415 & a .r* X ^ATES 0* * NOAA Technical Report NMFS Circular 415 A Basis for Classifying Western Atlantic Sciaenidae (Teleostei: Perciformes) Labbish Ning Chao September 1978 -*- ° U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service NOAA TECHNICAL REPORTS National Marine Fisheries Service, Circulars The major responsibilities of the National Marine Fisheries Service (NMFS) are to monitor and assess the abundance and geographic distribution of fishery resources, to understand and predict fluctuations in the quantity and distribution of these resources, and to establish levels tor optimum use of the resources. NMFS is also charged with the development and implementation of policies for managing national fishing grounds, development and enforcement of domestic fisheries regulations, surveillance of foreign fishing off United States coastal waters, and the development and enforcement of international fishery agreements and policies. NMFS also assists the fishing industry through marketing service and economic analysis programs, and mortgage insurance and vessel construction subsidies. It collects, analyzes, and publishes statistics on various phases of the industrv. The NOAA Technical Report NMFS Circular series continues a series that has been in existence since 1941. The Circulars are technical publications of general interest intended to aid conservation and management. Publications that review in considerable detail and at a high technical level certain broad areas of research appear in this series. Technical papers originating in economics studies and from management in- vestigations appear in the Circular series. NOAA Technical Report NMFS Circulars are available free in limited numbers to governmental agencies, both Federal and State. They are also available in exchange for other scientific and technical publications in the marine sciences. Individual copies may be obtained (unless otherwise noted! from D825, Technical Information Division, Environmental Science Information Center, NOAA, Washington, D.C. 20235. Re- cent Circulars are: 365. Processing EASTROPAC STD data and the construction of ver- tical temperature and salinity sections by computer. By Forrest R. Miller and Kenneth A. Bliss. February 1972, iv + 17 p., 8 figs., 3 app. figs. For sale by the Superintendent of Documents. U.S. Government Printing Of- fice. Washington. D.C. 20402. itiii Key to field identification of anadromous juvenile salmonids in the Pacific Northwest. By Robert J. MacConnell and George R. Snyder. •January 1972. iv + 6 p., 4 figs. For sale by the Superintendent of Documents. U.S. Government Printing Office. Washington, D.C. 20402. .177. Fishery publications, calendar year 1970: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. December 1972, iv + 34 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Office. Washington. D.C. 20402. 378. Marine flora and fauna of the northeastern United States. Protozoa: Ciliophora. By Arthur C. Borror. September 1973, iii + 62 p., 5 tigs. For sale bv the Superintendent of Documents, U.S. Government Printing Office. Washington, D.C. 20402. 367. Engineering economic model for fish protein concentration processes. By K. K. Almenas, L. C. Durilla, R. C. Ernst, J. W. Gentry, M. B. Hale, and J. M. Marchello. October 1972, iii + 175 p., 6 figs., 6 tables. For sale bv the Superintendent of Documents, VJ$. Government Printing Office, Washington, D.C. 20402. 368. Cooperative Gulf of Mexico estuarine inventory and study. Florida: Phase I. area description. By J. Kneeland McNulty, William N. Lindall. Jr., and James E. Sykes. November 1972, vii + 126 p., 46 figs., 62 tables. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington, D.C. 20402. 369. Field guide to the anglefishes (Pomacanthidae) in the western Atlantic. By Henry A Feddern. November 1972, iii + 10 p., 17 figs. For sale by the Superintendent of Documents, U.S. Government Printing Of- fice. Washington. D.C. 20402. 370. Collecting and processing data on fish eggs and larvae in the California Current region. By David Kramer, Mary J. Kalin, Elizabeth G, Stevens, James R. Thrailkill, and James R. Zweifel. November 1972, iv 4- 38 p., 38 figs., 2 tables. For sale by the Superintendent of Documents. U.S. Government Printing Office, Washington, D.C. 20402. 371. Ocean fishery management: Discussion and research. By Adam A. Sokoloski (editor). (17 papers, 24 authors.) April 1973, vi + 173 p., 38 figs . 12 tables, 7 app. tables. 379. Fisherv publications, calendar year 1969: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. April 1973, iv + 31 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Office. Washington. D.C. 20402. 380. Fisherv publications, calendar year 1968: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. May 1973, iv + 24 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Of- fice. Washington. D.C. 20402. 381 . Fisherv publications, calendar year 1967: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. July 1973, iv + 22 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington. D.C. 20402. 382 Fishery publications, calendar year 1966: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. July 1973, iv + 19 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Of- fice. Washington. D.C. 20402. 383. Fishery publications, calendar year 1965: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. July 1973, iv -I- 12 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Office, Washington. D.C. 20402. '.72. Fishery publications, calendar year 1971: Lists and indexes. By Thomas A. Manar. October 1972, iv + 24 p., 1 fcg. For sale by the Superintendent of Documents, U.F. Government Printing Office, Washington. D.C. 20402. 384. Marine flora and fauna of the northeastern United States. Higher plants of the marine fringe. By Edwin T. Moul. September 1973, iii + 60 p.. 109 figs. For sale by the Superintendent of Documents, U.S. Govern- ment Printing Office, Washington, D.C. 20402. 374 Marine flora and fauna of the northeastern United States. Annelida: OKgochaeta. By David G. Cook and Ralph O. Brinkhurst. May 1973. iii + 23 p.. 82 figs. For sale by the Superintendent of Documents, I' S. Government Printing Office. Washington, D.C. 20402. '.75. New Polvchaeta from Beaufort, with a key to all species recorded from North Carolina. By John H. Day. July 1973'. xiii + 140 p., 18 figs., 1 table. For sale bv the Superintendent of Documents. U.S. Government Printing Office. Washington. D.C. 20402. 385. Fishery publications, calendar year 1972: Lists and indexes. By Lee ('. Thorson and Mary Ellen Engett. November 1973, iv + 23 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Of- fice. Washington, D.C. 20402. 386. Marine flora and fauna ot the northeastern United States. Pyc- nogonida By Lawrence R. McOloskey. September 1973, iii + 12 p., 1 fig. For sale by the Superintendent of Documents. U.S. Government Printing Office. Washington, D.C. 20402. 176 Bottom-water temperatures on the continental shelf, Nova Scotia to New Jersey. Bv John B. Colton. Jr. and Ruth R Stoddard. June 1973, - '" p.. 15 tigs.. 12 app. tables. For sale by the Superintendent of Documents. U.S. Government Printing Office, Washington, D.C. 20402 387. Marine flora and fauna of the northeastern United States. Crustacea: Stomatopoda. By Raymond B. Manning. February 1974, iii + (i p.. 10 figs. For sale by the Superintendent of Documents, U.S. Govern- ment Printing Office. Washington, D.C. 20402. Continued <>n inside hack cover o u ,m ms r , f mm %. Vll \ T ( )V l NOAA Technical Report Circular- 415 A Basis for Classifying Western Atlantic Sciaenidae (Teleostei: Perciformes) Labbish Ning Chao September 1978 o 0. Q 60 U.S. DEPARTMENT OF COMMERCE Juanita M. Kreps, Secretary National Oceanic and Atmospheric Administration Richard A. Frank, Administrator National Marine Fisheries Service The National Marine Fisheries Service (NMFS) does not approve, rec- ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales pro- motion which would indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. CONTENTS Introduction 1 Materials 2 Methods . ., 2 A brief history of the study of western Atlantic Sciaenidae 3 Description and relationships 6 Groupings based on swim bladder morphology 6 Morphological patterns of swim bladders 8 Single-chambered swim bladder group 8 Sciaena pattern 8 Nebris pattern 9 Micropoganias pattern 9 Poganias pattern 9 Sciaenops pattern 10 Cynoscion pattern 10 Lonchurus pattern 10 Two-chambered swim bladder group 10 Stellifer pattern 10 Groupings of swim bladder patterns 11 Specific characters 13 Groupings based on otolith morphology 14 Morphological patterns of otoliths 14 Only sagitta enlarged 14 Sciaena pattern 14 Pogonias pattern 14 Larimus pattern 14 Nebris pattern 15 Cynoscion pattern 15 Menticirrhus pattern 15 Lonchurus pattern 18 Other forms of sagittae 18 Isopisthus pattern 18 Macrodon pattern 19 Micropogonias pattern 19 Both sagitta and lapillus enlarged 20 Stellifer pattern 20 Groupings of otolith patterns 21 Specific characters 21 Groupings based on external morphology 22 Patterns of pores and barbel systems 22 Structural patterns of snout pores 22 Two (or no) marginal pores 22 Five marginal pores 22 Five marginal and three upper pores 23 Five (or more) marginal and five (or more) upper pores 23 Structural patterns of mental pores and barbels 24 No pores 24 Four pores 24 Five pores 24 Six pores 25 Four (or five) pores and one barbel 25 Four pores and two barbels 25 Five pores and many barbels 25 Generic groupings based on external morphology 25 Specific characters ." 26 A synopsis of the suprageneric groups, genera, and species of western Atlantic Sciaenidae 27 Diagnoses and primary synonymy of the suprageneric groups, genera, and species 27 iii Sciaena group 27 I'mbrina group 29 ^fenticirrhus group 30 Larimus group 31 Xebris group 31 Micropogonias group 32 Seiaenops group 33 Pogonias group 34 Cynoscion group 34 Lonchurus group 37 Stellifer group 38 Freshwater Sciaenidae of America 41 Phylogenetic relationships of the genera and suprageneric groups 44 Relationships of the genera within the suprageneric groups 44 Relationships of the suprageneric groups 45 Key to the genera and species of western Atlantic Sciaenidae (with freshwater American genera) 47 Key to the species of Bairdiella 50 Cynoscion 51 Equetus 52 Larimus 52 Menticirrhus 53 Micropogonias 53 Ophioscion 53 Paralonchurus 53 Pareques 54 Sciaena 54 Stellifer 54 Umbrina 56 Acknowledgments 56 Literature cited 57 Systematic index 62 Figures 1. Terminology of pore systems on head of western Atlantic Sciaenidae 3 2. Terminology of otoliths 4 3. Inner ears of Sciaenidae 5 4. Proposed phylogentic relationships of genera and suprageneric groups of western Atlantic Sciaenidae . 6 5. Swim bladders of the Sciaena pattern 8 6. Swim bladders of two species of Pachyurus 8 7. Swim bladders of Menticirrhus saxatilus 8 8. Development of swim bladder in Nebris microps 9 9. Variation and development of swim bladders of Micropogonias 9 10. Ontogenetic development of swim bladders in Pogonias cromis 10 11. Ontogenetic development of swim bladder in Seiaenops ocellata 11 12. Swim bladders of the Cynoscion pattern 11 13. Swim bladders of species of Cynoscion 11 14. Relative size and development of swim bladders with the Lonchurus pattern 12 15. Swim bladders of the Stellifer pattern 12 16. Diversity of the swim bladder diverticula in the genus Stellifer 12 17. Groupings of western Atlantic Sciaenidae according to swim bladder patterns 13 18. Inner surface and lateral view of sagittae of the Sciaena pattern 15 19. Sagittae of the Pogonias pattern 16 20. Sagittae of the Larimus pattern 16 21. Sagittae in Nebris microps 17 22. Sagittae of the Cynoscion and the Menticirrhus patterns 17 23. Specific and ontogentic variations of sagittae in species of Cynoscion 17 24. Sagittae of the Lonchurus pattern 18 iv 25. Sagittae of the Isopisthus and the Macrodon patterns 18 26. Variation, possibly ontogentic, in sagittae of the Micropogonias pattern 19 27. Inner surface of sagittae and lapilli of the Stellifer pattern 20 28. Inner surface of sagittae and lapilli in species of Stellifer 20 29. Groupings of western Atlantic Sciaenidae according to otolith patterns 21 30. Groupings of western Atlantic Sciaenidae according to external morphology 22 31. Snout pore patterns of western Atlantic Sciaenidae 23 32. Mental pore and barbel patterns on the lower jaw of western Atlantic Sciaenidae 23 33. Diversity in snout pores with five or more marginal pores 23 34. Diversity in mental pore arrangements on the lower jaw 24 35. Diversity in mental pore patterns on the lower jaw in the species of Stellifer 24 36. Western Atlantic Sciaenidae with one mental barbel on the lower jaw 24 37. Four typical mouth positions and dentitions of western Atlantic Sciaenidae 25 38. Three levels of mouth position in the species of Stellifer 26 39. Sagittae of Micropogonias 33 40. United lower pharyngeal teeth of Pogonias cromis and Aplodinotus grunniens 35 41. Portions of lateral line scales of Plagioscion surinamensis 43 Table 1. Summary of morphological characters of the suprageneric groups of western Atlantic Sciaenidae 7 Digitized by the Internet Archive in 2013 http://archive.org/details/basisforclassifyOOchao A Basis for Classifying Western Atlantic Sciaenidae (Teleostei: Perciformes) 1 LABBISH NING CHAO 2 ABSTRACT Sciaenidae of the western Atlantic consist of 21 genera and at least 57 species, and are placed in 1 1 suprageneric groups: Cynoscion, Larimus, Lonchurus, Menticirrhus, Micropogonias, Nebris, Pogonias, Sciaena, Sciaenops, Stellifer, and Umbrina groups. The phylogenetic relationships of all western Atlantic genera are assessed on the basis of swim bladder, otoliths (sagitta and lapillus), and external morphology. The Stellifer group differs from all other western Atlantic sciaenids in having a two-chambered swim bladder and an enlarged lapillus. Phylogenetic and ontogenetic trends of the swim bladder range from a simple carrot-shape, to a more complicated structure with anterior projec- tions and horns, to a very complicated lateral diverticula system. The sagitta is usually oval or elongate in shape. The thickness and the impression of the sulcus on the inner surface of the sagitta are diagnostic among genera. External morphology is adaptive, especially in relation to feeding habits and habitat, but a trend is evident in that closely related genera often have similar body shapes, mouth positions, and other external features. Species of the genus Stellifer are exceptions in having diverse mouth positions and feeding habits. The synopsis section of the paper includes a diagnosis, a primary synonymy, and lists of types of nominal species for each taxonomic category. Four genera and 22 nominal species of New World fresh- water sciaenids are also included. Nomenclatural changes in regard to the genus Micropogon (to Micropogonias) and for the species Bairdiella chrysura (to B. chrysoura) is given. A tested key to species and genera of all western Atlantic sciaenids is included. The approximate range of dis- tribution and some counts are listed under each species. This paper is designed to serve as a basis for further revision of western Atlantic sciaenids. INTRODUCTION Sciaenid fishes are characterized by their large otoliths and, with few exceptions, by enlarged lateral-line canals on the head and pores on the snout and lower jaw, and by the extension of the lateral line to the tip of the caudal fin. They are also characterized by a large and often com- plex swim bladder and by the presence of well-developed drumming muscles. This paper treats all 21 genera and 57 species of Sciaenidae currently recognized in the western Atlantic Ocean. In addition, limited infor- mation is also provided for 4 genera and 22 nominal species of sciaenid fishes present in freshwater river sys- tems of the New World. Current concepts of sciaenid classification are largely based on morphology of the swim bladder, otoliths (sagitta and lapillus), and snout (rostral) and mental (mandibular) pores and/or barbels (Chu et al. 1963; Trewavas 1962, 1964, 1977; Robins and Tabb 1965; Gil- bert 1966; Lowe 1966; Lai Mohan 1969; Chao and Miller 1975). Other characters, such as the position of the 'A portion of a dissertation submitted to the School of Marine Science, College of William and Mary, Williamsburg, Va., in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Marine Science, May 1976. Contribution No. 817, Virginia Institute of Marine Science, Gloucester Point, VA 23062, U.S.A. 'Ichthyology Unit, Vertebrate Zoology Division, National Museum of Natural Sciences, Ottawa, Ontario, Canada K1A 0M8; present address: Fundagao Universidade do Rio Grande Base Oceanografica Atlantica, Caixa Postal 474, 96.200 - Rio Grande - RS, Brasil. mouth, dentition, body form, size of the second anal spine, marginal serration of the preopercle, arrangement of lateral-line scales, color of peritoneal lining, and num- bers of gill rakers, fin ray counts, and vertebrae, are im- portant in distinguishing species and have been used by many authors to assign species to genera or to assess the generic relationships of western Atlantic Sciaenidae. However, the generic boundaries have not been well defined and many nomenclatural problems remain. The purpose of this paper is to clarify generic bound- aries, define suprageneric groups, and describe the evolutionary trends of western Atlantic Sciaenidae primarily on the basis of the morphology of the swim bladder, otolith, and snout and mental pore and/or bar- bel systems. To classify the taxa above the generic level, the term suprageneric group is used here. Trewavas (1962) grouped eastern Atlantic sciaenids into tribes. Chu et al. (1963) grouped scaenids of the Chinese coasts into sub- families. Lai Mohan (1969) placed Indian species into both subfamilies and tribes. The suprageneric groups used here are comparable to the tribes of Trewavas (1962, 1977) and the subfamilies of Chu et al. (1963), but avoid the taxonomic problems inherent in using formal tax- onomic categories. Like those studies, the present as- sessment is also a regional study. Variations in swim bladder, otolith, and external morphology are often found at different taxonomic levels. Therefore, a com- parative study of sciaenids from several geographic regions, especially the eastern Pacific area, is necessary before allocating the western Atlantic sciaenid genera to formal taxonomic categories (tribes or subfamilies). The commendable study of Indo-West-Pacific Sciaenidae by Trewavas (1977), especially in its assess- ment of diagnostic characters, evolutionary significance and classification of sciaenid fishes, is an important and valuable contribution to this family. Regretably, I was not able to incorporate the information in this publica- tion here because it appeared after the present study was in press. MATERIALS Type-material and other preserved specimens were examined from the following institutions: AMNH — American Museum of Natural History, New York, N.Y. ANSP — Academy of Natural Sciences, Philadelphia, Pa. BMNH— British Museum (Natural History), London, England. CAS — California Academy of Sciences, San Fran- cisco, Calif. FMNH —Field Museum of Natural History, Chicago, 111. LACM — Los Angeles County Museum of Natural His- tory, Los Angeles, Calif. MCZ — Museum of Comparative Zoology, Harvard University, Cambridge, Mass. MNHN— Museum National d'Histoire Naturelle, Paris, France. NHMV — Naturhistorisches Museum, Vienna, Austria. NMC —National Museum of Natural Sciences, Ot- tawa, Canada. RMNH — Rijksmuseum van Natuurlijke Historie, Lei- den, Netherlands. SOSC — Smithsonian Oceanographic Sorting Center, Washington, D.C. UF —Florida State Museum, University of Florida, Gainesville, Fla. UMML — Rosenstiel School of Marine and Atmospheric Sciences, Miami, Fla. USNM — United States National Museum, Washington, D.C. VIMS — Virginia Institute of Marine Science, Glouces- ter Point, Va. ZMA — Institute voor Taxonomische Zoologie, Univer- siteit van Amsterdam, Amsterdam, Nether- lands. ZMK — Zoological Museum, Copenhagen, Denmark. Freshly frozen and Formalin-preserved specimens were also obtained from various biological surveys along the Atlantic seaboard of the United States. These surveys were conducted by the Virginia Institute of Marine Science on various research vessels; by the National Marine Fisheries Service (U.S.A.) laboratories at Woods Hole, Mass. on RV Albatross IV and at Sandy Hook, N.J., on RV Atlantic Twin; and by the National Science Foundation's RV Eastward, Beaufort, N.C. Specimens examined from Gulf of Mexico and Atlantic coast of South America were collected mainly by NMFS research vessels, the MV Oregon I and Oregon II, from 1962 to 1975. Primary type-specimens examined during the study are indicated under the synonymy of each species by their museum catalogue numbers and length of individuals. In other cases, except specifically noted, the type-specimens were not located. Numerous ad- ditional specimens have been studied, most of which are deposited at USNM, VIMS, and NMC. METHODS Comparative morphologies of the swim bladder, otolith, and pore systems were studied. Other charac- ters important in the recognition of species and genera are the arrangement and size of lateral-line scales, ser- rations of the preopercular margin, size of the second anal fin spines, color of the branchial chamber and peritoneal linings, meristics, and morphometries. The standard methods of Hubbs and Lagler (1964) were used for all counts and measurements except for some modifi- cations described by Chao and Miller (1975). In addition, scales perforated by lateral-line tubes or pores were counted as lateral-line scales from the upper end of the gill slit to the end of the hypural plate. On those species lacking well-defined lateral-line tubules or pores, counts were made on the number of transverse scale rows im- mediately above the lateral line. Vertebral counts were determined from radiographs and from cleared and stained specimens (Taylor 1967). The first caudal vertebra was identified by the absence of pleural ribs on the haemal process and usually by a short haemal spine just behind the elongate proximal pterygiophore of the anal fin. The urostylar centrum was included in counts. Otoliths of most species were extracted from freshly caught specimens but some came from preserved specimens. Inner surfaces of the right sagitta and lapil- lus (in the Stellifer group) were illustrated. Ontogenetic changes of otoliths and swim bladder were studied in some species. All available type-specimens were examined to assure the identities of species and genera of sciaenids studied in this paper. A primary synonymy is listed for each species to clarify nomenclatural problems. Morphological terminology used here follows Chu et al. (1963), Trewavas (1962, 1964), and Chao and Miller (1975). A modification of the terms used for the snout pores was suggested by E. Trawavas of BMNH, and for otoliths by J. Fitch of the California Department of Fish and Game. Snout pores along the margin of the rostral fold are termed marginal (snout) pores, and pores in front or above them are termed upper (snout) pores (Fig. 1). The surface of the sagitta, with a "tadpole-shaped" sulcus is the inner surface and the obverse side is the out- er surface (Fig. 2) which are the in situ position of the sagitta (Fig. 3). Also, the lateral margins of the sagitta are termed dorsal and ventral margins (Frizzell and Dante 1965). The "head section" of the sulcus is called the "ostium" and the "tail section" is called the "cauda" (Stinton 1975; J. Fitch, pers. commun.). — UPPER PORES MARGINAL PORES ROSTRAL FOLD MENTAL BARBEL MENTAL PORES Figure 1. — Terminology of pore systems on head of western Atlantic Sciaenidae. Snout; upper pores (above the dashed line) and marginal pores (below the dashed line). Lower jaw; mental pores and barbels(s). A. ventral view of the mouth, Menticirrhus saxatilis. B. tip of the snout, Stellifer lanceolatus. A BRIEF HISTORY OF THE STUDY OF WESTERN ATLANTIC SCIAENIDAE Prior to the binominal system of Linnaeus, Catesby in his 1743 edition of "The natural history of Carolina, Florida and the Bahama Islands," illustrated "Perca marina" ( = Micropogonias undulatus), "THE CROKER" (plate 3), from Chesapeake Bay and "Al- burnus Americanus" ( = Menticirrhus americanus), "THE WHITING" (plate 12), from Charleston, S.C. Ed- wards (1751) in the appendix of "A natural history of birds" illustrated (plate 210) a "Ribband Fish" ( = Equetus lanceolatus) from the Caribbean Islands. Lin- naeus (1758) in his 10th edition of "Systema Naturae," Pisces Thoracici, recorded the genus Sciaena with five species, none of them from the western Atlantic. However, two species of western Atlantic Sciaenidae were named as Chaetodon lanceolata ( = Equetus lan- ceolatus) based on Edwards' plate and Cyprinus ameri- canus ( = Menticirrhus americanus) from "the whiting" of Catesby. In the 12th edition of "Systema Naturae," 1766, Linnaeus added Labrus chromis ( = Pogonias cromis), Perca alburnus )= Menticirrhus americanus), Perca punctata (= Bairdiella chrysoura), Perca ocellata ( = Sciaenops ocellata), and Perca undulata ( = Micro- pogonias undulatus), although he did not place them with the Sciaenidae from Europe. Bloch and Schneider (1801) in their "Systema Ich- thyologiae" reported six genera and nine species of western Atlantic Sciaenidae: Johnius regalis ( = Cynos- cion regalis), J. saxatilis (= Menticirrhus saxatilis), Sciaena chromis ( = Pogonias cromis), Lonchurus depressus (= Lonchurus lanceolatus), Lonchurus ancy- lodon (= Macrodon ancylodon), Eques americanus ( = Equetus lanceolatus), Eques punctatus, Grammistes ac- cuminatus ( = Pareques acuminatus), and Bodianus stel- lifer (= Stellifer stellifer). A year later (1802) Lacepede in volume three of "Histoire Naturelle des Poissons" recorded Dipterodon chrysourus ( = Bairdiella chry- soura), Pogonias fasciatus (= P. cromis), and Cheilodip- terus acoupa ( = Cynoscion acoupa) and in volume four (1803) he added Lutjanus cayennensis (= Cynoscion acoupa), Lutjanus triangulum (= Sciaenops ocellata), Centropomus alburnus (= Menticirrhus americanus), Sciaena croaker ( = Micropogonias undulatus), and Leiostomus zanthurus of the western Atlantic and also a freshwater sciaenid Perca fourcroi (= Pachypops four- croi) from Surinam. The boundaries of the species, genera, and family Sciaenidae were not well defined, until Cuvier. In the first edition of "Regne Animal" (1817), Cuvier defined limits to the application of some generic names (e.g., Umbrina and Sciaena). Also four new genera of western Atlantic Sciaenidae were described: the scientific names Stellifer, Umbrina, Otolithes, and Ancylodon were derived from Cuvier's vernacular names by Oken (1817). In the second edition, Cuvier (1829) added a new generic name Corvina. Cuvier (1830, in Cuvier and Valenciennes) in volume five of "Histoire Naturelle des Poissons," described 7 genera and 21 species of western Atlantic sciaenids. There were four new genera, i.e., Larimus, Nebris, Lepipterus, and Micropogon, and five freshwater nominal species, Corvina oscula ( = Aplodino- tus grunniens), C. furcroea (= Pachypops fourcroi), and Lepipterus francisci ( = Panchyurus francisci), were in- cluded. Cuvier also described and illustrated the swim bladders of six western Atlantic sciaenid fishes. Giinther (1860) recorded 12 genera and 45 species of Sciaenidae from the western Atlantic including 3 genera and 9 species from freshwater. He used the shape of the swim bladder as one of the diagnostic characters in some of his generic descriptions. Jordan and Eigenmann (1889) listed 22 genera and 60 species, including 4 genera and 11 species of freshwater sciaenids. Jordan and Evermann (1898) included only 21 genera and 50 species of Sciaenidae from the Atlantic in the "Fishes of North and Middle America," including 3 genera and 5 species from freshwater. The studies of Desmarest (1823), Agassiz (1829), Castelnau (1855), Steindachner (1863), Bleeker (1865, 1873), Poey (1881), Berg (1895), and more recently of A. Miranda Ribeiro (1915), Meek and Hildebrand INNER SURFACE OUTER SURFACE am d m. A d. m.— r^— d.m. A' B* v. m.-' d.m.— Figure 2.— Terminology of otoliths. A, A', a: sagitta and B, B', b: lapillus of Bairdiella chrysoura; C, C, c: sagitta and D: relative size of lapil- lus in Leiostomus xanthurus. A, B, C, D: inner surface; A', B', C: outer surface; a, b, c: lateral view with inner surface down and anterior end to the left. a.m. = anterior margin; a. v. = anterior ventral margin; d.m. = dorsal margin; c. = cauda section of sulcus; m.g. = marginal groove; o. = ostium section of sulcus; p.d. = posterior dorsal margin; s. = "tadpole-shaped" sulcus (dotted area); v.m. = ventral margin. All subsequent figures of otoliths (sagittae and lapilli) are oriented as in this figure. (1925), Ginsburg (1929), Fowler (1942, 1954), Schultz (1945, 1949), Travassos and Paiva (1957), Lowe (1966), Cervigon (1966a), Cervigon and Holanda Lima (1972), Vazzoler (1969), Travassos and Rego-Barros (1971), Travassos (1973), Roux (1973), Jardim (1973), and Druz- hinin (1974) have contributed to the knowledge of tax- onomy and distribution of western Atlantic sciaenids in different geographic regions. At present, 21 genera and 57 species of sciaenids are recognized from the western At- lantic. In addition, there is one monotypic genus (Ap- lodinotus) recognized from the freshwaters of North America, and three genera (Pachypops, Pachyurus, and Plagioscionj from the freshwaters of South America. Few studies have been made on the relationships of genera and higher division of western Atlantic Sciaenidae. Gill (1962a, b, c, d) revised the genera and named six subfamilies for North American sciaenids, i.e., Corvininae, Haploidonotinae, Lariminae, Liostominae, Otolithinae, and Sciaeninae. Later, only five subfamilies of Sciaenidae were listed, the Cor- vininae was not mentioned (Gill 1864a, 1873). Genera and subfamilies were defined by external characters and the boundaries between them were not clear. Bleeker (1874) distinguished Isopisthus and Seriphus from the rest of the Sciaenidae on the basis of complete separation of the dorsal fins and longer anal fins, and included them as. h.s. B as as B 1 Figure 3.— Inner ears of Sciaenidae. A and A', with only sagitta enlarged, Pseudoaciaena crocea (Richardson), after Chu et al. ( I %.'): fig. 8A, B). B and B', with both sagitta and lapillus enlarged, Stellifer lanceolatus (Holbrook). A, B: dorsal view of right inner ear. A', B': lateral view of left inner ear. a = asteriscus; am. = ampulla; a.s. = anterior semicircular canal; c. = cere- bellum; h.,?. = horizontal semicircular canal; 1. = lapillus; la. = lagena; m. = mesencephalon; m.o. = medulla oblongata; o. = olfactory nerve; p.s. = posterior semicircular canal; s. = sagitta; sa. = sacculus; t. = telencephalon; u. = utriculus; v.l. = vagal lobe. in a small new family, "Scombropsoidei," which he felt, in some way, was a transition between the sciaenids and the sphyraenids. Later, Bleeker (1876) in part two of "Systema percarum revisum" named two phalanxes (tribes), the Sciaenini and the Hemisciaenini for the "familiar Sciaenoidei." He also described three sub- phalanxes (subtribes) in the Sciaenini, i.e., Equili, Aplodinoti, and Johnii, and two subphalanxes in the Hemisciaenini, i.e., Pseudosciaeni and Otolithi. Western Atlantic sciaenid genera were described under each sub- phalanx. The diagnostic characters for each taxon were determined from the external morphology, e.g., mouth position, dentition, scales, and fin rays. Jordan and Eigenmann (1889) and Jordan and Ever- mann (1896) divided the Sciaenidae into two sub- families, the Otolithinae and the Sciaeninae, based on the number of precaudal and caudal vertebrae, lower jaw position, dentition, and size of the second anal spines. In the present study, I did not find these characters useful above the generic level. The first comprehensive studies on the species and generic relationships of sciaenids were published by Trewavas (1962) for tropical west African sciaenids and Chu et al. (1963) for Chinese sciaenids. Trewavas (1962 and pers. commun.) grouped the 16 west African sciaenid species into five tribes, based mainly on swim bladder structure; and the diagnostic characters of each tribe were clearly defined. Chu et al. (1963) grouped 37 species of Chinese sciaenid fishes into seven sub- families based on both swim bladder and otolith struc- ture. Relationships among genera and subfamilies were also clearly defined. Because the authors lacked access to type-material, the true identity of a few species may be in doubt. Trewavas (1964), discussed the relationships of sciaenid fishes with a single mental barbel. She used the barbels and the structure of the swim bladder to assess the relationships of some barbeled sciaenid genera of the Xew World and suggested possible links of certain genera with Old World sciaenids. Similarly, Lai Mohan (1969) used morphology of the swim bladders and otoliths as diagnostic characters to define each of the 14 genera of Indian Sciaenidae. Although he placed the 14 genera in five subfamilies and recognized two tribes in each of two subfamilies, the definitions of some subfamilies and tribes remain unclear. Trewavas (1977) grouped 65 species of Indo-West- Pacific sciaenids in 27 genera and 10 tribes based on the acustico-lateralis (pore) system, swim bladder and drumming muscles, and saccular otolith. The subfamily Otolithinae was redefined as an Indo-Pacific subfamily with one species (Argyrosomus regium) in the Mediter- ranean and eastern Atlantic. She also suggested that the New World genera Sciaena, Umbrina, and probably Menticirrhus are the basis of subfamily Sciaeninae and within this of the tribe Sciaenini. Trewavas recom- mended that the grouping of sciaenid tribes into sub- families, I agree, must await further knowledge of the family as a whole. DESCRIPTION AND RELATIONSHIPS _Plagioscion Stellifer X^Ophioscion Bairdiella OdoMoscion Figure 4. — Proposed phylogenetic relationships of genera and supra- generic groups of western Atlantic Sciaenidae. Subfamily level lies between dashed lines A and B; tribal level lies between dashed lines C and D; X, Y, and Z represent clusters discussed in the text. The morphology of the swim bladders, otoliths, and snout and mental pores and barbels are diagnostic in the classification of sciaenid fishes. They were described and compared first, then combined to assess the phy- logenetic relationships of western Atlantic Sciaenidae. Within each of the characters, there are several struc- tural patterns which may reflect phyletic relationships among groups of genera. The variations within each pat- tern are important in understanding the boundaries of different taxa. At the species level, the ontogenetic development of certain characters may suggest possible trends in phylogenetic relationships in higher taxa of the family. The presence of drumming muscles in males only or in both males and females of certain taxa is also useful in assessing the relationships of sciaenids. Other charac- ters essential for the identification of genera and species may also be important to the study of their relationships and are described in the key at the end of this paper. Based on swim bladder, otoliths, and external mor- phology, phyletic relationships (or kinships) are propos- ed here for 21 western Atlantic and 2 American fresh- water genera of sciaenids (Fig. 4). Relationships among suprageneric groups were determined primarily by swim bladder morphology and secondarily by morphology of the otoliths (Table 1). External morphology was of some use as long as the adaptive and potentially convergent nature of these characters was kept in mind. By follow- ing the tactics used by Bolin (1947) for the marine Cot- tidae of California, the actual or "convenient" classi- fication level for tribes may be drawn as between dashed lines C and D on Figure 4. The subfamily level may be drawn between dashed lines A and B. Between dashed lines B and C, the level could be either subfamily or tribe. Groupings Based on Swim Bladder Morphology Sciaenid fishes have a physoclistous swim bladder with a single or two interconnected chambers. The usual- ly well-developed swim bladder is attached very firmly to the ventral surface of the third to sixth vertebrae, under the postcardinal vein. The associated swim bladder (drumming) muscles usually present in males, are sheathlike or present lateral to the swim bladder (not at- tached to) on the body wall, except in Pogonias, in which they are present on the swim bladder. The drumming muscles are composed of red striated fibers with abun- dant cytoplasm and conspicuous nuclei (Tower 1908; Jones and Marshall 1953). Tower concluded that contrac- tions of the drumming muscle set up vibrations in the bladder walls, thus producing sounds. Sound produc- tion in sciaenid fishes may function as recognition sig- nals associated with spawning, feeding, and/or aggres- sive behavior (Naval Ordinance Laboratory 1942, 1944; XI a 2 . - c g 8 Q, CD x g. ■a a> CO CO CD B. E o cd CD « bo C _o CD >> CO x) -a cd" o CO CD CUD bJD fi -a & -a • 5 o >• t ° 1 XI c _c CD -f 3 £ CT x: oj -° 3 o 3 x CD I s CD X ° .3 tfl - E x E o ^o >, 3 X 9 is o li X !h o o ■8 § -° ,s CD CD P cn O* CD o „- ■" CO be CJ c 'S X CD ° X >> -w X <" o £ X . t." X O 0J '^ CO e2 ® c E. £ I X , X O X ^0 c£ CO X c co X s 1- C0 c CD bo J x CD **- OS O en CD «- >- it « 8 S| D- 0J minal, or oblique body ob- teeth conical in narrow nds 3 XI 2 S fi O > X 3 o c x?*S M X C •!-> O CD li CD — tS as e XI >. O no 5 £? « X O X! X co X ■*- o .5 "g So j= o CD »-. cd t.' "D .2 " *(-! . * ,f. 3 +~ 3 3 t- . - CO CD -73 X? In 01 T3 O ? CO "C £ ° A ft XI C E k o *a CO - CO '"• O X " _r- CD "£ T3 J£ 3 c 3 o 3 B E 2.S x 1 X "° J3 £ C X > x <£ X ° 3 5 O CD 3 1 O CD B x .S 3 O X CD §1 3 00 tJ3 O C xl CD 6 S E S E S, E =S $ S CD E 2 E 3 -c OS a 0> 2 co CO ft. '3 CO El *QJ X X u X3 CO — CO X cc — < 03 X a tj > B CO E en CD en ft -£ O CO cn "cd I 2 ^T O X O "-i O XJ o X o x S CD c c a "p a o » CN a o o. o o, a. "? io co ■* c lO 1-H IO c Tf c io c -*f i- CM c X ^ c - ■"■■■ - — ~^——^ Cm o as CO CD CO c 3 © o _ „ _ "3 c CD b£ 1-1 „ & a CO CO CD .E CO CO CD .E CO CD S ft O 3 C K 5 CD C I 3 O C CD CUD Q- IS & B 3 C ft '& o c CD b£ 3 E D. '5 o c SI 3 E 3 co ■> E be 08 lO IO C CN IO lO c >o CO in C CM CO IO h a 3 m J3 ■s o & CD > CD o CO -0 3 CO xf CD a J2 ft -2" 1 & 3 CD CO > O ° C c X CD 3 § J ■o"2 CD (5. » J CD CO* « St _3 g> | x 2 CO ^ CD XI — IS CO -" _ >. as "^ "o CD X a & > £ O CO - 3 CO O CD "^ -A « > _ i "3 O CO =0 > ~ CO a3 cS 2 xi 'ti CO CO S 3 ao S co 3 CO co || co E -M 'So E CO S cO S3 3 -° 60 XI 3 O. CO ™ co Z o s * ^H «s CO CO 3| ft O pa8jB[u 3 B}}l3BS Ajuo m 1 5 b a E '•2 c 03 o X c _c c CO CUD •O D » CD CO XI GO CD > _« -3 'C O CD CO CD N ft 5 B CO 03 B 1. ■3 "3 ■3 c CO '55 a x ft « M 4> XI s OS CD CD CD 53 03 > „ X 03 3 -a CD CD 3 CD CD 1 CO CD CD •3- s (-i XJ -o S '§ CO CD XI 3 O '3 a > XI X CD "a E o E | s ° * CD C E -2 CO CD > X CD B T3 CD CD s en as" CD ■5 C O X "3 g CD " CD g -B -g v D O > a "g ^ -a i: _ cd C 05 ft CO ° ^2 .2 "-3 Q. CD CD CO CO O. "3 X! CO CD "3 S x § XI ■5 3 ■g X 5 CO CO CD c CO* CO CO "^ .13 08 ■t 3 c "a X CO Li ■ +j cn eG fO X X .2 "C en '5 a >. CO -£ pac Bqs-)o«BD jappB[q uiia\s a CO X '5 D, cc 1 S CD co c a as 3 s o o o. CO CO .2 §■ S CO C3 b ° .1 '3 CO 3 3 09 o cd CD c o at ■S a --I .a -a c CD Si Dijkgraaf 1947; Knudsen et al. 1948; Johnson 1948; Jones and Marshall 1953; Tavolga 1964). The sounds produced by different species of sciaenids also have different ranges of frequencies (Knudsen et al. 1948; Tavolga 1964; Fish and Mowbray 1970). This may result from the different morphology of swim bladders in different sciaenids. Comparison of sciaenid swim bladders in the present study is confined to external morphology and in situ position within the fish. Morphological patterns of swim bladders. — All western Atlantic sciaenids have well-developed swim bladders except the genera Menticirrhus and Lonchurus. The swim bladders in Menticirrhus atrophy during the juvenile stage and only a vestige remains in adults. In adult Lonchurus, the relative size of the swim bladder is much reduced. Based on the shapes of swim bladders in mature fish, western Atlantic sciaenids can be divided into two groups. One group has a single-chambered swim bladder (Figs. 5 to 14) and the other has a two-chambered swim bladder (Figs. 15, 16). Several distinct structural patterns of swim bladders are recognized here. Single -chambered 8 wim bladder group. Sciaena pattern. — A simple carrot-shaped swim blad- der without well-developed anterior horns or divertcula (Fig. 5), its anterior end falling short of, reaching but never passing the septum transversum. Swim bladders of eight genera have this pattern: Aplodinotus, Cteno- sciaena, Equetus, Leiostomus, Menticirrhus, Pareques, Sciaena, and Umbrina. In addition, species with simple swim bladder are also found in other genera. The swim bladder of Larimus fasciatus is simple (Fig. 5E) but L. breviceps has a pair of anterior projections (Fig. 5F). Pachyurus schomburgkii has a simple swim bladder and differs from the diverticulated swim bladder of P. bonariensis (Fig. 6). The generic status of Pachyurus is discussed in the synopsis section. The swim bladder of Menticirrhus degenerates or Figure 5. — Swim bladders of the Sciaena pattern. A, Sciaena trewavaaae; B, Leiostomus xanthurus; C, Equetus punctatus; D, Aplodinotus grunniens; E, Larimus fasciatus; F, Larimus breviceps. B Figure 6. — Swim bladders of two species of Pachyurus: A, P. bona- riensis; B, P. schomburgkii. Figure 7. — Swim bladders of Menticirrhus saxatilus: A, 58 mm SL; B, 75.5 mm SL; C, 185 mm SL; D, 182 mm SL. v. = vent. Positions of vent (v) on C and D are omitted. cf atrophies in the adult. Menticirrhus saxitilis has a well- developed simple bottle-shaped swim bladder in young specimens up to 90-100 mm SL (Fig. 7A, B), but only a trace of the swim bladder remains in the adult (Fig. 7C, D). Bearden (1963) reported the length and condition of the swim bladders of 420 M. americanus. His results showed a gradual atrophy of the swim bladder from specimens of 56-60 mm SL (5.6 c "o atrophied) to specimens of 125-130 mm SL (100*^ atrophied). I ex- amined 28 specimens from 50-108 mm SL. Five specimens 71.1-108 mm SL had completely atrophied swim bladders and the rest had only a trace of the swim bladder less than 7 mm in length. Menticirrhus ameri- canus and M. littoralis possess an atrophied swim blad- der even in juvenile specimens. A much simpler swim bladder pattern may also be present in the juvenile stage of certain species with complicated swim bladder pattern in adults (e.g., Pogonias and Sciaenops patterns). Nebris pattern. — The swim bladder has a pair of broad diverticula that originate on the anterolateral mar- gin of the main chamber, extend posteriorly and loop an- teriorly near the posterior end of the main chamber (Fig. 8). The diverticula are tapered and terminate anteriorly at the septum transversum. Nebris is the only genus with this pattern. Pachyurus bonariensis (Fig. 6A) also has a pair of diverticula which originate anterolaterally, but it is morphologically very different from Nebris. Micropogonias pattern. — The swim bladder of Micro- pogonias has a pair of tubelike diverticula that originate laterally from the posterior half of the main chamber (Fig. 9). The diverticula extend anteriorly as far as the septum transversum, but never reach the skull; they then curve posteriorly to the anterior border of the swim bladder medially. Micropogonias is the only genus with this pattern. Pogonias pattern. — A very complex lateral system of diverticula (Fig. 10) characterizes the swim bladder of Pogonias, the only genus posessing this pattern. A serial study of specimens from 24 mm TL to 1,090 mm TL in- dicates that juvenile fish possess a simple swim bladder (Fig. 10A). As the fish grow, the diverticula first develop anterolaterally (Fig. 10B), then posterolaterally (Fig. 10C-E). These diverticula are connected to each other (anastomose) and to the main chamber, except the por- tion with the drumming muscles (Fig. 10, shaded area), where they are attached by ligament only. The ontogenetic development of this swim bladder pattern may reveal the developmental pathway of other swim bladder patterns. Figure 8. — Development of swim bladder in Nebris microps. A, 125 mm SL (x2); B, 273 mm SL (xl. Dashed line represents location of septum transversum. Figure 9. — Variation and development of swim blad- ders of Micropogonias. Micropogonias undulatus: A, 67.1 mm SL; A', 180 mm SL; and M. furnieri: B, 160 mm SL; B ', 162 mm SL. v. = vent. A' Figure 10.