8 Lower region mm. Trans/ormation-Increasing on gut & between 8-11 Gill rakers: mm. 18.9 by midbrain onto extending hindbrain; ca. 7-9 Upper preanal myomeres (usually 16 15-18 Diagnostic through features: 11-12 Lower 39-41 total myomeres (usually 39); thereafter), 18 stage, flexion 12-13 Branchiostegals usually length anus between myomeres 0-1 preanal origin; A & BL through transformation stage; moderately narrow usually 45-50% LIFEmSTORY stages; postflexion eye; pigment little dorsolateral through by 8-11 mm. pigmentation on hindbrain CalCOFI area, well offshore from California & Worldwide; in Range: & %) in mean (range MORPHOMETRICS California Baja y-S & and night at at depth m 60-265 ca. bathypelagic, Habitat: Meso- PrF F Tr PoF Juv day during the m ;::500 Sn-AlBL 45-50 43-54 41-59 48-50 52-54 48 47 49 49 53 Spawning season: 10-13 BDIBL 10-12 12-20 15-18 18-20 pattern: hermaphroditism; ELH Protandric oviparous with planktonic 11 12 17 19 16 eggs & larvae 19-22 HLlBL 19-28 19-23 23-27 23-25 21 21 24 24 23 LITERATURE 37-55 53-57 HWIHL 42-51 42-54 31-49 48 48 55 48 41 1974 Ahlstrom SnLIHL 24-34 26-32 24-35 19-26 28-30 Ahlstrom et aI. 1984c 29 29 30 30 24 1964 Grey x 23-33 EDIHL* 25-34x 26-34x 26-34x 16-17x Jespersen & 1926 Uning 1919, 35-39 36-38 33-39 30-36 16-19 1988a 1986a, Ozawa 27x37 30x38 30x36 17x17 29x32 (Illustrator) ORIGINAL ILLUSTRATIONS 3-6 4-6 5-9 PILlBL 4-9 9-19 5 7 5 15 8 MacCall) Sumida Preflexion larva, 4.6 mm (B. L1BL P 0-0 0-0 0-4 9-13 6-7 2 larva, Flexion 4.7 mrn Watson) (W. 0 0 2 10 6 transformation; end the by round nearly becoming oval, Eye * of given second. axis is vertical first, horizontal axis (narrow) 266

279 Elongate fangjaw elongatum Gonostoma 4.6mm 4.7mm .~ .: : . . . . " .. ' . ,," ; . . . . . :'.: .'.: . .' . . .. ,.' " :" ... : ... ,,' " . . : ;" :,- :".'" ::' ,' .. .' . . ... " ". " ... ,: 9.8mm Figure Gonostomatidae 9. Preflexion larva, 4.6 mm (Tasaday I, tow 69); flexion larva, 4.7 mm (TC 8406, station 159, Tucker trawl); early transformation specimen, 9.8 mm (Ahlstrom 1974). 267

280 STERNOPTYCHIDAE: Hatchetfishes WATSON W. 49 about contains genera Sternoptychidae in ten species and slender (BD initially are ca. elongate 10-15% BL), nine (Nelson species 1994); the in of four genera occur with length ca. a preanal BL, 33-50% eyes narrow Current 1). Sternoptychidae (Table vicinity California (narrow axis ca. 35-60% little or no oflong axis), and species), larvae of Argyropelecus (all four The Dana- melanistic deep- become sternoptychines The pigment. while remain bodied, maurolicines the relatively Sternoptyx least species) two phos (at are and oculatus, transformation. during slender in CalCOFI plankton samples; collected regularly genera In most the gut becomes the wider, and shortens eye least at slightly, outside Valenciennellus tripunctulatus occurs largely transformation. very Pigment typically remains during larvae occa- the CalCOFI are area only and taken its in gut areas and head light and is largely internal the sionally stations. the most offshore at beginning the through of transformation. are cm), <10 (usually small sternoptychids Adult body Although a large literature exists concern- of meso- bioluminescent, and bathypelagic compressed, stages history life ing early the (e.g., of sternoptychids contains fishes. The family two subfamilies: Sternop- and Jespersen Ahlstrom 1931a; Sanzo 1919; Hming tychinae, the deep-bodied hatchetfishes; and Maurolic- 1984; Ahlstrom 1983, 1974; Badcock 1977; Belyanina elongate inae, that once were included in the fishes et al. 1984c; larval with problems 1988a), Ozawa the common Gonostomatidae. All share characters of identifications remain. Argyropelecus, especially and the of some least at within photophores the having problem genera. The identities have been Sternoptyx, photophore united in common glands, and groups of gradually acquiring both the the are Current California the in Argyropelecus larval of groups and photophore now but resolved, photophores within each the group during transformation Sternoptyx larvae still carmot be reliably. separated This juvenile (Table the to Sternoptychidae 2). stage with the simultaneous development of contrasts most The following species accounts are based photophores, all or on the photophore each within single, all group in gonostomatids and phosichthyids. examinations detailed and literature on specimens of35 of 25 A. hemigymnus, 32 A. Argyropelecus affinis, with oviparous, are Sternoptychids eggs planktonic lychnus, 52 Danaphos 35 sladeni, A. 37 oculatus, for described been have eggs planktonic larvae: and Sternoptyx and spp., 36 Valenciennellus tripunctulatus Argyropelecus hemigymnus (Sanzo 1931a) and Mauro- Meristic 3). obtained Sternoptychidae were (Table data licus muelleri (Sanzo 1931a; Mito 1961a, 1966), while from Haruta Kawaguchi Baird and (1971, 1986), planktonic larvae known for several species repre- are and Baird (1976), Badcock (1980), (1984a); and Fujii two but all senting of Ahlstrom (e.g., genera et the ten study. this during made given counts Myomere counts M and hemigymnus Argyropelecus 1984c). al. "Diagnostic under features" in following descrip- the to refer made counts tions muelleri eyes, examined a large with the unpigmented with hatch on specimens droplet and segmented yolk, and study this in and yolk sac, with an oil may not reflect the full range for each fins pectoral without from is information Ecological species. (1971, Baird (Sanzo mouth functional or 1931 a; Mito in a hatch others Presumably, 1966). the (1974), Clarke 1986), Baird and (1980). Badcock and state similar larvae known All development. of 268

281 Table 1. characters for the stemoptychid species reported to occur in the California Current region. Sternoptychidae Meristic 10+9 have fin rays. principal All caudal Vertebrae Fin rays Species BrR GR CV PrCV D Total A C PI P2 2 27-29 38-41 10-11 12-14 10-11 8-9 6 10-12+3--4 Argyropelecus affinis 18-22 10 10-11 25-28 11-12 8-9 36-39 11 9-10+5 hemigymnus A. 18-24 6 10 35-37 24-26 12 10-12 6 9-11+6 11 9 16-18 10 A. lychnus 11-12 9-11 35-38 11-12 24-26 9 6 10-11+6-7 17-21 A. sladeni 10 12-13 26-28 38-40 10-17 23-26 6-7 6 7-8+2-3 13-15 9-10 Danaphos oculatus 16-18 11-13 27-30 9-11 13-16 10-11 7'+7-8 5-6 diaphana Sternoptyx 6 6-9 17-19 29-31 12-15 9-11 5-6 9-11 11-13 7+7-8 S. 7-9 obscura 6 12-13 16-18 27-31 9-11 13-16 5-6 9-11 7+7-8 7-9 S. pseudobscura 6 12-17 21-22 32-35 7-10 22-25 12 6-9 8-9+5-6 14-17 9-10 Valenciennellus tripunctulatus Sternoptychidae 2. Order of photophore group development (first appearance of photophores in each group) in Table defined Stemoptychidae. are shown in Figure Stomiiformes 1 and groups in Table Photophore some 1. An Stomiiformes indicates simultaneous development, or insufficient specimens to determine sequence. .,&" Photophore group development sequence Species & I & PIO SAB, SO & BR & AN & SC & PO, SP, PRO, PAN AB, affinis Argyropelecus SP & PO & SC AN, SO, I, SAB PTO, & PAN PRO, & AB & BR, hemigymnus A. AN PAN PTO, AB, SO & BR & I & & & SC, PO & SP, PRO, SAB lychnus A. & & SAB AB SO & BR & I, AN, SC, PO, SP, PRO, PAN & PTO sladeni A. & SAB AN BR & AB, SO & I & & PO, SC, PRO oculatus Danaphos SO, AN, PO, SAN SC, AB & I & & BR & PTO, SP, PAN diaphana Sternoptyx PTO & PAN AB & BR, SAN & I, SP, AN, SC, PO, SO, obscura S. SO BR AB, PAN, PO & I, AN, & & SP & SC, PRO & SAB tripunctulatus Valenciennellus Table 3. Number of specimens (above) and size ranges Sternoptychidae mm, below) used in the preparation of the (in sternoptychid species descriptions. An ilL" indicates used in the description. literature Yolk-sac Eggs Flexion Preflexion Transformation Postflexion Species Juvenile 0 0 10 5 4 Argyropelecus affinis 11 5 3.7-6.9 8.5-10.8 9.4-11.5 10.4-14.6 14.9-17.8 La La 2 2 hemigymnus 11 A. 5 5 6.0-6.9 7.1-10.0 8.6-10.8 8.7-11.9 13.4-20.5 2 10 1 0 6 A. lychnus 8 5 4.6 6.5-10.0 9.3-9.7 4.5-7.1 8.7-12.6 14.2-19.1 A. sladeni 0 0 10 5 7 10 5 7.8-10.0 6.7-9.4 4.3-7.3 8.2-13.1 14.1-19.0 Danaphos 7 5 10 oculatus 0 7 5 4.5 4.5-7.9 8.3-13.2 12.8-18.1 17.5-23.6 26.2-34.6 10 0 12 5 Sternoptyx spp. 0 15 10 3.1-5.8 5.9-7.2 6.5-9.6 6.8-11.3 8.4-16.9 0 10 Valenciennellus 5 10 0 6 5 3.7-4.6 5.2-7.3 6.3-8.9 9.5-15.9 tripunctulatus 21.2-26.1 a Sanzo 1931a 269

282 STERNOPTYCHIDAE Argyropelecus a/finis HISTORY DESCRIPTION LIFE EARLY MERISTICS Range EGGS Mode diam.: Sliell Vertebrae: Yolk: No.ofOG: 39 Diam.ofOG: Total 38-41 surface: 11 Shell Precaudal 10-11 Pigment: 28 27-29 Caudal features: Diagnostic Fills: spines Dorsal 0 0 LARVAE Dorsal 8-9 rays 9 length: Hatching <3.7 mm 0 spines 0 Anal 12-14 Flexion ca. ca. mm 13 length: through mm 7-8 9-11 Anal rays through mm 10.5-11.5 ca. length: Transformation 14.5- ca. 6 6 Pelvic 15.0 mm* 11 10-11 Pectoral sequence: development Fin Caudal: P D, PI' & A , C C , I 2 2 10+9 10+9 Pigmentation: Preflexion-posiflexion-Internally on notochord at Principal 5-\ as in myomeres 0 last far forward as extending myomere, last Procnrrent: 11 some. gas bladder; around stomach; on Upper 10-12 Transformation-Over 4 3-4 frontals between eyes; subsequently increasing over fore- & Lower on on midbrain, opercular area, gut; lateral midline series begins at Gill rakers: 18-22 18-20 ca. 13-16 pigment condenses to one peduncle caudal Totsl internal mm; blotch. Upper 7-9 7-8 11-12 pigment 9-13 short 1-10 last at myomeres; Lower features: Diagnostic Internal 10 38-41 10-15 preanal myomeres (usually 12-13), length; preanal Branchiostegais 10 groups SC & AN in photophores 39); (usually myomeres total widely spaced. LIFE HISTORY %) in mean MORPHOMETRICS (range & Worldwide; Range: seaward the primarily area CalCOFI of in shelf continental Y-S PrF f PoF Tr Juv ca. primarily 100-600 mesopelagic, & depth m Habitat: Epi- Sn-AlBL 37-45 36-46 34-42 31-54 49-55 40 41 39 40 53 Possibly season: Spawning occur most frequently larvae year-round; from spring through fall 12-14 9-11 9-13 BDIBL 17-40 40-43 11 11 12 41 30 ELH pattem: Oviparous; planktonic eggs & larvae HLlBL 15-20 17-18 17-18 16-28 24-28 18 17 18 22 26 LITERATURE HWIHL 36-51 38-50 41-49 44-62 59-89 44 44 45 50 70 Belyanins 1984 SnLIHL 23-33 26-44 31-36 30-44 29-35 Matarese 1989 al. et 28 34 37 34 33 I 7-23 x 17-21x 18-19x 32-40x 20-37x ED/HLt ORIGINAL (Illustrator) ILLUSTRATIONS 34-42 37-58 33-38 32-51 35-48 20x45 19x38 18x36 33x42 37x45 MacCall) Sumida larva, mm 5.2 Preflexion (B. larva, rVlacCall) Sumida (D. mm 10.4 Flexion 3-4 3-4 2-5 PlLiBL 27-29 6-27 specimens, (B. mm MacCall) 11.5 Transformation Sumida 11.0 mm, 3 3 3 20 28 Watson) Juvenile, 16.0 mm (W. 0-0 0-0 0-0 0-7 6-9 L/BL P2 0 0 0 8 2 CLIHL 0-4 2-5 2-4 0-0 0-2 Larvae * transformation. part early of during shrink the 3 3 3 0 0.3 t horizontal is vertically elongate; Eye axis is given first, vertical second. 270

283 a/finis Argyropelecus Slender hatchetfish mm 5.2 10.4 mm mm 11.0 11.5 mm 16.0 mm Figure Sternoptychidae 1. Preflexion larva, 5.2 mm (CaICOFI 6301, station 130.45); late flexion stage larva, 10.4 mm (CaICOFI 8803, mm (CaICOFI 6201, station 90.80); late transformation specimen, 11.5 (CFRD 11.0 mm station transformation 78.3.58.3); early specimen, (FRONTS 16.0 mm 85, station 3, MOCNESS). juvenile, 8701, 75.2.52.3); station 271

284 STERNOPTYCIDDAE Argyrope/ecus hemigymnus DESCRIPTION MERISTICS HISTORY LIFE EARLY Range EGGS Mode Shell diam.: mm 0.92-1.04 Vertebrae: Yolk: Segmented 37-38 No.ofOG: I 36-39 Total Diam. of OG: 0.26-0.28 mm Shell Smooth surface: 11 11 Precaudal Pigment: None 25-28 26-27 Caudal Fins: Diagnostic features: Segmented yolk; secondary membrane inside chorion 0 Dorsal spines 0 8-9 8 rays Dorsal 0 0 LARVAE spines Anal 11-12 11 Hatching mm length: 2.5 Anal rays 6 6 10-11 Pelvic mm Flexion length: ca. length: ca. 7.8-12 Transformation 10-11 10 mm* Pectoral , C P D, C Caudal: & A , PI' sequence: Fin development I 2 2 10+9 Yolk-sac through posiflexion-None. 10+9 Pigmentation: Principal form; to begin Procurrent: Transformation-None before AN photophores 9-10 9-10 Upper eye; to region adjacent frontal on rnm; 10.2 ca. by stomach around 5 5 posttemporaVupper supracleithrum region; on opercle lower along Lower eye; by gas bladder below ca. 9 mm; subsequently increases over Gill rakers: 19-23 18-24 half heavily Juvenile-Anterior pigmented; Total head area. gut & on midline. discontinuous stripe along lateral Upper anterior no pigment; tail internal features: Diagnostic No midbrain Lower 10 eyes; transformation in stage; conical pigment choroid tissue below 10 Branchiostegals length myomeres long preanal preanal 14-18 transformation; before subsequently, before 11-14 transformation, 37-38) 36-39 (usually LIFE HISTORY myomeres. total area, Current California throughout Worldwide; Range: the of seaward & %) in mean (range MORPHOMETRICS shelf continental & m 100-700 ca. at principally rnesopelagic, depth Epi- Habitat: F YS PrF PoF Tr Juv year-round; most Possibly frequently 44-52 Sn-AlBL larvae occur Spawning season: 49-53 48-59 32-54 50-56 48 51 53 42 53 through winter from spring BDIBL 10-10 9-12 12-14 16-43 46-51 Oviparous; planktonic eggs & larvae pattern: ELH 10 II 13 30 48 19-20 HUBL 17-21 18-20 19-26 25-29 LITERATURE 20 19 19 22 26 37-47 46-49 HWIHL 41-45 41-78 46-109 1984 Belyanina 42 47 43 57 84 Borodin 1931 SnLIHL 32-33 33-41 33-37 32-43 31-41 1906 Brauer 36 33 35 36 36 Ehrenbaum 1909 13-14x 14-15x ll-14x 12-36x 32-38x EDlHLt Taning 1919 Jespersen & 36-41 35-38 37-42 42-52 41-52 et al. 1989 Matarese ..,t: ~A z:. .. 14x40 13x38 Olivar & 13x37 Fortuna 1991 ~VI\"T--' 36x48 1931a Sanzo 3-3 2-5 4-5 PILlBL 4-25 28-31 3 3 4 IS 29 (Illustrator) ILLUSTRATIONS ORIGINAL 0-0 P L1BL 0-0 0-6 0-0 6-9 2 0 0 0 2 8 6.9 larva, Preflexion MacCall) mm (B. Sumida CLlBL 4-9 2-7 5-9 0-7 0-0 mm Flexion larva, 10.8 (B. MacCall) Sumida 6 5 7 0 2 Larvae shrink during the early part of transformation. * t second. Eye is vertically elongate; narrow axis is given first, long axis 272

285 hemigymnus Argyropelecus Spurred hatchetfish mm 0.92-1.04 6.9mm mm 10.8 10.9 mm 9.9 mm 11.2 mm Figure Sternoptychidae 2. Egg, 0.92-1.04 mm; yolk-sac larva, 2.5 mm (redrawn from Sanzo 1931a); preflexion larva, 6.9 mm (CaICOFI mm; 8012, 83.3.80); flexion larva, 10.8 mm (CaICOFI 6801, station 90.37); early transformation specimen, 10.9 station mid-transformation 11.2 specimen, mm; late transformation specimen, 9.9 mm (modified from Sanzo 1931a). 273

286 Argyropelecus lychnus STERNOPTYCHIDAE EARLY LIFE HISTORY DESCRIPTION MERISTICS EGGS Mode Range diam.: Yolk: Shell Vertebrae: Diam.ofOG: No.ofOG: 36 35-37 Total surface: 11 Shell 11 Precaudal 25 Pigment: 24-26 Caudal features: Diagnostic Fins: o 0 Dorsal spines LARVAE 9 9 Dorsal rays mm <4.6 length: Hatching o 0 Anal spines ca. length: Flexion through mm mm 6.5-7.0 9-10 ca. 12 12 Anal rays ca. Transformation length: mm* 8.7-9.3 mm through ca. 12.5 6 6 Pelvic Fin P D, C & PI A, C sequence: development , 11 10-12 b z Pectoral z along Pigmentation: through pos!flexion-Internally Yolk-sac Caudal: notochord at last myomeres. 2--6 10+9 gas Transformation-Over 10+9 Principal around stomach; bladder; frontals over on eyes; between midbrain Procurrent: increasing & hindbrain by ca. 9 mm; subsequently on opercular 9-10 9-11 Upper gut on area, along supracleithrumlposttemporal region, and area; 6 6 Lower internal condenses. pigment caudal peduncle rakers: Gill Internal last at Diagnostic pigment features: 2-6 myomeres; 9-18 16-18 Total preanal postflexion stage, 9-11 15-17 (usually myomeres through Upper 35-37 35-36); in transformation (usually myomeres total stage), Lower very and length preanal transformation, before long relatively 10 10 Branchiostegals with tissue. conical narrow eye choroid HISTORY LIFE (range & MORPHOMETRICS %) in mean possible Pacific; Eastern Range: California in record; Ocean Indian PoF F Y-S PrF Juv Tr area, Current seaward of continental shelf 47-55 45-52 Sn-AlBL 55-61 43-56 28-57 Habitat: depth m 200-700 ca. at primarily pelagic, meso & Epi- 50 49 49 39 37 58 Possibly from occurrences larval most season: Spawning year-round; 16--61 12-15 10-13 9-11 BDIBL 57--69 winter through spring 64 11 11 13 33 7 18-32 18-19 19-23 18-22 30-35 HUBL plailktonic ELH pattern: Oviparous; eggs & larvae 23 20 18 20 33 13 21-34 59-82 28-39 28-46 45-81 HWIHL LITERATURE 29 50 32 37 52 69 28-33 37-40 29-39 28-40 28-31 SnLIHL Belyanina 1984 31 29 39 17 35 35 Matarese al. 1989 et 14-16x 12-17x 12-16x 31-41x 16-37x EDlHLt 40-48 36-40 34-44 28-39 38-47 (Illustrator) ORIGINAL ILLUSTRATIONS 29x42 23x50 14x32 15x38 35x43 15x39 Preflexion 5.6 MacCall) Sumida (B. mm larva, 33-36 5-33 2-3 3-5 3-5 PILlBL Flexion larva, 4 Watson) mm 9.3 4 2 (W. 0.4 20 34 MacCall) mm (B. Sumida 9.3 Postflexion larva, 0-0 0-0 0-8 10-11 0-0 PzLIBL (B. Sumida MacCall) 8.7 mm specimen, Transformation 0 0 10 2 0 0 Juvenile, Watson) (W. mm 19.1 4-10 CLIHL 0-3 0-0 3 3-8 0 3 6 7 0 * shrink during the early part of transformation. Larvae is vertically elongate; horizontal axis second. given first, vertical t Eye 274

287 Tropical hatchetfish /ychnus Argyrope/ecus 5.6mm 9.3 mm .. !. ;/ ./ / ... / ,'/ , ... ... ;'.····: .. ~~r~~\'?-:\-"'·· _\.~ .. \.-(~". ~======:::-"--c; -" 1>~ 9.3 mm 8.7 mm mm 19.1 Figure Stemoptychidae 3. Preflexion larva, 5.6 mm (CaICOFI 7501, station 107.60); late flexion larva, 9.3 mm (CaICOFI 5902, station (FRONTS 90.60); larva, 9.3 mm (CaICOFI 7712, station 113.35); transformation specimen, 8.7 mm postflexion 85, station 8, MOCNESS); 19.1 mm (CFRD 8803, station 74.8.55.2). juvenile, 275

288 STERNOPTYCIDDAE Argyropelecus sladeni EARLY LIFE HISTORY DESCRIPTION MERISTICS Mode Range EGGS Shell diam.: Vertebrae: Yolk: 36 No.ofOG: Total Diam.ofOG: 35-38 Shell surface: 11-12 11 Precaudal 25 24-26 Pigment: Caudal Diagnostic features: Fins: spines o Dorsal 0 LARVAE 9 rays 9 Dorsal length: Hatching o 0 spines Anal mm 8.5-9.4 ca. through mm 7.5 ca. length: Flexion 12 Anal rays 11-12 ca. 8.2-10.0 mm through ca. 13.0 Transformation mm* length: 6 Pelvic 6 , sequence: development Fin CI> & D P C & A 11 PI> 9-11 Pectoral 2 2 Pigmentation: Preflexion-posiflexion-None. Transformation-None Caudal: (white mm 10+9 before SC photophores form; by ca. 9.2-9.5 SC 10+9 Principal at present this size) on frontal region adjacent to eye, photophores Procurrent: anteriorly side on each posttemporallupper midbrain, lower of 10 along 10-11 Upper bladder, over gas around region, supracIeithrnm stomach; 6-7 6 Lower subsequently increases over gut area, laterally above on gut, Gill rakers: opercular over brain. area, & 17-21 Total tail; features: No internal Diagnostic in pigment pair of large 7-8 Upper stage; transformation midbrain over in anteriorly melanophores Lower 10-11 10 11-12 eye; mass choroid short tissue length; preanal below narrow 10 Branchiostegals preanal myomeres myomeres. total 12),35-37 (usually LIFE mSTORY %) MORPHOMETRICS (range & mean in continental area, CalCOFl in Worldwide; of primarily seaward Range: PrF F Y-S Tr PoF Juv shelf Sn-AlBL 36-43 37-43 38-42 depth Epi- m 100-600 at primarily mesopelagic, & 33-58 56-62 Habitat: 41 39 44 58 39 Possibly season: Spawning occurrences from larval most year-round; 10-12 21-52 BDIBL 12-14 14-17 52-57 fall spring through 11 13 15 40 54 ! HUBL 19-21 19-31 18-22 18-20 27-29 ELH & eggs planktonic Oviparous; larvae pattern: 19 20 20 25 28 40-50 37-46 40-111 HWIHL 39-49 84-117 LITERATURE 45 40 43 76 96 29-43 SnURL 24-36 29-37 29-32 31-36 Belyanina 1984 34 31 31 34 33 Matarese et a1. 1989 14-20x 20-37x 13-19x 14-19x 33-42x EDlHLt 35-47 34-44 44-51 37-49 41-52 ORIGINAL ILLUSTRATIONS (Illustrator) 16x40 16x38 16x40 32x44 38x48 larva, Preflexion Sumida (B. MacCall) mm 5.2 P L1BL 2-6 30-32 9-31 3-7 3-5 1 mm MacCall) Sumida (B. 9.4 larVa, Flexion 4 4 4 23 31 larva, Postflexion mm 10.0 (B. Sumida MacCall) 0-0 L1BL 0-0 0-9 P 0-0 8-11 2 8.2 mm (B. Sumida MacCall) Transformation specimen, 0 0 3 0 9 Juvenile, 13.8 Watson) (W. mm 1-3 0-4 0-0 CURL 0-3 1-3 2 2 1 3 0 early the during Larvae * transformation. of shrink part vertical first, t Eye is vertically elongate; horizontal axis is given second. 276

289 Lowcrest hatchetfish sladeni Argyropelecus S.2mm 9.4mm 10.0 mm 8.2 mm 13.8 mm Figure Sternoptychidae 4. Preflexion larva, 5.2 mm (CalCOFI 6501, station 130.40); flexion larva;9.4 mm (CalCOFI 7412, station 90.37); (CFRD postflexion 10.0 mm (CalCOFI 7501, station 107.60); transformation specimen, 8.2 mm larva, 8803, station 75.3.52.2); juvenile, 8803, station 68.8.55.8). 13.8 mm (CFRD 277

290 Danaphos oculatus STERNOPTYCIDDAE DESCRIPTION HISTORY LIFE EARLY MERISTICS EGGS Mode Range diam.: Shell Yolk: Vertebrae: No.ofOG: Diam.ofOG: 38-39 38-40 Total surface: Shell 12 12-13 Precaudal Pigment: 26-27 26-28 Caudal Diagnostic features: Fins: 0 0 spines Dorsal LARVAE 6 6 rays Dorsal length: Hatching 0 0 spines Anal ca. through 8 mm ca. length: Flexion mm 13 25 23-26 Anal rays length: Transformation mm 23.5 ca. through mm 16.5-18.0 7 6-7 Pelvic , , Fin sequence: development P C PI' D C 16 A, 10-17 h 2 2 Pectoral Pigmentation: of end until Preflexion-posiflexion-None postflexion Caudal: Transformation- gas over bladder. stage forms pigment when 10+9 10+9 Principal Only on gas bladder around & midbrain on posteriorly initially; Procurrent: of gut & region cover to increases mm; 21.5 ca. stomach by most 8 7-8 Upper ca. forms IImyomere of series by head dorsolateral mm; 23.5 ca. 3 2-3 Lower on ca. by 7-8 frrst myomeres along extends mm, 23.5 lateral rakers: Gill midline to caudal peduncle stage. juvenile in 13-15 Total myomeres II-IS 14-15 (usually preanal features: Diagnostic 2 2 Upper through postflexion stage, in II-12 37-39 stage), transformation II-13 Lower in (usually 38-39); eye less total myomeres than narrow 10 9-10 Branchiostegals Argyropelecus; transformation mm); at large size (~16.5 beginning long A base, with short 3 AN first only base; D photophores HISTORY LIFE base. common Range: area, Current California of seaward throughout Pacific Ocean; (range %) in mean & MORPHOMETRICS shelf the continental Y-S F PrF Tr PoF Juv Habitat: Epi- & depth m 183-914 at primarily mesopelagic, Sn-AlBL 50-54 48-53 40-44 40-49 34-39 throughout the year Spawning season: with Larvae collected highest 45 51 50 44 43 37 January-August catches 7-8 7-10 22-24 6-10 BDIBL 9-21 ELH pattern: Oviparous; planktonic eggs & larvae 8 8 8 8 14 23 17-21 17-21 16-18 12-17 HLlBL 16-21 LITERATURE 15 17 19 18 IS 19 38-56 39-50 58-76 43-60 35-56 HWIHL Ahlstrom 1974 47 44 48 36 43 65 a1. 1984c et Ahlstrom 17-28 20-25 26-38 SnLIHL 32-42 30-35 Matarese al. et 1989 6 31 21 24 36 33 15-18x 15-18x 20-33 x 13-21 x EDIHL* 39-54x ORIGINAL (Illustrator) ILLUSTRATIONS 35-53 32-43 34-40 31-43 39-50 30x47 17x38 16x44 16x35 25x38 43x43 Watson) (W. mm 4.5 larva, Yolk-sac MacCall) (B. mm Sumida larva, Preflexion 5.2 3-5 2-6 7-20 3-6 P]LlBL 20-23 4 4 4 13.2 larva, Flexion 13 (B. 22 mm MacCall) Sumida MacCall) Sumida mm (B. 16.5 larva, Postflexion 0-0 0-0 1-8 P L1BL 9-10 0-0 2 mm Transformation specimen, 19.2 Watson) (W. 0 0 0 0 3 9 vertically elongate, initially Eye * becoming nearly round in juvenile Narrow axis is given stage. first, long axis second. 278

291 Danaphos oculatus BottleUght 4.5 mm 5.2 mm 13.2 mm 16.5 mm mm 19.2 22.5 mm Sternoptychidae 5. Yolk-sac larva, 4.5 mm (CFRD 9104, station 71.7.75); preflexion larva, 5.2 mm (CaICOFI 7503, station 97.45); Figure mm mm 6912, station 77.65); postflexionlarva, 16.5 (CaICOFI (CaICOFI 7412, station 113.50); early transformation larva, 13.2 late flexion transformation late station 73.3.80, MOCNESS); 9104, specimen, 22.5 mm (Ahlstrom 1974). specimen, 19.2 mm (CFRD 279

292 STERNOPTYCHIDAE Sternoptyx Spp. HISTORY DESCRIPTION LIFE MERISTICS EARLY EGGS Range Mode Shell diam.: Vertebrae: Yolk: 27-31 No.ofOG: Total Diam.ofOG: 11-13 Precaudal Shell surface: 16-19 Pigment: Caudal Diagnostic features: Fins: Dorsal spines 0 LARVAE 9-11 Dorsal rays length: Hatching 0 <3 mm Anal spines 12-16 mm 6-7 ca. length: Flexion rays Anal Pelvic 5-6 mm 7-11 range within Variable length: Transformation ca. 9-11 Pectoral Fin development sequence: , C PI> CI> P D, A, z z Caudal: Pigmentation: Prejlexion-pos!flexion-Initially none; develops on 10+9 present the of any in absent or be following may area; gut & Principal head forebrain, Procurrent: of posterior margin any on areas specimen-anterior 7 margin Upper midbrain, hindbrain, under of at tip of lower jaw, on middle 7-8 Lower of gular membrane, at anterior margin gut, of on gas bladder, on inner surface membrane; rakers: Gill of PI base, on PI internal on in notochord 6-9 Total some S. diaphana. Trans/ormation-Increasing tail head & of on 2-3 Upper upper trunk, extending caudad & ventrad. gut areas; developing on 4-7 27-30 (usually 28-29); preanal Lower Myomeres features: Diagnostic 6 before transformation, body very deep during very length short Branchiostegals & internal pigmented; lightly transformation; after tail pigment of in diaphana, some HISTORY LIFE S. usually posteriorly. MORPHOMETRICS %) & (range mean in Mediterranean Worldwide polar seas; in California except Range: & of continental shelf seaward region, Current Y-S F PrF PoF Tr Juv 400-1500 bathypelagic, at ca. & m depth Meso- Habitat: Sn-AlBL 31-36 29-36 31-41 32-62 44-57 34 32 36 45 51 CalCOFl in Spawning year season: collected Larvae throughout the catches winter with area, somewhat larger BDIBL 14-18 19-22 23-32 22-79 73-97 20 15 47 26 83* & eggs Oviparous; pattern: ELH planktonic larvae HLlBL 19-25 20-25 21-31 20-27 26-37 22 23 23 26 31 LITERATURE HWIHL 42-56 50-62 46-82 52-116 52-122 49 55 60 94 79 Ahlstrom et 1984c aI. SnLIHL 34-46 27-45 30-49 25-48 19-40 & Baird 1980 Badcock 41 37 38 39 30 1983 Belyanina x 20-27 24-32x 22-39x 22-54x EDlHLt 33-62x aI. 1989 Matarese et 36-48 38-46 36-46 43-67 32-60 23x41 27x41 (Illustrator) ILLUSTRATIONS ORIGINAL 30x40 36x44 48x53 3-6 3-5 PILlBL 11-37 6-12 30-40 mm 5.2 larva, Pretlexion (W. Watson) 3 4 21 9 35 larva, 6.4 mm Flexion (H. Orr) M. 0-0 0-0 0-7 PzLIBL 8-12 0-0 (H. M. Orr) mm 11.2 specimen, Transformation 0 0 3 10 0 * (mean 91%) for S. diaphana, 84-97% (mean 75%) BDIBL= 73-77% for S. obscura. vertically elongate (usually more so for S. obscura than S. t Eye for diaphana), nearly round during transformation for S. becoming obscura; or diaphana juvenile for S. in narrow axis is given first, the long second. axis 280

293 Dollar hatchetfishes spp. Stemoptyx 5.2mm ~ .. ... : ... : 6.4 mm mm 8.8 11.2 mm Sternoptychidae Figure Preflexion larva (probably S. obscura), 5.2 mm (EASTROPAC II, station 75.036); early flexion larva (possibly 6. S. diaphana), 6.4 mm (NORPAC, station 156); postfiexion larva (probably S. obscura), 8.8 mm (Ahlstrom et al. 1984c; although neither are pectoral nor pro current caudal rays rays shown in this illustration, formation fin of both normally is nearing completion in specimens diaphana, of size); transformation specimen of S. this 11.2 mm (CaICOFI 5009, station 70.175). 281

294 STERNOPTYCIDDAE Valenciennellus tripunctulatus EARLY HISTORY DESCRIPTION MERISTICS LIFE Range EGGS Mode Shell diam.: Vertebrae: Yolk: 32-35 Total No.ofOG: 33 Diam.ofOG: surface: 12 Shell Precaudal 12 21-22 Caudal 21 Pigment: features: Diagnostic Fins: 0 0 spines Dorsal 7-10 7-8 LARVAE rays Dorsal 0 length: spines Anal Hatching 0 22-25 ca. mm through rays Anal mm 5 ca. length: Flexion 24 6.3-7.3 6-9 7 Pelvic ca. through 9.5 ca. length: Transformation mm 16.0 15-16 12-17 Pectoral development Fin sequence: P C" C A, PI' D & , z 2 Preflexion-flexion--None. Pigmentation: Posiflexion--None Caudal: 10+9 10+9 Principal initially, Transformation-- mm. ca. 8.5-9.0 over gas bladder by Initially Procurrent: 12.0 ca. over gas bladder only; on stomach by mm, 8-9 9 region; increases over little over midbrain, few in dorsolateral gut Upper 5-6 5 beginning form near series to end trunk, along stage. Lower of preanal 12-15 14-15 myomeres (usually features: Diagnostic Gill rakers: 14-17 15 Total transformation initially, decreasing 12-13 stage), 32-34 in total to 2-4 Upper gut with Sn-A/BL ca. slender 33-34); 55-60% 3 (usually myomeres 12-13 12 Lower myomeres postflexion stage; gas bladder far posterior (ca. through 9-10 10 over Branchiostegals transformation. before bladder gas only pigment 10-12); MORPHOMETRICS (range & mean in %) HISTORY LIFE warm Y-S in well water; offshore in Worldwide Range: area CalCOFI PrF F PoF Tr Juv mesopelagic, Primarily Habitat: at 100-700 m depth Sn-A/BL ca. 59-69 60-64 54-63 46-52 44-48 62 63 49 58 46 in area season: Spawning CalCOFI collected spring in Larvae BDIBL 10-12 9-11 10-13 13-25 24-27 10 II II 18 25 & ELH pattern: Oviparous; planktonic eggs larvae HLlBL 20-25 21-24 20-25 21-34 20-23 23 22 23 24 22 LITERATURE 46-54 HWIHL 36-51 46-54 40-71 54-70 48 46 49 51 63 Ahlstrom 1974 25-34 27-33 SnUHL 28-40 34-45 21-40 al. et 1984c Ahlstrom 30 31 38 35 31 1977 Badcock 21-23x EDIHL* IS-20x 17-24x 22-34x 29-44x 1964 Grey 39-40 37-42 35-45 38-47 29-48 Ozawa 1988a 22x40 19x39 27x42 21x39 38x42 ORIGINAL (Illustrator) ILLUSTRATIONS 3-8 2-5 PILIBL 7-11 4-7 4 5 5 9 23t Preflexion Watson) (W. mm 4.1 larva, P L1BL 0-0 0-0 0-0 0.2-8 .8-8 2 MacCall) (B. Sumida mm Flexion larva, 7.3 0 0 0 4 8 Postflexion larva, mm (B. Sumida MacCall) 8.6 26.7 mm (W. Watson) Juvenile, • Eye vertically elongate; narrow axis is given first, long axis second. all specimens fin rays broken in Pectoral but one of the juvenile t examined. 282

295 Constellationfish Valenciennellus tripunctulatus mm 4.1 7.3 mm 8.6mm 13.6 mm 26.7 mm Figure Sternoptychidae 7. Preflexion larva, 4.1 mm (TC 8405, station 41, Tucker trawl); late flexion larva, 7.3 mm (CalCOFI 6301, station 13.6 97.110); larva, 8.6 mm (CalCOFI 7205, station 27.139); transformation specimen, postflexion mm (Ahlstrom 1974); juvenile, 26.7 mm (CaICOFI 7205, station 31.145). 283

296 PHOSICHTHYIDAE: Lightfishes WATSON W. Photich- Phosichthyidae to as (commonly referred phosichthyids larvae the resemble Larval some of thyidae, a more but descriptive incorrect spelling-see gonostomatids set no sternoptychids; and single of species seven 1990) in Eschmeyer contains about 18 the at species all of separation allows larval characters genera. in in occur genera three species the Six Cal- level the However, all genera and most of family. COFI these, among 1); Phosichthyidae (Table region identified can species a combination using by be of and, to Vinciguerria a lesser of lucetia larvae only (especially morphometric eye and length preanal irregularis extent, in regularly occur Ichthyococcus shape), meristic (especially myomere counts), and characters. pigment of most CalCOFI of ichthyoplankton samples. Larvae at the the others are taken from time to time, usually outer their stations, CalCOFI but distributions are three histories Life are well known for six of the seaward primarily of area. CalCOFI the in the species V. CalCOFI Vinciguerria lucetia, region: nimbaria and and Ahlstrom (e.g., poweriae V. Counts phosichthyids are small (usually <10 cm), Adult 1958; Ozawa is available Limited information 1988a). compressed, biolumi- Ichthyococcus), (except slender 1988a). The (Ozawa elongatus Ichthyococcus for nescent bathypelagic. and meso- largely are that fishes literature the on based are accounts species following external differ little from gonostomatids in They 30 of on detailed examinations and (Vinciguerria), appearance. 27 lucetia, Vinciguerria irregularis,35 1 of specimens nonsuchae V. 20 specimens of poweriae, Woodsia and Phosichthyids eggs planktonic with oviparous, are (Table Although 2). Phosichthyidae elongatus 1 is for planktonic described been and larvae: have eggs to range reported and south as the northern east as far Ichthyococcus ovatus (Sanzo 1930) and several Vinci- part CalCOFI area (Rechnitzer and Bohlke of the guerria species (Ahlstrom and Counts 1958; Sanzo found no in larvae were 1958), CalCOFI samples and of known for five planktonic 1931a), larvae are while nimbaria V. is described not here. that species Larval Eggs 1984c). et Ahlstrom (e.g., genera seven the al. rarely in occur very samples CalCOFI and also are not are small a segmented with spherical and mm) «1 described here. Counts Silas and Ahlstrom See (1958), yolk. may oil present or absent, and the An droplet be and Rudomet- (1969), Ozawa (1973, 1988a), George of a double membrane, or not, may shell egg consist (1975), (1979) and et al. Miller Gorbunova kina (1981) on species. depending unpig- have larvae hatching, At for descriptions and illustrations of larval V. nimbaria. eyes, mented a large have and mouth, a functional lack Meristic data were obtained from literature the (Ahls- all development larval Throughout sac. species yolk and Bohlke trom and Counts 1958; Rechnitzer 1958; by characterized are (BD slender elongate and being Fujii from and 1984a) 1980; Mukhacheva 1964; Grey usually length :::;;10% BL), with a long preanal (usually this study. Myomere counts given during made counts on depending not, or trail, may gut The BL). ~66% following descrip- the under "Diagnostic features" in species. typically are oval, at least during the Eyes specimens made counts to refer tions the on examined preflexion stage, but usually become round by post- the in range and may not this study reflect the full for each flexion or transformation Pigment is light to stage. species. mainly on occurs it typically moderate when moderate; lower the the of body. half 284

297 Table Phosichthyidae characters for the phosichthyid species reported to occur in the California Current region. 1. Meristic All 10+9 rays. have caudal principal rays Fin Vertebrae prev Total D A CV P C 2 PI 2 Species BrR GR 29-30 45-47 14-16 7-8 16-18 elongatus Ichthyococcus 6+3-4 7-8 14-17 31-37 12-13 24-25 13-15 11-13 13-15 7-8 7-8 5-6+3 38-39 irregularis 1 8+17-19 12 13-16 13-17 22-24 16-19 39-43 10 lucetia 7-10+4-5 Vinciguerria 7 8-11+18-23 12 13-15 13-16 9-11 7 8-10+4 39-44 V. nimbaria 5-7+13-17 11 13-15 22-23 16-18 38-41 9-11 12-15 7 7-9+4-5 3-4+11-13 poweriae V. 11 11-12 7-8 19-20 42-45 4+4 14-16 23 9-10 Woodsia nonsuchae 3-5+13 17 2. Number of specimens (above) and size ranges Phosichthyidae mm, below) used in the preparation of the Table (in species phosichthyid An "L" indicates literature used in the description. descriptions. Yolk-sac Preflexion Flexion Postflexion Egg Species Transformation Juvenile 1 9 5 10 Ichthyococcus irregularis 0 0 5 3.3-7.9 7.8-9.4 10.0-21.7 4.1 21.6-30.2 La La 6 10 10 Vinciguerria 5 lucetia 4 5.7-7.0 7.3-18.9 2.2-5.5 14.8-21.6 27.4-36.2 La 10 3 0 7 2 poweriae V. 5 4.2-6.5 8.6-22.6 6.6-8.2 17.3-22.6 21.1-24.1 2 10 0 0 8 0 0 nonsuchae Woodsia 2.8-6.2 5.7-8.1 7.6-14.1 a Ahlstrom and Counts 1958 285

298 PHOSICHTHYIDAE Ichthyococcus irregularis DESCRIPTION HISTORY LIFE EARLY MERISTICS Range EGGS Mode diam.: Shell Yolk: Vertebrae: 38-39 No.ofOG: 39 Diam.ofOG: Total surface: Shell 25 Precaudal 24-25 13-15 Pigment: 14 Caudal features: Diagnostic Fins: 0 0 Dorsal spines LARVAE 11-13 11-12 Dorsal rays Hatching 3-4 ca. mm length: 0 0 spines Anal 7.8-7.9 length: Flexion mm 13-15 mm 13 ca. through 9.5 rays Anal mm 21-22 ca. 7-8 Transformation length: 7 Pelvic 7-8 , , PI 8 C & A D, Cl> sequence: development Pectoral P Fin 2 2 Yolk-sac--Small PI on bases; PI patch between ventral Pigmentation: Caudal: 10+9 10+9 bases. Prefiexion-posiflexion-Ventrolateral series along myosepta, Principal caudal onto extends size, larval with increases ca. by fmfold 6 mm; Procurrent: 5 5-6 Upper spreads onto isthmus bases, PI base & ventrum between on increases 3 Lower 3 of on pair mm; 4.9 by membrane branchiostegal by patch 6 mm; mandibular on snout; along bands symphysis extending longitudinal Gill rakers: 8 of gut. jaw by 8 mm; on trailing part of 8 Upper lower sides along PI with after gut trailing portion; lower elongate 17-19 17-19 features: Diagnostic Lower 12 12 Branchiostegals & PI of mm; 5-6 pigmented along lower part ca myosepta, on branchiostegal membrane, stripes on snout; ventrum between PI' on preanal 25-27 (usually total myomeres 37-40 25), (usually myomeres LIFE HISTORY 37-39). Range: Eastern Pacific Ocean; throughout California Current region, MORPHOMETRICS & (range %) in mean seaward shelf continental of Habitat: Meso- Y-S & PrF bathypelagic PoF F Tr Juv Sn-AlBL the year 70-74 69-73 66-74 season: Larvae Spawning occur throughout 74-81 72 72 71 71 78 eggs larvae planktonic Oviparous; pattern: ELH & 7-10 7-8 BDIBL 6-8 34-42 7 8 8 8 39 LITERATURE 15-20 16-20 13-18 HlJBL 25-34 16 14 17 18 31 52-63 29-59 46-55 HWIHL 45-92 48 52 57 51 66 SnLIHL 33-44 29-36 36-43 26-36 ILLUSTRATIONS ORIGINAL (Illustrator) 40 14 32 39 31 22-30x 19-22x 16-22x EDIHL* 27-38x Yolk-sac (B. Sumida MacCall) mm 4.1 larva, 28-43 22-29 18-25 34-43 MacCall) Sumida Preflexion (B. larva, mm 6.9 28x35 21x26 25x33 19x23 32x37 8.8 larva, Flexion mm Sumida MacCall) (B. Sumida MacCall) 18.4 Postflexion larva, mm (B. 2-5 3-5 3-5 23-27 PILlBL 4 4 3 25 6-13 6-13 6-13 PILLlBLt 0-0 * Eye axis oval; (-horizontal) is is given first, long (-vertical) axis narrow 9 9 8 0 second. 0-0 0-0 P L1BL 0-1 14-20 2 lower of elongate of fin. lobe t PILL to length refers 0 0 0 0.4 17 2-8 0-3 TGLlBL 3-17 0-0 0 2 5 12 0 286

299 Bulldog iightfish irregularis Ichthyococcus mm 4.1 mm 6.9 8.8 mm 18.4 mm Phosichthyidae 1. Yolk-sac larva, 4.1 Figure (CaICOFI 7203, station 117.80); preflexion larva, 6.9 mm (CaICOFI 5908, station 103.50); mm late flexion larva (composite illustration, principally of8.8 details mm CalCOFI 5105, station 103.70; from of head and gut added specimen 93.29). station mm specimen, CalCOFI 6204, station 117.90); postflexion larva, 18.4 mm (CaICOFI 7712, 9.4 from 287

300 Vinciguerria lucetia PHOSICHTHYIDAE EARLY LIFE mSTORY DESCRIPTION MERISTICS EGGS Mode Range diam.: mm Shell 0.58-0.74 0.4-0.5 mm diam.* Segmented; Yolk: Vertebrae: No. of None OG: Diam.ofOG: 40-41 39-43 Total Shell surface: Smooth 22-23 22-24 Precaudal Pigment: along embryo of sides series through mid-way Dorsolateral 18 16-19 Caudal in embryo. late and ventrally on tail development, extending onto head Fins: irregularly OG; Diagnostic Double shell membrane; no features: 0 0 Dorsal spines segmented yolk; ca. 0.6-0.7 diameter. mm 14-15 13-16 rays Dorsal 0 0 Anal spines LARVAE 15 13-17 rays Anal length: Hatching ca. 2 mm 7 7 Pelvic through mm mm 7 ca. 5.6 Flexion length: 10 10 Pectoral Transformation ca. 14.8 mmt 19-21.6 ca. length: mm through Caudal: , & Fin development sequence: P C" D, PI 10+9 A C , 10+9 2 2 Principal Yolk-sac-10-12 Pigmentation: on little side; each along dorsolaterally Procurrent: oftai!. pigment ventral margin Preflexion-posiflexion-Dorsolateral 8-9 7-10 Upper migrates by ca. ventrad, disappears 4 mm; usually 1 at cleithrurn after 4 4-5 Lower 4 mm; 1 above end of hindgut; 1-3 on ventral 1-4 tail; of margin ca. rakers: Gill of along 7.5 mm; along bases A base C rays by ca. 6-7 mm. by ca. 8-10 8-11 I Upper on dorsal surface over Transformation-Develops & head, gut, of 18-23 18-23 Lower vertebral caudal internally below peduncle. column in 12 12 Branchiostegals body; features: Diagnostic length ca. preanal slender Elongate, 70% BL; moderately narrow eye; Dray lover myomere 21-22 (vs. LIFE mSTORY myomere ray in V. nimbaria); A 19-20 1 opposite Dray 9-10 (vs. usually myomeres, preanal 25 nimbaria); V. in 10-11 opposite D Range: Eastern Pacific Ocean; throughout CalCOFI region, usually south 40-41 larvae on series melanophore dorsolateral total myomeres;

301 Panama lightfish [ueetia Vinciguerria 2.2mm mm 0.6-0.7 ... 13.1 mm a. 'd d a a . -B .,:',"- -Q····d· mm 15.0 mm 19.5 Figure Phosichthyidae 2. Egg, ca. 0.6-0.7 mm; early preflexion larva, 2,2 mm; late preflexion larva, 6,0 mm; postflexion larva, 13,1 mm; early specimen, transformation 15,0 mm; early juvenile 19.5 mm (Ahlstrom and Counts 1958), 289

302 PHOSICHTHYIDAE Vinciguerria poweriae MERISTICS DESCRIPTION HISTORY LIFE EARLY Range EGGS Mode Vertebrae: Shell mm diam.: 0.75-0.85 Yolk: Segmented Total 40 38-41 No.ofOG: 1 mm OG: of Diam. 0.17-0.19 22-23 Precaudal 23 Shell Smooth surface: 16-18 Pigment: 17 Caudal None Fins: shell inner thin no yolk; Segmented membrane; features: Diagnostic 0 0 spines Dorsal OG single 13-15 Dorsal 14 rays LARVAE 0 0 Anal spines 12-15 length: 13 Anal Hatching rays 7 Pelvic 7 Flexion length: ca. 6.5-8.5 mm 9-11 Pectoral 10 Transformation mm* 17 ca. through mm 22.6 ca. length: Caudal: , , sequence: P, development A, P D, C" Fin C 2 2 10+9 10+9 Principal Pigmentation: Prejlexion-jlexion-Unpigmented (ca. 40%) or 1 on Procurrent: only. one on usually tail, of side at lateral middle midline 7-9 8 Upper mm) or Posiflexion-None on each (specimens

303 poweriae Vinciguerria Highseas lightfish mm 0.8 5.0 mm 6.6mm 10.1 mm 22.9 mm Figure Phosichthyidae 3. Egg, ca. 0.8 mm (Ahlstrom and Counts 1958); preflcxion larva, 5.0 mm (CaICOFI 7804, station 90.150); flexion larva, mm (CaICOFI 7804, station 6.6 90.150); posttlexion larva, 10.1 mm (CaICOFJ 7804, station 93.200); transformation specimen, 22.9 mm (Ahlstrom and Counts 1958). 291

304 PHOSICHTHYIDAE Woodsia nonsuchae MERISTICS EARLY LIFE DESCRIPTION mSTORY Range Mode EGGS Vertebrae: diam.: Shell Yolk: 42-45 42 Total No.ofOG: Diam.ofOG: 23 23 Precaudal surface: Shell 19-20 19 Caudal Pigment: Fins: features: Diagnostic spines Dorsal 0 o 11-12 12 Dorsal LARVAE rays 0 Anal spines o Hatching length: <2.8 mm 14-16 14-16 rays Anal 6 Flexion length: mm 7.5-8 ca. through mm, ca 7-8 Pelvic >14.1 length: Begins Transformation rnrn, mm <20.3 9-10 Pectoral sequence: Fin development Cl> PI D C & , P A, & 2 2 Caudal: along Preflexion-Series Pigmentation: gular of margin ventral 10+9 10+9 Principal & heavy gut; along isthmus of margin along series membrane; ventral Procurrent: along of gut, continuing ventral internal tail; of margin dorsal margin 4 4 Upper series dorsal margin along notochord tip & on of notochord; around Lower 4 4 finfold. caudal C Pos{flexion-Disappears ca. on dorsal by part of Gill rakers: along series A rays; on not or present A base, on mm; 10 few midline 3-5 Upper of roof of mouth by ca. 11 mm. 13 13 Lower Moderately on lower half Diagnostic features: mainly heavy pigment, 17 Branchiostegals 17 of (usually myomeres preanal 29-31 body; myomeres total 41-44 29), (usually 42); preanal length ca. at if little, all; gut BL; 70-80% trails LIFE HISTORY 5) ahead (usually rnyomeres 4-5 insertion D elongate; C rays middle A origin. of subtropical, Tropical California of part Range: & into southern ranging in %) mean & MORPHOMETRICS (range of ar~a, well the seaward shelf continental Current Meso- Habitat: depth m 530-1335 ca. at bathypelagic, & PrF Y-S F PoF Tr Juv through autumn Spawning season: collected Larvae spring Sn-AlBL 66-80 72-77 76-83 74 74 79 & larvae Oviparous; pattern: eggs ELH planktonic BDIBL 8-11 8-11 8-12 9 9 9 LITERATURE HLlBL 20-30 21-28 23-29 24 24 26 al. 1984c et Ahlstrom HWIHL 35-35 31-41 29-35 1988a Ozawa 36 35 31 32-39 SnLIHL 33-42 (Illustrator) ILLUSTRATIONS ORIGINAL 35 38 40 EDIHL 19-25 16-22 mm MacCall) Sumida (B. 5.4 Preflexion larva, 22 20 19 Watson) (W. Flexion 5.7 mm larva, 1-5 PILlBL 3-5 2-3 3 4 3 L1BL P 0-0 0-0 0-1 2 0 0 0.2 292

305 Bigeye lightfish nonsuchae Woodsia mm 5.4 5.7 mm 11.5 mm 20.3 mm Figure Phosichthyidae 4. Preflexion larva, 5.4 mm (EASTROPAC II, station 46.054); flexion larva, 5.7 mm (TC 8405, station 58, Tucker al. 1984c); late transformation specimen, 20.3 mm (Ozawa 1988a). postflexion trawl); larva, 11.5 mm (modified from Ahlstrom et 293

306

307 STOMIOIDEA H. MOSER G. Fink monophyly of Although (1985) demonstrated distinctive and many have highly Stomioid larvae are he not fishes (see Stomiiformes above) stomioid did stom- Typically, structures. morphological specialized propose classification a for these or other formal an elongate body and gut; many in ioid larvae have Eschmeyer stomiiforms. listed (1990) the traditional six trails gut the groups some the In body. the from freely families in his classification but did not stomioid voluminous is but Fin striations. mucosal lacks gut superfamily (1974) Weitzman's recognize Stomioidea. the particularly rays, pectoral and pelvic late form rays, as a convenience guide the superfamily In we use this The in larval development. pectoral fins are positioned separating the six stomioid in other families from species a ventral in have most and low on the body and efficiently stomiiforms to summarize information large; are finfolds the Usually orientation. the however, used 1) and specimens meristics on (Table Stomioidea adult in form in their fins position and not out the species in preparing Stomioidea (Table descriptions finfold. 2). Stomioidea 1. Meristic characters for the stomioid species Table the California Current vicinity. All taxa have 10+9 in principal caudal fin rays except Bathophilus brevis which has 5+6. Sources for meristic information are listed in the introductory section of each family. Vertebrae rays Fin PrCV Total D A CV C Taxon PI P2 2 Chauliodontidae 9-14 56-62 5-7 9-14 3-5 53-56 Chauliodus macouni 6-10 8-10+3-5 Stomiidae 3-5 69-74 16-20 16-22 66-67 6 5 7+4-5 Stomias atriventer Astronesthidae 46-58 10-21 12-22 spp. Astronesthes 5-9 5-9 54-57 15-18 15-17 16-21 38-40 7-8 10-12+4-5 Borostomias panamensis 7 19-24 51-57 22-28 8-12 6-9 33-35 Neonesthes 8-10+4 capensis 6-8 Melanostomiidae 2 10-11 9-12 33-35 1+1 31-33 11-13 8-14 brevis Bathophilus 4+3 43-45 3-5 45-50 13-16 14-17 4-5 B. filifer 42-45 3--4 44--48 14-16 15-17 4-8 15-19 jlemingi B. 5-6+2-3 20-30 32-46 0-13 56-69 6-8 spp. Eustomias 11-15 13-17 63-65 21-26 8-12 50-53 7 6-8+4-5 boureei Flagellostomias 75-83 16-22 20-29 9-11 spp. Leptostomias 7-8 50-57 12-18 16-20 4-7 7-8 spp. Melanostomias 63-65 21-24 21-25 1+4 7-8+4-5 7-8 48-50 15 mitsuii Opostomias 17-24 0-3 15-24 49-65 6-7 spp. Photonectes 14-18 19-22 12-14 79-82 8-9+4 0 67-68 8-10 macro Tactostoma pus MaJacosteidae 54-58 21-25 25-29 4-8 9-ll+5-{j 50-53 5-7 6-7 Aristostomias scintillans Idiacanthidae 54-66 81-85 28--43 0 ldiacanthus antrostomus 6 11-14+7-10 295

308 Table Stomioidea 2. of specimens and size range (in mm, below) used in the preparation of the stomioid Number (above) An "L" literature used in the description. indicates descriptions. species Taxa descriptive summaries in chapter with indicated by an asterisk. introductions are Yolk-sac Preflexion Egg Posttlexion Taxon Flexion Transformation Juvenile Chauliodontidae La I 6 4 9 macouni Chauliodus I 3 4.7 10.0-16.4 18.1-21.3 23.7-48.7 42.3 32.6-35.7 Stomiidae La 2 6 6 6 3 atriventer Stomias 0 2.7-3.4 17.1-29.9 4.4-10.0 10.6-15.7 30.3-41.3 Astronesthidae I Astronesthes I 0 0 sp.* 0 0 0 5.9 8.1 0 0 3 Borostomias 0 panamensis* 0 0 0 13.7-17.2 1 0 0 capensis* 0 0 0 Neonesthes 0 15.2 Melanostomiidae 0 0 0 0 1 0 0 brevis* Bathophilus 9.3 9 4 4 0 0 3 B. filifer 2 3.7-8.1 3.0-3.9 8.3-12.0 12.4-22.3 30.4-43.3 0 4 6 1 8 0 3 B.jlemingi 5.6-11.0 15.0-24.4 12.0-13.5 3.7 39.4-45.7 0 0 0 0 0 0 sp. Eustomias * 18.6 0 0 0 0 0 0 Flagellostomias boureei* 36.4 0 0 0 0 1 0 0 Leptostomias sp.* 16.1 0 0 0 0 0 Melanostomias sp.* 0 16.0 0 0 0 0 0 mitsuii* Opostomias 0 15.0 0 0 0 0 1 0 0 sp. * Photonectes 15.9 La,b 3 9 7 5 3 3 pus Tactostoma macro 14.5-23.4 6.1-13.0 22.7-48.0 5.6-6.0 52.2-55.2 59.1-69.0 Malacosteidae 10 4 0 7 Aristostomias scintillans 0 0 3 6.6-11.8 13.0-17.4 17.1-47.0 44.6-60.7 Idiacanthidae 8 5 0 8 0 0 Idiacanthus antrostomus 3 4.5-17.1 21.2-32.3 32.0-67.4 77.9-85.1 Kawaguchi and Moser 1984 a b Kawaguchi and Moser 1993 296

309 CHAULIODONTIDAE: Viperfishes MOSER G. H. of consists Chauliodontidae eight in the genus species of perivitelline a characteristic be may this space, and species, One 1974). Novikova and (Parin Chauliodus 1977; (Belyanina chauliodontids all and Kawaguchi California macouni, occurs the Current region. It C. in Moser Sandknop 1984). Larvae Matarese 1984; and ranges from central to north Mexico California, Baja at hatch an sac. elongate 6-7 mm and have yolk Alaska of westward the north across and Gulf the Larvae are slender, elliptical in cross section, and have to zone transition Japan. Pacific Larvae C. northern of gut long, extremely an that extends the non-trailing macouni, the of the body. The is relatively small, head second most abundant larval stomioid in length entire CalCOFI surveys, extend southward to central Baja with the eyes. ovoid slightly and snout pointed a short, n.mi. 100-200 California and ca. at abundance in peak The median is small and developed on the best finfold the coast. from the warm All body. the of except the fins, Chauliodus sloani occurs in region posterior identified nearly all oceans and Loeb (1980) of appear caudal, are slow to develop and in late post- regions flexion larvae. Larvae reach a large size (some species of the larvae west mass to the its in the central water shrinkage >40 mm) and undergo considerable during area. study CalCOFI Larval transformation (Kawaguchi and Moser 1984). except is pigment for minute melanophores absent «30 cm), medium-size Viperfishes small to are of C. tip in larvae yolk-sac the at observed notochord of slender, They have fishes. rows five compressed pigment been has sloani by Mito (1961a). This not large deciduous scales embedded in shallow pockets, Chauliodus species. of other each reported for larvae with a more or pattern. pigment hexagonal less Coloration over iridescent blue a black or grey is mouth has huge fanglike teeth The background. large other stomii- are eggs Chauliodus than any larger on The premaxillary and mandible. dorsal fin is the Eggs identified. presently egg form C. of macouni forward well has and elongate first body ray an the on (2.7-3.1 may be confused with those of Pacific mm) anterior fins pelvic The tab. a fleshy in terminating are a have but shell sagax, Sardinops sardine, larger is well There midbody, and is posterior. the fin anal to of macouni C. oil globule. lack Larvae diameter an and ventral a large dorsal adipose fin and a small adipose and are both similar; somewhat are atriventer Stomias pattern photophore The is fin anus. to the anterior C. a larger reaches macouni but and slender elongate and postorbital with distinctive and suborbital organs straight is and gut The mm). 30 vs. mm 50 size (ca. the lateral and ventral series on prominent longitudinal nearly the entire elongate in both species but extends body. Small luminous organs are present each scale on length of relatively is and macouni C. in body the elsewhere present small organs the are and area on S. atriventer. Also, diameter in greater and shorter in in barbel chin short A body. and head some is present below the S. atriventer has serial pigment above and of relatively are Viperfishes species. predators common macouni following The pigment. lacks C. gut and meso- the oceans; world the of zones bathypelagic and on published literature and on description based is to migrate individuals smaller the species some in detailed observations macouni C. of of specimens 24 shallow epipelagic 1964a; (Morrow night at waters (Table 2). Stomioidea Meristic data were from obtained 1984, 1986b). Gibbs Matarese Miller (1972), Lea and the and (1989), al. et from made during this study. Ecological infor- counts (2.2- large have and oviparous are Chauliodontids and sources above the from obtained was mation from 3.6 mm) round planktonic eggs with segmented yolk Morrow (1968), and and (1964a), Fitch Lavenberg of C. macouni and no oil Initial yolk diameter globule. (1984, Gibbs 1986b). a large leaving diameter, shell the one-half ca. is eggs 297

310 CHAULIODONTIDAE Chauliodus macouni MERISTICS HISTORY DESCRIPTION EARLY LIFE Range EGGS Mode Vertebrae: Shell diam.: 2.7-3.2 mm 1.3-1.5 Yolk: Segmented; mm diam. Total 56--62 60 0 No.ofOG: Diam.ofOG: Precaudal 54 53-56 Smooth surface: Shell 4 Caudal 3-5 None Pigment: Fins: features: Large Diagnostic diameter; large perivitelline space; Dorsal spines 0 0 yolk; segmented no 00 rays 5-7 Dorsal 6 Anal 0 spines 0 LARVAE 9-14 12 rays Anal At Hatching length: least 4.7 mm 6-10 Pelvic 18-22 7 mm length: Flexion 9-14 Pectoral length: 33-49 mm Transformation ca. Caudal: , C Fin development sequence: , D, A, P & C PI I 2 2 10+9 10+9 Principal Pigmentation: None Procurrent: of length a maximum attaining elongate, Slender, Diagnostic features: Upper 8-10 9-10 length; small of head, >90% gut straight body elongate mm; 50 almost 4 3-5 Lower of except C late-forming, all far fins lack forward; arched ventrad; D Gill rakers: pigment. Upper Lower %) MORPHOMETRICS (range & mean in Branchiostegals 16-21 Y-S PrF F PoF Tr Juv HISTORY LIFE Sn-A/BL 86-92 93-94 93-97 80-83 87 90 94 95 of Gulf & Sea, Bering Japan, Northern Range: to central Baja 81 88 Alaska California BDIBL 4-6 4-4 4-5 14-15 4 5 4 4 14 7 Epi- Habitat: to bathypelagic to depths >4000 m HUBL 10-12 9-9 7-10 16-17 9 11 9 8 10 17 present Spawning season: Larva throughout year in CalCOFI samples HWIHL 33-40 40-41 37-49 38-42 abundance with February in highest 38 57 42 41 41 40 23-29 SnLIHL 30-32 30-39 22-26 larvae eggs & planktonic pattern: ELH Oviparous; 14 26 31 34 33 24 LITERATURE EDIHL* 20-24x 17-25x 19-24x x 22-23 28-31 29-34 28-40 22-23 Kawaguchi 1984 Moser & 52x33 22x32 22x30 21x32 22x22 22x26 al. 1989 et Matarese 1-2 PILlBL 2-2 1-14 13-14 0 2 2 3 4 13 (Illustrator) ORIGINAL ILLUSTRATIONS 0-0 0-0 PzLIBL 0-1 18-19 0 0 0 0.2 5 19 Arthur) Transformation 42.3 mm (N. specimen, Juvenile, 32.6 mm (N. Arthur) second. * Eye is oval; horizontal axis is given fust, vertical axis 298

311 mac()uni Chauliodus Pacific viperfish 2.9mm 15.0 mm mm 45.2 42.3 mm -~ ----- 32.6 mm Figure Chauliodontidae 1. Egg, 2.9 mm (Matarese and Sandknop 1984); preflexion larva, 15.0 mm; postflexion larva, 45.2 mm (Kawaguchi 32.6 juvenile, (CFRD 8701, station 62.2.56.0); mm mm, pigment not shown (CFRD 9104, 1984); transformation specimen, 42.3 Moser and station 70.0.62.5, MOCNESS). 299

