Guide to the Coastal Marine Fishes of California
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STATE OF CALIFORNIA THE RESOURCES AGENCY DEPARTMENT OF FISH AND GAME FISH BULLETIN 157 GUIDE TO THE COASTAL MARINE FISHES OF CALIFORNIA by DANIEL J. MILLER and ROBERT N. LEA Marine Resources Region 1972 ABSTRACT This is a comprehensive identification guide encompassing all shallow marine fishes within California waters. Geographic range limits, maximum size, depth range, a brief color description, and some meristic counts including, if available: fin ray counts, lateral line pores, lateral line scales, gill rakers, and vertebrae are given. Body proportions and shapes are used in the keys and a state- ment concerning the rarity or commonness in California is given for each species. In all, 554 species are described. Three of these have not been re- corded or confirmed as occurring in California waters but are included since they are apt to appear. The remainder have been recorded as occurring in an area between the Mexican and Oregon borders and offshore to at least 50 miles. Five of California species as yet have not been named or described, and ichthyologists studying these new forms have given information on identification to enable inclusion here. A dichotomous key to 144 families includes an outline figure of a repre- sentative for all but two families. Keys are presented for all larger families, and diagnostic features are pointed out on most of the figures. Illustrations are presented for all but eight species. Of the 554 species, 439 are found primarily in depths less than 400 ft., 48 are meso- or bathypelagic species, and 67 are deepwater bottom dwelling forms rarely taken in less than 400 ft. depth. The deepwater forms included are those taken in commercial trawling gear or that occasionally occur near the surface. An illustrated glossary is included to facilitate use of the identification keys and species descriptions. A comments section presents in detail reasons for certain taxonomic choices and to acknowledge personal communications. Original data presented include a ventral sensory pore pattern key for the skates, 170 geographic range limit extensions, and several depth range and maximum size records. Many of the family keys have been revised to incorporate recent taxonomic changes and to clarify previ- ously ambiguous terminology. TABLE OF CONTENTS ACKNOWLEDGMENTS 2 INTRODUCTION 3 HOW TO USE THE GUIDE The Keys 4 Measurements and Counts 4 Geographic Ranges 5 Depth Ranges 5 GLOSSARY AND INDEX TO ILLUSTRATED GLOSSARY_ 6 KEY TO THE FAMILIES 13 SPECIES PRESENTATION 32 APPENDIX. Taxonomic Comments and Personal Communica- tions Citations 211 LITERATURE CITED 216 GENERAL REFERENCES 222 INDEX TO COMMON NAMES 223 INDEX TO SCIENTIFIC NAMES 229 a GUIDE TO THE COASTAL MARINE ACKNOWLEDGMENTS This guide has been made possible through the works and studies of hundreds of fishery researchers and taxonomists, past and present. We cannot thank those of the past whose immortality lies in libraries, but we can credit most of those who have personally aided us. Unfortu- nately, space does not permit us to fully credit all that has been received from each contributor or to mention all who have helped us. First, we extend credit to fellow Department colleagues. The meticu- lously recorded data on range limits, depth, and sizes of California marine fishes offered to us by John Fitch has been of primary impor- tance. John also gave taxonomic advice, offered information on identifi- cation, and stimulated us to conduct the museum search for geographic and depth range limit extensions. Dan Gotshall and John Geibel had originally planned to collaborate on this publication but their heavy work loads precluded full collaboration. Dan Gotshall was able to assist in preparing the sections on the eelpouts, poachers, and pricklebacks; and John Geibel assisted in preparing the section on scombrids and printing photographs of some of the fish drawings. Dick Nitsos and Dick Parrish supplied data on depth of bottomfishes, and Paul Gregory and Jack Schott supplied pictures on which some of the fish drawings are based. Dan Collier and his wife Cynthia made photographic prints of most of the fish drawings. Jim Houk spent many hours in the muse- ums searching bottles for pertinent data, and helped in the literature search. Bill Craig and Dick Burge supplied information on ranges, and Pat Powell and the library staff aided in supplying literature and editing the reference section. Nancy Durell and Margaret Hughes typed the manuscript. Herb Frey edited the entire manuscript, and Harold Orcutt offered encouragement and suggestions to undertake this project. Many fish researchers and taxonomists have given us aid. Geographi- cally, from north to south, We wish to extend our thanks to the follow- ing for their suggestions and information so generously offered: Jay C. Quast, National Marine Fish. Service, Auke Bay, Alaska; Sigmund J. Westrheim, Fisheries Research Board of Canada, Nanaimo, B.C.; John L. Hart, Fisheries Research Board of