— Ontogenetic development of swim bladders in Pogonias cromis. A, 24 mm TL; B, 57.4 mm TL; C, 150 mm TL; D, 236 mm TL; E, 1,090 mm TL. C-E on the left sides of dashed midline represent dorsal view of the swim bladders (diverticula omitted); the shaded areas represent intrinsic drumming muscles; dashed line on top represents location of septum transversum. Sciaenops pattern. — The swim bladder of Sciaenops has a pair of tubelike anterior diverticula (Fig. HA). In large specimens, the complicated lateral diverticula and a pair of saclike projections are found dorsolateral^ in the anterior part (Fig. 11B, C). These diverticula are composed of numerous small labyrinthine chambers (Fig. 11D). Sometimes the lateral diverticula are com- pletely covered with fatty tissue (Fig. 11B). A pair of pits (cavities) is also present dorsolateral^ in the body wall, between the third and fourth pleural ribs, to receive the "saclike" dorsal projections of the swim bladder. Con- sequently, these projections extend closer to the body surface behind the base of the pectoral fins. This charac- teristic may be important in sound reception in large fish. Sciaenops is the only genus with this pattern. Cynoscion pattern. — A pair of prominent hornlike diverticula (horns) develops anteriorly from the antero- lateral corners in line with the sides of the main cham- ber (Fig. 12). The horns extend anteriorly passing the septum transversum, sometimes reaching to the back of the skull. The anterior border of the main chamber is al- ways behind the septum transversum. Four western At- lantic sciaenid genera have this pattern. Cynoscion, Iso- pisthus, Macrodon, and Plagioscion (Fig. 12A-D), and a single species, Atractoscion aequidens is present in the Old World (Trewavas 1962). The diversity in swim blad- ders among genera and species (Figs. 12, 13) with this pattern are limited to the extension of the horns, whether they are broad or narrow, straight or curved. These genera, except the primarily freshwater Plagioscion, have similar external features, such as an oblique ter- minal mouth, caninelike dentiton, and similar habitats. They form a distinct group among western Atlantic sciaenids. Lonchurus pattern. — The swim bladder has a pair of forked anterior diverticula, each with a long posteriorly directed branch and a short branch directed anteriorly (Fig. 14). Lonchurus, Paralonchurus, and Pachypops fourcroi have this pattern. The anterior branches of the swim bladder diverticula in Lonchurus and Paralon- churus are rather stout and short (Fig. 14A-C). They are thinner in Pachypops fourcroi (Fig. 14D), pierce the sep- tum transversum, and curve under the base of the skull. The relative size of the swim bladder in Lonchurus is much reduced (Fig. 14A, A', A"). Two-chambered swim bladder group. Stellifer pattern. — The swim bladder is divided into two chambers by an anterior constriction. The tip of the anterior chamber reaches into the head region and is yoke-shaped; the posterior chamber is carrot-shaped (Fig. 15). Bairdiella, Odontoscion, Ophioscion, and Stel- lifer have this pattern. A pair of diverticula is present on the posterolateral surface of the anterior chamber in Stellifer (Figs. 15B, 16). These genera usually have two pairs of enlarged otoliths (Fig. 2A, B, also see otolith sec- tion), a rather strong second anal spine and are relatively smaller fishes. This pattern of swim bladders is absent in the Old World sciaenids. 10 B Figure II. — Ontogenetic development of swim bladder in Sciaenops ocellata. A, 197 mm SL; B, 980 mm SL, shaded area represents fat tissue; C, 1082 mm SL, dorsal view of a portion of the swim bladder with the left side of the "saclike" projection; D, a cross section of the dorsal projection to show labyrinthine chambers. Dashed line represents location of septum transversum. ABC D Figure 12. — Swim bladders of the Cynoscion pattern. A, Aynoscion regalia; B, Isopisthus parvipinnis; C, Macrodon ancylodon; D, Plagi- oscion surinamensis: Dashed line represents location of septum transversum. Figure 13. — Swim bladders of species of Cynoscion: A, C. microlepi- dotus (393 mm SL); B, C. nothus (102 mm SL); C, C. virescens (219 mm SL); D, C. leiarchus (395 mm SL); D , C. leiarchus (191 mm SL). Groupings of swim bladder patterns. — Western At- itic sciaenids can be separated into two distinctive g oups (Fig. 17), based on the number of the swim blad- der chambers. The Stellifer pattern differs from all other sciaenids by having two chambers (Fig. 15). Single- chambered swim bladders are generally carrot-shaped with or without lateral diverticula (Figs. 5 to 11), except for the Cynoscion and Lonchurus patterns (Figs. 12 to 14). In the Cynoscion pattern the well-developed anterior horns are unique in developing laterally and following the contours of the main chamber forward (Fig. 12). The Lonchurus pattern has a pair of posterior branches from the base of the anterolateral^ developed diverticula (Fig. 14). 11 A' B Figure 14. — Relative size and development of swim bladders with the Lonchurus pattern. A, Lonchurus lanceolatus (87.7 mm SL); A , L. lanceolatus (103 mm SL); A", L. lanceolatus (114 mm SL, male); B, Paralon- churus elegans (180 mm SL, male); C, P. brasiliensis (157 mm SL, male); D, Pachypops fourcroi, after Cuvier and Valenciennes (1830:pl. 138, "Corb Fourcroy"), original size unknown, v. = vent. Figure 15. — Swim bladders of the Stellifer pattern. A. Bairdiella, Odonto8cion, and Ophioscion; B. Stellifer lanceolatus. Dashed line represents location of septum transversum. Figure 18. — Diversity of the swim bladder diverticula in the genus Stelli- fer. A, S. naso (75 mm SL); B, S. griseus (84.2 mm SL); B , S. griseus (108 mm SL); C, S. microps (129 mm SL); C , S. microps (105 mm SL); D, S. brasiliensis (76.6 mm SL); E, S. rastrifer (157 mm SL). I 'I -Pogonias Sciaenops . M/cropogon . Nebris . Larimus . Sciaena . . Menticirrhus -Cy nose ion ■ Lonchurus Stellifer Figure 17. — Groupings of western Atlantic Sciaenidae according to swim bladder patterns. Further grouping of swim bladder patterns (Fig. 17) relies on the similarity and complexity of swim blad- ders. The swim bladder of juvenile Menticirrhus (Fig. 7) is most similar to the Sciaena pattern, although only a vestige remains in adults. Species of Larimus usually have a pair of small anterior projections (Fig. 5F), ex- cept the northwestern Atlantic L. fasciatus, which lacks them (Fig. 5E). The swim bladder of Larimus appears to be a variation of the Sciaena pattern. These patterns form a group of simple swim bladders with or without small anterior projections (Fig. 17). The swim bladder pattern with well-developed divertic- ula can also be divided into two groups (or more) by the complexity of their lateral diverticula. Nebris have a pair of long diverticula originating from the anterolateral margin of the main chamber (Fig. 8) and extremely en- larged basally. In Micropogonias, the paired tubelike diverticula originate from the posterior half of the main chamber (Fig. 9). Another pattern of tubelike divertic- ula is found in Pachyurus bonariensis (Fig. 6A). It has a pair of anterior originated diverticula directed pos- teriorly. Sciaenops and Pogonias have much more complicated lateral diverticula in adults. Sciaenops has a simple short pair of diverticula in young specimens (Fig. 11A) and the "saclike" projections located dorsolateral^ develop in larger specimens (Fig. 11C). The complicated diverticula on the swim bladder of Pogonias also change ontogenetically (Fig. 10). The ontogenetic development of the swim bladder in Pogonias further indicates that the diverticula were developed from the simple swim bladder from anterior to posterior portions of the main chamber. The grouping of swim bladder patterns from simple to complex coincides with the ontogeny of the complicated swim bladder (e.g., Sciaenops and Pogonias patterns). This evidence may indicate a trend of swim bladder development, i.e., from simple to complex with the diverticula probably evolving from the anterior portion towards the posterior portion of the main swim bladder chambers. Specific characters. — Variations in swim bladder morphology are common within patterns. In the Stellifer pattern, the species of Stellifer have a pair of "bulblike" or "tubelike" diverticula at the posterolateral margins of the anterior chamber (Figs. 15B, 16). Slight variations are also found in species such as S. griseus and S. mi- crops (Fig. 16B, B', C, C). Bairdiella, Odontoscion, and Ophioscion lack diverticula on the anterior chamber of the swim bladder (Fig. 15A). The yoke-shaped anterior chambers in the Stellifer pattern terminate near the artic- ulation between the opercular and hyomandibular bones (Wintersteen and Chao'). In Bairdiella and Odon- toscion, the anterior swim bladder chambers curve for- ward abruptly and their tips end just behind the adduc- tor operculi muscle and medially to the dialator and levator operculi. In Ophioscion and Stellifer, the an- terior swim bladder chambers diverge more laterally, and their tips end subcutaneously and are frequently visible on superficial inspection. In the Cynoscion pat- tern, variations in anterior horn morphology were found among different genera (Fig. 12), species, and also onto- genetically within species (Fig. 13). There is little diver- sity in morphology among the swim bladders of the Sciaena pattern (Fig. 5). The anterior projections are present in Larimus breviceps and absent in L. fasciatus, indicating the possible extent of variation of swim blad- der morphology within a genus and/or a pattern. The drumming muscle is also useful in assessing rela- tionships within sciaenid fishes. The development of the drumming muscles seems to be correlated with the size of swim bladders and the sound producing function in some western Atlantic sciaenids. Generally, drumming muscles are present in male sciaenid fishes with a well- developed swim bladder (Smith 1905). In fishes with the Lonchurus pattern of swim bladders (Fig. 14), drum- ming muscles are well-developed in males of Paralon- churus but reduced in Lonchurus. Juvenile Men- ticirrhus have only a vestigial swim bladder and lack drumming muscles. Drumming muscles are present in both male and female Micropogonias. Pogonias has a unique drumming muscle among western Atlantic sciaenids, located intrinsically on the swim bladder in both sexes (Fig. 10). The drumming muscles of male Cy- noscion regalis are only well-developed during the spawning season and regress after spawning (J. Merriner, VIMS, pers. commun.). Possible relationships of western Atlantic sciaenids to sciaenids of other geographic areas may be revealed by the swim bladder shapes. Trewavas (1964) suggested a Wintersteen. J., and L. N. Chao. A new tribe of sciaenid fishes, with a revision of the genus Stellifer Oken, 1817. Manuscr. 13 possible link of Pachypops and Polyclemus (= Paralon- churus), the Lonchurus pattern swim bladder, with the tropical West African tribe Pseudotolithini. The swim bladder of Atractoscion aequidens (Cynoscionini of Trewavas) agrees with the Cynoscion pattern of the pre- sent study. The diverticula on the swim bladder of Pachyurus bonariensis (Fig. 6A) is also similar to that of Bahaba flavolabiata of the Indo-West Pacific (Chu et al. 1963). The complex swim bladder patterns of the Pogonias and Sciaenops patterns are common among many Indo-Pacific (Chu et al. 1963; Lai Mohan 1969) and some East Pacific sciaenids (e.g., Roncador stearn- sii). The branching diverticula of Pogonias and Sciaenops anastomose with each other, unlike the swim bladders of Indo-Pacific Otolithinae. Their relation- ships are not known at present. Groupings Based on Otolith Morphology The inner ear of sciaenids consists of three semicir- cular canals and a large sacculus, containing the sagitta (Fig. 3). The utriculus is an expanded area of the an- terior semicircular canal containing the lapillus near the junction with the horizontal semicircular canal. This sec- tion is greatly expanded in fishes with a large lapillus (Fig. 3B, B'). Posteriorly, the lagena is a small sac, located posterodorsally to the sacculus, containing the asteriscus (Parker 1908 and Chu et al. 1963). The relative sizes of the three otoliths, particularly the sagitta in all and the lapillus in some sciaenids, are usually larger and thicker than in most Perciformes. The discussion here is based on the general morphology of the sagitta and the lapillus. The sagitta lies obliquely in the sacculus with the smooth surface inside (medially) and is sloping from dorsolateral to ventromedial in situ (Fig. 3). The lapillus is enclosed in the utriculus obliquely, with the smooth surface inside and is oriented laterally. (Fig. 3B, B'). Morphological patterns of otoliths. — The sagitta of Sciaenidae is characterized by always having a distinct "tadpole-shaped" sulcus on its inner surface (Fig. 2). A marginal groove is usually present between the dorsal margin and the sulcus of the sagitta. The anterior por- tion of the sulcus usually is expanded into a pear or oval shape and is termed the "ostium," which always has a short spoutlike groove connecting with the anterior mar- gin of the sagitta (not shown in figures). The elongate narrow posterior portion is usually bent obliquely and is termed the "cauda" of the sulcus (Frizzell and Dante 1965; Stinton 1975). The sagitta distinguishes sciaenids from other perchlike families in that the ostium is merely outlined and never channeled as in other families, whereas the cauda is always channeled (or scooped out) in the Sciaenidae. The outer surface usually has crest- like or bloblike projections or granulations (Fig. 2B', C). Western Atlantic sciaenids can readily be divided into two groups based on their otoliths: a group with both the lapillus and the sagitta enlarged and a group with only the sagitta enlarged (Fig. 2). Based on the shape, thickness, and size of the sagitta and lapillus, and the morphology of the ostium and the cauda of the sulcus, 11 distinct patterns of otoliths are recognized here. Only sagitta enlarged. Sciaena pattern. — The sagitta is more or less oval, usually with a smoothly convex ventral margin and a straight or crenulate dorsal margin (Fig. 18). Laterally the outer surface of the sagitta is usually much thicker in the middle. The ostium of the sulcus is usually pear- shaped and its expanded part reaches the anterior mar- gin of the sagitta. The cauda is J-shaped with a relatively long and narrow distal end. A marginal groove is usually present between the sulcus and dorsal margin of the sagitta. Ctenosciaena, Equetus, Leiostomus, Pachyurus, Pareques, Plagioscion, Sciaena, and Umbrina have this pattern (Fig. 18). Variations in this sagitta pattern are mainly in the position of the ostium and the length and the curvature of the cauda, both of which vary among genera and species. In most species, the axis of the os- tium is parallel to that of the sagitta or inclined antero- ventrally (to the left of Fig. 18). The sagitta of Leiostomus xanthurus (Fig. 18D) differs from that of the Sciaena pattern in having the ostium inclined towards the dorsal margin and much thinner. Pachyurus schom- burgkii and Plagioscion surinamensis (Fig. 18K, L) also differ in the morphology of their sulci. Their relationship to Sciaena pattern is not certain at present. Pogonias pattern. — The sagitta is more or less semi- circular in shape with an evenly curved ventral margin and a straight but finely crenulate dorsal margin (Fig. 19). The ostium of the sulcus is broad and its expanded part does not reach the anterior margin of the sagitta. The cauda is J-shaped with a pointed distal end and does not reach to the margin. The marginal groove is present between the sulcus and the dorsal margin of the sagitta. Pogonias and Aplodinotus have this pattern. The relative size of the "tadpole-shaped" sulcus is larger in Pogonias than in Aplodinotus. The sagitta is more curved laterally in Pogonias than Aplodinotus, but the latter has more granulations on the outer surface (Fig. 19B', b). Larimus pattern. — The sagitta is slightly elongate and ovoid with a rather straight and slightly crenulate dorsal margin and a convex ventral margin (Fig. 20). The ostium of the sulcus is large, pear-shaped, and its ex- panded part does not reach the anterior margin of the sagitta. The J-shaped cauda is bend acutely with a pointed distal end. The marginal groove is not distinct. Larimus and Sciaenops have this pattern. The sagitta of Larimus is broader anteriorly than posteriorly (Fig. 20A, B); both ends are about equal in Sciaenops (Fig. 20C, D). The posterior portion of the sagitta is laterally very thick in Larimus (Fig. 20a, b). Sciaenops has a relatively thinner sagitta (Fig. 20c) and the shape of the sulcus is different from Larimus; therefore, a distinct otolith pat- tern could be recognized. 14 H K i J I Figure 18.— Inner surface (capital letters) and lateral view (lower case letters) of sagittae of the Sciaena pattern. A, a: Ctenosciaena gracilicirrh.ua; B, b: Equetus lanceolatus; C, c: Equetus punc- tatus; D, d: Leiostomus xanthurus; E, e: Pareques acuminatus; F, f: Paraques umbrosus; G, g: Sciaena trewavasae; H, h: Sciaena bathytatos; I, i: Umbrina coroides; J, j: Umbrina milliae; K, k: Pachyuru.8 schomburghii; L, 1: Plagioscion surinamensis. Nebris pattern. — The sagitta in Nebris is very thick and oval with a notch at the posterior margin (Fig. 21). The ostium of the sulcus is large and ovoid, its expanded part does not reach the anterior margin of the sagitta. The cauda is sharply bent with a very deeply grooved and enlarged distal portion. The marginal groove is absent. The outer surface of the sagitta is extremely elevated and very thick laterally (Fig. 21a). Cynoscion pattern. — The sagitta is elliptical and thin (Figs. 22A, 23), the dorsal margin is usually smoothly convex, and the ventral margin is slightly concave and finely crenulate. Laterally, the posterior portion of the sagitta is thicker than the anterior portion. The sulcus is usually elongate, the ostium ovoid or pear-shaped (Figs. 22A, 23). The expanded anterior part of the ostium may or may not reach the anterior margin of the sagitta. This varies among species and also within species, onto- genetically (Fig. 23). The cauda is long and bent with a short distal end (Figs. 22A, 23). The marginal groove is usually distinct. The outer surface usually has a granu- lar appearance, especially in large adult specimens (Fig. 23a-c). Cynoscion species have this pattern. In some species of Cynoscion, a notch is present in the middle of the dorsal margin (Fig. 23C, C). Menticirrhus pattern. — The sagitta is elliptical and thin (Fig. 22B, B', b), the dorsal margin is usually 15 B' Figure 19. — Sagittae of the Pogonias pattern. A, A', a: Pogonias cromis; B, B', b: Aplodinotus grunniens. A, B: inner surface; A', B': outer surface; a, b: lateral view. Figure 20.— Saggitae of the Larimus pattern. A, A', a: Larimua fasciatus; B, B', b: Larimus breviceps; C, C, c: Sciaenops ocellata (357 mm TL); D: S. ocellata (1,100 mm TL). A-D: inner surface; A -C : outer sur- face; a-c: lateral view. 16 Figure 21. — Sagitta in Nebris microps. A, inner surface; A', outer surface; a, lateral view. Figure 22. — Sagittae of the Cynoscion and the Menticirrhus patterns. A, A', a: Cynoscion nebulosus; B, B', b: Menticirrhus saxatilis. A, B: inner surface; A', B': outer surface; a, b: lateral view. A' Figure 23. — Specific and ontogenetic variations of sagittae in species of Cynoscion: C. regalia: A, a, 370mmTL; A , 516 mm TL; C. similis: B, b, 166 mm TL; B , 350 mm TL; C. virescens: C, c, 260 mm TL; C , 350 mm TL. A, A , B, B , C, C : inner surface; a, b, c: lateral view. 17 straight and finely crenulate. and the ventral margin is slightly convex. Laterally, the middle portion is the thickest. The sulcus is elongate, the ostium is pear- shaped (Fig. 22B) and its expanded part does not reach the anterior margin of the sagitta. The cauda is long and J-shaped. The marginal groove is present. The outer sur- face usually has crests and granulations. Menticirrhus species have this pattern. Lonchurus pattern. — The sagitta is moderately thin and elongate (Fig. 24), the ventral margin is usually smoothly convex, and the dorsal margin is slightly con- cave or crenulate. Laterally, the posterior portion of the sagitta is thicker than the anterior portion. The sulcus on the inner surface is elongate. The ostium is pear-shaped and bent toward the ventral margin of the sagitta an- teriorly. The cauda is J-shaped with a rather expanded distal end. close or reaching to the ventral posterior mar- gin of the sagitta. The marginal groove is absent. The outer surface has granulations (Fig. 24a-c). Lonchurus and Paralonchurus have this pattern. There are two dis- tinctive forms of sulcus on sagittae of this pattern. Lon- churus lanceolatus and Paralonchurus elegans have a rather narrow ostium (Fig. 24A, B). Paralonchurus brasiliensis has a relatively thicker sagitta and the os- tium is broader (Fig. 24C, c). Other forms of sagittae. Isopisthus pattern. — The sagitta is slightly elongate, rather broad and thick posteriorly (Fig. 25A, A', a). The sulcus is elongate; the ostium is round and its ex- panded part reaches to the anterior margin of the sagitta. The cauda is long and slightly bent at the distal end. The marginal groove is not distinct. The outer surface has crestlike elevations, especially thickened at the pos- terior half (Fig. 25a). Isopisthus has this pattern. Figure 24.— Sagittae of the Lonchurus pattern. A, A', a: Lonchurus lanceolatus; B, B', b: Paralon- churus elegans; C, C , c: P. brasiliensis. A-C: inner surface; A'-C: outer surface; a-c: lateral line. Figure 25.— Sagittae of the Isopisthus and the Macrodon patterns. A, A', a: Isopisthus parvipinnis; B, B', b: Macrodon ancylodon. A, B: inner surface. A', B': outer surface; a, b, lateral view. 18 Macrodon pattern. — The sagitta is relatively elongate and thin (Fig. 25B, b', b). The dorsal margin has a deep notch on the posterior half and the ventral margin has a projection in the middle. The anterior por- tion of the sagitta is slightly thicker than the posterior portion. The sulcus is elongate; the ostium is oval with a broader anterior portion and reaches to the anterior mar- gin of the sagitta. The cauda is rather straight and short with a disclike distal end (Fig. 25B). The marginal groove is absent. Macrodon has this pattern. Micropogonia8 pattern. — The sagitta in Micro- pogonias is very thick and shield-shaped, often with a shelf or flange on the outer surface or on the dorsal mar- gin (Fig. 26). The ostium of the sulcus is large and its ex- panded part does not reach the anterior margin of the sagitta. The cauda is oblique and bent only slightly towards the ventral margin with a round disclike distal end. The marginal groove is distinct. The sagittal shelf on the outer surface of M. furnieri and the lateral flange of M. undulatus vary with size and possibly onto- genetically (Fig. 26). Figure 26.— Variation, possibly ontogenetic, in sagittae of the Micropogonias pattern. Micropogonias fur- nieri: A, A , a, 235 mm TL; B, B , b, 295 mm TL; C, C , c, 330 mm TL. Micropogon undulatus: D, D , d, 141 mm TL; E, E , e, 250 mm TL; F, F , f, 350 mm TL. A-F: inner surface; A -F : outer surface; a-f: lateral view. 19 In summary, sagittae with the Sciaena, Larimus, and Nebris patterns are usually oval in shape and very thick in the middle. The Pogonias pattern is usually semicircular in shape and thin. Sagittae with the Cynos- cion, Menticirrhus, Lonchurus, and Macrodon patterns are usually more elongate and thin. The Isopisthus pat- tern is elongate and thick in the posterior portion. Sagit- tae with the Micropogonias pattern are irregular in shape and very thick. Both sagitta and lapillus enlarged. Stellifer pattern. — Although the anterior portion of the sagitta is reduced in size, it is still larger than in most Perciformes. The lapillus is much larger than in other sciaenids, about two-thirds or more of the sagitta size (Fig. 27). The ostium of the sulcus lacks its anterior por- tion, and the cauda is clearly bent in a J-shape reaching almost to the ventral margin of the sagitta. Bairdiella, Odontoscion, Ophioscion, and Stellifer have this pat- tern. There is considerable morphological variation within this otolith pattern (Fig. 27). Stellifer and Ophioscion have a parallelogram- shaped sagitta and a suboval lapillus, with both otoliths about the same size (Fig. 27A, A', B, B'). The ostium of the sulcus is short and reaches to the anterior margin of the sagitta. The cauda is bent obliquely towards the ven- tral margin of the sagitta. A marginal groove is evident along the dorsal margin of the cauda. The whole sagitta appears to be truncated at the middle of the ostium. The lapillus is large, sometimes larger than the sagitta (e.g., Stellifer, Fig. 28); a deep groove is present at the antero- ventral end of its inner surface with an open end to the ventral margin (Fig. 28A'-D'). The sagitta of Bairdiella is irregularly triangular shaped with a projection at the enterodorsal corner dor- A' B' C Figure 27.