312

313 Scaly STOMIIDAE: dragonfishes G. H. MOSER elongate an have sac. mm 3-4 at hatch yolk and includes species in the genus Stomias Stomiidae 11 cross Larvae are slender, a long with section round in 1969, and Fink 1986a; 1984, Gibbs 1964b; (Morrow that gut in is slightly shorter of relative length than that Fink 1986). the California Stomias atriventer occurs in Chauliodus. a with small, relatively is head The the Current region from central California to south snout moderately long and slightly ovoid eyes. The abundant is It California. of larval the most Gulf and anal is small. develop fins Dorsal finfold median stomioid in the an has and abundance CalCOFI surveys not do the pelvic fins in larvae postflexion but early peak ca. at coast. from n.mi. 300-350 the late until appear stage. Late-stage postflexion the in embryos and atriventer of species some other have S. dragonfishes small are to moderate-size (to Scaly dorsal melanophores that migrate ventrad to form fishes. ca. is body The compressed slender, cm), 40 gut the series above the paired that extend posteriad to covered pigmented of rows with 5-6 areas, hexagonal notochord These larvae. preflexion in tip subsequently a scale a gelatinous by covered structure, like with each less become median and a ventral series apparent Coloration life. in membrane is iridescent or black develops the gut. a Some species develop below fanglike green. The large mouth has moderate-size dark series along the body and have others midlateral and teeth. and anal fins Dorsal fin caudal the near are entire body over (Kawa- melanophores scattered the the are well posterior to midbody in most fins pelvic 1984). and guchi Moser of prominent photophores rows two are species. There the hexagonal areas each ventrally on each and side five of Stomias species been described: have Larvae one have barbel chin The photophores. small more or S. 1973); Rass and (Pertseva-Ostroumova colubrinus S. of 3/4 from than the head to a maximum ranges shorter atriventer ajfinis, S. and 1984); Moser and (Kawaguchi of body length, depending on the species; it terminates with are dragonfishes Scaly filaments. bulb a in S. pacificus (Ozawa and 1988a). Stomias S. nebulosus, relatively common midwater predators oceans, all of the in area CalCOFI the occurs of south colubrinus to at found usually young depths; bathypelagic meso- eastern tropical Pacific; its larvae easily distin- are migrate near- to of some species apparently individuals guished from those since atriventer, S. of they have night 1984, at waters surface (Gibbs 1986a). The body. ofthe half lower the covering melanophores following description based on published literature is pelagic Stomias eggs are round with is oviparous; of specimens S. 23 of and on detailed observations depending to 0.88 on mm, 1.5 ranging from diameters data Meristic 2). Stomioidea (Table atriventer were globule, oil a single have Eggs species. the segmented Ecologi- study. this during made counts from obtained (Kawaguchi membrane inner a distinct and yolk, and Gibbs cal information was obtained from (1984, 1984). Larvae Moser 1984; Matarese and Sandknop 1986a), and Fitch and Lavenberg (1968). 301

314 STOMIIDAE Stomias atriventer DESCRIPTION HISTORY LIFE EARLY MERISTICS Mode Range EGGS 0.88-0.92 mm Shell diam.: Yolk: Segmented; diam. Vertebrae: mm 0.70 1 No.ofOG: Diam. 72 0.20-0.25 mm OG: of 69-74 Total Shell Smooth surface: 66-67 67 Precaudal to continuing embryo, of forward series Paired Pigment: dorsum on 5 Caudal 3-5 brain outline Fins: Diagnostic single features: Small size, membrane, inner OG, 0 0 Dorsal spines segmented yolk 17 16-20 Dorsal rays 0 0 Anal spines LARVAE 19 16-22 rays Anal Hatching 2-3 mm length: 5 5 Pelvic 10-16 mm Flexion length: 6 Pectoral 6 30 mm Transformation ca. length: Caudal: sequence: Fin development & D & PI C & C P A, 10+9 & 10+9 Principal 2 I 2 series pigment prejlexion-Dorsal Yolk-sac-early Pigmentation: Procurrent: trunk & between series ventrolateral paired form to migrates gut 7 7 Upper short notochord posteriad extending tip. notochord above series to tip; 4 4-5 Lower prejlexion-jlexion-Series patch; Late on lateral to isthmus; gular Gill rakers: terminal & hindbrain, lateral region; postorbital jaw, lower section. gut Upper trunk; recessed in ventral gut sparse on above C, Posiflexion-Series Lower & A; on postorbital region. D, Branchiostegals attaining Diagnostic features: Slender, elongate, a maximum length of head small BL); 85-90% (ca. straight elongate mm; 30 ca. gut arched LIFE HISTORY (EL ca. 15% ofHL at postflexion stage); eyes ventrad; small elliptical above series paired stage; C at flexion near A form D & gut; below & of equator the from & California Range: Gulf to California Central to at form fins transformation. northern Chile %) in mean & (range MORPHOMETRICS m <100-1500 at depth Habitat: Meso- & bathypelagic y-S PrF F PoF present Spawning year throughout season: Larvae Juv surveys in Tr CalCOFI September & February in peaks with Sn-AlBL 74-82 84-90 87-89 84-86 87-91 76 86 88 89 85 & larvae Oviparous; planktonic eggs ELH pattern: 6-8 7-7 5-6 5-6 BDIBL 9-9 LITERATURE 6 6 7 7 9 15-18 14-16 13-14 HLlBL 9-13 11-12 Kawaguchi & Moser 1984 16 15 13 12 11 40-42 32-46 HWIHL 30-37 32-35 41-50 ORIGINAL (Illustrator) ILLUSTRATIONS 41 38 33 33 44 SnLIHL 17-20 26-32 33-40 32-38 26-31 30.3 (N. Arthur) mm Transformation specimen, 29 36 36 28 18 11-16x 33-44x 13-23 x EDIHL* 9-12x 16-19x 29-32 17-29 15-20 13-17 16-19 is Eye * second. vertical first, given is axis axis horizontal oval; 18x23 13x18 IOx15 18x18 39x31 2-4 2-2 1-2 0-0 9-11 PILlBL 0 3 2 2 10 0-0 0-0.4 L1BL 0-0 0-0 P 9-12 2 0 0 0 0.2 11 302

315 atriventer Stomias Blackbelly dragonfish 4.6mm 0.9mm 10.0 mm 22.2 mm 30.3 mm Figure Stomiidae 1. Egg, 0.9 mm (Matarese and Sandknop 1984); preflexion larva, 4.6 mm; flexion larva, 10.0 mm; postflexion larva, specimen, 22.2 (Kawaguchi and Moser 1984); transformation mm 30.3 mm (CaICOFI 5406, station 130.50). 303

316

317 ASTRONESTHIDAE: Snaggletooths G. H. MOSER includes 5 genera Astronesthidae ca. 35 species of and pigmentation heavy to melanophores, no or few from tropical midwater predators that occur principally in melano- punctate or myosepta, and finfolds the on are relatively rare subtropical waters. Astronesthids and phores along the body (Kawaguchi and Moser 1984; Ozawa 1988a). area survey CalCOFI Astronesthid larvae can be distin- in the from collections midwater by the forward placement other stomioids guished from (Table panamensis, Stomioidea 1). Only Borostomias of fin. lanternfish Larvae dorsal the of Myctophum the tropical Pacific species, and Neonesthes an eastern fin have the dorsal at midbody and a aurolaternatum capensis, from a subtropical are cosmopolite, reported confused with are and gut trailing astrones- sometimes California 1968; Lavenberg and (Fitch waters Current however, thid larvae; Hubbs et the in rare are larvae Astronesthid 1979). al. stalked has aurolaternatum M CalCOFI only series, occurring in time three samples astronesthids. a feature not found in known larval eyes, and mistakenly Rass Pertseva-Ostroumova (1973) 1951-1984, during off all California. Baja the an astronesthid larva phosichthyid as identified lchthyococcus. Astronesthids are small to moderate in size (up to moderately compressed, elongate, and cm) have 40 scaleless bodies. some black; is Pigmentation BL) species The three larvae (5.9-8.1 mm from CalCOFI is mouth The sides. silvery have surveys with often large, appear to be the same species, probably in the fang genus teeth. at, is origin fin dorsal The slightly like mm or Astronesthes. The 8.1 specimen (Figure of the anal fin posterior well midbody, to, in advance 1) is beginning notochord flexion. It is Astronesthidae moderately (BD depth all except origin. A present dorsal adipose in fin is slender BL). 11 % The trailing 64% ca. at body the from BL is deflected ventrad gut species some adipose a ventral and Radinesthes have at is length 70% BL. Head and broken off a preanal anterior fin prominent fin. anal are There the to two and small (EL 18% is HL) eye The BL. is 22% length photo- ventral photophore rows and numerous small elliptical. BL and the 62% at originates fin dorsal The may phores the head and body. The chin barbel over 75% at fin anal patch A BL. is of melanophores times several or head be shorter than the the head finfold. and Another hindgut adjacent over present the length a terminal is present bulb in some species. and the and the in finfold fin dorsal at present is patch Some species migrate from mesopelagic depths to mm 8.1 smallest however, mm), specimen (5.9 the surface waters at night and others apparently are specimen faded discernible is not patch a dorsal is and 1986c). benthopelagic (Gibbs 1 964a, 1984, in pterygiophores 12 myomeres, 58 ca. has It it. on the base longer fin base, and an anal fin dorsal slightly Presumably, Astronesthids are oviparous but plank- than dorsal. the not are Larvae described. been highly have eggs tonic diverse and have a great number of morphological those identified as Larvae to similar Borostomias complete specializations; no developmental however, panamensis by and (1973) Rass Pertseva-Ostroumova transformation specimens, has been de- series, with A EASTROPAC II (station 46.079). are available from are literature in the and scribed tentative identifications mm 13.7 Astronesthidae (Figure larva postflexion early larvae (Kawaguchi and Moser 1984). The described 1) (incomplete) (BD is slender with BL) 10% a trailing com- laterally 1) be grouped into two morphs: can BL) gut 79% length (preanal ventrad is deflected that pressed slightly body with elongate, sometimes trailing from at body the ca. and is elliptical eye The BL. 58% trailing with cross-section in rounded body 2) and gut at originates fin dorsal The HL). 19% (EL small 60% gut deflected from the body posterior or anterior to and BL the anal fin consists 77% BL. Pigmentation at Some the latter type have the species of midbody. midbrain of paired melanophores ventrolateral to the cartilaginous pedestals on fins anal have and and dorsal of melanophores dorsolateral and the hindbrain, a pair is one In finfolds. median large trailing gut the species with leaflike appendages and on ornamented the alternating melanophores lateral 3-4 in another nape, the at patterns range Pigmentation keel-like. is finfold preanal each side of the body, and melanophores on the trailing 305

318 gut. available mm) specimen advanced most The (17.2 2 ventrally tail the on hindgut; and trunk the between has rays 53 fin 13 anal forming and dorsal 11 and of the ventral the the margin anal a few on fin; above ray complements are The myomeres. incomplete fin the anal ca. midbody; and a few distally on finfold at those than lower are counts the and adult from B. and lower upper the of bases fin at rays and the Stomioidea study this in examined panamensis (Table 15.2 The rays. fin 9 principal has larva mm caudal B. 1). Identification 25 anal rays (complement not is tentative. panamensis as fin ca. rays, dorsal 55 myomeres. Dorsal fin ray length and complete), the increases posteriorly with the penultimate ray Pacific We no larvae of eastern Neonesthes have posi- (20% BL). The meristic counts and fm longest over the however, two astronesthids captured capensis; tions N adult of those match larva Laboratory of capensis NMFS Hawaii this Hancock Seamount on 8501 mm (Table Stomioidea 1). Recently, Evseenko and TC Suntsov appear to be that species. A 15.2 Cruise larva northeast of 25 a described (1995) from mm is Astronesthidae (Figure specimen postflexion 1) slender (BD 8% BL) with pigmentation elongate non- Hawaii that and morphology in is similar a moderately gut trailing BL). 81% (Sn-A lacks but larvae, our to tentatively identi- They eyes. 33% (EL large is eye The capensis. as N fied their specimen area and surround- HL) slightly ovoid. The corneal and bulged and eye the sclera ing outward a within are astronesthid Larval of in found descriptions species membrane funnel extends pigmented lightly shaped, above CalCOFI the comments are region limited to the dorsal fin The pupil. the of from outward the margin with illustrations is of posterior (Table specimens single to slightly well BL), (56% midbody origin 1). 2; Figure Astronesthidae Identification Stomioidea BL. 76% at origin fin anal the from separated The limited remain of astronesthid larvae will until severely anterior BL). (47% midbody to is pelvic bud fin are specimens transformation and larval complete series Pigmentation fore- is as follows: a melanophore on the were obtained from Gibbs data described. Meristic to ventrolateral a pair region; interorbital the in brain 1984, 1986c), Kawaguchi and (1984), Moser (1964a, of dorsal median melanophores, the a series hindbrain; and Ecological study. this during made counts from fin 6-7 anterior and ca. 5 posterior to (those the dorsal 1984, (1964a, Gibbs from obtained was information are fin dorsal the a single anterior melano- larger); to Lavenberg (1968). and Fitch and 1986c) of base; fin dorsal the 2 edge anterior the on phore 306

319 8.1 mm mm 13.7 mm 15.2 Astronesthidae I. Astronesthid larvae: Astronesthes sp., flexion Figure 8.1 mm (CaICOFI 6207, station 117.90; since the specimen stage, specimen; this in occur it would of a 5.9 mm larva [CaICOFI 6201, station 133.35] is shown where pigmentation faded, is dorsal the original 46.79; station II, Borostomias panamensis, postflexion stage, 13.7 mm (EASTROPAC Arthur); by illustration original N. Neonesthes capensis, postfiexion stage, 15.2 original with dorsal view of head (TC 8501, station 59; mm, illustration by N. Arthur); B. Sumida MacCall). illustration by 307

320 MELANOSTOMIIDAE: dragonfishes Scaleless H. MOSER G. Me1anostomiidae, contains family, stomioid largest the waters surface near to migrate sometimes that predators 16 (Morrow Gibbs and about and 200 species genera species may some at night; be benthopelagic (Morrow Gibbs are genera Eight 1985). Fink 1986d; 1984, 1964; Gibbs 1986d). 1984, Gibbs 1964; and found (Berry in the California Current and region Batho- Perkins 1966; Hubbs et al. 1979; pers. obs.). Melanostomiids oviparous with pelagic eggs. are philus represented B. species, three by is B. jlemingi, pus macro whose pelagic Tactostoma species only the is from ranges Bathophilusjlemingi brevis. B. and filijer, spherical been have eggs they described; are and large the to Current California the throughout Gyre Alaskan oil mm) with segmented yolk, a single (1.38-1.55 perivitelline large and globule and space (Kawaguchi Bathophilus filijer, an eastern central Baja California. tropical of be ranges north T. southern to macro pus may Pacific species, Baja Eggs Moser 1984, 1993). Bathophilus of Pacific Sardinops broadly in distributed is brevis sardine, California. those with confused Atlantic and Pacific is and subtropical and tropical the sagax, and similar in size in overlap they since are taken occasionally in the western part of the California larger is the however, characteristics; globule other oil Current region.· The monotypic vs. ca. 0.16- macropus T. in 0.30-0.40 (ca. pus macro Tactostoma mm is waters subarctic the transitional to the mm) of and endemic in than smaller are segments yolk and 0.18 in polygonal somewhat (vs. spherical are and sardine Baja the north Pacific; it ranges from northern Califor- of sardine). have larvae are elongate and Melanostomiid nia Gulf in the California Current to the northward a straight, muscular a pronounced with gut elongate Alaska monotypic The Japan. to northern westward and Flagellostomias boureei is widely distributed in the in terminus. is non-trailing except gut The Eustomias. tropical and subtropical waters The cross body to round laterally is compressed in of the world. Oposto- section; head the and the jaws and relatively are large temperate ranges mitsuii mias north Pacific. across the and small eyes elliptical. size larval Maximum are is Specimens Photonectes, Eustomias, of Leptostomias, mm typically 20-30 mm but some species exceed 30 and Melanostomias are collected rarely in midwater Tactostoma). (e.g., (e.g., mm 40 or Flagellostomias) CalCOFI these of representatives region; the in trawls usual The pattern consists of pigment one or more CalCOFI region from the eastern range the genera into of myomere each on melanophores the dorsal surface and central water mass. from Pacific the tropical myosepta. one more on or the hypaxial and of these genera should be Eustomias Specific identifications differs having in melanostomiids other from large considered taxonomy oftheir the since tentative eastern melanophores, dorsal isolated differs in Bathophilus Pacific resolved. been not has representatives of melanophores having on the ventral surface each 1939; myomere and (Beebe and Crane Kawaguchi are Melanostomiids fishes moderate-size to small 1984). Moser cm) with a variety of 40 shapes ranging body near (to compressed deep-bodied to slender from laterally and study early significant most The of melanostomiid and and scales lacks skin The section. cross in rounded black or dark in coloration, sometimes with silver or is (1939) Crane and Beebe was larvae by described who of rows iridescence. There are ventral two metallic of Leptostomias species: four larvae and illustrated photophores side numerous and prominent each on Photonectes Melanostomias bartonbeani, gladiator, the and head photophores small on dorsal The body. parvimanus, Flagellostomias boureei. They illustrated anal and fins are The caudal. the to close fins pelvic transformation but Eustomias and Bathophilus of larvae at are midbody or somewhat posterior to midbody; in to available not identifica- were series specific permit the Bathophilus they are elevated to side of the body. could be observed tion. larval pigmentation The genus. one but all in lacking is The adipose fin An of juveniles transformed newly of skin the through of length barbel ranges from small to several times the and Odontostomias micropogon Grammatostomias not may mayor and body ornamen- the terminal have thus providing a means of identifying jlagellibarba, are Melanostomiids tation. bathypelagic and meso- their larvae, although they were not available to Beebe 308

321 and stations Kawaguchi and Moser (1984) at the western margin of the survey area; most (1939). Crane specimens poor not usable of melanostomiid of the in are condition and summarized the characters larval developmental a for 18.6 An series. flexion mm larvae genera and illustrated and described of Batho- specimen sp. (Figure Melanostomiidae 1; Eustomias of mitsuii Opostomias brevis, philus and Tactostoma type larval the of typical is 2) Stomioidea Table this of macropus. Ozawa and Ozawa Aono and (1986) is CalCOFI extremely samples. The larva genus from (1988a) Melanostomias tentaculatus, Echiostoma added a rounded cross section. The slender BL) (BD with 5% Photonectes margarita, barbatum, P. albipennis, and gut diverg- and diameter, in small relatively is straight, Eustomias bifilis to the list of melanostomiid larvae long is The es the trunk at 77% BL. from head ventrad the from known Pacific. North compressed dorsoventral- (HL 23% BL) and markedly relatively small (EL 15% HL). and is oval eye The ly. Larvae of T. and jlemingi, B. filifer, Bathophilus protrude not does but pedunculate is eye The beyond collections macro pus are common enough in CalCOFI of is origin fin dorsal The head. the the lateral margin to permit descriptions detailed Identify- of larval series. 77% at origin fin the and anal BL. at ca. 84% BL Fin of larvae of other melanostomiids ing characteristics development ray and myomere are incomplete, howev- or larval whose adult stages occur rarely in the Cal- fin it appears the dorsal count will be in the mid- er region are below. briefly summarized COFI will be >40, and there will fin count anal the twenties, (maxi- myomeres. The dorsal finfold is large >60 be larva Bathophilus of preflexion mm 6.6 damaged A the the depth) twice ventral and depth trunk ca. mum brevis CalCOFI samples (CalCOFI was identified in Pigmentation finfold is smalL consists of a symphysial 80.200). specimen 6407, station postflexion A 15.7 mm blotch and a series spaced of seven, large, evenly the from 1984) Moser and Pacific western (Kawaguchi nape from the midline dorsal along the melanophores in is 9.3 A 1. shown Melanostomiidae Figure mm of specimens other the Most the fin dorsal origin. to midflexion specimen Stomioidea 2) from the (Table however and six had one spots, dorsal seven had survey area of the CalCOFI is west central water mass another The ten. had species) a different (probably of B. brevis larvae. It is deep-bodied representative smallest had noto- the at pigment apposing specimens BL) (BD gut and has an elongate voluminous 17% the larger present not larvae. in is tip but this chord terminal sigmoid a with BL) 94% (Sn-A section, postflexion most of larvae late the of characteristic not of Flagellostomias found boureei were Larvae 26% (HL Bathophilus species. The head is large other 36.4 mm A samples. CalCOFI in postflexion larva large (EL eyes BL) with slightly ovoid, moderately (Kawaguchi water central the from 1984) Moser and brevis Bathophilus HL). among unique is 18% representative of the survey area is CalCOFI mass west count stomioids in having an extremely low vertebral Melanostomii- the late larvae of this species (Figure of Other (33-35) and only 5+6 principal caudal fin rays. larva Table Stomioidea 2). The is relatively 1; dae 9.3 rays. caudal principal 10+9 have stomiiforms The (BD 17% BL) with an elongate straight deep bodied 34 larva has fin myomeres, 5+6 principal caudal mm diameter (Sn-A 85% BL) of moderate diverges that gut incomplete anal rays fin complements. and dorsal and to ventrad the trunk anterior from the anal The fin. just ventral and of paired dorsal consists pigmentation Body large (HL and is a head depth 17% BL) with head of melanophores, one melanophore on the dorsal series eye is dorsal profile; steep the jaws The large. are and surface ventral of each and fine speck- myomere, (EL small and narrow, elliptical, The HL). 12% dorsal on fins, anal and and the finfold, dorsal the ling far posteriad (dorsal fin origin 89% median fins are Head gut. the of and along regions ventral lateral the just are fin origin anal 82% BL). Pelvic fins BL, the pigmentation consists of a paired series ventral to (origin to BL). Meristics midbody for posterior 59% jaw, along melanophores serial isthmus, lower the 67. 7, P 26, A DIS, myomeres this were: larva 2 and mid- the over me1anophores paired hindbrain, or equal finfolds anal and dorsal both of Heights embedded ventrolateral to the hindbrain, melanophores melanophore large is one There depth. trunk exceed and gular, snout, the on speckling fine and opercular three per myomere along the dorsum and one to regions. Other pigment smaller ones in the hypaxial myosepta. the on melanophores scattered and dorsal is scanty: in Eustomias is represented CalCOFI standard on gut and posterior the below finfold the anal fins eight survey collections by nine specimens from 309

322 region; along the lower jaw, on minute melanophores (Sn-A does not and posteriad tapers that BL) 93% lateral to branchiostegal nares on the membrane, the (HL is large head The trunk. the from ventrad diverge above snout, the opercu- posterior the on and brain, the 21 % profile dorsal sloping a moderately with BL) and regions. lar and gular moderate and elliptical is eye The jaws. large with in dorsal is (EL 16% HL). The size fin origin (83% BL) Larvae Cal- not in found of Melanostomias were anal fin slightly forward of the origin (85% BL). COF! (Kawa- larva postflexion mm 16.0 A samples. midbody Dorsal finfold height at is equal to trunk western the from 1984) Moser and guchi is Pacific is ventral and depth less than slightly height finfold representative 2; ofthe genus (Figure Melanostomiidae depth. melanophore per large one is There trunk the dorsum and one myomere along Table or % 11 (BD slender is larva The 2). two minute Stomioidea with gut straight elongate BL) an BL) 93% (Sn-A of Epaxial ones in the hypaxial hypaxial myosepta. and relatively (HL small moderate diameter. The head is myosepta base below the dorsal fin have up to five snout a short with BL) 17% moderate and melanophores forming an size. jaw incomplete bar. Head pig- is eye The moderate HL). 22% (EL size in and oval ment consists of a melanophores of patch -shaped Y The is fin pelvic bud the above and melano- hindbrain and mid- minute BL). 66% (origin posteriad wen dorsal The not are complements fin fully and anal posterior gill ores ph on the arches. pigmentation Other ca. fonned there but are in the 18 and dorsal the in 15 anal the is dorsal and rays, fin bases and proximal on base. fin anal Dorsal myomeres. 56 ca. are There and The on the finfold below the terminal gut region. height is slightly less finfold trunk depth and the than and anal fin ray myomeres; has larva the 64 dorsal complements the and incomplete are series pigment dorsal The small. ventral finfold is are is pelvic fin bud not visible. discontinuous one pair with of dorsal melanophores on each of the anteriormost eight myomeres and one on dorsal fin the origin. to each myomere posterior of most the at found are sp. Photonectes Larvae small to Hypaxial myosepta have one three melano- seaward region of A pattern. sampling CalCOFI the phores. consists melanophores of: pigmentation Other 15.9 mm postflexion (CaICOFI station 6507, larva at the brain; the to and above ventrolateral along nares; the in is found representative of 90.120) larval form jaw, the lower gular and region, the below isthmus; more A the California Current region. advanced on liver; along the median finfold margins; scattered from in the same area postflexion illustrated larva is the dorsal and and anal fins on the finfold below the Melanostomiidae2. Notochord flexion occurs at Figure posterior gut region; and along melanophores minute ca. 10-15 larvae and mm 30 at metamorphose ca. mm the jaw, branchiostegal membrane, lateral to the lower SL. 15.9 mm larva is slender (BD 10% BL) with The nares, scattered and This fin. caudal on the larva an gut straight elongate % 91 (Sn-A moderate of BL) appears to be diameter. The is relatively small (HL head on based tentaculatus M BL) characters 17% by presented Aono Ozawa (1986) examina- on and and moderate HL) The jaws. and with 37% (SnL snout eye tion series of a transformation of that species from the The HL). 19% (EL smaH relatively and narrow is obs.). Atlantic Larvae (pers. of Melanostomias tral1s- fin pelvic is well bud posteriad (origin 67% BL). less slightly trunk depth than Dorsal finfold is height size, fonn Another mm. <20 typically a small at type and the pigment, is small. ventral dorsal and anal finfold The dorsal continuous has larva Melanostomias of fin are complements larvae however but of type have not been identified. not larva this the in fonned fully this has ca. specimen postflexion largest 16 19 and dorsal Adult reported Melanostomias valdiviae have been Pigmentation anal rays. There are ca. 60 myomeres. the from 1966) Perkins and (Berry region CalCOFI but consists series meIanophores. minute of The dorsal larvae are unknown. their consists of clumps of several small melanophores have myosepta hypaxial myomere; each several above only The in mitsuii Opostomias larval identified minute melanophores. In the largest larvae, melano- specimen flexion late mm 15.0 a is samples CalCOFI myomere per 7 ca. phores increase dorsal the in to from CalCOFI (Table Stomioidea 6204, station 60.140 series to and body series. 7-10 in the hypaxial Other 15.0 a to similar is larva This 2). illustrated larva mm pigmentation consists liver the below patch a linear of by Kawaguchi and Moser (1984) (Figure Melano- of section terminal the above line a gut. the and The 2). stomiidae deep- moderately is larva CalCOFI throughout is distributed Pigment ventral and dorsal the 15% (BD bodied straight elongate an with BL) gut 310

323 finfolds line a concentrated with body, the near except this but larva in the largest postflexion specimen anal, rays dorsal, at caudal The margin. finfold the and dorsal 15 are There rays. anal 20 and available ca. has pectoral in become base late pigmented and the fin paired of consists ca. myomeres. Body 79 pigmentation the above found brain, is Head pigment postflexion. series both and dorsal myosepta! epaxial and hypaxial gular jaw, region, region, opercular lower the on snout, one is large melanophore per myomere pigment. There of branchio- the the isthmus, the covering at and edge by in the dorsal series sometimes or one accompanied stegal single membrane. A P. Photonectes species, anteriormost more The minute melanophores. 10-12 margarita, is for the region (Berry and reported in than the larger are series dorsal the melanophores recent 1966; Hubbs et aL 1979); however, Perkins no are smaller In ones. posterior more there larvae work may species be one than more that suggests epaxial consists series hypaxial the and melanophores P. California Current region and marga- present in the in melanophore a single of the In myoseptum. each Scripps not may occur there (Cynthia Klepadlo, rita in melanophores 16.1 three larva are one there mm to Institution of comm.). pers. Oceanography, the larvae in and series hypaxial and epaxial the larger are lined with melanophores. completely myosepta melano- Leptostomias were not collected within the larvae scattered of consists pigmentation Other body liver the below phores a regular terminal the above CalCOFI line sampling area; however, they have and section stations on been captured the dorsal seaward most of the of west just gut. posterior The the region of finfolds flexion and melanophores minute have occurs ventral and CalCOFI stations. Although notochord mm) of the anal fin. The (10-15 size Lepto- small comparatively in region basal at a the on some are there head above the brain, on the opercular is pigmented from this reach may larvae postflexion region, stomias gill lower jaw, gular isthmus, and region, region, snout, 40 SL before metamorphosis. A 16.1 mm post- mm arches. Leptostomias larvae from the the region west of 2; Figure flexion larva Melano- (Table Stomioidea is 7210, station 31.135, sampling the CalCOFI CalCOFI until stomiidae 2), from identified be cannot area from of in the north Pacific Leptostomias The representative of Leptostomias species the region. taxonomy resolved. is is moderately deep-bodied anteriorly (BD 12% larva gradually posteriad to the caudal region. but tapers BL) litera- BL) 94% (Sn-A straight and and descriptions The foHowing are based on the elongate The is gut of specimens and of observations detailed on ture 22 a narrow to tapering anteriorly, diameter a large has large (HL Bathophilusfilifer, 22 30 and B.jlemingi, of specimens is relatively diameter posteriorly. The head and 20% BL) with a moderate snout (SnL 34% HL) specimens of Stomioidea (Table macropus Tactostoma 14% (EL is narrow and relatively small The jaws. eye literature data Meristic 2). the from obtained were posteriad (Morrow Gibbs 1964; Kawaguchi and 1984; 68% (origin and wen Moser is bud fin pelvic The HL). from aL and 1989) and et narrow. are finfolds ventral The Dorsal BL). Fujii 1984b; and Matarese observations study. this during made fonned not are complements fin anal and dorsal fully 311

324 15.7 mm mm 18.6 36.4 mm Melanostomiidae 1. Larval melanostomiids: Bathophilus brevis, postflexion stage, 15.7 mm (Kawaguchi and Moser 1984); Figure 90.140; original 'illustration by N. Arthur); Flagellostomias boureei, station Eustomias sp., flexion stage, 18.6 mm (CaICOFI 9210, postflexion stage, 36.4 mm (Kawaguchi and Moser 1984). 312

325 16.0 mm mm 15.0 mm 22.2 ... 16.1 mm Melanostomiidae 2. Larval melanostomiids: Melanostomias Figure postflexion stage, 16.0 mm; Opostomias mitsuii, postflexion stage, sp., 15.0 mm; Photonectes sp., postflexion stage, 22.2 mm (Kawaguchi and Moser 1984); Leptostomias sp., postflexion stage, 16.1 mm by N. Arthur). (CaICOFI 7210, station 31.135; original illustration 313

326 MELANOSTOMIIDAE Bathophilus filifer DESCRIPTION EARLY LIFE MERISTICS HISTORY Mode EGGS Range Shell diam.: Yolk: Vertebrae: 45-50 48 No.ofOG: Diam.ofOG: Total surface: 45 Shell 43---45 Precaudal Pigment: 3-5 4 Caudal features: Diagnostic Fins: 0 0 spines Dorsal 16 13-16 LARVAE rays Dorsal 0 Hatching 3 mm length: spines Anal 0 Flexion length: 8-12 mm 14-17 16 rays Anal Transformation length: ca. 25 mm 4--5 5 Pelvic 1 A, & D & C , C development Fin P"~ I P, sequence: , Pectoral z ventral series; postanal median ventral Pigmentation: Yolk-sac-Paired Caudal: series on to tail; scattered on head; fairly heavy of dorsal & ventral 10+9 10+9 tip Principal dorsal Prejlexion-Paired margins. at concentrated finfolds, series ProcllIrrent: 4 where from nape to near notochord tip the approximately series join; 4 Upper & ventral series; I per myomere in 1 to to ventrolateral several dorsal 3 3 Lower ventral on several 1 to hindbrain; below & above base; P, of region Gil! rakers: terminal gut series becomes section. Ventral Flexion-posiflexion- Upper & ventral series; dorsal by hypaxial in fewer obscured musculature; Lower specimens; some in isthmus on series hindbrain; over a patch faint on Brancilliostegals specimens. in some finfolds of features: voluminous Diagnostic gut gut; section terminal Elongate, HISTORY LIFE angle, slight at deflected in as flexure sigmoid acute late no deflected B. jlemingi; head large, ventrad; liver small, postflexion the Gulf Range: of & central tropical Eastern Pacific nortliward to to (length 5-6% BL), recessed anterior is gut; paired dorsal & ventral area the of part southernmost the & survey CalCOFI California pigment series; minute, pigment faint. melanophores Mesopelagic; possible vertical migration Habitat: %) MORPHOMETRICS (range & mean ill season: & during fall captured winter Spawning on Larvae mostly Y-S f PrF PoF Tr Juv surveys CalCOFI Sn-NBL ELM pattern: Oviparous; 73-80 eggs & larvae 90-93 planktonic 91-100 80--89 81-83 78 86 92 95 82 LITERATURE 7-9 10--12 BDIBL 10-12 11-12 12-13 11 II 8 11 12 Rass & Pertseva-Ostroumova 1973 14-19 20--25 21-22 17-22 HUBL 17-17 21 20 23 16 17 (Illustrator) ILLUSTRATIONS ORIGINAL 37-38 41-54 38-50 HW/HL 33---42 44---44 42 47 37 38 44 larva, Yolk-sac (N. mm Arthur) 3.0 11-14 SnLIHL 37--41 25--41 34--35 23-29 Arthur) Preflexion larva, 4.8 mm (N. 12 34 35 38 26 Arthur) (N. mm larva, Flexion 11.1 15-18x 15-23x 13-15x 38--46x ED/HL* 19-20x larva, Arthur) (N. 22.3 mm Postflexion 18-34 28-33 16-19 16--18 19-20 18x24 42x31 15xl7 16x17 20x20 is given first, vertical axis second. Eye * axis is horizontal oval; somewhat 0-1 2-6 4---4 2-3 P,LIBL t t all specimens. in broken rays I P juvenile 0.3 5 4 3 0-0 0-0 0--0 0.5-2 14-17 PzLlBL 0 0 0 1 16 5-6 5-5 LvLIBL 5-6 6 6 5 314

327 Jilifer Bathophilus Threadfin dragonfish ... . - : .. . . : ... : . : .. : ., .. -.-. . . . .. '- ~. . . "' --~ .--. . . .. -. . . '~'.~----- '. . . ... ".', ",', .. . . . .. . ~ ... , ----- ... 3.0mm . . '. ~. ... .. . . - '.' . ... 4.8mm mm 11.1 mm 22.3 Figure Melanostomiidae 3. Yolk-sac larva, 3.0 mm (CaICOFI 7210, station 157G.40); preflexion larva, 4.8 mm (CaICOFI 7202, station 153.30); larva, flexion 11.1 mm (CaICOFI 7205, station 150.110); postflexion larva, 22.3 mm (EASTROPAC II, station 46.84). 315

328 MELANOSTOMIIDAE flemingi Bathophilus MERISTICS EARLY LIFE HISTORY DESCRIPTION Mode Range EGGS Vertebrae: diam.: Shell Yolk: Total 47 44-48 No.ofOG: Diam.ofOG: Precaudal 42-45 43 surface: Shell Caudal 4 3-4 Pigment: Fins: features: Diagnostic Dorsal spines 0 0 Dorsal rays 14-16 15 LARVAE spines 0 Anal 0 ca. Hatching 3 mm length: 15-17 rays Anal 16 12-15 mm length: Flexion Pelvic 15 15-19 25-30 mm Transformation length: ca. 4-8 Pectoral 6 Fin development sequence: C, , , P A, C P, & 0 & 2 2 Caudal: Yolk-sac--On dorsal & ventral over scattered finfolds; Pigmentation: 10+9 Principal 10+9 Pre 1 per ca. series, paired ventral myomere; & flexion-Dorsal head. Procurrent: notochord at series at single embedded snout; on patch tip; median Upper 5-6 5 otic region median liver; below hindbrain; & ventrolateral series to Lower 2-3 3 per 1-3 by isthmus; on series paired with series dorsal mm, 9 rakers: Gill hindbrain; above by myomere; gular patch; patch scattered mm, 11 Upper over & & A branchiostegal associated 0 on margin of finfolds; Lower Flexion-Concentrated proximally on finfold, ventral membrane. Branchiostegals heaviest liver; throughout dorsal finfold; along lower jaw; below series recessed; above ventral & below terminal gut section. LIFE HISTORY ventral & per myomere in dorsal Postflexion-Many series; patch & concentrated at at midbrain; covers opercIe, on region; cIeithral nares. Alaska Range: to central Baja California; offshore to western part Gulf of terminal Similar to B. filifer deflected section but features: Diagnostic Current of California at greater angle present flexure sigmoid & larger liver larvae; late in bathypelagic to Meso- Habitat: than in (length B. filifer typically 6-8% BL) & less recessed; than B. pigmentation & heavier filifer. in melanophores larger abundance in Spawning larval Maximum season: CalCOFI collections in mean & MORPHOMETRICS in %) (range & summer spring & eggs larvae Oviparous; pattern: ELH planktonic Y-S PrF F PoF Tr Juv LITERATURE Sn-AIBL 79-91 88-93 90-94 82-83 85 76 90 93 83 Kawaguchi & Moser 1984 BDIBL 8-12 12-13 12-14 13-13 1989 al. et Matarese 10 8 12 13 13 HLlBL 21-23 20-22 19-22 17-18 (Illustrator) ORIGINAL ILLUSTRATIONS 16 21 22 20 17 HWIHL 33-41 37-41 34-45 53-63 Yolk-sac larva, 3.7 mm (N. Arthur) 43 39 38 40 58 Arthur) (N. mm 5.6 Preflexion larva, SnLIHL 31-44 35-41 32-41 26-29 13.0 (N. Arthur) mm Flexion larva, 17 35 38 36 27 larva, Postflexion mm 23.5 (N. Arthur) EDIHL* 13-17x 13-14x 12-14x 18-20x 18-23 16-18 14-17 18-20 37x33 15x21 13x17 13x15 19x19 first, given is axis horizontal oval; somewhat axis is Eye * second. vertical P,LlBL 0-0 0.3-5 3-5 3-4 23-39 0 4 3 4 31 P L1BL 0-0 0-0 0-9 0.3-1 2 23-26 0 2 0 0.7 25 LvLIBL 0-0 5-8 8-8 6-9 0 6 8 8 316

329 flemingi Batlwphilus Highfin dragonfish. mm 3.7 5.6mm - . . . . . . ~ ° ' ."" •••• •••• :; ' 0 . ' .. '", mm 13.0 mm 23.6 Figure Melanostomiidae 4. Yolk-sac larva, 3.7 mm (FRONTS 85, station 18); preflexion larva, 5.6 mm (CaICOFI 7507, station 103.80); postflexion flexion 13.0 mm (CaICOFI 7804, station 90.200); larva, larva, 23.6 mm (CaICOFI 6207, station 100.60). 317

330 MELANOSTOMIIDAE Tactostoma macro pus EARLY LIFE HISTORY DESCRIPTION MERISTICS EGGS Mode Range 1.38-1.55 mm Shell diam.: Vertebrae: Yolk: Segmented; 0.78-0.89 mm 81 79-82 Total diam. 67 No.ofOG: 1 67-68 Diam. 0.30-0.40 mm Precaudal OG: of Smooth surface: 14 Shell Caudal 12-14 Pigment: Fins: 0 0 Moderate space, perivitelline large Diagnostic diameter, shell reatures: spines Dorsal OG large 16 14-18 rays Dorsal 0 Anal 0 spines LARVAE 19 19-22 Anal rays Hatching length: ca. 4 mm 9 Pelvic 8-10 length: Flexion mm 13-24 0 0 Pectoral 50-55 mm ca. length: Transformation Caudal: sequence: development Fin , D A, C 10+9 C" P 10+9 & Principal 2 2 Pigmentation: dorsal & ventrolateral series Yolk-sac-Paired Procurrent: infracaudal & yolk-sac late in series with dense supra- continuous 8 8-9 Upper 4 at 4 50-60 of a maximum flexion-Dorsal series increases to Pre stage. Lower series as 7-9 mm & decreases thereafter; ventrolateral developing 2-3 Gill rakers: each myoseptum. Flexion--On lower on symphysis, melanophores jaw Upper isthmus, cIeithral region, base of Crays, & terminal gut PI base, Lower >25 absent in larvae series Posiflexion--Paired dorsal mm. section. Branchiostegals Diagnostic body, compressed laterally elongate, Extremely features: as count; round in cross section not in Chauliodus; high myomere LIFE HISTORY heavy caudal peduncle pigment; loss ofmelanophores in dorsal paired unpigmented. & series; narrow finfolds Pacific westward to Subarctic Japan & north & transitional Range: Current to in the California southward region California Baja central MORPHOMETRICS & in %) (range mean Mesopelagic migrating into epipelagic at night Habitat: y-S F PrF PoF Tr Juv during CalCOFI surveys are most abundant in season: Spawning Larvae 84-88 Sn-AlBL 80-82 & September 91-96 88-92 in with summer, July peaks 82-84 83-92 91 81 93 86 83 87 & eggs planktonic Oviparous; larvae ELH pattern: 6--8 6--7 BDIBL 5-6 6--7 6--7 5-6 6 7 6 6 6 7 LITERATURE 14-16 12-13 11-15 13-18 HLlBL 10-10 10-11 16 13 15 10 13 10 Amaoka et al. 1992 43-54 HWIHL 35-39 33-50 47-50 35-39 30-36 Kawaguchi 1993 Moser & 1984, 41 37 49 38 32 48 1989 al. et Matarese 27-34 20-25 11-19 27-36 27-33 SnLIHL 19-21 15 31 33 30 23 20 30-32x 14-25x 13-15x EDIHL* 16--24x 19-24x 12-15x somewhat is Eye * axis first, given is axis vertical horizontal oval; second. 20-30 17-19 14-17 23-30 19-24 19-24 14x18 18x23 31x27 13x16 20x21 21x21 2-4 0-1 2-3 2-2 0-0 PILlBL 0-2 0.4 3 2 2 0.5 0 P 0-0.8 L1BL 0-0 0-0 0-0.6 9-12 2-10 2 0 0 0.3 0.3 11 6 318

331 macropus Tactostoma Longfin dragonfish mm 4.9 mm 1.4 13.5 mm 34.2 mm --~----~/ ~------------- mm 44.0 . . . . . ~ . . . . . . . . . 49.4 mm postfiexion Figure 5. Egg, 1.4 mm; late yolk-sac larva, 4.9 mm; flexion larva, 13.5 mm; Melanostomiidae larvae, 34.2 mm, 44.0 mm; transformation specimen, 49.4 mm (Kawaguchi and Moser 1993). 319

332

333 MALACOSTEIDAE: Loosejaws MOSER H. G. about three Malacosteidae 15 genera and includes Presumably, malacosteids are oviparous; planktonic species in species; scintillans is the only Aristostomias unknown. eggs are Larvae are slender and elongate the genera and other although area, survey CalCOFI a head large compressed and with dorsoventrally species central the and water equatorial in found are in that may be several times longer trailing gut slender to west water mass the area. Larvae of of the CalCOFI the than body species 1981; in (Moser some the in n.mi.) (beyond offshore occur scintillans A. 200 small relatively Kawaguchi and Moser 1984). Eyes are region central area, primarily of the CalCOFI survey and Several types ovoid. of larval pigmentation are found between central in larvae have paired Baja Typically, and California Conception, Point malacosteids. melanophore ventral and dorsal been far have as as Adults California. collected north the on none with series Bering hypaxial paired ventral series become myosepta. the The Sea of margin eastern the to westward and developing musculature and are difficult the recessed in the central water mass. gut The see. to be may head the and pigmented is pigmented in various places (e.g., brain, snout, jaws, cm) «25 moderate-size to small are Malacosteids predators isthmus). an with fonn unidentified One extremely the meso- of and bathypelagic zones with (Kawaguchi lacks long gut pigment except on the gut extremely large jaws that extend well posterior the to and Moser 1984). fanglike. relatively Teeth are large and skull. small Eustomias larvae have a trailing gut is no floor There and a body shape similar to that but malacosteids jaw of in the mouth and specialized dorsal the the eyes are nearly round, the is flatter, head musculature permits an extraordinarily large jaw consists pigmentation The is and elongate relatively body com- opening. melanophores (5-10) fewer of are anal posteriad leaving and Dorsal far pressed. fins located lacks gut trailing the and midline, the in melanophores. a short caudal peduncle. The skin is black, smooth, and prominent scaleless. rows ventrolateral There two are evenly grouped photophores, consisting of of spaced or description following The scintillans is based A. of the stud small Numerous organs. light photophores (Kawaguchi on literature the and on 1984) Moser and present; are light Postorbital head. and body organs detailed examination and 3 of juveniles larvae 21 present are organs and preorbital suborbital or absent. ecological informa- (Table Stomioidea 2). Meristic and species; in A hyoid barbel is present in some A. 1964c; tion were obtained from the literature (Morrow half the length of the body scintillans it is more than and Fitch and Lavenberg 1968; Hart 1973; Kawaguchi and has a bulb-like terminus (Morrow 1964c; Gibbs Moser 1984; Goodyear and Gibbs 1986; Matarese et aL Fink 1985; Goodyear and Gibbs 1986). 1984; 1989) and from observations made during this study. 321

334 MALACOSTEIDAE Aristostomias scintillans LIFE HISTORY DESCRIPTION EARLY MERISTICS Mode EGGS Range Shell diam.: Yolk: Vertebrae: No.ofOG: 57 Diam.ofOG: 54-58 Total surface: Shell 50 50-53 Precaudal Pigment: 5-7 7 Caudal Diagnostic features: Fins: 0 0 Dorsal spines 21-25 23 LARVAE Dorsal rays m Hatching length: <6 0 0 Anal spines Flexion length: mm 13-17 28 25-29 rays Anal 45 ca. length: Transformation mm 6-7 6-7 Pelvic , , P Fin development sequence: C , D & A, PI C 4-8 4-8 Pectoral I 2 2 flexion-Short, Pre just a.pposing median series Pigmentation: dense, Caudal: dorsal first paired tip; posteriorly, notochord 10+9 10+9 to anterior form series Principal gradually then ventral paired anteriorly; augmented series are just Procurrent: to in smallest larvae, augmented anteriorly; posteriorly fonn 11 beginning 9-11 Upper region, a 5-6 5-6 pair posteriorly in the & otic pairs ventrolaterally Lower hindbrain, dorsolateral to dorsal series & a pair on snout; by 8.0 mm, rakers: Gill Flexion-- of about & consist pairs. 11-14 nape to complete Upper pairs between ones added smaller & series Additional in (large) dorsal Lower ones; ventral series extends forward larger head, recessed in trunk to Branchiostegals on 3-4 myomeres apart; series musculature above gut, spaced ca. lower on just & fonning isthmus, on gular region, above brain, patch HISTORY LIFE trailing gut, jaw; on proximally on Crays. Posiflexion--More added more & ventral Series, on continuous; eventually each series dorsal to central California Baja to region Current California to Sea Bering Range: gut, specimens. some in pattern ring-like vague fonning margin to water mass west & of central Diagnostic features: Elongate slender body & gut; trailing gut away head; long flattened elongate, origin; A at body from deflected Epi- large diurnal some apparently Habitat: mesopelagic; to migration; ovoid snout, dorsal eyes; pigment Eustomias & ventral paired series; in larvae captured neuston & ventral series lack larvae pigment. gut in CalCOFI surveys with peak Spawning season: Larvae taken year-round & (range MORPHOMETRICS mean in %) abundance in April-May Y-S PrF eggs pattern: larvae Oviparous; ELH F PoF planktonic & Tr Juv Sn-AlBL 115-144 LITERATURE 76-78 100 >100 126 77 Moser Kawaguchi 1984 & 7-8 6-7 6-8 BDIBL 13-19 7 7 6 16 Matarese 1989 et al. 20-22 16-19 HUBL 18-23 18-25 ORIGINAL (Illustrator) ILLUSTRATIONS 17 20 21 22 38-46 HWIHL 32-35 30-39 37-49 mm 9.0 (N. larva, Preflexion Arthur) 43 34 34 44 (N. 16.2 larva, Flexion Arthur) mm 27-50 SnLIHL 38-45 38-43 22-29 39 41 40 26 11-17x 9-12x 9-11 x 23-28x EDIHL* axis vertical first, is given horizontal axis oval; somewhat is second. * Eye 13-15 17-25 11-13 23-28 14x21 10x14 10xl2 26x26 3-4 3-3 2-3 PILlBL 6-31 4 3 2 19 P 0.1-2 L1BL 0-0 7-16 0-0 2 0 0 0.9 12 322

335 Shining Aristostomias scintillans loosejaw 9.0mm 34.7 mm Figure Malacosteidae L Preflexion larva, 9.0 mm (CaICOFI 6204, station 100.120); flexion larva, 16.2 mm (CaICOFI 6301, station 90.110); postflexion larva, 34.7 mm (Kawaguchi and Moser 1984). 323

336 ~ •

337 IDIACANTIDDAE: Blackdragons MOSER G. H. species of the genus The of about four family consists 1934; Gibbs 1964b, 1984). antrostomus, in A single species, 1. occurs Idiacanthus. from northern the California Current California south- eggs pelagic oviparous; are idiacanthids Presumably the to ward Pacific. eastern eastern The tropical unknown. are elon- and slender extremely are Larvae Pacific species tropical probably 1. is panamensis with gate gut that a slender the deflected is from body (Gibbs antrostomus 1. 1964b; Novi- with conspecific and in at the and trailing is flexion origin fin anal Current, California the of 1. 1967). Westward kova postflexion larvae. The head is flattened and the antrostomus the central water mass by replaced is stalks eyes elliptical are borne on long supported by a about equal is Dorsal finfold height cartilaginous rod. species 1. fasciola. is small. The body to finfold the and depth ventral well forming on the body; dorsal fin posterior begins exhibit Idiacanthids marked dimorphism with sexual fin and anteriorly added are rays length base is ca. species some in length cm 40 ca. females growing to at the end BL of the larval period. Rays are added 10% and males elongate extremely are Females with <7 cm. rapidly enlarges to fin the when metamorphosis during black, fanglike skin and large with teeth. jaws scaleless the length. At metamorphosis two-thirds the body The dorsal the with long-based are fins anal and dorsal re- cartilaginous supporting the rods stalks are eye of about to fin caudal the forward extending from just are larval sorbed and the pectoral fins sparse lost. The anal the length; body the of fin extends two-thirds melanophores on of a series of consists pigmentation the of the one-third fin about caudal to forward from isthmus, a series the hypaxial on and myomeres, the pair A length. body specialized spiny projections is of 1934; trailing the on spots scattered Novi- gut (Beebe present and to each dorsal anal ray. Pectoral anterior Kawaguchi and 1981; Moser and Weihs 1967; kova of prominent are at metamorphosis. fms lost rows Two al. 1989). Moser 1984; Matarese et photophores side; smaller are ventrally on present each head and in the distributed are photophores over and (1934) Beebe described (1967) Novikova and the photo- postorbital small A body. on rows vertical in larval development illustrated and metamorphosis present chin barbel has a leaflike the and is phore (1984) and Kawaguchi fasciola. summarized 1. Moser Males lack teeth, barbel terminus. a chin fins, paired development larval of antrostomus a illustrated and 1. large relatively have and postorbital a large and eyes is description following The larva. postflexion late The photophore. gut is developed and the poorly based observa- detailed on and literature published on extend are large and gonads the entire length the of tions supported body cavity. The vent has a fleshy structure, of Stomio- (Table antrostomus 1. of specimens 24 function the by modified first anal ray, that may in data Meristic 2). et Matarese from idea obtained were this al. and from counts made during (1989) study predators midwater are Females transmission. sperm was information Ecological 1). epi- extending with a broad bathymetric Stomioidea (Table range to from from and 1984), (1964b, Gibbs (1934), Beebe obtained depths and apparently are vertical migra- bathypelagic (1968). Fitch and Lavenberg bathypelagic are (Beebe tors. Males shortlived and 325

338 IDIACANTHIDAE Idiacanthus antrostomus HISTORY LIFE EARLY MERISTICS DESCRIPTION EGGS Mode Range diam.: Vertebrae: Shell Yolk: 81-85 No.ofOG: 81-85 Total Diam.ofOG: Shell surface: Precandal Pigment: Candal Diagnostic features: Fins: 0 0 spines Dorsal LARVAE 58 54-66 rays Dorsal Hatching length: 0 mm 0 <5 spines Anal 28-43 Flexion ca. length: 20-32 mm 35 Anal rays Transformation length: 67-76 6 mm 6 Pelvic , C D, 0 P A, 0 & Fin development sequence: C Pectoral I 2 2 8 embedded ca. of a series mm, 6.0 ca. flexion-At Pre Pigmentation: Candal: 10+9 10+9 Principal isthmus; at ca. dashes along mm, 9.0 a series of oblique dashes begins Procurrent: of each hypaxial myomere. Flexion- to fonn at margin posterior 12-13 11-14 Upper of dashes three groups, range to 5-13 fonn tend pos!flexion-Isthmus 7-10 dashes hypaxial 8); (usually Lower 8-9 expanded; when myosepta into spread rakers: scattered trailing gut. on Gill & elongate; gut elongate Body features: Diagnostic extremely slender Upper & flat; head postflexion; & Lower flexion in trailing slender, eyes elliptical (up on elongate to rods cartilaginous stalks by supported BL); 27% PI Branchiostegals , at lost diminutive males lack P poorly transfonnation; have 2 gland. luminous postorbital & developed large gonads, large gut, eyes HISTORY LIFE MORPHOMETRICS (range & mean in %) of margin to westward Pacific, eastern tropical to Temperate Range: northwestern central mass; also in temperate Pacific water PrF Y-S PoF F Juv* Tr migrate diurnally Habitat: Epi- to bathypelagic; females Sn-AlBL 80-95 129-139 68--69 93 134 89 69 season: In Spawning abundant surveys, CalCOFl larvae most during summer peaks in with August & November fall & 4-6 BDIBL 3-4 2-3 5-5 5 3 5 3 extreme larvae; & ELH sexual Oviparous; planktonic eggs pattern: 13-18 HUBL 13-15 10-15 9-10 dimorphism, with larvoid males 14 15 12 9 41-50 HWIHL 39-45 36-40 41-48 LITERATURE 46 41 38 45 42-50 SnUHL 17-44 44-53 23-25 Beebe 1934 35 49 46 24 Kawaguchi & Moser 1984 10-27x 11-12x EDIHLt 10013x x 20-21 1989 al. et Matarese 19-40 18-21 17-21 20-21 1967 Novikova 17x27 llx20 11x19 20x20 ILLUSTRATIONS ORIGINAL (Illustrator) 2-2 1-4 1-2 PILlBL 0-0 2 3 1 0 larva, mm (N. Arthur) 5.1 Preflexion L1BL P 0-0 0-0 0-0.2 6-10 2 mm 21.8 (N. Arthur) Flexion larva, 0 0.1 0 7 11-24 ESLlBL 12-21 19-23 0-0 20 20 16 0 Females. * t Eye axis is oval; horizontal second. is given first, vertical axis 326

339 Idiacanthus antrostomus Pacific blackdragon 5.1 mm 21.8 mm 55.0 mm Figure Idiacanthidae 1. Preflexion larva, 5.1 mm (CalCOFI 6110, station 80.120); flexion larva, 21.8 mm (CaICOFI 5010, station 70.130); postflexion larva, 55.0 mm (Kawaguchi and Moser 1984). 327

340 AULOPIFORMES WATSON W. The aulopiform fishes originally were apportioned body longitudinal the near are fins Pectoral fin. adipose and Myctophoidei suborders between the myctophiform commonly but are in axis ventral the nearer aulopoids, Okiyama 1984b). 1966; al. Alepisauroidei (Gosline et an that margin and set at oblique angle suggests a Rosen Mycto- to Myctophiformes the restricted (1973) There the in function paravane-like are alepisauroids. phidae a in others all placed and Neoscopelidae, and tissues no photophores and only other luminous occur on basis Aulopiformes, the new order, of several evermannellids. and scopelarchids some in of subsequently which most synapomorphies, putative and were questioned and/or rejected (e.g., Fink aulopiform The fishes apparently oviparous; are all 1982; structur- the However, 1992). Johnson Weitzman Okiyama 1984a). (e.g., hermaphroditism is common (Rosen arches gill upper the of al synapomorphies and a few only for species, known are eggs Spawned Johnson 1973; 1992) commonly are accepted as a basis others. a few for described been have ova maturing for recognition (some of a monophyletic Aulopiformes 1.0-1.5 ca. from range typically eggs Planktonic mm, to with none several oil globules, and with usually differ authors e.g., view, this from Okiyama 1984b; sculpturing on the chorion. Maturing ova suggest that The component families are variously 1986). Stiassny the range be may of size egg near the end lower volume this in classifications; in arranged different we which in classification (1990) Eschmeyer's follow the 1976; Okiyama 1984a). 0.5 (e.g., mm Bertelsen et al. for most unknown are hatched Newly aulopi- larvae Alepi- contains two order suborders, Aulopoidei and a differ- of an sauroidei example for 1994 Nelson (see that those unpigment- forms; known typically are have ent aulopiform classification). All four aulopoid a small to formed a partially eyes, ed mouth, and families Scopelarchi- Chlorophthalmidae, (Aulopidae, yolk sac. are moderate moderately very to Larvae eight the of alepisauroid seven and Notosudidae) dae, compressed, preanal slightly to elongate, with strongly families (Synodontidae, Giganturidae, Paralepididae, one-third to more than from length ranging less than Anotopteridae, Alepisauridae, Evermannellidae, and most of Omo- length. All aulopoids body two-thirds from recorded been have sudidae) California the pectoral and head the on spines lack alepisaruoids Current vicinity, and larvae of all families except (present in alepisaurids). girdle and paralepidids some Omosudidae Giganturidae, and Chlorophthalmidae, Larval pigmentation ranges from nearly absent to surveys. have been collected during CalCOFI complete, light. is relatively commonly most but nearly melanophore peritoneal Distinctive patches or bands The demersal, aulopoids to mesopelagic from range both of several families in the larvae characterize 1984a,b). Okiyama (e.g., suborders alepisauroids are primarily meso- and bathy- the while coastal (synodontids are residents). benthic pelagic Most are hover-and-strike or ambush predators. All are the The following chapters include only eight families collected been have larvae whose aulopiform with more or less elongate and compressed, usually and (tubular large eyes scopelarchids, giganturids, in in the Okiyama See area. study CalCOFI (1984a, large some usually mouths, with and evermannellids) chlorophthalmids, Johnson of 1988d) illustrations for indica, Gigantura Rofen and and Bertelsen (1991) for teeth. The armed only segment- with large fins contain dorsal also most have a small rays and ed species (1966b) for Omosudis lowei. Alepisauroidei Aulopoidei Suborders and families included: Aulopidae Synodontidae Paralepididae Scopelarchidae Anotopteridae Notosudidae Evermannellidae Alepisauridae 328

341 AULOPIDAE: Aulopids AMBROSE A. D. Complete of About 11 2 genera (Aulopus and Hime) early life history series including eggs species in known for only are Hime benthic (Okiyama japonica marine these 1974, tropical the in found are fishes subtropical waters and 1988b). 1984a, the Atlantic and mm), (1.18-1.34 of spherical are Pacific Eggs pelagic, without an (Parin transparent, Kotlyar oil globule, and with and 1989). AU three Atlantic species irregularly raised hexagonal meshes the on chorion are all Pacific bajacali, Aulopus and species, except A. Larvae surface. Chao japonica H of a Lee Hime. are as (1994) recognize and Hime are bajacali A. and elongate (BD a with BL), 12-20% (HL head large generic for synonym Only Aulopus. bajacali Aulopus and BL. 52-64% ca. length a preanal BL), 20-32% Kotlyar and (Parin area CalCOFI the in occurs 1984). the to anterior far is The anus Both fin anal origin. larvae tip southern the near collected been have Most species have which patch, pigment large single a the and Pacific both of from the Gulf Baja California expands larval development, dorsolaterally during on eastern the in of California; others have been taken the patch laterally on the taiL another and peritoneum, tropical adults Pacific near Cocos Island. The benthic Development is gradual direct, and with little addition- at 82-230 m depth, where they may be occur primarily throughout pigmentation al stage. H In pelagic the abundant. flexion notochord japonica ray deveiopment fin and aulopids Adult length, in moderate and slender are rela- are patches pigment the size, at occur a smaller the tively and smaller, number total origin of myomeres is in are fin dorsal The cm. 60 usually less than ca. than fewer in A. bajacali (41-43 vs. 47-49). the anterior third the and of a wen developed body is located over fin adipose the posterior end the of anal on The fonowing species detailed account is based (Mead base 1966). fin are and thoracic Pelvic fins examination mm); (3.8-7.5 of larvae preflexion 8 3 The and pupil are round and pectorals are lateraL eye (7.5-8.4 larvae larvae postflexion 6 mm); flexion directed fulcral laterally. and supramaxiUae Two (41.6-56.5 5 juveniles and mm); (8.8-12.3 on the caudal peduncle are characteristic. .. of mm) A scales A are: Meristic counts for the family D 14-22, Meristic 8-14, and Parin from obtained were data bajacali. this during made counts from and (1984) Kotlyar study 9, V 10-17 BrR and 36-53, (Okiyama PI P 11-14, 2 on Johnson (1992) and 1984a; Nelson 1994). from mm) SIO Marine (108-151 the Patterson 16 specimens Vertebrates Collection. and Johnson (1995) suggest that the aulopids may be information ecological Adult the group sister was based on Synodontidae and the of notes. SIO field collection Pseudotricho- notidae. 329

342 AULOPIDAE Au/opus bajaca/i MERISTICS LIFE HISTORY DESCRIPTION EARLY Range Mode EGGS Vertebrae: Shell diam.: Yolk: Total 47-49 48 No.ofOG: Diam.ofOG: 31-34 31 Precaudal Shell surface: 15-17 Caudal 17 Pigment: Diagnostic features: Fins: 0 0 Dorsal spines 14 rays 14-15 Dorsal LARVAE 0 0 spines Anal Hatching mm <3.8 length: 12-13 Anal rays 12 Flexion length: ca. 7.5-8.5 mm Pelvic 9 9 >13 mm Transformation mm, <41 length: 13-14 Pectoral 13 A, C, sequence: development Fin PI' P D, 2 Caudal: dorsolaterally Preflexion-j1exion-Patch Pigmentation: peritoneum; on Principal 10+9 10+9 Postflexion-Patches enlarging; in lateral patch region. on A tail Procurrent: ventrally region. melanophore occasionally present on hypural 8-10 Upper 8-10 brain; over jaws; lower & upper snout; Juvenile-On band vertical 8-10 8-10 Lower PI; opercle; proximally on behind usually patches proximally eye; on Gill rakers: ca. ventrally; pale dorsum, rays; 5 D first 9 on distally & covering Upper 2-4 3 narrowly more blotches irregularly-shaped extend also that dorsally 8-11 10 Lower on proximally ventrally. extensive more C, line; lateral below Branchiostegals 14 14 Diagnostic features: 47-48 single 13-14; PI 14-15; D total vertebrae; in tail on laterally patch & patch saddled-shaped large peritoneum on LIFE HISTORY A region. in MORPHOMETRICS (range & mean %) to Range: Baja Alijos, N) (ca. 24°50' Island, Rocas Cocos California specimens of tip southern near collected Baja most W; 83°26' N, 5°34' California Sur Y-S PrF F PoF Tr Juv bottom soft on Demersal Habitat: m depth at 82-230 Sn-AlBL 52-57 55-56 59-62 61-64 54 56 60 62 Larvae Spawning in collected season: in January-March, CaiCOFI tows BDIBL 13-15 12-14 13-16 15-18 June, peak a November-December with 14 14 13 16 20-22 HLlBL 20-22 24-26 29-32 larvae planktonic pattern: ELH Oviparous; 21 21 25 31 HWIHL 56-84 70-79 76-80 41-43 LITERATURE 77 69 73 42 12-24 SnLIHL 23-28 21-36 23-27 1974, 1984a Okiyama 18 26 28 25 32-40 EDIHL 33-35 29-32 ILLUSTRATIONS (D1ustrator) ORIGINAL 22-29 35 34 30 26 5-9 larva, Preflexion 3.8 C. (R. mm 4-5 Walker) PILlBL 6-7 15-17 7 5 7 16 P L1BL 0-0 0-0 0-1 23-26 2 0 0 0.2 25 330

343 bajacali Aulopus Aulopus 3.8 mm 12.3 mm Figure Aulopidae 1. Preflexion larva, 3.8 mm (CaICOFI 6611, station 147.20); postflexion larva, 12.3 mm (Okiyama 1974). 331

344 SCOPELARCHIDAE: Pearleyes M. E. AND WATSON w. SANDKNOP Scopelarchidae four in species 18 contains genera range from deep- species (Johnson 1984a). Larvae Okiyama and Johnson 1986); five (Johnson 1984a; head a large with robust and moderately bodied to species representing all genera, four occur the in elongate and slender small head. a moderately with Scopelarchidae California Current (Table vicinity 1). The mouth snout wedge- and long is is the large, well beyond the limits of the All five species range the and shaped, eye is usually with its longer narrow, Benthal- CalCOFI study the within area; study area, forward. axis inclined "vertical" obliquely Preanal bella dentata widely distributed others while length is about one-third to one-half of body length. is the The from gut southward range southern California or Baja somewhat at tube "S"-shaped or a straight is California (Johnson 1974). Larvae five all of species deep abdominal of the moderately to very bottom the in occur collections, most CalCOFI ichthyoplankton is musculature hypaxial the from separated cavity and frequently abundance highest in and summer during a more mem- perivisceral translucent broad less or by at autumn offshore and stations. this membrane are the brane. Clearly visible within patches melanophore or saddles large characteristic the over in except gut, Benthalbella which unpig- is scopelarchids Adult small to medium-size (ca. are to this In mented, or nearly so, as a larva. addition bathypelagic predators 10-30 cm) meso- and in all seas (Johnson except 1974, Arctic and Mediterranean the larval pigment, Scopelarchoides and Rosenblattichthys to They elongate, are moderately moderately 1984a). melanophores may few a have and dorsal the on strongly to half about length preanal with compressed, B. on the tail. Larval ventral margins, laterally and dorsal short-based The length. body of fin, two-thirds Rosenblattichthys dentata melanophores and have may of only is composed anal like which long-based the fin fin; fin rays develop preco- pectoral pectoral on the adipose inserts ahead of midbody. An segmented rays, in Rosenblattichthys. ciously of is located posteriorly, above about the last third fin well and is with The fin. anal the large armed mouth particu- Larval scopelarchids are easily recognized, as numer- more smaller, as well long lanceolate teeth large mouth head large to moderate their with by larly directed are eyes tubular The teeth. conical ous, eyes, narrow and space translucent wide usually a characteristic and forward, obliquely or upward have the between and gut the and musculature, hypaxial the of tissue, the pearl organ, from which the patch pearly the characteristic pigment patches over gut. They are name the family derives. At common least one for be in to larvae fish other any with confused unlikely and bioluminescent is infans, B. Scopelar- species, Larger CalCOFI larvae scopelarchid study area. the choides kreffti be as bioluminescent well may (Johnson a combination of of be can use by species to identified 1974). meristic, pigment characters, but and morphometric, larvae cannot always be identified below the smallest are Scopelarchids synchronous hermaphrodites of genus. level the 1974). at development larval and Eggs (Johnson undescribed. Recently larvae hatching are hatched literature on The based are descriptions following or (Scopelarchus scopelarchids as tentatively identified of 1984a) and on detailed observation (Johnson 1974, unpigmented, long, mm 3.6-4.0 are Scopelarchoides) 25-33 of each species (Table specimens Scopelarchidae from Meristic 2). obtained were data ecological and with oval unpigmented eyes, partially open to formed preanal (ca. length short mouth, small 30% BL), and Johnson (1974); additional counts were made meristic study. this during are one all for known but larvae Older sac. yolk 332