Canada, St. Andrews, N.B.; Alex E. Peden, British Columbia Provincial Museum, Victoria, B.C.; William G. Pearcy, Oregon State University, Corvallis, Ore.; W. I. Follett and Earl S. Herald, California Academy of Sciences, San Fran- cisco; Leonard J. V. Compagno, Systematic Zoology Dept., Stanford University; John S. Stephens, Occidental Colleges Los Angeles; Boyd W. Walker, University of California at Los Angeles; Robert J. Laven- berg and Camm Swift, Los Angeles County Museum of Natural His- tory; Shelly R. Johnson, University of Southern California; Elbert H. Ahlstrom, National Marine Fisheries Service, La Jolla; Carl L. Hubbs, Laura Hubbs, Richard H. Rosenblatt, Robert L. Wisner, and Layton P. Taylor, Jr., Scripps Institution of Oceanography; and Lo-Chai Chen, California State University at San Diego. Museum staff members and curators were most cooperative and help- ful. Special thanks must be extended to the sage of reference, grammar, and literature, Lillian J. Dempster, California Academy of Sciences, for FISHES OF CALIFORNIA 3 her enthusiastic search for descriptions and references. The entire staff of the California Academy of Sciences Ichthyology Department, espe- cially Pearl M. Sonoda and William N. Eschmeyer, made it possible to conduct primary museum work in development and testing identifica- tion keys. John Bleck, University of California at Los Angeles, and Joe Copp and Donald Dockins, Scripps Institution of Oceanography, gave us much aid in the museum search for range and size extensions. To all the above and to those not listed, we offer our sincere gratitude for all the help you have given us. Daniel J. Miller Robert N. Lea INTRODUCTION All the shallow water inshore marine fishes recorded as occurring in California waters between the Oregon and Mexican borders of Cali- fornia are included in this bulletin, as well as most marine fishes intro- duced into the Salton Sea. In all, 144 families are represented by the 554 species described. Three of these species have not been recorded from California but are included to enable identification since their presence in California waters has not been confirmed. Five California species are undescribed, two in family Stichaeidae, and one each in families Scorpaenidae, Agonidae, and Gerreidae. Of the 554 species, 439 most frequently are found in waters shallower than 400 ft., 48 are pri- marily meso- or bathypelagic (about V4 of all California meso- and bathypelagic forms), and 67 bottom dwelling forms rarely taken above 400 ft. This guide is primarily for identification purposes, using a brief de- scription of color, meristic counts, and body proportions and structures to enable keying out a fish to family and to species within a family. The following additional data are given, if available: geographic rarige, maximum size, depth range, a comment on rarity, and vertebral counts. A fish is considered rare if 20 or less ever have been taken in California; uncommon if the fish is seldom seen either because of scarcity or be- cause it is unavailable even though actually present in large numbers; and common if it is easily available and appearing frequently in sport or commercial catches or in tide pool, mud flat, or kelp canopy collec- tions. Illustrations of fish primarily were based on literature, but the origins were not cited because in most cases the published figure was changed. In most cases drawings were compared either with fresh or preserved material, if available. The fish drawings, family key figures, and skate ventral pore patterns were prepared by the senior author. Drawings of some diagnostic characters and all the shark's teeth were prepared by Cathy Short, California Department of Fish and Game. 4 GUIDE TO THE COASTAL MARINE HOW TO USE THE GUIDE The Keys This guide is an accumulation of artificial diagnostic keys based on external features of shape, proportion, color, and numbers of fin rays, gill rakers, lateral line pores, and scales. Internal characters have been avoided except for gill rakers and teeth which can be examined without dissection. The term "artificial" refers to a key based on characters not necessarily indicative of taxonomic or phyloienetic order. Many taxo- nomic characters are skeletal and internal, and depict evolutionary trends from primitive to more specialized or advanced forms. Even though the keys are not based necessarily on evolutionary criteria, the sequence of families is generally in phylogenetic order; i.e. from the primitive hagfishes to highly specialized molas. The sequence we fol- lowed was that of Greenwood et al (1966); however, in several in- stances closely related families containing only one or several species have been grouped on a single page without adherence to this sequence. The keys work on the principle of an either/or choice using "a" and "b" alternatives. If a fish does not fit the description of "a" go on to "b", etc. In each case where there is a jump to another section of the key, there is a citation of the section from which you were referred, facilitating backtracking through the key.