— Inner surface of sagittae and lapilli of the Stellifer pattern. A, A : Ophioscion punctatissimus; B, B': Stellifer lanceolatus; C, C: Bairdiella chrysoura; D, D : Odontoscion dentex. A-D; sagittae; A'-D': lapilli. Figure 28.— Inner surface of sagittae and lapilli in species of Stellifer: A, A': S. colonensis; B, B': S. microps; C, C: S. ras- trifer; D, D': S. venezuelae. A-D: sagittae; A'-D': lapilli. B B' C c D D' 20 sally (Fig. 27C). The ostium of the sulcus is very short. The cauda is inverse L (or J)-shaped and broadened at the middle. The marginal groove is well defined. The lapillus is smaller than the sagitta and is irregular in shape (Fig. 27C). The groove along the anteroventral end of the lapillus opens to the ventral margin. The sagitta of Odontoscion is subtriangular in shape. The ostium of the sulcus is reduced to a small oval- shaped piece at the anterior margin (Fig. 27D). The cauda is long and reaching to the ventral margin. The marginal groove is present on the sagitta. The lapillus is ovoid and its anteroventral groove is distinct (Fig. 27D'). Groupings of otolith patterns. — Based on the mor- phology of otoliths, western Atlantic sciaenids can readi- ly be divided into two groups (Figs. 2, 3, 29; Chao and Miller 1975). The Stellifer pattern differs from all other sciaenids by having a large lapillus that is more than one-half the size of the sagitta (Figs. 27, 29). The remain- ing morphological otolith patterns can be divided into two groups (Fig. 29), one having a more or less ovoid or oval-elongate sagitta and the other having a shieldlike sagitta as found in Micropogonias (Fig. 26). Within the former group, the Cynoscion, Menticirrhus, Lonchurus, and Macrodon patterns usually have thin, elongate sagit- tae (Figs. 22 to 25) and the rest have a broader and thicker sagitta. In the general outline of the sagitta, the — Micropogon — Neb r is c c Larimus Sciaenops Sclaena Pogonias . Leiostomus . Macrodon Cynoscion Menticirrhus Lonchurus .Isopisthus .Stellifer Figure 29. — Groupings of western Atlantic Sciaenidae according to otolith patterns. Cynoscion, Menticirrhus, and Lonchurus patterns are most similar to each other and have the posterior por- tion of their sagittae slightly thicker than the anterior portion (Figs. 22, 24). The Isopisthus pattern has the pos- terior portion of its sagittae further thickened (Fig. 25a). The sagitta of Macrodon is unique in having the anterior portion of the sagitta thicker than the posterior portion (Fig. 25b). Among the remaining otolith patterns, the Nebris pattern (Fig. 21) is unique in the general outline of its sagitta and the morphology of the sulcus (Fig. 29). The Larimus pattern has a "rectangular" sagitta (Fig. 20), which differs from the more generalized oval-shaped sagitta of the Sciaena and Pogonias patterns (Figs. 18, 19, 29). Leiostomus xanthurus also differs from other Sciaena patterns by having its sagitta (Fig. 18D, d) nar- rower and thinner. It is probably an intermediate form between the elongate thin and ovoid thick forms of sagit- tae. Specific characters. — The sizes of the lapillus and sagitta in the Stellifer pattern vary among the genera (Fig. 27). The trend of reducing the size of the sagitta and increasing the size of the lapillus is evident from Ophios- cion to Stellifer, and Bairdiella, than to Odontoscion (Fig. 27). Cynoscion has the greatest number of species (12) of any western Atlantic sciaenid genus, and the mor- phological variation in their sagittae is great (Fig. 23). The otoliths in Menticirrhus are relatively smaller than in other genera of sciaenids (Smith 1905). Micropogonias has the most distinctive sagitta, with the sagitta further ossified and a lateral flange developed along the dorsal margin (Fig. 26). Nebris is unique in having an oval, thick sagitta (Fig. 21) with a notch in the posterior mar- gin that does not resemble that of any other western At- lantic sciaenid. Although the Pogonias pattern (Fig. 19) has a semicircular sagitta, the sulcus is similar to that of the Sciaena pattern (Fig. 18). Variations among the genera with the Sciaena pattern are mainly in the posi- tion and shape of the sulci. The sagittae of Larimus and Sciaenops are slightly rectangular (Fig. 20), but the shapes of the sulci differ. Within a given taxon, the morphology of the otoliths appears more variable than that of the swim bladder. Chu et al. (1963) relied heavily on the relative position of the "head" (ostium) and "tail" (cauda) sections of the "tadpole-shaped" sulcus to diagnose the subfamilies of Chinese Sciaenidae. In some species of western Atlantic sciaenids, the relative position of the ostium and cauda shift slightly as the fish grows (Figs. 20, 23, 26). This feature should be viewed cautiously when used to group taxa above the generic level. Schmidt (1968) indicated that the sagitta of sciaenid fishes are large, extraordinarily compact, and often ap- pear rather shapeless. The ratio of otolith length to stan- dard length varies between 1 to 18 and 1 to 28; the ratio of otolith length to otolith thickness may be as high as 1 to 0.45. He also noted many typical features of sciaenid otoliths that make up a "sciaenid type." Fossil materials of sciaenid otoliths are important in assessing phyletic relationships of modern sciaenids. 21 John Fitch of the California Department of Fish and Game is currently studying this aspect of sciaenid rela- tionships. Nolf (1976) described sciaenid otoliths of Pliocene and Miocene (neogenes) from Trinidad and in- dicated that some modern sciaenid species had a wider range of distribution in the past. Groupings Based on External Morphology Prior to Trewavas (1962) and Chu et al. (1963), most recent authors used external morphology to classify sciaenid fishes (see section "A brief history of the study of western Atlantic Sciaenidae"). Characters used were body shape, presence or absence of the mental barbel and/or pores, size of the second anal fin spine, and posi- tion of the mouth. Unfortunately these characters are ex- tremely adaptive, evolutionary plastic, and conver- gent. They are more indicative of feeding adaptations and habitats than of phylogenetic relationships (Fig. 30). Even so. these characters do show that similar external morphology is often found in closely related taxa. Exter- nal morphology is also diagnostic at the generic and specific levels. The following discussion assesses the limits and usefulness of these characters. Patterns of pore and barbel systems. — The pores at the tip of the snout and lower jaw (mandible) of sciaenids are the openings of the well-developed cavernous lateral- line canals on the head (Fig. 1). The snout (rostral) pores can be divided into upper pores, those present at the tip of snout, and marginal pores, those present along the edge of the rostral fold (Fig. 1). Usually, there are three or five distinct upper pores, although some genera lack upper pores and some may have more than five as adults. Typically there are five marginal pores, one median and two pairs of lateral pores (Fig. 31). Some genera have only two marginal pores. The mental (mandibular) pores are most often five (four to six), one median and two pairs of lateral pores (Fig. 32), whereas some genera com- pletely lack mental pores. One or more mental barbels may be present in many western Atlantic sciaenid fishes. The number of pores and barbels is correlated with the mouth position and feeding habitats of the species (Chao 1976; Chao and Musick 1977). Variations are broad among the genera and species (Figs. 33 to 36). Structural patterns of snout pores. — There are four snout pore patterns recognized in western Atlantic Sciaenidae (Fig. 31). Two (or no) marginal pores. — This pattern has no upper pores and the rostral fold is thin and entire without notches (Fig. 31 A). Cynoscion, Macrodon, Nebris, and Plagioscion have this pattern. They have a very oblique, large mouth and the lower jaw projects in front of the up- per, except in Plagioscion, which has a slightly oblique and terminal mouth. Fishes with this snout pore pattern and mouth position are midwater to upper water column feeders. Five marginal pores. — This pattern has no upper pores and the rostral fold has notches at the openings of the lateral pores (Fig. 31B). Aplodinotus and Larimus have this pattern. Aplodinotus has a moderate-sized mouth, terminal or slightly inferior, and is a typical lower water column to bottom feeder. Larimus has a large mouth, very oblique, and feeds in midwater to the upper water column. LARGE CANINES, BODY ELONGATE WITHOUT BARBEL SCIAENIDAE- WITHOUT LARGE CANINES WITH BARBEL MOUTH VERY OBLIQUE MOUTH OBLIQUE GENERA _ Cynoscion Macrodon _ Isopisthus L arimus ~l_ Nebris MOUTH TERMINAL i: MOUTH INFERIOR TO SLIGHTLY OBLIQUE BACK NOT 1 Bairdiella Stellifer Ophioscion Sciaena Leiostomus Pareques —Equetus body elongate Sr.mf>nnps ELEVATED BACK ELEVATEO no apical pore Ctenosciaena WITH AN APICAL TWO BARBELS Umbrina . \_Menticirrhus .Lonchurus \ . Paralonchurus ' BARBELS IN TUFT barbels not r- Micropogon in tuft ~]_Pogonias Figure 30.— Groupings of western Atlantic Sciaenidae according to external morphology. 22 Five marginal and three upper pores. — The rostral fold of this pattern is indented at the openings of the marginal pores (Fig. 31C). Bairdiella, Ctenosciaena, Menticirrhus, Odontoscion, Ophioscion, Paralonchurus, and Stellifer have this pattern. The position and size of the mouths in this group of fishes suggest that some feed in midwater and some on the bottom. In bottom feeders such as Menticirrhus and Paralonchurus, the ros- tral folds are very deeply indented below the marginal pores (Fig. 33C, D). For the midwater or lower midwater feeders, Stellifer lanceolatus and Ophioscion punctatis- simus, the rostral folds are only slightly indented (Fig. 33A, B). Ophioscion punctatissimus also has two minute pairs of upper pores present dorsolateral^ to the outer lateral marginal margins pores (Fig. 33B). -e=^_3^ Five (or more) marginal and five (or more) upper pores. — This pattern usually has five distinct rostral and five marginal pores (Fig. 31D). Sometimes an extra pair or Figure 31. — Snout pore patterns (diagramatic) of western Atlantic Sciaenidae. A, no upper and two marginal pores; B, no upper and five marginal pores; C, three upper and five marginal pores; D, five upper and five marginal pores. H Figure 32.— Mental pore and barbel patterns (diagramatic) on the lower jaw of western Atlantic Sciaenidae. A, no pore and no barbel; B, four pores and no barbel; C, five pores and no barbel; D, six pores and no barbel; E, four pores and one barbel; F, four pores and two barbels; G, five pores and many barbels, not in tuft; H, five pores and many barbels, anterior three pairs of barbels in tuft. E F G H Figure 33. — Diversity in snout pores with five or more marginal pores. A-D, three upper pore pattern; E-H, five upper pore pattern. A, Bairdiella chrysoura; B, Ophioscion punc- tatissimus (with additional four minute pores); C, Paralonchurus elegans; D, Menticir- rhus americanus; E, Pogonias cromis; F, Sciaenops ocellata; G, Micropogonias un- dulatus; H, Pareques umbrosus (with seven marginal pores). 23 two pairs of minute pores are developed lateral to the upper and marginal pores (Fig. 33H). The rostral fold is smooth or slightly notched below (or at) the openings of the marginal pores. Equetus, Leiostomus, Lonchurus, Micropogonias, Pachyurus, Pareques, Pogonias, Sciaena, Sciaenops. and l r mbrina have this pattern, have a horizontal or inferior mouth, and are typically bottom feeders. Variation within this snout pore pattern is main- ly in the presence of additional minute pores (Fig. 33E- H), which may also vary ontogenetically. Structural patterns of mental pores and bar- bels. — There are seven mental pore and barbel patterns recognizable in western Atlantic sciaenids (Fig. 32). Variations are rather common within genera, especially in speciose genera such as Stellifer and Bairdiella. No pores. — This pattern has neither pores nor bar- bels on the lower jaw (Fig. 32A) and is found in Cynos- cion, Isopisthus, Macrodon, and Plagioscion. All fishes with this pattern have a large oblique mouth and are up- per water or midwater feeders, although Plagioscion has a nearly terminal mouth. Four pores. — This pattern has two pairs of laterally set pores (Fig. 32B), which may vary in arrangement and size among different genera (Figs. 34A-D, 35A). Nebris, Odontoscion, Larimus, and some Stellifer species have this pattern, have a terminal or oblique mouth, and feed in the midwater to upper water column. Five pores. — This pattern has one median pore located at the center of the lower jaw symphysis and two E F G H Figure 34. — Diversity in mental pore arrangements on the lower jaw. A-D, four mental pore pattern; E-H, five mental pore pattern. A, Nebris microps; B, Larimus fasciatus; C, Odontoscion dentex; D, Stellifer sp. B (see p. 55, key); E, Aplodinotus grunniens; F, Sciaenops ocellata; G, Ophioscion punctatus; H, Equetus punctatus. B Figure 35. — Diversity in mental pore patterns on the lower jaw in the species of Stellifer; A, Stellifer sp. B (see p. 55, key), four mental pores; B, S. stellifer, five mental pores; C, S. rastrifer, six mental pores; D, S. colonensis, six mental pores. c D Figure 36.— Western Atlantic Sciaenidae with one mental barbel on the lower jaw. A, Ctenosciaena gracilicirrhus (lacks an apical pore); B, Menticirrhus and C, Umbrina (with an apical pore). 24 lateral pairs (Fig. 32C). In fact, the median pore (open- ing) often consists of a pair of closely set pores inside. The arrangement of the five pores varies among genera (Figs. 34E-H, 35B). Aplodinotus, Equetus, Leiostomus, Ophioscion, Pachyurus, Pareques, Sciaena, Sciaenops, and some species of Bairdiella (B. ronchus and B. sanc- taeluciae) and Stellifer (S. microps, S. stellifer, and S. uenezuelae) have this pattern. Six pores. — This pattern has two median pores and two pairs of lateral pores (Figs. 32D; 35C, D). Bairdiella chrysoura, Stellifer colonensis, S. griseus, and S. rastri- fer have this pattern. Two median pores may be closely set and appear as the five-pored pattern. Fishes with this pattern of mental pores have a terminal to slightly inferior mouth and feed from lower midwater to the bottom. Four (or five) pores and one barbel. — This pattern has two pairs of lateral pores and a barbel at the anterior tip of the lower jaw (Fig. 32E). Ctenosciaena, Menticirrhus, and Umbrina have this pattern. The morphology of the barbel varies among these three genera (Fig. 36). Cteno- sciaena has a thin barbel tapering at the end. Menti- cirrhus and Umbrina have a short rigid barbel with an apical pore. They have subterminal to inferior mouths and are lower midwater to bottom feeders. Four pores and two barbels. — This pattern has two pairs of lateral pores and a pair of thin barbels originat- ing on the medial side of each posterior mental pore (Fig. 32F). Lonchurus is the only genus with this pattern and has an inferior mouth, typical of bottom feeders. Five pores and many barbels. — This pattern has five pores, one median and four lateral, together with 3 to 16 pairs of minute barbels at the symphyses of the man- dibles and along the rami of the lower jaws (Fig. 32G, H). Micropogonias, Paralonchurus, and Pogonias have this pattern. The arrangement of the barbels differs among the genera, especially in Paralonchurus. Three pairs of minute barbels form a tuft situated laterally to the me- dian pore, and 12 to 16 pairs of small barbels distributed along the rami of the lower jaw (Fig. 32H). Micro- pogonias and Pogonias lack the tufts of barbels at the symphysis of the lower jaw (Fig. 32G). Fishes with this pore and barbel pattern have an inferior mouth, and are typically bottom feeders. Generic groupings based on external morphology. — Western Atlantic sciaenids can be readily separated into two groups by the presence or absence of mental barbels (Fig. 30). The group without barbels can be further di- vided by mouth position and body shape (Fig. 30). Cynos- cion, Isopisthus, and Macrodon differ from other sciae- nids without barbels in having an elongate body, an oblique mouth with a pair of enlarged canines at the tip of the upper jaw, and a protruding lower jaw (Fig. 37A). Those species that lack canines can be divided into Figure 37.— Four typical mouth positions and dentitions of western Atlantic Sciaenidae. A, Macrodon ancylodon; B, NebrU microps; C, Odontoscion dentex; D, Micropogonias undulatus. 25 groups by their mouth positions and body shapes (Fig. 30). Nebris and Larimus have a very large and oblique mouth (Fig. 37B). Bairdiella and Odontoscion both have a terminal mouth (Fig. 37C). The genus Stellifer consists of species with oblique (S. lanceolatus), subterminal (S. rastrifer), and inferios (S. microps), mouths (Fig. 38). Ophioscion, Leiostomus, Pareques, Equetus, and Sciae- nops have inferior mouths. Sciaenops differs from these fishes by having an elongate body. Sciaena is character- ized by a nearly terminal and horizontal mouth. Sciaenids with mental barbel(s) usually have an in- ferior mouth and can be divided into different categories by the number and/or position of barbels (Fig. 30). Ctenosciaena has a thin tapered barbel and a subter- minal mouth (Fig. 36A), unique among western Atlantic sciaenids with barbels. Menticirrhus and Umbrina have an apical pore at the tip of the single rigid barbel (Fig. 36B. C). Lonchurus has a pair of long slender barbels at the symphysis of the lower jaw (Fig. 32F). Paralon- churus has 3 pairs of minute barbels in a tuft at the sym- physis of the lower jaw lateral to the median mental pore and 12 to 16 pairs of small barbels along the rami of the lower jaw (Fig. 32H). Micropogonias has 3 to 4 pair of small mental barbels and Pogonias has 12 to 13 pairs that never form a tuft at the symphysis of the lower jaw (Fig. 32G). The combination of morphological characters describ- ed above reflects habitat or similarities in feeding habits (Chao and Musick 1977). The genera listed at the top of Figure 30 generally feed on the upper water to mid- water column and the genera listed near the bottom feed in the lower water column to the bottom. Within the dif- ferent patterns of snout pores, mental pores, and barbels of western Atlantic Sciaenidae, three different feeding modes may be recognized (Fig. 30). The midwater to up- per water column feeders have only marginal pores (two or five) on the snout, and have zero to four mental pores without barbels on the lower jaw. The midwater to lower water column feeders have five marginal pores and two to five upper pores on the snout, and four to five mental pores without barbel(s) on the lower jaw. The third group, bottom feeders, have five marginal and three to Figure 38. — Three levels of mouth position in the species of Stellifer: A, Stellifer lanceolatus, oblique; B, S. rastrifer, subterminal; C, S. microps, inferior. five (or more) upper pores on the snout, and have four to five mental pores and one to many barbels on the lower jaw. The number of pores and barbels seems to increase in feeding niches that are closer to the bottom. Specific characters. — Western Atlantic sciaenids with two marginal snout pores and with neither upper pores nor mental pores (Fig. 31A) such as Cynoscion are upper water-midwater column feeders. Although Nebris has a very oblique mouth (Fig. 37B) and snout pore ar- rangement similar to Cynoscion, it also has four mental pores (Fig. 34A) and a rounded body with a flat ventral surface. This suggests that Nebris is a bottom dweller feeding from the bottom upward. Sciaenid fishes with five marginal pores and no upper pore on the snout are midwater to upper midwater column feeders, such as Larimus, which has four mental pores similar to Nebris (Fig. 34B). Larimus, however, has a compressed body that may not be adapted to a bottom habitat as in that of Nebris. The pattern of five marginal pores and three upper pores (Figs. 31C, 33A-D) is common among different genera, such as Bairdiella, Odontoscion, Ophioscion, Stellifer, and Menticirrhus. Species of Menticirrhus all have four pores and one barbel with an apical pore on the lower jaw, and are bottom feeders. Although the number of mental pores among the species of Bairdiella varies from five to six, they all have a terminal mouth and are midwater to lower midwater feeders. Species of Stellifer not only have a variable number of mental pores, from four to six (Fig. 35), but also have different mouth posi- tions, from inferior to slightly oblique (Fig. 38). The species with a slightly oblique mouth, such as S. sp. B (see p. 55, key), S. stellifer, and S. rastrifer, are appar- ently midwater column feeders. But each species has a different number of mental pores (Fig. 35): S. sp. B four, S. stellifer five, and S. rastrifer six. Therefore, their feed- ing habitats are probably correlated with the positions of their mouths. In addition, other genera of western Atlan- tic sciaenids, Ctenosciaena, Paralonchurus , and Sciaena, also have a similar snout pore pattern. Among them, Ctenosciaena and Sciaena both have terminal mouths and probably feed in midwater down to the bottom. Ctenosciaena also has one barbel and four pores on the lower jaw (Fig. 36A). Based on these variations, the three 26 upper and five marginal snout pore pattern is probably the more generalized among western Atlantic sciaenids with different feeding niches. Sciaenid ilshes with five or more upper and marginal pores (Figs. 31D, 33E-H) are mostly bottom feeders with an inferior mouth. They usually have five pores on the lower jaw with or without mental barbels. Lonchurus has four pores and two long tapered barbels on the lower jaw (Fig. 32F). When comparing the pore and barbel systems of western Atlantic Sciaendae with the Indo-Pacific (Chu et al. 1963), East Atlantic, and East Pacific sciaenids, the five marginal and three to five upper pores on the snout seem to be generalized in most sciaenids. Also the five pores (or six pores with five openings) on the lower jaw is typical in sciaenids. Sciaenids with a single mental bar- bel are found in the Indo-West Pacific, on both sides of the Atlantic, and in the eastern Pacific (Trewavas 1964). However, sciaenids with more than one barbel are found only along the Atlantic and Pacific coasts of America, ex- cept that Daysciaena albida (Cuvier) of the Indian Ocean has a pair of short barbels. The external characters described above are some- times variable within a genus and yet similarities are found among different genera. Neither the differences nor the similarities necessarily reflect phylogenetic rela- tionships. These external morphological characters and those mentioned in the key may support the groupings on other criteria. Their usefulness in assessing relation- ships of western Atlantic sciaenids is usually not