345 Table Scopelarchidae Meristic characters for the scopelarchid species in the California Current vicinity. 1. have All 9 pelvic fin rays, 10+9 caudal fin rays, and 8 branchiostegal rays. principal Vertebrae Fin rays Prev CV Total D A C, PI Species 17-21 6-8 54-55 31-35 20-23 21-25 13-16+13-16 dentata Benthalbella 21-24 17-18 32-33 49-51 9-10 13-14 23-26 14+ volucris Rosenblattichthys 13-14+15-17 20-23 45-48 6-7 20-23 26-30 16-20 nicholsi Scopelarchoides 18-22 21-26 29-30 7-9 12-15+13-15 44-49 17-18 anaUs Scopelarchus 24-29 7-8 46-51 18-21 13-16+ 13-16 29-33 17-18 guentheri S. Number the (above) of specimens scopelarchid and size ranges (in mm, below) used in preparation of Scopelarchidae2. Table species descriptions. Transformation Postflexion Flexion Preflexion Yolk-sac Juvenile Species 2 10 5 6 1 2 dentata Benthalbella 51.9-56.5 4.2-12.7 13.3-19.3 30.8-49.5 4.1 4.0, 67.3 4 10 9 2 6 0 volucris Rosenblattichthys 8.9-12.7 14.2-39.2 45.8-57.7 49.0-58.1 3.4-8.6 4 10 5 4 10 0 Scopelarchoides nicholsi 9.0-12.4 12.0-18.1 4.0-9.3 27.2-44.4 54.4-69.8 2 6 5 5 8 0 anaUs Scopelarchus 14.2-23.7 9.8-12.6 27.6-31.5 4.6-7.3 33.7-47.2 3 8 5 3 6 0 guentheri S. 9.2-12.5 15.8-31.2 33.7-41.7 55.6-81.5 3.6-8.7 333

346 SCOPELARCHIDAE Benthalbella dentata MERISTICS EARLY LIFE HISTORY DESCRIPTION EGGS Range Mode diam.: Shell Vertebrae: Yolk: No.ofOG: Diam.ofOG: 54-55 Total 54-55 Shell surface: 20-23 Precaudal 31-35 Pigment: Caudal Diagnostic features: Fins: 0 spines Dorsal o LARVAE 6-8 rays Dorsal 7 ca. Hatching length: 4 mm Anal spines o 0 17-21 i8-20 length: ca. 13-19 Flexion mm rays Anal 57 Transformation 9 Pelvic 50-55 mm through ca. ca. mm 9 length: 21-25 23-24 development sequence: Fin C PI' P C A, & D Pectoral , b 2 2 to many Yolk-sac Pigmentation: few through postflexion-None except Caudal: 10+9 scattered on 10+9 membrane. PI on Traniformation-LittIe blade fin or Principal occiput & snout at beginning of stage, increasing & Procurrent: over spreading after ca. jaw 53 laterally mm; 13-16 scattered lower eyes; around & snout on Upper 13-16 15-16 trunk on ca. 52 & peritoneum on mm; tail by posteriorly beginning Lower dorsally beginning just caudad PI of level behind & & spreading rakers: Gill o Upper 0 stage. of to ventrad enclose gut end by P Diagnostic 13-18+37-42=54-55; Myomeres features: ofD ahead o Lower 0 2 8 PI before on 8 except unpigmented origin; transformation. Branchiostegals mean & MORPHOMETRICS (range in %) HISTORY LIFE Y-S PrF Mexico Guadalupe, Tr F PoF of Gulf Isla (57°46'N) Alaska de to Juv Range: W 150°101' of east primarily N), (29°12.1-13.8' Sn-AlBL 37-40 57-61 40-52 41-48 46-50 45 38 46 48 59 63 primarily 500-1000 m depth Mesope\agic, Habitat: BDIBL 11-18 17-20 10-10 12-13 14-19 larvae collected October-July with Spawning In CalCOFl season: area, 15 10 18 16 13 II maximum March-April during 17-18 18-26 22-23 HURL 22-24 19-22 23 23 23 18 21 22 synchronous Oviparous, are larvae hermaphrodites; ELH pattern: 42-49 35-62 49-60 HWIHL 33-36 planktonic 51 55 29 45 35 35 23-43 SnUHL 38-45 39-44 36-36 LITERATURE 35 42 36 41 33 * 5-28 x I 13-16x 12-18x 20-20x EDlHLt 1982b Belyanina 26-46 25-28 23-25 24-33 Johnson 1974, 1984a 19x34 14x28 14x26 20x24 1988c Ozawa * * 4-8 3-8 15-16 4-5 PILlBL ILLUSTRATIONS (Illustrator) ORIGINAL 5 4 16 6 II * 1-14 0-0 0-0 0-1 14-18 PzLIBL MacCall) Preflexion (B. mm Sumida 9.2 larvae, mm, 6.5 0 0 0.1 16 8 12 Flexion 18.5 larva, mm (B. Sumida MacCall) Sumida MacCall) mm (B. Postflexion larva, 42.5 damaged, Both yolk-sac specimens ED SoL, * & PIL could not be measured; juvenile specimen damaged, eyes missing. t Eye oval; "horizontal" axis second. is given first, "vertical" axis 334

347 Northern pearleye dentata Benthalbella mm 6.5 9.2mm 18.5 mm 42.5 mm .. .. : ... .t • • 53.0 mm Figure Scopelarchidae 1. Preflexion larvae, 6.5 mm (CaICOFI 6902, station 67.60), 9.2 mm (CaICOFI 7803, station 97.60); flexion larva, mm (CFRD 7804, station 80.70); postflexion larva, 42.5 (CaICOFI 8812, station 27.9.26.2); transformation specimen, 53.0 mm mm 18.5 (Johnson 1974). 335

348 SCOPELARCHIDAE Rosenblattichthys volucris DESCRIPTION EARLY LIFE MERISTICS HISTORY EGGS Range Mode Shell diam.: Yolk: Vertebrae: No.orOG: 49-51 50 Diam.ofOG: Total 18 17-18 surface: Shell Precaudal Pigment: 32 32-33 Caudal Diagnostic features: Fins: 0 0 Dorsal spines 9-10 LARVAE 9 rays Dorsal length: <3.2 mm Hatching 0 0 spines Anal length: Flexion ca. 9-14 23 mm 21-24 rays Anal mm 28-60 ca. length: Transformation 9 9 Pelvic 24-25 23-26 Fin sequence: development D C" Pectoral P, PI' A, & C lo few to Preflexion--None Pigmentation: PI of surface inner on distally Caudal: 10+9 10+9 base above patch peritoneal large blade; PI anterior scattered & on Principal 60% of gut. Flexion-On ventral to between myomeres half margin Pl"ocurrent: 14 mm; 9.5 by 34-38 14 by Upper 34-38 myomeres between margin dorsal on 13-14 & myomeres 22-24 by between 12.7 mm; 1 midlaterally at mm 12 Lower bases 45-47 by 2 1 or of 12.1 Posiflexion-Near mm. myomere rakers: Gill 0 mm; 14.7 by rays C bases middle lower near 2 upper middle 1 of 0 or Upper I rays by bases at mm; 14.S mm. 22.8 C by 3 & 2 rays A of 0 0 Lower I 8 mid- & dorsolaterally at nape; Transformation-Over & on 8 hindbrain; BrallclJiostegais margin dorsal on 16-38; ca. at myomeres myomere ca. at 38-50; HISTORY internally on first 7 LIFE pterygiophores between 31-32 mm; peritoneal A & ventrally. posteriorly patch spreads features: Large head with Diagnostic snout long & large mouth; deep vicinity California from southern Current California to central Rallge: body; myomeres stage, 12-18+33-36 during preflexion 23-27+25-27 N) 33-25° (ca. Sur (ca. 6° Pacific eastern equatorial & Baja California single 50-51); (usually 47-52 total large postfiexion by stage, to 3° S) N pigment peritoneal dorsal each 2 blotches to on up patch; & ventral on peduncle, 2 on C. 1 blotch caudal margins; laterally Mesopelagic Habitat: & (range MORPHOMETRICS %) ill mean season: Spawning year the throughout collected larvae area, CalCOFI In August-January during average 011 catches largest with PrF F PoF Y-S Tr Iuv ELH are pattern: Oviparous, hermaphrodites; larvae synchronous 37-52 55-62 Sn-AlBL 62-64 59-64 62-62 planktonic 44 62 57 63 62 LITERATURE 12-21 21-28 17-27 16-18 17-17 BD/BL 16 24 24 17 17 31-34 30-39 20-34 HL/BL 27-31 31-32 Belyanina 1982b 24 32 29 35 32 Johnson 1984a 1974, 47-77 43-53 HWIHL 27-35 28-32 30-44 47 54 31 39 30 ILLUSTRA nONS (Illustrator) ORIGINAL 24-48 33-34 44-57 46-55 SnLIHL 31-31 50 37 50 33 31 Preflexion Orr) M. (H. mm 7.0 larva, 11-35x 7-12x 6-12x 17-24x EDIHL* IS-20x mm (H. 13.0 M. Flexion Orr) larva, 15-21 19-24 23-48 20-24 17-23 Transformation (H. specimen, 37.0 mm Orr) M. 19x33 9x21 IOx18 19x21 19x20 9-16 5-16 11-22 21-23 22-23 PILIBL axis second. * oval; "horizontal" axis is Eye given first, "vertical" 17 12 8 22 22 0-0 1-14 P,LIBL 0-0 15-19 19-20 0 0 19 7 18 336

349 Ch.ubby peadeye volucris Rosenblattichthys ~c-,-~~-?f~ .~ 7.0mm 13.0 mm mm 14.5 mm 26.0 37.0 mm --------,--- 38.9 mm Figure Scopelarchidae 2. Preflexion larva, 7.0 mm (CalCOFI 6607, station 107.65); flexion larva, 13.0 mm (CaICOFI 6706, station 107.50); 37.0 postflexion 14.5 mm, 26.0 mm (Johnson J984a); transformation specimens, larvae, mm (CFRD DSJ-50, station 11), 38.9 mm (Johnson 1974). 337

350 SCOPELARCHIDAE Scopelal'choides nicholsi MERISTICS EARLY LIFE HISTORY DESCRIPTION Range Mode EGGS Vertebrae: diam.: Shell Yolk: 46-47 45-48 Total No.ofOG: Diam.ofOG: 17 16-20 Precaudal Shell surface: 26-30 28-30 Caudal Pigment: Fins: Diagnostic features: 0 spines Dorsal 0 6-7 rays Dorsal 7 LARVAE Anal 0 spines 0 mm length: <4.0 Hatching 21-22 20-23 rays Anal Flexion through mm 12-12.4 ca. mm lellgth: 9-9.3 Pelvic 9 9 Transformation length: ca. mm 26-28 ca. through mm 40-45 20-23 21-22 Pectoral sequence: developmellt Fin C , , C A, P2' D, PI I 2 Caudal: over peritoneal Prejlexion-posiflexion-Large Pigmentation: patch 10+9 10+9 Principal upper part margin of 0-2 (usually level on ventral PI; of of at gut 1) Procurrent: O-few stage); flexion during form (buds buds P2 of level at gut 13-14 13 Upper on ofmyomeres in (usually 1) vicinity ventral margin of tail 34--39; 15 15-17 Lower of 1 laterally on each side over middle hypurals by 8-9 mm. Gill rakers: forms Transformation-Series on laterally articular & dentary Upper 0 0 posteriorly beginning & at anteriorly; dorsally mm 26 ca. spreading 0 Lower 0 around eye 32-36 mm; dorso- between & on & and snout over brain Branchiostegals 8 8 tail on ventrolaterally beginning ca. at peduncle caudal on 30-32 mm & spreading cephalad; pair of peritoneal 2 of level at P patches added HISTORY LIFE 26-27 mm. at Diagnostic features: Myomeres 45-48 (usually 47-48), 12-13+34--36 stage, preflexion in in 18-19+29-30 single stage; postflexion Rauge: Cedros, N) 27° (ca. California to Vicinity of Isla equatorial Baja pigment peritoneal pair smaller then mm, 26 ca. until anteriorly patch Panama west off & W) N, off Peru to & Colombia (5°_9° 136° waters P of single melanophores usually present patches added at level of 2; (ca. 6°_12° S) over middle hindgut tail, lateraliy & of margins ventral on posteriorly of hypurals. occurs depth range at Habitat: Epi- & mesopelagic; of in shallower part the day during deeper night, & (range in %) MORPHOMETRICS mean season: Spawning larvae year; throughout the occur to reported Larvae y-S in June collected February, in CalCOFI November area & f PrF PoF Tr Juv larvae are pattern: ELH synchronous Oviparous, hennaphrodites; Sn-AlBL 45-50 36-47 47-52 52-53 50-54 41 47 49 planktonic 53 53 12-16 BD/BL 15-20 18-21 15-20 15-17 LITERATURE 15 18 20 17 16 18-25 HLIBL 22-28 21-26 25-29 25-27 1982b Belyanina 22 25 23 27 26 1974, Johnson 1984a HW/HL 56-64 54-67 52-74 41-54 38-43 58 61 59 47 40 (Illustrator) ILLUSTRATIONS ORIGINAL SnLIHL 24-41 35-49 35-44 34--39 27-34 30 43 40 36 30 (H. M. Postflexion 16.8 larva, Orr) mm EDIHL* 12-18x 10-14x 11-17x 13-18x 18-20x M. 33.4 Transformation Orr) specimen, mm (H. 23-36 19-23 20-27 17-23 20-22 14x29 Ilx21 13x23 15x20 19x21 "vertical" first, axis given is axis "horizontal" oval; Eye * second. 3-6 P1LIBL 4--5 4-5 8-16 12-16 4 5 4 12 15 P LIBL 0-0 0-0.1 0-3 2 11-18 16-17 0.Q2 0 1 14 17 338

351 Scopelarchoides nicholsi mm 7.5 16.8 mm 23.0 mm 33.4 mm 47.5 mm Figure Scopelarchidae 3. Preflexion larva, 7.5 mm (Johnson I 984a); postflexion larvae, 16.8 mm (CaICOFI 6611, station 150.60), 23.0 mm (Johnson transformation specimen, 33.4 mm 1984a); (EASTROPAC I, station 13.255); juvenile, 47.5 mm (Johnson 1974). 339

352 SCOPELARCHIDAE Scopelarchus analis DESCRIPTION EARLY LIFE MERISTICS mSTORY EGGS Mode Range Shell diam.: Vertebrae: Yolk: 44-49 46--48 No.ofOG: Total Diam.ofOG: 17-18 17-18 Shell Precaudal surface: 29-30 Caudal 29 Pigment: Fins: Diagnostic features: Dorsal spiues 0 0 7-9 rays Dorsal 8 LARVAE <4.8 mm Hatching length: Aual spines 0 0 Flexion 10-13 ca. mm 22-25 21-26 length: Anal rays Pelvic 9 9 33-55 mm through length: ca. 25 mm Transformation 20 18-22 PI' & P C C" sequence: development Fin Pectoral D, A, 2 2 Pigmentation: Preflexion-posiflexion--None except choroid tissue Caudal: on 10+9 10+9 below before ca. 9 mm; 1 peritoneal patch above anterior middle eye Principal by ofP, level to forward shifting mm, Procurrent: by 10 of pair mm; 15 ca. gut 13-14 12-15 patches above hindgut dorsolateral between P of peritoneal midway Upper 2 13-15 14-15 anus & Lower & Transformation--Dorsolateral at ca. 20 mm. ventrolateral Gill rakers: spread 25 mm, by tail of to lateral midline adjacent fonn stripes Upper 0 0 cephalad of across bar as connected become caudad, end caudal & on snout & jaws over midbrain by 25 by mm; ca. 28 mm; peduncle 0 0 Lower on P, after 25 mm; peritoneal sections 8 8 Branchiostegals 33-55 30 mm. fuse by mm; by stage, preflexion early 9+36-37 Myomeres features: Diagnostic in LIFEmSTORY HL 46--47); (usually 45-49 postflexion total stage, by 19-22+27-30 ~O% no 22-23); (usually ~5 rays A 20-23%); (usually BL pigment through peritoneal patch anterior one then mm, 9.8 ca. before in southern Circumglobal warm water; in CalCOFI Range: area off added mm. of pair dorsolateral mm, 19 patches 20 by posteriorly California peninsula California Baja & (range MORPHOMETRICS & mean in %) Habitat: Mesopelagic y-S F PrF the throughout collected Larvae Spawning season: year PoF Juv Tr synchronous 43-49 ELH pattern: Oviparous, are larvae hennaphrodites; Sn-NBL 36--43 49-56 50-56 57-65 46 38 53 54 61 planktonic 13-14 BDIBL 10-15 16-18 13-16 16-17 14 13 17 15 17 LITERATURE 21-22 20-23 HLlBL 20-23 20-25 25-28 22 22 21 22 26 Belyanina 1982b HWIHL 41-43 44-47 45-63 41-53 41-43 1974 Johnson 41 46 52 47 42 32-38 28-41 SnLIHL 38-46 31-37 33-34 ORIGINAL ILLUSTRATIONS (Illustrator) 37 35 43 34 34 , 7-11 x 11-13x 11-15x EDIHL* 10017x 18x larva, Preflexion 4.9 mm (R. C. Walker) 27-38 23-27 26-30 21-25 22-23 mm Walker) C. (R. 11.7 larva, Flexion 12x33 l2x28 9x24 13x23 18x23 18.8 larva, Postflexion (R. mm Walker) C. P,LlBL 4-5 3-8 3-6 11-16 18-24 5 4 5 13 21 is given first, "vertical" axis second. * Eye oval; "horizontal" axis P L1BL 0-0 0-0 0-2 5-11 11-23 2 0 1 0 17 8 340

353 BlackbeHy pearleye analis Scopelarchus 4.9mm 11.7 mm .:}~f.::~~~,~~~ __ .-----'--"~~ ___ : ... ------ ~ 7-1V-~~ 18.8 mm 21.0 mm 22.0 mm 30.5 mm Figure Scopelarchidae 4. Preflexion larva, 4.9 mm (CaICOFI 6204, station 100.160); flexion larva, 11.7 mm (CaICOFI 6207, station (Johnson 90.140); larvae, 18.8 mm (CaICOFI 5207, station 97.90), 21.0 mm postflexion 1974); transformation specimens, 22.0 mm, 30.5 mm (Johnson 1974). 341

354 SCOPELARCHIDAE Scope/archus guenther; MERISTICS DESCRIPTION HISTORY LIFE EARLY Range EGGS Mode Vertebrae: Shell diam.: Yolk: Total 49 46-51 No.ofOG: Diam.ofOG: 17 17-18 Precaudal surface: Shell 29-33 Pigment: Caudal 33 Fins: features: Diagnostic 0 spines Dorsal 0 8 7-8 rays Dorsal LARVAE length: <3.5 mm Hatching 0 0 Anal spines 25-27 Flexion ca. length: rays mm 9-12 24-29 Anal 9 9 Pelvic mm mm ca. length: Transformation 50-55 through 33 18-21 Fin development sequence: C , PI> P D, A, Pectoral Cz, 20-21 * I z Caudal: Preflexion-flexion-8ingle mid- above patch peritoneal Pigmentation: 10+9 10+9 Principal 10 mm; pair slightly shifts usually 6 mm, ca. by gut ca. by cephalad Procurrent: dorsolateral 8.4 peritoneal of by patches posteriorly mm. 14-15 13-16 mm, Upper Posiflexion-Few externally over hypural area by 15-16 15 13-16 Lower ventrolateral & dorsolateral adjacent cephalad spreading as to stripes by at ca. 25 mm; internally on hindbrain lateral 28 midline beginning Gill rakers: Upper 0 0 mm. over spreading midbrain by 33 mm, Transformation-Externally 0 Lower 0 on mm; 42 ca. by over forebrain membrane; gular & jaw, upper snout, 8 8 Branchiostegals peritoneal sections fuse. mm; D on by 42 Myomeres Diagnostic features: stage, during pretlexion 10-14+33-36 LIFE HISTORY in posttlexion 18-22+26-31 total stage, HI.. Ql % BL (usually 46-50; pigment present midway ~4 (usually 25-27); peritoneal 19-20%); A added posteriorly by gut over by 6 mm, anteriorly dorsolateral pair to Circumglobal Range: primarily in warm water; in CalCOFI off area 8.4 laterally on caudal peduncle after mm; mm. 15-16 southern & Baja California peninsula California & %) in mean MORPHOMETRICS (range mesopelagic & epipeJagic Lower Habitat: Y-S Larvae collected Spawning throughout year the season: F PoF PrF Juv Tr Sn-AlBL pattern: ELH synchronous hermaphrodites; larvae are Oviparous, 37-43 43-51 50-57 53-58 50-53 47 39 52 52 55 planktonic 11-12 11-14 BDIBL 13-16 14-19 16-18 LITERATURE 11 13 16 18 15 17-21 18-21 HLlBL 23-26 20-23 22-27 1982b Belyanina 19 19 21 24 24 1984a 1974, Johnson 48-59 38-46 HWIHL 40-53 33-36 43-49 43 53 49 34 45 ORIGINAL (Illustrator) ILLUSTRATIONS SnLIHL 33-43 38-48 37-48 30-36 25-29 45 36 42 27 33 (H. Pretlexion M. Orr) larva, 6.5 mm 9-14x 12-17x 9-13 x 24-29x 16-17x EDlHLt (H. mm Flexion 11.5 larva, Orr) M. 24-40 26-30 23-26 27-34 23-24 mm, larvae, 18.5 22.5 mm (H. M. Orr) Posttlexion 1Ox24 11x28 15x33 17x23 26x31 mm 33.5 specimen, Transformation M. Orr) (H. 4-6 3-5 4-10 PILlBL 11-15 20-21 4 4 13 20 6 common counts North in the * PI 20-21 eastern most are Pacific; 8-10 0-7 0-0 0-0 PzLIBL 10-11 1974). (Johnson 18-19 has guentheri S. elsewhere PI 0 0 10 9 3 second. t Eye oval; "horizontal" axis is given first, "vertical" axis 342

355 guentheri Scopelarchus Staring peadeye mm 6.5 mm 11.5 mm 18.5 22.5 mm 35.5 mm 48.5 mm Figure Scopelarchidae 5. Preflexion larva, 6.5 mm (CaICOFI 6608, station 117.65); flexion larva, 11.5 mm (CaICOFI 6712, station 117.35); mm 113.50); (CaICOFI 6607, station 107.45), 22.5 mm (CaICOFI 6901, station transformation specimens, 35.5 mm 18.5 larvae, postflexion (CaICOFI 6210, station 130.80), 48.5 mm (Johnson 1974). 343

356 NOTOSUDIDAE: Paperbones SANDKNOP w. WATSON AND E. M. notosudid of the approximately species, 19 Two elongate, slightly notosudids Larval sac. yolk com- are report- been of the have genera, three two representing compressed with a more becoming pressed posteriorly, Ahliesaurus Current vicinity. ed from the California slightly head that is slightly compressed to depressed, brevis, primarily an Indo-Pacific ranges species, growth. depressed with eyes are more becoming The the of CalCOFI seaward edge the near to eastward becoming more rounded during horizontally elongate, while central off California, Baja Scopelo- study area quickly short initially The transformation. snout throughout area CalCOFI the saurus ranges harryi and long a short, The wedge-shaped. and gut becomes is across the North Pacific BL, increas- between N 20°-60° about 40-S0% ca. length (preanal tube straight to ing transformation S0-60% just before or during the (Bertelsen al. 1976). Larval notosudids, apparently et There pectoral stage). no spines on the head or are in commonly relatively occur harryi, S. CalCOFI all and largely limited is light Larval pigmentation girdle. ichthyoplankton primarily seaward samples, at the or finfold trans- peduncle caudal the to before fin and spring during stations and summer. which 'also has a Ahliesaurus formation, except in (ca. notosudids are small to medium-size Adult 1976). midlateral melanophore series (Bertelsen et al. IS-SO cm) fishes that are meso- and or bathypelagic or continental on epibenthic (Bertelsen slopes insular Larval notosudids are unlikely to be confused with et al. 1976; Kreffi 1984). They are primarily predators Primary CalCOFI the in larvae other any area. study on pelagic crustaceans and small fish (Bertelsen al. et characters body with are some- elongate, the slender 1976). Notosudids long and slender, with a slightly are snout, wedge-shaped long head, depressed what cylindrical and head depressed moderately to to eyes, elongate short preanal horizontally relatively compressed trunk tapering to a strongly compressed of band to limited largely pigment and length, a peduncle. The head is long with a long wedge- caudal melanophores Among the the on peduncle. caudal elongate, mouth, snout, large and horizontally shaped be S. to likely collect- is most harryi species, notosudid is Preanal body of S0-60% about length oval eyes. easily is distin- It area. ed within the CalCOFI study length. The fins only segmented rays. The contain by from count myomere higher its A. brevis guished midbody is near fin dorsal short-based the longer- and shorter length (ca. vs. (S7-61 preanal 30-40% 42-S0), dorsal The posterior. far is fm anal based fin adipose vs. BL vs. ca. SO%, lengthening to ca. 4S-S0% ca. The rays. fin anal few last the is above trunk, cheeks, 60% during transformation and early juvenile stages), and scales. are covered tail with large cycloid Teeth by lacking midlateral melanophores that the and in developed older are rakers gill and and larvae well characterize A. flexion the least at from brevis stage resorbed be or adults in are slowly juveniles, but may onward. to ready individuals in absent are both at maturity; lost (Bertelsen 1984, 1986). Kreffi 1976; al. et spawn on based is description following The detailed brown, There are Notosudids tissues. luminous no are yolk-sac, larvae (2 4.3-4.6 29 mm; of 10 examination brown dark head, a darker with usually black gill to preflexion, S.0-12.S mm; 7 flexion, 14.3-20.2 mm; 7 Some dark margins on the and scale pockets. covers, 40.0- postflexion, 22.3-41.S mm; 3 transformation, scales. silvery have species of mm) (37.S-S0.S 4 juveniles and mm) 4S.2 harryi. S. Notosudids are synchronous hermaphrodites (Bert- CalCOFI from larval Since A. brevis were identified no elsen et al. 1976). eggs Planktonic are unknown, but here, but given not is description a separate samples, Bertelsen al. (1976) reported maturing ovarian et that flexion shown are larvae in stage postflexion and eggs brevis with a single oil 0.SS-O.6S are A. of mm 1. (1976) al. et Bertelsen to Refer Figure Notosudidae S. harryi ca. 4.3 mm, are globule. Recently hatched additional and Ozawa (1978) for descriptions and and with mouth, open an eyes, unpigmented a small information. 344

357 8.8 mm 24mm Figure Notosudidae 1. Ahliesaurus brevis: flexion larva, 8.8 mm; postflexion larva, 24 mm (Bertelsen et al. 1976). 345

358 NOTOSUDIDAE Scopelosaurus harry; LIFE HISTORY DESCRIPTION EARLY MERISTICS EGGS Range Mode Shell diam.: Vertebrae: Yolk: 58-61 No.ofOG: Total 59 Diam.ofOG: Shell 26-30 surface: Precaudal Pigment: Caudal 29-32 features: Diagnostic Fins: 0 o Dorsal spines 10 LARVAE 10-12 Dorsal rays o ca. 4 mm Hatching length: 0 spines Anal 16-19 14--20 mm 18 Flexion length: ca. Anal rays 9-10 Pelvic 37.5 9 Transformation length: 40-50 mm, shrinking to ca. mm 12 10-14 , sequence: C Fin , A, C Pectoral PI development P & Ad, D & I 2 2 on Caudal: preflexion---fJ-few Yolk-sac Pigmentation: & margin ventral & 10+9 10+9 Principal middle ventral near fmfold none before of 8-9 mm), (usually tail ca. Procurrent: larval increasing with margin finfold nearer dorsal on located length; 12 12 & ventral margins few extending Upper of last tip, myomeres & notochord 11-12 12 onto dorsal forming finfold, caudal bands Lower & mm; 11 ca. ventral by rakers: Gill notochord tip by laterally 7 mm. ca. on Flexion-posiflexion-Series 0-1 o mm; 14 ca. after ofhypurals distal along margin Upper laterally mm, 33 by 17-20 18 Lower & peduncle caudal embedded on along ventral margin caudal of 10 10 Branchiostegals on few mm; after rays A on few peduncle; 22 with decreasing C rays, growth. in BL 31-43% HISTORY LIFE slender; Diagnostic preanal length Elongate, features: myomeres eyes; oval elongate horizontally mm; <25 ca 57-61 larvae to (9-12+47-50 through early postflexion stage, gradually shifting Pacific North Range: 20°-60° N; throughout CalCOFI study ca. between to late 21-22+39-40 by postflexion stage); pigment largely limited area peduncle. caudal mesopelagic, Juveniles Habitat: primarily continental on epibenthic adults %) in mean & (range MORPHOMETRICS & insular slopes spring in larvae collected & area, CalCOFI In season: Spawning summer PrF Y-S F PoF Tr Juv (March-August) 31-34 Sn-AlBL 32-40 40-43 43-45 41-46 48-51 32 42 36 44 43 50 ELH Oviparous, synchronous hermaphrodites; planktonic larvae pattern: 5-9 8-9 BDIBL 5-6 6-6 7-8 5-6 LITERATURE 9 6 5 6 5 7 14--15 HLlBL 14--18 16-18 16-19 16-19 24--26 1976 et al. Bertelsen 14 17 15 17 25 18 Matarese et al. 1989 32-44 29-49 27-33 HWIHL 23-30 24--35 23-26 Okiyama 1984a,b 36 38 29 30 27 25 1988c Ozawa 1978, SnLIHL 11-30 36-42 38-42 37-41 33-36 25 39 39 39 34 * ORIGINAL ILLUSTRATIONS (Illustrator) 18-22x 23-41 x 14--18x 12-15x EDlHLt 16-17x 11-21 8-10 8-11 8-11 11-15 mm, 5.8 larvae, Preflexion (N. mm 12.8 Arthur) 47x28 26x15 15xlO 20xlO 14xl0 17x13 Postflexion larva, 33.8 Arthur) mm (N. 1-1 1-2 1-3 1--'3 PILlBL 2-3 4--6 2 1 2 1 3 5 * measurement precluding damaged, specimens yolk-sac of snout Both 0-0 0-0 0-2 L1BL P 0-0 3-4 6-7 2 length. 0 0 0 1 3 7 is second. axis vertical first, given t Eye axis horizontal oval; 346

359 harryi Scopelosaurus Scaly paperbone : ... •. .•... , ...•... , ... ~ ... : ... ,~ ... ~ : ... : .. ·····:IA·.::·(r, .. ::'.~ .. :;.~'.''::':/, .. : ~3~~·~ 5.8 mm 12.8 mm 33.8 mm Figure Notosudidae 2. Preflexion larvae, 5.8 mm (CalCOFI 7205, station 42.143),12.8 mm (CalCOFI 7205, station 39.139); postflexion larva, 33.8 mm (CalCOFI 9107, station 88.60). 347

360 SYNODONTIDAE: Lizardfishes G. MOSER E. G. STEVENS! AND H. subfamilies: family Synodontidae includes three The that depending (5-12, blotches gut lateral species) on Synodontinae (lizardfishes), Harpadontinae (Bombay larval period. the body and gut throughout remain The with Bathysaurinae. Bathysaurinae, 2 and ducks), Larvae rounded. is somewhat head the and are elongate in the Cal- is represented species in a single genus, species 40 mm) ca. to (up size a large reach most of (Hubbs 1979; et al. mollis Bathysaurus by region COFI a gradual to the juvenile undergo transformation and by et 1985). Sulak Larvae were discussed Johnson al. >60 mm in length with stage. Transparent individuals Okiyama (1988), Matarese al. et and (1989). (1974), juvenile morphology and lateral body blotches are 35 species contains about and genera two Synodontinae et et 1978; al (Jones species some for known Sumida at is represented the CalCOFI in survey area and by 1979; al. Moser 1981; 1988; Ozawa 1984; Okiyama al. 1989). Matarese et genus Synodus (Table species Syno- least four in the for limit northern The 1). dontidae most is these of Synodus eggs may be confused with those the S. of only and Sur California Baja Bahia Magdalena, also eggs their since Pleuronichthys genus pleuronectid lucioceps ranges northward into California waters sculpturing be can Eggs shell. the on polygonal have and Robertson Lavenberg (Allen and 1994; Bussing shell by and by the shape identified diameter their 1995). Larvae of nearshore of S.lucioceps occur in waters six- usually are Pleuronichthys in Those polygons. California. the as far north as Monterey Bay, and sided whereas those in Synodus are uniform cm) 60 Synodontines are medium-size (up to ca. five-sided less regularly arranged. The and usually benthic predators found worldwide in and tropical elongate their with heads, elongate rounded gut, larvae entering sometimes bays, and waters coastal subtropical series and paired black gut the on laterally spots are of brackish and cylindrical, elongate, are They waters. nearshore from easily larvae. other separated fish sharply and head a broad with tapering pointed slightly a distinctive feature also are spots peritoneal Paired of Jaws triangular beyond well large, are snout. extend and Saurida, Harpodon, Trachinocephalus, larval depressible teeth. The head the and bear sharp eye, and juvenile Pseudotrichonotus. scales. body are covered with cycloid, tightly adhering deeply is fin caudal The forked. The dorsal fin is The description is based lucioceps Synodus of on anterior is well posteri- the anal midbody, to somewhat 1981; the literature (Moser Okiyama 1984; et Matarese ad, are abdominal, originating the pelvic and well fins of larvae (5 23 examination detailed on and 1989) al. foreward dorsal. of is present. Most the An adipose fin yolk-sac, 3.0-3.8 7 preflexion, 3.7-9.2 mm; 5 mm; are species laterally dorsally red or and to brown gray mm), 15.4-41.9 6 postflexion, mm; 8.1-11.1 flexion, pale and ventrally; some species have a pattern of specimens transformation 4 and mm) 3 (54.5-63.8 (Anderson et al. bars or spots, lateral saddles, dusky and ecological juveniles Meristic data mm). (67.3-88.6 1986). Cressey 1966; Fitch and Lavenberg 1971; information were (Fitch literature the from obtained Lavenberg and Lea 1972) and from 1971; and Miller Synodontines eggs planktonic spherical spawn made observations during Most study. this of the colorless, that diameter) in mm are an lack (0.95-1.48 from localities specimens used in the description were globule, and have a moderately large perivitelline oil of Punta where Sur, California Baja Eugenia, north The space. surface egg shell the covered with a of is other than Synodus species S. lucioceps are rare. The from network; polygon diameters range polygonal southern characters of and morphological pigment on depending 0.034 0.063 to species the mm, (south larvae Sebastian Bahia of the for used Viscaino) at (Zvyagina 1965; Sumida et al. 1972). Larvae hatch consistent with those from are more north- description 2.5-3.8 mm and develop a series of large rapidly only where localities ern lucioceps is found. S. Fisheries Fonnerly, ! at La Jolla Laboratory, National Service, Marine California. Southwest Fisheries Science Center, La Jolla, 348

361 Table Synodontidae Meristic characters for the Synodus species in the California Current vicinity. 1. species All have 10+9 caudal fin rays and 8 pelvic rays. principal Fin Vertebrae rays D A Total PrCV CV C Species PI 2 12-13+11-12 12-13 10-11 10-12 47-48 8 39-40 S. evermanni 11-12 7-9 59-61 8-9 15-16+13-15 51-53 11-13 lacertinus S. 12-15 60-63 9-10 11-13 12-15+12-14 13-15 50-55 S. lucioceps 11-14 10-12 9-10 59-62 11-14+10-13 12-13 50-52 S. scituliceps 11-12 8-9 10-11 11-12+10-11 55-58 13-14 46-50 sechurae S. 349

362 SYNODONTIDAE Synodus lucioceps MERISTICS EARLY LIFE HISTORY DESCRIPTION Mode Range EGGS Shell diam.: 1.2-1.5 mm Vertebrae: diam. 0.9-1.1 Homogeneous; Yolk: mm 0 60-63 No.ofOG: 61 Total Diam.ofOG: Covered with Precaudal polygons surface: 50-55 51 Shell irregular 9 9-10 Candal Paired Pigment: dorsal series that migrate ventrad hatching before Fins: homogeneous size; Large features: Diagnostic polygonal yolk; 0 0 spines Dorsal surface on shell sculpturing 11-13 Dorsal rays 11 LARVAE spines 0 0 Anal 12-15 mm 3.0 12 length: Anal rays ca. Hatching Flexion 8-11 mm length: 8 8 Pelvic length: 50-64 Transformation 13-15 mm 13 Pectoral development Fin Caudal: P D, PI' , , A, C C sequence: 2 I 2 10+9 10+9 ventrolateral Principal from Pigmentation: Yolk-sac-6 near patches PI base to Procurrent: notochord tip; scattered over head Pre yolk sac. & flexion- 12-15 13 from increase 1 postanal intestine 6; 4 to Upper Ventrolateral blotches over 12-14 12 Lower median blotch ventral hypural 1 at & Flexion-postflexion-7 region. intestine; over blotches ventrolateral at base A on margin; hypural on Gill rakers: Upper blotch on hindbrain & cerebellum at ca. 40 mm; lateral 25 mm; ca. to C. anterior Transformation-8 lateral blotches, some diamond Lower Branchiostegals region; brain covered. shaped; opercular in Slender gut; rounded head elongate body; features: Diagnostic LIFE HISTORY ventrolateral blotches increasing preflexion larvae; from round large 7 at to hatching 4 at blotch; postanal ventral single stage; flexion transformation in blotches lateral diamond-shaped pigment; hypural of California Range: San Bay, California, to Gulf Francisco size large specimens; (50-64 transformation. at mm) continental shelf & bays Habitat: Sand or mud bottoms of & (range MORPHOMETRICS mean %) in surveys, fall Spawning season: In CalCOFI are most larvae abundant in during September-November a peak & with winter Y-S PrF F PoF Tr Juv prolonged planktonic ELH pattern: larval Oviparous; larvae; & eggs 71-76 61-67 Sn-AlBL 66-72 67-80 70-74 71-74 64 69 74 73 72 73 period BDIBL 12-15 10-16 10-12 7-9 9-12 10-11 LITERATURE 13 13 11 8 11 11 15-19 13-18 HLlBL 15-19 15-17 16-22 20-23 et Matarese 1989 al. 18 16 16 19 16 21 Moser 1981 58-88 61-71 HWIHL 54--68 45-60 44-53 41-59 1984 Okiyama 68 72 61 51 47 48 11-26 14-19 13-22 SnLIHL 16-22 22-24 21-29 (Illustrator) ILLUSTRATIONS ORIGINAL 19 20 16 19 23 24 43-46x 25-26x 25-28x EDIHL* 24-39x 21-25 20-22 mm Egg, ca. 1.4 (H. M. Orr) 36-40 30-50 30-35 26-34 mm (B. Sumida MacCall) Yolk-sac 3.6 larva, 32x41 44x38 25x33 26x28 23 21 Preflexion mm M. Orr) 6.1 larva, (H. Postflexion mm, lateral & ventral views (H. larva, Orr) 34.0 M. 3-6 6-10 6-8 6-8 12-13 PIUBL 8-13 Postflexion mm M. Orr) (H. 41.9 larva, 7 4 7 12 11 7 mm MacCall) Sumida (B. Transformation specimen, 63.7 UBL P 0-0 0-0 0-0 7-14 0-4 14-16 2 0 0 0 1 11 15 * transformation; during round becoming oval, is somewhat Eye horizontal second. axis is given first, vertical axis 350

363 lucioceps Synodus California lizardfish 3.6mm 1.4 mm 6.1 mm 10.5 mm 34.0 mm ~::=""lE"""::::"":L~::::Z::::::'::::'~~~ mm 34.0 41.9 mm mm 63.7 3.6 Figure Egg, 1.4 mm (CaiCOFI); yolk-sac larva, 1. mm (CaICOFI 7510, station 97.28.8); preflexionlarva, 6.Jmm Synodontidae (CaICOFI 6610, station 120.35); flexion larva, 10.5 mm (Okiyama 1984a); postflexion larvae, 34.0 mm, lateral and ventral views (CFRD transformation 60-17); specimen, 63.7 mm (SK 88396, Torrey Pines (SIO Ref. CoIl., Bahia Ascuncion, Baja California Sur), 41.9 mm Beach, California). 351

364 P ARALEPIDIDAE: Barracudinas AMBROSE A. D. 12 genera Paralepididae about 56 species includes aI1d Paralepidids hermaphroditic and presumed to be are occur species paralepidid nine least At (Nelson 1994). with oviparous unknown are eggs but eggs, planktonic area Paralepididae (Table CalCOFI in the but 1), larvae L. ringens of L. neles and 1984a). Larvae (Okiyama ringens, of only four (Lestidiops neles, Lestidiops without and eyes unpigmented hatch with a functional macrura) Arctozenus are risso, Stemonosudis and mouth; presumably species paralepidid other the hatch sam- collected regularly in CalCOFI ichthyoplankton paralepidids Larval of development. at a similar stage pacificum Lestidiops of Sudis atrox and ples. Larvae body, elongate slender, are short their for noted trunk adjacent in occasionally collected are central Pacific gut the during of elongation marked larvae, early in atlantica Magnisudis masses. and equatorial water pntogeny, and numerous peritoneal pigment patches lengthens. the that increase in number as gut sphyraenopsis Lestidiops of Larvae rare. are larvae Other is have larval Larvae samples. CalCOFI in identified been not pigment typically on top of the head, at the bases in dorsal the of and the caudal and anal fins, johnfitchi are undescribed. Macroparalepis of generally forms pigment body Dorsal region. peduncle transformation but early-forming dorsal during pigment small medium-sized to are Adults (ca. 15 - 1 00 > of certain species in a few characteristic are blotches that fishes compressed cm) elongate, slender, laterally (e.g., genera Uncisudis). Advanced Stemonosudis, zones. occur in the epi-, meso-, and bathypelagic SL) with an of 15-40% larvae (ca. a large head have Adults of s. unknown (e.g., still are species some Eyes profile. a straight and snout elongate are pointed is positioned macrura). A short-based single dorsal fin to slightly strongly oval, depending on species, but behind and long-based fin anal The midbody. is become round by transformation. the behind well originates fin. adipose A dorsal dorsal above the present is fin last anal fin rays. A ventral distinguish used be may characters Numerous the to adipose fin preceding the anal some in present is fin region. of the in species and genera CalCOFI larvae Lestidiops, Sudis). and Stemonosudis, genera (e.g., larvae except Spines in are all lacking paralepidid are with position their and abdominal fins Pelvic preopercular angle. spine a large has at the which Sudis the feature. taxonomic is a useful fin dorsal to respect have larvae macrura prolon- Stemonosudis a distinctive fins Pectoral are low on the body. The snout is pointed Magn- Arctozenus jaw. lower the of and risso gation depressible sharp, large, with terminal mouth and and larvae have peritoneal pigment sections isudis atlantica fixed on bones. and Gill rakers dentary teeth palatine latter the however, posteriad; size in decrease that is on multiple in spines or teeth to reduced are series the region CalCOFI the in with paralepidid only a total raker the number to bony plates; gill refer counts of 60's 1). (Table Paralepididae vertebral in the count bony Some species have no scales, except plates. neles and Lestidiops caudal heavy have larvae pigment (subfamily along tribe and Sudinae the line lateral pacificum larvae Lestidiops lack pigment. lip lower scales cycloid have species other Lestidini); (tribe jaw lower the of but lack possess tip the on pigment Magnisudis). Arctozenus and Light Paralepidini, e.g., Lestidiops pigment on the dorsal body margin. Larval in two present are organs genera and (Lestidium ringens have pigment as well as pigment lip lower on Lestrolepis). Adults of the tribe Lestidini have been by margin body dorsal the preflexion. late and vertically drifting observed swimming rapidly early of life paralepidid The descriptions following observed prey (Houot the are Fishes 1958). principal (M and atlantica) are history stages based literature on in radiographs paralepidids juvenile adult and during of on detailed examinations 36 and 25 between of speci- this study. Barracudinas are important for whales food the other species (Table Paralepididae 2). of each mens and such as salmons and tunas (Fitch and fishes are first time. the for described neles L. of Larvae paralepidids recent Among 1968). Lavenberg fishes, during Meristic data were obtained from counts made related to thought to be most Anotopteridae are closely Ege this study and from the literature (Hubbs 1916; (Rofen 1966c). Parr 1931; Harry 1953; Rofen 1930, 1953, 1957; 352

365 1966a; Shores Miller and Lea 1972; Ozawa 1969; Post (1987) has suggested that the subfamily Okiyama 1984a; Matarese et 1988e; 1987; Post 1986d, Sudinae should be a separate family, that tribes Para- al. Caudal 1992). 1989; and Amaoka et al. vertebral Lestidini lepidini be raised to subfamily rank, and and counts the in provided been not have usually literature be atlantica Paralepis and risso Notolepis renamed that is also excluded from the and for this family this count Magnisudis respective- atlantica, and risso Arctozenus list sources of meristics in of this study. Principal listed ly. "Lestidiops pacificum" is by Post (1972) as information Ege ecological (1953), were Rofen Lestidiops jayakari pacifica. Rofen (1966a) and (1986b, Post and (1984a), Okiyama 1987). (1966a), as al. (1992) listed it a distinct species. Amaoka et that Paralepididae meristic characters for the paralepidid species Selected occur in the California Current region. Table 1. species have 10+9 principal caudal fm rays. All Fin rays Vertebrae PrCV Total D A Species P C PI 2 2 28-34 35-42 72-86 8-13 8-12 12-13+12-13 10-13 risso Arctozenus 28-31 39-42 8-10 100II 9-10 84-88 15-17+15-17 neles Lestidiops II-12 26-30 35-41 76-89 9-10 pacificum 8-9 L. 14-15+14-15 41-45 82-91 26-33 II-12 8-II 8-13 13-20 13-20+ ringens L. II-12 49-51 96-100 10-12 27-30 8 18-20+19 L. sphyraenopsis 51 98 12 12 29 johnfitchi Macroparalepis 9-12 20-24 15-18 9-10 31-41 60-69 13-17+14-17 Magnisudis atlantica 85-97 29-34 7-9 33-38 100II 9 17+17 macrura Stemonosudis 13-15 27-30 52-55 21-22 10-12 16-18+ 15-17 8-9 atrox Sudis the 2. of specimens (above) and size ranges (in mm, Paralepididae used in Number preparation of the Table below) "L" indicates literature used in the description. An paralepidid descriptions. Yolk-sac Flexion Postflexion Preflexion Transformation Juvenile Species 6 0 10 3 4 6 Arctozenus risso 34.0-46.5 5.5-10.8 12.4-23.7 23.6-33.0 60.1-96.1 2 10 6 6 4 5 Lestidiops neles 11.7-17.6 26.3-27.2 3.7-10.2 42.4-55.2 63.5-87.0 3.7-3.9 4 2 10 0 4 5 L. pacificum 11.5-14.5 7.8-7.8 14.7-25.7 28.5-41.2 49.0-80.5 10 6 10 0 5 5 ringens L. 16.5-21.5 3.0-14.8 23.3-37.5 39.1-53.1 63.5-86.1 La La 2 0 3 7 Magnisudis atlantica 23.3-26.6 11.2-16.3 34.0-73.0 9 7 macrura 0 Stemonosudis 7 5 5 23.7-29.8 10.2-22.8 3.4-9.8 34.7-52.8 56.0-83.4 10 6 10 5 Sudis atrox 5 0 11.0-18.2 19.8-21.4 3.2-5.8 7.4-13.3 27.8-45.4 a Ege 1930 353

366 P ARALEPIDIDAE Arctozenus risso LIFE HISTORY DESCRIPTION EARLY MERISTICS EGGS Mode Range diam.: Shell Yolk: Vertebrae: 72-86 No.ofOG: 80 Diam.ofOG: Total Shell surface: 35-42 36 Precaudal Pigment: Fins: o Diagnostic features: 0 spines Dorsal 8-13 9 rays Dorsal LARVAE o 0 Anal spines 28-34 30 length: Hatching rays Anal mm Flexion length: 12-24 ca. 8-12 10 Pelvic ca. gradual Transformation length: 11 34 - >46 mm, Pectoral 10-13 development Fin D, , A, C sequence: Ad , Ph C , P Caudal: I 2 2 10+9 Pigmentation: 10+9 j1exion-l peritoneal Flexion-l-4 patch. peritoneal Pre Principal patches, decreasing in size posteriad. of snout; Postflexion-On tip Procurrent: 12-13 12 Upper & below notochord lower jaw; infraorbital region; cranium; above posterior from part 12 4-9 12-13 of A through caudal peduncle; peritoneal Lower mm, patches; by ca. 25 few first D & of base on bases A rays. of Gill rakers: dorsum from 3-9 to C. Juvenile-Along entire D Transformation-On Upper dorsum, peritoneal 18-36 8-12 patches. Lower 8 features: In preflexion larvae, head pigment absent, 1 Diagnostic 8 Branchiostegals in ca. 50% of 72-86 HW HL, vertebrae; peritoneal pigment patch, decreasing patches, pigment peritoneal 1-9 larvae, flexion-postflexion HISTORY LIFE in D origin a to posterior vertical through P size in posteriad; 2 specimens. & juvenile transformation waters; in the in eastern Pacific Range: temperate & tropical Worldwide 0 from Columbia British (55 to at least north central Baja California N) MORPHOMETRICS %) in mean & (range 0 (28 N) Y-S bathypelagic meso-, Epi-, Habitat: PrF & F PoF Tr Juv Sn-AlBL 32-42 29-35 29-33 season: Spawning occur CalCOFl in year-round larvae Preflexion 71-73 44-68 30 31 36 72 57 ichthyoplankton collections. 8-11 8-9 BDIBL 8-10 7-8 7-7 planktonic larvae ELH pattern: Oviparous; 9 9 9 7 8 17-23 16-19 HLlBL 18-22 24-25 24-25 LITERATURE 17 18 20 24 25 44-51 21-24 26-36 35-49 HWIHL 16-23 Ege 1930 48 43 31 23 18 1989 et al. Matarese 26-41 33-50 46-50 SnLIHL 49-52 50-52 1984a Okiyama 36 41 48 51 51 1966a Rofen 36-42x 19-28x 26-37x 15-18x EDIHL* 12-17x 17-21 19-23 18-22 11-16 14-17 40x19 23x20 32x21 14x13 16x15 axis * Eye slightly to moderately elongate is (horizontally); horizontal Eyes second. vertical given first, 96.1 in specimen juvenile are mm 2-3 2-2 2-4 7-8 not & damaged included. 2 2 3 8 5 on 96.1 transformation & 79.3 t PI measured on 40.4 mm specimen & mm 0-0 0-0 0.8-2 2-3 4-4 of these specimens. remainder on damaged PI specimens; juvenile 0 1 0 2 4 t juvenile mm 96.1 & on damaged 60.1 P on P measured specimens; 2 2 other juvenile specimens. 354

367 risso Arctozenus Ribbon barracudina mm 9.5 ~----~=-~~4 ~~~~~-------~~ mm 18 mm 23 30mm ."1, .. 45mm Figure Paralepididae 1. Preflexion larva, 9.5 mm; flexion larva, 18 mm; postflexion larvae, 23 mm, 30 mm; transformation specimen, 45 mm (Ege 1930). 355

368 PARALEPIDIDAE Lestidiops neles EARLY DESCRIPTION mSTORY LIFE MERISTICS Mode EGGS Range Shell Yolk: diam.: Vertebrae: Diam.ofOG: No.ofOG: 84-88 86 Total surface: Shell 42 39--42 Precaudal Pigment: Fins: features: Diagnostic o spines Dorsal 0 9 8-10 Dorsal rays LARVAE o 0 Anal spines Hatching length: ca. 3.5 rom 30 28-31 Anal rays 22 ca. through mm II rom length: Flexion ca. 9 9-10 Pelvic through ca. 60 mm ca. 40 Transformation length: mm II IO-II Pectoral , , C , PI' C P A, D, Ad Fin development sequence: Caudal: I 2 2 Yolk-sac-Forming 10+9 Pigmentation: finfold, & margin body on eyes, on 10+9 Principal ventral primarily on end angle to of of tail. Prejlexion-Posterior Procurrent: 15-17 margins A of origins lower jaw; on body 17 & future D & fmfolds near Upper peritoneal tail; of end on heavy patches from 0 to 4. increasing fin; 15-17 17 Lower row row along lower jaw; brain; on Flexion-postflexion-Over rakers: Gill postfiexion. patches, row by infraorbital 5-8 peritoneal preopercle; 7-9 Upper Transformation-juvenile---On fin & heavy rays, on C Ad, gradually 27-31 Lower dorsum, over scattering patches. peritoneal 7-8 8 8-9 Branchiostegals lip pigment absent features: flexion-preflexion Diagnostic In larvae, into finfold; in postfiexion larvae to & caudal pigment extends LIFE HISTORY behind stage, juvenile anus well D, a vertical from the insertion of the ED & 10% ca. patches, pigment peritoneal 7-8 to 14% from decreases Range: Vicinity of Cabo San Baja Lucas, to at least Costa Rica California of tip posterior maxilla HL; by transformation. of over nostrils WN) & MORPHOMETRICS (range mean in %) Epi- Habitat: to rnesopelagic F Y-S PrF PoF Tr Juv collected CalCOFI in & larvae preflexion Yolk-sac season: Spawning October samples in January & 27-32 Sn-AlBL 51-69 68-72 22--44 69-72 65-67 29 28 62 70 66 71 ELH larvae planktonic Oviparous; pattern: 4-5 5-6 7-10 BDIBL 6-7 6-7 5-6 LITERATURE 4 8 7 7 5 6 13-16 HLlBL 19-22 13-15 22-23 15-17 18-19 Harry 1953 16 14 14 20 19 22 32-54 25-33 15-18 15-17 HWIHL 33--46 20-20 ILLUSTRATIONS ORIGINAL (Illustrator) 46 40 30 20 17 16 10-19 53-60 55-58 54-57 34--49 14-36 SnLIHL larva, 3.7 Yolk-sac (M. Vona) rom T. 40 25 13 56 57 56 10.1 mm Vona) (M. T. larva, Preflexion 14-14x 11-14x 26--42x 21-30x 9-IIx 33--42x EDIHL* rom (M. Vona) T. Flexion 17.6 larva, 13-14 23-35 18-24 8-10 25-36 10-14 14xl4 36x31 12xl2 IOx9 38x30 26x22 i ~ Eye slightly elongate axis (horizontally); horizontal * vertical fust, given is I-I 2-3 2-2 2--4 5-7 4-6 I second. 2 3 2 I 6 5 specimen & two in mm) (3.9 smallest yolk-sac largest damaged t PI I-I 0-0 0-0 4-5 2--4 0-0 specimens damaged of rom); 43.6 & (42.4 specimens transformation PI 0 0 0 3 5 mean. or range in included not PI & P t damaged in largest juvenile (87.0 mm) & are not included in 2 mean. range or 356

369 Lestidiops neles 3.7mm 10.1 mm 17.6 mm 78mm FigureParalepididae 2. Yolk-saclarva, 3.7mm (CalCOFI 6001, station 157.40); preflexion larva, 10.1 mm (CalCOFI 7202, station 157.10); flexion 17.6 mm larva, (CalCOFI 7210, station 157.50); juvenile, 78 mm (Harry 1953). 357

370 PARALEPIDIDAE Lestidiops pacificum EARLY LIFE HISTORY DESCRIPTION MERISTICS Range Mode EGGS Vertebrae: Shell 82 diam.: 76-89 Total Yolk: 36 35-41 No.ofOG: Precaudal Diam.ofOG: Shell surface: Fins: Pigment: o Dorsal spines 0 featnres: 10 Diagnostic rays 9-10 Dorsal o Anal spines 0 LARVAE 30 Anal 26-30 rays 8-9 Hatching length: Pelvic 8 11 11-12 Flexion length: ca. 11.5-15 mm Pectoral 28-42 mm Transformation length: ca. Caudal: 10+9 , Ad C D, A, & , 10+9 P PI' Fin sequence: C development Principal I 2 2 Preflexion-On lip; internal behind eye; along ventral Pigmentation: Procnrrent: 15 14-15 from margin body increase patches peritoneal develop; will A where Upper 15 1 to 4. fore- between row 14-15 Flexion-pos(/1exion---Crescent-like & Lower posterior row; dorsally; infraorbital Gill rakers: midbrain along row jaw; lower to 5 & the below above A; & anus margin body midventral between Upper Lower 36 peduncle the & along the ventral body margin in caudal notochord Transformation- 4-12. 8 patches region; peritoneal increase to Branchiostegals 8 over cranium; at base isthmus; ventral lobe juvenile-{)n snout, ofD, extending along dorsum; ofC; patches. gradually 11-12 peritoneal LIFE HISTORY larvae, pigment In Diagnostic from features: preflexion-flexion absent finfold, & caudal eye, behind pigment embedded dorsal body margin Range: In the northeastern ca. from south Pacific, 38° & ca. N east of caudal in & below & above rows pigment at peduncle; notochord 173° Gulf collected also W; of Panama, off Chile, & in north of New A & postflexion stage, pigment row tail margin between anus along Zealand. pigment & crescent-shaped row of between fore- & midbrain; in transformation-juvenile 11-12 peritoneal pigment patches, specimens, meso-, Epi-, Habitat: & bathypelagic ventral half of than C half, & anus more heavily pigmented dorsal D. under collected specimens Preflexion season: Spawning & June in August mean MORPHOMETRICS (range Ufo) in & larvae Oviparous; pattern: ELH planktonic Y-S PrF F LITERATURE PoF Tr Juv Sn-AlBL 25-25 54-60 27-44 57-62 57-60 Amaoka et 1992 al. 34 25 60 57 59 1953 Ege and Rass 1973 Pertseva--Ostroumova 5-8 BDIBL 9-9 7-8 7-9 5-6 9 8 6 6 8 Rofen 1966a 14-16 HIJBL 16-18 13-15 16-17 19-21 ORIGINAL (Illustrator) ILLUSTRATIONS 14 17 17 15 20 33-38 35-39 HWIHL 22-25 24-32 31-40 Arthur) (N. mm Preflexion larva, 7.8 36 34 38 27 23 (N. Arthur) Flexion larva, 11.5 mrn 31-36 26-30 SnlJHL 35-48 43-45 42-48 mrn Postflexion larva, 14.7 (N. Arthur) 28 33 42 44 45 31-35 EDIHL 23-32 20-23 17-21 30* 33 28 21 19 specimen flexion an pre one Only * eye. with 2-2 2-3 2-3 2-5 PILlBL 4-7 2 2 3 2 6 0-0 3-4 L1BL P 0-0 0-2 6-7 2 0 0 0.5 6 3 358

371 Lestidiops pacificum mm 7.8 mm 11.5 14.7 mm ·'"'··· ._-_···...",.,.c···,.·i!"'· ···.·.·o·_···~···· ..•...••...•...•..•..•.•• 0 .. 33.0 mm Figure Paralepididae 3. Preflexion larva, 7.8 mm (Cato I, station A 1); flexion larva, 11.5 mm (Climax II, station A 5); postflexion larva, 14.7 (Climax II, station A 5); transformation specimen, mm 33.0 mm (Amaoka et al. 1992), pectoral fin should have rays. 359

372 P ARALEPIDIDAE ringens Lestidiops MERISTICS DESCRIPTION HISTORY LIFE EARLY Range Mode EGGS Vertebrae: diam.: Shell Yolk: 82-91 Total 86 No.ofOG: Diam.ofOG: 41 41-45 Precaudal Shell surface: Pigment: Fins: spines Dorsal 0 o features: Diagnostic 8-13 Dorsal 12 rays Anal spines 0 LARVAE o Anal 28 rays 26-33 Hatching length:

373 Slender barracudina ringens Lestidiops 5.6mm 9.4 mm ... ~ ... -'f4f, ... ~ ~/ ~:r: .. ~ ... :~ ... :' .. ~ ... ~ ... ' .. ~ ... ::~ ... §!;,~E~.~;: ... . : .. ::.:= ... :: ... : ... = .. = ... = ... _~.= .. ~--=~_. =._'==--. =- .. :::.--=:~~_~ .. _.i __ ~ ___ :_:_~ 16.5 mm 28.5 mm 58.0 mm Figure Paralepididae 4. Preflexion larvae, 5.6 mm (CalCOFI 6706, station 103.40), 9.4 mm; flexion larva, 16.5 mm; postflexion larva, 28.5 (Moser 1981); late transformation specimen, mm 58.0 mm (CalCOFI 6208, station 60.80). 361

374 PARALEPIDIDAE Magnisudis atlantica MERISTICS EARLY LIFE mSTORY DESCRIPTION Mode Range EGGS Vertebrae: diam.: Shell Yolk: Total 68 60-69 No.ofOG: Diam.ofOG: Precandal 31-41 surface: Shell 31 Pigment: Fins: Diagnostic features: Dorsal spines 0 0 9-12 Dorsal rays 10 Anal 0 spines 0 LARVAE length: Hatching Anal rays 20-24 22 length: Flexion mm ca. 8.5-10.5 9-10 9 Pelvic 15-18 16-17 Pectoral ca. length: Transformation mm, 23-47 gradual Caudal: A, sequence: development Fin Ad P C Ph , C D, , , I z z 10+9 10+9 Principal increasing Pre flexion-l peritoneal patch to 2. Pigmentation: Procurrent: snout, origins; above Pos{/lexion-On cranium, & A jaws, & D & 17 13-17 Upper stage, A; nape, on postflexion late by of end near notochord below 17 14-17 Lower patch between Ad 3 peritoneal & A, & patches decreasing in size rakers: Gill body margin; tail dorsal Transformation-juvenile-Along posteriad. 7-10 row expands; region. patch on crescent-like infraorbital 8 Upper 25-32 30 Lower number Vertebral features: Diagnostic to (60-69) be appears only 8 8 Branchiostegals to separate early larvae from those reliable character of Arctozenus rays in the family A rays numerous most & PI (72-86); (15-18) risso LIFE HISTORY 1 peritoneal (20-24) are patch, increasing among the fewest; pigment stage), 2 (preflexion-flexion to postflexion-juvenile late 3 by & stage; anus at final position (64-72% SL) by 13-15 mm (early postflexion Range: Alaska southeast from Pacific eastern in Pacific; & Atlantic 0 stage). Chile; of distribution known broadest to N) paralepidid any (59 MORPHOMETRICS & mean in %) (range approach coasts in adults Habitat: Epi-, meso-, & bathypelagic; large the temperate & zones polar PrF* Y-S F* PoF Tr Juv Spawning larvae Mainly March-June in western Atlantic but season: found year-round; larvae rare in CalCOFI ichthyoplankton collections 35-38 Sn-AlBL 39 40-71 67-72 64-71 37 56 70 69 ELH pattern: larvae planktonic Oviparous; BDIBL 13-14 11-13 12-14 12 12 13 LITERATURE HLlBL 23-33 34-36 31-35 28 35 33 Ege 1930 HWIHL 21-23 23-39 19-25 Rofen 1966a 31 22 22 1984a Okiyama SnLIHL 48-59 54-56 48-54 Ozawa 1986d 53 51 55 1989 aI. et Matarese EDlHLt 13-16x 14-27x 11-18x 14-19 14-16 11-17 * (1930). Ege Adapted from 21x17 15x15 15x14 t (horizontally); Eye horizontal axis elongate given to slightly moderately 7-7 7-7 fust, vertical second. 5 7 7 25.7 mm on & larva postflexion mm 16.3 measured PI 26.6 & on t 2-2 2-3 specimens transformation of remainder these damaged PI only; on 3 3 2 specimens. & P P measured on 34.0 § 38.6 mmjuvenile specimens only; damaged z z specimens. juvenile of remainder on 362

375 Duckbill barracudina atlantica Magnisudis mm 5.8 9.8 mm 14.3 mm 28.0 mm 54.5 mm Figure Paralepididae 5. Preflexion larva, 5.8 mm; flexion larva, 9.8 mm; postflexion larva, 14.3 mm; transformation specimen, 28.0 mm; juvenile, 54.5 mm (Ege 1930). 363

376 P ARALEPIDIDAE Stemonosudis macrura MERISTICS DESCRIPTION EARLY LIFE HISTORY Range Mode EGGS Shell diam.: Vertebrae: Yolk: 92 85-97 No.ofOG: Total Diam.ofOG: 31 29-34 surface: Shell Precaudal Pigment: Fins: features: Diagnostic o 0 spines Dorsal 7-9 8 rays Dorsal o spines 0 Anal LARVAE 33-38 35 Anal rays Hatching length: 9 length: Flexion ca. 10-23 mm 9 Pelvic II 10-11 Pectoral ca. Transformation length: gradual mm, 35-53 Caudal: Fin development sequence: PI' D, Ad C P C A, , , h 2 2 10+9 10+9 Principal Preflexion--On base snout, jaw, oflower prolongation Pigmentation: Procurrent: behind eye; by ca. 7 mm, 3-5 cranium on peritoneum; by of patches 17 17 Upper ca. along patches 2-3 & margin tail ventral along rows 4-5 mm, 8 17 17 margin. tail dorsal Lower Flexion-postflexion-Above below end of & rakers·: Gill peritoneal patches 15. to 4 from increases notochord; 0 Upper anus between margin body along Transformation-juvenile-Row & 19 Lower row along base of A; 2 rows A; lateral caudal peduncle region; on 8 8 Branchiostegals dorsal along row increases 16-17 development; with margin body peritoneal patches. HISTORY LIFE the of features: Diagnostic prominent prolongation cartilaginous A juveniles lower present through transformation stage; jaw with 16-17 peritoneal pigment patches melanophores & a row along of punctate the of from Pacific, eastern the in Pacific; & Indian waters Warm Range: each side 33-38 vertebrae; 85-97 margin; body dorsal of rays. A Point Conception, California, to Chile mean %) & (range MORPHOMETRICS in to mesopelagic Habitat: Epi- in Y-S CalCOFI ichthyoplankton larvae occur Preflexion season: Spawning PrF F PoF Tr Juv ca. up to mm 150 juveniles from only known year-round; collections Sn-AlBL 25-58 24-29 53-59 53-57 52-63 26 42 58 56 56 ELH pattern: Oviparous; planktonic larvae 4-7 8-11 BDIBL 4-5 3-5 3-5 LITERATURE 9 4 6 4 4 HLlBL 16-22 15-19 16-19 13-17 13-15 1957 Ege 18 17 17 14 15 Okiyama 1984a 17-26 27-41 HWIHL 13-17 10-15 15-23 1988e 1986d, Ozawa 32 21 14 13 18 1966a Rofen 33-53 24-34 SnLIHL 49-55 41-53 50-54 and Pertseva-Ostroumova Rass 1973 41 30 52 50 52 EDIHL 21-35 13-21 10-14 9-11 9-13 27 17 12 10 10 63-407, largest in specimen examined forming still plates Bony * (SIO mm). 135 3-5 2-3 1-2 PIUBL 3-5 1-3 4 2 I 2 4 P UBL 0-0 0-0 1-3 0-3 2-4 2 0 0 2 3 3 364