above the species level. A SYNOPSIS OF THE SUPRAGENERIC GROUPS, GENERA, AND SPECIES OF WESTERN ATLANTIC SCIAENIDAE Diagnoses and Primary Synonymy of the Suprageneric Groups, Genera, and Species Based on the structural patterns of swim bladder, otolith, and pore and barbel systems, western Atlantic sciaenids can be grouped into 11 suprageneric groups: Sciaena, Umbrina, Larimus, Sciaenops, Cy- noscion, Micropogonias, Lonchurus, Menticirrhus, Neb- ris, Pogonias, and Stellifer groups. The South American freshwater sciaenid genera Pachypops and Pachyurus are not included in these groups, due to lack of comparative material. The taxonomic position of these 11 supra- generic groups is between genus and family (between A and D, Fig. 4). Tribes and subfamilies are not utilized for western Atlantic Sciaenidae until comparisons can be made with sciaenids of other regions. The following descriptions of western Atlantic sciaenid groups are arranged in order from simple to more com- plex swim bladder patterns. In the description of each group, the diagnoses of the suprageneric group and in- cluded genera are given and the species are listed for each genus. A primary synonymy is given for each genus and species for nomenclatural purposes. The generic syn- onymy relies heavily on Jordan (1917-1923) and Norman (1966). Available type-specimens are listed in the syn- onymy of each nominal species. Where no type- specimens are listed, I am unaware of the existence or location of any. Sciaena GROUP Diagnosis. — Swim bladder in a simple carrot-shape, without diverticulum (Sciaena pattern, Fig. 5); sagitta more or less oval and thick (Fig. 18), lapillus not en- larged; snout with five marginal pores and three to seven upper pores (Figs. 31C, D; 33H); lower jaw with five pores and no barbels (Fig. 32C). There are three genera in this group: Equetus, Paraques, and Sciaena. Leiostomus has similar diagnostic characters except that the sagitta of Leiostomus is thinner and narrower (Fig. 18D). However, Leiostomus is probably more closely related to the Sciaena group than to any other western Atlantic sciaenid groups. Equetus Rafinesque Eques Bloch 1793, pt.7:90 (type-species: Eques ameri- canus Bloch, by monotype = Chaetodon lanceolatus Linnaeus, preoccupied by Eques Linnaeus 1758:459, a genus of lepidopteran insects). Equetus Rafinesque 1815:86 (substitute name for Eques, therefore taking the same type-species: Eques ameri- canus Bloch). Diagnosis. — Body oblong, compressed, back much ele- vated, rapidly tapering to a narrow caudal peduncle; sides with broad oblique bands; anterior portion of dor- sal fin very long and filamentous. Mouth small and in- ferior, teeth in villiform bands; gill rakers few, short, and slender. Vertebrae 10 + 15 = 25; no free interneurals an- terior to first dorsal fin (McPhail 1961). Swim bladder simple, carrot- or bottle-shaped (Fig. 5C). Sagitta oval, the expanded part of the ostium of sulcus reaching to an- terior margin (Fig. 18B, C); snout with five to seven up- per pores; lower jaw without barbel. Endemic to the tropical western Atlantic. Two species: E. lanceolatus and E. punctatus. Mainly inhabit coral reefs, sometimes shallow coastal waters. Equetus lanceolatus (Linnaeus) "Ribband Fish" Edwards 1751, pi. 210, middle figure, Caribbean Islands (nonbinominal). Chaetodon lanceolatus Linnaeus 1758:277, Caribbean Islands (after Edwards, pi. 210). "Serranas" Parra 1787, pi. 2, upper figure, Cuba (non- binominal). Eques americanus Bloch 1793, pt. 7:91, pi. 347, fig. 1, West Indies. 27 Eques balteatus Cuvier 1829:175 (after Edwards and Bloch). Sciaena edwardi Gronovius (Gray) 1854:53, (West) In- dian Sea (after Edwards). Equetus punctatus (Bloch and Schneider) "Sen-anas" Parra 1787, pi. 2, lower figure, Cuba (non- binominal). Eques punctatus Bloch and Schneider 1801:105, pi. 3, fig. 2 (based on Parra). acuminatus and E. umbrosus can be separated by their pigmentation pattern and counts (also see "Key to the species of Pareques"). The sagittae of these two species differ from each other by features of the sulcus (Fig. 18E, F). Paraques umbrosus is a more northerly species distributed from Chesapeake Bay to Florida and also recorded from Texas over sandy mud bottom of inshore and offshore waters; P. acuminatus is a more southerly species from Florida to Brazil in coral reef and rocky bot- tom coastal waters. Sciaena Linnaeus Pareque8 Gill Pareques Gill in Goode 1876:50 (type-species: Gram- mistes acuminatus Bloch and Schneider 1801, by original designation). Diagnosis. — Body oblong, compressed, anterior profile steep, back tapering rapidly to a narrow caudal pedun- cle; sides with longitudinal stripes; anterior portion of dorsal fin not filamentous. Mouth small, inferior; teeth in villiform bands; gill rakers short and blunt. Vertebrae 10 + 15 = 25; three free interneurals anterior to first dor- sal fin (McPhail 1961). Swim bladder simple, carrot- shaped. Sagitta oval, the expanded part of the ostium reaching to anterior margin (Fig. 18E, F), the cauda long, J-shaped. Snout with five to seven upper pores (Fig. 33H); lower jaw without barbel. Tropical eastern Pacific and western Atlantic, about seven species. Three western Atlantic species: P. acuminatus, P. umbrosus, and one undescribed. Inhabits sandy and muddy bottoms of high salinity waters and coral reefs. Pareques acuminatus (Bloch and Schneider) "Chaetodon, lineis fuscis, . . . ." Seba 1761:72, pi. 26, fig. 33, no locality (nonbinominal). Grammistes acuminatus Bloch and Schneider 1801:184 (after Seba). Eques lineatus Cuvier 1830:169, Brazil (syntypes: MNHN 7475, 135 mm SL; MNHN 43, 137 mm SL). Eques pulches Steindachner 1867:349, Barbados (type: presumably in NHMV, but not found). Sciaena Linnaeus 1758:228 (type-species: Sciaena umbra Linnaeus, by subsequent designation of International Commission for Zoological Nomenclature 1972, Opin- ion 988). Diagnosis. — Body elongate, compressed, dorsal profile slightly elevated or evenly arched; anterior portion of dorsal fin not elongated. Mouth horizontal to inferior; teeth in bands; gill rakers short and slender, widely spac- ed. Vertebrae 25 (11 + 14 in S. bathytatos and 10 + 15 in S. trewauasae). Swim bladder simple, carrot-shaped (Fig. 5A). Sagittae oval, the expanded part of the ostium reaching to anterior margin (Fig. 18G, H). Snout with three to five upper pores (Fig. 31C, D); lower jaw without barbel. Two species in the western Atlantic, S. bathy- tatos and S. trewauasae, inhabiting deeper subtropical waters, and one species in the eastern Atlantic and Mediterranean, S. umbra. Some of the six East Pacific nominal species of Sciaena probably should be assigned to other genera (Chao). 4 Remarks. — The nomenclature of the genus Sciaena was not stabilized until S. umbra Linnaeus 1758 was desig- nated as the type-species of the genus under the Plenary Powers of the International Commission for Zoological Nomenclature (1972, Opinion 988). A neotype was desig- nated by Trewavas (1966). The history of the case and synonymies were discussed by Trewavas (1963). This genus is the type-genus of the tribe Sciaenini (Trewavas 1962). The genus Sciaena is still not well defined and it may contain more species than currently recognized (Chao and Miller 1975). Pareques umbrosus (Jordan and Eigenmann) Eques acuminatus umbrosus Jordan and Eigenmann 1889:440, Charleston, S.C. (holotype: CAS 18958, 162 mm SL or USNM 25981, 165 mm SL, see remarks). Sciaena bathytatos Chao and Miller Sciaena bathytatos Chao and Miller 1975:267, fig. 9, Caribbean Sea off Colombia (holotype: USNM 211514, 208 mm SL). Remarks. — Jordan and Eigenmann (1889) described the subspecies E. acuminatus umbrosus based on a specimen from Charleston sent by C. Leslie. Both specimens of CAS 18958 and USNM 25981 were from the same source and fit the original description. Both specimens were also previously labeled as E. acuminatus. At present, I am not able to decide which one is the holotype. Eques Sciaena trewauasae Chao and Miller Sciaena trewauasae Chao and Miller 1975:262, fig. 8, 'Chao, L. N. A basis for classifying eastern Pacific Sciaenidae (Tele- ostei: Perciformes). Manuscr. 28 Caribbean Sea off Colombia (holotype: USNM 211513, 155 mm SL). Leiostotnus Lacepede Snout with three upper pores (Fig. 31C). Eastern Pacific and western Atlantic, two species. The western Atlantic species, C. gracilicirrhus, inhabits offshore waters over hard sandy bottom. Leiostomus Lacepede 1803, 4:439 (type-species: Leio- stotnus xanthurus Lacepede, by monotypy). Liostomus Gill 1862c:93 (invalid emendation of Leio- stomus Lacepede 1803, therefore taking the same type- species: Leiostomus zanthurus Lacepede). Diagnosis. — Body oblong, compressed, back elevated, sides with oblique stripes and a dark humeral spot behind upper end of gill slit; anterior portion of dorsal fin not elongated. Mouth small and inferior; teeth in villi- form band; gill rakers short and slender. Vertebrae 10 + 15 = 25. Swim bladder simple carrot-shape (Fig. 5B). Sagitta oval and thin, the expanded part of the ostium reaching to anterior margin and its axis inclined toward the dorsal margin (Fig. 18D). Snout with five upper pores; lower jaw without barbel. Northwestern Atlantic. Monotypic: L. xanthurus. Found over sandy and mud bottoms in estuarine and inshore waters. Leiostomus xanthurus Lacepede Leiostomus xanthurus Lacepede 1803, 4:439, pi. 10, fig. 1, Carolina (syntypes: MNHN 7599, two specimens, 148 and 163 mm SL). Mugil obliquus Mitchill 1814:16, New York. Sciaena multifasciata LeSueur 1822:255, east Florida (length of the type-specimen not reported). Leiostomus humeralis Cuvier 1830:141, pi. 110, New York (holotype: MNHN 9733, one dried stuffed speci- men, 146 mm SL). Umbrina GROUP Diagnosis. — Swim bladder simple carrot-shaped, without diverticula (Sciaena pattern, Fig. 5); sagitta oval (Sciaena pattern, Fig. 18); snout with five mar- ginal pores and three to seven upper pores (Fig. 31C, D); lower jaw with four pores and one barbel, an apical pore present in Umbrina. Two genera in this group: Cteno- sciaena and Umbrina. Ctenosciaena Fowler and Bean Sciaena (Ctenosciaena) Fowler and Bean 1924:15 (type- species: Sciaena dubia Fowler and Bean, by mono- typy). Diagnosis. — Body oblong, moderately compressed, head moderate and blunt with well-developed cavernous canals. Mouth subterminal to inferior, snout protrud- ing; lower jaw with a barbel tapering to a fine point without an apical pore at the tip (Fig. 36A). Vertebrae 10 + 15 = 25. Sagitta oval, the expanded part of the ostium slightly in contact with the anterior margin (Fig. 18A). Ctenosciaena gracilicirrhus (Metzelaar) Umbrina gracilicirrhus Metzelaar 1919:72, fig. 24, Vene- zuela (holotype: ZMA 113.103, 105 mm SL). Sciaena (Ctenosciaena) dubia Fowler and Bean 1924:16, Wilkes Exploring Expedition, locality unknown (holo- type: USNM 83309, 111 mm SL). Umbrina Cuvier Sciaena Linnaeus 1758:288 (of Artedi 1738, in part; in- cludes S. cirrosa; type-species: Sciaena umbra Lin- naeus by subsequent designation of International Commission for Zoological Nomenclature 1972, Opin- ion 988). Umbrina Cuvier 1817:297 (type-species: Sciaena cirrosa Linnaeus, by original designation). Attilus Gistel 1848:109 (type-species: Sciaena cirrosa Linnaeus, by original designation). Diagnosis. — Body moderately elongate; back slightly arched, ventral profile nearly straight. Head oblong, snout thick and protuberant. Mouth small, horizontal to inferior; teeth in bands, outer row of teeth in upper jaw moderately enlarged. Gill rakers short. Vertebrae 11 + 14 = 25. Swim bladder simple carrot-shape, without diver- ticula. Sagitta oval, the expanded part of the ostium reaching to anterior margin (Fig. 18 I, J). Snout with five to seven upper pores (Fig. 31D); lower jaw with four pores and a short, thick barbel, the latter with an apical pore (Fig. 36C). Eastern Pacific and both sides of the Atlantic, about nine species. Four species in the western Atlantic: U broussonetii, U. canosai, U. coroides, and U. milliae. Inhabits inshore and offshore waters with sandy or mud bottoms and around coral reefs. Remarks. — The name Chromis in the footnote of Lacepede 1802, volume 3, p. 546 was from the sketch of Le Pere Plumier, a missionary in Martinique, for the "grygry" or "grogro," which is Umbrina coroides Cuvier 1830. The name Chromis is older than Umbrina but later than Cromis Browne (after Jordan 1917). Also see Gil- bert (1966) for more detailed synonymy. Umbrina broussonetii Cuvier Umbrina broussonetii Cuvier 1830:187, Jamaica (holo- type: MNHN 7471, 151 mm SL). Umbrina canosai Berg Umbrina canosai Berg 1895:56, Montevideo, Uruguay to Mar del Plata Argentina (no type-specimens nor speci- fic localities mentioned in the original description). 29 L'mbrina coroides Cuvier Umbrina coroides Cuvier 1830:187, pi. 117, Brazil (holo- type: MNHN 5343. 174 mm SL). Umbrina milliae Miller Umbrina milliae Miller 1971:303, fig. 1, Colombia (holo- type: USNM 204932, 193 mm SL). Menticirrhus GROUP Diagnosis. — Swim bladders atrophy as the fish grow, only a vestige remains in adult. Drumming muscles ab- sent in both sexes. Sagitta oval, elongate, and relatively small, the expanded part of the ostium not reaching to anterior margin {Menticirrhus pattern, Fig. 22B). Snout with three upper pores and five marginal pores, rostral fold deeply indented (Fig. 33D); lower jaw with a short, rigid barbel, knoblike and five pores, the median at tip of barbel (Fig. 36B). Anal fin with only one weak spine. One genus in this group: Menticirrhus. Menticirrhus Gill Menticirrhus Gill 1862b:86 (type-species: Perca alburnus Linnaeus = Cyprinus americanus Linnaeus, by origi- nal designation). Cirrimens Gill 1863:17 (type-species: Umbrina ophio- cephalus Jenyns 1842:45, by original designation). Menticirrhus (Umbrula) Jordan and Eigenmann 1889: 424 (type-species: Umbrina littoralis Holbrook, by subsequent designation of Jordan and Evermann 1898: 1469). Diagnosis. — Body elongate, rounded; dorsal profile slightly arched, ventral profile nearly flat. Head conical, low and broad, snout projecting. Mouth small and in- ferior; teeth in villiform bands. Gill rakers short and tuberclelike. Vertebrae 10 + 15 = 25. Other diagnostic characters as in the group diagnosis. Tropical and tem- perate eastern Pacific and western Atlantic, nine species. Three species in the western Atlantic: M. americanus, M. littoralis, and M. saxatilis. Inhabits shallow coastal waters with sandy bottoms, juveniles often found in es- tuaries. Menticirrhus americanus (Linnaeus) "Alburnus Americanus" Catesby 1743, "Carolina whit- ing," pi. 12, lower figure, Charleston (pre-Linnaean and nonbinominal, see remarks). Cyprinus americanus Linnaeus 1758:321 (based on Catesby; not Cyprinus americanus of Linnaeus 1766: 530, a cyprinid, see remarks). Perca alburnus Linnaeus 1766:482, South Carolina (holo- type: Linnaean Society of London, no. Ill, a dried left side of skin, 282 mm TL, see remarks). Umbrina arenata Cuvier 1830:190, Brazil (syntypes: MNHN 7500, 76.4 mm SL; MNHN 7472, a dried and stuffed specimen, in ethanol, 341 mm TL). Umbrina martinicensis Cuvier 1830:186, Martinique (holotype: MNHN 7498, 193 mm SL). Umbrina gracilis Cuvier 1830:189, Brazil (lectotype: MNHN 44, 156 mm SL, see remarks). Sciaena alburnus (non-Linnaeus) Gronovius (Gray) 1854:108 (Gronovius' specimen; BMNH 1853.11.12.75, a dried skin, 228 mm TL, see remarks). Umbrina phalaena Girard 1859:167, Brazos Santiago, Texas (syntypes: USNM 154721, 88.3 mm SL; other syntypes not located). Umbrina januaria Steindachner 1877:170, Rio de Janeiro (type: not located). Remarks. — The description of "Alburnus Americanus" by Catesby (1743) stated that "under the lip having five or six fleshy barbels, resembling teeth hanging to it on the outside. It had one small fin on the middle of the back." The lower figure of plate 12, "Alburnus" agrees with this description. Except for these features, the general appearance of the figure is a species of Men- ticirrhus but it is impossible to identify it with any of the Menticirrhus species. The name Cyprinus americanus Linnaeus 1758 is based on Catesby. But in Linnaeus 1766, the 12th edition of "Systema Naturae," he treated the Catesby reference as a synonym of Perca alburnus which is based on the D. Garden skin (Linnaean Society of London, no. Ill, 282 mm TL). He also employed the name Cyprinus americanus (= Notemigonus bosci Cuvier and Valenciennes, from Jordan and Eigenmann 1889) within the Cyprinidae on page 530 (1766) based on Garden material but with no reference to his 1758 usage or to Catesby. Linnaeus (1766) appeared to be acknowledging that he had been misled by the plate (Catesby 1743, "Alburnus Americanus," pi. 12) and was clarifying his intentions both for the name C. americanus and for the binomen to use for the sciaenid (Perca al- burnus). A. Wheeler (BMNH, pers. commun.) pointed out that Linnaeus was acting in effect as "first reviser" for Cyprinus americanus and also for Perca alburnus and suggested that M. alburnus (Linnaeus 1766) should be the correct name for this species. Although the earlier usage was in favor of M. alburnus (prior to Jordan and Eigenmann 1889), I am in favor of the currently stabiliz- ed usage of M. americanus. Furthermore, Jordan and Eigenmann (1889) stated that Cyprinus americanus was the name Linnaeus gave to the whiting of Catesby and that there could be no doubt that the fish Catesby had in mind was Menticirrhus americanus. A specimen, MNHN 9037, was catelogued as syntype of Umbrina gracilis Cuvier 1830, in MNHN, Paris and was identified by Irwin (1970) as M. littoralis (Hol- brook). It is not a type of U. gracilis, because the locality and the donor of the specimen do not agree with the original description of Cuvier (M. Bauchot, MNHN, Paris, pers. commun.). Thus, MNHN 44, is selected as lectotype. Wheeler (1958) wrote "Sciaena alburnus (non-L.) 30 Gronovius (Gray), 1854" as a synonym of Menticirrhus americanus (Linnaeus 1758). This also implied that S. alburnus Gronovius (Gray) was not intended to be Perca alburnus Linnaeus 1766. In the original description of Sciaena alburnus, Gronovius (Gray) 1854, quoted "Perca pinnis dorsalibus distinctis muticis, . . . ." from Lin- naeus (1766), 12th edition genus 168, no. 6. According to Wheeler (1958), a figure of S. alburnus and a specimen (BMNH 11.12.75, a dry skin, 288 mm SL) with Garden's collection number were present in Gronovius' manu- script. Linnaeus (1766) also wrote Perca alburnus as "Habitat in Carolina, D. Garden, Whiting." Thus, in my opinion, Sciaena alburnus Gronovius (Gray) was in- tended to be Perca alburnus Linneaus. on first gill arch. Vertebrae 11 + 14 = 25. Other diagnos- tic characters as in the Larimus group. Tropical eastern Pacific and western Atlantic, about seven species. Two western Atlantic species: L. breviceps and L. fasciatus. Inhabits estuarine and inshore open waters with sandy and muddy bottoms. Larimus breviceps Cuvier Larimus breviceps Cuvier 1830:146, pi. 140, Saint Domi- nique and Brazil, (syntypes: MNHN 7578, 139 mm SL, St. Dominique; MNHN 7636, 156 mm SL, Brazil). Monosira stahli Poey 1881:326, pi. 6, Puerto Rico (holo- type: 190 mm long, not located). Menticirrhus littoralis (Holbrook) Vmbrina littoralis Holbrook 1855:142, South Carolina (holotype: ANSP 11567, 198 mm SL). Larimus fasciatus Holbrook Larimus faciatus Holbrook 1855:153, pi. 22, fig. 1, Charleston (type presumably lost). Menticirrhus saxatilis (Bloch and Schneider) Johnius saxatilis Bloch and Schneider 1801:75, New York (holotype: HU 8792, Humboldt-Universitat, Ber- lin, a stuffed skin in poor condition, according to Irwin 1970). Sciaena nebulosa Mitchill 1815:408, pi. 3, New York. Menticirrhus focaliger Ginsburg 1952:97, St. Joseph Bay, Fla. (holotype: USNM 144101, 104 mm SL). Menticirrhus atlanticus McFarland 1963:98, Mustang Island, Texas (nomen nudum). Larimus GROUP Diagnosis. — Swim bladder carrot-shaped, with or without a pair of small tubelike diverticula at the an- terior end (Sciaena pattern, Fig. 5E, F). Sagitta with broader anterior portion and narrower posterior portion, the expanded part of the ostium not reaching to anterior margins (Larimus pattern, Fig. 20A, B). Snout with five marginal pores and no upper pores (Fig. 31B), lower jaw with four pores and no barbel (Fig. 34B). Larimus is the only genus in this group. Larimus Cuvier Larimus Cuvier 1830' 145 (type-species: Larimus brevi- ceps Cuvier, by monotypy). Amblyscion Gill 1864b:165 (type-species: Amblyscion argenteus Gill, by monotypy). Monosira Poey 1881:326 (type-species: Monosira stahli Poey, by monotypy). Diagnosis. — Body short and robust, compressed; dorsal profile slightly elevated, ventral profile convex. Head with prominent cavernous canals; snout short. Mouth large and oblique, teeth small, conical in one to two rows on both jaws. Gill rakers, long and slender, more than 25 Nebris GROUP Diagnosis. — Swim bladder with carrot-shaped main chamber and a pair of diverticula, which almost extend to the tapering end of the main chamber then loop back (Fig. 8) reaching the septum transversum anteriorly (Nebris pattern). Sagitta oval, with a notched posterior margin, very thick (Fig. 21); ostium large, its expanded part not reaching to anterior margin; cauda J-shaped, with broad and deeply grooved distal end; snout with only two marginal pores, without upper pores; lower jaw with four minute pores and no barbel. One genus in this group: Nebris. Nebris Cuvier Nebris Cuvier 1830:149 (type-species: Nebris microps Cuvier, by monotypy). Cheironebris Fowler 1944:179 (type-species: Nebris oc- cidentalis Vaillant, by monotypy). Diagnosis. — Body elongate, rounded in cross section, tapering posteriorly. Head extremely cavernous, interor- bital space very broad, eye very small, more than 10 times in head. Mouth very large, oblique, nearly ver- tical; teeth small, conical in narrow bands on upper jaw, and two rows of small teeth in lower, the outer row slight- ly smaller than the inner. Gill rakers long and slender. Vertebrae 12 + 13 = 25. Other diagnostic characters as listed for the group. Tropical eastern Pacific and western Atlantic, two species. One species in the western Atlan- tic, N. microps. Found over sandy to muddy bottoms in coastal and estuarine waters. Nebris microps Cuvier Nebris microps Cuvier 1830:149, pi. 312, Surinam (type: in Berlin Museum, not examined). 