377 Sharpchin barracudina Stemonosudis macrura 3.6mm 8.8mm III 'I I " 49.0 mm 60.0 mm Figure Paralepididae 6. Preflexion larvae, 3.6 mm, 8.8 mm (Ozawa 1986d); flexion larva, 15.0 mm; transformation specimen, 49.0 mm, 60.0 mm (Ege 1957). peritoneal pigment patches faded; juvenile, 365

378 PARALEPIDIDAE Sudis atFox HISTORY DESCRIPTION EARLY LIFE MERISTICS Range Mode EGGS Vertebrae: Shell diam.: Yolk: 52-55 Total 54 No.ofOG: Diam.ofOG: Precaudal 27 27-30 Shell surface: 27 25-28 Caudal Pigment: Fins: Diagnostic features: Dorsal spines o 0 LARVAE Dorsal 12 rays 10-12 length: Hatching o Anal spines 0 21-22 through mm rays Anal 21 11-13 Flexion mm length: ca. 5-6 ca. 8 gradual mm, 20 length: Transformation 8-9 Pelvic Pectoral 14 13-15 & Ad P PI' D & A, C C sequence: development Fin , b 2 2 Caudal: Pre flexion- along peritoneum. Three patches Flexion- Pigmentation: 10+9 10+9 Principal tip, snout 5 peritoneal preoperculum; patches. Posiflexion-On On Procurrent: hindbrain, midbrain, jaw, lower spine, preopercular caudal & 16-18 Upper 17 margin; body vertical dorsal Transformation-juvenile-On peduncle. 15-17 Lower 17 ; at of C; line base 6 rays; PI dorsal along row of P base at line 2 patches. peritoneal rakers: Gill 5-10 Upper of vertebrae (52-55); in preflexion number Low features: Diagnostic 21-28 Lower 3 peritoneal & eyes, 3 large preopercular larvae, elliptical spines, 8 Branchiostegals 8 pigment series longitudinal larvae, in flexion spots; of fine serrations above eye larvae, pigment patches; in postflexion 5 peritoneal large & HISTORY LIFE preopercular spine with serrated edges & a few row & hooks retrorse along serrations fine of transformation-juvenile in mandible & snout; large, elongate specimens, PI & 6 peritoneal pigment patches. Range: Tropical western Atlantic at Pacific Pacific; & from in eastern least Chile to N 32° MORPHOMETRICS (range & in mean %) meso-, Epi-, Habitat: & bathypelagic Y-S PrF F PoF Tr Juv Spawning through Early-stage larvae occur mainly from February season: July samples ichthyoplankton CalCOFI in Sn-AlBL 44-57 65-71 57-65 62-67 65-69 52 61 68 66 65 Oviparous; larvae, in primarily upper ELH planktonic pattern: m 250 9-11 BDIBL 13-16 10-13 13-15 12-15 11 11 14 14 13 LITERATURE 21-24 22-32 HLlBL 35-44 30-36 35-41 22 26 40 39 32 Belyanina 1982a 37-67 HWIHL 57-77 23-30 24-28 23-27 Berry 1966 Perkins & 66 27 49 25 26 Okiyama 1984a 32-54 SnLIHL 26-39 52-57 51-53 48-53 1988e Ozawa 1986d, 31 41 54 52 51 1966a Rofen EDIHL* 32-40x 14-21x 23-33x 15-16x 15-18x 1969 Shores 15-21 13-18 14-21 14-20 12-17 36x18 29x16 16x17 16x17 16x15 is elongate moderately (horizontally); horizontal axis to slightly Eye • 4-8 4-5 5-13 11-18 15-22 vertical given second. first, 5 9 5 15 19 0-0 0-0 3-7 6-7 7-7 0 0 6 7 7 366

379 Hideous barracudina atrox Sudis ~) 2.4 mm .".''-? 5.4 mm 9.2mm 21.5 mm Figure Paralepididae 7. Preflexion larva, 2.4 mm; flexion larva, 5.4 mm; postflexion larva, 9.2 mm (Ozawa 1986d); late transformation specimen, 21.5 mm (Berry and Perkins 1966). 367

380

381 ANOTOPTERIDAE: Daggertooth D. A. AMBROSE anotopterid single the is pharao species Anotopterus Anotopterids are hermaphroditic (Okiyama 1984a); rarely in known collected (Nelson 1994). Larvae are are The smallest available larva eggs unknown. lack are Adults region. CalCOFI elongate, the slender, pointed with head body, a slender has mm) (8.5 a large developed adipose a well have fin, a dorsal fin but prolongation a fleshy snout, at the tips of jaws, a both a small anal above fill. The mouth is large, with and rays, fin undeveloped tooth, canine palatine a gut strong teeth, the largest of which the are fanglike pigment peritoneal lacks and midbody past extends that projection elongate An teeth. palatine the is present on patches. By about 25 forming in all fins are rays mm, lower was 147 specimen recorded jaw largest tip. The between the the along runs tube a sinuous and dorsum weighed and long cm (Eschmeyer et al. kg 1.6 only head adipose and fin. Development is direct. A. have was thought to 1983). antitropical an pharao Pacific and Atlantic the in distribution al. et (Hubbs account species following The detailed on is based mm since specimen has 41 a single however, 1953); examination of an 8.5 mm preflexion (CaICOFI larva collected at been (SIO W 139°8.0' S, 0°5.8' 60-23) 11 station speci- transitional mm a 25.1 00), 0.1 8403, from species the recorded has (1979) Fourmanoir and men 70-105), 1966c; (Rofen (SIO literature and Okiyama data from Meristic 1984a). were obtained scom- of contents the from Caledonia New stomach brids, as as northwest of Madagascar. Rofen well Rofen 1953; et al. (Hubbs literature and Miller 1966c; that (1966c) observed the larger adults inhabit colder information was obtained 1972). Lea Ecological al. et and (1966c) Rofen from mainly Eschmeyer and waters toward the poles, whereas, the young (1983). smaller adults inhabit more regions. temperate Anotopteridae most be to thought is to related closely 1984b). the Paralepididae (Okiyama 369

382 ANOTOPTERIDAE Anotopterus pharao DESCRIPTION HISTORY LIFE EARLY MERISTICS Mode EGGS Range Shell diam.: Vertebrae: Yolk: 79-80 No.ofOG: 78-83 Total Diam.ofOG: 49-53 Shell surface: 52 Precaudai Pigment: Caudal features: Diagnostic Fins: o 0 Dorsal spines LARVAE o 0 rays Dorsal mm length: Hatching <8 o 0 Anal spines Flexion length: > <25 14-17 mm, 14 14 mm rays Anal mm 9-11 25 ca. length: Transformation 10 Pelvic 12-16 14 sequence: PI' development P C, A, Ad, Fin Pectoral 2 projection, jaw Pigmentation: Pre flexion-By 8.5 mm, on lower Caudal: 10+9 on scattered snout lateral & patch 10+9 posterior to eye, brain, Principal of edges at margin of tail, & ventral gut, along dorsolaterally along Procurrent: finfold Ad regions; along in half & absent A dorsum; 14-17 of posterior Upper end from 25 body, & of 14-16 over Transitional-By ca. tail. mm, scattered Lower concentrated along dorsum. Gill rakers: length o BL; >60% Diagllostic featllres: Total vertebrae 78-83; preanal Upper 0 0 o head large of tips at prolongation with Lower both pointed snout; a fleshy 7-9 fanglike pigment peritoneal D; large 7 teeth on palatines; no jaws; lBranchiostegais by between patches absent; dorsum ca. 29 mm, sinuous tube along head & Ad. HISTORY LIFE %)* (range & mean in MORPHOMETRICS temperate Primarily Range: the world of waters polar & Y-S PrF Habitat: mesopelagic; least ocean open at Epi- to F PoF Tr Juv Spawning Sn-AlBL Preflexion 61-66 season: larvae collected in CalCOFI region during 64 69 69 September & May March, January, 8-9 BD/BL Oviparous; planktonic larvae pattern: ELH 8 8 7 25-30 HLIBL LITERATURE 27 35 32 HWIHL Maul 1952 22 19 1948 Nybelin 52-60 SnLIHL Okiyama 1984a, 1988g 56 66 59 Rofen 1966c EDlHLt 14 13 10 14.2 * Morphometries include measurements from illustrations of mm 2-2 PILIBL 2 1966c). larva (Okiyama 1984a) & 50 mm juvenile (Rofen preflexion 4 3 8.5 from missing Eye t iarva. mm pref1exion 0-0 L1BL P2 0 0.8 370

383 Daggertooth pharao Anotopterus 14.2 mm 50.0 mm Figure Anotopteridae 1. Preflexion larva, 14.2 mm (Okiyama 1984a); juvenile, 50.0 mm (Rofen 1966c). 371

384

385 EVERMANNELLIDAE: fishes Sabertooth . A. AMBROSE D. includes 7 or 8 species Evermannellidae 3 genera and pigment sections, melanophores along the myosep- neal Evermannella 1994). Swinney (Johnson 1984b; 1982, a little Teeth behind midbody. ta, and the anus located ahlstromi Development larvae. small in very appear is direct and the Califor- and from E. indica are reported of E. ahl- 30 the only larvae by Current region; however, completed is juvenile to transition gradual nia coalesce mm SL. The peritoneal pigment patches and stromi identified been samples. have CalCOFI in gut enclose completely by the SL. mm 35-45 ca. 18 are Sabertooth length (maximum small fishes the of predators between mesopelagic cm) ocean open and externally an possess They ca. 40° N S. 40° have pigment Evermannella peritoneal three Larval of musculature, tail the visible division tripartite base. fin the on melanophores and patches caudal eyes tubular six or semi-tubular in species, seven the of melanophores are peduncle more numerous in Caudal a large anteriormost monstrous jaws, massive head, than E. indica E. larvae. ahlstromi in fangs, palatine present) (when scales and body, a deep are counts to the lateral line Meristic region. restricted are descriptions following The on detailed based PI 26-37, A 10-13, and 11-13, family for are D the mm), 6 of 7 preflexion larvae (3.4-4.9 examinations 45-54. be to thought vertebrae fishes Sabertooth are larvae flexion larvae (4.9-7.9 mm), 8 postflexion group the to a sister Alepisauridae plus Omosudidae (23.0- 4 transformation mm), (10.1-22.4 specimens 1982, (Johnson but 1984b; see Patterson and Johnson 29.0 and 4 juveniles (35.6-56.1 mm) mm), of E. 1995). mm), ahlstromi and 4 postflexion 4 (8.8-18.3 larvae 4 mm), (20.4-26.0 and specimens transformation Evermannellids are synchronous hermaphrodites E. of mm) (30.0-58.5 E. of Larvae indica. juveniles described whose eggs are unknown. Larvae have been for ahlstromi than 10.2 smaller mm are described here for all genera (Schmidt Rofen 1966d; Johnson 1918; time. first obtained the were the from data Meristic Johnson and Glodek 1975; Wassersug and 1976; literature and Glodek (Johnson 1975; Johnson 1982). Ozawa Larvae are moderately 1986c). Johnson 1982; a pointed snout, large elongate and compressed with Ecological information was obtained from Johnson (1982). oblong perito- mouth, unpaired large to moderate eye, 373

386 EVERMANELLIDAE Evermannella ahlstromi MERISTICS HISTORY EARLY LIFE DESCRIPTION Range Mode EGGS Vertebrae: Shell diam.: Yolk: Total 47-49 48 No.ofOG: Diam.ofOG: Precaudal surface: Shell Caudal Pigment: Fins: Diagnostic features: spines Dorsal 0 0 rays Dorsal 10-l2 II LARVAE Anal spines 0 0 mm <3.4 length: Hatching Anal rays 29-32 30-31 mm ca. 4.9-7.9 Flexion length: Pelvic 9 23-30 mm ca. length: Transformation 9 12 Pectoral 12 Fin C , D, development sequence: , P & A, C PI I 2 2 Caudal: laterally hindbrain; Preflexion-On Pigmentation: on isthmus; 10+9 10+9 Principal internally at dorsolaterally patches gut; 1-2 3 base; PI on Procurrent: dorsal melanophores row a in 3-4 anus; to posterior & gut on Upper on on dorsum; edge distal anteriorly middorsally; of finfold 8-13 Lower 2-7 dorsolaterally on posterior tail myosepta; scattered at mid-dorsum; rakers: Gill 17-27 ventrally tail. along Flexion-On lower jaw tip; snout; line Upper o o & on mid- &. hindbrain; spreading into midbrain; fore- between o Lower o from 3 dorsal patches myosepta & from tail Posiflexion- row. ventral 8 Branchiostegals 8 on operculum; On upper jaw & angular; vertical row 7 fins; all on 2 in patches in epaxial myosepta, 5 midlateral myosepta, in hypaxial LIFE HISTORY line at base myosepta; blotches of linearly Transitional-Heavy C. covering Juvenile-Generally bands. muscle 3 with associated body; blotches numerous. iridescent more off waters transitional Pacific; Eastern Range: Baja equator- California, 0 A 47-49 total vertebrae; D 10-l2, usually II; Diagnostic features: & to ially ca. 145 W, & in transitional waters between equatorial central pigmentation peduncle caudal 31; usually by transforma- light; 29-32, & Pacific South North the of waters gill & projecting beyond cover. elongate filaments gill stage, tion adults mesopelagic ocean; Open Habitat: in %) MORPHOMETRICS & (range mean season: occur Spawning Larvae in samples from CalCOFI February-April July-October & Y-S PrF F PoF Tr Juv upper ELH pattern: Oviparous; planktonic larvae usually collected in Sn-AlBL 47-55 50-63 51-59 47-52 53-60 52 57 56 49 100m 55 BDIBL 13-16 14-18 17-20 17-18 20-22 LITERATURE 14 16 18 17 21 HLlBL 24-29 23-31 29-33 23-26 28-29 1984b 1982, Johnson 27 30 26 28 24 Johnson Glodek & 1975 29-43 HWIHL 50-57 25-38 32-36 38-42 51 36 26 39 34 ORIGINAL ILLUSTRATIONS (Illustrator) 29-44 SnLIHL 34-46 30-43 28-33 23-26 36 39 36 30 24 Preflexion larva, 3.4 mm Walker) (R C. EDIHL* 14-18x 15-19x l2-16x 17-20x 22-26x C. (R Walker) mm Flexion 7.5 larva, 22-33 31-47 18-24 23-25 23-27 larva, Postflexion Walker) C. 10.1 mm (R. 15x28 17x38 14x21 18x24 23x26 3-5 4-6 II 12-13 3-ca. >14 horizontal vertically; elongate Eye * second; vertical first, given axis 4 5 12 are measurements eye entire of pigmented portion. not just capsule 0-0 0-2 7-9 9-10 >9 on damaged specimens. juvenile all P & PI t 2 0 0.5 8 9 374

387 Evermannella ahlstromi mm 3.4 7.5 mm ~~ ... :J~~ \Ij 10.1 mm .:)':.,'.: ... ;;. /: ... -:. .. ' 'i .:. ... : ... ':' '" ... , 17.7 mm 26.6 mm Figure Evermannellidae 1. Preflexion larva, 3.4 mm (EASTROPAC II, station 46.052); flexion larva, 7.5 mm (CaICOFI 7205, station mm; 20.125); larvae, 10.1 mm (CaICOFI 5910, station 107.90), 17.7 postflexion transformation specimen, 26.6 mm (Johnson and Glodek 1975). 375

388 EVERMANNELLIDAE Evermannella indica EARLY LIFE HISTORY DESCRIPTION MERISTICS EGGS Mode Range Vertebrae: Shell diam.: Yolk: No.ofOG: 50 Total 48-52 Diam.orOG: Shell 17 surface: 17 Precaudal Pigment: 33 33 Caudal features: Diagnostic Fins: 0 0 Dorsal spines LARVAE 12 12-l3 Dorsal rays Hatching 0 length: 0 mm <3.7 spines Anal 27-31 29 mm mm-6.3 <4.4 ca. length: Flexion Anal rays 9 Pelvic 20-30 mm 9 Transformation length: ca. 11-12 12 Pectoral sequence: development Fin PI C A, & P D, C , b 2 2 Caudal: isthmus, between occiput; Pigmentation: Preflexion-On snout tip, & 10+9 10+9 Principal orbits; at 3 peritoneal PI patches; at midlateralline myosepta on base; Procurrent: peduncle. caudal on trunk; of on ventrally; tail on Flexion-Row Upper above at shoulder & anus; myosepta midlateral trunk epaxial on Lower rows myosepta; vertical myosepta; proximally tail on on D finfold at proximally midbody; near Gill finfold rakers: A on midtail; Post- base. C on Upper o o on heavy myosepta; trunk & & on head flexion-Increasing dorsum o o Lower caudal on ventrum Transi- operculum. on series vertical peduncle; 8 8 Branchiostegals with associated linearly, tional-On blotches large fins; all forming body numerous; more Juvenile-Lateral blotches the 3 muscle bands. LIFE HISTORY iridescent. total A 48-52 usually 12-l3, D vertebrae; 12; Diagnostic features: filaments usually 29; 27-31, peduncle pigmentation heavy; gill caudal Pacific & Indian & Atlantic, the of waters equatorial Central Range: not cover. projecting beyond gill Pacific eastern from absent Oceans; waters equatorial generally %) MORPHOMETRICS (range & mean in m hauls upper ocean; Open Habitat: 100-800 depth, usually taken in exceeding 400 m Y-S PrF PoF F Tr Juv Spawning season: Larvae & small juveniles taken throughout the year but none identified in CalCOFI samples Sn-AlBL 50-55 54-62 52-60 57 55 53 upper in collected larvae planktonic Oviparous; pattern: m 100 ELH BDIBL 17-23 16-17 17-22 19 17 19 LITERATURE HUBL 25-36 23-26 24-30 29 24 27 1984b Johnson 1982, HWIHL 28-39 33-45 33-45 1975 Glodek & Johnson 34 39 39 1986c Ozawa SnL/HL 31-41 31-35 24-28 1966d Rofen 31 36 27 EDIHL* 14-20x 17-21x 23-25 x 21-25 23-26 29-32 first, elongate second; * vertical vertically; given axis horizontal Eye are measurement eye capsule, not just pigmented portion. of entire 16x22 18x25 24x31 transformation all on damaged specimens. P t & juvenile 2 8-9 PILlBL 8-10 >9-17 9 9 8-9 P L1BLt 2 9 >9 >9 376

389 indica Evermannella Sabertooth . II .0._ \ , .p~, ~ ... ... ..' . ... ~- '''''~ 4.4 mm mm 6.3 , , \ \ I I , , , / mm 9.1 14.6 mm 28.0 mm Figure Evermannellidae 2. Flexion larva, 4.4 mm; postflexion larvae, 6.3 mm, 9.1 mm, 14.6 mm (Ozawa 1986c); juvenile, 28.0 mm (Johnson 1982). 377

390

391 ALEPISAURIDAE: Lancetfishes AMBROSE A. D. and two The includes family one genus Alepisauridae length preanal teeth, a and canine prominent flexion, ferox (Okiyama A. and brevirostris Alepisaurus species, larvae to preflexion in BL 37% ca. increases that from specks 1984a). 81 % BL in transformation specimens. Fine ca. Pacific north A. the from absent is brevirostris have fins pre- of pigment are present ferox A. been by OIl collected the pectoral (Francis Larval 1981). E west and 179°04' of N 29°48' of south forms pigment peritoneal stage; flexion during only the however, adults range in (Boehlert and Mundy 1992); evermanneHids, the Unlike with postflexion stage. the eastern Pacific from Alaska to whom closely be may they related, alepisaurids do Chile. Lancetfishes the peritoneal pigment sections (Johnson have not are distinct large length (maximum ca. em), pelagic to 200 Like the scopelarchids and giganturids, the 1982). bathypelagic tropical and temperate in seas. predators have a translucent alepisaurids space between the gut possess They a compressed, elongate body covered the of and hypaxial completion to prior musculature, and a large mouth with jaws pointed with pores, Transformation flexion stage. keel along the a fleshy teeth, dagger-like each side of ferox A. In gradual. is present from preopercular four are larvae, by the spines extending a sail-like dorsal fin peduncle, caudal stage flexion fin anal the near forms pigment and adipose behind head almost the small the fin, and a to Scales, fin far located fin anal short-based posteriorly. postflexion the by origin stage. larvae A. brevirostris light organs, and a swim bladder are spines, absent. anal lack the pigment and spines at preopercular both Meristic counts for the family D 29-49, A 11-19, are alepisaurids Preflexion origin. confused be may fin and PI 12-16, P 1984a). 7-10, V 47-51 (Okiyama scopelarchids; however, alepisaurids with have a more 2 are of adults is mostly iridescent. They The body eye. rounded noted for new many and appetite voracious their con- stomach their been described have species from following species description is based on The mm a 5.2 of 3 larva, preflexion detailed examinations (Fitch 1968). Lavenberg and tents flexion mm), (6.9-8.4 larvae 10 postflexion larvae Alepisaurids are synchronous hermaphrodites whose 5 mm), transformation specimens (6.4-15.5 (15.8- eggs unknown. are Larvae have for described been ferox. A. mm) (30.6-52.2 juveniles of 5 and mm), 22.3 were data (Fujii literature from obtained Meristic Alepisaurus as brevirostris Alepisaurus both species: 1984a; Okiyama Heemstra and Smith 1986; 1984d; (Rofen mm sp., 10.0 , ferox A. 6.9-17.2 1966b) and was 1989). al. information Ecological et Matarese (Okiyama (Okiyama and 6.4 1984a) mm 1988f). mm from Esch- and (1968) Lavenberg and Fitch obtained elongate, have a large head (HL Larvae are moderately (1983). aL meyer et until eyes oblong mid-post- somewhat BL), 24-43% 379

392 ALEPISAURIDAE Alepisaurus ferox EARLY LIFE DESCRIPTION HISTORY MERISTICS Range Mode EGGS diam.: Shell Yolk: Vertebrae: 50 47-52 No.ofOG: Total Diam.ofOG: surface: Shell 23 19-26 Precandal Pigment: 27 24-31 Candal Diagnostic features: Fins: Dorsal o spines o LARVAE Dorsal rays 30-45 length: Hatching <5 mm o Anal spines o length: Flexion 6-8.5 mm ca. l3-18 rays Anal length: ca. 8-10 mm 16-30 Transformation Pelvic 12-15 P D, A, PI' C, sequence: development Fin Pectoral 2 flexion-Fine jaw lower Pigmentation: Pre Flexion-On specks on PI. Caudal: 10+9 10+9 gular Principal angle. Postflexion-Dorsally on head; & operculum, region, on mid- A origin; dorsolaterally below Ad; near patch peritoneum; Procurrent: Transjormation-Spreading dorsally & laterally Upper trunk. on laterally tail. dorsum & along Juvenile-Body Lower iridescent. becoming Gill rakers: spines; on ridges bony Preopercular features: Diagnostic pigment head; 2--{) pigment blotches peritoneal origin; A absent. near Upper 16-24 Lower 7-8 MORPHOMETRICS %) in mean & (range Branchiostegals HISTORY LIFE PrF Y-S PoF F Tr Juv tropical & Temperate Range: Pacific eastern the in worldwide; waters Sn-NBL 45-49 43-78 62-81 60-66 37 47 62 59 68 Chile to Islands Aleutian from the BDIBL 18-36 16-20 19-26 26-35 to surface Near depth) Habitat: m (1829 bathypelagic 17 18 28 23 30 29-33 HLlBL 31-35 31-43 35-40 Spawning season: 31 24 37 33 37 39--{)4 HWIHL 45-59 38-55 40-46 larvae ELH pattern: Oviparous; planktonic 52 42 50 51 48 SnLIHL 43-48 35-51 36-43 31-38 LITERATURE 32 44 43 40 35 18-23x EDIHL* 24-31 x x 27-33 23-34x Okiyama 1988f 1984a, 16-29 19-35 23-34 28-33 (Illustrator) ILLUSTRATIONS ORIGINAL 27x27 20x23 29x37 30x30 27x28 12-l3 9-24 PILlBL 21-27 20-25 5.2 Pretlexion larva, mm (N. Arthur) 11 12 14 22 24 (N. Arthur) mm Flexion 8.4 larva, L1BL P 0-3 0-0 2-5 6-8 2 specimen, Transformation Arthur) mm 20.5 (N. 0 0 0.4 7 4 stage; Eye off-round through mid-postflexion * horizontal axis given first, second. vertical 380

393 /el'ox Alepisaul'us Longnose lancetfish mm 5.2 mm 6.4 mm 8.4 \ ... \ ... . ·Ilt':~[ftft/f.;;';'·~··········· 10.0 mm 20.5 mm Figure Alepisauridae I. Preflexion larva, 5.2 mm (CaICOFI 7205, station 20.135); flexion larvae, 6.4 mm (Okiyama 1988±), 8.4 mm (Okiyama (CaICOFI station 27.135); postflexion larva, 10.0 mm 7210, 1984a); transformation specimen, 20.5 mm (SIO 70-310). 381

394 MYCTOPHIFORMES: Lanternfishes H. G. MOSER Neoscopelidae includes two families, Myctophiformes ctenosquamates (Myctophiformes + Acanthomorpha), with a total Myctophidae, and and >240 35 of genera based on numerous synapomorphies, of which the 1994). (Nelson Hulley 1994; All species are or pelagic of the gill arches is the most reductive restructuring benthopelagic that occupy fishes deep-sea habitats. The important (Johnson unequivocal place that 1992). Neoscopelidae is a small family (six three in species evolution has attracted myctophiforms occupy in teleost genera) consisting of photophore-bearing (e.g., species in conflicting resulting hypothe- research much interest, benthopelagic Neoscopelus) that inhabit continental 1985; (Rosen confusion considerable and ses Stiassny slope waters and structurally reduced forms (Scope- Johnson 1986). characters used assessed (1992) the in lengys), lacking photophores, that inhabit deep oceanic previous for hypotheses and argued cogently cteno- are midwaters. a highly diverse and Myctophids monophyly squamate based on loss the of the fifth speciose of species) (>240 primarily luminous, fatriily and toothplate pharyngobranchial internal third the oceans. all midwater, fishes that inhabit levator in muscle myctophiforms and in acantho- morphs. Monophyly is more of Myctophiformes Prior to Rosen's (1973) analysis ofthe interrelation- hypothesis (1973) Rosen's although problematic was ships consid- of euteleost fishes, Myctophiformes was corroborated Stiassny by con- tentatively and (1986) aulopiforms included that group the a much ered larger firmed (1992). Johnson by to them (Gosline 1971). Rosen be (1973) considered Families included: Neoscopelidae Myctophidae 382

395 NEOSCOPELIDAE: Blackchins G. H. MOSER Neoscopelidae consists species in of The family six Neoscopelids but oviparous be to assumed are three genera. eastern in is found the Only Scopelengys planktonic eggs have not been identified. The larvae Pacific. in regularly taken are tristis Scopelengys Adult deep bodied and robust with a somewhat massive are and that has a strong sigmoid flexure. The head gut north as southern far Califor- as deep midwater trawls jaws teeth are large; the jaws bear strong sharp that are tropical eastern the in found larvae however, nia; are anteriorly. enlarged to fins first the are pectorals The captured are Pacific the southern- only rarely in and most part second A area. survey CalCOFI of the rays become and elongate, extending posteriad develop species both of Scopelengys Larvae anus. beyond the water the inhabits clarkei, S. central nominal species, the over blotch pigment a large have grit. survey CalCOFI area (Butler and of S. Larvae the of west mass and Ahlstrom 1976). Bekker Shcherbachev the a stripe through clarkei tristis (1990) S. while eye develop an lack larvae pigmented are but stripe eye on the Scopelengys nominal the of status the that suggested lower the on and nape and head jaw posteriorly species needs review. 1988e; Butler (Okiyama 1974, 1984a, and Ahlstrom S. clarkei is deeper caudal peduncle the Also, 1976). in «30 cm) Neoscopelids in size medium to small are tristis (Butler and Ahlstrom 1976). S. compared to with Jaws are large, and head compressed a body. of of S. species tristis some of larvae resemble Larvae bear of villiform the and orbit, extend back the to the a more have Lampanyctus genus myctophid but small are eyes The teeth. in large and Scopelengys in placement posteriad a more fin, of the anal massive Scopelengys Neoscopelus. cov- body elongate an has the lack and gut, Br2 larval in present photophore highly deciduous with cycloid scales. ered large, myctophids. Larvae Neoscopelus those from of differ rows ventrolateral has Neoscopelus photophores of and snout, of Scopelengys in having a relatively shorter Scopelengys luminous tongue; the lacks on light organs longer preopercular have and fins pectoral smaller gut, organs. above lies Fins are large; the dorsal fin origin spination 1988e). (Okiyama the pelvic fin and the anal is well fin origin behind the pectorals dorsal fin insertion. The large extend posteri- The description following on is based tristis S. of An ad fin the is anus to or anal fin origin. adipose literature the Okiyama 1976; Ahlstrom and (Butler in present. The benthopelagic Neoscopelus is silvery of detailed examination 14 on and 1988e) 1984a, 1974, and with musculature, well ossified has coloration, firm mm; 3.7-5.6 preflexion, (5 6.9-7.7 3 flexion, larvae bathypelagic In a large gas bladder. contrast, the (24.8 2 juveniles and mm) 8.0-16.4 6 postflexion, mm; a gas . Scopelengys is weakly ossified, flabby, and lacks and informa- ecological and data Meristic mm). 27.5 and bladder (Fitch and Lavenberg 1968; Butler Ahls- (Fitch tion were obtained from the literature and 1984). The trom 1976; Nafpaktitis 1977; Hulley 1976; Lavenberg 1968; Butler and Ahlstrom ctenoid scales, is known monotypic Solivomer, with observations Nafpaktitis 1977; Hulley 1984) and from the Philippines. only from made during this study. 383

396 NEOSCOPELIDAE Scope/engys tristis LIFE DESCRIPTION mSTORY EARLY MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: 31 29-32 No.ofOG: Diam.ofOG: Total 12-13 13 Shell surface: Precaudal 18 17-19 Pigment: Caudal features: Diagnostic Fins: 0 o Dorsal spines 11-13 11 LARVAE Dorsal rays o 0 Hatching length: Anal spines 12-14 mm ca. length: Flexion 6.0-8.0 rays Anal 13 17 mm Transformation length: 8 > Pelvic 8 15 14-17 , D, Pectoral C PI' , development Fin P C A, sequence: I 2 2 have larvae Caudal: & gut above PrejIexion--Smallest a blotch Pigmentation: 10+9 10+9 median embedded 7 in ca. a postanal an mm, 4.6 by series; ventral Principal linear blotch in snout an give the post-orbital blotch & embedded Procurrent: 6-9 Upper eye. through a streak of Flexion-pos(jlexion-Postanal appearance 7-8 to reduced series Lower or elongate. becomes gut above blotch absent; 1-5, gut moderate Body deep & robust; features: Diagnostic in length, rakers: Gill 1-2 sigmoid; strongly robust, head needle-like, teeth large; jaws & larger Upper 6-9 & round of Lower eyes jaws; small; PI forms early in preflexion stage tip at 8 similar 8 becomes lacks clarkei S. through eye; streak pigment large; & Branchiostegals posterior mandibular, has after & pigment nape & head, stripe eye stage. flexion LIFE HISTORY mean in MORPHOMETRICS & (range %) Indian, Pacific, Tropical Range: central from Oceans; Atlantic & California to eastern the in Pacific Chile Y-S PrF PoF F Tr Juv Habitat: Bathypelagic Sn-AlBL 61-61 48-56 60-66 62--65 64 53 61 63 Larvae Spawning season: were captured throughout the year on cruises EASTROPAC 19-21 22-23 BDIBL 20-24 19-20 21 23 22 20 planktonic Oviparous; pattern: ELH larvae 27-32 33-35 HUBL 32-33 27-31 29 32 34 29 LITERATURE 38-44 HWIHL 44-54 40-45 39-39 41 49 42 39 & Ahlstrom 1976 Butler SnLIHL 32-34 26-36 36-41 36-36 Okiyama 1988e 1974, 1984a, 33 30 38 36 24-28 EDIHL 20-21 14-19 13-13 ORIGINAL (Illustrator) ILLUSTRATIONS 26 17 20 13 25-34 14-25 36-40 PILlBL Preflexion larva, 3.7 mm (N. Arthur) * 19 38 30 mm (N. Arthur) 5.6 larva, Flexion 0-0 L1BL P 2-9 0-2 2 t 0 1 5 broken. rays PI * rays P t broken. 2 384

397 Scopelengys tristis Pacific blackchin 3.7mm 5.6mm 13.9 mm Figure NeoscopeJidae 1. Preflexion larva, 3.7 mm (EASTROPAC II, station 47.005); flexion larva, 5.6 mm (EASTROPAC II, station 47.040); postflexion larva, 13.9 mm (Butler and Ahlstrom 1976). 385

398

399 MYCTOPHIDAE: Lanternfishes l AHLSTROM G. H. H. MOSER AND E. of body. consisting structures complex are Photophores in fishes ubiquitous most the are Lanternfishes the a modified cup-like scale containing photogenic tissue and world ocean may have the greatest biomass of aU of overlain by a scale modified as a lens. Photophores Hulley their (1994) total oceanic fishes. suggested structure on arranged similar the head. Myctophid are this tons million but is likely a 600 exceeds biomass have fundamental pattern a photophores (Figure least estimate. conservative There are at 80 species but most species 1) some (to genera and Myctophidae Current California the in genera 23 and representing have the within degree) basic a unique arrangement We have identified waters (Wisner 1976). adjacent This has pattern. to hypothesis lead photophore that the aU 23 larvae of approximately 50 species representing Other intraspecific recognition. pattern may function in Mycto- northeastern in genera the Pacific (Table organs are: luminous of small kinds photo- secondary among are larvae Lanternfish 1). most the phidae phores on the head and body, supra- and infracaudal abundant encountered larvae in CalCOFI samples; fonn various of and glands dimorphic) sexually (often Triphoturus mexicanus and Stenobrachius leucopsarus associated specialized complexity, the with photophores 11 and rank the of sixth and abundance in seventh total on scales or patches luminous eyes, and bases the of in mycto- are series time the taxa abundant most 30 et elsewhere 1977; and al. on the body (Nafpaktitis fins phids. HuHey 1994). Myctophids are cm) small to medium-size (3-35 Almost aU myctophids undergo did vertical migra- body fishes deep-sea and head, a compressed with tions, probably associated foraging with on planktonic of large eyes, and to large moderate with bands jaws lanternfishes many night, At crustaceans. migrate most in is terminal mouth teeth. set closely small, The of to from daytime depths upward layer mixed the completely from excluded is maxillary and the species species where surface 300-2000 m. Some the come to gape. There the a single is an by fonowed fin dorsal may be captured by neuston nets. they dipnetted or supported a cartilaginous fin plate. The anal by adipose to tend species Deep-living undergo or no vertical little fin the to posterior dorsal slightly is under or origin fin of some For and migration. degree the species, pattern rays eight have and abdominal are fins pelvic the base; migration for juveniles and adults different vertical is range Pectoral fins from large and in most species. Larvae 1994). HuHey 1977; al. of (Nafpaktitis et weB to developed and or even formed small weakly generally found in the upper mixed are myctophids at absent in some species. There is a rudimentary spine Myctophinae of the subfamily however, larvae layer; base the the of the ray, anal first the ray, dorsal first 500 some m for species) have deeper distributions (to the and ray, pectoral upper ray. pelvic outermost Color than do those and of the Lampanyctinae Nichols (Loeb live or green, iridescent blue, from ranges specimens of 1993). Smith and Moser 1984; or species shallow-living in silver to dark brown black in species. The body is covered with deep-living an have are presumably oviparous Myctophids and scales; ctenoid have species a few cycloid rounded are their planktonic eggs eggs; however, planktonic in bladder juveniles but may is scales. A present gas between coHected infrequently. The great disparity the invested or reduced become farty adults. in tissue with of planktonic eggs and high larval apparent paucity Lanternfishes commercially harvested only off are of disintegration by explained be may abundance the their sub-antarctic; the in and Africa South however, during eggs the as us by identified Eggs capture. enonnous biomass may mark them for much greater in myctophids diameter from ca. 0.70 mm to range exploitation in the future (Nafpaktitis et al. commercial diameter, moderately a yolk, segmented have 0.90 et 1977; Hulley al. 1994; Paxton 1995). 0.1- large perivitelline a single oil globule (ca. space, mm It chorion. wrinkled a fragile, and 0.3 diameter), of luminous organs, the Lanternfishes have a variety during the tow is that the likely thin chorion is broken of which are the rows or groups paired prominent most lateral regions of the disintegrated is of and on the ventral and photophores embryo the subsequently and 1 Formerly, at National Service, Fisheries Marine La Laboratory, Jolla California. Southwest Fisheries Science Center, La Jolla, 387

400 passed meshes. disintegration the through Similarly, of Punta the part, southern south in member common extrusion and of yolk-sac explain could their larvae a number of however, Sur; California Baja Eugenia, near absence from the samples. are reported from the region (see Berry species other and Perkins 1966; Wisner 1976; Hubbs et al. 1979; Larvae taxo- of lanternfishes are among most the are: D. andersoni, D. these Among 1983). Bekker nomically and studied extensively tractable of fish all fulgens, D. garmani, is trachops, and a form that D. at hatch They larvae. 2.0 in size at ca. mm and range perspicillatuslmollislfulgenslrafenesquii D. the of part from transformation depending on mm, 30 to 10 the pers. (B. complex princi- The comm.). species Mundy, species. array a vast have larvae Myctophid of morpho- pal distributions most these species are to the of of logical and pigment characters that identification permit south the to or west of the of the California core useful analyses of systematic in of are and species Current where to a lesser extent, D. and, theta D. et al. 1984a; Paxton et and subfamilies genera (Moser of pacificus are predominant. Our identifications larval for al. 1984). Head, gut, and distinctive are shape body (a D. (a pacificus D. and form) stout slender theta and most a recognizable have genera species morpho are based abundance, relative distribution, on form) moderately are body species most slender, Although and extensive transformation Identification series. of shape markedly robust to attenuate highly from ranges Diaphus the of peripheral from localities larvae are Eyes and elliptical deep-bodied or in compressed. survey CalCOFI discovery of the awaits area distin- Myctophinae round or nearly round in most the and resolution guishing larval characters and the of taxo- Lampanyctinae. Many the narrow-eyed myctophine of within genus. the nomic problems choroid a well developed mass of have species tissue surface on the ventral and eye several genera of the detailed The following descriptions on based are gut the Typically, eyes. stalked have slightly sig- is observations and species 41 of specimens 11--46 of on the has extends midbody, to and distinctive moid, published (Table Myctophi- where applicable literature however, gut length can transverse mucosal folds; dae are identified other eight of Larvae 1). species (preflexion range from extremely short Lampanyctus) or the west to rarely south captured of the regular elongate and trailing free from the body (Mycto- to pattern are CalCOFI and sampling not described aurolaternatum). phum be may pectoral The fms large shown however, are stages separately; larval single in higher have species some shaped; a and distinctly Figure 2 (Benthosema Myctophidae fibulatum, larvae in count ray pectoral than in some and adults M and Lampadena lychnobium, Myctophum luminosa, rays. have The elongate, lower pectoral ornamented asperum, 3 Myctophidae Figure in and spinosum) (M fin pelvic the usually is is it although form, to last selen- M and obtusirostrum, M M brachygnathum, in some precocious Usually, median the species. finfold Meristic data were obtained from several litera- ops). Tarletonbeania is well developed and in Loweina and (Moser sources ture and 1976; Wisner 1970; Ahlstrom is it photo- Br2 the genera, but two all In voluminous. Zahuranec et al. 1984a) and from counts Moser 1980; develops the larval period and in during phore many made information was during Ecological study. this genera in Lampanyctinae) in 11 and Myctophinae (3 obtained Wisner Nafpaktitis et (1976), from primarily develop other photophores during the larval period. (1994). al. and (1977), Hulley genus for the large Except most Diaphus, have species melanophore a unique that allows their identifi- pattern revised those species of Lam- (1980) Zahuranec pattern a recurring and cation of be loci can pigment with reduced fins or absent pectoral and panyctus for recognized 1984a). al. most (Moser genera et placed 17 species in Nannobrachium, a generic name Identification proven to of has species Diaphus larval ! previously by used Larvae (1887). GUnther I five of forms extremely difficult. Two be of Diaphus larvae Nannobrachium and guide this in described are species have been described (Moser et al. 1984a): a slender two science belong these of to species new to (Zahura- form with numerous persistent postanal ventral melano- referred two are species These press). in nec, by to postanal fewer phores and a stout form with melano- descriptive names "no and pectorals" (Nannobrachium two these Within flexion. before coalesce that phores Nannobrachium the Zahuranec's of use since "niger") few morphs, distinguishing available are characters for nomina produce would names manuscript (1980) nuda. species. member of Diaphus theta is the most common can ascertained these two species identities of be The part central and northern the genus the in the of from synonym lists in Zahuranec (1980). pacificus is CalCOFI most survey area and D. the 388

401 All Table in the California Current vicinity. species species have selectedmyctophid Meristic characters for 1. Myctophidae principal caudal fin rays. 10 superior and 9 inferior Vertebrae Fio rays A CV D PreV Total Species Lampanyctinae 12-14 15-16 6-7+6-8 18-19 11-13 32-35 13-15 longipes Bolinichthys 8 6-7+6-7 20-21 13-16 13-15 8 13-15 16-17 35-38 Ceratoscopelus townsendi 9-10 11-13 16-17 5-7+5-6 8 16 13-15 31-33 Diaphus pacificus 10-12 12-14 12-14 18-20 6-8+6-8 34-36 15-16 8 D. theta 8+8 20-22 14-15 13-15 15-17 15-17 35-37 8 Lampadena luminosa 13-14 14-16 36-38 20-22 14-17 16 8-9+8-9 8 urophaos L. 16-18 19-22 13-14 13-15 14-16 35-37 8-10+9-11 8 Lampanyctus acanthurus 17-19 21-23 14-15 6-7+6-7 37-39 16 13-14 8 L. nobilis 17-20 13-15 15-16 19-21 35-38 12-15 6-7+7-8 8 parvicauda L. 34-36 16-19 13-15 8 19-20 12-14 14-15 7+6-8 L. steinbecki 35-37 17-18 19-21 13-15 13-14 14-16 7-8+7-8 8 tenuiformis L. 18-20 16-18 11-13 34-35 13-15 6-7+5-6 8 15-17 Lobianchia gemellarii 16-20 10-13 8 11-14 17-19 31-34 6-8+6-8 13-15 idostigma Nannobrachium 20-23 14-16 16-19 6-8+6-8 13-16 7-8 15-16 36-38 N. "niger" 15-19 18-21 12-15 6-7+6-7 8 o 13-16 33-35 N. "no pectorals" 12-17 20-22 17-19 12-15 36-39 6-8+6-8 8 15-16 N. regalis 16-19 10-13 12-16 20-22 8 35-38 15-16 7-8+7-8 N. ritteri 27-31 16-18 10-12 12-15 12-15 6-8+6-8 6 12-13 Notolychnus valdiviae 11-13 18-20 21-22 21-24 11-14+10-14 16 8 35-38 Notoscopelus resplendens 10-13 15-18 14-17 21-22 15-16 8 35-38 7-9+7-9 Parvilux ingens 14-16 20-22 8-10 13-15 8 15-16 35-38 6-8+7-8 Stenobrachius leucopsarus 11-14 20-22 11-13 39-41 15-17 8-10+8-9 18-20 8 minimus Taaningichthys 14-17 18-19 13-16 8-10 5-7+6-7 33-35 15-17 8 mexicanus Triphoturus 13-15 17-19 16-18 5-6+5-6 33-35 8-10 8 15-16 T. nigrescens Myctophioae 14-17 18-20 19 8 12-14 31-32 8-9+7-8 13 Benthosema fibulatum 19-22 13-16 18-19 8 12-14 31-33 7-9+7-8 13-14 B. panamense 8+7-8 16-19 18-19 12-14 13-15 15 8 33-35 B. suborbitale 16-19 9-11 35-40 22-25 8 5-7+5-7 14-15 13-17 Centrobranchus nigroocellatus 18-20 8-9+8-9 10-12 14-18 12-15 31-35 8 13-14 Diogenichthys atlanticus 10-13 18-20 15-17 10-12 7-9+7-9 29-32 8 12-13 laternatus D. 6-8+6-7 18-20 17-20 12-15 13-16 8 32-34 14-16 Electrona risso 17-20 5-6+5-6 23-26 38-41 6-8 10-12 11-14 14-16 Gonichthys tenuiculus 6-9+6-9 18-21 11-14 19-21 12-15 8 35-37 15-17 atratum Hygophum 18-21 34-37 18-21 12-14 8-9+8-9 13-15 8 15-16 Hproximum 21-25 7-9+7-8 21-23 13-15 38-40 13-15 8 16-17 reinhardtii H 6-7+6-7 10-13 19-21 13-17 37-39 8 9-13 17-19 rara Loweina 13-15 19-22 10-11 16-17 8-9+8-9 35-38 8 15-17 J.fyctophum asperum 12-16 9-12 8+7-8 21-27 25-26 8 42-46 18-19 M aurolaternatum 20-21 12-13 18-19 16-18 8-9+7-9 15 8 35-36 brachygnathum M 12-14 22-25 17-20 14-17 15-16 8 8-9+8-9 37-39 lychnobium M 18-21 21-23 12-14 13-16 36-39 8 15-16 7-9+7-9 M nitidulum 17-19 19-21 12-13 15-16 17-19 obtusirostrum M 8 35-36 8-9+7-8 19-20 34-35 16-18 17-19 15-16 13 8 8+7-8 M selenops 18-20 37-40 12-15 12-14 22-24 8 15-16 8-9+8-9 M spinosum 36-38 21-23 11-13 19-24 14-16 Protomyctophum crockeri 13-17 8 7-9+7-8 22-24 21-25 14-17 11-13 37-39 14-16 thompsoni P. 8 7-9+6-8 37-40 19-21 15-20 22-23 13-15 15-17 Symbolophorus californiensis 8 8-10+7-8 36-38 21-22 13-16 18-22 13-18 15-16 evermanni S. 8 8-9+7-9 11-14 39-42 22-24 crenularis Tarletonbeania 11-15 17-20 17-18 8 5-7+5-7 389

402 Table Myctophidae Number of specimens (above) and size ranges (in mm, below) used in the preparation of the myctophid 2. descriptions. species indicates literature used in the description. An "L" Flexion Posttlexion Species Preflexion Transformation Juvenile 12 8 Lampanyctinae 7 4 4 5.0--6.4 6.7-11.7 Bolinichthys longipes 3.1-4.8 10.6-13.0 15.4-20.7 townsendi 11 Ceratoscopelus 6 13 6 5 6.2-7.8 7.8-18.3 2.6-6.1 16.6-20.1 17.7-20.3 pacificus 5 5 Diaphus 3 3 3 2.1-3.9 4.1-4.4 4.6-7.5 8.6-9.6 9.8-10.3 7 7 3 3 D. theta 3 2.9-5.4 5.6-6.4 6.9-10.5 11.0-13.7 14.7-15.9 urophaos 17 6 14 Lampadena 6 3 6.8-8.3 8.6-15.6 17.4-21.0 2.9-6.7 19.6-22.6 acanthurus 7 5 12 Lampanyctus 2 2 6.2-14.4 2.5-5.0 5.2-5.9 15.0-18.4 25.0-26.9 3 6 5 L. 0 nobilis 1 6.2-18.3 5.2-6.4 3.5-5.2 21.5 6 3 8 3 L. parvicauda 3 7.2-13.5 3.5-5.2 5.8-6.6 14.5-16.4 25.0-28.1 5 4 6 L. 0 steinbecki 0 6.1-10.0 4.5-5.8 3.0-3.9 tenuiformis 4 7 L. 3 0 0 3.2-4.2 4.3-4.9 5.0-16.1 3 Lobianchia gemellarii 5 7 3 2.9-4.5 5.0-5.6 6.2-13.5 11.8 14.7-16.2 7 9 3 idostigma Nannobrachium 0 3 2.2-5.3 5.3-6.0 6.3-11.7 20.3-22.4 N. "niger" 8 3 5 1 0 3.3-6.1 7.5-13.9 6.6-7.3 20.7 "no 3 6 N. pectorals" 6 0 0 3.0-5.3 5.6-6.6 6.5-14.4 11 15 9 regalis N. 1 2 3.4-5.8 8.1-16.4 6.0-7.9 21.2 27.4-31.3 7 4 8 3 N. ritteri 3 6.1-7.1 7.2-15.6 3.4-5.8 15.7-17.6 22.4-26.5 15 8 9 Notolychnus valdiviae 5 3 6.2-10.6 2.8-4.4 4.4-6.2 10.1-10.8 12.5-13.9 12 15 Notoscopelus 7 resplendens 1 2 5.0-6.5 6.8-19.1 2.4-4.9 21.0 26.1-26.3 ingens Parvilux 2 6 3 0 0 4.5-6.5 10.1-17.6 7.6-7.8 4 8 9 4 leucopsarus Stenobrachius 3 3.1-6.4 6.6-7.9 8.4-16.1 15.7-18.4 19.6-21.0 Taaningichthys minimus 3 3 10 0 0 8.8-20.3 7.0-8.5 4.9-6.8 7 9 Triphoturus 5 mexicanus 2 3 2.7-6.1 6.3-7.6 8.0-13.5 12.5-15.4 16.7-17.8 3 T. nigrescens 6 8 3 5.6-6.4 3.2-5.6 6.6-14.0 15.0 16.2-17.8 Myctophinae La La La La panamense Benthosema 0 4 B. suborbitale 6 5 2 4 5.2-6.5 2.6-5.0 6.6-9.6 9.7-10.8 11.0-16.1 La La La La nigroocellatus Centrobranchus 0 La La La La La Diogenichthys atlanticus La La La La D. laternatus 0 La La La La risso Electrona 0 La La La La Gonichthys tenuiculus 0 390

403 Continued. Table Myctophidae 2. Flexion Postflexion Preflexion Transfonnation Juvenile Species La La La La 0 atratum Hygophum 6 4 7 0 3 Hproximum 2.7-5.8 6.1-7.6 7.7-11.8 12.8-16.2 La La La La La H reinhardtii La La La La 0 Loweina rara 6 3 7 2 3 aurolaternatum Myctophum 11.7-13.8 15.4-25.8 3.8-10.6 27.6-29.0 25.9-28.2 La La La La 0 M nitidulum La La La La La Protomyctophum crockeri La La La La 0 P. thompsoni La La La La 0 californiensis Symbolophorus 6 3 6 1 0 S. evermanni 6.1-7.5 3.4-6.1 7.6-16.9 24.5 La La La La La crenularis Tarletonbeania Moser and Ahlstrom a 1970 SUGL Bu Suo 9 PCR ------- Ant VLO SAO -----6------1 ~ P~I ~ :l~-------~----------~ , 6 0,: """"" 0: 'i o Vn" ~ C R l1""~ ~ : ... ... : So Br Pre Op 3 2 VO PO ~ I:NG L .. ~ ~ ~ ~=--":-=-,4,--,-~ : AOp AOa : · A ·, : ... 0·· .. ... Figure Myctophidae 1. Generalized photophore pattern and terminology for Myctophidae (from Fujii 1984c). Ant-antorbital organ; Cp-cheek AG-anal anal organs; AOp-posterior anal organs; Br-branchiostegal organs; Bu-buccal organ; AOa-anterior organs; organ; organ; INGL-infracaudalluminous gland; Dn-dorsonasal organs; PLO-suprapectoral organ; PG-thoracicorgans Op-opercular or pectoral organs; Pol-posterolateral organ; Prc-precaudal organs; PYG-subpectoral organs; SAG-supraanal organs; So--suborbital organ; gland; Suo--supraorbital organ; VLG-supraventral SUGL-supracaudalluminous Yn-ventronasal organ; YG-ventral organ; organs. 391

404 8.7mm mm 12.8 9.5 mm .• 9.0mm Figure Myctophidae 2. Postflexion myctophid larvae (top to bottom): Benthosemajibulatum, 8.7 mm; Lampadena luminosa, 12.8 mm; 9.0 Myctophum 9.5 mm; Myctophum spinosum, lychnobium, mm (Moser and Ahlstrom 1974). 392

405 6.8mm mm 7.5 7.6mm 7.8mm Figure Myctophidae 3. Postflexion Myctophum larvae (top to bottom): M. asperum, 6.8 mm; M. brachygnathum, 7.5 mm; M. obtusirostrum, mm M. selenops, 7.8 (Moser and Ahlstrom 1974). 7.6 mm; 393

406 MYCTOPHIDAE longipes Bolinichthys EARLY LIFE HISTORY DESCRIPTION MERISTICS Mode Range EGGS diam.: Shell Vertebrae: Yolk: 33-34 32-35 No.ofOG: Diam.ofOG: Total 16 surface: 15-16 Shell Precaudal Pigment: 18-19 18 Caudal features: Diagnostic Fins: 0 0 Dorsal spines 12-l3 11-l3 LARVAE rays Dorsal <3.0 mm Hatching length: Anal 0 0 spines l3-15 mm 5.0-6.5 14 Flexion length: ca. rays Anal ca. 1O.5-l3.0 mm length: Transformation Pelvic 8 8 12-14 12-l3 C , D & A & Fin development sequence: , P PI & C Pectoral I 2 2 series ventral median in 4-6) Preflexion-3-9 (usually Pigmentation: Caudal: Principal extending posteriad from above preanal arch 10+9 gas 1 above gut; 10+9 of as small as larvae in region otic in Procurrent: 1 embedded 4.0 mm; bladder by 1, mm. 5.0 mm, postanal series 4.4 to Flexion-By 6-7 7 coalesces Upper 6-8 7 Lower caudal in 2, occasionally gut of arch preanal 1 above region; peduncle spinal column in above 1-3 of caudal series remains; embedded rakers: Gill Upper 5 4-5 as Posiflexion-Alllarvae peduncle region in larvae mm. small as 5.3 12-15 l3-14 spinal >7.0 mm 2 or 1 2-3); (usually 1-9 column, have above series Lower 9 9 Branchiostegals cerebellum at ca. midline mm. dorsal in to posterior 8.0 body; head Diagnostic features: Moderately slender size jaw & nearly round; snout LIFEmsTORY midbody moderate; eyes blunt; gut & to large diagnostic but sparse pigment sigmoid slight with curvature; (see , Ceratoscopelus Br2 photophores form townsendl); ca. 5.0 mm; Br at Pacific & Range: from 30° Eastern N to Pacific 20° S; also in l southwest larvae 0PI' PO, form during transformation; & similar to somewhat Ocean Indian in are but townsendi Ceratoscopelus of more a larger, have stouter, those round eye, have embedded pigment in in the otic region (lacking C. mesopelagic to Epi- Habitat: townsendi & early developing photophores larvae lack <10.5 mm), townsendi C. in larvae. present Spawning Larvae captured throughout season: year MORPHOMETRICS (range & mean in %) Oviparous; planktonic eggs & ELH larvae pattern: LITERATURE Y-S PrF F Tr PoF Juv Sn-AlBL 58-62 53-58 59--63 60--62 1984a al. Moser et 60 55 61 62 59 1964 Pertseva-Ostroumova BDIBL 14-18 18-23 20-24 19-25 20 15 (D1ustrator) ILLUSTRATIONS ORIGINAL 22 22 21 22-25 24-27 HUBL 27-29 26-29 mm mm, 3.1 larvae, Preflexion (G. Mattson) 4.7 25 24 27 29 28 Flexion larva, (G. mm 5.2 Mattson) HWIHL 53--66 53-62 54-62 52-62 Mattson) larva, Postflexion (G. mm 10.6 56 60 58 59 58 Juvenile, 16.6 mm (G. Mattson) l3-26 22-26 SnUHL 21-24 20 24 23 19 17 35-44x 33-38x 34-40x EDIHL* 31-35x oval; axis * Eye slightly horizontal second. vertical axis first, given is 35-46 36-40 32-35 36-43 39x41 35x38 38x41 33x33 33 4-7 6-8 9-11 PILlBL 6 10 7 20 24 2-10 0-2 L1BL 0-0 P 2 0 0.5 6 l3 12 394

407 Popeye lampfish longipes Bolinichthys mm 3.1 4.7 mm mm 5.2 10.6 mm 16.6 mm Figure Myctophidae 4. Preflexion larvae, 3.1 mm (CalCOFI 6611, station 147.60), 4.7 mm (CaICOFI 6207, station 80.160); flexion larva, 60-251); (510 station 150.80); postflexion larva, 10.6 mm 5011, juvenile, 16.6 mm (NORPAC, station 78 Deep). mm (CaICOFI 5.2 395

408 MYCTOPHIDAE Ceratoscopelus townsendi EARLY LIFE DESCRIPTION HISTORY MERISTICS Range Mode EGGS Yolk: diam.: Shell Vertebrae: No.orOG: 36 35-38 niam.ofOG: Total 16 surface: Shell 16-17 Precalldal Pigment: 20 20-21 Caudal Diagnostic features: Fins: 0 0 Dorsal spines LARVAE 13-15 14 rays Dorsal length: mm <2.2 Hatching 0 0 spines Anal 13-14 ca. 6.1-7.8 mm Flexion length: 13-16 rays Anal mm 17-20 ca. Transformation length: 8 8 Pelvic 13-14 & C Fin sequence: C development , A, D 13-15 PI' P , Pectoral I 2 2 ca. (ave. dashes Preflexion---6-11 Pigmelltation: median in postanal 8) Caudal: series; developing gas bladder by ventral above pair mm; 10+9 10+9 5.0 on Principal terminal section Pl"ocurrent: to coalesces series Flexion-Postanal gut. of at A I stage; series 6 or 2 (usually 1-4 of of embedded 3) end by insertion 6-7 Upper 6-7 6 caudal region. above vertebral column in peduncle Posiflexion- Lower some larvae >10.5 mm (present in most in Embedded otic region in rakers: Gill! 4 4--5 medially some >16.0 mm); I above hindbrain in larvae larvae Upper mm. >12.5 10-12 11 Lower 9 slender ca. Diagnostic gut to features: Moderately midbody, body; 9 Branchiostegals eyes relatively head sigmoid; slightly small; elliptical, slightly large, with sliver lunate of choroid tissue in larvae >5.0 mm; snout acute, LIFE HISTORY at form photophores Br2 mm; larvae in blunted >14.5 becoming mm, Vn at 7.8 mm, & at 8.7 mm, PO, at 9.0 mm; upper OP 7.0 PLO Atlantic, of waters temperate to warm in distributed Widely Range: larvae to Bolinichthys similar transformation; during first I form PO Oceans Indian & Pacific smaller a eyes, narrower & but relatively with slender, more longipes acute snout, more (lacking photophores larval distinct & B. in to mesopelagic Habitat: Epi- anterior on pigment lacks longipes); region of gut mid-lateral on & margin ventral of gut (present in Diaphus theta, Stenobrachius the throughout season: Larvae Spawning occur area; CalCOH in year hypural lacks Triphoturus); & leucopsarus, in (present melanophore highest abundance in a peak with summer, the in is August Diaphus). larvae & eggs planktonic Oviparous; pattern: ELH %) (range & mean in MORPHOMETRICS LITERATURE Y-S PrF F PoF Tr Juv 1992 al. et Amaoka 55-61 40-53 Sn-AlBL 60-61 58-62 60-61 & 1988 Araujo Badcock 46 58 60 60 60 1982a Belyanina aI. 1989 Matarese et 13-16 10-14 BDIBL 14-19 19-21 18-21 14 11 17 20 20 1979 aI. Miller et Moser & 1974 Ahlstrom 23-26 16-23 24-27 HL/BL 28-31 31-33 Moser et al. 1984a 19 24 25 29 32 Ozawa 1986b, 1988d 55-74 52-58 46-58 HW/HL 41-53 45-46 Shiganova 1977 62 54 56 46 47 20-25 SnLIHL 23-25 20-22 19-27 19-21 ORIGINAL (Illustrator) ILLUSTRATIONS 23 24 20 23 21 26-34x 22-25 x 28-30x EDIHL* 23-24x 24-32x Preflexion (G. mm, 3.1 larvae, mm Mattson) 5.4 33-45 33-37 23-24 27-40 24-25 mm (G. Mattson) Flexion larva, 6.9 29x40 29x36 23x24 28x33 23x23 Postflexion Mattson) (G. 7.8 larva, mm 20.6 mm (G. Mattson) Transformation specimen, 5-8 7-8 8-15 PILIBL 25-28 t Juvenile, (G. mm 20.3 Mattson) 12 7 7 27 P 2-9 LIBL 0-1 0-0 14-14 2 t 0 0.5 5 14 second. axis vertical first, given * Eye is axis horizontal elliptical; slightly PI rays broken. t broken. t P 2 rays 396

409 Dogtooth lamp fish townsend; Ceratoscopelus mm 3.1 ... - --------.~ ..--.-...- 5.4 mm 6.9mm 7.8mm 16.6 mm 20.3 mm Figure Myctophidae 5. Preflexion larvae, 3.1 mm (CalCOFI 5908, station 93.90),5.4 mm (CalCOFI 5910, station 100.60); flexion larva, 6.9 (CalCOFI mm 6604, station 103.90); postflexion larvae, 7.8 mm (CalCOFI 6610, station 123.50), 16.6 mm (Moser and Ahlstrom 1974); transformation specimen, 20.6 mm (NORPAC, station 80); juvenile, 20.3 mm (CalCOFI 4907, station 111.58). 397

410 MYCTOPHIDAE Diaphus pacijicus* LIFE EARLY mSTORY DESCRIPTION MERISnCS Range Mode EGGS diam.: Yolk: Shell Vertebrae: 31-33 32 No.ofOG: Diam.ofOG: Total Shell surface: 16 Precaudal 16 16 Pigment: 16-17 Caudal Diagnostic features: Fins: 0 0 Dorsal spines 14 13-15 LARVAE rays Dorsal length: mm <2.0 Hatching 0 spines 0 Anal Flexion 12 length: ca. 4.0-4.5 mm 11-13 rays Anal Transformation length: ca. 8.0-10.0 rom 8 8 Pelvic 10 9-10 Pectoral , & A & D CI> P PI' C Fin sequence: development 2 2 Pre flexion-Initially, up Pigmentation: 7 dashes in postanal median to Caudal: 10+9 10+9 2.8 1 by to (coalescing series ventral region, hypural 1 at future mm), Principal 1 anteriorly on ventral margin gut of section; & a pair on terminal gut Procurrent: 2.8 by 1 embedded 5-7 6 mm, above developing Flexion- bladder. gas Upper 6 Lower to 4 in vertical series at posterior edge posiflexion-Same 5-6 up pattern; insertion. 1 at A hypural; of rakers: Gill 7 Deep Diagnostic features: becoming somewhat compressed, & stout; 6-8 Upper with sliver; choroid lunate slightly elliptical eyes large, relatively head 14 13-15 Lower notochord flexion Branchiostegals & transformation at small size; caudal melanophore of early coalescense of postanal series; melanophore genus; typical , Br gut of PO form POI> & photophores anteriorly on ventral margin LIFEmSTORY 2 s , , P0 Br mm; ca. 6.6 rom; lower PVO, P0 7.5 ca. & P0 by , by 3 2 4 3 at AOa & series, VO VLO, Op, some upper the first to form ca. from 15° to N 25° S Pacific tropical Eastern Range: transformation. Epi- Habitat: mesopelagic to MORPHOMETRICS (range & mean in %) season: year-round Possibly Spawning Y-S F PoF PrF Tr Juv ELH pattern: Oviparous; planktonic eggs & larvae 56-61 44-56 Sn-AlBL 57-64 61-64 62-65 58 60 50 64 62 LITERATURE 15-23 BDIBL 23-27 25-27 28-30 26-27 19 25 29 27 26 Moser 1974 Ahlstrom & et Moser 1984a a1. 22-26 28-32 27-30 HLlBL 31-33 30-31 24 30 29 32 31 ORIGINAL nONS ILLUSTRA (Illustrator) 59-65 HWIHL 59-63 58-60 47-56 50-53 61 62 59 52 52 Sumida (B. rom 2.8 MacCall) mm, 2.1 Preflexion larvae, 23-27 22-33 21-24 23-25 27-31 SnLIHL (B. Sumida MacCall) larva, Late flexion 4.3 rom 26 29 29 23 24 Postfiexion larva, 7.5 rom MacCall) Sumida (B. 30-32x 23-34x 30-39x 21-23x 21-23x EDlHLt Juvenile, 9.8 rom MacCall) (B. Sumida 38-44 27-39 23-24 22-24 34-38 31x36 28x32 35x40 22x23 22x24 Identification • relative based distribution, is larvae pacijicus D. of on 7-12 5-8 6-8 13-16 abundance, meristics transformation series; see & on extensive 6 15 7 9 16 family for more detailed introduction explanation. to 0-0 0.9-1 3-14 18-20 15-16 second. axis is oval; Eye slightly t horizontal given first, vertical axis 0 0.9 9 16 19 398

411 Pacific headUghtfish pacific us Diaphus 2.1 mm mm 4.3 7.5 mm 9.8mm Figure Myctophidae 6. Preflexion larvae, 2.1 mm (EASTROPAC II, station 76.01a), 2.8 mm (CaICOFI 7210, station 157.35); late flexion larva, mm 4.3 (CaICOFI 5612, station 173.G.20); postflexion larva, 7.5 mm (CalCOFI 5612, station 167.G.40);juvenile, 9.8 mm (CaICOFI 7210, station 157.35). 399

412 Diaphus theta* MYCTOPHIDAE DESCRIPTION EARLY LIFE mSTORY MERISTICS EGGS Range Mode Shell diam.: Yolk: Vertebrae: No.ofOG: Diam.ofOG: 35 34-36 Total Shell surface: 15-16 16 Precaudal Pigment: 19 18-20 Caudal features: Diagnostic Fins: 0 0 Dorsal spines LARVAE 13 12-14 Dorsal rays Hatching length: <3.0 mm 0 0 Anal spiues Flexion length: ca. 5.5-6.8 mm 13 12-14 rays Anal 11-14 ca. length: Transformation mm 8 8 Pelvic , PI' CI> 10-11 10-12 P D & A & C sequence: development Fin Pectoral 2 2 median dashes 15 to up Prej1exion-Initially, Pigmentation: in postanal Caudal: in preflexion dashes) (10-14 coalescence series, little with ventral 10+9 10+9 Principal terminal on a pair have larvae earliest larvae; the 1 in section, gut Procurrent: region region caudal (1 pairs gut lateral 2 & & 1 ca. at midgut 7 6-8 Upper anteriorly); ca. the mm, 4.4 moved by has anterior lateral gut pair 7 6-8 Lower anteroventrad; by ca. 5.2 mm, 1 present above the gas bladder. Gill rakers: coalesces anteroventral pair on gut postanal 6-7 Flexion-9-11 in series; 6-7 Upper series coalesces to ca. 1 on ventral midline. to Posiflexion-Postanal 15 14-16 Lower blotches; embedded 5-6 9 hypural posterior on series vertical in 1-4 9 Branchiostegals edge. Slender body becoming Diagnostic compressed & somewhat features: LIFE mSTORY development; head relatively small; snout relatively less slender with elliptical with lunate choroid tissue; slightly eyes relative short; eye 20° Range: Common in California Current region throughout & S to increases development; gut moderate in size with length, slightly waters westward to northern subarctic-transitional Japan coalesces melanophores gut in of on anteroventral pair sigmoid; during midline ventral persists series pigment postanal stage; flexion Habitat: to Epi- mesopelagic coalescence the in throughout early (vs. period larval deep-bodied, big-headed pacificus); 6-9 form photophores Br2 by D. by PO mm; s CalCOFI present throughout the year in Larvae area; Spawning season: 7.5 mm; , , & YO"~ PO, mm. P0 10.5 ca. by by 8.2 mm; P0 AOa, 2 3 in in peak with spring-summer, the July abimdance highest MORPHOMETRICS (range & mean in %) Oviparous; pattern: ELH eggs & larvae planktonic Y-S F PoF PrF Tr Juv LITERATURE 45-53 48-55 Sn-AlBL 56-61 61-64 60-63 al. et Matarese 1989 48 62 58 52 62 & Ahlstrom 1974 Moser Moser et al. 1984a 10-14 13-17 25-26 BDIBL 19-23 24-26 12 15 21 25 25 1964 Pertseva-Ostroumova 23-27 HUBL 15-21 21-25 27-28 27-30 (Illustrator) ILLUSTRATIONS ORIGINAL 25 19 23 28 27 HWIHL 50-64 52-57 54-59 53-67 47-54 3.2 Preflexion larva, mm, 4.4 mm MacCall) Sumida (B. 57 60 57 54 50 5.8 larva, Flexion mm (B. MacCall) Sumida 19-20 22-27 23-25 9-28 SnLIHL 18-21 larvae, (B. 8.6 mm Postflexion Sumida MacCall) mm, 7.8 21 24 23 19 19 14.7 Sumida MacCall) Juvenile, (B. mm 25-31x 25-28 x 29-32x 30-30x EDlHLt 28-32x 32-41 30-34 33-35 29-37 28-32 27x35 27x32 30x30 30x34 31x33 of D. theta larvae * based on distribution, relative Identification is transformation extensive series; on & see meristics abundance, 4-8 7-11 11-12 6-8 13-16 introduction explanation. more detailed for family to 6 15 12 7 9 axis second. t oval; horizontal slightly axis is given first, Eye vertical 0-0.6 0.6-11 12-15 0-0 16-18 0.2 0 6 14 17 400

413 California headlightfish theta Diaphus 3.2mm 4.4 mm 5.8mm 7.8 mm 8.6 mm 14.7 mm Figure Myctophidae 7. Preflexion larvae, 3.2 mm (CaICOFI, station 5908, station 87.75),4.4 mm (CaICOFI 5406, station 70.70); flexion larva, mm; postflexion larvae, 7.8 mm, 8.6 mm 5.8 (CaICOFI 4907, station 59.117); juvenile, 14.7 mm (CaICOFI 6608, station 87.70). 401

414 MYCTOPHIDAE Lampadena urophaos DESCRIPTION EARLY LIFE HISTORY MERISTICS Range Mode EGGS Shell diam.: Yolk: Vertebrae: No.ofOG: 35-38 Total 37 Diam.ofOG: Shell surface: 16 16 Precaudal Pigment: 21 20--22 Caudal Diagnostic features: Fins: 0 0 spiues Dorsal LARVAE 14--16 Dorsal rays IS <2.9 Hatching mm length: 0 0 spines Anal mm 6.8--8.3 ca. length: Flexion 13-14 13-14 Anal rays 17-21 mm ca. Transformation length: Pelvic 8 8 15-16 , D 14--17 P .. p .. A & C & sequence: Fin development C Pectoral 2 2 bladder, above I Prejlexion-Initially, gas developing Pigmentation: Caudal: midway of gut, & I on ventral I preanal arch between above margin 10+9 10+9 Principal anus 4.2 at ca. tip; notochord & the apposing midline dorsal on I mm, Procurrent: I 8-9 insertion, 8 I at future D at ca. 6.2--6.5 mm, one at ventral midline; Upper just to PI base, & 1 embedded ventrolaterally on 9 posterior embedded Lower 8--9 surface of hindbrain. Flexion-At 6.5-7.0 mm, on dorsal midline at Gill rakers: 4 3-5 on below caudal peduncle, pair & A base, & above D origin, Upper & nape 8-9 mm, at gut; above embedded midline 2 in 7.0--8.3 9-11 midline dorsal on Lower 9 anterior to Ad, pairs dorsally at Ad & D insertion, & 9--10 & at cerebellum Branchiostegals series above gut Pos!flexion-Additional 2 more in embedded midline in melanophores dorsum & postanal ventrum paired along midline & HISTORY LIFE some specimens. to gut depth; body Moderate Diagnostic features: slightly midbody, Range: North Eastern Pacific between ca. 25° & 42° N; in northwest eyes elliptical slightly large, relatively moderate; size head sigmoid; Pacific at a 10°-30° N; in central Pacific at 0°-35° N; off Australia; dense large, with pigment distinctive sliver; choroid with the occurs in subtropical Atlantic subspecies 6.5-7.2 by form photophores Br2 7.2-8.0 by PLO mm; melanophores; 8.5-9.5 mm; Vn by POI & 13.5 mm. by PO mm; s mesopelagic to Habitat: Epi- %) in mean & (range MORPHOMETRICS CalCOFl area; throughout occur Larvae season: Spawning in year the in summer the highest abundance & August a peak with fall, in Y-S PrF F PoF Tr Juv Oviparous; pattern: ELH eggs & larvae planktonic 45-63 Sn-A/BL 63--66 62-67 62--63 59--60 64 55 64 63 60 LITERATURE 12-17 21-27 23-26 BDIBL 18-22 20--21 13 20 21 24 24 1983 Fahay Matarese al. et 1989 HLlBL 27-29 17-25 27-31 27-29 29--33 & Ahlstrom 1972 Moser 21 27 28 28 30 Moser 1981 59-88 57--63 47--65 59--64 HWIHL 39--51 et 1984a al. Moser 68 61 58 62 46 19--28 17-24 21-29 SnLIHL 20--21 18-20 ILLUSTRATIONS (Dlustrator) ORIGINAL 21 22 23 21 19 29--42x 25-30x EDIHL* 31-37x 26-36x 20--25 x Mattson) Preflexion larvae, 3.7 mm, 5.6 mm (G. 32-38 35-40 26-40 25-32 20--25 (G. Mattson) mm 6.8 larva, Flexion 34x37 32x35 28x29 32x35 22x23 Mattson) (G. mm 8.8 larva, Postflexion Mattson) mm 20.9 (G. Transformation specimen, 7-8 8-12 11-14 22-23 16-17 PILlBL 17 9 7 13 22 P 13-15 0--0 0.6-1 3-10 L1BL 14--15 2 elliptical; * Eye axis horizontal slightly first, second. axis given is vertical 0 I 6 IS 13 402

415 urophaos Lampadena Sunbeam lampfish ~~--~-~~ ~~~~'. ~~ ~~~~~~~"""'---'---"--'----'---'--~'---- . -. ----------- .. '- _.- .. -----,~. mm 3.7 5.6mm ) - ... ... ~--'---- . ,-- ... . . .-'.;~-:;~:.-'.~> ';~;' --.1---..::::.::::...>_\_. _ ;~S:(.,\!~t""· ... __ -- .. -:1" .. 6.8 mm 8.8 mm ~~~4fK~ )·1r1t¥·'4{":~=][i: =~= _· ---.Y/ ' ---=---.r ~ - ~X,;~- -~c-:. l d'/://J' 13.4 mm :?0~~~ ~:{9!~ -..,--,...,...,..,,-,.,...,,~-:" ~ \ \ .. );i ~.~~ .. 1 ~~_.:c" ~~ 20.9mm Figure Myctophidae 8. Preflexion larvae, 3.7 mm (CalCOFI 5908, station 107.80), 5.6 mm (CaICOFI 5910, station 103.60); flexion larva, mm 127.80), 5909, station 120.40); postflexion larvae, 8.8 (CaICOFI (CaICOFI 6507, station 13.4 mm (Moser 1981); 6.8 mm mm (CaICOFI 5908, station 107.40). transformation specimen, 20.9 403

416 MYCTOPHIDAE Lampanyctus acanthurus LIFE HISTORY DESCRIPTION MERISnCS EARLY EGGS Range Mode Shell diam.: Vertebrae: Yolk: Total 35-37 No.orOG: Diam.o(OG: 36 surface: 14-16 Precaudal 15 Shell Pigment: 21 19-22 Caudal features: Diagnostic Fins: spines Dorsal 0 0 13-14 13 LARVAE rays Dorsal Hatching length: <2.5 mm Anal spines 0 0 rays 5.0-6.0 mm Anal 17 16-18 ca. length: Flexion mm length: ca. 15.0-18.4 Transformation Pelvic 8 8 13-15 14-15 Pectoral & P2' Cl> sequence: development Fin & PI C & A D 2 flexion-l Calldal: Pre section gut ofterrninal surface dorsal on Pigmentation: 10+9 10+9 3.5 Principal at 1 & bladder gas developing above 1 mm, smallest larvae; by in of 1 in have specimens current: Pro tip midline above lower jaw; by 4.5 mm, most Upper 8-10 9 Ad Flexion- 1 anterior to region, & 1 at tip. jaw upper cerebellum, Lower 9-11 10 >8.0 mm have larvae 1 embedded most postjlexion-As above; at Gill rakers: upper edge near cleithrum trunk myosepta outlined of PI base; anterior 5 ca. by mm. 5-6 Upper 16.0 11 10-12 Lower large armed with with jaws, featllres: Diagnostic Large, head deep Branchiostegals at body eyes; round large tips; jaw teeth largest teeth; curved strong, tapered & compressed initially, becoming somewhat robust in HISTORY LIFE short postflexion stage; gut pigment & sparse sharply initially; sigmoid 7.0 at form procurrent Br2 photophores ca. high mm; but diagnostic; C ray count (8-10+9-11). N, 35° Range: North Pacific central water east mass between ca. 20° & the margin to western of Current California the in MORPHOMETRICS (range & mean %) Habitat: Mesopelagic Y-S PrF f PoF Juv Tr year Spawning season: Larvae captured throughout Sn-AJBL 40-60 64-67 61-68 55-60 55-56 52 66 65 58 56 ELH & eggs planktonic Oviparous; pattern: larvae 15-26 BDIBL 25-31 25-29 19-25 17-19 LITERATURE 22 28 28 22 18 HLlBL 23-33 33-38 28-36 30-33 27-28 29 35 32 32 27 46-53 HW/HL 50-67 45-57 36-44 42-43 57 51 49 40 42 ILLUSTRATIONS ORIGINAL (Illustrator) SnLlHL 30-43 36-38 16-18 20-24 29-36 37 37 33 22 17 Sumida 4.2 mm, 3.2 larvae, Preflexion MacCall) mm (B. 34-46 EDIHL 32-37 24-27 27-35 21-22 Flexion larva, Sumida 5.2 mm (B. MacCall) 37 34 32 25 22 mm MacCall) Sumida Postflexion larva, 10.8 (S. specimen, Transformation Sumida MacCall) mm 25.0 (B. PILlBL 6-8 9-10 9-13 7 10 11 15 35 P LIBL 0-1 5-10 11-19 9-13 2 * 2 rays transformation P * specimens. in broken 0.2 8 16 11 404

417 acanthurus Lampanyctus SpinytaH lampfish mm 3.2 5.2 mm 10.8 mm 25.0 mm Figure Myctophidae 9. Preflexion larvae, 3.2 mm (Southtow, station 13-76),4.2 mm (Tasaday XI, station A2); flexion larva, 5.2 mm (Tasaday station A2); postflexion larva, XI, 10.8 mm (Southtow, station A4); transformation specimen, 25.0 mm (Tasaday XI, station A2). 405

418 MYCTOPHIDAE Lampanyctus nobilis LIFE mSTORY DESCRIPTION EARLY MERISTICS Mode Range EGGS Shell diam.: Vertebrae: Yolk: 38 37-39 No.ofOG: Total Diam.ofOG: 16 Shell surface: 16 Precaudal Pigment: 21 21-23 Caudal Diagnostic features: Fins: spines 0 0 Dorsal LARVAE Dorsal 14--15 14--15 rays <3.5 0 Hatching 0 mm length: spines Anal 17-19 5.0-6.5 mm 18 length: Flexion ca. rays Anal 8 8 20 mm Transformation length: ca. Pelvic 13-14 14 Fin Ph &~, A Pectoral , D P C sequence: development & I 2 anterior 1 & above Preflexion-posiflexion-l Caudal: to Pigmentation: 10+9 10+9 at lower forebrain; 1 above cerebellum; Principal jaw symphysis; on midline of in Procurrent: gular region between urohyal & symphysis, usually 1 larvae 6 6--7 mm, 2--4 in larger larvae; blotch above gas bladder; 1 embedded <4.0 Upper 12 to mass; beginning at ca. gut mm, series on medially anterior 7 6--7 Lower trunk of entire rakers: P to myosepta anterior myosepta larvae, late ; in Gill 2 between outlined bases. & P PI 3--4 3 Upper 2 10 Lower 9--11 round nearly large, head; features: Diagnostic large jaws Large eyes; Branchiostegals teeth with prominent patch tooth & of upper tip at jaw; gut short, early sigmoid; sharply tapered in body larvae, robust in late larvae; gular distinctive pigment pattern with median series; photophores HISTORY LIFE Br2 form at ca. 10 mm. Ocean; extreme except tropics in Circumglobal Range: at Pacific eastern (range & mean in %) MORPHOMETRICS area CalCOFI survey of western margin Epi- mesopelagic to Y-S Habitat: PrF PoF F Tr Juv Sn-AlBL year scason: Larvae captured throughout Spawning 43-54 58--61 57--64 50 59 62 55 Oviparous; pattern: ELH planktonic eggs & larvae BD/BL 21-25 23-28 26--32 23 26 30 20 LITERATURE 29--33 HLlBL 33-34 2&-39 31 34 35 27 Miller et al. 1979 HWIHL 41-58 42--44 40--50 1984a al. et Moser 43 49 45 41 SnLIHL 22--40 23-37 36--44 (Illustrator) ORIGINAL ILLUSTRATIONS 36 39 31 22 29--36x EDIHL* 31-33x 25-34x (N. 3.5 larva, Preflexion mm Arthur) 32-35 35--41 25-37 Arthur) mm (N. Flexion larva, 6.2 34x39 32x34 30x32 22 9.6 mm (N. Arthur) Postflexion larva, 18.3 specimen, Transformation Arthur) mm (N. 4--10 PILlBL 6--8 10--11 7 7 11 13 P L1BL 0--0 0--0 0.3-8 2 Eye * second. vertical axis first, is given axis horizontal elliptical; slightly 0 0 5 12 406

419 Noble lampfish nobilis Lampanyctus \, "" .. i ". ... ,"' . 3.5 mm .. -~,--- .. 6.2mm 9.6mm 18.3 mm Figure Myctophidae 10. Preflexion larva, 3.5 mm (Cato II, station A 7); flexion larva, 6.2 mm (Clarke-Hawaii, station 71-6-17); postflexion larva, mm (Moser 9.6 et al. 1984a); transformation specimen, 18.3 mm (Clarke-Hawaii, station 71-6-5). 407

420 Lampanyclus parvicauda MYCTOPHIDAE DESCRIPTION EARLY LIFE HISTORY MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: No.ofOG: Diam.ofOG: 36--37 35-38 Total surface: Shell 15-16 16 Precaudal 19-20 Pigment: 19-21 Caudal features: Diagnostic Fins: 0 0 spines Dorsal LARVAE 14 13-15 Dorsal rays length: Hatching mm <3.0 0 0 Anal spines 5.7-7.0 mm Flexion ca. length: 17-20 18 Anal rays 14.5-16.5 mm ca. length: Transformation 8 8 Pelvic & 13 , Fin C PI> 12-15 A P D C!, sequence: development & Pectoral 2 2 Pigmentation: Preflexion-posiflexion-Initially, 1 on ventral midline Caudal: a pair cIeithrum, to posterior just gut of ca. 10+9 gut section, on terminal 10+9 Principal midline 1 in mm, 4.0 ca. 8 series; midline in postanal ventral by Procurrent: forebrain, to each lobe to anterior 1 posterior of at 1 cerebellum, 7 6--7 Upper gas dorsum where Ad will form, at lower jaw symphysis, above 7-8 7 Lower 1 at postanal only & 2-5 in postanal ventral ventral series; bladder, Gill rakers: base Ad flexion postflexion larvae. & in at it 1 apposing with midline 3 3-4 Upper 11 mm, trunk at posiflexion-Beginning to anterior Late ca. myosepta 10 9-11 Lower ventrally & dorsad. progressing become outlined, initially P Branchiostegals 2 In features: Diagnostic early preflexion larvae the gut than is straighter in other species of the genus; body highly compressed, becoming LIFE HISTORY of a in late postflexion stage; snout somewhat blunt; presence robust in distinct pattern pigment genus; for unusual series pigment postanal to California Eastern Pacific from ca. 20° N Range: 15° S & the Gulf of stages; flexion-postflexion eye slightly elliptical with lunate choroid sliver; photophores 11.0 form at ca. mm. Br2 Habitat: Mesopelagic %) in mean & (range MORPHOMETRICS Larvae throughout captured season: Spawning year Y-S PrF F PoF eggs & ELH pattern: Oviparous; Juv planktonic Tr larvae Sn-AlBL 40--54 55-59 59-63 55-57 56--60 LITERATURE 44 57 fil 56 59 21-26 11-22 Moser al. et 1984a 26--31 23-26 BDIBL 18-20 16 23 24 28 19 ILLUSTRATIONS ORIGINAL (Illustrator) 17-26 28-29 HLlBL 31-35 28-31 28-30 21 28 30 33 29 mm (N. Arthur) Preflexion 4.2 3.7 mm, larvae, HWIHL 42-44 40--55 41-42 45--60 44-45 5.7 Flexion larva, mm (N. Arthur) 42 46 43 53 44 (N. Arthur) mm 15.0 Transformation specimen, 23-32 31-35 26--28 SnLIHL 29-36 20--23 Arthur) mm (N. 25.0 Juvenile, 26 33 34 27 22 31-38x 28-3 Ox 20--28x EDIHL* 22-24 18-20 23-29 34-40 29-33 second. vertical first, is given axis horizontal oval; axis slightly * Eye 34x37 29x31 24x25 23 19 8-9 10--13 4-8 8-10 9-23 P1LlBL 6 8 11 16 9 P L1BL 0--0 0--0.6 2-9 8-10 6--12 2 0 0.2 5 9 9 408

421 parvicauda Lampanyctus Slimtail lampfish GiE~;:;~0~:E~,: 3.7mm ~~jI 4.2 mm S.7mm 7.5 mm 25.0 mm Figure Myctophidae II. Preflexion larvae, 3.7 mm (CaICOFI 7201, station 137.35), 4.2 mm (CaICOFI 7202, station 150.55); flexion larva, 5.7 mm (EASTROPAC II, station OP.023); postflexion larva, 7.5 mm (Moser et ai. 1984a); transformation specimen, 15.0 mm (Scope, station 9-2-Cl); juvenile, 25.0 mm (CaICOFI 5205, station 137.50). 409

422 MYCTOPHIDAE Lampanyctus steinbecki* MERISnCS EARLY LIFE HISTORY DESCRIPTION Mode Range EGGS Vertebrae: diam.: Shell Yolk: Total 34-36 35 No.ofOG: Diam.ofOG: Precaudal 15 14-15 surface: Shell Caudal 19-20 Pigment: 20 Fins: Diagnostic features: Dorsal 0 spines 0 12-14 12-13 Dorsal rays LARVAE spines Anal 0 0 length: Hatching mm <3.0 rays 17-18 16-19 Anal mm ca. Flexion length: 4.0-6.0 Pelvic 8 8 20 Transformation mm ca. length: Pectoral 13-15 14 C" Fin development sequence: C PI' D & A, P & 2 2 Caudal: upper flexion-Embedded Pigmentation: Pre in otic region; at & lower 10+9 10+9 Principal lateral midline 2 at above PI base; on inner surface of jaw 1 or tips; Procurrent: on above gas bladder; by 3.7 mm, trunk; ventral base PI & adjacent Upper 7 7 terminal gut section; on some PI midline below gut, on opercIe, on 7 Lower 6-8 Flexion-Blotch bases. basibranchial rays embedded region; at near rakers: Gill usually in preopercular region; series; lateral midline 2-3 in embedded 3-4 3-4 Upper Posiflexion-Lateral stage. flexion series above spinal column in late 9-11 Lower 10 spinal to basibranchial region; up to 6 in lateral midline series & 9 in Branciliostegais column series; on some. in P2 larvae Early strongly Diagnostic features: & deep-bodied tapered, HISTORY LIFE becoming & stout; highly eyes large; snout blunt; head robust & pattern basibranchial blotch that with absent in pigment distinct is tenuiformis larvae; BT2 photophores form at ca. 5.0 mm. L. putative Range: of waters subtropical Pacific; tropical eastern & Widespread in the Ocean possibly in Indian & mean in %) (range MORPHOMETRICS Habitat: Epi- to mesopelagic y-S PrF PoF F Tr Juv throughout captured Larvae season: Spawning year Sn-A/BL 43-60 62-71 60-70 52 66 Oviparous; planktonic & larvae pattern: ELH eggs 66 BDIBL 24-34 38-41 36-41 LITERATURE 28 39 40 HUBL 29-33 34-38 32-37 aI. et Moser 1984a 31 35 35 HWIHL 55-64 58-75 60-81 (Illustrator) ORIGINAL ILLUSTRATIONS 58 66 68 SnLIHL 34-42 25-33 30-37 (N. Arthur) Preflexion larva, 3.5 mm 38 34 30 5.0 mm (N. Arthur) larva, Flexion EDIHLt 30-40x 41-42x 35-48x 32-46 45-47 37-52 35x40 42x46 41x44 to a third to & L. L. species, related closely is steinbecki L. * tenuiformis & Perkins 1966; jestivus, reported to occur in the CalCOH area (Berry 8-13 6-7 14-15 species, these of 1976; Bekker 1983). The taxonomic status Wisner 7 II 15 latter the particularly is distinct larval two, unresolved. We identify two LIBL P 2-9 0-0 10-13 2 the & distribution L. steinbecki & L. tenuiformis based on as types 0 5 12 of relative abundance of adults & identifications our larvae. Confirmation & collection of complete transformation series the taxonomic awaits species of the complex. resolution second. t Eye slightly oval; horizontal axis is given first, vertical axis 410

423 Longtin lampfish Lampanyctus steinbecki 3.5 mm 5.0 mm 6.6 mm Figure Myctophidae 12. Preflexion larva, 3.5 mm (CaICOFI 6207, station 90.150); flexion larva, 5.0 mm (CaICOFI 6207, station 90. 120); postflexion larva, 6.6 mm (Moser et al. 1984a). 411

424 MYCTOPHIDAE Lampanyctus tenuiformis* EARLY LIFE HISTORY DESCRIPTION MERISTICS Range EGGS Mode diam.: Shell Yolk: Vertebrae: 36 No.ofOG: Diam.ofOG: Total 35-37 15 14-16 Shell surface: Precaudal 20 Pigment: 19-21 Caudal features: Diagnostic Fins: 0 0 spines Dorsal 13-14 LARVAE 13-14 rays Dorsal mm <3.0 Hatching length: 0 Anal spines 0 17-18 length: ca. 4.2-5.0 mm Flexion 17-18 Anal rays mm Transformation <20 length: 8 8 Pelvic 14 development Fin C , D sequence: 13-15 PI' P , C Pectoral & A & I 2 2 Pigmentation: L. steinbecki. Flexion-Similar Pre flexion-Similar to Caudal: 10+9 but in 1 on only usually region, basibranchial none 10+9 steinbecki L. to Principal 1 on upper midline, lateral I (large) on PI rays; finfold just anterior Procurrent: 7 to anus. & none masked; embedded pigment Posiflexion-Most 7-8 above Upper 7 7-8 spinal column. Lower & tapered, strongly deep-bodied larvae Early features: Diagnostic rakers: Gill 4-5 4 body & stout large; less eye & blunt; snout head becoming stout; Upper 10-12 10 as & steinbecki; larvae not snout flexion L. in as blunt postflexion Lower spinal & column; above lack pigment region basibranchial in Branchiostegals to anus; anterior fmfold prominent in melanophore photophores Br2 mm. 5.0 ca. form at LIFE HISTORY MORPHOMETRICS (range & mean in Ufo) & circumglobal in tropics subtropics; Apparently Range: taxonomy unsettled y-s PrF F PoF Juv Tr mesopelagic Epi- Habitat: to 61-65 50-57 Sn-NBL 57-66 55 63 62 year Spawning throughout Larvae season: captured 35-40 26-31 BDIBL 24-38 planktonic eggs & larvae Oviparous; pattern: ELH 29 37 32 HLlBL 29-34 26-38 35-36 LITERATURE 32 32 35 50-58 48-64 HWIHL 48-63 54 57 55 34-39 SnLIHL 35-37 20-35 36 36 29 (Illustrator) ILLUSTRATIONS ORIGINAL 37-40x 35-48x 26-43 x EDlHLt 40-52 39-43 26-48 larva, Arthur) (N. mm 3.8 Preflexion 38x41 41x44 38x40 (N. 4.7 Arthur) mm Flexion larva, Postflexion Arthur) larva, 7.1 mm (N. 9-14 6-8 8-9 PILlBL 7 12 8 P L1BL 0-0 0.9-3 3-10 2 comments on L. tenuiformis species complex in the description of * See 0 7 2 steinbecki larvae. L. horizontal t Eye slightly oval; second. axis is given first, vertical axis 412

425 tenuiformis an"ctus 'J Lamp ... ... , . \ ... ~.\ .. 3.8 mm 4.7mm --"-'. mm 7.1 71 statIOn .. -10-7); flexion larva, . . 47mm(CaiC (Clarke- Hawa11, station 20.145); OFI7205, . larva 3.8 mm 4 129) . Myctophidae 13. pre(flC~~~FI station 2. . 7'205, Figure. 1 va 7.1 mm tfl exlOn ar , pos 413

426 MYCTOPHIDAE Lobianchia gemellarii EARLY LIFE HISTORY DESCRIPTION MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: No.ofOG: 34-35 35 Diam.ofOG: Total 16 surface: SlJell 15-17 Precaudal 19 18-20 Pigment: Caudal Diagnostic featnres: Fins: 0 0 Dorsal spines 16-18 17 LARVAE Dorsal rays ca. length: Hatching mm 2 0 spines 0 Anal 5.0-6.0 ca. mm Flexion length: 14 13-15 rays Anal 8 Transformation mm 12-14 ca. length: 8 Pelvic 12 & C A, 1'2 D C" P, sequence: development Fin 11-13 , Pectoral 2 inner lower on blotch mm, 3.0-4.0 flexion-At Pre Pigmentation: Caudal: 10+9 10+9 surface PI to anterior blotch embedded deeply blotch, PI of base base, Principal cluster on gut ventral midline posterior cleithral symphysis, pair just to Procurrent: embedded 6-7 6 & rays, PI on basally scattered gut, of terminal section on Upper at blotch above gas bladder; insertion. mm, 1 in A at midline >4.0 5-6 6 Lower Flexion-Basally Posiflexion- gut. below midline ventral on rays; A rakers: Gill at anterior on myosepta, beginning & gut region Superficial embedded Upper 4-6 gut; above region epaxial on mm, 7.0 11-15 basal on blotches two and, by Lower region PI surface inner to added some C; of some in 1'2 on base; of Brallciliostegais specimens. with eyes oval Stout body large deep, broad head, features: Diagnostic LIFE HISTORY early- 5 rays upper with PI bilobed choroid ventral lunate with sliver; forming & upper teeth rays elongate; prominent in small larvae; Br2 4 Tropical-subtropical cosmopolite Range: & 7 mm; form by 6 mm; POI photophores PO AO~, by VOl' AOa" s 11 01' & VLO, by mm. Habitat: Epi- to mesopelagic MORPHOMETRICS (range & mean ill %) season: year throughout captured Larvae Spawnillg F PrF Y-S PoF Tr luv ELM pattern: Oviparous; planktonic eggs & larvae 48-56 54-58 61-65 Sn-A/BL LITERATURE 60-62 56 52 63 61 61 1983 15-24 24-29 Fahay BDIBL 30-38 25-27 27 20 35 26 33 & Ahlstrom 1974 Moser aI. 1984a et Moser 32-33 27-33 HUBL 27-38 32-33 1988d 1986b, Ozawa 32 30 34 32 35 1964, Pertseva-Ostroumova 1974 63-71 69-72 HW/HL 48-52 57-76 Sanzo 1931b 71 51 66 68 50 Shiganova 1977 25-40 25-29 27-34 SnLIHL 21-23 1918 Hilling 28 30 35 23 22 x 32-33 21-31x 25-33x 21-23 * ED/lll., (Illustrator) ILLUSTRATIONS ORIGINAL 37-40 40-50 23-33 32x38 27x30 29x44 22 22 Arthur) Preflexion (N. mm 4.2 larva, Late Arthur) flexion larva, 5.6 mm (N. 9-13 14-21 21-30 PILIBL Transformation specimen, 11.8 mm Arthur) (N. 18 18 27 10 15 0-0.7 0-0 16-18 6-17 LIBL P2 0.2 11 17 0 16 axis vertical Eye slightly oval; horizontal * is given axis second. first, 414

427 Lobianchia gemellarii 4.2mm 5.6mm 6.7 mm 11.8 mm Figure Myctophidae 14. Preflexion larva, 4.2 mm (CaICOFI 7205, station 24.139); late flexion larva, 5.6 mm (CaICOFI 7205, station J 1.8 mm (Clarke-Hawaii, station 70--9-6). transformation Ahlstrom 1974); postflexion specimen, 24.139); larva, 6.7 mm (Moser and 415

428 MYCTOPHIDAE Nannohrachium idostigma DESCRIPTION EARLY LIFE HISTORY MERISTICS Range Mode EGGS Shell diam.: Vertebrae: Yolk: 31-34 32 No.ofOG: Diam.ofOG: Total surface: 14 Shell 13-15 Precandal Pigment: 17-19 18 Caudal features: Diagnostic Fins: 0 0 spines Dorsal LARVAE 13 11-14 rays Dorsal 0 mm <2.2 length: Hatching 0 Anal spines 5.3-{).3 ca. length: mm Flexion 17-18 16-20 rays Anal 8 Transformation mm <20 7 mm, II. > length: Pelvic 8 11-12 sequence: development Fin & D A & PI & P , C 10-13 C & Pectoral I 2 2 I Pre on snout by 2.7 mm; flexion-Pair above eye by Pigmentation: Candal: 10+9 10+9 3.5 section gut terminal on pair mm; & above gas bladder by 4.0 rnm; Principal some 4.5 I posterior to eye & at nape (in specimens) by rnm. Procurrent: 7 Flexion-I to upper margin of anterior base; I 6-8 Upper embedded PI 7 in arch series above gas gill region; embedded bladder. upper Lower 6-8 to posterior preopercle 1 on mm, 8.6 ca. Postflexion-By one at the Gill rakers: 4 4-5 Upper gut region. & postorbital myosepta becoming outlined in region 10-12 11 Diagnostic features: large Head large jaws & & compressed; Lower 9 9-11 teeth moderately large of tapered uniform size; body nearly & Branchiostegals compressed & strongly becoming stout short postflexion stage; gut in LIFE HISTORY straighter & becoming flexed, moderate in length; pigment pattem distinct with supra- & postorbital above pigment no melanophores; brain; photophores Br2 mm. 6.3 ca. at form ca. S Range: In Pacific east 25° to N 33° of from W 135° (range %) in mean & MORPHOMETRICS Epi- to mesopelagic Habitat: PrF F Y-S PoF all during captured Larvae season: Spawning seasons CalCOFI on surveys Juv Tr 58-{)8 60-{)5 planktonic 33-59 Sn-AlBL eggs ELH larvae Oviparous; pattern: & 54-55 45 62 63 55 LITERATURE BDIBL 14-26 23-27 27-31 19-20 19 25 29 20 al. 1984a Moser et 17-31 HLlBL 30-32 30-35 30-30 31 24 32 30 (Illustrator) ILLUSTRATIONS ORIGINAL 41-{)8 HWIHL 42-49 46-52 46-49 46 55 50 48 Mattson) (G. larvae, Preflexion mm 4.4 mm, 3.5 26-36 34-38 33-38 SnLIHL 21-23 mm Mattson) (G. Flexion larva, 5.5 32 35 22 36 Postflexion 10.8 mm larva, Mattson) (G. 29-41 x 28-34x 21-30x EDIHL* 16-18 (G. mm Mattson) 21.0 Juvenile, 33-44 32-34 22-33 36x38 31x33 26x29 18 is axis horizontal elliptical; slightly Eye axis * second. vertical first, given 4-12 10-12 9-13 9-10 8 11 11 9 5-{) 0-5 7-13 11-11 1 10 5 11 416

429 Nannobrachium idostigma -------~----~ ':.\ .. '., .' .' . ____ " .. , ,):(,,<.:/,'//' i.··· .. ,.~:~:-==.~= .. ,: ••••.. , ;;7 _ ... __ ... . ••...•.. 3.5 mm 4.4mm 5.5 mm 7.2mm ~~~/ ~ .•... 10.8 mm 21.0 mm Figure Myctophidae 15. Preflexion larvae, 3.5 mm (CaICOFI 6604, station 137.55), 4.4 mm (CaICOFI 6607, station 130.80); flexion larva, 6207, station 130.80); 1984a), station 130.55); postflexion larvae, 7.2 mm (Moser et al. mm 10.8 mm (CaICOFI 5.5 (CaICOFI 6110, juvenile, 21.0 mm (EASTROPAC I, station 13.167). 417

430 MYCTOPHIDAE Nannobrachium "niger"* DESCRIPTION MERISTICS HISTORY LIFE EARLY Mode Range EGGS diam.: Shell Vertebrae: Yolk: No.ofOG: 36-38 37 Total Diam.ofOG: Shell surface: Precaudal 15-16 16 Pigment: 20-23 21 Caudal featnres: Diagnostic Fins: 0 0 Dorsal spines 14 LARVAE 14-16 rays Dorsal Anal spines mm 0 0 <3.0 length: Hatching 18 Flexion length: ca. 6.2-7.3 mm rays Anal 16-19 7-8 ca. Pelvic Transformation mm 20 length: 15 13-16 sequence: Fin development A & & P & C C & Pectoral D PI b 2 2 Preflexion--In larvae, I in midline anterior to smallest Caudal: Pigmentation: 10+9 10+9 Principal a pair forebrain, midbrain, of region anterolateral on of tip near a pair Procurrent: lower to upper snout, I on anterior jaw symphysis, I embedded 6-8 Upper 7 margin of I on of PI base, inner surface ventral 2 on I or & base, PI 6-8 8 Lower region anterior below midline I proximally of on gut; by 4.0 mm, rakers: Gill lower blade bladder. Flexion--Some have PI I & I above gas 4-5 4 Upper on region, basibranchial at posterior & gut, of region lateral (minute) 10-12 II Lower distal & bases on mm, 9.0 PI Posiflexion--By . ca. P on of regions 2 Branchiostegals to anterior series Trunk have some & rays paired D. Transformation-- myosepta outlined. LIFE HISTORY features: jaws Stout body with large head; very long snout; Diagnostic patches large, large teeth; paired tooth with at tip of snout with curved preopercular of short gut spines; & anterior series canines; posterior of margin western Pacific; north sUbtropical & Tropical Range: California unique early pigment larvae; pattern; preflexion in Br2 photophores Current region flexion. in form early Mesopelagic Habitat: mean in %) MORPHOMETRICS (range & Larvae captured during all seasons Spawning season: surveys CalCOFI on but most frequently during surmner Y-S F PrF PoF Tr Juv ELH Oviparous; planktonic eggs larvae pattern: & Sn-AlBL 39-61 62-68 68-75 51 64 72 65 LITERATURE 17-28 BDIBL 26-32 33-42 23 29 36 28 & Ahlstrom 1974 Moser HLlBL 24-39 37-41 39-45 Moser et al. 1984a 32 39 42 31 HWIHL 36-51 41-41 43-56 ILLUSTRATIONS (Illustrator) ORIGINAL 44 41 48 63 SnLIHL 39-61 52-56 43-56 mm Arthur) (N. larva, Preflexion 3.7 50 51 53 35 Head of3.7 mm preflexion larva, dorsal view (N. Arthur) 24-33 x 23-25x 18-24x EDlHLt Arthur) mm (N. Flexion larva, 7.1 27-28 26-38 19-27 (N. mm 20.7 Arthur) Transformation specimen, 29x31 24x28 21x23 19 3-10 9-11 PILlBL 13-16 in introduction explanation See family. * to 7 10 14 10 axis horizontal oval; axis slightly t Eye second. vertical first, is given P L1BL 0-0 3-6 12-26 2 0 5 17 12 418

431 Nannobrachium "niger" 3.7mm .. ~ I 11" 7.1 mm 8.7mm . . l(l,·I/ «( .. ~. ({(t. 20.7 mm Figure Myctophidae 16. Preflexion larva, 3.7 mm, lateral view and dorsal view of head (CaICOFI 7205, station 27.137); flexion larva, and 7.1 (CaICOFI 7205, station 21.133); postfiexion larva, 8.7 mm (Moser mm Ahlstrom 1974); transformation specimen, 20.7 mm (Clarke- 71-3-4). Hawaii, station 419

432 MYCTOPHIDAE Nannobrachium "no pectorals"* MERISTICS EARLY LIFE mSTORY DESCRIPTION EGGS Range Mode Vertebrae: Shell diam.: Yolk: 33-35 33 No.ofOG: Total Diam.ofOG: 13-16 Shell surface: Precaudal 14 18-21 Pigment: Caudal 19 Diagnostic features: Fins: 0 0 spines Dorsal 12-15 LARVAE Dorsal rays 13 <3.0 length: mm Hatching 0 spines Anal 0 15-19 17 Flexion length: ca. 5.6-6.6 mm Anal rays length: Transformation mm <20 8 8 Pelvic 0,20-21 0 Pectoralt A & C & P Cl> , D development PI' Fin sequence: 2 2 Caudal: Pigmentation: tip on lower 5.0 & 4.0 Prejlexion-Between of mm, 10+9 10+9 Principal blade. Flexion-By peppering 5.6 mm, & on PI ca. on jaw, snout, Procurrent: of mm, PI 6.0 ca. by blade; 1 in midline surface lateral on peppering 6-7 6 Upper of pair midbrain, pair forebrain, at anterolateral anterior to surface 6 6-7 Lower 1 embedded in otic of embedded vertical series snout, region, tip near Gill rakers: to anterior developing on inner & PI lateral base, & peppering Upper 4 4 surfaces PI of base. Postflexion-By 6.5 mm, on nape, embedded 11-12 11 Lower mass, gut to anterior 7.5 P on peppering ; by paired series & mm, 2 Branchiostegals to D anterior specimens). (some large with body Stout PI huge base head; & blade; Diagnostic features: LIFEmSTORY on & patches tooth paired teeth large; jaws relatively snout long; rays peppered with base & PI larvae; early short gut jaw; upper in melanophores; minute Br2 ca. 6.5 mm; PI lost at at form photophores of central western to mass Pacific water California Range: North margin transformation. Current region %) in mean & (range MORPHOMETRICS Habitat: Mesopelagic throughout captured season: Larvae Spawning year Y-S PrF F PoF Tr Juv Sn-AIBL larvae ELH pattern: Oviparous; & eggs planktonic 37-53 54-62 60-66 44 59 63 LITERATURE BDIBL 16-25 23-28 29-35 26 19 32 Moser 1981 HLlBL 20-28 30-35 31-39 et al. Moser 1984a 25 33 36 HWIHL 43-67 46-61 46-51 ILLUSTRATIONS (Illustrator) ORIGINAL 53 47 55 SnLIHL 30-40 35-41 36-43 Preflexion larva, 4.0 mm (N. Arthur) 36 39 40 mm 6.0 (N. Arthur) larva, Flexion 34-40x EDlHLt x 29-31 22-30x view (N. of 6.0 dorsal larva, flexion mm Arthur) Head 33-34 35-43 24-33 37x39 30x33 26x29 explanation introduction * See in to family. 8-19 PILlBL 20-23 15-28 rays; adults. in absent 20-21 PI PI to t Larvae with up 12 22 22 oval; slightly Eye t second. vertical axis first, given is axis horizontal P L1BL 0-0 0-7 12-17 2 0 4 15 420

433 Nannobrachium pectorals" "no 4.0mm 6.0mm ... 9.4 mm Figure Myctophidae 17. Preflexion larva, 4.0 mm (CaICOFI 7205, station 3 Ll41); flexion larva, 6.0 mm, lateral view and dorsal view postflexion of (CaICOFI 7205, station 24.137); head larva, 9.4 mm (Moser 1981). 421

434 MYCTOPHIDAE Nannobrachium regalis EARLY HISTORY DESCRIPTION LIFE MERISnCS Mode EGGS Range Yolk: diam.: Shell Vertebrae: 36-37 No.orOG: niam.ofOG: 36-39 Total surface: 15 Shell 15-16 Precaudal 21 Pigment: 20-22 Caudal Diagnostic featllres: Fins: 0 0 Dorsal spines LARVAE 15 12-17 DOl"sal rays <3.2 length: Hatching 0 0 spines Anal 6.0-8.0 Flexion ca. length: 18 17-19 rays Anal Transformation length: ca. 20.0 mm 8 8 Pelvic Fin C, 1'" & 1'1 & A & D 13-14 C sequence: development 12-15 Pectoral b ca. flexion-By Pre to anterior midline 1 in mm, 4.3 Pigmentation: Caudal: (most lobe midbrain each to 1 anterior & 4.7 by specimens); 10+9 forebrain 10+9 Principal peppering on bladder; 5.4 mm, by above at lower jaw tip & gas mm, Procnrrent: P1 blade & 1 at gut above 7 preanal arch of gut; embedded mm, 5.8 by 6-8 Upper 7 at margin near upper c1eithrum snout tip, Flexion-On base; 6-8 ofP1 at Lower 1 posterior eye; to anterior just & nostril, on to Ad; at blotch orbit; Gill rakers: 4 above series gut P inner surface & gas bladder; base; 1 in on of 3-5 Upper 1 peppering cerebellum; above midline on 1'2 Postflexion- blades. 9-11 10 Lower 10 9-11 region; gut in outlined becoming Myosepta some midline 1 in develop Branchiostegais D. anterior to head Diagnostic features: Initially, & small, body slender, short & gut HISTORY LIFE deep become body & head mm, 5.0 at ca. beginning strongly sigmoid; & jaws & head large with armed elongate large snout compressed; Subarctic-transitional Bering including Pacific, North Range: of region upper jaw tip; 1'1 & on teeth with patch tooth teeth; large with larger Bahia off & Japan Bay, Sagami at captures southernmost Sea; distinctive pattern & blotch Ad with pigment pigmented; & large P 2 California eastern & Baja Pacific, Magdalena, the in Sur, western mm. stripe through eye; Br, photophores form at ca. 7.5 respectively in MORPHOMETRIes (range & mean %) to Habitat: Epi- mesopelagic PrF Y-S occur Larvae season: Spawning area; PoF F in year CalCOFI the throughout Tr Juv abundance peak with spring-summer, the in highest July in 29-57 58-67 Sn-AJBL 64--70 56-57 41 62 56 68 59 Oviparous; ELH pattern: planktonic eggs & larvae 11-28 26-30 BDIBL 28-34 20--20 18 LITERATURE 27 31 20 23 15-33 32--43 HL/BL 34--39 28-31 1992 Amaoka et al. 22 36 30 39 33 Matarese et al. 1989 40--48 34--64 50--66 HW/HL 38--49 Ahlstrom 1974 & Moser 60 45 44 40 44 1984a al. et Moser SnLIHL 22--43 42--46 23-26 34--47 44 33 42 24 35 (Illustrator) ILLUSTRATIONS ORIGINAL 24--29x 30-40x 17-23x EDIHL* 15-15 33--46 27-31 19-25 Preflexion Mattson) (G. mm larvae, 3.8 mm, 5.8 20x21 36x40 27x29 15x16 15 Flexion larva, 7.2 mm (G. Mattson) (G. Mattson) of 9.1 mm flexion larva, dorsal view Head 3-10 10--17 12-17 14--15 L/BL P1 19.9 mm (G. Postflexion Mattson) larva, 14 8 13 14 11 Juvenile, 28.0 mm (G. Mattson) 4-22 15-27 0-0.7 12-13 LIBL P2 t 0.1 12 20 12 is vertical slightly elliptical; horizontal axis Eye given first, * axis second. specimen. t 1', rays broken in transfonnation 422

435 Pinpoint lampfish regalis Nannobrachium ~ ... " 3.8mm 7.2mm 9.1 mm 13.0 mm 19.9 mm Myctophidae 18. Preflexion Figure 3.8 mm (CaICOFI 6407, station 77.8.60),5.8 mm (CaICOFI 6407, station 76.70); flexion larva, larvae, 7.2 mm (CaICOFI 4907, station 61.87); head of 9.1 mm postflexion larva, dorsal view (CaICOFI 4908, station 72.56); postflexion larvae, station 13.0 (Moser and Ahlstrom 1974), 19.9 mm (CaICOFI 6806, mm 90.65); juvenile, 28.0 mm (CaICOFI 4911, station 31.65). 423

436 MYCTOPHIDAE Nannobrachium ritteri* DESCRIPTION HISTORY LIFE EARLY MERISTICS EGGS Range Mode Shell Vertebrae: diam.: Yolk: 35-38 Total 36-37 No.ofOG: Diam.ofOG: 15-16 15 Shell surface: Precaudal Pigment: 20-22 21 Caudal Diagnostic features: Fins: 0 0 spines Dorsal 14 12-16 rays Dorsal LARVAE <3 Hatching 0 spines length: Anal mm 0 Flexion length: ca. 6.0-7.1 18 16-19 rays Anal mm 8 Pelvic ca. length: Transformation mm 16.0-18.0 8 II-12 10-13 Pectoral , D C C sequence: development Fin , P PI & A & & I 2 2 Pigmentation: ca. Preflexion--At Caudal: to anterior midline in I mm, 4.0 10+9 10+9 Principal 4.5 hindbrain forebrain; by ca. & mm, 1 embedded in midline below Procurrent: anterior mm, 5.0 by midbrain; to midline to 1 in anterior 1 embedded 7-8 Upper 7 1 on more above gas bladder, base, inner surface of PI 1 gut, 1 or 7-8 8 Lower section, above terminal gut cleithrum & 1 above nape. Flexion--l at rakers: Gill to anterior upper becoming PI base. Posiflexion-Myosepta outlined 4 4-5 Upper at gut, spreading dorsally in & ventrally; 1 anterior to Ad some 9-II 10 on more ventral midline below gut; specimens; P on rays. 1 or Lower 2 Branchiostegals small rounded head, slender body, Diagnostic features: Initially, & & head & body become deep gut; strongly short, sigmoid compressed, LIFE HISTORY preflexion stage; in postflexion stout body stage; during gut elongates upper large relatively large, jaws with large teeth, especially at eyes than jaw tip; snout somewhat elongate but blunter N. in regalis; Range: to ca. south Pacific northeastern the of Subarctic-Transitional zone midline brain; especially pigment, diagnostic of Br2 photophores on California Baja Magdalena, Bahia Sur 7.0 mm. form at ca. rnesopelagic Epi- Habitat: to 0/0) in mean & (range MORPHOMETRICS area; CalCOFI in year Spawning throughout occur Larvae season: the winter highest abundance in the & Y-S in a peak March with spring, PrF F PoF Tr Juv pattern: ELH & eggs planktonic larvae Oviparous; Sn-AlBL 53-59 58-63 29-50 55-57 53-56 38 61 55 54 56 LITERATURE II-22 BDIBL 23-28 28-31 21-24 18-20 17 25 29 23 19 Ahlstrom 1965 15-28 HLlBL 27-31 27-33 27-28 29-30 Bolin 1939 21 29 31 27 30 et Matarese 1989 a1. 49-65 HWIHL 47-67 50-56 43-50 39-44 1974 Ahlstrom & Moser 58 54 52 42 47 1984a a1. Moser et 17-33 SnLIHL 36-39 31-42 27-33 18-21 26 38 35 30 20 ILLUSTRATIONS (Illustrator) ORIGINAL 29-44x 36-38x 26-35x EDIHLt 21-24x x 20-21 29-46 37-42 29-35 21-24 20-21 Mattson) 4.7 larva, mm (G. Preflexion 36x39 30x32 37x40 21x21 22x22 7.0 mm (G. Mattson) larva, Flexion Postflexion larva, 7.6 mm Mattson) (G. 7-9 6-8 PILlBL 8-12 9-10 9-10 dorsal of postflexion 7.6 larva, view (G. mm Mattson) Head 7 10 8 10 10 (G. Mattson) 14.4 mm Postflexion larva, L1BL P 0-2 6-10 0-0 8-10 100II 2 (G. Mattson) Juvenile, 22.5 mm 0 8 0.8 9 II on * species, L. fernae, occurs related the northwestern margin A closely not the CalCOFI area (Wisner 1976); we have larvae identified the of of L. L. but presume they are similar to fernae ritteri larvae. t Eye slightly oval; horizontal axis second. is given fust, vertical axis 424

437 ritteri Nannobrachium Broadfin lampfish 4.7mm 7.0mm 7.6mm 10.1 mm .•... mm 14.4 . -' ... \ Ci )~~" " Figure Myctophidae 19. Preflexion larva, 4.7 mm (CaICOFI 6801, station 93.90); flexion larva, 7.0 mm (CaICOFI 6201, station 83.65); 6806, and dorsal view of head (CaICOFI and station 70.70), 10.1 mm (Moser Ahlstrom 1974); view postflexion 7.6 larvae, mm, lateral juvenile, 90.160); 22.5 mm (CaICOFI 4909, station 82.67). specimen, 14.4 mm (CaICOFI 6204, station transformation 425

438 Notolychnus valdiviae MYCTOPHIDAE LIFE HISTORY DESCRIPTION EARLY MERISTICS EGGS Mode Range Yolk: Shell diam.: Vertebrae: Diam.ofOG: No.ofOG: 30 27-31 Total Shell surface: 13 12-13 Precaudal Pigment: 17 16-18 Caudal features: Diagnostic Fins: 0 0 Dorsal spines LARVAE II 10-12 Dorsal rays mm <2.8 length: Hatching 0 0 spines Anal Flexion 4.4-6.2 length: ca. mm 13-14 12-15 Anal rays ca. Transformation length: 10.0-10.8 mm 6 6 Pelvic sequence: development Fin C , D & A C & PI & , P 13-14 12-15 I 2 2 Pectoral 2.8 ca. Preflexion--By Pigmentation: just midgut on I laterally mm, Caudal: anterior to preanal arch mm, 3.2 ca. by gut; of I) (usually ventral 1-4 10+9 10+9 Principal I above developing gas dashes; by ca. 3.8 mm, midline postanal Procurrent: terminal section. Flexion--Usually 2 or 3 on & gut pair on bladder 7 6-8 Upper gut; lateral ca. by I margin. mid-hypural at 5.0 mm, a streak or 7 6-8 Lower 3 or ventral midline series, in postanal Postflexion--2-7 (usually 4) Gill rakers: A gut on 3 laterally to up base; largest of in either to displaced side 2 2 Upper laterally larvae. Transformation--Blotch gut. on 8 8-9 Lower count (27-31); Low features: low Diagnostic total vertebral uniquely 9 9 Branchiostegals slender body; gut short, larger anterior section tapers ray count P (6); 2 slight gradually, initially size, in moderate head curvature; sigmoid LIFE mSTORY elongate somewhat becomes snout rounded; somewhat blunt & at tip; irregularly eyes narrow, becoming oval; a crescent of choroid-like & tropical temperate in Circumglobal Range: waters tissue on dorsal surface eye of by 4.0 mm & on ventral surface by ca. 6.0 mm; teeth minute; pigment sparse but diagnostic, particularly the Epi- Habitat: mesopelagic to is streak; the pineal region mid-hypural interorbital the in organ visible postflexion late in larvae; transforms at small size; Dn, Br , 2 season: Spawning Larvae occur throughout year area; in CalCOFI highest , VLO, _ Br2 form in PVO!> photophores PV0 & PO form fust; adult S I 2 & summer, with a peak in August in abundance spring the eye. position below eggs planktonic larvae pattern: ELH Oviparous; & %) & mean MORPHOMETRICS in (range LITERATURE PrF PoF F Y-S Juv Tr 1983 Fahay 38-43 51-52 51-52 40-44 44-53 Sn-AlBL Moser Ahlstrom & 1974 48 52 41 42 51 et Moser al. 1984a 21-22 13-16 II-15 19-20 16-23 1964 Pertseva-Ostroumova BDIBL 19 19 15 22 13 Hming 1918 21-26 27-29 27-28 25-31 18-22 HUBL ORIGINAL ILLUSTRATIONS (Illustrator) 23 20 28 28 27 44-55 47-62 46-49 50-56 48-55 HWIHL 2.8 larva, mm Preflexion (G. Mattson) 54 47 51 48 53 Mattson) (G. mm 5.4 Flexion larva, 22-25 27-34 32-39 34-40 23-36 SnLIHL Mattson) (G. larva, mm 7.2 Postflexion 32 36 36 23 30 mm Mattson) (G. Transformation 10.7 specimen, 24-29x 21-29x x I 8-25 19-20x 21-22x EDIHL* 9.7 Juvenile, mm (G. Mattson) 20-22 22-29 27-38 36-42 30-49 19x21 21x26 23x33 25x42 26x39 axis narrow or with long variably Eye oval inclined * trapezoidal, usually 5-8 5-7 7-16 18-20 16-19 forward; axis narrow given fust, long axis second. 18 is 19 12 7 6 7-9 8-9 0-0 0.9-8 0-0 0 0 5 8 8 426

439 Topside lamp fish valdiviae Notolychnus mm 2.8 5.4 mm mm 7.2 mm 8.7 10.7 mm o 9.7 mm Figure Myctophidae 20. Preflexion larva, 2.8 mm (CalCOFJ 6501, station 80.160); flexion larva, 5.4 mm (NORPAC, station 91); 10.7 postflexion 7.2 mm (CalCOFI 6201, station 80.200),8.7 mm (Moseret al. 1984a); transformation specimen, larvae, mm (EASTROPAC mm I, 12.084); juvenile, 9.7 station (CalCOFI 6607, station 110.60). 427

440 MYCTOPHIDAE Notoscopeius respiendens EARLY LIFE HISTORY DESCRIPTION MERISnCS Mode EGGS Range diam.: Shell Yolk: Vertebrae: 37-38 Diam.ofOG: 35-38 No.ofOG: Total 16 surface: 16 Shell Precaudal Pigment: 21-22 21 Caudal features: Diagnostic Fins: 0 0 Dorsal spines 22-23 21-24 LARVAE rays Dorsal 0 mm Hatching <2.4 length: spines Anal 0 5.0-6.5 mm 19 18-20 ca. Flexion length: Anal rays 20.0 mm Transformation length: ca. 8 8 Pelvic Fin sequence: development C & & PI , 11-13 P 12 A & D C Pectoral b 2 2 at Prejlexion-Initially, 3.0 ca. at cleithrum; & tips jaw Pigmentation: Caudal: 10+9 at Principal ca 10+9 region; otic in & bladder gas developing above embedded mm, mm, 3.8-4.0 tandem above terminal gut section & pair above Procurrent: 2 in 4.8 between cerebellum; mm, 1 in midline at nape, 1 anterior & 5.0 11-14 12 Upper more 10-14 13 or on I ventrolaterally midbrain, to 1 midgut, above trunk Lower midline on a series each of on lateral beginning at midbody, dashes Gill rakers: 6 Flexion- Upper base, & ofD 6--7 beginning of a midline series on A base. side 14 13-15 larvae midbrain in some above & beginning of pair Transverse series Lower 10 9-10 hypaxial region above A base in some larvae. on Pos!flexion-Paired Branchiostegals to on dorsum extends Ad in some larvae; by ca. 8.5 mm, series 1 at angle jaw of in gular more 1 or have some mm, 9.5 ca. by some; in LIFE HISTORY mm, & anterior to forebrain; by ca. 12.5 & on P rays & A region 2 margin; hypural by 14.5 mm, on edge ofbranchiostegal membrane. temperate Range: Pacific in eastern waters; from to tropical in Worldwide features: procurrent C ray (21-24); high Diagnostic count High D ray Chile to California central + count (11-14 strongly 10-14); initially slender, with short, sigmoid body & in acute snout compressed; & deep become head gut; mesopelagic to Epi- Habitat: stage, preflexion becoming bulbous; eyes large, nearly round; teeth posterior on teeth hooked well developed; anteriorly part jaw; oflower & fall in CalCOFI abundant in summer Larvae most season: Spawning form 4.2 at PO Br2 photophores mm; at complex pattern; pigment s in peak with area, August Vn 9.2; PLO at 16.2 mm. at 6.2; ELH pattern: Oviparous; planktonic eggs & larvae MORPHOMETRICS (range & mean in %) LITERATURE Y-S PrF PoF F Tr Juv Badcock & Merrett 1976 Sn-AlBL 38-58 58-63 56--60 57-57 1983 Fahay 44 61 58 59 57 1989 a1. et Matarese 1981 Moser BDIBL 24-29 19-26 24-28 19-21 22 26 26 & 1972, Ahlstrom 1974 Moser 24 20 Moser 1984a a1. et 28-32 HLlBL 23-31 30-34 30-30 1988d 1986b, Ozawa 27 31 30 29 30 43-60 HWIHL 49-69 48-53 39-41 nONS ORIGINAL (Illustrator) ILLUSTRA 51 57 49 53 40 SnLIHL 33-37 29-37 21-29 17-17 mm, mm (G. Mattson) 3.1 larvae, Preflexion 4.8 35 33 19 25 17 Mattson) 6.5 (G. mm, Postflexion mm larvae, 16.2 29-40x EDIHL* 28-40x 35-37x 25-25 Mattson) mm 21.0 (G. Transformation specimen, 35-45 39-43 28-41 Juvenile, 26.3 mm (G. Mattson) 35x4l 36x41 36x37 28 25 7-8 6--11 6--8 12-13 PILlBL given first, vertical axis second. Eye slightly oval; horizontal axis is * 7 8 10 12 12 P 0-0 L1BL 0-5 6--15 13-14 2 0 2 12 14 13 428

441 Patchwork lampfish respiendens Notoscopeius mm 3.1 4.8mm mm 6.5 mm 16.2 mm 21.0 26.3 mm Figure Myctophidae 21. Preflexion larvae, 3.1 mm (CaICOFI 6304, station 120.120), 4.8 mm (CaICOFI 6107, station 90.70); postflexion 6207, 21.0 mm (CaICOFI station station 113.45), 16.2 mm (LACM 9567-6); transformation specimen, larvae, mm (CaICOFI 6.5 6610, to 90.110); 26.3 mm, parasitic copepod attached juvenile, ventrum (CaICOFI 6208, station 90.150). 429

442 MYCTOPHIDAE Parvilux ingens MERISTICS EARLY LIFE mSTORY DESCRIPTION Mode Range EGGS Vertebrae: Shell diam.: Yolk: Total 35-38 No.ofOG: Diam.ofOG: 37 15-16 Shell 15 surface: Precaudal 21-22 22 Caudal Pigment: Fins: Diagnostic features: Dorsal spines 0 0 LARVAE 14-17 16 Dorsal rays length: Hatching spines 0 0 Anal 15-18 length: rays Anal mm 6.6-7.8 ca. 17 Flexion Pelvic mm length: > 17 Transformation 8 8 Pectoral 10-13 12 P C , PI' D, C A, development Fin , sequence: I z z 5.6 to Caudal: Pigmentation: Pre flexion-By by mm, 1 midventral hindbrain; 10+9 10+9 end Flexion-By I in stage, of more Principal 1 or mm, 6.5 gas bladder. above Procurrent: midline anterior to midbrain, 1 in midline posterior to cerebellum, & 7-9 midline Posiflexion-Apposing gut. anterior embedded I on to Upper pair 8 7-9 8 posterior slightly dorsal peduncle, largest Lower caudal to ventral; specimens of caudal peduncle; myosepta 2 on dorsal may have 1 or midline Gill rakers: 5 4-6 outlined in gut region becoming late in Upper stage. postflexion 11-15 in in 12 longer becoming stage, preflexion features: Gut short Diagnostic Lower Branchiostegals 38%-54% (EL stages; extremely large, nearly round eye subsequent distinct HL); apposing with peduncle caudal pigment pattern, stage; melanophores in postflexion form LIFE in photophores Brz mSTORY larvae. postflexion California Current region from central Baja Range: north to California in mean %) MORPHOMETRICS (range & ca. to zone transition in westward & Cape California, Mendocino, 0 W 145 y-S PrF PoF F Tr Juv Mesopelagic Habitat: 36-42 Sn-AlBL 52-58 45-50 47 39 55 & summer in CalCOFI during spring Spawning season: Larvae captured area 22-22 15-20 BDIBL 21-22 18 22 21 ELH pattern: larvae & eggs planktonic Oviparous; 26-28 HLlBL 16-25 25-27 21 27 26 LITERATURE HWIHL 56-73 55-57 47-57 64 53 56 al. 1989 et Matarese 26-27 24-28 SnLIHL 24-27 1974 Ahlstrom & Moser 26 26 26 et Moser 1984a al. 41-46x EDIHL* x 40-45 37-46x 46-54 47-48 38-46 ILLUSTRATIONS ORIGINAL (lliustrator) 43x50 42x47 41x42 Preflexion larvae, 6.5 mm, 4.5 mm Arthur) (N. 5-6 4-4 7-10 PILlBL Flexion larva, 7.8 mm (N. Arthur) 4 6 8 0-0 0-0 0-7 PzLIBL 0 3 0 horizontal * Eye slightly oval; axis second. vertical is given first, axis 430

443 ingens Parvilux Giant lampfish mm 4.5 mm 6.S ~ -~--~~-~ 7.8 mm ... ". '. ". ~.:.l.~Il? ••... ~ ~~":)..: ,"' ... 14.4 mm Figure Myctophidae 22. Preflexion larvae, 4.5 mm, 6.5 mm (CaICOFI 7205, station 31.131); flexion larva 7.8 mm (CaICOFI 9008, station 83.110); postflexion larva, 14.4 mm (Moser and Ahlstrom 1974). 431

444 Stenobrachius leucopsarus* MYCTOPHIDAE DESCRIPTION EARLY LIFE HISTORY MERISTICS EGGS Mode Range Shell Yolk: diam.: Vertebrae: No.ofOG: Diam.ofOG: 36 35-38 Total surface: Shell 15 15-16 Precaudal Pigment: 21 20-22 Caudal Diagnostic features: Fins: 0 0 spines Dorsal LARVAE 14 13-15 rays Dorsal ca. 2.0 Hatching mm length: 0 0 spines Anal Flexion mm 6.5-8.0 length: 15 14-16 rays Anal mm 16-19 length: Transformation 8 8 Pelvic CI> PI> , P C & D & A Fin development sequence: 8-10 9 2 2 Pectoral Pigmentation: Preflexion-Postanal 19-22; with series midline ventral Caudal: & just posterior to section, at midgut, on pairs gut at terminal lateral 10+9 10+9 Principal stage, of end at mm; 6.0 at ca. bladder gas above embedded base; PI Procurrent: the & midline in coalesces pair anteroventrad gut migrates anterior 7 6-8 Upper postanal base Flexion- coalesce. PI to & 8 series begins the below 7-8 Lower Postanal series coalesces to 1-3 insertion atA one the stage, of at end Gill rakers: 1-2 of hindbrain & at nape. Postflexion- not 1 at midline embedded; 5 5-6 Upper some 8.0 mm; in region larvae embedded 1 (minute) in by otic 13 12-14 Lower mm, 1-6 in paired series 10 dorsum anterior to D in alternating on > 9 9-10 Branchiostegals D insertion, Ad origin & insertion, & a at 1 or more arrangement, series anterior to D. Transformation-Paired series on lateral midline LIFEIDSTORY lateral midline augmented. & dorsum on slightly gut but mexicanus Triphoturus to Similar features: Diagnostic N in California Current Pacific 29° North Range: to Gulf of from ca. eye wider, more postanal shorter, melanophores 11-14), vs. (19-22 & Japan to westward & Alaska (ca. early form photophores mm); Diaphus 6.8 have <17 Br2 theta have & hypural stage flexion to prior melanophores midline postanal to Epi- Habitat: mesopelagic Brl form in late larvae; Op & Vn, form POI,PO mm; at >4.4 pigment S early in transformation. season: occur Spawning Larvae in year the throughout area; CalCOFI March with peak in in abundance winter-spring, highest (range MORPHOMETRICS & in %) mean Oviparous; ELH planktonic eggs & pattern: larvae PoF F PrF Tr Juv Y-S LITERATURE 55-56 53-57 54-58 36-49 56-60 Sn-AlBL 54 44 56 58 56 Ahlstrom 1965, 1972a 21-23 17-24 8-13 14-19 19-21 BDIBL 1960 Fast 11 16 22 20 22 Matarese al. 1989 et Moser 1974 Ahlstrom & 23-27 21-23 26-28 lllJBL 27-30 16-21 et al. 1984a Moser 22 18 28 25 29 46-61 56-65 HWIHL 40-48 40-49 50-60 ORIGINAL ILLUSTRATIONS (Illustrator) 43 60 55 56 45 23-28 23-26 14-28 20-23 19-28 SnLIHL Preflexion larva, 3.7 mm (N. Arthur) 24 24 21 21 24 Juvenile, Arthur) 21.0 mm (N. 29-37x 22-25 27-30x 23-33x 23-28 EDlHLt 27-40 35-41 32-39 28x34 28x35 25 24 34x37 * closely related northern species, A nannochir, extends into the S. the CalCOFI not identified area (Wisner 1976); of have we part northern 7-10 7-8 3-7 6-8 6-11 to nannochir S. they but are leucopsarus similar larvae presume S. 5 7 9 8 7 larvae. 8-10 1-9 0-0.5 0-0 10-11 slightly oval; horizontal axis t second. axis is Eye first, vertical given 10 9 0.3 6 0 432

445 Northern lampfish leucopsarus Stenobrachius , I ···~··I··.' ~) 3.7mm mm 5.3 mm 7.8 12.5 mm 21.0 mm Figure Myctophidae 23. Preflexion larvae, 3.7 mm (CaICOFI 6501, station 73.51),5.3 mm; flexion larva, 7.8 mm; postflexion larva, 12.5 (Ahlstrom 1972a); juvenile, mm 21.0 mm (CaICOFI 6608, station 90.28). 433