31 Micropogonias GROUP Micropogonias undulatus (Linnaeus) Diagnosis. — Swim bladder carrot-shaped with a pair of tubelike diverticula extending anteriorly from the mid- dle of sides to septum transversum. (Micropogonias pat- tern. Fig. 9.) Sagitta very heavy, irregular in shape (Micropogonias pattern, Fig. 26), the expanded portion of the ostium not reaching to anterior margin, the cauda oblique with an expanded distal end. Snout with five marginal pores and five to seven upper pores (Fig. 31D); lower jaw with five pores and three to five pairs of minute barbels (Fig. 32G). One genus in this group, Micro- pogonias. Micropogonias Bonaparte Micropogon Cuvier 1830:213 (type-species: Micropogon linaetus Cuvier, in part = Um brina furnieri Desmarest, see remarks; preoccupied by Micropogon Boie 1826: 977, Aves). Micropogonias Bonaparte 1831, 52:170 (substitute name for Micropogon, therefore taking the same type-species, M. lineatus Cuvier in part = U. furnieri Desmarest, see remarks). Micropogonoides Trewaves (1977) 33:271 (lapsus cala- mi). Diagnosis. — Body elongate, dorsal profile elevated, ven- tral profile nearly straight. Head conical, preopercular margin with 10 to 14 spines, 2 to 5 strong spines at the angle. Mouth inferior, teeth in villiform bands, outer row teeth in upper jaw slightly enlarged. Gill rakers short. Vertebrae 10 + 15 = 25. Other diagnostic characters as listed for the group. Tropical eastern Pacific and western Atlantic, about five species. Two species in the western Atlantic: M. furnieri and M. undulatus. Inhabits es- tuarine rivers and coastal waters with sandy to muddy bottom. Micropogonias furnieri (Desmarest) Umbrina furnieri Desmarest 1823:182, pi. 17, fig. 3, Ha- vana, Cuba (presumed holotype: MNHN 4968, 109 mm SL, see remarks). Micropogon lineatus Cuvier 1830:215, pis. 119 and 138, in part, swim bladder (lectotype: MNHN 4968, 109 mm SL, Cuba; paralectotype: MNHN 7459, 159 mm SL, Montevideo, see remarks). Micropogon argenteus Cuvier 1830:218, Surinam (holo- type: RMNH 685, 300 mm SL, see remarks). Corvina crawfordi Regan 1903:627, Montevideo (holo- type: BMNH 1903.10.16.7, 210 mm SL, see remarks). Micropogon patagonensis MacDonagh 1931:409, Richo de San Bias, Argentina (holotype: Museo de la Plata, no. 2. V. 31.1, 465 mm SL, not examined, see remarks). Micropogon barretoi MacDonagh 1934:70, Punta Piedras, Argentina (holotype: Museo de la Plata, no. 12. VII. 33.15, 423 mm SL, not examined, see remarks). Ophioscion woodwardi Fowler 1937:311, figure on p. 312, Haiti (holotype: ANSP 68257, 108 mm SL). "Perca marina" Catesby 1743, "Croker," pi. 3, upper figure, Chesapeake Bay (pre-Linnaean and nonbi- nominal). Perca undulata Linnaeus 1766:483, South Carolina (syn- types: the Linnaean Society of London no. 112, right side of a dried skin, 212 mm SL, and no. 113, left side of a dried skin, 241 mm SL). Sciaena croker Lacepede 1803, 4:309, Carolina (type: presumably in MNHN, but not found). Bodianus costatus Mitchill 1815:417, New York. Sciaena opercularis Quoy and Gaimard 1824:347, baie de Rio de Janeiro (type: reported missing by Roux 1973, also see remarks). Micropogon lineatus Cuvier 1830:215, in part (paralecto- type: MNHN 7457, two specimens, 62.8 and 138 mm SL, New York; MNHN 7459, 273 mm SL, Montevideo, also see remarks). Remarks. — The fish genus name Micropogon Cuvier 1830 is preoccupied by Micropogon Boie 1826, Aves. Al- though the bird genus name has only been used in syno- nymys in recent years, the fish genus name has gained wide and stable used since Cuvier 1830. C. G. Gruchy (NMC) and the author had looked into this matter and found the inherent nomenclatural and systematic prob- lems of the bird genus and its family have not been solved. To avoid further complications, we decided to resume the valid substitute name Micropogonias (Bona- paret 1831) for fish genus. Jordan (1917) designated M. lineatus Cuvier = Perca undulatus Linnaeus as type-species of Micropogonias. His reason was that Cuvier's (1830) description on p. 215 mentioned the New York specimens (MNHN 7457, M. undulatus) first. Cuvier's (1830) description of M. lineatus fits both M. furnieri and M. undulatus. The syn- types obtained from New York (MNHN 7457) by Mil- bert are apparently M. undulatus (there is also a specimen of Leiostomus xanthurus, 87.2 mm SL, in the same jar). The syntype from Cuba (MNHN 4968) was labeled as "type de Umbrina fournieri Desmarest," and the specimen illustrated on plate 119 appears to be M. furnieri. The oblique streaks of M. undulatus usually are not as distinct, particularly the area between the soft portion of dorsal fin and lateral line. MNHN 7459 from Montevideo contains two syntypes, of which the smaller specimen (159 mm SL) appears to be M. furnieri and the larger one (273 mm SL) appears to be M. undulatus. In this case, MNHN 4968 (109 mm SL) from Cuba is selected as the lectotype of M. lineatus and it is likely to be the holotype of Umbrina furnieri. Therefore, the type- species of Micropogonias is M. furnieri (Desmarest). Some morphometric and meristic characters of the presumed holotype, MNHN 4968, follow: Morphometric characters (in mm): TL 110; SL 109; snout to anal fin origin 72.6, to dorsal fin origin 40.5, to pectoral fin origin 34.7, to pelvic fin origin 36.0; max- imum depth 28.2; head length 33.6; snout length 9.7; 32 eye diameter 8.5; interorbital width 8.1; pectoral fin lengths 19.5 and 22.5; pelvic fin lengths 16.0 and 19.2; length of anterior (spinous) portion of dorsal fin base 20.2; length of posterior (soft) portion of dorsal fin base 42.7; length of anal fin base 12.5; length of second anal fin spine 13.4; depth of caudal peduncle 9.2. Meristic characters: D X+I, 27; A. II, 8; pectoral rays 18 and 17; lateral gill rakers on the first right gill arch 8 (upper arm) + 15 (lower arm); medial (or inner) gill rakers on the first right gill arch 3 (upper arm) + 10 (lower arm); pored lateral-line scales, left side 52, right side 48; transverse lateral-line scales, from posterior (soft) dorsal fin origin to lateral line 9 (vertical series) and 12 (oblique series), from lateral line to anal fin origin 12 (vertical series) and 15 (oblique series). The distinguishing characters of the two species oi Micropogonias (see also "Key to the species of Micropogonias") used here are valid only for adult specimens. The sagitta morphology of M. furnieri from northern Brazil (Fig. 26A-C) is distinct from that of M. undulatus from Chesapeake Bay (Fig. 26D-F). But the sagittae of specimens which key out as M. undulatus from the Bay of Campeche, Mexico, are hardly dis- tinguishable from specimens keyed out as M. furnieri from the Caribbean coast of Colombia (Fig. 39). Other nominal species of this genus were discussed by Jordan and Evermann (1898), Devincenzi (1924), MacDonagh (1934), and Roux (1973). Among them Corvina crawfordi Regan, Micropogon patagonensis McDonaugh, and M. barretoi McDonaugh were described from the mouth of Rio de La Plata or south; morphometric characters were mainly used by the authors for diagnoses. Based on ob- servations of M. undulatus from Chesapeake Bay region, Virginia (Chao 1976), such morphological variation is fre- quent and not a valid basis for species diagnoses. The true identity of M. argenteus Cuvier and Sciaena oper- cularis Quoy and Gaimard is uncertain at present. Sixteen morphometric and meristic characters were compared in 22 specimens of southern specimens (M. furnieri) from Colombia to Uruguay and 24 specimens of the northern species (M. undulatus) from Chespeake Bay to Bay of Campeche. The result of discriminate function analyses indicated one of the Campeche specimens (NMC 76-522, 116 mm SL) was intermediate. All northern specimens can be readily separated from southern ones by color pattern and number of scales (6 to 7 in M. furnieri and 8 to 9 in M. undulatus) in a vertical series from dorsal fin origin to lateral line (also see sec- tion on "Key to the species of Micropogonias"). Whether they are species or subspecies of one species requires fur- ther study. I suggest that the northern species (or form) M. undulatus is present on Atlantic and Gulf of Mexico coasts of U.S.A., south to the Bay of Campeche, Mexico. The southern species (or form) M. furnieri is present in the Antilles and on South American coasts. Sciaenops GROUP Diagnosis. — Swim bladder carrot-shaped with a pair of short tubelike diverticula at the anterior lateral corners of the swim bladder (Sciaenops pattern, Fig. 11). In large specimens, a pair of "saclike" projections which consist of numerous labyrinth chambers (Fig. 11C, D) are pre- sent on the dorsal surface of the swim bladder; a pit is present between the third and fourth pleural ribs on both sides of the body wall to receive the paired "saclike" projections. Sagitta rectangular, the expanded portion of the ostium not reaching to anterior margin (Larimus pat- tern, Fig. 20). Snout with five upper and five marginal pores (Fig. 33F); lower jaw with five pores and no barbel (Fig. 34F). Sciaenops is the only genus in this group. Figure 39.— Sagittae of Micropogonias. A, A', a: M. undulatus, 205 mm TL, Bay of Campeche, Mexico; B, B , b: M. furnieri, 240 mm TL, coast of Caribbean Sea, Colombia. A, B: inner surface; A , B : outer surface; a, b: lateral view. 33 Sciaenop8 Gill Pogonias cromis (Linnaeus) Sciaenops Gill 1864a:30 (type-species: Perca ocellata Linnaeus, by monotypy). Diagnosis. — Body elongate, robust, back slightly elevated, ventral surface flat. Head long and low; snout long. Mouth inferior; teeth villiform in bands, those of outer row in upper jaw and inner row in lower jaw slightly enlarged. Gill rakers short. Vertebrae 10 + 15 = 25. Other diagnostic characters as listed for the group. Sub- tropical northwestern Atlantic. Monotypic: S. ocellata. Inhabits estuarine, surf zone and inshore waters with sandy bottoms. Sciaenops ocellata (Linnaeus) Perca ocellata Linneaus 1766:483, South Carolina (syn- types: Linnean Society of London, no. 106, one dried right side skin, 394 mm TL, and no. 107, one dried right side skin, 399 mm TL). Lutjanus triangulum Lacepede 1803, 4:181, figure in vol. 3, pi. 24, fig. 3 (type locality: Sumatra, an East Indian Island, is apparently an error; type specimen: presumably in MNHN but not found). Sciaena imberbis Mitchill 1815:411, New York. Pogonias GROUP Diagnosis. — Swim bladder with very complicated diver- ticula along the side of main chamber (Fig. 10). Sagitta semicircular, the "tadpole-shaped" sulcus not reaching to the margins (Fig. 19A). Snout with five upper pores and five marginal pores (Fig. 31D); lower jaw with five pores and 12 to 13 pairs of small barbels (Fig. 32G). Monotypic: Pogonias. Pogonias Lacepede Pogonias Lacepede 1802, 3:137 (type-species: Pogonias fasciatus Lacepede = Labrus cromis Linnaeus, by monotypy). Pogonathus Lacepede 1803, 5:120 (type-species: Pogona- thus courbina Lacepede = Labrus cromis Linnaeus, by subsequent designation of Jordan 1917:65). Diagnosis. — Body oblong and deep; dorsal profile ele- vated, ventral profile nearly straight. Head large; snout conical, not conspicuously projecting in front of upper jaw. Mouth moderate, horizontal, and slightly inferior; teeth in villiform band on both jaws. Lower pharyngeals very large, fully united, with coarse, molariform teeth (Fig. 40A, a). Gill rakers very short. Vertebrae 10 + 14 = 24. Complexity of swim bladder increases with size (Fig. 10); other diagnostic characters as in group diagnosis. Tropical and temperate western Atlantic. Monotypic: P. cromis. Inhabits coastal waters especially in areas with large river runoffs over sandy and sandy-mud bot- toms. Labrus cromis Linnaeus 1766:479, Carolina. Pogonias fasciatus Lacepede 1802, 3:137, figure in vol. 2, pi. 16, fig. 2, Brazil (syntypes: MNHN 7461, one dried stuffed specimen, 103 mm TL; MNHN 7460, one dried stuffed specimen, 181 mm SL). Pogonathus courbina Lacepede 1803, 5:120, Reio de La Plata (from the manuscript of Commerson). Mugil grunniens Mitchill 1814:15, New York. Mugil gigas Mitchill 1814:16, New York. Sciaena fusca Mitchill 1815:409, New York. Cynoscion GROUP Diagnosis. — Swim bladder with a pair of well-developed horns (Fig. 12) from anterolateral corners of main cham- ber, tapering forward to the back of skull. Sagitta elongate oval with smooth or notched margins. Snout with only two (or no) marginal pores and without upper pores (Fig. 31A); lower jaw without obvious pores or bar- bels (Fig. 32A). Four genera in this group: Cynoscion, Isopisthus, Macrodon, and a South American fresh- water genus, Plagioscion (see section on "Freshwater Sciaenidae of America"). Cynoscion Gill Cestreus Gronovius (Gray) 1854:49 (type-species: Ces- treus carolinensis, by monotypy, preoccupied by Ces- treus McClelland 1842). Cynoscion Gill 1872b:81 (type-species: Otolithus regalis Cuvier = Johnius regalis Bloch and Schneider, by original designation). Apseudobranchus Gill 1863:18 (type-species: Otolithus two-roe Cuvier = Cheilodipterus acoupa Lacepede 1800, by original designation). Archoscion Gill 1863:18 (type-species: Otolithus analis Jenyns 1842, by original designation). Cynoscion (Buccone) Jordan and Evermann 1896:394 (type-species: Cestreus praedatorius Jordan and Gil- bert in Jordan and Eigenmann 1889, by original desig- nation). Symphysoglyphus A. Miranda Ribeiro 1915, Sciaenidae: 43 (type-species: Otolithus bairdi Steindachner, by monotypy). Paralarimus Fowler and Bean 1924:18 (type-species: P. patagonicus = Cheilodipterus acoupa Lacepede, by monotypy). Cynoscion (Eriscion) Jordan and Evermann 1927:506 (type-species: Cynoscion nebulosus Cuvier, by mono- typy)- Diagnosis. — Body elongate, compressed, predorsal fin outline nearly straight, ventral evenly arched. Head con- ical, snout pointed. Mouth large, oblique, lower jaw pro- jecting; teeth sharp in narrow ridges, tip of upper jaw usually with a pair of large canines at least twice the size of other teeth (no obviously enlarged teeth in C. stein- dachneri), tapering from base to tip. Preopercular mar- 34 Figure 40.— United lower pharyngeal teeth of Pogonias cromis (A, a) and Aplodinotus grunniens (B, b). A, B: dorsal view (anterior end to the top of the figure); a, b: lateral view (anterior end to the left of the figure). gin membranous or ciliate. Gill rakers short to long and slender. Vertebrae usually 13(12) + 12(13) = 25, except 15+12 = 27 in C. nothus. Swim bladder with a pair of an- teriorly directed lateral horns, some straight and some curved (Fig. 13). Sagitta oval, the expanded part of the ostium usually not reaching to anterior margin in most species (Fig. 23); cauda usually short and expanded at the end. Eastern Pacific and western Atlantic, about 23 species. There are four northwestern Atlantic species: C. arenarius, C. nebulosus, C. nothus, and C. regalis; and eight Caribbean and southwestern Atlantic species: C. acoupa, C. jamaicensis, C. leiarchus, C. microlepidotus, C. similis, C. steindachneri, C. striatus, and C. vires- cens. Inhabits estuarine and inshore waters. Remarks.— In Klein (1749), p. 23 and 24, under "Ces- treus," species of "Mugil" were described and "Sciaena" was described in a footnote. Type-species of Sciaena: Mugil cephalus Linnaeus = "Cestreus dorso repando" Klein, was subsequently designated by Walbaum (1793), but was invalid because of Linnaeus' use of Sciaena in 1758. Cynoscion acoupa (Lacepede) Cheilodipterus acoupa Lacepede 1800, 2:540, Cayenne (holotype: MNHN 5502, 262 mm SL, Surinam). Lutjanus cayennensis Lacepede 1803, 4:196 (syntypes: MNHN 5502, 262 mm SL, Surinam; MNHN A.5617, two specimens, 522 and 567 mm SL, Martinique; MNHN A. 4562, two specimens, 278 and 288 mm SL, Brazil). Otolithus rhomboidalis Cuvier 1829:173, Cayenne (based on "Lutjan de Cayenne," Lacepede). Otolithus toe-roe Cuvier 1830:72, pi. 103 (syntypes: MNHN 5500, 175 mm SL, Surinam; MNHN 4616, 347 mm SL, Cayenne; MNHN A. 4518, two specimens, 493 and 485 mm SL, Cayenne). Paralarimus patagonicus Fowler and Bean 1924:18, Pat- agonia, Argentina (holotype: USNM 83222, 136 mm SL). Cynoscion maracaiboensis Schultz 1949:160, fig. 20, Maracaibo, Venezuela (holotype: USNM 12742, 251 mm SL). 35 Cyrwscion arenarius Ginsburg Cynoscion arenarius Ginsburg 1929:83, Texas (holotype: USNM 89385, 245 mm SL). Cynoscion jamaicensis (Vaillant and Bocourt) Otolithus jamaicensis Vaillant and Bocourt 1915:156, pi. 6, fig. 1, Jamaica (holotype: MNHN A. 557, 205 mm SL). Archiscion petranus A. Miranda Ribeiro 1915, Sciaeni- dae: 42, Brazil (type: not located and not listed in P. Miranda Ribeiro 1953). Cynoscion leiarchus (Cuvier) Otolithus leiarchus Cuvier 1830:78, Cayenne, Brazil (syntypes: MNHN 5503, two specimens, 152 and 234 mm SL; MNHN A.2690, 187 mm SL; MNHN A.5422, a dried stuffed specimen, 112 mm SL). Remarks. — The identity of the nominal species, Otolithus obliquatus (Valenciennes in Sauvage 1879), was discussed by Jordan and Evermann (1898) and Ran- dall and Cervigon (1968). The higher soft dorsal and anal ray counts in two type-specimens (selected by Randall and Cervigon 1968, MNHN 7632: lectotype 209 mm SL, D. X+I, 30, A. II, 12 and paralectotype 182 mm SL, D. X+I, 28, A. II, 11) were also noted in this study. The ver- tebrae total 25 in both specimens and the soft portion of their dorsal fins is covered with small cycloid scales beyond the basal half of the fins. These specimens (MNHN 7632) and two specimens of Leiostomus xan- thurus (MNHN 7600, 106 and 126 mm SL) of Cuvier (1830:143) were collected by M. Plee from Martinique. They are only known from the West Indies by these descriptions. Therefore, I suggest that 0. obliquatus is a synonym of C. regalis (Bloch and Schneider) and the type-locality is probably an error. Cynoscion similis Randall and Cervigon Cynoscion microlepidotus (Cuvier) Otolithus microlepidotus Cuvier 1830:79, Surinam (type: presumably in Berlin, not examined). Otolithus bairdi Steindachner 1879:40, pi. 1, fig. 2, Santos (presumed holotype: NHMV 51130, 152 mm SL, donated by Steindachner). Cynoscion nebulosua (Cuvier) Labrus squetaegue var. maculatus Mitchill 1815:398, New York (not Labrus maculatus Bloch 1793). Otolithus nebulosus Cuvier 1830:79, locality unknown (holotype: MNHN 7527, 233 mm SL). Otolithus carolinensis Valenciennes 1833:475, South Carolina (holotype: MNHN 7507, 335 mm SL). Otolithus drummondi Richardson 1836:70, New Orleans. Cynoscion nothus (Holbrook) Otolithus nothus Holbrook 1855:134, pi. 19, fig. 1, South Carolina (type: presumably lost). Cynoscion similis Randall and Cervigon 1968:170, fig. 2, Isla de Margarita, Venezuela (holotype: USNM 201382, 284 mm SL). Cynoscion steindachneri (Jordan) Cestreus steindachneri Jordan in Jordan and Eigenmann 1889:372, Curuga, Brazil (holotype: MCZ 10922, 318 mm SL). Cynoscion striatus (Cuvier) "Guatucupa" Marcgrave 1648 (translated by Taunay 1941):177, Brazil (nonbinominal). Otolithus striatus Cuvier 1829:173 (after Marcgrave). Otolithus guatucupa Cuvier 1830:75, Montevideo (syn- types: MNHN 7517, two specimens, 358 and 366 mm SL). Cynoscion virescens (Cuvier) Otolithus virescens Cuvier 1830:72, Surinam (type: pre- sumably in Berlin Museum, not examined). Otolithus microps Steindachner 1879:38 (type: presum- ably in NHMV, but not found). Cynoscion regalia (Bloch and Schneider) Johnius regalis Bloch and Schneider 1801:75, New York. Roccus comes Mitchill 1814:26, New York. Labrus squeteague Mitchill 1815:396, pi. 2, fig. 1, New York. Cestreus carolinensis Gronovius (Gray) 1854:49 (holo- type: BMNH 1853.11.12.42, a dried skin, 344 mm TL, not of Otolithus carolinensis Valenciennes). Otolithus thalassinus Holbrook 1855:132, pi. 18, fig. 2, Charleston, S.C. Otolithus obliquatus Valenciennes in Sauvage 1879:209, Martinique (lectotype: MNHN 7632, 209 mm SL, also see remarks). Macrodon Schinz Ancylodon Bosc 1816:497 (type-species: "Lonchures" Schneider = Lonchurus ancylodon Bloch and Schnei- der, by original designation; preoccupied by Ancylo- don Illiger 1811, Mammalia). "Ancylodons" Cuvier 1817:299, vernacular name. Ancylodon Oken 1817:1182 (type-species: Lonchurus an- cylodon Bloch and Schneider, by original designation; not Ancylodon Illiger). Macrodon Schinz 1822:482 (substitute name {or Ancylo- don Bosc, therefore taking the same type-species: L. ancylodon Bloch and Schneider). 36 Nomalus Gistel 1848, p. VIII (substitute name for Ancy- lodon Bosc, therefore taking the same type-species: L. ancylodon Bloch and Schneider). Sagenichthys Berg 1895:52 (substitute name for Ancylo- don Bosc, therefore taking the same type-species: L. ancylodon Bloch and Schneider). Diagnosis. — Body elongate, moderately compressed; dor- sal profile slightly arched, ventral profile evenly arched. Head pointed, compressed. Mouth large, oblique; lower jaw projecting in front of upper jaw; teeth in narrow ridge, upper jaw with a pair of very large arrow-headed canines (Fig. 37A); lower jaw with several enlarged ar- row-headed canines at the tip, exposed externally when mouth closed. Preopercular margin membranous. Gill rakers slender. Vertebrae 13+12 = 25. Swim bladder with a pair of anteriorly directed lateral horns (Cynoscion pat- tern, Fig. 12C). Sagitta oval, lateral margins concave or notched (Fig. 25B); the expanded part of the ostium reaching to anterior margin; cauda nearly straight with an expanded end (Macrodon pattern). Tropical eastern Pacific and western Atlantic, two species. One western Atlantic species: M. ancylodon. Inhabits inshore waters, over sandy to muddy bottoms. Macrodon ancylodon (Bloch and Schneider) Lonchurus ancylodon Bloch and Schneider 1801:102, pi. 25, Surinam. Ancylodon jaculidens Cuvier 1830:81, Cayenne (syn- types: MNHN 7451, 212 mm SL; MNHN 7454, 144 mm SL). Ancylodon atricauda Giinther 1880:12, Rio de La Plata (type: "5 inches," not found in BMNH). Stellifer mecatoris Delsman 1941:71, fig. 10, Mouth of Amazon River (type: not seen). Isopisthus Gill Isopisthus Gill 1863:18 (type-species: Ancylodon parvi- pinnis Cuvier, by monotypy). Paraplesichthys Bleeker 1876 (from Kaup's manuscript 1862) :335 (type-species: Isopisthus parvipinnis Gill = Ancylodon parvipinnis Cuvier, by monotypy). Diagnosis. — Body elongate, compressed, dorsa! profile nearly straight, ventral profile evenly arched. Head pointed, rather compressed. Mouth large, oblique, lowe jaw projecting; teeth in narrow band, tip of upper jaw with a pair of enlarged caninelike teeth. Preopercular margin membranous with fine cilia. Anterior (spinous) and posterior (soft) portions of dorsal fin well separated by a gap. Gill rakers moderately long. Vertebrae 11 + 14=25. Swim bladder with a pair of tubelike an- teriorly directed lateral horns (Cynoscion pattern, Fig. 12B). Sagitta nearly oval, the expanded part of the os- tium reaching to anterior margin (Fig. 25A); cauda oblique, slightly bent at the end. Tropical eastern Pacific and western Atlantic, two species. One western Atlantic species: /. parvipinnis. Inhabits inshore waters over sandy to muddy bottoms. Isopisthus parvipinnis (Cuvier) Ancylodon parvipinnis Cuvier 1830:84, pi. 105, Cayenne (syntypes: MNHN 745, three specimens, 100-140 mm SL). Isopisthus affinis Steindachner 1879:43, pi. 2, fig. 2, Port Alegre, Brazil (syntypes: NHMV 15190, two speci- mens, 168 and 172 mm SL). Isopisthus harroweri Fowler 1916:402, fig. 3, Colon, Pan- ama (holotype: ANSP 45236, 145 mm SL). Lonchurus GROUP Diagnosis. — Swim bladder carrot-shaped with a pair of anterior diverticula, each diverticulum with a short an- teriorly directed branch and a long posteriorly directed branch (Lonchurus pattern, Fig. 14); sometimes the swim bladder is reduced in size. Sagitta oval elongate to rectangular, the "tadpole-shaped" sulcus elongate; the expanded part of the ostium reaching to anterior margin and the cauda long, J-shaped (Lonchurus pattern, Fig. 24). Snout with five marginal pores and three to five up- per pores (Fig. 31C, D); lower jaw with two to many pairs of barbels (Fig. 32F, H). Two genera of western Atlantic Sciaenidae in this group: Lonchurus and Paralonchurus . Lonchurus Bloch Lonchurus Bloch 1793, pt. 7:143 (type-species: Lon- churus barbatus Bloch, by monotypy). Lonchiurus Bloch (French translation in Laveau), 1797, pt. 10:119 (invalid emendation of Lonchurus). Diagnosis. — Body long and rounded; dorsal profile slightly arched, ventral profile nearly flat. Pectoral fin very long. Head conical, low and broader than body; snout projecting. Mouth inferior; teeth conical, in bands. Gill rakers short. Vertebrae 11 + 18=29. Swim bladder much reduced in size, but retaining typical Lonchurus pattern (Fig. 14) with anterior branches of diverticula more prominent, posterior end of the main chamber tapering into a fine point (Fig. 14A). Drumming muscles thin, present only in male. Snout with five upper pores; lower jaw with four pores and two barbels (Fig. 32F), longer than eye diameter. Monotypic: L. lanceolatus. Found over sandy and muddy bottom along the coast and in estuarine waters. Lonchurus lanceolatus (Bloch) Perca lanceolata Bloch 1788:383, fig. 3, India (= West Indies). Lonchurus barbatus Bloch 1793:144, pi. 360, fig. 1, Suri- nam. Lonchurus depressus Bloch and Schneider 1801:102, Surinam. 