446 MYCTOPIDDAE Taaningichthys minimus* MERISTICS EARLY LIFE HISTORY DESCRIPTION Range Mode EGGS Vertebrae: Shell diam.: Yolk: 39-41 Total 40 No.ofOG: Diam.ofOG: 18-20 Precaudal 19 Shell surface: Caudal 20-22 21 Pigmeut: Fins: features: Diagnostic 0 0 spines Dorsal 11-13 12 Dorsal rays LARVAE 0 Anal spines 0 mm <4.9 length: Hatching 11-14 rays 7.0-8.5 13 Anal ca Flexion length: Pelvic 8 8 21.0 mm Transformatiou leugth: ca. IS-17 16-17 Pectoral , Fin sequeuce: developmeut & D P C Ph C A, h 2 2 Caudal: Pigmentation: elongate 4.9 apposing mm, Preflexion--By median 10+9 10+9 Principal region, gut caudal peduncle blotches in 1 above terminal future Procurrent: section, I above gas bladder, 1 region, otic in 1 embedded 1 at nape, 8-10 9 Upper spinal or more column; by embedded above anterior 1 in 6.1 mm, 8-9 Lower 8 1 above medulla; by 6.8 mm, embedded & above cerebellum midline Gill rakers: Posiflexion--By column. spinal of ca. extends full length series 4-S Upper 4-S median ofD, side each on series mm, 10.0 paired of beginning spot 11-14 12-13 Lower series postflexion late in & stage, on hypural margin; insertion, A at Branchiostegals origin D from extends series paired & paired ventral series at A C to peduncle. caudal & posterior LIFE HISTORY features: vertebral Diagnostic body; High total slender count (39-41); straight; & eyes small; relatively head gut relatively long, slender & distinct region peduncle in caudal particularly pigmentation, round; in Worldwide Range: subtropics southern & northern column. spinal above embedded Habitat: rnesopelagic to Epi- in MORPHOMETRICS %) mean & (range year-round Possibly Spawning season: Y-S PrF F PoF Tr Juv eggs planktonic Oviparous; pattern: ELH larvae neustonic & Sn-AlBL 59-62 63-66 60-65 60 65 63 LITERATURE 9-12 BDIBL 13-15 13-18 1983 11 14 Fahay 15 1989 al. et Matarese 18-22 HLlBL 23-24 21-26 1972 & Ahlstrom Moser 23 20 23 Moser et al. 1984a 51-53 HWIHL 47-58 46-56 52 52 SI ORIGINAL ILLUSTRATIONS (Illustrator) SnLIHL 20-27 21-26 21-26 24 24 23 mm 4.9 larva, MacCall) Preflexion Sumida (B. EDIHL 28-31 27-28 26-29 Sumida MacCall) larva, Flexion 7.2 mm (B. 30 28 27 Sumida mm (B. MacCall) Postflexion larva, 16.9 P,LlBL 3-4 Juvenile, mm (B. Sumida MacCall) 21.S 4-14 4 4 9 P L1BL 0-0 0-0 0.8-10 2 bathyphilus & Two paurolychnus other * Taaningichthys species, T. T. 0 0 5 but larvae occur in the CalCOFI area; we have not identified their their would total vertebral counts (35-36) lower distinguish them from T. minimus (39-41). 434

447 Taaningichthys minimus 4.9mm 7.2mm 16.9 mm 21.5 mm Figure Myctophidae 24. Preflexion larva, 4.9 mm (CaICOFI 7205, station 20.141); flexion larva, 7.2 mm (CFRD Ref. ColI., Hawaii Inst. mm Mar. Miller Neuston Net Series 3); postflexion larva, 16.9 BioI., (CaICOFI 7205, station 27.133); juvenile, 21.5 mm (SID 63-544). 435

448 MYCTOPHIDAE Triphoturus mexican us EARLY LIFE mSTORY DESCRIPTION MERISTICS EGGS Mode Range diam.: Shell Yolk: Vertebrae: No.ofOG: 33-35 Diam.ofOG: 34 Total Shell surface: 16 15-17 Precaudal 18 Pigment: 18-19 Caudal features: Diagnostic Fins: 0 0 spines Dorsal LARVAE 14 13-16 Dorsal rays ca. Hatching 2.0 mm length: 0 spines 0 Anal 6.3-8.0 mm length: Flexion 15-16 14-17 Anal rays 12.5-15.4 Transformation mm ca. length: 8 8 Pelvic P 8-10 Fin development sequence: CI> A, C , D, PI> 9 Pectoral 2 2 in 11-14 mm, postanal flexion-At Pre Pigmentation: 2.0-3.5 ventral Caudal: on pair midline series, gut ventrolateral posterior to PI 1 or just base, 10+9 10+9 Principal along gut, midlateral on 2 pairs some & section, gut terminal on pair Procurrent: Ad fmfold; just posterior of 1 locus in most larvae to margin ventral 6 5-7 Upper in midline to coalesces 1 on ventral gut on mm; >3.7 pair anterior 6 6-7 Lower bladder by 4.3 mm; >4.0 above embedded gas mm; most larvae Gill rakers: 3-5 to 1 or 2 by end of stage. Flexion-At lower postanal series reduced 5 Upper gut. gut lateral symphysis; jaw above embedded & coalesced pair(s) 12 10-14 Lower at base oflower at mm, jaw, 9.0 Posiflexion-By delicate series angle 10 8-11 Branchiostegals 10.5 inferior procurrent caudal rays, & A insertion; by of mm, 1 at to PI base. anterior embedded mSTORY LIFE length gut slender; features: Diagnostic Moderately slightly moderate, of choroid tissue; Br2 sigmoid; eye elliptical with lunate mass from San Francisco, Range: northern population Pacific; Eastern ventral larvae; late in form glands luminous caudal & photophores including Mexico, southern to California to California, of Gulf westward throughout present pigment finfold midlateral lacks period; larval 0 127 W; southern population from Guatemala to Chile T. nigrescens); fewer postanal ventral in pigment series (present (11-14 vs. Stenobrachius leucopsarus in than midline melanophores mesopelagic to Habitat: Epi- 19-22); S. theta finfold ventral lack pigment. leucopsarus & Diaphus area; Larvae Spawning occur throughout the year CalCOFl season: in %) MORPHOMETRICS (range & mean in August-September peak fall, to with abundant from spring larvae eggs planktonic Oviparous; pattern: ELH Y-S PrF F PoF & Tr Juv 44-61 Sn-AlBL 61-66 62-66 56-59 60-61 LITERATURE 64 64 61 55 58 Ahlstrom 1965, 1972a 16-18 17-21 21-22 BDIBL 9-15 17-19 19 18 12 18 21 1984a al. et Moser HLlBL 24-27 25-29 31-34 19-25 29-30 (Illustrator) ILLUSTRATIONS ORIGINAL 21 27 29 32 25 46-55 40-41 49-53 46-52 HWIHL 40-42 (N. Preflexion larva, 3.6 mm Arthur) 49 51 41 41 .51 Arthur) (N. mm Juvenile, 16.7 SnLIHL 15-28 22-28 24-30 23-23 21-24 22 26 27 23 22 21-28x 21-25x 17-26x EDIHL* 15-18x 16-19x horizontal oval; Eye * second. vertical first, given is axis axis 28-36 26-28 15-16 20-31 16-18 24x33 24x27 16x16 20x25 17x17 8-9 8-12 3-6 8-10 PILlBL 5 9 10 8 9 9-10 1-6 L1BL 0-0 0-0.6 P 10-11 2 4 0 0.3 9 10 ! I 436

449 Mexican lampfish us mexican Triphoturus 3.6mm - ... _ ... - .. 4.4 mm ... . .. ~ ... -----~ ------ ... ", ", -", -', ... -"'" ... . ... ... .. - ...•.. mm 6.8 mm 10.5 16.7 mm Figure Myctophidae 25. Preflexion larvae, 3.6 mm (CaICOFI 5904, station 100.75),4.4 mm; flexion larva, 6.8 mm; postflexion larva, 10.5 mm (Ahlstrom 1972a); juvenile, 16.7 mm (CaICOFI 6610, station 97.35). 437

450 MYCTOPHIDAE nigrescens Triphoturus LIFE EARLY HISTORY DESCRIPTION MERISTICS Mode Range EGGS Vertebrae: diam.: Shell Yolk: No.ofOG: 33-35 34 Total Diam.ofOG: 15-16 surface: Shell 16 Precaudal 17-19 18 Pigment: Caudal Diagnostic features: Fins: 0 0 Dorsal spines 13-15 14 LARVAE rays Dorsal length: ca. Hatching 2 mm spines 0 0 Anal mm 5.5-6.5 length: Flexion 16-18 17 rays Anal ca. length: Transformation 15 mm 8 8 Pelvic 8-10 9 Pectoral , C PI' D, A, P C" Fin development sequence: 2 2 Pre Pigmentation: lateral on dashes 11 to up mm, 3.0-4.5 flexion-At Caudal: 10+9 10+9 Principal midline, at 1 embedded PI base, 7 embedded ventral median ca. dashes, postanal margin of ventral finfold, on pair on terminal gut & Procurrent: on laterally & Upper 5-6 6 atcleithral midline 1 in mm, 4.5-5.5 at midgut; section 5-6 to reduced series midlateral jaw 6 symphysis, symphysis, 1 at lower Lower gut; to 2-4. rakers: Flexion-Pair 2-6, at midlateral postanal series reduced & Gill 2-4 3 at one rays; & of 2 on 1 or insertion Upper A A along series delicate bases 7-11 9 Vertical caudal peduncle. Posiflexion- Lower at base of series margin ventral Branchiostegals base. of C rays; by 10 mm, 1 or 2 above caudal & at Ad peduncle gut slender; Moderately Diagnostic features: length, in moderate LIFE HISTORY sigmoid; eye slightly elliptical with lunate ventral choroid slightly ventral photophores pigmentation; finfold & tissue; midline lateral T. at transformation; form larvae; late in form glands luminous caudal Range: Indian & Pacific sUbtropical & tropical across Widespread Oceans mexicanus pigment. midline lateral lacks but similar Habitat: Mesopelagic mean MORPHOMETRICS (range & in %) Larvae year throughout captured season: Spawning PrF Y-S PoF F Tr Juv ELH larvae pattern: & eggs planktonic Oviparous; Sn-AlBL 53-62 61-64 56-64 55-57 63 58 61 57 56 LITERATURE BDIBL 12-18 17-21 19-22 17-18 15 19 20 17 18 1981 Moser Moser 1984a et al. HLlBL 21-26 24-30 27-29 27-28 1988d Ozawa 1986b, 23 27 28 28 27 53-55 HWIHL 53-62 50-59 36-40 ORIGINAL ILLUSTRATIONS (Illustrator) 57 54 55 41 38 SnLIHL 18-29 28-32 25-31 20-20 mm 4.1 larva, Preflexion Arthur) (N. 25 29 30 19 20 Flexion Arthur) (N. mm 5.6 larva, EDIHL* 22-30x 22-25 x 18-26x 16-16x mm (N. Arthur) Juvenile, 16.2 29-36 26-29 22-28 17-18 25x32 23x28 22x26 15x18 16x17 is axis horizontal axis oval; Eye * second. vertical first, given 9-10 7-11 6-8 PILlBL 7 9 9 6 5 L1BL P 0-0 0.4-0.7 1-6 7-9 2 0 0.6 3 9 8 438

451 Trip/wturus nigrescens Higbseas lampfish 5.6mm 8.5 mm 16.2 mm Figure Myctophidae 26. Preflexion larva, 4.1 mm (NORPAC, station 78 Deep); flexion larva 5.6 mm (CaICOFI 5009, station 70.280); 16.2 postflexion 8.5 mm (Moser 1981); juvenile, larva, mm (Cato II, station A5). 439

452 Benthosema panamense MYCTOPIDDAE LIFE HISTORY EARLY DESCRIPTION MERISTICS EGGS Mode Range Yolk: Shell diam.: Vertebrae: Diam.ofOG: No.ofOG: 32 31-33 Total Shell surface: 14 13-14 Precaudal Pigment: 18-19 18 Caudal features: Diagnostic Fins: 0 0 Dorsal spines LARVAE 13 12-14 rays Dorsal Hatching length: <2.5 mrn 0 0 spines Anal Flexion ca. 4.1-5.8 mm length: 21 19-22 Anal rays Transformation length: ca. 10.5-12.3 mm 8 8 Pelvic C" & C P D, A, & P, 14-15 Fin development sequence: 13-16 2 Pectoral 2 flexion-By Pre elongate of pair ventrolateral mm, 2.5 Pigmentation: Caudal: 2 pairs section, terminal gut on cleithrum; at dashes pair dorsolateral 10+9 10+9 Principal 5-9 in postanal median ventral series; by 3.0 & lateral midgut, on Procurrent: 1 on mm, lateral a single only have some mm, 4.0 by tip; jaw lower 8 7-9 Upper forms postanal above Flexion-Patch 1. & to series coalesced pair gut 7 7-8 Lower bladder along & cleithral dash angles dorsad cleithrum; by ca. rakers: Gill each 5.0 base, anterior to P, anterior nostril, in ceratohyal mm, to 8-10 9 Upper surface of hindbrain, & 1 to several on ventrolateral 19 embedded region, 17-21 Lower vertical series in hypurals. edge of on Posiflexion-By ca. posterior 8 8 Branchiostegals ca. by some); (in base A on a series mm, 6.0 1 medially mm, 7.7 above cerebellum; by 8.0 mm, a series embedded along haemal spines. HISTORY LIFE features: Moderately stout body; gut to ca. midbody, Diagnostic slightly in blunt snout rounded; size, in moderate head sigmoid; northeastern 10° & N 28° between N, Pacific Range: Subtropical-tropical after wider elliptical, eye lunate postflexion stage; stage; preflexion California of including Gulf Dn Br2 & choroid tissue; postanal series coalesces to 1 melanophore; at forming photophores ca. & PVO, larvae; 4.0 mrn; POs in 6 mm at Habitat: forms dense aggregations coast; near mesopelagic to Epi- AO, upper larvae. 8 mm Op in 7 mm larvae; PO,; VO" & in at night surface MORPHOMETRICS (range & in %)* mean CalCOFI were region in The few occurrences season: Spawning throughout year F Y-S PrF PoF Tr Juv ELH pattern: Oviparous; planktonic eggs & larvae 48-53 Sn-AlBL 55-61 54-62 51 58 58 57 LITERATURE 21-25 14-17 19-23 BDIBL 16 21 23 25 Moser 1970 & Ahlstrom 1984a al. et Moser 21-24 28-31 27-33 HLlBL 23 30 30 32 61-78 59-68 52-63 HWIHL on (1970). Ahlstrom & * Calculations based Moser in data 71 62 57 53 first, vertical axis second. given is axis horizontal oval; somewhat Eye t SnLIHL 24-3 Ox 26-29x 25-29x EDlHLt 27-34 32-38 35-43 27x31 27x35 28x39 29 3-5 3-6 0-3 CLIHL 0.7 4 4 o 440

453 Benthosema pa namense 4,7 mm 7,7mm 441

454 MYCTOPHIDAE Benthosema suborbitale DESCRIPTION EARLY LIFE HISTORY MERISTICS EGGS Mode Range diam.: Yolk: Shell Vertebrae: No.ofOG: Diam.ofOG: 33-35 34 Total Shell surface: 15 15 Precaudal 18-19 Pigment: 19 Caudal features: Diagnostic Fins: 0 0 spines Dorsal LARVAE 12-14 13 Dorsal rays length: ca. 2.0 Hatching mm 0 0 spines Anal 17 Flexion mm 5.2-6.5 length: 16-19 rays Anal mm 10.0 Transformation length: ca. 8 8 Pelvic 14 , 13-15 , development Fin sequence: C PI' A, C D, P Pectoral I 2 2 symphysis Preflexion-Palr anterior to c1eithral just at Pigmentation: Caudal: 10+9 10+9 ca. in midline. coalesces later 4 mm, lower Flexion-at jaw symphysis Principal mm; by ca. 5.5 to two embedded blotches anterior PI base, one near Procurrent: & the base. fin of bottom near other top 8 8 Upper 7-8 with narrow eyes 7 lunate mass Short, deep body; features: Diagnostic Lower surface; gut short, terminal tissue of choroid section on ventral Gill rakers: ventrad, acutely deflected Sn-A 3 BL in preflexion & flexion <50% 3-4 Upper mm; first & 5.0 second middle Br photophore ca. at stages; forms 10-11 11 Lower anterior blotches embedded scanty; ca. pigment PO's form at 9 mm; Branchiostegals to has which risso Electrono to similar base; PI relatively gut longer PI blotches anterior to PI base. on pigment & lacks but rays LIFE HISTORY MORPHOMETRICS (range mean in %) & apparently & except central Warm Range: eastern circumglobal, water; Pacific equatorial Ocean y-S PrF F PoF Tr Juv rnesopelagic to Epi- Habitat: Sn-AlBL 43-46 46-50 49-53 51-55 55-58 44 48 52 53 56 Possibly Spawning season: year-rouud 15-19 19-27 BDIBL 25-27 24-31 24-26 & eggs planktonic Oviparous; pattern: larvae ELH 29 23 17 25 26 23-25 23-26 HUBL 25-27 28-29 26-28 LITERATURE 24 25 27 28 27 60-68 66-80 68-85 HWIHL 55-57 50-57 1976 Merrett & Badcock 63 72 75 54 56 Fahay 1983 SnLIHL 23-36 28-40 25-33 22-24 18-23 Moser Ahlstrom & 1974 31 30 20 33 23 1984a al. et Moser 18-23 x 17-25x EDIHL* 19-25x 23-30x 28-49x 1988d 1986b, Ozawa 41-48 42-52 32-33 33-42 31-49 1974 Pertseva-Ostroumova 1964, 20x46 22x39 21x45 26x33 38x39 1977 Shiganova 6-11 11-15 4-6 PILlBL ORIGINAL ILLUSTRATIONS (Illustrator) 5 9 19 32 13 0.3-4 0-0 L1BL P 0-0 8-9 2 larva, 4.5 mm Arthur) Preflexion (N. 0 2 0 9 13 Flexion larva, Arthur) 6.5 mm (N. Postflexion Arthur) (N. mm 7.3 larva, Arthur) Transformation specimen, 9.7 mm (N. Arthur) Juvenile, 14.5 mm (N. vertical axis second. first, given is axis horizontal narrow; elliptical, Eye * 442

455 Benthosema suborbitale mm 4.5 7.3 mm mm 9.2 9.7mm 14.5 mm Figure Myctophidae 28. Preflexion larva, 4.5 mm; flexion larva, 6.5 mm (CaICOFI 7205, station 31.141); postflexion larvae, 7.3 mm (Clarke-Hawaii, station 71-6-29),9.2 mm (Moser and Ahlstrom 1974); transformation specimen, 9.7 mm (CaICOFI 7210, station 31.141); juvenile, 14.5 mm (CaICOFI 7210, station 31.145). 443

456 Centrobranchus nigroocellatus MYCTOPHIDAE DESCRIPTION EARLY LIFE HISTORY MERISTICS EGGS Range Mode diam.: Shell Yolk: Vertebrae: No.ofOG: Diam.ofOG: 38 Total 35-40 surface: Shell 14-15 14-15 Precandal 24 Pigment: 22-25 Candal Diagnostic featnres: Fins: 0 0 Dorsal spines LARVAE 10 9-11 rays Dorsal mm length: <2.8 Hatching 0 0 Anal spines 5.4-6.3 Flexion length: ca. mm 18 16-19 Anal rays mm 12 Transformation ca. length: 8 8 Pelvic sequence: C Fin development 14-15 A, & D & P C PI' & 13-17 Pectoral I 2 2 flexion-At <4.0 mm, anterodorsal Pigmentation: PI base, on Pre to Candal: 10+9 jaw orbit, & upper on axilla, near trunk of lower margin posterior tips, 10+9 Principal & all but & forebrain, gut; lateral to tenninal mid- anteromedial to Procurrent: stage. of mm, 6.0 ca. Flexion-By postorbital by lost end pigment 6 5-7 Upper series Br rays, outline & of 6 5-7 liver. surface ventral on patch Lower margin largest larvae, on posteroventral Posiflexion-On & of orbit rakers: Gill posteriorly lower jaws. & upper on 0 0 Upper deep- features: Initially moderately slender, becoming Diagnostic 0 0 Lower narrow 7-8 bodied & highly compressed; head large 7 with elliptical eyes; Branchiostegals choroid tissue extremely elongate, tenninal gut unpigmented; conical only deflected; snout becomes bulbous; large ftnfolds; section slightly HISTORY LIFE postorbital, branchiostegal on pigment lost; except pigment, early membrane & liver fonns in postflexion stage; Br2 photophores fonn Wann-water cosmopolite; apparently absent from low Range: oxygen distinguishes C. nigrocellatus lack mm; 5.0 ca. of on pigment tail at waters tropical of Pacific eastern andreae. those from of C. mesopelagic; to Epi- Habitat: night at surface at & MORPHOMETRICS (range %)* mean in year-round Spawning Possibly season: PrF Y-S F PoF Tr Juv Oviparous; planktonic eggs & larvae ELH pattern: Sn-AlBL 46-65 63-66 68-70 64 56 69 57 LITERATURE 12-29 28-36 BDIBL 31-38 32 21 35 19 Ahlstrom 1970, & 1974 Moser 1984a aI. Moser et 29-32 25-31 32-32 HUBL Pertseva-Ostroumova 1974 1964, 31 28 26 32 HWIHL 40-48 41-47 40--50 45 44 44 48 based Calculations * Moser & Ahlstrom (1970). on in data SnLIHL axis elliptical, very narrow; horizontal t axis is given ftrst, vertical Eye second. 14-17x 14-16x 12-14x EDlHLt 20--27 29-40 26-31 16x33 13x23 14x29 20x21 P L1BL 2 21-32 7-28 CLIHL 22-31 26 20 27 o 444

457 nigroocellatus Centrobranchus Roundnose lanternfish 3.9mm :~. '.-::--.. -""" mm 7.3 o o 17.7 mm Figure Myctophidae 29. Preflexion larva, 3.9 mm; flexion larva, 5.8 mm; postflexion larvae, 7.3 mm, 12.0 mm; juvenile, 17.7 mm (Moser and Ahlstrom 1970). 445

458 MYCTOPHIDAE Diogenichthys at/anticus MERISTICS DESCRIPTION EARLY LIFE mSTORY EGGS Range Mode Shell diam.: Yolk: Vertebrae: No.ofOG: 31-35 33 Total Diam.ofOG: 13-14 surface: Shell Precaudal 14 Pigment: Caudal 18-20 19 Diagnostic features: Fins: spines Dorsal 0 0 LARVAE 10-l2 11 rays Dorsal length: mm <2.9 Hatching 0 Anal spines 0 Flexion ca. length: 14-18 16 6.0-6.9 mm rays Anal Pelvic 8 length: Transformation mm ca. 13.5-14.5 8 12-15 14 P & D A, & Pectoral & p) C C)' sequence: development Fin 2 2 Pigmentation: ventrolateral Preflexion--By ca. 3.0 mm, pair just Caudal: 10+9 10+9 Principal posterior 2 pairs gut, terminal on pair dorsolateral c1eithrum, to on lateral midgut, ca. 1 postanal series at ventral margin; & laterally 3 in Procurrent: 8-9 8 Upper arch preanal above gut; of surface dorsal 3 on of first mm, 5.0 at ca. 2 on 8 8-9 Lower of symphyseal barbel; 1 or gut & up laterally more 4 to by end Gill postanally stage. of rakers: C; on rays of base at (large) Flexion--l 2 2 Upper 1 & pair above hindbrain; ofP) on anterior part pair below embedded 11-13 12 to form begin at bases paired mm, 7.0 ca. Posiflexion--At Lower base. series Branchiostegals 6-8 7 midline postanal in 12 & gut on 6 laterally to up rays; A of ventral in larvae. one posterior to D & 1 posterior to Ad in largest largest larvae; LIFE mSTORY Diagnostic features: Moderately slender somewhat becoming body, compressed; gut midbody, slightly sigmoid; head size moderate; to acute, becoming relatively shorter; eye elliptical, becoming snout Range: into cosmopolite; warm-water distributed Unevenly extends ventral tissue, wider in later stages, no choroid although the scleral California Current region from central water mass the through may envelope extend ventrad early from the eye 5.0 mm; stage; postfiexion forms at ca. symphyseal barbel mesopelagic Habitat: Epi- to Br2 gut; trunk arch preanal above photophores of melanophore on at mm; 7.0 ca. at P0 mm; 6.0 POs at ca. 8.5 mm; AOa) at ca. form 2 occur Spawning season: Larvae in year throughout with CalCOFI area, mm. 11.0 ca. abundance April in peaks November & %)* MORPHOMETRICS (range & mean in planktonic eggs Oviparous; pattern: ELH & larvae F Y-S PrF PoF LITERATURE Tr Juv Sn-AlBL 57-60 50-56 56-62 1965 Ahlstrom 53 58 58 56 60 1983 Fahay Moser 1970 Ahlstrom & 11-13 BDIBL 17-27 16-18 12 17 23 26 25 1984a al. et Moser Fortufio & Olivar 1991 27-29 HLlBL 21-25 28-33 1986b, 1988d Ozawa 22 28 30 28 29 1974 Pertseva-Ostroumova Shiganova 1977 53-56 48-60 47-64 HWIHL 55 55 55 63 TAIling 1918 59 SnLIHL & Ahlstrom (1970). in data on based Calculations * Moser 12-18x 18-21x 21-30x EDlHLt first, very narrow Eye in small larvae; horizontal given axis elliptical, is t 33-40 36-39 34-37 second. vertical axis 14x37 26x36 19x37 28x35 39 P L1BL 2 20-31 BbLIHL 6-22 13-33 13 24 22 o o 446

459 Longfin lantern Diogenichthys atlanticlls fish mm 3.6 .• - .. -- =--~.,...,,--,-,,-.-- mm 5.1 6.0mm 7.2mm 12.8 mm 14.5 mm 16.0 mm Figure Myctophidae 30. Preflexion larvae, 3.6 mm, 5.1 mm; flexion larva, 6.0 mm; postflexion larvae, 7.2 mm, 12.8 mm; transformation mm 16.0 (Moser and Ahlstrom 1970). mm; juvenile, specimen, 14.5 447

460 MYCTOPHIDAE Diogenichthys latematus HISTORY DESCRIPTION MERISTICS LIFE EARLY Range Mode EGGS diam.: Vertebrae: Shell Yolk: 29-32 No.ofOG: Total 31 Diam.ofOG: 12-13 12 Precaudal Shell surface: 18-20 Caudal 19 Pigment: features: Diagnostic Fins: spines Dorsal 0 0 10-13 Dorsal rays 11-12 LARVAE Anal mm <2.9 0 spines Hatching length: 0 15-17 rays Anal 16 Flexion ca. 5.2-6.1 mm length: Pelvic 8 ca. mm Transformation length: 10.5-12.3 8 10-12 11 A, Ph Pectoral P C C], sequence: development Fin & D , z z Caudal: ventrolateral mm, 3.0 ca. flexion-By Pre just pair Pigmentation: 10+9 10+9 Principal terminus pair on dorsolateral cleithrum, to posterior on 2 pairs gut, of Procurrellt: midgut, lateral mm, & 3-5 in postanal median ventral series; at ca. 4.5 7-9 8 Upper anterior on margin postanal jaw. of Flexion-5-9 lower in series, 7-9 8 Lower embedded. becoming in postanal Pos(flexion-By ca. 7.0 mm, 10-12 Gill rakers: on each isthmus on pair base, p] on region anterior on 1 (large) series, 3-4 Upper 3 side ventrolateral & of urohyal region, dorsolateral pairs hindbrain, on 10-11 10-11 Lower base at 1 (large) paired series on of lower & Crays, of beginning Brallchiostegais 7 7 mm, gut on laterally >7.7 larvae in 2 more base; A of side each or 1 (some specimens); series. 11-12 in postanal LIFE HISTORY atlanticus; D. lacks barbel to Morphology similar Diagnostic features: but melanophore lacks jaw; on trunk of/ower tip large at pigment has of gut that is present in D. atlanticus; Br preanal arch above Point 33° Pacific from ca. Eastern Conception, California to ca. Range: z pair to form at ca. 4.5 mm; P0 photophores the only other develop Pacific common in eastern tropical minimum waters with low oxygen S; 5 larvae. in to Epi- Habitat: mesopelagic %)* in mean & (range MORPHOMETRIes CalCOFI Larvae season: Spawning in most present year-round area; August & abundant during February-April Y-S PrF PoF F Tr Juv & eggs planktonic Oviparous; pattern: ELH larvae 50-53 Sn-AlBL 52-54 53-58 51 53 56 59 LITERATURE 10-14 BDIBL 14-16 17-26 15 11 23 26 1965 Ahlstrom HLIBL 17-24 23-28 27-32 Castillo 1979 20 26 29 30 1970 & Ahlstrom Moser 61-68 HW/HL 57-65 50-64 Moser et 1984a al. 63 60 56 50 1986b, Ozawa 1988d SnLIHL 15-22x 21-21x EDlHLt 21-26x on based Calculations Ahlstrom * (1970). & Moser in data 34-44 35-38 29-38 larvae; horizontal very is given elliptical, Eye t narrow first, in small axis axis second. vertical 19x39 21x36 24x33 32 P]LlBL PzLIBL 448

461 hth"s laternatus . . Diogenes lanternfish 'or DlOgemc 3.6mm ---~. ~---------.,~- .. /:::::~~~.: ... <~'~~~'--~~~::~" ~~~~~~~~.~.?~.~~:; ; ~.~. ... ,/i· ... ·;.i ·· mm 5.8 6.6mm 11.3 mm I' 3 . 58mm, a mm, . I . . flexIOn arva, 31. PreflexlOn hidae 6 ruv, . Figure Myctop larvae, 6. , (Moser 11 3 mm . postflexion . . juvemle, mm; 12.3 6 mm 1970). and Ahlstrom 449

462 Electrona risso MYCTOPIDDAE LIFE HISTORY DESCRIPTION EARLY MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: Diam.ofOG: 32-34 33 No.ofOG: Total Shell surface: 15 14-16 Precaudal 17 Pigment: 17-20 Caudal Diagnostic features: Fins: o Dorsal spines 0 LARVAE 12-15 14 Dorsal rays < Hatching mm 3.8 length: o 0 Anal spines length: 19 Flexion ca. mm 6.0-7.0 18-20 Anal rays mm 9.5-10.0 ca. Transformation length: 8 8 Pelvic & D A, , C C sequence: development Fin P 16 13-16 Pectoral b 2 2 flexion-None. mm, Pre Flexion-By ca. 6.0 a pair at Pigmentation: Caudal: lower 10+9 PI on a patch & tip jaw developing above mm, 7.0 by blade; 10+9 Principal gas bladder. >9.0 side each 1 on have mm larvae Posiflexion-Some Procurrent: foregut. of 7 6-8 Upper features: to gut slightly Diagnostic sigmoid, extends Stout; ca. 6 6-7 Lower foregut midbody; somewhat becoming relatively saccular; head thick, Gill rakers: at large scanty; but eye pigment broad; narrow; 8-9 transforms & large Upper Br2 photophores 10.0 mm); to form at ca. 5.8 begin size small (ca. 18-21 Lower to similar transformation; at form to first the series PO mm; 8 7-9 Branchiostegals suborbitale & has gut, a shorter Benthosema pigment, which c1eithral on rays. PI lacks pigment LIFE HISTORY (range MORPHOMETRICS in %)* & mean Indian, the in Atlantic, Pacific, Recorded from disjunct localities Range: water central from region Current California enters Oceans; Southern & Y-S PrF F PoF Juv Tr mass Sn-A/BL 47-55 53-58 57-62 58-60 mesopelagic Habitat: Epi- to 52 56 59 59 seasons during present all Spawning season: Larvae in CaiCOFI surveys 16-19 19-21 19-30 BDIBL 29-31 20 24 17 31 pattern: ELH larvae & eggs planktonic Oviparous; 27-29 26-29 28-33 HUBL 32-34 27 28 33 30 LITERATURE 61-72 58-67 HWIHL 68-72 53-58 64 70 66 55 Dekhnik 1966 Sinyukova & SnLIHL Fahay 1983 al. 1989 et Matarese 7-13x 16-19x 18-27x 35-46x EDlHLt Moser Ahlstrom 1970 & 33-41 15-16 35-42 38-49 1984a al. Moser et llx15 18x38 22x38 41x44 in * Moser & Ahlstrom (1970). data on based Calculations second. vertical first, given is axis axis narrow; elliptical, t Eye horizontal 450

463 risso Electrona Chubby flashlightfish 3.9mm 6.3 mm 7.9mm 9.1 mm 9.9mm Figure Myctophidae 32. Preflexion larva, 3.9 mm; flexion larva, 6.3 mm; postflexion larvae, 7.9 mm, 9.1 mm; juvenile, 9.9 mm (Moser and Ahlstrom 1970). 451

464 tenuiculus Gonichthys MYCTOPHIDAE LIFE EARLY HISTORY DESCRIPTION MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: No.ofOG: Diam.orOG: 39 38-41 Total Shell surface: 15 14-16 Precaudal Pigment: 24 23-26 Caudal Diagnostic features: Fins: 0 0 spines Dorsal LARVAE 11 10-12 Dorsal rays Hatching mm <2.5 leugth: 0 0 Anal spines Flexion length: ca. 5.2-6.9 mm 18 17-20 rays Anal 11.5-14.0 length: ca. mm Transformation 7 6-8 Pelvic C P Fin sequence: C, & P" development A, & D & 13 11-14 2 2 Pectoral Pigmentation: flexion-By 3.0 mm, 1 at mid-postana! ventral Pre Caudal: a pair on gut lateral margin, arch, preanal to embedded a blotch & 10+9 10+9 Principal anterior midline of brain (posterior to cerebellum, gut; 3 on to Procunrellt: posterior & forebrain), & at tips of upper anterior & to optic lobes, to 5-6 5 Upper lower the opposing (l midline dorsal at 3 (on) stage, of end by jaws; 5-6 5 Lower blotch postanal ventral 1 & margin, I at preanal arch of gut above Gill rakers: between these), a second at postanal ventral margin anterior to first, 5 4--6 Upper peppering margin on finfold, ventral jaw of dorsal upper & 10 8-12 Lower augmented, Flexion-Above nostril; one at lower & lost. jaw 8-9 9 Branchiostegais embedded P, otic region; on anterodorsal blade; on in head peppering side each on series of beginning base; P, to A dorsal on 1 base; of LIFE HISTORY anterior midline insertion; Dbase & 1 at D to opposing postanal blotches a heavy form bar. Postflexion-At margin posterior on nape; ca. Range: Pacific S 22° & N 32° Eastern between on orbit; of a liver; on region; gular at membrane; branchiostegal gut lateral series; on A rays. to night mesopelagic; at surface at Habitat: Epi- features: Diagnostic deep-bodied highly & becoming slender, Initially gut sigmoid compressed; sharply with terminal section deflected Spawning area, Larvae occur throughout the year in CalCOFI season: ventrad; head large & jaws with relatively small oval eye; elongate with from December to May highest abundance conical tissue pigmented choroid at tip; large median finfold; Bf2 photophores ca. at form 6.5 mm; Br" Bf form to upper Op the first , 2 larvae planktonic Oviparous; pattern: & eggs ELH transformation. at LITERATURE %)* in mean & (range MORPHOMETRICS 1979 Castillo Y-S PrF PoF F Tr Juv Moser 1974 1970, Ahlstrom & al. et Moser 1984a 54--64 Sn-A/BL 40-53 57-66 49 62 59 50 21-32 12-19 data in * 29-34 (1970). Ahlstrom Moser on BDIBL based Calculations & 26 15 33 22 small larvae; horizontal t axis Eye elliptical to oval, very is in narrow given axis second. vertical first, 22-27 29-36 31-37 HL/BL 25 33 35 29 41-52 34-44 36-42 HWIHL 48 42 38 39 SnL/HL 17-22x 13-15x 14-18x EDlHLt 29-39 21-27 18-21 19x34 14x19 15x24 20x22 P LIBL 2 5-19 8-18 18-21 CLIHL 16 19 11 o 452

465 Slendertail lantern fish tenuiculus Gonichthys 3.0mm 4.9mm mm 6.2 -.-y~ ~~ ,. .: / ;' . ~~,/ ~~r/,~... ~ ___ --/ ~~~ 7.7 mm mm 12.2 14.4 mm postflexion Figure 33. Preflexion larvae, 3.0 mm, 4.9 mm; flexion larva, 6.2 mm; Myctophidae larvae, 7.7 mm, 12.2 mm; juvenile, 14.4 mm (Moser and Ahlstrom 1970). 453

466 Hygophum atratum MYCTOPHIDAE DESCRIPTION EARLY LIFE HISTORY MERISTICS EGGS Mode Range Sheil diam.: Yolk: Vertebrae: Diam.ofOG: No.ofOG: 36 35-37 Total Shell surface: 16 15-l7 Preealldal Pigment: 20 19-21 Calldal features: Diagnostic Fins: 0 0 spines Dorsal LARVAE 12-13 11-14 rays Dorsal mm length: <2.4 Hatching 0 0 spines Anal 7.8-9.7 ca. length: mm Flexion 19-20 18-21 Anal rays 11.8-14.5 ca. mm Transformation length: 8 8 Pelvic P, Fin D, A, & C, development P sequence: C 13-14 12-15 lo lo Pectoral flexion-By 3.0 mm, ventrolateral pair just posterior Pigmentation: Pre Calldal: c1eithrum, to on terminus dorsolateral pair of 2 lateral gut pairs, gut, 10+9 10+9 Pl"incipal 2 ventral on postanal tandem margin, on isthmus, in 2 widely-spaced ProcllrJrent: lower 1 at mm, 5.0 by margin; dorsal at I & hypural 1 at tip; jaw 8 6-9 Upper posterior on dash end of stage, flnfold; anlagen; some by ventral on 8 6-9 Lower isthmus divided into pair, lateral 1-3 more on gut, & a postanal pair Gill rakers: 4-5 on each upward side in ventral margin. extending myosepta from 4-6 Upper on isthmus; 1 on opercle; up to Flexion-lor 2 more median dashes 15-16 14-17 Lower gut; beginning on 8 laterally on of Posiflexion-Up base. A to series 9 8-9 Rranchiostegals on 8 laterally up dashes; myoseptum 3 postanal to A on 15 to up gut; base. LIFE HISTORY thin, featllres: Very slender body; gut elongate, Diagnostic & nearly eyes straight; head flattened; narrow, elliptical short stalks; conical on distributed from Eastern ca. 30° N to in Range: widely S; 30° Pacific jaw lower genus; of tissue; choroid ventral typical pigment isthmus minimum oxygen low with waters Pacific tropical eastern CaiCOFI melanophore (absent in H reinhardtii larvae in present to (up fewer area); 3) with compared dashes myosepta postanai H surface at mesopelagic; to Epi- Habitat: night at reinhardtii (up to first the photophores VO. PO" POlo Brio 10); to & fewer (35-37) transformation; at appear total vertebral lower count throughout present Larvae season: Spawning area, with in year CaiCOFI with (18-21) A rays compared 21-25). & (38-40 reinhardtii H November in abundance peak in MORPHOMETRICS (range & %)* mean pattern: larvae ELH planktonic Oviparous; eggs & y-s PrF Tr Juv F PoF LITERATURE 54-63 62-66 52-60 Sn-AlBL 66-67 1970 Ahlstrom & Moser 61 55 65 66 al. et Moser 1984a 19-22 9-14 9-10 BD/BL 10-16 10 12 9 21 on (1970). Ahlstrom & * Calculations based Moser data in 23-25 25-29 19-27 26-29 HLIBL t elliptical, very narrow; horizontal axis is Eye given first, vertical axis 23 27 26 24 second. 47-58 42-48 35-44 47-56 HWIHL 40 45 53 53 SnLIHL 11-22x 11-13x 18-34x 11-13x EDlHLt 20-22 22-34 22-44 21-23 14x29 12x22 12x2! 29x30 P,L/BL 7-8 0-8 0-27 CLIHL 0-4 8 6 5 454

467 Thickhead ianternfish atFatum Hygophum .~-~.~~ ; b)L±,,""';jj\~ 2.3 mm 5.2 mm 8.4 mm mm 13.8 mm 13.2 . ). flexion larva, . m , I nd ventral arva, Preflexion 4 . 2 VIews , . . mm (latera a mm, e 23 I 5 mm; 13.8 larva, postflexion 84m' Figure Myctophldae 3. nd Ahlstrom 1970) . . (Moser 13 2 mm , a juvem e, . 455

468 MYCTOPHIDAE Hygophum proximum HISTORY DESCRIPTION LIFE EARLY MERISTICS EGGS Range Mode diam.: Shell Yolk: Vertebrae: No.ofOG: 36 34-37 Diam.ofOG: Total Shell surface: 15-16 15-16 Precau!lal Pigment: 18-21 21 Caudal features: Diagnostic Fins: 0 spines Dorsal 0 12-14 LARVAE 13 rays Dorsal 2 mm ca. Hatching length: 0 Anal 0 spines Flexion mm 6.1-7.6 length: 18-21 19-20 rays Anal 8 Transformation ca. 12 mm length: 8 Pelvic Fin development C sequence: , , , P A & C D, P 14 13-15 Pectoral 1 1 2 2 on pair ventrolateral mm, 3.0 flexion-By Pre Pigmentation: gut Caudal: 10+9 c1eithral junction, 1 or 2 lateral posterior on pair to along gut, & pairs 10+9 Principal terminal section of gut; between 3.0-4.0 mm, hypaxial on pair region Procurrent: 8-9 to anus, 1 or more in dorsal just ventral finfolds (some & 8-9 posterior Upper 8-9 8-9 just on pair & isthmus by symphysis; c1eithral to anterior specimens), Lower on 6.0 mm, a median embedded dash isthmus. on Flexion-None Gill rakers: 4-5 4-5 another Postflexion-By 8.0 mm, hypaxial region or mid-lateral gut. Upper embedded isthmus. dash median on 13 12-15 Lower Diagnostic body; compressed Moderately head features: Brallchiostegals & deep, choroid conical with eyes narrow moderately gut ventral tissue; LIFE HISTORY at & sigmoid; pigment; pigment dashes on isthmus slightly sparse mm. 6.7 ca. at Sr photophores appear symphysis; c1eithral 2 Widespread in tropical Range: subtropical Pacific & Indian Oceans & & mean in %) MORPHOMETRICS (range Epi- mesopelagic to Habitat: PrF Y-S PoF F Tr Juv year-round Spawning season: Possibly Sn-A/BL 57-63 61-64 57-63 53-57 61 62 61 55 planktonic & larvae eggs ELH Oviparous; pattern: 12-14 BDIBL 14-17 18-27 21-26 LITERATURE 13 16 23 23 21-24 HLIBL 25-26 22-28 28-29 aI. et 1979 Miller 23 25 25 29 Moser & 1974 Ahlstrom Moser et aI. 1984a 55-63 55-73 57-65 HWIHL 49-51 Fortuno Olivar & 1991 61 59 61 51 1986b, Ozawa 1988d SnLIHL 20-29 30-33 29-35 14-18 Pertseva-Ostroumova 1974 31 31 24 16 (Illustrator) ILLUSTRATIONS ORIGINAL EDIHL* 14-17x 19-20x 21-29x 37-38x 30-38 31-40 29-33 37-43 4.7 larvae, 3.3 mm, flexion Pre (N. Arthur) mm 15x33 25x35 19x31 38x39 (N. Arthur) Flexion larva, 6.3 mm, lateral view 10-12 4-7 7-8 LIBL P1 mm, flexion view (N. Arthur) larva, 6.3 ventral 5 11 7 26 mm 12.8 Juvenile, (N. Arthur) LIBL P 0-0 0-0 0-2 2 0 0 0.5 10 in small larvae; horizolltal axis is given elliptical, * Eye narrow very first, second. axis vertical 456

469 Hygop/tum proximum mm 4.7 6.3 mm mm 6.3 8.9mm 12.8 mm Figure Myctophidae 35. Preflexion larvae, 3.3 mm, 4.7 mm; flexion larva, 6.3 mm, lateral and ventral views (CaICOFI 7205, station (EASTROPAC I, station 12.221). 1974); (Moser and Ahlstrom postflexion juvenile, 12.8 mm 20.127); larva, 8.9 mm 457

470 MYCTOPHIDAE Hygophum reinhal'dtii MERISTICS EARLY LIFE HISTORY DESCRIPTION Mode EGGS Range Shell Vertebrae: diam.: Yolk: 38-40 Total No.orOG: 39 Diam.ofOG: 16-17 Shell surface: 17 Precaudal 21-23 Caudal 22 Pigment: Diagnostic features: Fins: spines Dorsal 0 0 Dorsal LARVAE 14-15 13-15 rays length: 0 spines Anal Hatching 0 mm <3.4 21-25 22-23 8.8-10.3 Anal ca. length: Flexion mm rays Pelvic 8 8 mm 14.9-16.4 ca. length: Transformation 13-15 14 Pectoral Fin P & D A, PI> & C CI> sequence: development 2 2 Calldal: pattern Pre except atratum Pigmentation: to flexion-Similar of H 10+9 10+9 Principal none at lower jaw tip (most specimens), end of & by stage, 1-4 more ProClirrent: Flexion-7-12 in outlining hypaxial myosepta. postanal in series 7-9 8 Upper series. postanal in Posiflexion-5-1O series. postanai 7-8 Lower 8 features: Nearly identical Diagnostic to H in morphology atratum; Gill rakers: undergoes notochord flexion & transformation at slightly size; larger 4 4-5 Upper atratum); (38-40 vs. lower H in 35-37 vertebral count total higher 13-15 14 Lower jaw pigment present is atratum H in H. most in present not 8-9 Branchiostegals 9 postanal in reinhardtii more region; CalCOFI from melanophores (maximum series pigment myosepta H. 4 in of maximum vs. 12 of LIFE HISTORY higher (18-21). atratum atratum); ray A H count (21-25) than in (range MORPHOMETRICS %)* in mean & Range: Oceans; Pacific & Atlantic subtropical southern & Northern central water mass into CaiCOFI area intrudes from eastward Y-S PrF F PoF Tr Juv to Epi- mesope!agic; Habitat: at surface at night Sn-A/BL 55-63 62-67 60-65 58-60 59 62 throughout CalCOfI Spawning Larvae 64 the year in season: occur area; 59 53 in highest abundance in the summer & fall, with peaks August & BDIBL 7-10 7-8 8-15 19-21 November 7 9 11 20 20 HLIBL 21-27 23-24 24-25 24-24 & eggs Oviparous; planktonic larvae ELM pattern: 23 23 25 24 26 HWIHL 47-60 42-45 39-48 54-60 LITERATURE 43 51 42 62 57 SnL/HL 1983 Fahay Moser & 1974 Ahlstrom 1970, 8-15x 9-9 x 9-16x EDlHLt et Moser 25-27 al. 1984a x 22-36 20-21 19-25 31-32 Fortuflo Olivar & 1991 Ozawa 1988d 1986b, 9x20 lOx26 13x22 26x32 43 Shiganova 1977 in Ahlstrom Moser & (1970). * Calculations based data on Eye first, vertical axis axis given horizontal t narrow; elliptical, very is 4-9 CLIHL 6-9 7-9 0-4 second. 7 8 8 2 o 458

471 Slender lanternfish reinhardtii Hygophum 3.4 mm ,--.--..r/~-.:.-...~~_ ' ... ~ ... ------"'!l;~~~~~@:::3~ .. ~············~:§?!X,.::> ...--- 6.7mm :I! ... .. ... ~ ... ~ .. :~:~ ...• ~JI!.. ••• _".,. ¥. .. ~ 7.4mm 12.8 mm 14.9 mm 16.0 mm Figure Myctophidae 36. Preflexion larvae, 3.4 mm, 6.7 mm; flexion larva, 7.4 mm, lateral and ventral views; postflexion larva, 12.8 mm; transformation 14.9 mm; juvenile, specimen, 16.0 mm (Moser and Ahlstrom 1970). 459

472 MYCTOPHIDAE Loweina rara mSTORY DESCRIPTION EARLY LIFE MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: 37-39 Diam.ofOG: No.ofOG: 38 Total 17-19 Shell surface: 18 Precaudal 21 Pigment: 19-21 Caudal features: Diagnostic Fins: 0 0 Dorsal spines 11 LARVAE 10-13 rays Dorsal mm Hatching length: <2.8 0 0 spines Anal 8.4-10.8 15 length: ca. Flexion mm 13-17 Anal rays Transformation length: ca. mm 20.0-21.0 8 8 Pelvic 9-13 10-11 Pectoral , A, Fin PI' C P sequence: D & C development b 2 2 bar Pigmentation: 4.0 mm; a heavy transverse flexion-By between Pre Caudal: 10+9 fore- 10+9 base, PI to anterior a blotch embedded an midbrain, & blotch Principal on midgut, embedded above a blotch surface dorsal of terminal section Procurrent: 6 & of gut, apposing dorsal ventral & region, mid-postanal in blotches 6-7 Upper mm, dorsal blotch; by 6.0 more a median blotch 6 another anterior 6-7 Lower embedded melanophore at nape, isthmus, in & a blotch an embedded Gill rakers: 2 ray; PI lower elongate of by stage, of 2 end spatulate on swellings Upper embedded blotches at each expanded other 7 6-7 isthmus PI base & toward Lower 8-9 lost blotches postanal & cavity gill in of 8 ventral outline to border Branchiostegals in some blotches (large) Pos!'flexion-Numerous specimens. finfold; several embedded in postanal hypaxial myosepta; LIFE lllSTORY voluminous PI base. several on & Compressed body features: Diagnostic with voluminous dorsal regions Range: Atlantic, Pacific, & Indian Oceans; of Tropical-subtropical oval wide relatively head; ventral large finfolds; ventral no eyes; ca. to N 30° S from Pacific 30° in eastern large with gut elongate tissue; choroid large section; terminal PI with A & distinctive far posteriad; ornamented lower ray; D elongate night to mesopelagic; at Habitat: Epi- surface at pigment transverse pattern, including bar between fore- & midbrain in Tarletonbeania crenularis). (lacking area; CalCOFI in Spawning throughout season: year present Larvae in abundance highest March %)* mean & (range MORPHOMETRICS in pattern: ELH larvae & eggs planktonic Oviparous; Y-S F PrF PoF Juv Tr LITERATURE Sn-AlBL 73-81 66-74 54-66 70 59 77 59 1983 Fahay et 1989 al. Matarese 13-17 19-23 23-30 BDIBL 20 15 26 Moser 1974 & Ahlstrom 1970, 26 Moser et al. 1984a 22-30 HLlBL 28-34 29-32 Olivar Fortuno 1991 & 26 30 32 28 HWIHL 48-60 41-48 35-52 44 44 53 39 (1970). data on based & Calculations * Ahlstrom Moser in SnLIHL vertical axis given is second. first, oval; t Eye horizontal axis x 20-23 19-23x 19-20x EDlHLt 29-37 25-30 23-32 22x33 20x28 20x27 21 PILlBL P L1BL 2 460

473 Dwarf fish lantern Loweina rara 4.8mm 8.5 mm 24.5 mm Figure Myctophidae 37. Preflexion larva, 4.8 mm; flexion larva, 8.5 mm; postflexion larva, 17.6 mm; transformation specimen, 20.0 mm; juvenile, 24.5 mm (Moser and Ahlstrom 1970). 461

474 MYCTOPHIDAE Myctophum aurolaternatum EARLY LIFE HISTORY DESCRIPTION MERISTICS Mode EGGS Range Shell Vertebrae: diam.: Yolk: Total 43-44 No.ofOG: 42-46 Diam.ofOG: Shell surface: 18---19 18 Precaudal Pigment: 25-26 25 Caudal Diagnostic features: Fins: spines Dorsal 0 0 11 LARVAE 9-12 Dorsal rays length: mm Hatching <3.8 0 Anal 0 spines 21-27 24 length: 11.7-13.8 ca. Flexion Anal rays mm 8 Pelvic 8 Transformation length: ca. 27-30 mm 13-15 12-16 Pectoral A, C" PI & C & D, P development Fin sequence: 2 2 Caudal: dorsum mm, 2 (apposing) on flexion-By & Pre 4.0 Pigmentation: 10+9 Principal 10+9 ventrum near notochord tip, 2-6 gut, on laterally 4-6 on dashes paired Procurrent: isthmus, c1eithrum, to posterior just dashes ventrolateral paired 1 2-4 Upper 8 8 surface of gut, on tip & lateral lower of several section terminal on to 7-8 7-8 Lower jaw, of edge lower on several & at ventral margin, postanally 2-3 gill rakers: Gill cover; by on dashes 6.0 mm, 3-4 isthmus, 2-5 at postanal ventral Upper of laterally 7-10 preflexion, by end several fmfold; ventral on margin, on gut, Lower 2-3 at blotches apposing midbrain, of margins lateral on Branchiostegals notochord (dispersed to hypural region). tip at postanal Flexion-6-9 A of on series jaw; upper ventral margin, margin anterior series on LIFE HISTORY rays; on trailing gut. peppering Pos!flexion-Postanal & lateral gut series augmented, peppering basally regions lower & upper on & of series C; to anterior finfold on rays; branchiostegal on C. eastern Range: Tropical-subtropical Pacific & Pacific Indian Oceans; in slender (42-46); body features: High total vertebral count Diagnostic 25° S 17° & N between gut gut; & elliptical flattened; head stage; flexion at trailing becomes D on moderately long stalks; eyes forms at margin of finfold; Epi- Habitat: at night depth m 300 to surface at rnesopelagic; to 30 mm). distinctive large size (up to ca. attains pigment pattern; Larvae Spawning season: collected January-April & September- (range & in %) MORPHOMETRICS mean November in CalCOFI area pattern: Y-S larvae & eggs planktonic Oviparous; PrF ELH F PoF Tr Juv LITERATURE Sn-A/BL 50--71 65-114 103-133 53-54 86-116 89 60 119 101 54 & Ahlstrom 1974 Moser BDIBL 6-8 6-9 6-11 12-13 19-20 7 7 8 12 Moser 1984a et al. 19 1986b, 1988d Ozawa 20--26 HLlBL 20--22 19-23 21-21 26-27 22 23 21 21 27 (Illustrator) ILLUSTRATIONS ORIGINAL 39-54 HWIHL 38-43 47-48 37-44 41-47 55 40 41 48 43 Preflexion larvae, 5.8 mm, 4.0 (B. MacCall) Sumida mm SnUHL 20--27 20--27 18-22 18-26 14-18 Sumida MacCall) mm (B. 12.5 Flexion larva, 22 23 23 20 16 Postflexion larva, 25.8 MacCall) Sumida mm (B. 11-19x 12-15x 12-14x EDIHL* 14-15x 27-30 mm MacCall) Sumida (B. 25.9 Juvenile, 15-30 18-21 17-19 17-18 14x22 13x19 13x18 14x18 29 * first, vertical given axis second. is axis horizontal elliptical; Eye 3-5 4-5 6-8 16-17 PILlBL 4 4 7 16 9 P L1BL 0--0 0--0 0.5-2 10--11 2 0 0 2 0.8 11 ESLIHL 24-63 53-64 31-59 22-25 0--0 52 58 46 24 0 462

475 Golden lanternfish aurolaternatum Myctophum 4.0mm 5.8 mm mm 12.5 25.8 mm 25.9 mm Figure Myctophidae 38. Preflexion larvae, 4.0 mm (EASTROPAC II, station 47.005), 5.8 mm (CaICOFI 7210, station 157.45); flexion (EASTROPAC II, station 46.187); juvenile, 25.9 mm postflexion larva, 25.8 mm 12.5 larva, mm (CaICOFI 7210, station 157.50); (EASTROPAC I, station 12.018). 463

476 MYCTOPHIDAE Myctophum nitidulum HISTORY EARLY LIFE DESCRIPTION MERISTICS Mode EGGS Range diam.: Shell Yolk: Vertebrae: No.ofOG: 38 Diam.ofOG: Total 36-39 15 Shell surface: 15-16 Precaudal 21-22 Pigmeut: 21-23 Caudal features: Diagnostic Fins: 0 0 Dorsal spines 12-14 LARVAE 13 rays Dorsal mm Hatching length: <3.1 0 0 spines Anal ca. length: Flexion 6.5-7.0 mm 19 18-21 Anal rays 11 ca. length: Transformation mm 8 Pelvic 8 Fin development sequence: , P" 13-16 C 14-15 D P A, & C & Pectoral I 2 2 1 jaw, lower of tip on mm, <4.0 larvae flexion-In Pre Pigmentation: Caudal: 10+9 10+9 ca. ventral anterior on 2 pairs midline, ventral postanal along midway Principal on ca. of gut, a dorsolateral pair terminal gut, section of 4 surface Proem'reut: surface (ring-like) on medial peppering 7-9 8 of PI base, & at blade; PI on Upper jaws, series on isthmus & upper mm, lower 4.0 1 on nostril, at & ca. 7-9 7 Lower region posterior on hindbrain, on & opercie, of orbit, ventrolaterally Gill rakers: dorsal on midline by margiu; ventral on one the to posterior slightly 5-8 6 Upper head above PI base, a series ventrally on cleithrum, a 6.0 1 on 13-19 16 mm, Lower ray, pairs on ventral series associated 2 branchiostegal each with 9 9-10 Brancliiostegals surface of & terminal section of gut, 1 at A insertion. Flexion-Dne midline more nostril, 1 or series more most brain; above on in HISTORY LIFE of Posiflexion-Beginning vertical series at posterior augmented. margin. hypural in in Oceans; Indian & Atlantic Pacific, Range: Tropical-subtropical body robust broad massive head & gut; Diagnostic features: Stout with 0 0 N & 31 S ca. between Pacific 32 precocious tissue; PI choroid oval on short stalks; conical ventral eyes large fan-shaped with base forms P, blade; & early; & highly complex surface at night at to Epi- Habitat: mesopelagic; pigmentation; distinctive mm. Br2 photophores form at ca. 7.0 season: area; in year Spawning throughout present CalCOFI Larvae the %)* in mean & (range MORPHOMETRICS highest abundance the in August in peak with fall, & summer late PrF Y-S F PoF Tr planktonic eggs & ELM larvae pattern: Oviparous; Juv 48-57 Sn-AlBL 59-68 51-54 62-67 LITERATURE 54 63 65 53 BDIBL 20-27 Castillo 1979 14-19 24-30 20-21 22 27 17 21 Fahay 1983 Ahlstrom 1974 1970, & Moser 32-37 33-37 HLIBL 24-33 27-28 Moser et aI. 1984a 35 29 35 27 1991 Olivar & Fortufio 60-78 56-68 HWIHL 56-64 49-54 Ozawa 1986b, 1988d 63 60 67 51 SnLIHL in on Calculations * data based Moser & Ahlstrom (1970). 14-18x 13-15x 13-16x EDlHLt 22-31 very horizontal axis Eye given somewhat oval, is narrow in small larvae; t 23-26 24-37 21-24 first, vertical axis second. 14x24 15x23 15x30 27 P,UBL 8-11 7-13 4-9 CUHL 0-0 6 9 10 o 464

477 Pearly lanternfish Myctophum nitidulum --- -~ ,. ... . -- .. .--_ '\. 3.8mm . -"-- ... , \ ~ / / ... , ./, .. . ... ----_ ... 5.8mm 7.0mm 0 0 1IIf:lJ!;o ~l~\~"\~~~~~ mm 18.8 Figure Myctophidae 39. Preflexion larvae, 3.8 mm, 5.8 mm; flexion larva, 7.0 mm; postflexion larva, 11.7 mm; juvenile, 18.8 mm (Moser 1970). and Ahlstrom 465

478 MYCTOPHIDAE Protomyctophum crockeri MERISTICS EARLY LIFE lllSTORY DESCRIPTION EGGS Mode Range Shell Vertebrae: diam.: Yolk: Total 36-38 37 No.ofOG: Diam.ofOG: 14-16 Precaudal 15 surface: Shell 21-23 Pigment: 22 Caudal Diagnostic features: Fius: 0 0 spines Dorsal LARVAE 11-13 12 Dorsal rays Hatching length: mm 3.8 0 0 < spines Anal 21-22 length: Flexion 7.2-8.8 ca. 19--24 mm rays Anal mm 15.2-17.6 ca. length: Transformation Pelvic 8 8 13-17 15-16 Pectoral D ; P A; PI; C" c;; sequence: development Fin 2 Caudal: flexion-By Pre ca. 5.0 of side each on I (large) mm, Pigmentation: 10+9 10+9 Principal one at gut enlarges & is more midgut. The Flexion--pos{/lexion- Procurrent: ca. anterior; by gas bladder. mm, 10.0 patch above 7-9 8 gut; foregut somewhat Diagnostic features: Slender body; short Upper 7-8 7 Lower conical; small relatively head & in pointed stages; flexion preflexion rakers: Gill elliptical snout; choroid ventral moderately narrow lacks eye, tissue; 5 Upper 4-6 patch & above gas bladder; Br2 gut on laterally only scanty, pigment 16 14-18 Lower series, at ca. 8.0 PO form Br3 & upper OP the first photophores mm; 8-9 at form to transformation. 9 Branchiostegals mean & (range MORPHOMETRICS in %)* LIFE lllSTORY Y-S Pacific Range: Subarctic North & widespread in California Current, south PrF F PoF Tr Juv zone across N; 25° northern transition Japan westward to to Sn-AlBL 35-39 35-52 34-39 37 41 36 51 57 Epi- to mesopelagic Habitat: BDIBL 7-12 11-13 12-20 season: year occur Larvae throughout Spawning area; CalCOFI in highest 12 10 15 24 26 abundance in peak with winter, the December in HUBL 17-20 17-19 19-25 18 18 21 29 31 larvae Oviparous; planktonic ELH eggs & pattern: HWIHL 50-61 59-65 52-67 55 62 59 50 49 LITERATURE SnLIHL et Matarese al. 1989 18--23x 21-29x x 20--23 EDlHLt 1970 Ahlstrom & Moser 40--46 41-46 38-44 et Moser 1984a al. 19x43 21x43 1988d Ozawa 25x42 28x39 28 PILlBL * (1970). Ahlstrom & Moser in data Calculations based on L1BL P is larvae; to oval, very narrow in small horizontal t Eye axis elliptical 2 given first, vertical second. axis 466

479 crockeri Protomyctophum 4.1 mm mm 7.6 12.4 mm 16.6 mm mm 15.5 mm 17.8 . m , 12.4mm,16. . flexion larvae, tflexion larva, m· 76mm;pos specimen, transformation 6 mm, 4.1 70) . larva . Myctophldae. an (Moser Ahlstrom 19 d PreflexlOn 40 . . mm 17.8 J·uvenile, m· ure Flg 5 467 m , 15.