37 Paralonchurus Bocourt Paralonchurus Bocourt 1869:21 (type-species: Paralon- churus petersi Bocourt, by monotypy). Polycirrhus Bocourt 1869:22 (type-species: Polycirrhus dumerili Bocourt by monotypy, not Polycirrhus Grube 1850. Annelida). Polyclemus Berg 1895:54 (substitute name for Polycir- rhus Bocourt, non-Grube, therefore taking the type- species: P. dumerili). Paralonchurus (Zonoscion) Jordan and Evermann 1896: 401 (type-species: Polycirrhus rathbuni Jordan and Bollman 1889, by monotypy). Paralonchurus (Zaclemus) Gilbert in Jordan and Ever- mann 1896:402 (type-species: Paralonchurus goodei Gilbert MS, by monotypy). Diagnosis. — Body elongate, slightly rounded; dorsal profile slightly elevated, ventral profile nearly flat. Pec- toral fin long or short. Head broad, nape slightly convex; snout projecting. Mouth small and inferior, teeth in con- ical bands. Gill rakers short. Total vertebrae 25 or 29 (see "Key to the species of Paralonchurus"). Swim bladder well developed (Fig. 14B, C). Drumming muscles pre- sent only in male. Sagitta elongate, the expanded part of the ostium reaching to anterior margin; cauda J-shaped (Fig. 24B, C). Snout with three upper pores (Fig. 31C); lower jaw with 12 to 16 pairs of minute barbels, anterior 3 pairs form a tuft at the symphysis of lower jaw, other bar- bels along the rami of lower jaw (Fig. 32H). Tropical eastern Pacific and western Atlantic, about six species. Two species from the western Atlantic in this genus: P. brasiliensis and P. elegans. Inhabits estuarine and in- shore waters with sandy to muddy bottoms. Paralonchurus brasiliensis (Steindachner) Genyanemus brasiliensis Steindachner 1875:476, Para, Santos (type: presumably in NHMV, but not found). Micropogon ornatus Giinther 1880:13, pi. 7, fig. A, Rio de La Plata (holotype: BMNH 1879.5.14.289, 175 mm SL). Paralonchurus rathbuni Puyo 1949:215, fig. 114 (not Polycirrhus rathbuni Jordan and Bollman 1889:162). Paralonchurus elegans Boeseman Paralonchurus elegans Boeseman 1948:3, Surinam (holo- type: RMNH 390, 200 mm SL). Remarks. — Paralonchurus elegans and Lonchurus lanceolatus have very similar sagittae (Fig. 24) and both have a long black pectoral fin. Paralonchurus brasiliensis has a total vertebrae count of 29 as in L. lanceolatus. Paralonchurus brasiliensis and P. elegans both have well- developed swim bladders and many minute barbels on the chin. The interrelationship of these three species cannot be determined without comparative material of the eastern Pacific species of this group. The reduced size of the swim bladder in L. lanceolatus (Fig. 14A) has been viewed as an important diagnostic character (Giinther 1860). Paralonchurus brasiliensis has been considered to belong in a separate genus, Polyclemus as by Berg (1895). The type-species of Paralonchurus (P. petersi) and Polyclemus (P. dumerili) are eastern Pacific species. Study of these species will be necessary to clarify the generic status of these fishes. Stellifer GROUP Diagnosis. — Swim bladder separated into two chambers by a constriction; the anterior chamber yoke-shaped located in front of the septum transversum and the pos- terior one simple carrot-shaped (Stellifer pattern, Fig. 15). A pair of diverticula sometimes present on the an- terior chamber posterolateral^. Otoliths with both sagit- ta and lapillus enlarged (Stellifer pattern, Fig. 27). Snout with five marginal pores and three to five upper pores (Fig. 31C, D); lower jaw with four to six pores and without barbels (Fig. 32B-D). Four genera of western At- lantic sciaenids in this group: Bairdiella, Odontoscion, Ophioscion, and Stellifer. At present, the generic bound- aries of Bairdiella and Odontoscion are not clear, espe- cially when the eastern Pacific species and genera of this group are considered (Wintersteen and Chao, see foot- note 3). Bairdiella Gill Bairdiella Gill 1862a:33 (nomen nudum). Bairdiella Gill 1862b:83 (type-species: Bodianus argyro- leucus Mitchill 1815 = Dipterodon chrysourus Lace- pede 1803, by original designation). Corvula Jordan and Eigenmann 1889:377 (type-species: Johnius batabanus Poey, by original designation, see remarks). Vacuoqua Jordan and Evermann 1927:506 (type-species: Corvula macrops Steindachner, by original designa- tion, see remarks). Diagnosis. — Body oblong, compressed; dorsal profile slightly elevated, ventral profile nearly straight. Head with or without strong spines at angle of preopercular margin. Mouth terminal, slightly oblique; upper jaw teeth conical in a narrow band, outer row moderately en- larged; lower jaw teeth on a narrow ridge, median row slightly enlarged. Gill rakers moderately long and stiff. Vertebrae 12+13 or 11 + 14=25. Swim bladder in two chambers, without diverticula on anterior chamber (Fig. 15A). Sagitta triangular; ostium incomplete; lapillus more than half the size of sagitta (Fig. 27C). Snout with five marginal and three upper pores; lower jaw with four to six pores, the median pair of pores in six-pored form very closely set (Fig. 32B-D). Tropical eastern Pacific and western Atlantic, about eight species. Four species of western Atlantic Sciaenidae included here: B. bata- bana, B. chrysoura, B. ronchus, and B. sanctaeluciae (also see remarks). Inhabit inshore waters, estuaries, and coral reefs over sandy to muddy bottom. 38 Remarks. — The genera Corvula Jordan and Eigenman 1889, and Vacuoqua Jordan and Evermann 1927 are listed under Bairdiella for the convenience of the present study. Bairdiella batabana and B. sanctaeluciae differ from B. chrysoura and B. ronchus by lack of strong spines (or serrations) at the preopercular margin and the vertebral arrangement is 12+13 instead of 11 + 14. These characters are diagnostic within the suprageneric group Stellifer. Therefore, B. batabana and B. sanctaeluciae may be recognized as a separate genus (or subgenus) from other Bairdiella species. Jordan and Evermann (1927) distinguished the eastern Pacific sciaenid fish Corvina macrops Steindachner, from Johnius bata- banus Poey by its deeper, symmetrical body and the silvery coloration of Bairdiella and related species. These characters are not diagnostic within the genus Bair- diella. Study of East Pacific Bairdiella and related species will be necessary to clarify the generic status of Bairdiella (see footnotes 3 and 4). Bairdiella batabana (Poey) Johnius batabanus Poey 1860, 2:184, Batabano, Cuba (holotype: MCZ 21957, 189 mm SL). Corvula sialis Jordan and Eigenmann 1889:379, Key West, Fla. (holotype: USNM 26575, 132 mm SL). Bairdiella chrysoura (Lacepede) Perca puncatata Linnaeus 1766:482, South Carolina (pre- occupied by Perca punctatus Linnaeus 1758:291, = Bodianus fuluus punctatus; syntypes?: Linnean Society of London, no. 108, a left side dried skin, 144 mm TL, and no. 109, a left side dried skin, 153 mm TL). Dipterodon chrysourus Lacepede 1803, 4:166, South Carolina (type: presumably in MNHN, but not found). Bodianus argyroleucus Mitchill 1815:417, pi. 6, fig. 9, New York. Bodianus exiguus Mitchill 1815:419, New York. Bodianus pallidus Mitchill 1815:420, New York. Homoprion xanthurus Holbrook 1855:170, pi. 24, fig. 2, South Carolina (not Leiostomus xanthurus Lacepede). Bairdiella ronchus (Cuvier) Corvina ronchus Cuvier 1830:107 (syntypes: MNHN 95, two specimens, 123 and 149 mm SL, St. Dominique; MNHN 5345, two specimens, 144 and 156 mm SL, Surinam; MNHN 5345, a dried stuffed specimen, 247 mm SL, Maracaibo?). Corvina subaequalis Poey 1875:58, Cuba (holotype: 245 mm TL, not located, also see Robins 1964). Sciaena bedoti Regan 1905:391, pi. 6, fig. 1, Cuba (syn- types: BMNH 1905.3.18.2, 155 mm SL; Museum d'Histoire Naturelle, Geneve 678.1, 222 mm TL, also see Robins 1964). Bairdiella verae-crucis Jordan and Dickerson 1908:17, fig. 1, Vera Cruz (holotype: USNM 61676, 194 mm SL). Corvina fulgens Vaillant and Bocourt 1915:164, pi. 6, fig. 2, U.S.A. (syntypes: MNHN A. 975, two specimens, 149 and 150 mm SL). Bairdiella sanctaeluciae (Jordan) Corvula sanctaeluciae Jordan 1889:649, Port Castries, St. Lucia (holotype: USNM 4173, 105 mm SL). Odontoscion Gill Odontoscion Gill 1863:18 (type-species: Corvina dentex Cuvier, by original designation). Diagnosis. — Body oblong, compressed; profile evenly arched. Head conical; snout short, blunt; eye large. Pre- opercular margin smooth without strong spines. Mouth large, terminal, slightly oblique; enlarged conical teeth in a widely spaced row on both jaws, lower jaw with some tiny teeth outside the large ones, and two larger canine- like teeth at the tip (Fig. 37C). Gill rakers relatively long. Vertebrae 12+12=25. Swim bladder with two chambers, anterior one yoke-shaped in front of septum transversum and posterior one carrot-shaped, without diverticula (Fig. 15A). Sagitta and lapillus both enlarged, only a small oval-shaped ostium left on the sulcus (Fig. 27D). Snout with five marginal pores and three upper pores (Fig. 31C); lower jaw with four pores and no barbel (Fig. 34C). Tropical eastern Pacific and western Atlantic, three species. One species in the western Atlantic: O. dentex. Inhabits coral reefs and offshore waters over sandy bottoms. Odontoscion dentex (Cuvier) Corvina dentex Cuvier 1830:139, pi. 109, Saint Domi- nique (holotype: MNHN 144, 114 mm SL). Ophio8cion Gill Ophioscion Gill 1864b:164 (type-species: Ophioscion typicus Gill, by monotypy). Ophioscion (Sigmurus) Gilbert in Jordan and Evermann 1898:1452 (type-species: Corvina vermicularis Gun- ther, by monotypy). Diagnosis. — Body robust, dorsal profile elevated, ven- tral profile slightly convex. Head broad, conical top with prominent cavernous canals; bony interorbital width 3.5 or more in head. Mouth small, inferior, teeth conical in bands. Gill rakers short. Vertebrae 10+15=25. Swim bladder with two chambers, anterior one yoke-shaped and posterior one carrot-shaped, without diverticula (Fig. 15A). Both sagitta and lapillus enlarged; ostium incom- plete (Fig. 27A). Snout with three large and two to four minute upper pores, and five marginal pores, rostral fold deeply indented (Fig. 33B). Tropical eastern Pacific and western Atlantic, about 11 species. Three nominal species of this genus from western Atlantic are listed 39 here: 0. adustus, 0. panamensis, and 0. punctatissimus. Inhabits inshore waters over sandy to muddy bottoms. Ophioscion adustus (Agassiz) Corvina (Sciaena) adusta Agassiz 1831:126, pi. 70, Mon- tevideo (type: see remarks). Corvina gilli (?) Steindachner 1867:335, La Plata (type: not located, see remarks). Ophioscion panamensis Schultz Ophioscion panamensis Schultz 1945:134, fig. 8, Fox Bay, Colon, Panama (holotype: USNM 122612, 52 mm SL). Ophioscion punctatissimus Meek and Hildebrand Ophioscion punctatissimus Meek and Hildebrand 1925: 644, pi. 68, Panama (holotype: USNM 81766, 131 mm SL, Canal Zone). Remarks. — The soft dorsal and anal fin ray counts of Corvina adusta Agassiz (1831) on page 127 (D. X+I, 28; A. IT, 9) are different from the Spix figure on plate 70 (D. X + I, 19 or 20; A. II, 7). The descriptions of Ophioscion adusta (Agassiz) in Jenyns (1842:42), Gunther (1860:289), Berg (1895:52), and Schultz (1945:128) resemble the text description of Agassiz. Jordan and Eigenmann (1889:403) interpreted the specimen (MCZ 22417, 180 mm TL from Fonteboa, Brazil = 0. punc- tatissimus) as O. adusta of Spix plate 70 and suggested that a new name should be proposed. Jordan and Ever- mann (1898:1447) added Stellifer microps (MCZ 1031, Fonteboa, Brazil) and 0. punctatissimus (MCZ 21728, Jeremie, Haiti) in their description of 0. adusta. The de- scriptions of 0. adustus (Meek and Hildebrand 1925:639) and 0. panamensis (Schultz 1945:134) resemble plate 70 of Spix. Furthermore Schultz (1954) interpreted 0. adusta by a specimen from Uruguay (USNM 86710, 120 mm SL) and suggested that Ophioscion woodwardi Fowler (= Micropogonias furnieri) is probably a junior synonym of 0. adusta (the text description of Agassiz 1831). The descriptions of 0. adusta in A. Miranda Ribiero (1915, Sciaendae, p. 23), Devincenzi (1924:239), and Dahl (1971:260) are probably 0. punctatissimus or Stellifer species. Jordan and Eigenmann (1889) reported that the nominal species Corvina gilli (Steindachner 1867:335) is similar to 0. adusta of Spix's plate 70 (Agassiz 1831). The description of C. gilli Steindachner from La Plata River seems to be of a sciaenid fish except for the low soft dorsal fin ray count (16). Its identity is unknown. The type-specimen of Corvina adusta was "in Museo Monacensi, 10" longum" (Agassiz 1831). It was probably destroyed in Munich during a bombing attack in April 1944 (Trewavas 1950). The length of the fish (10 in) is also too big for Ophioscion or Stellifer species. In my opin- ion, therefore, the text description of C. adusta (Agassiz 1831) is most probably a species of Micropogonias, al- though the minute mandibular barbels were not men- tioned by these authors (also see section on Micro- pogonias). in addition, four pairs of minute barbels are present on the lower jaw of 0. woodwardi Fowler (holo- type: ANSP 68257). The plate of C. adusta (Agassiz 1831) probably fits best the description of O. panamen- sis (Schultz 1945). But the type-specimens of 0. pan- amensis (holotype and 13 paratypes in USNM) are all juveniles (23.3 to 52 mm SL). Their identities are doubt- ful and no specimens have been reported except the types. For the present, 0. panamensis and 0. punctatis- simus are recognized here and included in the "Key to the genus Ophioscion.'" Stellifer Oken "Stelliferes" Cuvier 1817:283, vernacular name. Stellifer Oken 1817:1182 (type-species: Bodianus stelli- fer Bloch, by original designation). Stelliferus Stark 1828:459 (type-species: Stellifer capen- sis Stark = Bodianus stellifer Bloch, by monotypy). Homoprion Holbrook 1855:168 (type-species: Homoprion lanceolatus Holbrook, by subsequent designation of Gill 1862b:83). Bairdiella (Nector) Jordan and Evermann 1898:1432 (type-species: Bairdiella chrysoleuca Gunther 1867, by original designation). Stellifer (Zestis) Gilbert in Jordan and Evermann 1898: 1439 (type-species: Stellifer oscitans Jordan and Gil- bert, by original designation). Stellifer (Zestidium) Gilbert in Jordan and Evermann 1898:1439 (type-species: Stellifer illecebrosus Gilbert, by original designation). Stellifer (Stellicarens) Gilbert in Jordan and Evermann 1898:1439 (type-species: Stellifer zestocarus Gilbert, by original designation). Diagnosis. — Body robust, back elevated. Head broad, rather flat on top with apparent cavernous canals; in- terorbital wide, bony interorbital width 3.5 or less in head. Mouth moderate to large, inferior to oblique (Fig. 38); teeth conical or villiform in bands. Gill rakers usual- ly long and slender, some moderate to short. Vertebrae 10+15 = 25. Swim bladder in two chambers; the anterior chamber yoke-shaped in the head with a pair of divertic- ula, tubelike or bulblike (Fig. 16), and the posterior chamber simple carrot-shaped. Both sagitta and lapillus enlarged, the ostium on sagitta incomplete (Fig. 28). Snout with five marginal pores, the median one rounded, separate from the rostral fold and arranged rhom- boidally with three upper pores at the tip of the snout (Fig. IB); lower jaw with four to six pores, no barbel (Fig. 35). Tropical eastern Pacific and western Atlantic, about 24 species. Eleven species of western Atlantic sciaenids belong to this genus: S. brasiliensis, S. colonensis, S. griseus, S. lanceolatus, S. microps, S. naso, S. rastrifer. S. sp. A, S. sp. B, S. stellifer, and S. venezuelae. In- habits inshore waters of sandy bottoms, also in estuaries and around coral reefs. 40 Stellifer brasiliensis (Schultz) Stellifer venezuelae (Schultz) Ophioscion brasiliensis Schultz 1945:128, fig. 6, Santos, Brazil (holotype: USNM 87742, 77 mm SL). Stellifer colonensis Meek and Hildebrand Stellifer colonensis Meek and Hildebrand 1925:623, pi. 46, fig. 1, Mindi, Panama (holotype: USNM 81729, 99 mm SL). Ophioscion venezuelae Schultz 1945:131, fig. 7, Vene- zuela (holotype: USNM 121749, 140 mm SL). Remarks. — Two additional new species of Stellifer (see footnote 3) are included in the section on "Key to the species of Stellifer" as Stellifer sp. A and Stellifer sp. B. Freshwater Sciaenidae of America Stellifer griseus Cervigon Stellifer sp. Cervigon 1966a:509, fig. 209, Morro de Puer to Santos, Venezuela. Stellifer griseus Cervigon 1966b:l, fig. 1, North Penin- sula de Araya (holotype in Museo de Historia Natural La Salle, Venezuela, MHNLS 1.875, 120 mm SL, not examined; paratypes: USNM 200782, two specimens, 108 and 123 mm SL). Stellifer lanceolatus (Holbrook) Homoprion lanceolatus Holbrook 1855:168, pi. 23, Beau- fort S.C. (type: presumably lost). Stellifer microps (Steindachner) Coruina microps Steindachner 1864:205, pi. 2, fig. 2, Guiana (type: presumably in NHMV, but not found). Ophioscion costaricensis Caldwell 1958:117, fig. 1, Tortu- guero, Costa Rica (holotype: UF 5831, 117 mm SL). Stellifer naso (Jordan) Stelliferus naso Jordan in Jordan and Eigenmann 1889: 395, Cachoeira, Brazil (syntypes: USNM 130630, 5 specimens, 68-75 mm SL). Stellifer rastrifer (Jordan) Stelliferus rastrifer Jordan in Jordan and Eigenmann 1889:393, Santos, Brazil (holotype: MCZ 10815A, 128 mm SL). Stellifer stellifer (Bloch) Bodianus stellifer Bloch 1790, pt. 4:55, pi. 231, fig. 1, "Cape of Good Hope" (type-locality is probably in error; since Bloch's specimen was obtained from a Dutch auction, the specimen very likely came from Surinam rather than Africa; also see Jordan and Eigenmann 1889:394). Corvina trispinosa Cuvier 1830:109, Cayenne and Brazil. Stellifer mindii Meek and Hildebrand 1925:626, pi. 66, fig. 2, Mindi Reef, Panama (holotype: USNM 81730, 93.4 mm SL). Four genera, Aplodinotus, Pachypops, Pachyurus, and Plagioscion, and 22 nominal species of sciaenids are listed here from Atlantic river systems of America. Of these, Aplodinotus is the only North American fresh- water inhabitant, and the other three genera are en- demic to continental South America. Because of lack of sufficient material for the South American genera, the diagnoses of these genera are based mainly on the literature (mainly on Gunther 1860; Liitken 1875; Jor- dan and Evermann 1896; Jordan and Eigenmann 1889; Eigenmann 1912; Starks 1913; Campos 1942; Lowe 1966; Ringuelet et al. 1967) and type-specimens. The exact limits of these genera are not clear at present. Aplodinotus Rafinesque Aplodinotus Rafinesque 1819:418 (type-species: Aplo- dinotus grunniens Rafinesque, by monotypy). Haploidonotus Gill 1862d:102 (invalid emendation of Aplodinotus, Rafinesque 1819, therefore taking the same type-species: Aplodinotus grunniens Rafinesque). Eutychelithus Jordan 1876:242 (type-species: Corvina richardsoni Cuvier, by monotypy = Aplodinotus grun- niens Rafinesque). Diagnosis. — Body oblong, snout blunt, back elevated and compressed. Mouth small, horizontal to inferior; lower pharyngeals very large, completely united (Fig. 40B). Swim bladder simple carrot-shaped (Fig. 5D). Sa- gitta more or less semicircular, the expanded part of the ostium not reaching anterior margin (Fig. 19B). Snout without upper pores (Fig. 31B); lower jaw without mental barbels. Vertebrae 10+14=24. Freshwater North and Central America. Monotypic: A. grunniens. Remarks. — The original description of Amblodon Rafinesque (1819:421) included two species of buffalo fish (Catostomidae), A. bubalus and A. niger. The unique lower pharyngeal teeth of Aplodinotus grunniens were wrongly attributed to these fishes. Rafinesque's (1820:24) subsequent replacement of Aplodinotus with Amblodon does not make Amblodon an available name in the Sciaenidae. Aplodinotus grunniens Rafinesque Aplodinotus grunniens Rafinesque 1819:419, Ohio River (no type mentioned in the description). 41 Sciaena oscula LeSueur 1822:252, pi. 13, Lake Erie (holo- type: MNHN A. 5696, 308 mm SL). Sciaena grisea LeSueur 1822:254, Ohio River (type: "18 to 24 inches in total length," presumably in MNHN, but not found). Corvina richardsoni Cuvier 1830:100, Lake Huron (type: presumably in MNHN, but not found). Amblodon concinnus Agassiz 1854:307, Tennessee River (lectotype herein designated: MCZ 21970, 251 mm SL). Amblodon lineatus Agassiz 1854:307, Osage River (syn- types: MCZ 569, two specimens, 198 and 160 mm SL). Amblodon neglectus Girard 1859:167, Rio Grande del Norte, Rio Bravo (holotype: USNM 639, 84.3 mm SL). Remarks. — Two catalogue numbers, MNHN A. 5696 and MNHN 7536, are indicated as "syntypes of Corvina os- cula LeSueur": MNHN A. 5696, a dried stuffed specimen, 308 mm SL, collected by LeSueur from Lake Erie was examined; MNHN 7536 from New Orleans was not found. In Agissiz's (1854) original description no types were mentioned. The MCZ specimens of A. con- cinnus and A. lineatus catalogued as types are con- sidered the types. Pachypops Gill Pachypops Gill 1862b:87 (type-species: Micropogon trifilis Miiller and Troschel 1848, by original designa- tion). Diagnosis. — Body moderately elongate, dorsal profile elevated and descending nearly straight under soft por- tion of dorsal fin, ventral profile nearly straight. Head oblong, snout convex and projecting; suborbital region much swollen and translucent. Mouth small, inferior, teeth villiform in narrow bands; maxillae almost en- tirely concealed under the suborbitals. Preopercular margin slightly serrate. Snout with five upper pores and five marginal pores; lower jaw with five to six pores and three to many barbels. From Cuvier (1830:pl. 138) and Trewavas' (1964) description, the swim bladders of Pachypops fourcroi and P. trifilis have a pair of anterior appendages each dividing into a longer posterior branch and a shorter anterior one; the anterior branch extend- ing in front of the septum transversum (Lonchurus pat- tern, Fig. 14). The sagitta of a Pachypops fourcroi (153 mm SL) is similar to the sagitta of Pachyurus schom- burgkii (Fig. 18K). Four nominal species are listed here: P. adspersus, P. camposi, P. fourcroi, and P. trifilis. Pachypops adspersus (Steindachner) Pachyurus (Lepipterus) adspersus Steindachner 1880: 123, Rio San Antonio, Brazil (type: presumably in XHMV, but not found). Pachypops camposi Fowler Pachypops camposi Fowler 1954:252, Rupununi River, British Guiana (holotype: ANSP 39773, 117 mm SL, labeled as P. steindachneri). Pachypops fourcroi (Lacepede) Perca fourcroi Lacepede 1803, 4:398, no locality (holo- type: MNHN 7539, 135 mm SL). Pachypops trifilis (Miiller and Troschel) Micropogon trifilis Miiller and Troschel 1848:622, British Guiana (type: not located). Pachyurus Agassiz Pachyurus Agassiz 1829:125, 127 (type-species: Pachy- urus squamipennis Agassiz, by monotypy). Lepipterus Cuvier 1830:152 (type-species: Lepipterus francisci Cuvier, by monotypy). Diagnosis. — This genus may be divided into at least two groups (or genera) based on the swim bladder. One with a simple carrot-shaped swim bladder (P. schomburgkii, Fig. 6B, the other with a pair of tubelike diverticula ex- tending from the anterior corners of swim bladder, taper- ing back to the end of the main chamber (P. bonariensis, Fig. 6A). Because of lack of study material, only com- mon characters are presented. Body moderately elongate, dorsal profile slightly elevated, ventral profile nearly straight or slightly arched. Head conical; snout blunt, swollen, and translucent. Eye moderate to large. Mouth horizontal, inferior or terminal; teeth villiform in bands. Snout with five marginal pores and usually without upper pores, some with three to five upper pores; lower jaw with five pores and no barbel. Sagitta moderate to large. Second spine of anal fin moderate to strong. Eight nominal species are included in this group: P. biloba, P. bonariensis, P. francisci, P. grunniens, P. lundii, P. paranensis, P. schomburgkii, and P. squamipennis. Pachyurus biloba Cuvier Corvina biloba Cuvier 1830:112, no locality (holotype: MNHN 7683, 75.6 mm SL). Pachyurus bonariensis Steindachner Pachyurus (Lepipterus) bonariensis Steindachner 1870: 126, La Plata, Argentine (NHMV 15181 may be one of the syntypes, 188 mm SL, La Plata, Argentine; CAS specimen, Indiana University 11353, 152 mm SL, Buenos Aires, collected by W. B. Scott and labeled cotype is apparently a mistake). Pachyurus francisci (Cuvier) Lepipterus francisci Cuvier 1830:152, pi. 113, "Riviere de Saint-Francois," Brazil (type: not located). 