480 MYCTOPHIDAE Protomyctophum thompsoni EARLY LIFE HISTORY DESCRIPTION MERISTICS EGGS Mode Range Shell diam.: Vertebrae: Yolk: 37-39 No.ofOG: Diam.ofOG: Total 38 Shell 15 surface: 14-16 Precaudal 23 22-24 Pigment: Caudal Diagnostic features: Fins: 0 0 Dorsal spines LARVAE 12 11-13 Dorsal rays mm <2.8 length: Hatching Anal spines 0 0 length: mm ca. 7.9-9.9 Flexion 21-25 23 Anal rays length: Transformation mm 17.2-18.8 ca. 8 Pelvic 8 14-17 Fin seqllence: development D A, PI> C P C 15 & , Pectoral b 2 2 Pigmentation: series. in median Preflexion-lO-14 postanal ventral Caudal: 10+9 coalesced laterally 1-3 6-11; foregut. on series Flexion-Postanal to 10+9 Principal maximum a series reduced Procm-rcnt: Posiflexion-Postanal at 9 of from 8 7-9 of Upper flexion bladder gas above mm; 17.0 by none to stage post beginning 7 6-8 Lower 13.0 mm; beginning at vertical 13.5 mm, 1-4 embedded in ca. ca. by series region. hypura\ lateral in Gil! rakers: 3-4 4 similar to P. crockeri but Diagnostic features: Morphology Upper 14 13-16 crockeri); P. (absent different; pigmentation series present in postanal Lower 9 8-9 Branchiostegals 1 in crockeri); P. than more (no Br2 meianophores gut lateral 1-3 series at ca. OP upper & photophores PO mm; 8.5 photophores appear HISTORY LIFE at form to first the transfonnation. in (range & %)* mean MORPHOMETRICS zone Pacific, North Sea; Bering including of Range: Subarctic-transitional N 38° in California Current south to ca. Y-S PrF F PoF Tr Iuv mesopeiagic Habitat: Epi- to 33-43 Sn-AlBL 35-48 36-40 52-52 40 41 38 52 collected Larvae Spawning season: January-February, May, & October area CalCOFI in 12-20 12-13 8-13 BD/BL 26-27 15 10 13 26 Oviparous; ELM pattern: planktonic eggs larvae & 18-21 17-21 19-26 HLIBL 28-29 19 20 21 29 LITERATURE HWIHL Matarese et 1989 aI. 55-65 52-62 SnLIHL 54-64 54-54 1970 Ahlstrom & Moser 58 59 58 54 et 1984a Moser aI. 20-22x 20-24x 21-28x EDlHLt 42-44x Ozawa 1988d 37-42 41-44 37-42 46-46 1967 Pertseva-Ostroumova 1964, 21x42 22x39 23x40 43x46 (1970). Calculations * in Moser & on Ahlstrom data based elliptical, vertical given is horizontal narrow; first, Eye t second. axis axis LIBL P 2 468

481 Northern flashlightfish thompsoni Protomyctophum - ... ... . mm 4.2 _ .. _"-' ;!!.!/W":j ~ - ... : .,... : .. ... g, ----- .. "- -. --=------- 8.8mm 11.0 mm 13.8 mm 18.6 mm mm 17.7 Figure Myctophidae 41. Preflexion larva, 4.2 mm; flexion larva, 8.8 mm; postflexion larvae, 11.0 mm, 13.8 mm; transformation specimen, 18.6 mm; juvenile, 17.7 mm (Moser and Ahlstrom 1970). 469

482 Symholophorus californiensis MYCTOPHIDAE LIFE EARLY HISTORY DESCRIPTION MERISTICS EGGS Mode Range Yolk: Shell diam.: Vertebrae: No.ofOG: Diam.ofOG: 37-40 39 Total surface: Shell 15-17 16 Precaudal Pigment: 22-23 22-23 Caudal features: Diagnostic Fins: 0 0 Dorsal spines LARVAE 14 13-15 Dorsal rays length: Hatching mm <3.8 0 0 spines Anal Flexion length: ca. 8.5-10.0 mm 20 19-21 rays Anal Transformation length: ca. mm 24 8 8 Pelvic P & & C PI> sequence: development Fin D , C A, 17-18 15-20 I 2 2 Pectoral Pigmentation: mm, ventrolateral Pre just flexion-By ca. 4.0 pair Caudal: to anterior gut, on laterally pairs 3-4 c1eithra, on pairs 2 dorsolateral 10+9 10+9 Principal 11 of extending series gut, up to median in a ventral section terminal Procurrent: from above preanal arch region; of gut to mid-postanal a mm, 5.0 by 9 8-10 Upper anterior edge oflower jaw on pair P & ; by 7.0 mm, peppering up on 7 2 7-8 Lower 3 pairs lie vertically isthmus, to on c1eithral pair on pair c1eithra, on rakers: Gill ventrolateral region (some tip jaw lower at hindbrain, pair of 6 6--7 Upper margin outlining dorsal series specimens), distally base, & some PI of 16 15-17 Lower on & gut PI end of stage, lateral Flexion- rays; postanal series lost. by 9 9-10 Branchiostegals ventral margin of PI base; 8.0 mm, series ca. along posiflexion-By PI blade. bar on LIFE HISTORY Early becoming slender, preflexion Diagnostic larvae features: long, moderately robust in postflexion stage; gut somewhat nearly across to ca. 25° S; westward Current the Pacific; North Range: California large, elliptical flattened; head eyes straight; & slightly stalked; conical Japan to zone transition tissue; precocious, choroid paired fins pigmented; distinctively large, photophores PI base aliform; much of early pigmentation lost; Br2 mesopelagic; Habitat: Epi- to at surface at night at form ca. transformation. mm; POI & PO 9.0 first to form at s in throughout present CalCOFI Spawning season: Larvae the year area; MORPHOMETRICS & mean in %)* (range with summer, spring~arly April-May in peak highest abundance in PoF Y-S PrF F Tr Juv eggs & larvae ELH pattern: planktonic Oviparous; 62-67 50-65 63-66 Sn-AlBL LITERATURE 64 64 57 58 8-13 13-17 16--23 BDIBL Ahlstrom 1965 15 11 20 19 Matarese et al. 1989 Moser 1974 & Ahlstrom 1970, 28-30 18-28 24--32 HUBL Moser et 29 al. 1984a 29 24 28 Ozawa 1988d 55-67 52-64 56--60 HWIHL 1974 1964, Pertseva-Ostroumova 61 59 44 58 SnLIHL in * based on data Calculations Moser & Ahlstrom (1970). elliptical t Eye small to oval, very narrow in is larvae; horizontal axis 13-16x 16--17x 17-26x EDlHLt vertical first, given second. axis 22-33 25-27 25-39 17x26 20x26 14x30 21 7-8 6--10 4--9 CLIHL 7 8 2 7 470

483 California lanternfish californiensis Symbolophorus 3.9mm 5.0 mm mm 7.2 mm 11.5 23.8 mm 23.6 mm Figure Myctophidae 42. Preflexion larvae, 3.9 mm, 5.0 mm, 7.2 mm (lateral andventral views); postflexion larvae, 11.5 mm, 23.8 mm; juvenile, 23.6 mm (Moser and Ahlstrom 1970). 471

484 MYCTOPIDDAE Symholophorus evermanni* DESCRIPTION EARLY LIFE HISTORY MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: 37 No.ofOG: Diam.ofOG: Total 36-38 15-16 Shell surface: 15 Precaudal 21-22 21-22 Pigment: Caudal features: Diagnostic Fins: 0 0 spines Dorsal LARVAE 13-16 15 rays Dorsal 0 <3 mm Hatching length: Anal spines 0 mm 19 18-22 length: Flexion 6.1-7.5 Anal rays length: mm 20 ca. Transformation 8 8 Pelvic 15-16 Fin development sequence: & 13-18 , D, PI' C P , A C Pectoral I 2 2 postanal Pigmentation: flexion-By 3 mm, up to 2 on Pre ventral Caudal: margin, pair on terminal section of gut, 0-2 10+9 along gut 10+9 pairs Principal symphysis; side), & pair at c1eithral one by missing on (sometimes 5 Procurrent: 8-9 pair have some & symphysis jaw 9 isthmus. on mm, pair at lower Upper 8 7-9 basally & Flexion-posiflexion-Series postanal on PI rays; lacking on Lower gut regions. rakers: Gill 5-7 6 Aliform Diagnostic features: flattened; Upper PI base; head large, slightly 15-16 early stalked eyes in preflexion to postflexion stages; elliptical narrow 14-17 Lower preflexion stage, becoming oval in postflexion eyes in conical stage; Branchiostegals 7.0 Br2 form at ca. photophores choroid tissue; pigment sparse; ventral mm. LIFE HISTORY & in %) (range MORPHOMETRICS mean Range: Pacific complex a species apparently, Ocean; Indian & Tropical Y-S PrF F PoF & mesopelagic Epi- Habitat: Tr Juv Sn-AlBL year-round 56-62 62-64 59-64 Possibly season: Spawning 48-59 62 63 59 55 ELH larvae planktonic Oviparous; pattern: 12-13 BDIBL 13-17 16-23 17-20 12 15 20 19 LITERATURE 26-29 28-29 27-29 HUBL 25-30 28 28 28 28 Beckley & 1994 Olivar 67-71 61-70 HWIHL 56-67 43-44 1988d 1986b, Ozawa 65 69 62 44 Pertseva-Ostroumova 1974 1964, SnLIHL 15-35 29-35 23-33 16-21 25 33 28 19 (Illustrator) ILLUSTRATIONS ORIGINAL 12-14x 17-22x 14-15x EDIHLt 22-26 32-32 27-32 28-31 mm larvae, Preflexion 5.2 (N. 3.8 Arthur) mm, 13x30 14x32 20x30 24 larva, flexion Late Arthur) (N. view lateral mm, 7.5 mm, Late flexion larva, 7.5 ventral view Arthur) (N. 11-12 6-10 12-16 PILlBL mm (N. Postflexion larva, Arthur) 20.0 8 11 15 13 Arthur) mm (N. Juvenile, 24.5 L1BL P 0.9-10 0-0 0--0 11-13 2 0 0 12 5 S. putatively larvae appear to be All evermanni; another * type, a single undescribed extends into Symbolophorus the species from area CalCOFI Pacific 1993) & its larvae may be similar to (Gago equatorial the North of S. evermanni. those larvae; very t Eye elliptical, small in horizontal axis is given first, narrow vertical axis second. 472

485 evermanni Symholophorus Evermann's lanternfish mm 3.8 >,;'.:' .•. · ... : .• _·_.· ..• _ .•..• ---~-·-··~~.;.~.7 ... ... "::,:~";;'.:.~" ~ mm 5.2 mm 7.5 ~ -:.>-- -- ~- :::. ----------- -- 20.0 mm 24.5 mm Figure Myctophidae 43. Preflexion larvae, 3.8 mm, 5.2 mm (CalCOFI 7205, station 24.137); late flexion larva, 7.5 mm, lateral and ventral (CalCOFI 7205, station 24.137); postflexion larva, 20.0 mm 7205, (CalCOFI station 31.135); juvenile, 24.5 mm (EASTROPAC I, views station 11.098). 473

486 Tarletonbeania crenularis MYCTOPHIDAE DESCRIPTION EARLY LIFE HISTORY MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: No.ofOG: Diam.ofOG: 39-42 41 Total surface: Shell 17 17-18 Precaudal Pigment: 24 22-24 Caudal features: Diagnostic Fins: 0 0 spines Dorsal LARVAE 12-13 11-14 rays Dorsal mm <3.0 length: Hatching 0 0 Anal spines mm 7.8-10.5 ca. length: Flexion 18 17-20 Anal rays length: Transformation 19.3-21.7 mm ca. 8 8 Pelvic Fin development sequence: D C" PI' P , C A, & 13 11-15 Pectoral 2 2 dorsal Pigmentation: ca. 3.0 mm, fleXion-By & ventral apposing Pre Caudal: of gut terminal 10+9 dorsal blotches posteriad on tail, blotch on surface 10+9 Principal ventral gut; to anterior & one blotch above foregut, section, embedded Procurrent: blotch postanal foregut above blotch in & most lacking embedded 6 5-7 Upper on large mm; by ca. 7.0 mm, peppering >5.0 PI blade & larvae 5-7 6 Lower voluminous median finfold. Flexion-By ca. 9.0 melanophores on Gill rakers: rows & foregut of surface ventral on of 2 pairs of beginning mm, 6 4-7 Upper nape; on midbrain & forebrain; transverse pair at (single) some at 11 10-12 Lower cerebeIJum. 8 8-9 Branchiostegals features: Diagnostic Morphology similar to larvae of D rara; Loweina choroid rara; L. in as posteriad far as not origin on present tissue HISTORY LIFE peppering on PI rays (absent rara); inL. (absent eye of ventral surface paired above brain (transverse melanophores median rara); L. in & in N; Range: Northeast Pacific; south in California Current to ca. 28° bar counts for total vertebrae (39-42 vs. 37-39) & in L. rara); higher Gulf Sea Bering & Alaska of photophores Br2 rara; L. with compared 13-17) vs. (17-20 rays A at ca. form mm. 10.0 Habitat: rnesopelagic; to night at surface at Epi- %)* in MORPHOMETRICS & (range mean CaiCOFI area, in Spawning season: Larvae occur throughout the year abundance in highest February-March with PrF F Y-S Tr PoF Juv eggs larvae planktonic Oviparous; & ELH pattern: 49-60 60-65 65-72 Sn-AlBL 54-54 54 63 67 54 57 LITERATURE 8-16 18-22 BDIBL 21-29 22-23 26 13 22 20 20 Ahlstrom 1965 1939 Bolin 23-29 18-25 26-31 HUBL 30-30 Matarese et aI. 1989 26 21 28 30 29 Ahlstrom & 1970, 1974 Moser 48-61 HWIHL 53-69 39-40 44-64 et Moser aI. 1984a 54 59 51 40 43 Pertseva-Ostrournova 1964, 1974 SnLIHL 19-30x x 22-23 17-21x 22-23 EDlHLt based data (1970). Moser Ahlstrom on in Calculations & * 21-30 26-32 25-36 t second. vertical frrst, given is axis horizontal oval; axis Eye 22x29 23x31 19x25 24 22 3-9 7-9 0-4 0-0 CTLIHL 2 7 8 o o 474

487 Blue lanternfish crenularis Tarletonbeania 3.7mm 18.9 mm 21.7 mm 21.6 mm Figure Myctophidae 44. Preflexion larvae, 3.7 mm, 7.5 mm; flexion larva, 10.5 mm; postflexion larva, 18.9 mm; transformation specimen, 1970). 21.7 mm; juvenile, 21.6 mm (Moser and Ahlstrom 475

488 GADIFORMES A. D. AMBROSE classification a topic is Gadiform of intense presently (Monod 1968). Additional apomorphic features are investigation; the absence given in Markle (1989). system- among consensus of atists (1994) Nelson (1989). Cohen in documented is 482 species in and the families twelve included about of gadiform fishes characteristics history life Early commer- order. Gadiformes contains many important Markle been and Fahay by summarized have (1984). cial quarter over one fishes that comprise of the to spawning oviparous, are fishes Gadiform spherical fish 1994). They are catch (Nelson marine world's with usually eggs, oval coils gut The globule. an oil marine benthopelagic fishes predominantly cool of early the in ontogeny and this, combined with rounded than waters. More half in live of all species gadiform to an head, overall tadpole-like appearance contributes the deep sea beyond commercial fishing many depth; first-feeding of the and larvae. In yolk-sac gadiform are small, apparently rare, or widely dispersed are anus the larvae rather finfold the through laterally exits al. et (Cohen species confined is 1990). One to medially, than larvae. teleost other in is usual as Some popula- freshwater some has another and freshwater usual with forms a caudal fin also differ from the (Nelson 1994). tions range from moderate- Gadiforms pattern some in developing secondary caudal rays elongate very to ly strongly compressed to slightly and before some primary typically fins pelvic The rays. a vertebral high a relatively with have Most count. however, are first fins form; the they form second in to short preanal length (ca. <50% BL); long-based dorsal and last merlucciids gadines (cods and haddock). the in fins, when and anal fins, usually multiple; pelvic features The of body shape, and morphology, anus present, inserted below or front in and pectorals the of with specific fin in development pelvic combination with to up 11 rays; no true spines in the fins (spine- initial for useful most be to appear characters familial identification. dorsal fin); first the in anteriorly occur may rays like the entire forming the margin premaxilla the upper of and the branchiostegal rays numbering 6-8. jaw; represented are Gadiformes by 16 genera 6 in barbels Mental are found in most gadids, macourids, families Current vicinity. The in California the Gadi- and morids. Despite its absence of in over all a third are not included dae Gadus guide in this because few gadiforms, distinctive caudal skeleton (isocercal the macrocephalus larvae have proximus Microgadus and tail) has received much attention. presence The "X" of been in collected and surveys CalCOFI life their early and "Y" elements (usually bones) has interpreted been well known (Dunn stages and Matarese history are as for Gadiformes (Markle 1989). The synapomorphic have Melononid 1989). al. et Matarese 1984; larvae neural "X" and "Y" bones are located anterior to the been in collected not tows CalCOFI are also and parahypural of the first pre-ural vertebra and the spine (see guide 1989). al. et Matarese this excluded from Families included: Bregmacerotidae Macrouridae Moridae Merlucciidae 476

489 GADIFORMES AMBROSE A. D. a topic presently is Gadiform of classification intense are features apomorphic Additional 1968). (Monod absence the investigation; system- among of consensus in Markle (1989). given is in documented Cohen (1989). Nelson (1994) atists about and in families twelve included 482 species the fishes gadiform of Early life history characteristics commer- important many contains Gadiformes order. have (1984). Markle and by summarized been Fahay that one quarter fishes cial comprise over of the are Gadiform oviparous, spawning spherical to fishes are 1994). (Nelson catch fish marine world's They gut coils oval eggs, usually with an oil globule. The of cool fishes benthopelagic marine predominantly early with in ontogeny and this, rounded the combined More waters. than all half of gadiform species live in to overall tadpole-like head, an appearance contributes sea many deep the depth; fishing beyond commercial of In yolk-sac and first-feeding gadiform the larvae. rare, small, apparently dispersed widely are or are the anus exits laterally the through rather larvae finfold (Cohen et al. One species is confined to 1990). than medially, as is usual in other teleost larvae. Some has some freshwater popula- and freshwater another fin also differ from the usual a caudal with forms Gadiforms from range 1994). (Nelson tions moderate- pattern in developing some secondary caudal rays before some primary rays. The pelvic fins typically compressed to ly very elongate and slightly to strongly they are to form; however, fins form second in the first a have Most count. vertebral with a relatively high (ca. length preanal short BL); long-based <50% dorsal last haddock). gadines (cods and the in and merlucciids when fms, and anal fins, usually multiple; pelvic The features shape, anus morphology, and body of present, inserted below or in front of pectorals and the in with specific pelvic development combination fin with to up rays; no true spines in the fins 11 (spine- initial for useful most be to appear characters familial dorsal identification. the in occur may rays like anteriorly fm); first the premaxilla forming the entire margin of the upper by represented Gadiformes are 16 genera 6 in rays the 6-8. numbering branchiostegal and jaw; Gadi- in families vicinity. the Current California The Mental are found in most gadids, macourids, barbels and morids. Despite its in over absence a third all of not few Gadus guide because dae are this included in caudal skeleton gadiforms, the distinctive (isocercal and macrocephalus have larvae proximus Microgadus presence The attention. much received tail) has "X" of been life their and surveys CalCOFI in collected early and "Y" elements (usually bones) has been interpreted are well known stages (Dunn and Matarese history as Gadiformes for (Markle 1989). The synapomorphic have Melononid 1989). al. et Matarese 1984; larvae to the neural "X" and "Y" bones are located anterior been are also and tows CalCOFI in collected not parahypural of the first pre-ural vertebra and the spine (see guide 1989). al. et Matarese this excluded from Families included: Bregmacerotidae Macrouridae Moridae Merlucciidae 476

490 BREGMACEROTIDAE: Codlets G. MOSER STEVENS! G. E. AND H. in Bregmacerotidae contains approximately species 12 single and globule, oil on, a yolk, homogeneous the 1984a; Bregmaceros genus m:onotypic (Houde 1984a). Judging narrow space perivitelline (Houde Nelson 1994). Larvae of two species occur in the size at from the codlet mm), 1.5 (ca. hatching small south pattern survey CalCOFI Eugenia, of Punta Baja may eggs than reported, possibly <0.7 mm. smaller be bathymaster, is a Sur. these, One B. of California have many to similar are Larvae that gadiforms other distribution extends into the Gulf neritic whose species head, coiled large jaws, a massive gut, and a robust Gulf the of California and southward Panama. of to of early preflex- strongly patterns Pigment tail. tapered species, other the of in sp. Bregmaceros called Larvae «3.0 species mm) are diagnostic for and ion larvae guide, this represent and pattern pigment a unique have general groups: three 1) form those with scant or that an apparently species undescribed to endemic is a ventral pigment, with series postanal moderate head head or lacking, 2) present pigment those heavy with Two other species, B. eastern the Pacific. tropical large and ventral postanal blotches, and 3) dorsal and waters tropical inhabit macclellandi, B. and atlanticus a bar those with moderate at the and pigment head south CalCOFI the of Houde 1984a). (Houde area midpoint tail. For those which develop the of species of Bregmaceros (1984a) summarized characteristics B. size, small very a at ray occipital the e.g., and larvae discussed the taxonomic confusion within diagnos- macclellandi, occurrence of the the ray can be the family. early stage larvae, especially when combined for tic is transitional with Pigmentation patterns. pigment small are Codlets cm), in slender, planktivorous, «12 fishes pelagic Some to difficult are species and larvae mm 3.0-6.0 identify of tropical and subtropical seas. at (Houde 1984a). Larger larvae may be this stage enter may and regions coastal to restricted are species patterns, and meristics, pigment specific by identified distributions. whereas others oceanic have estuaries, development size at the The unusual their by recognized easily are Codlets fins. the occipital ray. of first fin dorsal consists an elongate occipital ray. of The anal dorsal fin and the second bathymaster is B. fin are long-based of The following description of specimens (5 39 based observations on detailed lobed. and The caudal fin strongly is developed well 1.8-2.2 mm; 10 preflexion, 2.3--4.2 mm; 6 yolk-sac, dorsal the from separate fins Pelvic fins. and and anal flexion, 4.5-5.1 mm; 5 mm; 10 postflexion, 5.0-12.6 are and have jugular rays; the outer three 5-7 are rays juvenile/adults, 3 mm; 14.6-24.4 transformation, separate, the past well extending elongate, and thick, The line the anal fin. of lateral extends along the origin and mm), sp. Bregmaceros literature. the on 32.0--41.0 dorsal margin detailed observations of 32 speci- is of the body. Scales are relatively large. from described are 12 (4 yolk-sac, 1.9-2.2 report- mm; preflexion, 2.3--4.6 epipelagic; are species Most mens some however, vertical mm; diurnal undergo and mesopelagic be to ed 5 flexion, mm; 6.2-14.0 postflexion, 11 4.3-6.0 migrations. data were obtained from several litera- commercially is species One mm). Meristic harvested Cavinato and sources ture (D'Ancona (Cohen et Belyanina 1965; al. 1990; Nelson 1994). Markle 1984) and 1974; Fahay and Houde 1984a; from at a relatively small mature become Bregmacerotids counts during study. this made original Ecological size (typically information primarily obtained Houde are eggs planktonic Their mm). <30 was from (l984a). reported to be small «1.0 mm), with chori- a smooth La National Laboratory, Jolla Marine at Formerly, ! Service, Fisheries California. La Jolla, Southwest Fisheries Science Center, 477

491 BREGMACEROTIDAE bathymaster Bregmaceros MERISTICS EARLY HISTORY DESCRIPTION LIFE Mode Range EGGS diam.: Shell Vertebrae: Yolk: 49 48-51 Total No.ofOG: Diam.ofOG: 14 13-15 Shell surface: Precaudal 34-36 35 Pigment: Caudal Diagnostic features: Fins: 0 0 Dorsal spines 49 LARVAE 44-51 Dorsal rays 0 0 Hatching mm <1.8 length: Anal spines 48 45-52 4.3 length: Flexion mm 5.0-5.1 through mm Anal rays 6 6 ca. 14-24; Transformation length: gradual Pelvic 18 17-19 Fin development sequence: , PI P A, D, C, Pectoral 2 Caudal: shield Preflexion-At <2.0 mm, above developing gas Pigmentation: 32-36 Total of midline 5-8 I at in jaw, angle postanal series, a ventral bladder, Procurrent: of gut; of gut, & a blotch above preanal arch surface on series ventral Upper end 3.0 mm, most in reduced series postanal stage, of by larvae; by Lower postanal lacking. series I embedded otic Flexion-I nape; at in rakers: Gill ventral gut series lacking; by end of stage, of snout; region; pair at tip Upper & an embedded blotch above larvae some have margin I at hypural Lower colunm. posterior spinal anterior Pos(flexion-Usually above a pair 7 7 Branchiostegals midbrain cerebellum; added column; spinal above & above blotches & margin, hypural at nape, brain, above augmentation mm, by 12.0 stomach; column; juvenile of beginning pigment HISTORY LIFE around shield spinal ventrally & on tail. dorsally series compressed more stage, preflexion In head Diagnostic features: & Range: Tropical-subtropical eastern Pacific postanal ventral series longer than snout pigment Bregmaceros sp.; in sparse pattern then early, ray lost occipital forms later but distinctive; neritic Coastal; Habitat: ca. (during early postfiexion stage, at mm) 5.0 in Bregmaceros than flexion 4.0 sp. (during mm). stage, at ca. Spawning season: Larvae captured throughout year MORPHOMETRICS (range %) in mean & ELH larvae Oviparous; pattern: planktonic eggs & PrF Y-S LITERATURE F PoF Tr Juv Sn-AlBL 42-46 43-48 41-47 36-46 32-38 D'Ancona 1965 & Cavinato 36-37 45 45 44 39 37 36 1984a 1981, Houde BDIBL 25-29 24-29 25-31 15-16 14-36 15-17 28 27 ORIGINAL (Illustrator) ILLUSTRATIONS 27 21 15 16 HLlBL 20-28 20-27 23-27 17-26 15-19 16-18 4.2 larva, Flexion (B. mm Sumida MacCall) 23 24 25 22 17 17 HWIHL 48-114 45-72 53-66 48-69 62-77 64-73 76 60 65 57 69 71 23-36 SnLIHL 20-32 25-29 22-34 17-28 16-21 31 27 25 25 22 19 EDIHL 38-52 28-38 40-54 25-36 24-28 29-30 43 48 34 29 26 29 5-11 6-14 4-10 6-10 PILlBL 10-14 13-14 10 6 8 12 7 13 P L1BL 0-0 0-4 3-7 8-34 37-47 51-54 2 0 0.4 5 20 43 52 ORIJBL 0--0 0--0 0--0 2-10 11-26 21-23 0 0 0 6 22 17 478

492 Codiet Bregmaceros bathymaster ... " ", ,;", 2.1 mm mm 4.2 mm 8.3 9.5 mm 16 mm Figure Bregmacerotidae 1. Preflexion larva, 2.1 mm (Houde 1984a); flexion larva, 4.2 mm (CalCOFI 5612, station 151 G.130); postflexion (Houde 1984a); (D' Ancona and Cavinato 1965),9.5 mm mm transformation specimen, 16 mm (D'Ancona and Cavinato 1965). larvae, 8.3 479

493 Bregmaceros sp. BREGMACEROTIDAE DESCRIPTION EARLY LIFE mSTORY MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: No.ofOG: Diam.ofOG: 53 52-54 Total Shell surface: 15 13-15 Precaudal Pigment: 38 37-41 Caudal Diagnostic features: Fins: 0 0 Dorsal spines LARVAE 49-53 Dorsal rays <1.9 mm Hatching length: 0 0 Anal spines length: Flexion 4.3-4.6 6.0 ca. through mm mm 53-54 50-60 Anal rays at ca. Transformation length: begins 14 mm, gradual 6 6 Pelvic D, PI P C, A, development Fin sequence: , 2 18-20 Pectoral ventral Pigmentation: Preflexion--At <2.0 at blotch mm, postanal Caudal: above 25-29, many scattered & & ventral on anterior myomere 36 34-36 Total 1 to mass, a blotch anterior region of brain, of covering gut surfaces Procurrent: on lateral surface several & base, at jaw angle; at >2.0 PI of mm, 1 Upper region, another on blotches 2 median dorsal caudal blotch ventral in Lower midline snout, on region, otic in embedded a patch tail, of scattered rakers: Gill entire over orbit, of tip, region, posterior margin jaw lower at gular Upper Flexion--Posterior & P rays. ventral blotch brain, opercle, nape, on 2 Lower begin to form 6.0 ca. Pos{/lexion--By mm, patches expands dorsally. 7 7 Branchiostegals on D & A rays Crays of future lobed portions of fms; by 8.0 & on begin covered fully mm, longitudinal series tail; to form on trunk & mSTORY LIFE ca. by 14.0 mm. in than robust & gut B. wider & more mass Head features: Diagnostic Pacific Northeastern Range: tropical-subtropical ca. stage, at flexion earlier (during forms ray occipital bathymaster; stage mm) in 4.0 bathymaster (during postflexion than at ca. B. offshore to Coastal Habitat: 5.0 ventral posterior with blotches, tail ventral 2 & dorsal 2 mm); blotch enlarging. winter & Spawning primarily captured Larvae season: during fall MORPHOMETRICS (range mean in Ufo) & ELH pattern: Oviparous; planktonic eggs & larvae PrF F Y-S Juv Tr PoF LITERATURE 33-49 30-39 33-44 42-47 Sn-AlBL 40 34 42 45 16-31 27-33 19-26 BDIBL 27-36 22 30 21 30 ORIGINAL ILLUSTRATIONS (Illustrator) 16-30 23-28 23-33 20-24 HLlBL 27 25 22 22 Yolk-sac larva, 2.2 MacCall) Sumida (B. mm Preflexion larva, (B. MacCall) 2.2 Sumida mm 3.5 mm, 64-80 66-77 38-62 54-77 HWIHL mm 4.3 MacCall) Sumida (B. larva, Flexion 70 48 65 71 mm Sumida 7.2 (B. MacCall) Postflexion larva, 18-29 10-20 15-32 25-34 SnLIHL MacCall) mm 14.0 Sumida (B. Transformation specimen, 25 28 24 13 38-42 33-48 30-43 23-39 EDIHL 41 39 35 29 7-9 6-8 6-12 0-0 PILlBL 8 8 7 0 17-41 5-17 17-24 P 0-0 L1BL 2 11 19 32 0 1-4 1-7 6-17 ORUBL 0-0 10 4 3 0 480

494 Codlet Bregmaceros sp. 2.2mm 2.2 mm 3.5 mm 4.3 mm 7.2 mm 14.0 mm Figure Bregmacerotidae 2. Yolk-sac larva, 2.2 mm (CaICOFI 7202, station 157.55); preflexion larvae, 2.2 mm, 3.5 mm (CaICOFI 7210, larva, mm 7210, station 157.10); postflexion (CaICOFI 7.2 (CaICOFI 7210, station 157.20); flexion 157G.55); larva, 4.3 mm station transformation specimen, 14.0 mm (CaICOFI 5612, station 173G.20). 481

495

496 MACROURIDAE: Grenadiers D. A. AMBROSE occur in the CalCOFI a least is berryi Mesobius and migrations At 13 macrourid species exclusively Macrouridae 1), but larvae pectoralis and dweHer. midwater A. area (Table acrolepis are only 5 C. of acrolepis, (Albatrossia pectoralis, Coryphaenoides C. (Cohen important commercially et al. species 1990). allied closely most are and berryi, Mesobius lepto/epis, Macrourids Merluc- spp.) have the with Nezumia the within the Gadifonnes and dominate ciidae bentho- identified been samples. CalCOFI in ichthyoplankton fauna fish pelagic larvae are collected in the scaphopsis Caelorinchus numbers of in sea deep the of individuals Cohen 1973). and (Marshall species and anguli- Coryphaenoides Gulf of of Larvae California. ceps, Malacocephalus and armatus, C. C. yaquinae, reproduction, specu- is known about it is but Little are laevis unknown, those while are C. of Jilifer eggs broadcast near the bottom and that lated are in identified been not have but described CalCOFI as they drift toward the surface (Marshall and develop N. of collections. Larvae liolepis, Nezumia convergens, Iwamoto 1973; Merrett 1978, 1989). the few For and to stelgidolepis N. only identified are genus. ca. described, species are eggs 1.0-4.0 mm in diameter with mm), a single (most <2 oil and globule a chorion grenadiers are medium to of (ca. these large Adults hexagonal with sculpturing (Fahay and Markle 1984). 150 30-> TL), compressed moderately and elongate cm Few the captured are macrourids larval despite abun- with trunk, short head, large a rather long fishes, and dance it has slope most continental in adults of areas; tail tapering to a fine point with no cauda! fin (Cohen to that larval development and descent been suggested et of aL 1990). The high, short first dorsal fin consists bottom et al. the is rapid (Cohen 1990). two rays followed by 7-13 segmented rays. spinous long, low second dorsal fin is separated from the The trunk, short head, a large have macrourids Larval first by a gap longer than half the base of the first tapering long fin pelvic early tail, markedly formation, rays dorsal fin and its rays are shorter than the opposite fin pectoral that fin dorsal a second and base, stalked the of the of end the to continuous is fin, anal which the than shorter and is lower fin. anal Larval pigment but appear may adults, in thoracic are fins Pelvic tail. region. heavy on the is abdomen and frontal head stages. the pointed, is snout The larval in jugular the from to juvenile larval stage, During transformation to mouth inferior, and the scales cycloid subterminal mouth pectoral the fin shortened, is greatly length base 7 and 6 or number rays Branchiostegal with spinules. orientation becomes less snout and oblique, be- the is barbel mental A 12-16. are vertebrae abdominal more comes pronounced. all present on except Mesobius berryi. A ventral abdominal the between wen organ is developed light distinguish may be used to the characters Numerous pelvic in Caelorinchus scaphopsis and bases fin larvae region, and of species in the CalCOFI genera to Malacocephalus laevis. It is anterior in rectum the including some adult diagnostic characters as such in and berryi Mesobius aU species of Nezumia, three numbers the of morphology and rays branchiostegal of absent in Albatrossia and Coryphaenoides. but is The retia may Genera with those into grouped mirabilia. be are mirabilia retia and swimbladder developed in wen and rays 7 branchiostegal (Malacocephalus, Mesobius, are in Meso- and Albatrossia reduced but most genera and those Nezumia) 6 with rays branchiostegal bius. The anus is fin anal in immediately before the Coryphaenoides). (Albatrossia, Caelorinchus, and Coryphaenoides, Albatrossia and slightly to anterior Mesobius berryi larvae have dorsal and ventral (more the in Caelorinchus and midway Mesobius, fin anal tail. Nezumia along patches pigment numerous) the fin the and fin anal the between pelvic N. converg- in ventral have 2 rows margin tail and pigment medio- ens and liolepts, the in fin N. to and closer pelvic lateraHy the along with tail. 6 bran- those Among Malacocephalus stelgidolepis. N. Adults of most and snout- the have Albatrossia rays, chiostegal shortest macrourids are bathybenthal, ranging from the outer (ca. length anus ofBL) yolk-sac the 20% from to stage to shelf continental plain. Coryphaenoides abyssal the formation, initial and anal fin the fewest gas glands and acrolepis vertical Ji/ifer are known to C. make (2). Caelorinchus have a low number fin of pelvic 483

497 l rays a high number of pyloric caeca (15-26),4 gas (7), the from obtained were data ecological and Meristic the anal size fin glands, formation at smallest literature 1966; Fitch (Lavenberg Novikov and 1970; pelvic (6.5 mm), elongate rays, fin and pigment absent Hubbs Miller and Lea 1972; Iwamoto and Stein 1974; of the tail. Coryphaenoides half posterior the from and Iwamoto 1977; Iwamoto 1978, 1979; Stein 1980a; basipterygia. et 1982; Pearcy and Stein al. 1989; Cohen et have acrolepis C. Matarese larvae have robust base pigment laterally and on the pectoral fin on the 1990). al. pectoralis, series of Albatrossia Larval 12-14 pyloric caeca, and of the tail, anterior two-thirds Caelorinchus scaphopsis, and early larval stages of (1980b) described larvae filifer C. 4 gas Stein glands. Coryphaenoides leptolepis, Mesobius berryi, and oval having as of pigment extending from a patch an Nezumia the first time. Cohen et al. are described for to about fin the dorsal first 12th position anterior to the able of were (1990) differentiate adults between not to fin dorsal second also C. has ray. the highest filifer consider C. leptolepis and C. C. liocephalus, and we rays of a single be first dorsal fin species. (13-16). C. to leptolepis number total evenly larvae leptolepis pigment heavy spaced have Since notochord undergo not do macrourids larval trunk spots over the not extending posterior to the lOth larval separate to used was anal formation fin flexion, glands, 6 gas stage, juvenile the by ray fin anal a and developmental stages. (PrAF) "Preanal fin formation" dorsal The number relatively wide space between fins. fin refers to the interval prior to initiation of anal be and useful gas glands may rays fin pelvic of in to and refers (AF) formation" fin "anal formation other of larvae stage later differentiating three or fins. developing anal formed fully larvae with region: from species Coryphaenoides C. this denomina- as Preanal length and head length are used (usually 8), gas glands 4; C. anguliceps-P2 7-9 tors because macrourid for the morphometrics section and glands 10-12 (usually 11), gas armatus-P 5--6; 2 tails. larvae captured in nets often have damaged glands 6. yaquinae-P (usually 8-11 C. 10), gas 2 (1990) spelling of Caelorinchus is used, Eschmeyer's on The incorrect but used commonly the to opposed as following descriptions are based 6-19 Coelorinchus. specimens 2). Macrouridae (Table taxon each of 484

498 Table Macrouridae Meristic characters for the macrourid species that occur in the California Current region. 1. gill Total rakers Pyloric st arch) (inner I BrR glands 10 Species Gas P caeca PI 2 5-14 Albatrossia pectoralis 12-16 9-11 6 16-21 2 6-8 9-12 15-26 9-11 4 scaphopsis 17-20 Caelorinchus 6 7 5-13 12-14 10-13 19-22 Coryphaenoides 6 4 acrolepis 8-9 7-9 4 9-12 9-14 6 19-24 7-9 C. anguliceps 7-14 10-13 5-6 10-12 18-22 10-12 armatus 6 C. 8-14 13-16 8-12 4 18-23 6 9-11 Jilifer C. 18-22 6 9-16 II 6 10-12 leptolepis 9-10 C. ll-12 8-12 8-11 10 6 16-22 6 yaquinae C. 11-14 ll-14 8-9 50-100+ 2 7 16-22 laevis Malacocephalus 12-14 7 10-13 2 10-12 12-17 7-9 berryi Mesobius 10-12 21-32 7-9 18-22 9-11 2 7 Nezumia convergens 25-37 9-12 10-13 7 2 20-25 10-12 N. liolepis 8-12 2 10-13 20-26 8-11 24-58 7 stelgidolepis N. Macrouridae 2. Number of specimens (above) and total length Table (in mm, below) used in preparation of the range macrourid descriptions. The symbol species of indicates length n>n the end a minimum the tail was damaged. because Juvenile Yolk-sac PrAF AF Transformation Species 14 0 0 4 Albatrossia pectoralis >22.1 7.5-15.3 5.8-7.1 I 2 0 5 0 scaphopsis Caelorinchus 6.5->20 >25->28 3.7 0 8 8 0 0 acrolepis Coryphaenoides 10.5-23.5 7.6-9.9 2 4 0 0 0 leptolepis C. 5.8-6.3 7.4->42 2 6 I 3 3 berryi Mesobius 14.2->23.4 5.1-5.4 73 6.0-1l.2 148-213 .. 12 0 0 3 Nezumia sp(P). 5.8->13.2 >15.7-49.6 >94 485

499 MACROURIDAE Albatrossia pectoralis LIFE HISTORY DESCRIPTION EARLY MERISTICS Range EGGS Mode diam.: Yolk: Shell Vertebrae: 14 No.ofOG: Diam.ofOG: Precaudal 13-15 surface: Shell Fins: II First II Pigment: dorsal spines Diagnostic features: First rays 7-9 dorsal 9-II Total first dorsal LARVAE 7 Pelvic 6-8 Hatching length: 5 mm ca. 19 16-21 Pectoral Transformation length: 12-16 Pyloric caeca , & 2 2 20 & A 10, PI P sequence: development Fin Gas glands 2 posterior Gill rakers (inner first arch): Yolk-sac--Dn lower on eye; to snout, brain; Pigmentation: 2 above jaw, nape; over c1eithrum; Upper gut; 0-2 & scattered on lateral surface of 100II end 5-12 PI base; blotch ventrally at midtail; scattered near of tail. on Lower 5-14 on Preanalfinformation-Expanding Total operculum & lateral surface of until gut; from end of anteriorly tail; spreads patch spreading midtail 6 6 Branchiostegals 15 mm BL, ca. along BL; mm 12.5 by filled at both gaps by ca. sides expanding of lateral surface & tail & ventral tail margins on dorsal LIFE HISTORY anteriorly. Analfinformation-Evenly spaced along & ventral dorsal tail margins. from Japan to the Okhotsk & to Bering Seas, east Range: Pacific North 10 gas 6-8; P 6; total 9-11; features: rays Branchiostegal Diagnostic 2 of south Gulf Alaska, to northern Baja California the glands yolk-sac larvae; AF through yolk-sac for BL 20% ca. 2; Sn-A ventral & along pigment at midtail; blotch the with larvae PrAF early young 140 Habitat: Adults near bottom to 1200 m depth; bathypelagic & dorsal ventral through PrAF late in AF of margins larvae. & tail the settle to bottom when about 50-60 cm long & mean in %) MORPHOMETRICS (range season: Spawning with year Throughout peaks in fall & winter the pattern: Oviparous; Y-S planktonic larvae PrAF ELH Tr AF Juv 20-22 17-23 Sn-AlBL* LITERATURE 21 20 48-65 BO/Sn-An 61-73 58 58 67 51-66 50-68 HUSn-An ILLUSTRATIONS ORIGINAL (Illustrator) 61 57 69 53-100 HWIHL 71-83 Vona) larva, 5.8 mm Yolk-sac T. (M. 69 77 52 PrAF 15.3 mm (M. Vona) mm, larvae, 7.5 T. 21-33 20-32 SnLIHL 25 26 25 38-49 EOIHL 34-42 of 9 in tails of broken determined be 14 not could length Body * because 42 37 34 & specimen. PrAF specimens in the only AF 6-18 13-24 specimens in 8 of 14 PrAF rays & not measured. P were damaged t PILIHL 2 14 20 18 44-133 3-21 P L1HLt 2 >42 14 80 26-36 38-73 PIBLIHL 31 58 49 486

500 Albatrossia pectoralis Giant grenadier 5.8 mm 7.5 mm 15.3 mm Figure Macrouridae 1. Yolk-sac larva, 5.8 mm (CaICOFI 6310, station 87.35); PrAF larvae, 7.5 mm (CaICOFI 5512, station 87.40), 15.3 mm (CaICOFI 8404, station 60.52.5). 487

501 MACROURIDAE Caelorinchus scaphopsis MERISTICS DESCRIPTION HISTORY LIFE EARLY Range Mode EGGS Vertebrae: Shell diam.: Yolk: No.ofOG: Precaudal Diam.ofOG: Shell surface: Fins: II Pigment: spines II First dorsal 7-9 First rays dorsal 8 Diagnostic features: 9-11 10 Total first dorsal LARVAE 7 Pelvic 7 length: 17-20 Hatching Pectoral 15-26 Transformation length: Pyloric caeca 4 4 , Gas PI A, 2D, ID, glands P sequence: development Fin 2 Gill arch): 1st (inner rakers posterior head; Pigmentation: Preanalfinformation-On top eye; to of 1-2 Upper on preoperculum & operculum; on body anterior to gut; side ventral 8-10 Lower half surface lateral & ventral on gut; of anterior of of tail. Anal fin 9-12 Total formation-Anterior 18 mm, on tail to about A ray 25-32; by ca. on 6 6 P PI of base . & Branchiostegals 2 features: Branchiostegal rays 6; 7; lD rays 9-11; total Diagnostic P 2 in PrAF ofSn-AL 70% & HISTORY LIFE gas BD ca. glands 15-26; caeca pyloric 4; larvae; AF a small A forms at mm); body length (by 6.5 anus not from lower absent pigmentation 2 rays; ID & P elongate A; to adjacent California Range: & Southern northern Gulf of California tail. & snout through jaw formation, present anteriorly on A from Benthopelagic Habitat: depth m 296 to 183 MORPHOMETRICS (range & mean in %) Gulf February from the in Spawning collected season: Early-stage larvae Y-S of California PrAF AF Tr Juv Sn-AlBL* ELH pattern: Oviparous; planktonic larvae 29-37 36 34 LITERATURE BD/Sn-An 66--71 53-64 69 69 58 HUSn-An 57-80 70--76 57 76 73 HWIHL 48-73 48-51 ILLUSTRATIONS ORIGINAL (Illustrator) 84 59 50 SnLIHL 23-27 26--28 mm Vona) T. 3.7 (M. larva, PrAF 18 24 27 AF >18.0 T. mm Vona) larvae, 6.5 mm, (M. EDIHL 29-37 26--30 45 32 28 2 in broken Tails * 15-23 specimens; transformation both & specimens AF 5 of >26-->36 PILlHLt 29 17 included not morphometrics for in damaged specimens. both transformation measured. not & specimens PI damaged on t 54-171 >50-->84 P2L1HL~ P 5 damaged on 1 of t AF specimens & on both transformation 45 128 2 measured. not specimens; PIBLIHL 30--55 10--13 53 39 12 488

502 Shoulderspot grenadier scaphopsis Caelorinchus 3.7mm ~ mm 6.5 ... '. -"" ... -.;;:; . . ', 18.0 mm ... _ ""=.=-c==--_ -::.: ... - .. ... Figure Macrouridae 2. PrAF larva, 3.7 mm (CaICOFI 5702, station 121G.40); AF larvae, 6.5 mm (CaiCOFI 5702, station 118.7G.32), 18.0 mm, tail broken (CaICOFI 5702, station 12lG.40). 489

503 MACROURIDAE acrolepis Coryphaenoides MERISTICS DESCRIPTION mSTORY LIFE EARLY Range Mode EGGS Vertebrae: diam.: Shell Yolk: 12-16 No.ofOG: Precaudal Diam.ofOG: Fins: surface: Shell II First dorsal spines II Pigment: rays 8-11 10 dorsal First 9 or features: Diagnostic Total first dorsal 10-13 8-9 Pelvic 8 LARVAE 19-22 Pectoral Hatching length: 12-14 Pyloric caeca HL 9.4-9.8 Transformation length: mm 4 4 development Fin P glands Gas 10, A, 20, PI sequence: , 2 Lower formation-Over top of Pigmentation: & fin head; on snout Preanal rakers first arch): Gill (inner peritoneum; anal & dorsal of bases along over internally jaw; lower 2 0-2 Upper fin Anal tail. of absent posterior finfolds; one-third laterally on the II or 10 5-11 Lower & tail; transformation-Increasing laterally on body formation through Total 5-13 posterior one-third lateral pigmented. remains tail of lightly 6 6 Branchiostegals 8-9; P 1D 10-13; features: Diagnostic Branchiostegal rays 6; total 2 BL ca. 30% 12-14; for PrAF gas glands 4; Sn-A pyloric caeca LIFEmSTORY light on lateral through pigmentation stages; basipterygia robust; AF 20% of body. posterior the Japan & Seas; Bering Range: Okhotsk to North northern from Pacific mean & %) in MORPHOMETRICS (range south Mexico North American coast to Isla de Guadalupe, along Benthopelagic Habitat: m depth, occasionally bathypelagic 600-2500 Y-S PrAF AF Tr Juv occur primarily in CalCOFI (PrAF) larvae Spawning season: Early-stage Sn-AlBL 29-33 31-34 30 32 collections in ichthyoplankton January & February; off Oregon, Stein & Pearcy (1982) reported ripe females captured in April, September & 51-60 BO/Sn-An 53-61 October, spent females in October 56 57 HUSn-An 51-66 55-67 larvae pattern: planktonic Oviparous; ELH 60 60 HWIHL 61-82 48-71 LITERATURE 69 59 18-24 SnLIHL 19-28 1980b Stein 22 24 Stein & Pearcy 1982 EOIHL 37-49 28-43 et Matarese al. 1989 44 37 17-21 15-16 PILIHL* 20 on 3 of 8 PrAF & 5 of 8 AF specimens; not fins damaged Pectoral * 15 measured. 94-146 L1HLt P 2 specimens & all 8 AF specimens; t Pelvic fins damaged on 6 of 8 PrAF 120 measured. not 49-75 PIBLIHL 46-65 57 54 490

504 Pacific grenadier acrolepis Coryphaenoides mm 1.8 HL mm HL 3.8 HL mm 6.7 9.8 mm HL Figure Macrouridae 3. PrAF larva, 1.8 mm HL; AF larvae, 3.8 mm HL, 6.7 mm HL; transformation specimen, 9.8 mm HL (Stein 1980b). 491

505 MACROURIDAE Coryphaenoides fllifer DESCRIPTION EARLY LIFE mSTORY MERISTICS Mode EGGS Range Shell diam.: Yolk: Vertebrae: No.ofOG: Diam.ofOG: Precaudal surface: Shell Fins: Pigment: II II dorsal First spines features: Diagnostic 11-14 12 rays First dorsal 13-16 Total first dorsal LARVAE 9 9-11 Pelvic Hatching length: 18-23 Pectoral Transformation length: 14.1-14.6 mm HL 8-12 Pyloric caeca 4 P 2D, & A & ID & sequence: development Fin PI 4 glands Gas 2 of region frontal Analfinformation-On lower Pigmentation: on head; (inner Rakers first arch): Gill & jaw peritoneum; on dorsolaterally regions; suborbital & gular Upper 1D & ray; 2D 12th about to posteriorly extending surrounding Lower stage, Transformation-8imilar to earlier along lateral scattered line. 8-14 Total posteriorly extending pigmentation except dorsal only 2D 7th ca. to 6 6 Branchiostegals ray. 6; rays 1D features: Diagnostic Branchiostegal 13-16; P 9-11; gas LIFEmSTORY 2 basipterygia; glands 4; prominent Sn-A patch pigment <30% BL; oval 0 extending from anterior region posteriorly of to 0 about the 12th 2D 1D Bering Range: Sea N) (32 California southern to N) (66 to ventro-Iaterally & line. lateral the ray depth Habitat: Bathybenthal, 2065-2904 m %) MORPHOMETRICS (range & mean in ripe & eggs Pearcy (1982) reported with females Stein season: Spawning y-S PrAF AF Juv Tr Oregon off & August June, in January, July, Sn-AlBL larvae Oviparous; pattern: ELH planktonic LITERATURE BD/Sn-An Stein 1980b HLlSn-An Stein & Pearcy 1982 et al. 1989 Matarese HWIHL SnLIHL EDIHL 492

506 Threadfin grenadier filifer Coryphaenoides mm HL 6.4 mm HL 10.5 14.6 mm HL Figure Macrouridae 4. AF larvae, 6.4 mm HL, 10.5 mm HL; transformation specimen, 14.6 mm HL (Stein 1980b). 493

507 MACROURIDAE Coryphaenoides /epto/epis HISTORY DESCRIPTION LIFE MERISTICS EARLY EGGS Range Mode diam.: Shell Vertebrae: Yolk: No.ofOG: 12 12 Diam.ofOG: Precaudal Shell surface: Fins: II II Pigment: First spines dorsal 8-10 First rays dorsal Diagnostic features: first Total dorsal 10--12 9-10 LARVAE Pelvic 19 18-22 length: Hatching Pectoral II mm, >6.2 length: Transformation HL mm II <13.6 caeca Pyloric Fin development sequence: P , A, 6 ID, PI 2D, 6 glands Gas 2 Gill (inner arch): first rakers Preanal fin formation-Above, below, & posterior to Pigmentation: eye; heavy to posteriorly on from dorsally scattered trunk; midbrain Upper I 8-15 Lower from PI base. about fin formation-On Anal myomere 7; absent 9-16 Total extending eye; to posterior & below head, of region frontal increasing 5th Transformation-juvenile- ray. D to ca. over posteriorly trunk 6 6 Branchiostegals just from trunk on spots scattered Evenly I D to anterior to posterior 10th about the A ray. HISTORY LIFE total 6; 9-10; P 10--12; lD Diagnostic features: Branchiostegal rays z through from robust extremely basipterygia AF glands gas 6; (55°-59° California southern of south to N) Range: Alaska Southeast stages; Sn-A ca. transformation AF BL from PrAF through 30--40% of of2D; origin & lD stages; relatively wide space between insertion m depth 2400--2800 Bathybenthal, Habitat: pigmentation heavy over trunk in evenly spaced extending not spots juvenile. to 10th A ray by posterior occur larvae in CalCOFI ichthyoplankton season: Spawning tows PrAF March & February in (range MORPHOMETRICS %) & mean in Oviparous; planktonic ELH pattern: larvae PrAF Y-S AF Tr Juv LITERATURE 31-37 Sn-AlBL* 34 40 1980b Stein 1989 Matarese et al. BD/Sn-An 63-63 55-70 63 64 ORIGINAL (Illustrator) ILLUSTRATIONS HLlSn-An 62-66 49-60 64 56 T. Vona) mm 5.8 (M. PrAF larva, HWIHL 63-75 42-82 69 54 25-31 SnLIHL 25-28 * was Tail included specimens; AF 4 of 3 on damaged not in the 27 28 morphometrics. EDIHL 39-41 16-38 4 AF specimens; not included in the fin was damaged on 3 of t Pectoral 40 23 morphometrics. 14-19 in damaged were fms Pelvic specimens t examined. all PILlHLt 16 20 PzLIHLt 51-67 32-51 PIBLIHL 59 41 494

508 /epto/epis Coryphaenoides Ghostly grenadier mm TL) (5.8 mm 1.2 HL 6.2 HL mm 15.2 mm HL Figure Macrouridae 5. PrAF larva, 5.8 mm TL (1.2 mm HL) (CaICOFI 7203, station 70.65); AF larva, 6.2 mm HL; transformation specimen, 15.2 mm HL (Stein 1980b). 495

509 MACROURIDAE Mesobius berryi EARLY LIFE HISTORY DESCRIPTION MERISTICS Mode EGGS Range diam.: Shell Yolk: Vertebrae: 11 No.ofOG: 11 Diam.ofOG: Precaudal surface: Shell Fins: II Pigment: II First dorsal spines 9 features: Diagnostic 8-10 dorsal rays First 11 10-12 dorsal Total 8 LARVAE 7-9 Pelvic 5 mm NL ca. Hatching length: 13 12-14 Pectoral length: Transformation 12-17 caeca Pyloric , Fin ID, sequence: 2 P A, development 2 PI 2D, Gas glands 2 midbrain on patches Yolk-sac-Small Pigmentation: & hindbrain, & first rakers (inner arch): Gill on patches 2 eye; to posterior patches about base; 8 p] dorsal Upper from row numerous extending posteriorly along nape along tail, more Lower II 10-13 fin Preanal tail margin; ventro-lateral row along tail. ventral Total formation-On of 7 top gut, gradually spreading over on abdomen; 7 Branchiostegals oflower angle posterior & lower branchiostegal region; on snout, jaw, jaw. formation-Uniformly fin Anal HISTORY LIFE & dorsally saddles spaced more Transformation- ventrally along tail; ventral saddles numerous. & body persists. on spotting head juvenile-Bold California Range: Northeastern Pacific from southern California & Baja Diagnostic features: Branchiostegal rays 7-9; 7; 12-14; P PI gas 2 Islands Hawaiian the to Island & Christmas BL <10% to in yolk-sac larvae, glands 2; 25% ca. Sn-A decreasing ca. increasing larvae, yolk-sac in HL 50% in length base PI juveniles; captured Habitat: Bathypelagic; as as trawls midwater in 0-313 shallow pigment patches 70% before transformation; regularly spaced to ca. m; deep most taken in hauls to at least to m 650-1000 m as as 0-2700 of body, & ventral absent near end tail. dorsal along depth %) MORPHOMETRICS (range & mean in Spawning season: Yolk-sac & PrAF larvae collected in CalCOFI & October July, April, January, in tows ichthyoplankton y-s AF PrAF Tr Juv ELH larvae planktonic Oviparous; pattern: 24-26 Sn-A/BL* 18-23 6-9 25 21 16 7 8 LITERATURE BD/Sn-An 47-54 52-75 78-85 100-118 50 63 82 86 111 Iwamoto & 1977 Hubbs 1984 & Fahay Markle HUSn-An 60-72 52-53 77-91 115-138 64 53 84 131 129 (Illustrator) ORIGINAL ILLUSTRATIONS 62-77 66-73 57-63 HWIHL 42-46 60 68 70 53 44 mm Vona) (M. T. 5.4 larva, Yolk-sac 21-28 15-27 SnLIHL 15-17 21-25 larvae, PrAF Vona) 6.7 mm, 11.2 mm (M. T. 24 16 22 22 23 40-44 35-46 36-39 EDIHL 25-35 43 37 41 36 31 Tail on 1 on & specimens 3 AF not of 2 * 3 juveniles; damaged of 20-27 10-18 3-9 in included morphometrics. 25-51 P]LlHLt 24 15 20 6 38 t specimens; not measured. damaged PI 3 of 6 PrAF on was larvae; yolk-sac of larger the P only forming was P t two 2 2 8-35 38-70 0-3 25-62 P L1HLt 2 & 3 AF 5 PrAF of specimens. of 1 damaged 1 on 26 2 43 54 45 35-71 69-80 66-79 PIBLIHL 7-8 74 53 71 7 7 496

510 Mesobius berryi Midwater grenadier 5.4 mm 6.7mm 11.2 mm Figure Macrouridae 6. Yolk-sac larva, 5.4 mm (CaICOFI 7210, station 31.133); PrAF larvae, 6.7 mm (CaICOFI 6304, station 90.100), 23.4 11.2 (CaICOFI 7807, station 90.190); AF larva, mm mm, tail broken (Hubbs and Iwamoto 1977). 497

511 sp(p). MACROURIDAE Nezumia DESCRIPTION LIFE EARLY HISTORY MERISTICS EGGS Mode Range diam.: Shell Yolk: Vertebrae: No.ofOG: Diam.ofOG: 13-14 Precaudal surface: Shell Fins: Pigment: II II First spines dorsal Diagnostic features: 8-11 rays dorsal First 10-13 dorsal Total LARVAE 8-12 Pelvic length: Hatching 18-26 Pectoral Transformation length: 21-58 Pyloric caeca , development sequence: P ID, Fin A, 2D, PI 2 2 Gas 2 glands Pigmentation: Preanal anteriorly head, formation-Over fin & arch): rakers (inner first Gill on on ventral abdomen; dorsally; on branchiostegal region; heavy Upper by mm, 13 ca. & dorsal finfold of posterior one-third of tail; midline Lower series a double increasing anteriorly forming ventral midline & along 7-12 Total tail. laterally along absent ID; under formation-Heavy fin Anal 7 7 Branchiostegals end & of tail. along midline dorsal Diagnostic 7; total 1D 10-13; P Branchiostegal 8-12; rays features: HISTORY LIFE 2 for Sn-A <20% BL glands specimens >6 gas 2; caeca 21-58; pyloric pigment absent on PI base & along dorsal rum through AF stage; liolepis--Central Range: N. converge~ulf of California to Chile; N. of & 13 ventral ca. mm, series along dorsal midline by tail; midline (ca. 17° southern Mexico California (ca. 37° N) to Gulf of California & midline of tail & tail. on laterally series double Island, Columbia British stelgidolepis--Vancouver N. N); southern to S) 18° (ca. Peru in %) (range MORPHOMETRICS & mean convergens-Benthopelagic 600 1865 to Habitat: N. N. in m; Y-S AF PrAF Juv Tr stelgidolepis--Benthopelagic liolepis--Benthopelagic, 768 to 1655 rn; N. in depth rn 277-909 Sn-AlBL* 15-22 15 17 Spawning season: tows larvae occur in CalCOFl ichthyoplankton PrAF & in winter spring primarily BD/Sn-An 64-85 81-88 85 80 63 ELH larvae pattern: Oviparous; planktonic 100-115 43-83 HLlSn-An 109 72 97 LITERATURE 42-58 63-94 HWIHL 78 48 39 Matarese et a1. 1989 SnLIHL 18-21 20-32 20 25 25 ILLUSTRATIONS (Illustrator) ORIGINAL EDIHL 35-56 22-29 41 25 27 PrAF larvae; mm >13.2 rum, 8.8 Vona) (M. T. 11-20 16-34 PIUBLt >15.7 (M. T. Vona) rum larva, AF 15 22 52 >34->48 UBq 47-146 P 2 * 2 in damaged juvenile 12 Tail of 93 >58 PrAF specimens & on only the determined. be not could length body & specimen 46-65 25-40 PIBLIHL & on morphometrics. in include not t specimens PrAF 12 of 1 damaged PI 31 56 7 on all juvenile damaged on 6 of 12 PrAF specimens & P t AF & 2 specimens; minimum ranges are listed for AF specimens. & juvenile 498

512 Nezumia spp. mm 8.8 --~-- -~- ~------ 13.2 mm 15.7 mm Figure Macrouridae 7. PrAF larvae, 3.3 mm (CaICOFI 6401, station 93.30), 13.2 mm, tail broken (CaICOFI 7205, station 30.52); AF larva 15.7 mm, tail broken (CaICOFI 5106, station 150.25). 499

513 MORIDAE: Codlings A. D. AMBROSE morid species occur Five the CalCOFI area (Table in position base fin vertical in change little is there but Moridae adults. papilla from larvae to A pronounced genital (Microlepedium 1) but larvae of only two in develops postflexion (when larvae. A mental barbel rastrelliger) collected Physiculus are verecundum and early or present) develops in the late larval pelagic CalCOFI ichthyoplankton in regularity any with juvenile period. slightly pectoral fm base is only 1he samples. are larvae nematopus Physiculus in collected delayed. is stalked and caudal fin formation In the microlepis Antimora Larval California. of Gulf and divides forms, fin dorsal a single late Physiculus, then johnsonii Halargyreus johnsonii are unknown. H on pigment in development. Larval is heavy the gut only adults the from reported been recently have and brain and prominent on the pelvic rays, but light (Logan Pacific et al. 1993). eastern rays. Transformation the on is gradual and pectoral of in medium to small are codlings length Adult the except changes, striking no with decrease direct for compressed, (ca. 17-75 cm), elongate and moderately fin M pelvic of in rays verecundum. number in to and with the tail tapering a slender caudal peduncle, Numerous used be may to characters the distinguish 1972; with a separate caudal fm (Fitch and Barker and the genera of region. in species M CalCOFI larvae dorsal The 1990). 1983; Eschmeyer et al. al. et Cohen verecundum fewest has the fewest myomeres (44-46), fin consists and a section anterior of a short-based rays and (39-42), fin dorsal second the in a relatively A single section. posterior long-based separate long- to preanal distance (ca. 38-53% BL) compared long all except H johnson anal based fin is present in ii, 24-33% have larvae Physiculus BL). (ca. Physiculus has two. In which fins. pelvic jugular have species All early fins a heavy and pelvic elongate forming, patch is rays fin pelvic of verecundum, M the number gut the on ventrally pigment of where the light organ five from ontogeny during reduced in postflexion two in adults. All fms consist solely to larvae will form. has rastrelliger soft of along pigment of a line P. of this lacks nematopus P. tail; the margin ventral the scales, rays. All otophysic an species cycloid have rays P. P. rastrelliger and has 7 pelvic pigment. spine the first neural connection, fused the skull, and to more P. rastrelliger also has caudal nematopus has 6. in morid only is the johnsonii H bones. hypural five Although nematopus P. (22-26). fin rays than (27-32) the region without a mental barbel. California Current larval are johnsonii H and microlepis A. unknown, Physiculus has a light organ on a dark belly in the they should be readily by identified presumed their A. microlepis area. scaleless in inferior is mouth The of high relatively tadpole-like shape and number others. the all in verecundum but M Adult subterminal other collected primarily in midwater trawls. The are precaudal myomeres (A. microlepis, 24-25 and H conti- are morids outer the from ranging benthopelagic, also the fewest has microlepis A. 22-23). ii, johnson region. the in morid any of dorsal (4-5) rays fin first slope nental shelf to lower and some (A. microlepis) of communities in those are important members descriptions morid following The of life early habitats. of history examinations detailed based are stages on each (Table species of between 14 and 30 specimens and natural history of the early life is known Little the few species described, of eggs are rastrel- P. and verecundum M of morids. For Moridae 2). Larvae 0.52-1.16 oil with globule diameter, in mm a single liger are described for the first time. Meristic data were and (Fahay chorion smooth a 1984; and from Markle obtained from counts made during this study and Barker and Fitch 1985). al. et Fahay 1972; found and (Fitch literature the Kitagawa and (1972) Barker 1984; Markle advanced mm in eggs a 67 specimen ovarian Paulin 1989). source principal M of of The Fitch was information ecological (1972). Barker and morids verecundum. Larval are noted for being stocky body, with head, a rounded elongate anteriorly tapering as listed Laemonema (1990) al. verecunda et. Cohen and relatively volumi- and early forming pelvic fins, Microlepedium verecundum; for the preferred name retain of the (1990) dorsal and anal the fins. and however, we follow Eschmeyer portions nous posterior guide. identification this for name latter in Pelvic fins are greatly reduced in relative size adults, 500

514 Table Moridae Selected meristic characters for the morid species that occur in the California Current region. 1. Fin Vertebrae rays Species GR lD 2D lA Total PrCv CV P 2A C Aror 2 PI microlepis 24-25 4-5 57-58 Antimora 50-55 36-42 36-42 0 20 6--7 18-20 33 35-37 17-26 21-29 51-58 6--9 47-60 22-23 30-35 39-53 15-20 5-6 Halargyreus 38 23-32 johnsonii 39-42 39-42 39-42 44-46 7-9 0 33-35 11-13 verecundum 19-22 2 16--19 Microlepedium 29-32 62-76 0 11-14 38-41 51-58 9-11 57-67 62-76 nematopus Physiculus 6 22-26 14-19 21-25 50-55 52-62 38-42 56-66 rastrelliger 56--66 0 8-13 24-28 P. 12-14 27-32 24-32 7 Moridae 2. Number Table specimens (above) and size range (in mm, below) used in the preparation of the morid of descriptions. Postflexion Flexion Preflexion Species Transformation Juvenile 1 10 Microlepedium verecundum 2 0 9.4 18.5 2.7-7.5 33.2-36.1 6 10 9 nematopus 5 0 Physiculus 2.4-9.8 8.3-10.9 10.7-13.5 38.0-59.8 5 2 10 0 4 lliger P. rastre 9.6--10.5 3.3-8.4 12.7-18.3 66.7-88.0 501

515 MORIDAE Microlepedium verecundum DESCRIPTION HISTORY LIFE MERISTICS EARLY EGGS Range Mode Vertebrae: diam.: Shell Yolk: 44-46 Total No.ofOG: Diam.ofOG: Shell surface: 11-13 Precaudal Pigment: Caudal 33-35 Diagnostic features: Fins: Dorsal 0 spines LARVAE 7-9+39-42 Dorsal rays <2.7 Hatching Anal length: 0 spines mm length: Flexion mm <18.5 rays mm <9.4 Anal 39-42 - mm <33.2 2 Transformation Pelvic length: 19-22 Pectoral , P PI' sequence: development Fin A, & 20 C" C 10, 2 2 Caudal: P gut, nape, & PI Pigmentation: Preflexion--On brain, fin bases; & 2 29-32 broad stage early some Total tail of end near at midtail, bar in ca. by larvae; & Gill rakers: branchiostegal region; snout, operculum, infraorbital, 6 mm, on absent from posterior half of tips isthmus; & Upper jaw Flexion--On tail. 5-6 11-13 Lower gradually expanding posteriad along tail. Posiflexion-juvenile- Occasionally Branchiostegals line rays; rays P of ends & 20 few first of ends on 7 2 dorsal along tail. of midline lateral margins ventral & & HISTORY LIFE features: Heavy pigment bar Diagnostic midtail; by flexion stage, at on dorsal & ventral lateral on margins, midtail at concentrated myosepta; 39-42; along 20 rays & P to rays 5, decreasing midline, 2 Cabo ca. near area Oceanic Range: 15° to south N) 28° (ca. Lucas San stage, midbody; about anus postflexion through adult; 2 by HL ca. Clarion (ca. 115° W) N & offshore to Isla of 30% ca. BO & forming 33% early as or not rays BL P BL; 2 Physiculus. in as elongate to down fished trawls midwater in Collected Habitat: with m 180 ca. ca. to 3100 m bottom depth %) (range & mean in MORPHOMETRICS icthyoplankton CalCOFI season: Spawning larvae occur in Preflexion collections from January through September Y-S F PrF Tr PoF Juv planktonic Oviparous; pattern: ELH larvae Sn-A/BL 38-53 39-42 48 49 52 41 LITERATURE BOIBL 32-38 23-24 36 28 27 24 25-39' HLlBL 28-30 34 33 36 29 HWIHL 67-87 45-51 ORIGINAL (Illustrator) ILLUSTRATIONS 76 71 56 48 SnLIHL 15-26 17-18 3.3 mm, 5.9 mm (B. larvae, Sumida MacCall) Preflexion 19 21 24 17 MacCall) mm 9.4 (B. Sumida Flexion larva, EOIHL 37-47 29-29 41 39 26 29 5-11 PILlBL 8 14 20 24 P L1BL 0-27 2 >18 13 36 37 502

516 Micro[epedium verecundum 3.3 mm 5.9mm 9.4mm Figure Moridae 1. Preflexion larvae, 3.3 mm (CalCOFI 5706, station 157.20), 5.9 mm (CalCOFI 5706, station 148.30); flexion larva, 9.4 mm (CalCOFI 7205, station 157.10). 503

517 MORIDAE Physiculus nematopus DESCRIPTION HISTORY MERISTICS LIFE EARLY Range Mode EGGS Vertebrae: diam.: Shell Yolk: 51-58 Total No.ofOG: Diam.ofOG: 11-14 Precaudal surface: Shell 38-44 Pigment: Caudal Fins: Diagnostic features: spines Dorsal 0 9-11+57--67 LARVAE Dorsal rays Hatching length: 0 Anal spines rays Anal length: 62-76 mm 11.0 ca. through 8.4 ca. Flexion 6 Pelvic Transformation length: 21-25 Pectoral Fin development sequence: A, & 20 C & P PI' C 10 , 2 2 h Caudal: brain, gut, jaw Pigmentation: & operculum; Preflexion-On angle, 22-26 Total tail; of ca. by ventral patch near midtail; finfold pigment near end mm, 5 isthmus, to form on rakers: Gill at end of pelvic rays, midtail patch spreads 4--6 Upper bar, & a second tail mm, 7 ca. by first; the to anterior forms patch 10--14 above anterior to anus & row heavy & below notochord along Lower patch 7 on & P along Flexion-pos!flexion-Spreading tail. of Branchiostegals lateral most 2 tail dorsal to ventral Juvenile-On notochord. & fins, branchiostegal region, peritoneum, body. & scattered over LIFE HISTORY features: In C on pigment stage, preflexion early Diagnostic & patch 0 stages, postfiexion through pigment late from midtail; at preflexion Panama (ca. Range: California of 31 Northern N) to Gulf Gulf of (ca. 0 anus; to anterior patch ventral heavy tail ventral absent along margin; N) 7 elongate 22-26; rays 6; P early rays; forming, P caudal total rays 2 2 30% BL. ca. at anus m; 42 from specimens from ranging depths at Captured 1271 Habitat: to greater than about depths bottom; 366 m well probably captured off are %) MORPHOMETRICS (range & mean in in taken often juveniles layer scattering near or Y-S ichthyoplankton CalCOFI in Spawning occur larvae Preflexion season: F PrF PoF Tr Juv from April February collections through Sn-AlBL 24-31 24-31 26--32 29-31 27 28 29 30 ELH pattern: Oviparous; planktonic larvae BOIBL 19-22 19-21 18-23 18-21 LITERATURE 21 20 20 19 HLlBL 16--26 24-26 22-26 27-30 Fahay 1984 Markle & 22 24 25 29 HWIHL 71-100 66--80 66--74 52--64 ORIGINAL ILLUSTRATIONS (Illustrator) 79 73 70 56 SnLIHL 11-25 18-25 19-23 21-25 larva, Preflexion mm 2.4 T. Vona) (M. 19 21 21 23 Postfiexion larva, (H. M. Orr) mm 14.1 EOIHL 38--53 33-42 34-40 22-25 41 37 38 23 6--14 12-16 are * Pelvic minimum values because the rays usually were fin lengths PILlBL 15-23 16--33 10 18 15 21 broken. P L1BL* 20-40 8--30 17-43 2 29-39 22 26 26 34 5-11 PIBLlBL 6--7 5-7 2-5 7 6 6 4 504

518 Physiculus nematopus 2.4mm 9.2mm 14.1 mm ORR 14.1 mm Figure Moridae 2. Preflexion larva, 2.4 mm (CFRD 7203 27/3 #9, 30°11' N, 114°11' W); flexion larva, 9.2 mm (Fahay and Markle 1984); postflexion 14.1 mm (CaICOFI larva, 5604, station 103G.40), ventral view (Fahay and Markle 1984). 505

519 MORIDAE Physiculus rastrelliger DESCRIPTION HISTORY LIFE EARLY MERISnCS Range EGGS Mode diam.: Shell Vertebrae: Yolk: 50-55 Diam.ofOG: No.orOG: Total 12-14 Shell surface: Preeaudal 38-42 Pigment: Caudal Diagnostic features: Fins: 0 spines Dorsa! 8-13+52-62 LARVAE Donal rays Hatching length: spines Dorsal 0 8.5-12.5 mm length: ca. Flexion Anal 56-66 rays length: Transformation 7 Pelvic 24-28 Pectoral 2D C, P" sequence: development Fin C" & ID A, & PI' jaw gut, nape, brain, Preflexion-On Pigmentation: & angle, Caudal: 27-32 mm, finfold near end of tail; by 5 cleithrum; along in operculum; Total rows dorsal & ventral internal & heavy isthmus, on anus, to anterior rakers: Gill 7-10 notochord to at midtail. ventral tail margin Flexion-Line along Upper 17-22 near reduced extending almost to trunk; row along dorsal tail margin, Lower on Branchiostegals tail. Posiflexion-On base of P, & PI' on P, rays, & of 7 end anterior tail region. Juvenile-On fins, in dorsum peritoneum, scattered & allover body. region, branchiostegal LIFE HISTORY late Diagnostic features: By row preflexion stage, pigment present 0 0 anus; to anterior patch heavy & Crays total tail ventral along margin Northern Range: (ca. Panama of Gulf to N) N) 41 (ca. California 7 similar 27-32; rays 7; morphology P, to P. voluminous nematopus; fins. at 183-366 m; depth range 128-523 m Habitat: Near bottom, usually %) & (range MORPHOMETRICS in mean ichthyoplankton in occur larvae CalCOH Spawning season: Prefiexion April samples primarily from December through Y-S PrF F PoF Tr Juv Oviparous; pattern: ELM planktonic larvae 27-31 Sn-AlBL 27-30 26-30 28-33 30 27 30 29 LITERATURE 21-23 17-21 BDIBL 21-24 15-18 22 22 19 17 18-21 22-24 27-30 24-32 HUBL 23 19 28 28 nONS ILLUSTRA ORIGINAL (Illustrator) 79-86 64-84 HWIHL 81-100 56-58 82 90 74 57 (M. Vona) T. mm 8.4 mm, 3.3 larvae, Preflexion SnLIHL 16-26 12-20 18-21 21-25 12.7 larva, Postflexion (M. T. Vona) mm 16 19 21 25 EDIHL 41-50 41-45 34-39 23-27 47 43 37 25 the usually because values minimum are lengths Fin * broken. rays were 7-13 15-25 19-29 P,LlBL* 18-20 18 24 8 19 12-26 P,LIBL* 10-21 20-46 17-21 18 15 33 19 5-8 6-7 4-6 PIBLlBL 3-3 7 7 5 3 506

520 rastrelliger Ius . codling Hundre - athom d f: PhYSICU mm 3.3 8.4 mm 12.7 mm . larva, preflexion . late 84mm(C 133.40), station 6712, CalCOFI alCOFI 5602, station 127.50); FigureMon ~e . arva, 12.7 mm (Ca 133.35). ·d 3 Preflexionlarva, 3.31~;:'i 6904; station Iy postflexlOn I - m7

521 MERLUCCIIDAE: Hakes A. D. AMBROSE silvery, steel-grayish the the back and lighter on on family is variously Merlucciidae considered a separate vertical diel make hakes all belly. and Almost sides a subfamily or recent classifica- The Gadidae. of most disperse daytime the when migrations, benthic schools ca. 13 one includes (1994) with tion genus in Nelson Merluccius in feed to night at column. water the species in the Merlucciidae. Merluccius productus, the common remains at bottom when the in hake apparently most the is whiting, or hake Pacific productus the northeastern Pacific. Several possible subpopula- Hake exhibit seasonal (Bailey spawning 1982). al. et identified northeastern the tions have been along and latitudes moving to higher migrations generally seasons and back during to lower the inshore warm the al. (Bailey Georgia of et in (1) coast: Pacific Strait (2) in Puget Sound (Utter and Hodgins 1971), 1982), latitudes and cold deeper water during the season. migratory population extending from (3) a large coastal than live Males grow more slowly which females, aL Hake are 1990). et males longer (Cohen than Canada to hake found Baja California, and (4) a dwarf (Alheit (MacGregor Baja southern off California commercially and ecologically important fishes 1971; and Vrooman and 1977). in hake Pitcher 1995). Giant the Paloma Gulf northern have of been described by California oviparous spherical, are Hake smooth, spawn and Mathews as (1985) Merluccius however, hernandezi; 0.80-1.20 with diameter in mm a buoyant about eggs al. this species is not widely recognized (Cohen et 1983). of oil globule (Fahay most Larvae single 1990). Mar, from ranges angustimanus Merluccius Del described are Markle and (Fahay 1984). wen species Califor- Gulf the including Colombia, to California of with a non-functional round Larvae hatch mouth, 1990). at larvae Only (Ginsburg 1954; Cohen et nia well developed pigment on the eyes, and unpigmented coastal migratory population of to attributable the tadpole their by identified are Larvae body. general productus Merluccius CalCOFI identified been in have the located anus shape, the loop, a single with gut in samples. body, and one to three distinc- of the the half anterior tive pigment bars the to disperse which on tail begin are hakes medium large Merluccid (maximum to Counts (1955) and late flexion by the Ahlstrom stage. and ca. cm) on 70-140 predators demersal pelagic size in described multiple M productus ossification the slope the eastern of the continental shelf and upper the would fins anal and second which of dorsal centers and oceans, Indian southwestern and Atlantic, Pacific, to the divided fins in other gadoids. correspond Zealand. The large head is ca. off New southern M to juvenile the is graduaL In Transformation mouth the dorsally, ridge with BL 25-33% a V-shaped the have all productus, pectorals the except fins fun teeth, the is eye and large. developed with large well is by ca. rays 20 mm. Larvae of M of complements is short, The first dorsal fin high, and triangular while M and angus us are timan undescribed. hernandezi is fin separate second, the long and deeply at notched midlength. dorsal second the resembles fin anal The based on is description species following The placed Pectoral fin. fins are long, slender and posterior from morphometries Counts and (1955). Ahlstrom becomes the pelvic fins. The caudal to progressive- fin literature obtained from (Ginsburg Meristic data were skeleton includes and growth with forked ly caudal the Matarese aL et Cohen 1989; al. et 1981; 1954; Inada are family the Meristics 1981). (Inada bones Y X- for 1990) from obtained was information ecological and 12-18; PI 35-46; A 34-45; 1,7-12; ID 48-58; V 2D et Cohen (1990). and (1982) aL Baileyet al. Br (usually usually body The 7. is and 7); 6-9 P 2 508

522 7 Table Meristic characters for the merluccid species in the California Current vicinity. All species have 1. Merlucciidae rays. branchiostegal Fin rays Vertebrae Gill rakers Total 10 A 2D PrCv P CV U L 2 PI Species 49-52 36-40 14-17 1,9-12 7 36-40 27-29 3-5 angustimanus 21-23 12-14 Merluccius 47-52 36-42 37-42 hernandezi 14-20 total M 1,9-12 14-17 23-25 29-31 52-55 6-8 37-44 37-44 3-6 13-18 productus M 509

523 MERLUCCIIDAE Merluccius productus mSTORY EARLY DESCRIPTION LIFE MEruSTICS Mode EGGS Range Shell diam.: 1.07-1.18 mm Yolk: Vertebrae: mm 1.0 ca. Homogeneous; mm) 1.12 (ave. 53 52-55 Total diam., may appear granular 1 24 ofOG: No. 23-25 mm Precaudal 0.27-0.34 OG: of Diam. 29 0.30 (ave. mm) 29-31 Caudal Shell surface: transparent Smooth, Fins: & head yolk, OG, On embryo of dorsum Pigment: Dorsal spines Late-stage Diagnostic features: separated eggs-pigment into 11+40 4 areas 9-12+37-44 rays Dorsal & 1 ventrally on tail; myomeres >50. o along dorsum Anal spines 0 40 37-44 rays Anal LARVAE 6-8 Pelvic 2.4 mm ca. Hatching length: 16 14-17 Pectoral Flexion length: ca. 8.5 mm Caudal: 6+2-3 mm 30-35 Transformation length: ca. Principal , P A, PI C, 20 & 10, sequence: development Fin Procurrent: 2 Pigmentation: to late-stage embryos. Pre flexion- Yolk-sac-Similar 16-18 Upper of PI base; 1 on occipital region; interorbital region; Medial surface 14-17 Lower ventrolaterally peritoneum; on dorsolaterally ventrally on scattered gut; rakers: Gill rows at midtail; occasionally lateral 4 myomeres ca. bar anus; behind 4 3-6 Upper & tail bar. Flexion-posif!exion-Increasing over head; between anus 16 13-18 Lower bar & mandible; tail becomes 7 on bar premaxilla behind eyes; 7 Branchiostegals along dorsum. Transformation- 16.5 mm, continuous by ca. indistinct; juvenile--On in fin on snout; increasing body; scattered gradually LIFEmSTORY juvenile. membrane by 52-55; A 37-44; preanal length <46% features: Diagnostic Vertebrae (ca. Northern to N) 48° ca. Range: Vancouver Island, British Columbia bar a single BL; at midtail in pigment preflexion & flexion larvae, N, California of Gulf the 23° including indistinct postflexion becoming stage. during m, 1000 mainly concentrated over continental Habitat: Surface to ca. MORPHOMETRICS (range & mean in %)* 500 & slope between 45 & m depth shelf y-S F PrF to April or June Spawning season: January- January with peak in Tr PoF Juv & Baja off California southern California water February in deep Sn-AlBL 42-45 41-46 41-43 36-40 eggs larvae planktonic Oviparous; pattern: ELH & 22-30 BOIBL 17-20 19-24 16-17 LITERATURE 23-27 26-28 28-29 26-29 HLlBL Ahlstrom Counts & 1955 HWIHL Morphometries average of ranges & are values * taken Ahlstrom from the SnLIHL included in the Counts (1955). The following size ranges are preflexion, stages: developmental mm; 3.0-8.4 flexion & postflexion 28-38 EDIHL 36-43 25-29 27-30 combined, & 8.5-19.9 mm; transformation, 20.0-25.9; juvenile, mm. 40.0-46.9 33-35 0-0 0-3 4-28 510

524 productus Merluccius Pacific hake mm 1.07-1.18 mm 2.4 mm 4.3 10.1 mm 15.8 mm 36.0 mm Figure Merlucciidae 1. Egg, 1.07-1.18 mm; yolk-sac larva, 2.4 mm; preflexion larva, 4.3 mm; flexion larva, 10.1 mm; postflexion larva, the 15.8 juvenile, 36.0 mm (Ahlstrom and Counts 1955; mm; fine stippling represents shape and not pigment). 511

525 OPHIDIIFORMES AMBROSE A. D. is unsettled. Rosen and The classification this group of viviparity to define the two ophidiiform of absence ophidioid (1969) considered it a suborder suborders, Patterson The Ophidioidei. and Bythitoidei (Ophidioidei) 1971) treated (1968, Gosline while Gadiformes, the of by families Carapidae and are differentiated Ophidiidae the of (longer it as a suborder of the Perciformes. than We the dorsal fin for follow Cohen fin anal length the and who carapids (1978) assigned it to a separate order. and equal to or shorter for ophidiids) and the Nielsen supramaxillary in of the bone (present presence Ophidiiforms are primarily marine benthic inhabitants carapids). absent and in ophidiids The bythitoid and of depths wide-ranging but occupy habitats bladder swim the by separated are families of in (present sea; the salinities depths greatest the to (intertidal have precaudal for and Aphyonidae) fresh in also absent and Bythitidae hiding or water) and a propensity burrowing vertebrae (Svetovidov 1961). various in substrates in Aphyonidae). in «23 >25 Bythitidae, elongate, less or more These have fishes tapering by represented Ophidiiformes is in genera 18 4 pelvic mental each or jugular present), fins (when with of 3 families California Current vicinity. Larvae the in rays occasionally one or a minute with two and soft of have families these CalCOFI in identified been fins anal and Dorsal spine. spines, without single, are Carapidae, ichthyoplankton samples: and Ophidiidae, the with joined and to extending bases often with long Aphyonidae The Bythitidae. are this in included not caudal more are pterygiophores fin anal and Dorsal fin. because guide their larvae have not been reported from vertebrae. There adjacent the than numerous two are of tows. plankton biology the discussed (1969) Nielsen nostrils on each side 1994). head of the (Nelson group from taken embryos late-stage illustrated and this 1974). Eagle and (Nielsen tissue ovarian or Cohen and Nielsen (1978) presence the used and Suborders Ophidioidei included: families Ophidiidae Carapidae Bythitoidei Bythitidae 512

526 OPIDDIIDAE: Cusk-eels AMBROSE A. D. were included in Brotulidae and Cusk-eels formerly are oviparous and have no Cusk-eels larval ophidiids. Cohen by reviewed been genera The Ophidiidae. have organ, copulatory related the unlike specialized Nielsen is Ophidiidae repre- family The (1978). and Bythitidae. Some ophidiids eggs produce pelagic oval least at by 14 region sented in the California Current in surface the at float which in raft a mucilaginous seven species. Larvae (Cherublemma these of of Mito 1929a; (Sparta least at 1966; stages early their L. Chilara emmelas, taylori, Lepophidium negro pinna, stage Ophidion are eggs scrippsae Fahay 1992). Late Ophidion stigmatistium, scrippsae, and tenta- 0. sp., 1 about off-round, oil without diameter, in mm an identified been have grandis) Spectrunculus tively in globule, (Ambrose neuston in individually float and the meso- of Larvae samples. CalCOFI ichthyoplankton the the in species known of Larvae al. et 1983). and deep-living pelagic Brotulataenia nielseni, hatch vicinity at ca. 2-3.5 mm with Current California filamentosa as niger, Lamprogrammus and Dicrolene unpigmented eyes, small and mouth, non-functional shallow-dwelling Lepophidium as the more well preflexion; during loops gut The buds. fin pectoral larval size at gut looping, as well relative the as Ophidion microlepis, Otophidium and galeoides, loop, features. taxonomic useful of the are position indefatigable unknown. are Chilara Larval taylori CalCOFI the while pattern, sampling throughout range Larvae are elongate slender (BD 7-24% and relatively length preanal BL), ca. with With of BL. 25-50% San Francis- of Ophidion scrippsae are not found north growth, larvae mouth become more compressed, the Point co Bay. A single preflexion larva from the migrate becomes less oblique, and the pelvic fins Conception vicinity has been identified tentatively as anteriorly. 78.2.59.4). station 8701, Spectrunculus grandis (CFRD and nigropinna Lepophidium larvae stigmatistium L. branchiostegal early the region on Pigment appears mainly California, associated Baja off commonly occur Lepophidium All 2-10 have larvae for species. several emmelas Cherublemma bays. near refugia tropical with base 1-2 large spots along the and of the anal finfold larvae been have occasionally sp. taken and Ophidion of end of tail. the the Larvae dorsally spots near the of pattern near CalCOFI end from the southernmost have small of Chilara and Ophidion stellate patterns San Lucas, Baja California Sur. Young Brotula Cabo al. on the body (Gordon et melanophores laterally also trawls midwater in collected been have clarkae in 1984). Fahay (1992) noted the Lepophidium that area. this from to posterior and the ossify vertebrae from anterior Ophidion, vertebrae In early. pectoral form rays fin shallow cusk-eels small (15 cm for Adult some are and the middle the toward ends both form from species) water 1.5 medium to length (about for some m late. form rays fin pectoral C. and C. taylori emmelas origin anal fin benthopelagic deep-sea forms) with the nektonic extended an have stage. prejuvenile the dorsal origin fin dorsal the behind well and and anal continuous with the caudal fin. Scales are fms study area, fishes the Among larval in the CalCOFI or and cycloid (in regular rows small at angles oblique Embassichthys the pleuronectid, bathybius, superficially pair is of each anterior nostril The other). to each Spectrunculus those resembles identified as tentatively possess Males on unique a the snout. higher grandis S. larvae (see grandis et al. Materese 1989). gas the from formed apparatus sound-producing and a less 65) vs. (>75 myomeres more have pro- ribs, and vertebrae, which is used in bladder, perhaps or activities (Rose 1961; Courtenay mating courtship nounced gut loop than E. larvae. Bythitid bathybius 1971). Cusk-eels for an important food source are pigmentation larval tends that than clumped more be to marine birds, and fish (Fitch and Lavenberg mammals, Several volume). this Bythitidae, (see ophidiids of also volume) this Stichaeidae, (see species stichaeid 1968). gut resemble some ophidiids before the superficially Gordon early life the summarized (1984) al. et postanal the and pigmentation the however, has looped; several of history literature and presented illustrations families. total two count will differentiate the myomere 513

527 Among C. most northern ophidiids, the taylori has the could using separated be probably indefatigable meristic distribution and the (Table counts highest Ophidiidae meristic differences (Table 1). Ophidiidae C. 1). head largest the has emmelas and greater (usually number lowest the BL) 22% than L. negro myomeres has (55-58). a higher of pinna following are descriptions based on literature The myomere count (75-77) than stigmatistium (68-71) L. and detailed examinations on of of 1-34 specimens usually has two dorsal pigment spots near the end and each (Table Ophidiidae 2). Postflexion Brotula species of the L. for one to compared tail, stigmatistium. clarkae, Lepophidium stigmatistium, of series larval Ophidion scrippsaeand Ophidion sp., stages larval early and lemma the Cherub of sp. both have Ophidion tail; dorsally the on however, most pigment Ophidion emmelas, Lepophidium pinna and negro are described time. the for first is similar to (70-76 Ophidion 65-69), vs. vertebrae of number a higher has sp. sp. Ophidion midlateral pigment (not present myoseptal 0. in was described by Lea (1980, unpub- which imitator distribution scrippsae), and a more southern (south of lished dissertation). Meristic data were obtained PhD 1899; Garman 1891; (Alcock sources literature from identified by may be clarkae Brotula Lucas). San Cabo patch the low total vertebral count (55-56), an oval of and 1974; Cohen 1962; 1961, Robins Cohen Nielsen Lea 1978; Lea and Robins 1978; melanophores most covering et stellate Ambrose al. and 1980; the of cheek, which are present by postflexion. The mouth barbels Rasonarivo and Aboussouan 1984; Machida 1983; and counts 1986; made Matarese from 1989) al. et larvae unknown of Brotulataenia nielseni, Dicrolene this study. was information during Ecological obtained Lepophidium niger, filamentosa, Lamprogrammus from primarily Lea and (1980) 1962). (1961, Robins Otophidium microlepis, Ophidion galeoides, and 514

528 Table Ophidiidae Meristic characters for the ophidiid species in the CalCOFI area. A minute, hidden pelvic spine may be 1. but present included in the counts. All have 2 pelvic fin rays, except they are absent in Brotulataenia nielseni and is not 5-11 as a juvenile. Dicrolene filamentosa have the lower only rudimentary buds which have niger Lamprogrammus may rays (Cohen and longer than the upper rays fin and Nielsen 1978). pectoral free counts rudiments. raker Gill include rays fin Vertebrae Total CV D C PrCV A GR BrR P, Species 26-28 79-89 8 55-56 I~S-II 40-41 6+5 4-6+17-21 15 8 clarkae Brotula 69-73 85-91 62-72 23-26 3-4+11-14 12-13 9 7 nielseni Brotulataenia 4+5 55-58 99-113 81-93 24-26 4-5+12-19 42-44 13-14 7 emmelas Cherublemma 187-229 150-181 22-26 18-19 68-72 86-91 4+5 1-4+7-9 7 taylori Chi/ara 84-90 22-23 13-16 100-104 5+17 8 filamentosa Dicrolene 4+4 17 ca. 110 ca. 90 12-14 8 niger Lamprogrammus 4+5 51-62 66-77 117-141 97-121 21-26 9-15 14-16 7 Lepophidium microlepis 21-24 116-121 3+5-13 4+5 75-77 138-148 59-61 15-16 7 L. negro pinna 68-71 52-56 122-130 3-4+13-15 99-109 4+5 19-22 14-16 7 L. stigmatistium 1-3+4-5 4+5 21-23 97-114 61-64 123-143 47-49 14-17 7 Ophidion galeoides 50-54 20-23 124-153 99-126 2-4+4-8 4+5 65--69 14-16 7 scrippsae 0. 3-4+9-12 4+5 70-76 135-163 112-139 25-28 54-60 7 15-16 Ophidion sp. 2+4 4+5 18-19 88-96 106-115 45-49 59-64 7 13-15 indefatigabi/e Otophidium 4+4 22-23 53-56 71-79 103-148 73-113 3-4+8-9 18-23 8 Spectrunculus grandis 2. Number of specimens (above) and size range (in Ophidiidae below) used in preparation of the ophidiid species Table mm, descriptions. An ilL" literature used in the description. indicates Postflexion Flexion Transfonnation Preflexion Juvenile Yolk-sac Species 2 4 0 0 0 Brotula clarkae 40.0-44.3 50.0 63.5-80.6 10 4 10 5 5 0 Cherublemma emmelas 14.7-42.0 45.0-58.1 60.8-75.6 2.9-10.5 11.8-13.3 La La La La 5 5 taylori Chilara 94.8-104.7 71.7-84.4 10 6 7 0 0 5 Lepophidium negro pinna 21.2-30.5 4.2-10.6 10.5-14.5 3.0-3.6 7 0 5 6 10 L. stigmatistium 13.3-17.7 45.6-61.1 1O'\-12.S 3.9-10.0 2.4 La La La La 5 5 scrippsae Ophidion 58.5-71.8 97.6-110.5 1 5 0 Ophidion sp. 54.0-88.0 36.5 12.7 7.0 2.7 Lb C Lb,l 0 0 L 0 grandis Spectrunculus 7.4 et al. 1983 a Ambrose b Matarese et al. 1989 C Jordan and Thompson 1914 515

529 OPHIDHDAE Bl'otuia clarkae MERISTICS EARLY LIFE HISTORY DESCRIPTION Mode EGGS Range diam.: Vertebrae: Shell Yolk: 55-56 Total 55 No.ofOG: Diam.ofOG: Shell Precaudal 15 surface: 15 40--41 40 Pigment: Caudal features: Fins: Diagnostic spines Dorsal 0 0 108-118 115 Dorsal rays LARVAE 0 Hatching length: 0 spines Anal Flexion length: 78-89 Anal 86 rays length: Transformation <63 Pelvic* through mm ca. 50 development Fin Pectoral 26-28 sequence: 27 Pigmentation: Caudal: & mid- over scattered mm, 40 Posiflexion-By 6+5 6+5 oval Principal hindbrain; under eye; most patch (stellate melanophores) on of cheek; base along dashes several over scattered evenly peritoneum; ProCllrrent: of row of & A pterygiophores; scattered medially on D; D under middle Upper 0 0 0 Lower 0 spreading D, PI on distally mm, 50 Transformation-By anteriad. Gill rakers: heaviest body, entire over at scattered mm, 65 Juvenile-By rays. 22-26 23 Total edges unpaired fins. of 4--6 Upper 5 55-56; total 27); Diagnostic features: Vertebrae (usually PI rays 26-28 17-21 18 Lower & 3 above on 12 barbels, 3 below Crays 6+5; branchiostegal rays 8; 8 each side oval 8 patch of Branchiostegals of mouth; long PI & P, rays (>12% of BL); Barbelst cheeks; stellate melanophores on meianophores large dashed tail. posterioiaterally on LIFE HISTORY %) MORPHOMETRICS (range & mean ill. Cabo Range: N) (6° Peru to N) (24°30' Sur California Baja Lazaro, San Usually Habitat: collected 40-75 m deep on mud & broken shell bottom PrF Y-S PoF F Tr JlIV season: Spawning Sn-A/BL 46--46 43--48 46 44 44 Oviparous; late postflexion & transformation stage larvae pattern: ELH BDIBL 16-19 17-19 in midwater 17 17 18 HLIBL 22-23 24--26 LITERATURE 22 23 25 HW/HL 31-33 30-33 Hubbs 1944 32 30 32 SnLIHL 18-21 20-25 (Illustrator) ILLUSTRATIONS ORIGINAL 20 20 22 EDIHL 22-22 17-20 larva, Walker) C. (R. mm Posttlexion 44.3 22 22 18 21-21 PILIBL 13-14 21 14 20 ray length. unequal of branches two * One with three t below on each side-12 total. Three above mouth & P LIBL 13-15 15-16 2 14 13 16 516

530 Brotula clarkae -.-.---." .. .. 44.3 mm Figure Ophidiidae 1. Postflexion larva, 44.3 mm (SIO 71-343). 517

531 OPHIDIIDAE Cherublemma emmelas MERISTICS DESCRIPTION HISTORY LIFE EARLY Range EGGS Mode Vertebrae: diam.: Shell Yolk: Total 55-58 No.ofOG: 57 Diam.ofOG: surface: 14 Shell 13-14 Precaudal Pigment: Caudal 42-44 43 Fins: features: Diagnostic 0 spines 0 Dorsal LARVAE Dorsal rays 99-113 105 <3 mm length: Anal 0 0 spines Hatching 81 & 85 81-93 Anal Flexion length: ca. II mm rays mm ca. length: Transformation 45-60 Pelvic* 25 24-26 development Fin D sequence: , C & A & Ph Pectoral P 2 Caudal: still mm, 2.9 Preflexion--By patches eye, on forming Pigmentation: 4+5 4+5 Principal above, under, tail, near gut; ventrally along anus, & to anterior patches Procurrent: by tip; at except tail of half posterior on ventrally & dorsally heavy 0 0 Upper on & bladder, gas ca. 4.5 mm, jaw, PI base, cheek, snout, lower 0 Lower 0 ca. branchiostegal cleithral symphysis; by 7 mm, on region, & Gill rakers: third of midlaterally tail; by ca. posterior mm, spreading along 8 18-22 20 Total on head, body, & tail, gut Flexion-juvenile-Increasing laterally on 4-5 & A on especially peritoneum & margins. Upper D 5 12-19 IS Lower Diagnostic features: total vertebral Low Dray (55-58) & (99-113) 7 7 Branchiostegals counts; gut loop formed by >ca. 5.6 mm; head relatively large (HL ca. mm; 22% BL); in preflexion larvae, pigmentation on PI 4.5 base by LIFEmSTORY in flexion larvae, pigmentation on D & A & row posteriolaterally on on tail; increasing on peritoneum pigment postflexion larvae & juvenile specimens. Southern Range: N) (ca. California of to Peru (ca. Gulf 26° of portion I, 14.076) station EASTROPAC S, 9° %) (range & mean in MORPHOMETRICS at ca. bottom m 102-740 Habitat: Relatively rare; on green mud depth Y-S PrF F PoF Tr Juv Preflexion season: Spawning larvae taken flexion & during stage Sn-NBL February tows ichthyopJankton CaiCOFI in 38-54 43-49 36-44 37-40 38-41 44 46 39 39 38 Oviparous; larvae, extended pelagic juvenile pattern: planktonic ELH BD/BL 12-27 21-26 17-24 14-20 15-20 stage 19 24 19 17 18 HUBL 17-27 23-29 22-27 24-28 25-29 LITERATURE 22 26 24 26 27 HWIHL 46-50 50-72 31-44 28-39 32-34 1926 Trotter 60 48 37 34 33 SnLIHL 16-30 30-35 20-34 21-24 17-21 ILLUSTRATIONS ORIGINAL (Illustrator) 25 25 33 23 19 EDIHL 21-44 20-21 14-17 11-20 16-18 6.3 2.9 larva, Preflexion mm (N. Arthur) mm, 29 14 21 IS 17 Arthur) (N. mm 12.2 Flexion larva, 2-6 4-7 PILlBL 7-24 13-14 12-15 4 5 13 13 13 One * length. of branches two with ray unequal P L1BL 3-3 4-6 5-13 2 7-9 7-8 3 5 9 8 8 518

532 Cherublemma emmelas 2.9mm 6.3 mm 12.2 mm 33.0 mm Figure Ophidiidae 2. Preflexion larvae, 2.9 mm (IATTC 90002, station MAB #1),6.3 mm (CalCOFI 7202, station 157.10); flexion larva, larva, 33.0 5502, station 150.25); postflexion (CalCOFI mm (Trotter 1926, as Cherublemma lelepris). 12.2 mm 519

533 OPIDDIIDAE Chi/ara taylori mSTORY DESCRIPTION EARLY LIFE MERISTICS EGGS Mode Range Shell diam.: Yolk: Vertebrae: 86-91 No.ofOG: 88 Diam.ofOG: Total 18-19 Shell surface: 18 Precaudal Pigment: 70 Caudal 68-72 features: Diagnostic Fins: 0 0 Dorsal spines 187-229 202 LARVAE Dorsal rays 3.5 mm length: ca. Hatching 0 0 spines Anal mm 21-30 ca. 161 length: Flexion 150-181 Anal rays 70-84.5 mm Transformation length: ca. Pelvic* 24 , P sequence: development Fin D, C, PI 22-26 A, Pectoral 2 Yolk-sac-Along Pigmentation: Caudal: tail. of 30% dorso-posterior & ventrum 4+5 4+5 Principal angle Preflexion--posiflexion-On region; double oflower jaw & gular extending base along internally along ventrum row brain & onto of Procurrent: 0 ca. caudal finfold; by snout; on lateral on mm, 11 tail midline, 0 Upper 0 0 increasing anteriad; by Lower ventrally ca. 36 mm, decreasing & generally scattered. more Gill rakers: 11 9-12 Total & Dray (86-91) vertebral total (187-229) features: High Diagnostic 3 1-4 half of the visceral cavity by counts; loop forms in the posterior gut Upper 8 7-9 ca. on on line caudal brain; Lower under finfold; pigment SL; mm 14 snout; 7 tail ventral extending along a double 7 Branchiostegals in midline row; row along lateral of tail. HISTORY LIFE (range %) mean in & MORPHOMETRICS 0 (47 to Washington Elizabeth, Cape Range: N) Santa Maria, Bahia Baja 0 Y-S PrF F PoF California Sur N) (25 Tr Juv Sn-AlBL eelgrass, in mud, also sand, prefers tail-burrowing, Nocturnal, Habitat: & 33-44 31-44 29-32 25-29 29-33 27-30 39 39 31 27 29 31 rock depth on bottoms from <1 m to 283 m rubble 7-11 6-10 BDIBL 7-8 7-8 8-9 9-10 Early-stage larvae Spawning season: occur in CalCOFI ichthyoplankton 9 8 7 7 9 10 in September off Baja from mainly tows a peak with July-October 10-15 8-15 HLlBL 11-13 11-14 12-14 15-16 California 12 13 12 12 13 16 HWlHLt 37-39 31-37 ELH extended pelagic Oviparous; planktonic pattern: larvae, juvenile 34 38 stage 14-27 19-40 26-31 SnLIHL 21-27 19-21 18-22 20 27 29 25 20 20 LITERATURE 23-41 22-26 EDIHL 29-41 21-25 21-24 22-23 24 36 30 23 23 23 1983 al. Ambrose et novaculum) Harry (as Ophidion 1951 2-6 2-3 3-3 PILlBL 2-3 5-6 7-8 3 3 3 et al. 1989 Matarese 3 6 7 L1BL 0-0 P 0-1 0-0 1-3 5-7 2 7-8 0 0.6 0 2 6 8 One ray with two branches of unequal length. * t specimens. Head width not measured in yolk-sac through postflexion 520

534 Spotted cusk-eel Chilara taylori 5.0mm 4.8mm 7.3 mm 14.4 mm Figure Ophidiidae 3. Yolk-sac larva, 5.0 mm; preflexion larvae, 4.8 mm, lateral and ventral views, 7.3 mm, 14.4 mm (Ambrose et al. 1983). 521

535 OPHIDIIDAE Lepophidium negro pinna HISTORY EARLY LIFE MERISTICS DESCRIPTION Range Mode EGGS Yolk: diam.: Shell Vertebrae: 75 No.orOG: Total 75-77 Diam.ofOG: surface: Shell 15-16 16 Precaudal Pigment: 59 59-61 Caudal features: Diagnostic Fins: 0 o spines Dorsal LARVAE 143 138-148 rays Dorsal Hatching length: 3 ca. mm 0 o Anal spines 117-119 nun length: 116-121 Flexion ca. 10.5-15 rays Anal ca. 31 mm length: Transformation > Pelvic* , sequence: development Fin D C, PI> 23 & A P 21-24 Pectoral z Pigmentation: Yolk-sac-Row onto snout; below eyes extending Caudal: 4+5 yolk; scattered 2 dorsally on gut; 1 anterior on to anus; 4 ventrally 4+5 Principal Procurrent: of dorsally near end 2 tail. Preflexion-On angle oflower along tail; PI; o above 0 gas loop; gut of at bottom jaw; ventral body midline below Upper o 0 bladder; Lower ventrally patches 4-6 1 tail; along (small) occasionally of tail dorsal two the patches; between main ventral anterior Gill rakers: 9-16 melanophores absent Flexion-Along ventral midline of occasionally Total branchiostegals; on gut; jaw lower Posiflexion-On A rays. of base on 3 Upper 5-13 &onC. Lower 7 7 gut ca. loop by 75-77; vertebrae Total features: Diagnostic formed Branchiostegals usually centrally 2 cavity; visceral 6.8 nun & positioned in the near tail. of pigment patches dorsally end HISTORY LIFE %) in MORPHOMETRICS (range & mean & Baja of coast along N), California 28° Sur, Cedros Isla Range: (ca. Gulf inshore to California of mainly between collected larvae Peru; Y-S F PrF PoF Tr 113 & 130 CalCOFI lines Juv 38-44 Sn-NBL Habitat: Sandy mud, 22-139 m depth 28-39 36-42 36-47 40 41 38 33 occur Spawning season: larvae Early-stage mainly from August-October BDIBL 11-15 13-15 13-16 6-9 ichthyoplankton tows in CalCOFI 14 14 7 13 15-20 18-20 14-15 HLlBL 15-21 Oviparous; planktonic larvae pattern: ELH 14 18 19 17 43-52 49-73 HWIHL 68-82 42-46 LITERATURE 62 47 45 75 20-36 27-36 15-19 SnLIHL 25-32 1984 al. et Gordon 31 31 28 18 EDIHL 38-43 21-37 21-24 17-20 (Illustrator) ILLUSTRATIONS ORIGINAL 21 28 40 19 0-2 4-5 4-5 PILlBL 5-7 Yolk-sac larva, 3.2 mm (N. Arthur) 4 5 6 0.8 13.3 mm (N. Arthur) Flexion larva, (N. Arthur) mm 30.0 Postflexion larva, 0-1 2-6 0 7-9 PzLIBL 4 0.4 0 8 length. * One ray with two branches of unequal 522

536 Specldefin cusk-eel pinna negro Lepophidium \ ,. , '. " . ... . ~ 3.2mm mm 5.1 13.3 mm mm 30.0 Figure Ophidiidae 4. Yolk-saclarva, 3.2 mm (CaICOFI 6607, station 117.40); preflexion larva, 5.1 mm (Gordon et al. 1984); flexion larva, 13.3 (CaICOFI mm 5709, station 120.40); postflexion larva, 30.0 mm (CaICOFI 5410, station 117.40). 523

537 OPHIDIIDAE Lepophidium stigmatistium MERISTICS LIFE HISTORY DESCRIPTION EARLY Range Mode EGGS Vertebrae: diam.: Shell Yolk: Total 68-71 69 No.ofOG: Diam.ofOG: 14-16 16 Shell surface: Precandal 52-56 53 Caudal Pigment: Fins: Diagnostic featnres: 0 0 spines Dorsal 122-130 126 LARVAE Dorsal rays Anal mm <2.4 length: Hatching 0 spines 0 length: Flexion mm 10-13 ca. 103 99-109 rays Anal Transformation length: ca. Pelvic* <45 mm 19 19-22 , Pectoral PI C, & P sequence: development Fin A & D 2 Caudal: Pigmentation: 5 ventrally anus; above gut of angle at Yolk-sac-I 4+5 4+5 Principal in I dorsally above series. last along tail; ventral tail one Pre flexion- Procurrent: On angle of lower jaw; 2 under 5-7 gut; gut; 3 internally 2 or above 0 0 (occasionally smaller ones along main ones). ventrally Upper tail between 0 0 Lower on posteriolaterally region; branchiostegal Flexion-posiflexion-On Gill rakers: gut; increasing on gut ventrally Juvenile-Dorsally & along A base. 18 17-19 Total along on & on head & body; ventrolaterally on tail; dorsolaterally D 3-4 4 margins. A Upper 14,15 13-15 Lower Total features: Diagnostic vertebrae 68-71; gut loop completed by ca. 7 7 Branchiostegals in centrally visceral dorsally patch pigment one cavity; forms mm & 6 tail. of near end LIFE mSTORY MORPHOMETRICS (range & mean in %) N), (ca. 28° Cedros Isla Range: along coast of Baja California Sur & into Gulf the Y-S PrF California of F PoF Tr Juv Habitat: Demersal, Sn-AlBL rn 20-238 depth 36-42 34-44 37-43 35-37 44 39 38 39 36 Spawning August- collected in larvae Early-stage mainly season: BDIBL 13-16 15-17 14-17 12-12 December 15 10 16 16 12 HLlBL 19-21 19-22 19-21 21-22 Oviparous; ELH pattern: larvae planktonic 19 20 20 20 21 HWIHL 51-75 45-54 42-49 33-35 LITERATURE 61 51 72 44 34 SnUHL 28-36 23-37 24-39 18-22 32 44 33 32 19 EDIHL 20-30 21-24 18-23 18-20 39 25 22 20 19 ORIGINAL ILLUSTRATIONS (Illnstrator) 4-5 4-5 5-5 PILlBL IO-II 5 0 5 5 10 Yolk-sac larva, Arthur) 2.4 mm (N. larva, Preflexion (N. mm 5.1 Arthur) P L1BL 0-3 3-6 7-8 6-7 2 Flexion larva, Arthur) mm 1l.2 (N. 4 0 1 8 7 15.9 mm (N. Arthur) larva, Postflexion One * length. ray with two branches of unequal 524

538 Blotchfin cusk-eel stigmatistium Lepophidium 2.4mm 5.1 mm 11.2 mm 15.9 mm Figure Ophidiidae 5. Yolk-sac larva, 2.4 mm (CalCOFI 6909, station 133.30); preflexion larva, 5.1 mm (CaICOFI 6310, station 133.23); flexion 11.2 mm larva, (CaICOFI 6310, station 137.30); postflexion larva, 15.9 mm (CaICOFI 6509, station 133.30). 525

539 OPHIDIIDAE Ophidion scrippsae MERISTICS DESCRIPTION HISTORY LIFE EARLY Mode Range EGGS Vertebrae: 1.06 1.00 diam.: mm x Shell Yolk: Homogeneous; ca. 65-69 x (0.96-1.00 67 mm) 1.00-1.06 Total 0.72-0.84 diam. mm Precaudal No.ofOG: 0 15 14-16 Diam.ofOG: 50-54 Shell Smooth, Caudal transparent surface: 52 Fins: Scattered mainly Pigment: along dorsum of embryo spines Dorsal 0 0 Off-round features: Diagnostic pigment with embryo slender long egg; 124-153 Dorsal rays 136 posterior scattered dorsum along to gut length just short of head; Anal spines midbody 0 0 Anal rays 99-126 109 Pelvic* LARVAE Hatching 20-23 21 25 ca. length: mm Pectoral length: Flexion Caudal: mm 9.6-11.5 ca. 4+5 4+5 length: mm 30-55 ca. Transformation Principal , sequence: P Fin PI C, A, D, development Procurrent: 2 0 0 Upper near row Pigmentation: margins scattered over body; Yolk-sac-Finely 0 0 Lower tail; & gut on flexion-Ventrally Pre finfolds. anal & dorsal of near angle to PI; at of lower jaw; Gill rakers: posterior anus; on body dorsally 6-II 8 Total middle base. PI above internally jaw; lower of & tip near frequently 3 2-4 Upper on laterally extending Flexion-On branchiostegal region; dorsally gut Lower 4-8 5 posterior on gut. A; Pos!flexion-Increasing laterally on tail & 7 7 Branchiostegals Transjormation-juvenile-On C. D from absent generally & A & D margins. HISTORY LIFE at loop 65-69; formed gut vertebrae Total features: Diagnostic yolk-sac mm 7.5-8.5 in the central portion of the visceral cavity; in 0 in preflexion stages, pigment row through dorsal & anal fmfolds; on Tosca, California (ca. Arguello, 34 Baja N) to Punta Range: Point 0 absent body, on laterally blotches stellate stages, flexion-postfiexion N) 24 (ca. California midline. lateral from mud in burrows also bottoms, sand on Usually Habitat: m 3-70 ca. from in mean & (range MORPHOMETRICS %) depth occur Spawning in Early-stage season: larvae Y-S ichthyoplankton CalCOFI PrF F PoF Tr Juv July tows mainly through peak. in August-September off October with a California Baja Sn-AlBL 47-53 45-48 42-46 36-42 35-37 36-40 45 46 50 39 36 38 larvae reared laboratory settled larvae; ELH pattern: Oviparous; pelagic BDIBL 5-10 16-17 9-13 13-17 12-13 12-16 to at bottom the stage, postfiexion (early mm 12 ca. after days 36 II 7 16 15 13 14 hatching) mm (day 46) the & sand burrowed by in 18 ca. HUBL 16-20 16-18 19-23 16-20 18-19 18-20 18 17 21 19 19 19 LITERATURE HWlHLt 37-43 38-42 38 39 Ambrose a1. 1983 et SnLIHL 20-25 21-34 26-33 21-31 20-24 21-28 28 23 25 30 22 23 EDIHL 29-36 10-37 22-30 21-28 21-24 19-23 ray with * two branches of unequal length. One 23 31 26 24 22 21 on yolk-sac Head through postfiexion specimens. t width not measured 3-8 PILlBL 6-9 0-3 10-11 9-12 6-10 6 7 2 8 II II P L1BL 0-0 0-0 0-2 3-8 9-II 9-II 2 0 0 5 10 10 526

540 Basketweave cusk-eel scr~J7sae l}J7hidion ... .. .. .. .. • • '" .. At.", ... .. 3.8 mm x mm 1.0 1.1 4.8mm 10.6 mm 15.0 mm mm 27.4 Figure Ophidiidae 7. Egg, 1.0 x 1.1 mm (Matarese and Sandknop 1984); yolk-sac larva, 3.8 mm; preflexion larva, 4.8 mm; flexion larva, 27.4 10.6 postflexion larvae 15.0 mm, mm; mm (Ambrose et al. 1983). 527

541 OPHIDIIDAE Ophidion Sp. MERISTICS DESCRIPTION EARLY LIFE HISTORY Mode EGGS Range Vertebrae: Shell diam.: Yolk: Total 70-76 No.ofOG: 72 Diam.ofOG: Precaudal 15-16 16 Shell surface: Caudal 54-60 56 Pigment: Fins: features: Diagnostic Dorsal spines o 0 135-163 rays Dorsal LARVAE spines Anal 0 o length: Hatching <2.7 mm II 2-139 rays Anal length: mm Flexion <12.7 <54 length: Transformation Pelvic" mm 25-28 Pectoral sequence: C & P P" Fin & D & A development z Caudal: above hindbrain; under snout; Yolk-sac-On Pigmentation: below & 4+5 4+5 Principal gut; ca. 2 externally in area; 17 tail; 17 ca. along ventrally PI ca. Procurrent: to Pre dorsally, extending aoteriorly almost flexion- aous, eyes white. Upper o 0 flexion-Anterior braochiostegal to on gular & eye; regions; ventral o Lower 0 gut; on row mid- & ventro- row internal hindbrain; & on laterally Gill rakers: Pos{/lexion- tail. body; midlateral row along along dorsolaterally 12-15 Total 13 laterally Juvenile- Fading on gut tail. body of & except near end on 3 Upper 3-4 end near heaviest body, over scattered Finely of blotching dorsal tail, 9-12 10 Lower SL. mm 90-100 ca. by 7 7 Branchiostegals 25-28; rays PI 70-76; loop Total features: Diagnostic gut vertebrae by cavity visceral the in centrally formed is well & mm forms at >7 HISTORY LIFE patches along 13 pigment numerous by ventrum; & dorsum mm; ca. on pigment midline; internal pigment heavy lateral stage, flexion body. on laterally Range: Gulf Sao the to N) 23° (ca. Sur California Baja Lucas, of Cabo California Paoama; not known from the of Gulf %) MORPHOMETRICS (range & mean in to 18-32 On Habitat: m; adults collected muddy saody substrates at ca. m 40 Y-S thao deeper PrF F PoF Tr Juv Spawning season: Sn-AlBL 37-38 40 39 39 33 38 planktonic ELH pattern: Oviparous; larvae, extended pelagic BDIBL II-15 stage traosformation 10 14 12 12 13 HLlBL 20-22 LITERATURE 18 19 17 19 21 HWIHL 30-36 72 42 42 38 32 SnLIHL 18-22 10 25 27 21 20 (Illustrator) ILLUSTRATIONS ORIGINAL EDlHLt 18-21 48x28 25 29 19 19 mm Arthur) Yolk-sac larva, 2.7 (N. Preflexion larva, 7.0 mm (N. Arthur) PILlBL 100II 22 4 0 5 10 mm (N. Arthur) Flexion 12.7 larva, PzLIBL 13-17 0 2 0 7 15 * ray with two braoches of unequal length. One fIrst, vertical on yolk-sac specimen; horizontal axis is given t Eye oval second. axis 528

542 Ophidion sp . .- .' . ~(8..;:"~:[Q- mm 2.7 7.0mm 12.7 mm Figure Ophidiidae 6. Yolk-sac larva, 2.7 mm (IATTC 90012, station MSB #1); preflexion larva, 7.0 mm (IATTC 90006, stationASL #2); flexion larva, 12.7 mm (CaICOFI 5503, station 157.10). 529

543 OPHIDIIDAE (?) is grand Spectrunculus EARLY DESCRIPTION HISTORY LIFE MERISTICS EGGS Mode Range Yolk: Vertebrae: Shell diam.: 71-79 Total No.ofOG: Diam.ofOG: 18-23 Shell surface: Precaudal 53-56 Pigment: Caudal features: Diagnostic Fins: 0 Dorsal spines LARVAE 103-148 rays Dorsal Hatching length: 0 Anal spines 73-113 mm Flexion length: >15.6 Anal rays length: Transformation <52 mm Pelvic* 22-23 development sequence: Fin Pectoral fore- Preflexion--On Pigmentation: & jaw; midbrain; lower Caudal: 4+4 ventrally on tail; gut 2 patches 3 patches dorsolaterally loop; on at Principal dorsum; heavy on end on tail. Procurrent: of Posiflexion--4 patches dorsum. 0 8; gutloop Total vertebrae Diagnostic branchiostegals 71-79; features: Upper 0 of mm cavity; visceral the portion posterior the in Lower formed <7.4 at 3-4 pigment patches body; & 2 ventrolaterally on heavy dorsolaterally rakers: Gill 3-4 at end Upper of tail. Lower 8-9 MORPHOMETRIeS (range & mean in Branchiostegals 8 %) LIFE HISTORY Y-S F PrF Tr PoFt Juvt 36-38 Pacific & Sn-AlBL Range: Found widely in Atlantic 43 37 43 at Habitat: or near the on bottom Adults 800-4300 m depth 9-10 BDIBL 10 21 25 season: Spawning HLlBL 9-12 II 17 16 ELH larvae Oviparous; pattern: planktonic HWIHL, 65 LITERATURE SnLIHL 14-45 29 32 35 Rasonarivo & 1986 Aboussouan 38-44 EDIHL Jordan Thompson 1914 & 41 14 II et Matarese 1989 al 1980 Hureau & Nielsen 3-4 P1LIBL 4 6 9 P L1BL 0 2 * in rays Two thick each. 0 5 7 t Measurements taken & postflexion mm 29.8 of illustrations from mm 15.6 (1989). al. et Matarese in specimens prefiexion 56 mm from illustrations of taken Measurements specimen from Jordan t & Thompson (1914). specimen. , Head width only measured on 7.4 mm prefiexion 530

544 Giant cusk-eel grandis Spectrunculus mm SL 15.6 mm SL 29.8 56.0 mm Figure Ophidiidae 8. Preflexion larva, 15.6 mm; postflexion larva, 29.8 mm (Matarese et al. 1989); juvenile, 56.0 mm (Jordan and Thompson 1914). 531

545 CARAPIDAE: Pearlfishes A. D. AMBROSE A few their have exterilium larvae, named for species are reported in the eastern Three carapid species appendage (Gordon et al. 1984); however gut trailing (Table Carapidae Pacific 1990). Markle and Olney 1; none of these is found in the eastern Pacific. and Sur California Echiodon Baja from ranges exsilium En- of Galapagos the to California Gulf Islands. the vermicularis cheliophis is N and found between 30° in collected been have larvae Only vexillifer stage the western of the in Pacific, Gulf in S 30° and Aden, elongate CalCOFI ichthyoplankton samples. These are eastern Pacific including in the the Gulf of California ofBL), large «20% with larvae a moderate-sized head and Galapagos Islands. the ius dub may be Encheliophis (ca. eyes a large a gut rosette, nasal HL), 30% that of Gulf the but from are records confirmed widespread, length preanal a short TL), mm «5 (ca. early coils from California to Columbia and Hawaii. Only larvae in 150% and a tapering tail HL), that frequently ends ichthyo- CalCOFI in found been have exsilium Ec. of a broken is moderate, typically Pigmentation filament. collections, plankton numbers at the southern in modest becoming more sparse with development. of sampling the pattern. end «30 cm) carapids are small to medium-sized Adult The forms early (bud visible even in yolk- vexillum and temperate in occur which fishes specialized highly sac placement is of exsilium) systematic Ec. and its with waters, and slope shelf marine tropical the highest Echiodon, the to the vexillum is just anterior In use. seas tropical abundance 1994; in diversity and (Nelson while ray; in first dorsal fin Encheliophis, there is a and Markle Although are species 1990). Olney some of the gap origin the and vexillum the between wide exsilium), Ec. most are inquilines in (e.g., free-living vertebrae 8-11 are there Ec. exsilium, In dorsal fin. the body cavities bi- as of living invertebrates such the but 4 only of vexillum origin anterior to the valves (e.g., En. dub ius), holothurians (e.g., En. vertebrae for En. vermicularis En. dub ius. Larvae of vermicularis), elongate are Carapids asteroids. and and to unknown. The number of vertebrae anterior are fin anal the compressed, with laterally longer rays than dorsal for origin fin (11-13) En. dubius than is smaller fin and fin The rays. origin dorsal opposing the anal for vermicularis En. (16-19). dorsal of fin The number beneath the are far forward, usually anus the pectoral the anterior rays to 30 for 31st vertebra probably is < fin is rarely absent (e.g., which En. vermicularis). The dubius. En. for more or vermicularis En. 35 and supramaxillary is absent as are scales. Vertebrae Dentition is useful to separate genera of carapids defined number about 85-145. As is by Cohen and and (Markle have Ec. 1990). Olney larvae exsilium or Carapinae subfamily the only (1978), Nielsen pectoral than have been reported in more rays fin pearlfishes are represented within the study CalCOFI at metamorphosis 1-4 may adults; resorbed be rays area. (Table Carapidae 1; Markle 1979; Markle and Olney 1990). Olney and oviparous; the few known eggs are Carapids are and ellipsoidal, pelagic, globule oil a single possess necessi- carapid of morphology unique The larvae al. et (Gordon may 1984). developmental Early stages fish larval standard the from a change tates description contained in a mucilaginous raft. Carapid larvae are be format. head Since the length damaged, easily is tail relatively well known and have proven to be sys- of st 31 reference used been have as vertebra the and Markle (Olney and Markle 1979; and tematic value length body than rather comparison for points (Markle Olney distinct two pass Pearlfishes 1980). through Olney and 1980; anal use We 1990). Olney and Markle larval stages. In the planktonic vexillifer stage, larvae larval developmental stages. fin separate to formation is a long, which vexillum a unique possess thread-like interval to refers (PrAF) formation" fin "Preanal the the to anterior process tenuis or second The fin. dorsal prior to initiation of fin formation and "anal fin anal is stage the which in benthic the is stage vexillum the body initial- gone, the head is relatively small, and refers or developing with formation" (AF) larvae to fully fins. anal formed 1981). ly lengthens, then subsequently shortens (Trott 532

546 The following on detailed descriptions are based time. Encheliophis came from larval The for the first examinations mm of mm 1 yolk-sac (3.5 0.4 TL, larva IA off samples TIC were data Meristic Panama. HL), mm 0.6-1.3 TL, mm (4.8-13.8 8 PrAF HL), and the literature (Arnold 1956; from obtained Rosenblatt of 10 AF larvae (25.6-97.3 mm TL, 1.8-4.5 mm HL) 1961; Trott 1970, and 1990) Olney and Markle 1981; (3.1-5.4 Echiodon TL, mm larvae 2 PrAF exsilium; study. this counts from during information Ecological 1 AF larva 0.5-1.0 mm HL) of Encheliophis sp.; and 1981), (1970, obtained Trott (1956), Arnold from was Gordon and Markle and (1984), al. et Encheliophis mm Olney (60 ius. dub (1990). of HL) mm 2.6 TL, described are these of taxa Yolk-sac PrAF larvae and Carapidae 1. Meristic characters for adults and larvae of carapid Table in the CalCOFI area. Counts of dorsal fm rays species are to the 31st vertebra (D30), anal fm rays to the 31st vertebra (A to ), vertebrae to the dorsal fin origin (DFO), anal rays 30 the fin origin (AFO), and vertebrae to the vexillum origin dorsal (VVO). Species DFO AFO WO PI D30 A30 Adults 19-21 38-42 10-11 47-50 8-10 exsilium 0 Echiodon 47-54 35-43 16-20 ius 11-13 17-22 dub Encheliophis 0 16-19 0 19-27 En. 36-43 17-24 0 vermicularis Larvae 21-25 39-42 50-51 10-11 9-11 exsilium Echiodon 8-11 20 37 48-52 12 18 Encheliophis dub ius 4 533

547 CARAPIDAE Echiodon exsilium MERISTICS EARLY LIFE mSTORY DESCRIPTION Range EGGS Mode Shell diam.: Vertebrae: Yolk: > to 130 No.ofOG: tapers Total Diam.ofOG: uncountable filament Shell surface: 21-25 Precaudal Pigment: Caudal Diagnostic .features: Fins: LARVAE spines Dorsal 0 38-42 TL mm Hatching length: <3.5 rays Dorsal 14 mm TL ca. formation fin Anal length: (DJo) 31st vertebra to Transformation length: stage Vexillifer to TL; 130 mm smallest ca. Anal 0 spines ca. adults 47-50 TL 85 mm rays Anal sequence: D & A, PI & development Fin C (A30) vertebra 31st to 0 Pelvic anterior eye; to posterior jaw; lower Yolk-sao-On PI to Pigmentation: 19-21 near Pectoral & vexillum bud; scattered on yolk; 1 ventrally midtail; dorsally end at over (prAF)--On formation fin region; tail. of Preanal gular Caudal: Principal brain bladder; internally anterior & ca. to vexillum to & gas posterior ca. by of 33% ca. posterior in midlaterally primarily tail; of tail; 66% Procurrent: 0 TL, only laterally on tail; by ca. 10.5 mm, only near end of 8 Upper mm & ca. over mm, 14 cranium (AF)-At tail; Analfin formation internal Lower 0 brain; on ventrally Gill by rakers: on peritoneum. dorsoposteriorly mm, 40 vertebrae Diagnostic anterior 8-11 features: to vexillum origin; 9-11 0 Upper rudiments 3+6 A rays rays vertebra; D origin; D to Lower 31st to anterior 39-42 prevexillum length> 7 1st Dray; anterior vexillum to 1.6xHL; ca. just Branchiostegals lower have of region, & tip on on pigment gular larvae PrAF jaw, larvae gut internally on snout; AF & have pigment ventrally on LIFE mSTORY stage. peritoneum on later in dorsoposteriorly Sur to Panama, including the Baja from Range: California Pacific Eastern MORPHOMETRICS & mean in %) (range Gulf Galapagos the of California & Islands on than Habitat: Free-living adults collected deeper outer shelf, usually Y-S PrAF AF Tenuis Juv 64 bottoms sand m over muddy 133-169 Sn-A/HL 130-149 168 146 138 Spawning season: in occur Larvae CalCOFI the thorughout collections year BDIHL 71-82 68-90 55 78 77 specialized Oviparous; vexillifer planktonic stage larva, ELH pattern: HUBL 10-13 4-8 stage larva tenuis benthic 13 12 6 45-59 HWIHL 34-51 LITERATURE 53 59 41 SnLIHL 20-29 20-33 Markle & Olney 1990 14 25 26 Olney 1979 Markle & EDIHL 33-40 24-34 1970 Trott 36 36 29 (Illustrator) ILLUSTRATIONS ORIGINAL 16-29 16-25 PIUBL 18 23 21 Vona) T. (M. Yolk-sac 3.5 mm larva, 169-194 PrVLIHL 167-197 vexillifer Vona) 7.3 (M. larva, PrAF T. mm 179 190 183 Vona) AF vexillifer larva, 25.3 mm (M. T. 534

548 Echiodon exsilium _- ... ~------ .. - ... - - ... ... .. -~"'"-~ -~~ 3.5 mm 7.3 mm 25.3 mm Figure Carapidae 1. Yolk-sac larva, 3.5 mm (CalCOFI 7202, station 137.35); PrAF vexillifer larva, 7.3 mm (IATTC 90014, station MSB mm #1); vexillifer larva, 25.3 AF (CalCOFI 5503, station 157.10). 535

549 CARAPIDAE Encheliophis dubius* EARLY LIFE HISTORY DESCRIPTION MERISTICS Mode EGGS Range Shell !liam.: Vertebrae: Yolk: Total No.ofOG: Diam.ofOG: 19-21 Shell surface: Precalldal Pigment: Caudal Diagnostic features: Fins: Dorsal spines 0 35-43 LARVAE rays Dorsal Hatching mm <3 length: .) (D vertebra 31st to 3 < ca. 60 Anal fin formation: 0 mm Alia! spines 47-54 mm small as 57 length: TL Transformation adults as Anal rays D sequence: development Fin , , & P A, C 31st vertebra (A .) to 1 Pelvic Preanal Pigmentation: 0 fin formation-By ca. 3 mm TL, over brain eyes, anteriodorsally gut, 16-20 between on under brain posterior to eyes, & Pectoral by 2 ventrally near end of tail; on ca. 5.5 mm, & vexillum, isthmus, Caudal: anterior (minute) Analfinformation-l tail. on ventrally A of base to Principal Protun-ellt: rays, a few specks dorsally near end vexillum, on one of speck tail, & midbrain. fore on dorsally o Upper o ca. origin; Lower 18 A rays to Diagnostic features: 4 vertebrae to vexillum 37 ca. rays to 3lstvertebra; prevexillum lAx D < ca. length D origin; rakers: Gill o maxilla free & movable; Upper of anteriad vexillum well HL; first D ray; o & fine jaw row region; gular along Lower lower from absent pigmentation 6-7 A. of Branchiostegals base LIFE HISTORY MORPHOMETRICS (range & %) mean in Y-S PrAF Gulf Pacific from but widespread, be eastern in May confirmed AF Range: Tenuis luv California Hawaii from & Colombia to of Sn-AlHL 133-154 144 144 Habitat: onward stage Molluscan-inhabiting from tenuis BDIHL 78-92 larvae Early stage collected July February & in from Spawning season: 85 102 Panama HLlBL 16-19 17 4 stage tenuis Oviparous; stage, vexillifer pelagic benthic pattern: ELH 39-67 HWIHL 53 40 LITERATURE 17-28 SnL/HL 22 29 & 1990 Markle Olney 32-50x ED/HLt (as jordan i) 1981 1970, Trott Encheliophis 24-35 ORIGINAL ILLUSTRATIONS (Illustrator) 41x30 27 16-21 LIHL P 1 PrAF larvae, 504 mm mm, 3.1 (R. C. Walker) 18 8 C. AF larva, 60.0 mm (R. Walker) LIHL P 0 2 ° ° PrVLIHL 122-125 since larvae of the * PrAF larvae are attributable to Encheliophis sp., 123 135 sympatric congener En. unknown. are vermicularis PrAF in (horizontally) elongate moderately Eye t axis horizontal larvae; axis given first, vertical second. 536

550 dub Encheliophis ius Pacific pearlfish mm 3.1 mm 5.4 '/, ' 60.0 mm Figure Carapidae 2. Encheliophis sp. PrAF larvae, 3.1 mm (IATTC 91006, station T8 0-5), 5.4 mm (IATTC 90022, station AAB 1 #2); mm Encheliophis ius AF larva, 60.0 dub (IATTC 90021, station ASBTlO). 537

551 BYTHITIDAE: Brotulas AMBROSE D. A. late BL) mm 3.5 (ca. embryos intraovarian stage from bythitid genera recognized least at 23 Cohen (1986) similar which have trophotaenia to those described by at with least which of half species, unde- 200 are and for (1975) Cohen Wourmes O/igopus longhursti. the These were fishes formerly included in scribed. mm C. The smallest rubrirostris available (7.3 BL) Brotulidae. three in the occur species bythitid At least already has visceral the in posteriorly looped gut the area study CalCOFI Bythitidae (Table 1). cavity. The smallest BL) 8 mm B. marginata (ca. is marginata Brosmophycis are rubrirostris Cataetyx and eyes, with larva a yolk-sac well-formed still pigmented commonly north-temperate species whose larvae occur mouth, and straight gut. Its centrally positioned gut CalCOFI in the Baja central to south area study B. marginata does not until 9-10 mm BL. form loop California, Mexico. O/igopus diagrammus prefers notochord and C. rubrirostris have a prolonged flexion Island, water southward from warmer San Clemente In transformation. until persists that stage transition the not have California; its larvae in identified been stage, head has marginata B. a more and rounded ichthyoplankton CalCOFI collections. and snout, C. rubrirostris and has a deeper body smaller eyes those relatively than the juveniles or of are body in similar to the shape bythitids Adult Pigmentation is more or less continuous adults. along anterior most live-bearers, are but cusk-eels the have diagrammus 0. of ventrum and dorsum the embryos, and intromit- on the snout, lower males nostril have an and tail the mostly posteriorly on gut 1978). Sizes range from organ tent (Cohen and Nielsen rubrirostris, C. of and intertidal less than 5 cm for some cryptic freshwa- the and of margins body B. along patches alternating in longer to ter forms than cm 70 for some deep-sea marginata. benthopelagic species. O/igopus Cataetyx and have a pelvic single thoracic fin ray, based as well as dorsal and following on detailed are The descriptions examination (Table species each of 10-28 of specimens Brosmophycis fin. caudal anal fins contiguous with the rays has are two thoracic pelvic joined which fin nearly of C. rubrirostris and Bythitidae 2). A larval series early stages larval that fin a caudal and tip the at 0. is separate from the and marginata B. diagrammus of described data Meristic were time. are first for the and fms. dorsal anal obtained 1957; (Best literature from 1964a; Cohen small the Gordon et al. (1984) summarized amount Miller and Grinols and Greenfield 1966; Follett 1970; Cohen 1972; Lea Matarese 1978; Nielsen et and larval on All available al. the information of bythitids. preanal have area study in larvae known lengths our study. this during made counts from and 1989) Ecolog- about with 40-50% of body length and are born a from information came et al. Eschmeyer ical primarily and (1983) (1991). Gibbs only known are young diagrammus 0. gut. straight 538

552 Table Bythitidae characters for the bythitid fishes in the CalCOFI area. A minute, hidden pelvic spine is not 1. Meristic the the The rudimentary gill rakers were excluded from counts. counts included in Cataetyx rubrirostris and O/igopus of diagrammus. Fin rays Vertebrae Total PrCV Species A D CV Gill rakers C BrR PI P2 63-65 16-17 47--49 20-26 98-110 72-81 Brosmophycis 2 8-9+8 marginata 15-16 4-5+ 7 60-63 14-15 46--49 100-114 4-5+4 25-26 76-86 0+3 rubrirostris Cataetyx 8 49-53 24-29 75-91 37-41 3-5+4-5 95-115 12 0-2+3 O/igopus diagrammus 8 Bythitidae 2. Number of specimens (above) and size range (in mm, below) used in preparation of the bythitid species Table descriptions. (SIO yolk-sac larvae were removed from a pregnant female diagrammus 53-169, 184 mm). O/igopus Preflexion Transfonnation Yolk-sac Flexion Juvenile Species 10 8 2 5 Brosmophycis marginata 3 8.5-16.7 16.5-35.6 7.9-8.7 37.5-41.0 81.2-125.0 7 6 0 10 rubrirostris Cataetyx 5 7.3-lO.3 12.5-30.6 24.5-57.8 72.4-103.8 0 0 0 5 O/igopus diagrammus 5 3.4-3.6 31.2-60.0 539

553 BYTHITIDAE Brosmophycis marginata MERISTICS DESCRIPTION EARLY LIFE HISTORY Range Mode EGGS Shell diam.: Vertebrae: Yolk: 64 Total No.ofOG: 63-65 Diam.ofOG: 16 Shell surface: 16-17 Precaudal 48 Pigment: 47-49 Caudal Diagnostic features: Fins: spines Dorsal 101 98-110 LARVAE Dorsal rays < 75 72-81 Length at birth: ca. 8 mm Anal spines Flexion length: 17-36 ca. mm rays Anal 2 81 mm 2 ca. Transformation 37 to < length: Pelvic Pectoral D Fin sequence: development P Ph & 20-26 A, C, 2 Caudal: Pigmentation: lip; on larva; preflexion mm 9.8 to Yolk-sac-8imilar 8+8 8-9+8 cleithrum; gas bladder; above 4 more 3 patches laterally on gut; or Principal heavy & tail ventrally; along margins alternating patches dorsally less Procur