42 Lepipterus corvina Reinhardt 1851:30, Rio das Velhas, Brazil (syntypes: ZMK 1 and 2; 336 and 274 mm SL). mm SL). Pachyurus grunniens (Schomburgk) Corvina grunniens Schomburgk 1860:136, pi. 2, Esse- quibo River, Comacca Island, British Guiana (not of Aplodinotus grunniens Rafinesque). Pachyurus lundii Reinhardt Pachyurus lundii Reinhardt 1855:108, Rio das Velhas, Brazil (syntypes: ZMK 1 and 2; 336 and 274 mm SL). Pachyurus paranensis Daneri Pachyurus paranensis Daneri 1956:6, fig. 1, Santa Fe, Argentina (type: not located). Pachyurus schomburgkii Gunther of Pachyurus schomburgkii Gunther 1860:282, rivers Brazil (holotype: BMNH 49.11.8.22, 197 mm SL). Pachyurus natteri Steindachner 1863:10, pi. 3, Rio Branco, Rio Negro, Brazil (type: presumably in NHMV, but not found). Pachyurus nattereri Campos 1942:17 (valid emendation of Steindachner's spelling). Pachyurus squamipennis Agassiz Pachyurus squamipennis Agassiz in Spix and Agassiz 1831:128, pi. 71, Brazil. Remarks. — Agassiz's spelling of the species name is squamipennis and should be used although it has often been spelled as squamipinnis. The type-specimen of this species probably was destroyed in Munich during a bombing attack in April 1944 (Trewavas 1950). Plagio8cion Gill Plagioscion Gill 1862b:82 (generic description, no spe- cies or type indicated, compared with Corvina of Cu- vier; type-species: Sciaena squamosissima Heckel, by subsequent designation of Jordan and Eigenmann 1889:380). Diplolepis Steindachner 1863:2 (type-species: Diplolepis squamosissimus Steindachner = Sciaena squamosis- sima Heckel, by monotypy, preoccupied by Diplolepis Geoffroy 1762, cynipid insect and Diplolepis Fabricius 1805, calcid insect, quoted from Neave 1939-40; spelled Diplolepsis on p. 3). Diagnosis. — A freshwater genus of Sciaenidae from South American rivers, some species occasionally in es- tuaries, rarely marine. Body elongate, dorsal profile evenly arched, ventral profile nearly straight. Head slightly conical, snout protuberant. Mouth moderate, terminal, slightly oblique, no enlarged caninelike teeth at tip of upper jaw. Scales along lateral line concealed by many smaller scales in some species, appearing as one enlarged scale (Fig. 41). Gill rakers rather long and slender. Swim bladder with a pair of hornlike divertic- ula, originating from anterior quarter of main chamber and hooked at the distal ends (Cynoscion pattern, Fig. 12D). Sagit*a suboval, expanded part of the ostium reaching to anterior margin and the cauda deeply curved in a J-shape (Fig. 18L). Nine nominal species are in- cluded in this genus: P. auratus, P. heterolepis, P. mac- donaghi, P. microps, P. monacantha, P. pauciradiatus, P. squamosissimus, P. surinamensis, and P. ternetzi. Figure 41.— Portions of lateral line scales of Plagioscion surinamen- sis. The large-scale view shows numerous small scales covering a segment of lateral line; in the small-scale view, the lateral line pores are seen through the small scales. Plagioscion auratus (Castelnau) Johnius auratus Castelnau 1855:12, pi. 4, fig. 2; Ucayala, Brazil (holotype: MNHN 7622, 203 mm SL). Plagioscion heterolepis (Bleeker) Johnius heterolepis Bleeker 1873:456, Surinam (syn- types: RMNH 6042, two specimens, 123 and 129 mm SL, these type-specimens are similar to the Stellifer group in external appearance. Its true identity is un- certain at present). 43 Plagioscion macdonaghi Daneri Plagioscion macdonaghi Daneri 1954:179, fig. 1, Rio de La Plata (holotype in Coleccion Nacional del Institute Nacional de Investigacion de las Ciencias Naturales, Brazil, no. 4197, 147 mm TL, not examined). Plagioscion microps Steindachner Plagioscion microps Steindachner 1917:657, pi. 1, fig. 1, Amazon (presumed syntype: NHMV 15180, 206 mm TL, collected by Steindachner in Surinam, may be one of the originally described specimens, 208 and 214 mm TL). Plagioscion monacantha (Cope) Coruina monacantha Cope 1867:402, near Paramaribo, Surinam (holotype: ANSP 11519, 167 mm SL). Sciaena magdalenae Steindachner 1878:22, pi. 1, fig. 1, Magdalena River (types: 270-540 mm TL, presumably in NHMV, but not found). Plagioscion pauciradiatus Steindachner Plagioscion pauciradiatus Steindachner 1917:660, figure on p. 661, Paramaribo (type: presumably in NHMV, but not found). Plagioscion squamosissimus (Heckel) Sciaena squamosissima Heckel 1840:438, Amazon. Sciaena rubella Schomburgk 1860:133, British Guiana. Johnius courvina Castelnau 1855:11, pi. 5, fig. 1, Rio Crixas, Rio Arognay, Brazil (holotype: MNHN 7503, 366 mm TL). ■Johnius amazonicus Castelnau 1855:12, pi. 4, fig. 1, Amazon (syntypes: MNHN 7504, two specimens, 96.3 and 145 mm SL). Plagioscion surinamensis (Bleeker) Pseudosciaena surinamensis Bleeker 1873:458, Surinam (holotype: RMNH 5995, 88.1 mm SL). Plagioscion ternetzi Boulenger Plagioscion ternetzi Boulenger 1895:523, Paraguay (syn- types: BMNH 1895.5.17.1-2, two specimens, 88.4 and 263 mm SL). Phylogenetic Relationships of the Genera and Suprageneric Groups The morphological patterns of swim bladders, otoliths, and external features of western Atlantic sciaenid fishes (Table 1) are considered here to assess phylogenetic rela- tionships. In the following discussion, relationships will be discussed among genera within the groups, then among suprageneric groups. The adaptive characters are indicated and their phyletic significance are also dis- cussed. The dendrogram (Fig. 4) indicates similarities of these sciaenid genera but may not express the actual phylogeny. Study of sciaenids in other areas, par- ticularly the eastern Pacific (see footnote 4), is needed to test the relationships hypothesized here. Relationships of the Genera Within the Suprageneric Groups Sciaena GROUP Members of this group all have a simple swim bladder (Sciaena pattern, Fig. 5) and a thick sagitta (Sciaena pattern, Fig. 18). The shape and position of the sulcus varies among the genera and species. Equetus and Paraques are very similar in having an elevated dorsal profile and dark stripes on the side. They are mainly cor- al reef and coastal dwellers. Sciaena usually lives in deeper offshore water (Chao and Miller 1975) than other sciaenids. Members of the Sciaena group have enclosed inferior mouths except for Sciaena with a subterminal and slightly inferior mouth. Habitat diversification of this group is greatest among all western Atlantic supra- generic groups of sciaenids. Umbrina GROUP This group is most similar to the Sciaena group, in having a simple swim bladder (Sciaena pattern, Fig. 5) and a thick sagitta (Sciaena pattern, Fig. 18). These fishes also have a barbel at the tip of lower jaw (Fig. 36A, C). The mental barbel of Ctenosciaena is flexible and tapered at its tip without an apical pore, whereas that of Umbrina is rigid and short with an apical pore. Also, the mouth of Ctenosciaena gracilicirrhus is subterminal, and that of Umbrina is inferior. These morphological dif- ferences may be attributed to an adaptation of C. gracilicirrhus to deeper waters habitats than Umbrina. The sagittae of C. gracilicirrhus, S. trewavasae, and U. milliae are most similar among fishes of Sciaena pattern sagittae (Fig. 18A, G, J) suggesting a relationship between the Sciaena and Umbrina groups (Fig. 4). Leiostomus is similar to members of the Sciaena group, but it has a much thinner sagitta (Fig. 18D). In addition, the ostium of its sulcus is inclined toward the dorsal margin (to the right of Fig. 18) . The kinship of this genus is tentatively put with the Sciaena and Umbrina groups. Pogonias GROUP The North American freshwater Aplodinotus grun- niens and western Atlantic Pogonias cromis have a similar thin and semicircular sagitta (Pogonias pattern, 44 Fig. 19). Both of their lower pharyngeal bones are fully united into a triangular plate (Fig. 40). The uniting suture of the lower pharyngeals is apparently weaker in P. cromis. Although the swim bladder of A. grunniens is a simple carrot-shape (Sciaena pattern, Fig. 5D) and that of P. cromis has very complicated diverticula (Pogonias pattern, Fig. 10) in adults, they may still be most closely related to each other (Fig. 4). Cynoscion GROUP Genera of this group have a similar swim bladder (Cynoscion pattern, Fig. 12). Macrodon and Cynoscion are most similar to each other in external morphology, except that the large canines in Macrodon have arrow- headed tips and several large canines are present at the tip of lower jaw (Fig. 37A). Isopisthus differs from all other western Atlantic sciaenids in having a space between the anterior (spinous) and posterior (soft) por- tions of dorsal fin, but its general body shape resembles Cynoscion. The South American freshwater genus Plagioscion has a terminal horizontal mouth, and lacks enlarged canines. Its body shape is deeper and is not as fusiform as other genera of this group. This may be an adaptation to the freshwater habitat, whereas, the fast swimming and fusiform Cynoscion, Isopisthus, and Mac- rodon may be adapted to open water habitats. The sagit- tae of the genera in the Cynoscion group differ from each other (Figs. 23, 25). Macrodon ancylodon has a thin elongate sagitta (Fig. 25B) similar to Cynoscion (Fig. 23), but the general outline and the sulcus differ. In Plagios- cion surinamensis (Fig. 18L) and Isopisthus parvipinnis (Fig. 25A), the sagittae have a similar thickened pos- terior half. Sciaenidae usually have a thick sagitta; therefore, this similarity is not indicative of generic rela- tionships. Plagioscion may be more closely related to the Cynoscion group than to other sciaenid fishes (Fig. 4). Lonchurus GROUP Paralonchurus and Lonchurus show a sequence of reduction in swim bladder size (Fig. 14). Lonchurus lanceolatus has the most reduced swim bladder (Fig. 14A) and weak drumming muscles only in male. It is a coastal and estuarine dweller. Its habitat is similar to that of Menticirrhus which has only a vestige of the swim bladder remaining in adults. The reduced swim bladder of L. lanceolatus also shows a basic two-horned struc- ture, which resembles that of the Cynoscion group (Fig. 12). Paralonchurus usually has a well-developed swim bladder, but the relative size in P. elegans is slightly reduced (Fig. 14B), especially in female specimens. They inhabit deeper inshore waters than does Lonchurus. In comparing the sagitta (Fig. 24) and external morphology of Lonchurus and Paralonchurus, the similarities among L. lanceolatus, P. brasiliensis, and P. elegans indicate that they are most closely related to each other. Other eastern Pacific Paralonchurus species are most important to define the boundaries and relationships of this supra- generic group (or tribe). In addition, the South American freshwater scianid genus Pachypops may also belong to this group, based on the swim bladder of P. fourcroi (Fig. 14D). The few species examined from this genus all have inferior mouths and three to many pairs of barbels. Stellifer GROUP Genera of this group all have a two-chambered swim bladder (Fig. 15) and an enlarged lapillus (Fig. 27). The genus Stellifer has a pair of diverticula posterolateral^ on the anterior chamber of the swim bladder (Fig. 16). Morphological variation of the swim bladder diverticula is most important in assessing species relationships within Stellifer. Ophioscion has a sagitta and lapillus shaped similarly to Stellifer (Fig. 27A, B), and the body shape, cavernous head, and swim bladder position are similar. Ophioscion lacks diverticula on the anterior chamber of the swim bladder, but is otherwise similar to Stellifer (Fig. 4). Bairdiella and Odontoscion (Fig. 37C) have a terminal mouth and lack apparent cavernous canals on the head. Odontoscion inhabits clear-water reefs and has large eyes and large caninelike teeth (Fig. 37C0. Genera of this group are mainly estuarine. Osteological studies of Topp and Cole (1968) on Sciaenops and Mohsin (1973) on four Cynoscion species from the Gulf of Mexico are also useful in assessing rela- tionships within the genera. Other presently monotypic suprageneric groups of western Atlantic sciaenids may contain more genera from other geographic regions, es- pecially the eastern Pacific. Relationships of the Suprageneric Groups The Sciaena and Umbrina groups are similar in swim bladder (Sciaena pattern, Fig. 5) and otolish (Sciaena pattern, Fig. 18) morphology. Their external mor- phology is similar except that the Umbrina group has a mental barbel (Fig. 36A, C). Members of both groups are mainly bottom feeders with inferior mouths. The genus Sciaena of the Sciaena group and the genus Ctenosciaena of the Umbrina group have entered deeper waters and both have a relatively larger subterminal mouth to feed in midwater. The cluster of Sciaena and Umbrina groups are in turn most closely related to the Larimus group (Fig. 4). The species of Larimus have a basic simple swim bladder (Fig. 5E, F). A pair of anterior projections found on the swim bladder of L. breviceps appear to be a modification of the Sciaena swim bladder pattern. The sagitta of Larimus species (Fig. 20A, B) has a unique out- line, and the ostium is much larger and does not reach to the anterior margin of the sagitta. The lack of upper pores on the snout (Fig. 31B) and four minute mental pores (Fig. 34B) in Larimus are adaptive characters cor- related with its large oblique mouth and upper water column feeding habits (Fig. 30; Table 1). Among the suprageneric groups, members of both Pogonias and Sciaenops have a complicated swim blad- 45 der as adults (Figs. 10, 11), but the structural patterns are different and their sagittae also differ (Figs. 19, 20C, D). Although Pogonias and Sciaenops have a similar inferior mouth and inhabit inshore coastal and estuarine waters, the body shape of Sciaenops is more elongate and less compressed than Pogonias. This may be an adaptation of Sciaenops to the shallow-water surf zone habitat. They are probably most closely related to the Larimus, Sciaena. and Umbrina groups, but these relationships could be drawn at the tribal level (between lines C and D. Fig. 4). The suprageneric groups included in this clus- ter (Z. Fig. 4) show a basic carrot-shaped swim bladder with diverticula present or absent. Their sagittae are basically suboval and most members are bottom feeders. In this cluster of suprageneric groups, the trend of swim bladder development in western Atlantic sciaenids is clearly demonstrated from the simple Sciaena pattern with independent drumming muscles in the male to the complex Pogonias pattern with intrinsic drumming muscles in both sexes. Furthermore, the ontogenetic development of the swim bladder in Pogonias cromis (Fig. 10) and Sciaenops ocellata (Fig. 11) may also re- flect the phylogenetic development of the swim bladder, from simple (primitive) to complex (advanced) with diverticula developing on the swim bladder from the anterior to the posterior end. The Menticirrhus and Lonchurus groups resemble each other in the trend of reduced swim bladders (Figs. 14, 17) in adults. The sagittae are elongate and thin (Figs. 22B, 24). Both groups have inferior mouths and barbels. Their body shapes are elongate and rounded. The flat ventral side of the body is an adaptation for in- habiting the bottom habitats or specifically the surf zone (Table 1). Whether these similarities should be inter- preted as phylogenetic relationships or merely ecological convergence is not clear at present, but their close kin- ship is apparent. The Menticirrhus group also has a pored single barbel (Fig. 36B) similar to that of Um- brina (Fig. 36C), but this probably should be viewed as ecologically convergent for bottom feeding rather than as phylogenetically important. Other suprageneric groups of western Atlantic sciaenids may not be as closely related as the groups al- ready discussed. Further clustering of the taxa (between lines A and B, Fig. 4) is more difficult due to the limita- tions of a regional study. There are gaps, especially in assessing the relationships of the Micropogonias and Nebris groups. Nebris microps has a pair of well- developed anterior diverticula on its swim bladder (Fig. 8) and a very thick oval sagitta (Fig. 21). Micropogonias species have a pair of posteriorly originating tubelike diverticula on their swim bladders (Fig. 9) and rather thick, shieldlike and irregular sagitta (Fig. 26). At pre- sent, I think that Nebris is more closely related to the Z cluster (Fig. 4) than is Micropogonias. The clustering is based mainly on the position of the diverticula. The swim bladder diverticula of Micropogonias are developed more posteriorly than in the Nebris group. Both of them have a sagitta different from the members of the Z clus- ter (Fig. 4; also see Table 1). Grouping of the Cynoscion group with the Menti- cirrhus and Lonchurus groups (Y cluster, Fig. 4) is based on the basic shape of the main chamber of the swim bladder (in the Cynoscion and Lonchurus groups) and their elongate thin sagittae (in all). Although the mem- bers of the Lonchurus group have a pair of posteriorly directed diverticula on their swim bladders (Fig. 14), the main chambers have two well-developed anterior branches resembling those of the Cynoscion group (Fig. 12). Especially in the reduced form of Lonchurus lanceolatus (Fig. 14A), the anterior branches remain un- changed. This cluster (Y) is a convenient grouping and it may not reflect phyletic relationships (also see Table 1). The X and Y clusters on Figure 4 are more closely related to each other phyletically than to the Stellifer group, because they all have a single chambered swim bladder and only the sagitta enlarged (Table 1). The swim bladder of the Stellifer group has an additional chamber (Fig. 15) in front of the main carrot-shaped chamber. This yoke-shaped anterior chamber is located in front of the septum transversum and its anterior ends reach the skull. The sagitta of the Stellifer group is reduced in size and the lapillus is enlarged (Fig. 27). The sequences of sagitta reduction and lapillus enlargement are evident among modern genera of the Stellifer group. The unique swim bladder and otolith characters of the Stellifer group have not been reported in sciaenid fishes of geographic regions other than the New World (Win- tersteen and Chao, see footnote 3). The Indo-west Pacific sciaenid genera, Johnius and Wak are characterized by a "T-shaped" swim bladder but lack a constriction (Chu et al. 1963). Their sagittae also show a reduction at the an- terior portion of the ostium. Possible phyletic links of the Stellifer group with Johnius and Wak are not clear at present. In conclusion, western Atlantic Sciaenidae can be readily divided into two groups. One group is charac- terized by a swim bladder with two chambers and with two pairs of enlarged otoliths (lapillus and sagitta). The other group is characterized by a single chambered swim bladder and only one pair of enlarged otoliths (sagitta). Further divisions are based mainly on swim bladder structure, secondly on the morphology of the sagittae, and thirdly on external morphology. Swim bladder structure may be graded from simple to complex. Such a gradation is also reflected in the onto- genetic changes of several species such as Pogonias cromis (Fig. 10) and Sciaenops ocellata (Fig. 11). The primitive condition of the swim bladder (simple Sciaena pattern) exists in juveniles and adults of many species. Further development of the swim bladder is determined by the development of the diverticula from the anterior to the posterior end of the main chamber. Whether the reduction in size or loss of the swim bladder in adults of Menticirrhus and Lonchurus groups (Figs. 7, 14) is at- tributed to phylogenetic relationships or ecological con- vergence is undetermined at present. The general morphologies of the sagitta and its sulcus were used to determine relationships of different taxa es- pecially at the generic level. In the primitive condition. 46 the sagitta is thick and the sulcus opens to the anterior margin of the sagitta, i.e., the expanded part of the os- tium reaches the anterior margin (Sciaena pattern, Fig. 18). This is the most generalized pattern in many groups of western Atlantic sciaenids. The external morphology of the western Atlantic Sciaenidae, especially the mouth positions and body shapes, are more diverse than in any other perciform family in the region. These characters are more or less adaptive and plastic, but may be used to supplement other characters in assessing the relationships of the western Atlantic sciaenids. Biochemical characters of parvalbumins have been studied by Sullivan et al. (1975) and Rao et al. 5 to assess the systematic relationships of some sciaenid species 'Rao, K. V. R., C. L. Mahajan, L. Pennell, B. Sullivan, J. Bonaventura, and C. Bonaventura. The structure and evolution of parvalbumins. II. Differential rates of evolution among sciaenid isoparvalbumins. Manu- script. from the Atlantic American coast. Biochemical data should contribute much to the current knowledge of phy- logenetic relationships of western Atlantic Sciaenidae. Four genera of Sciaenidae — Aplodinotus, Pachyurus, Pachypops, and Plagioscion — may live in freshwater for their entire life history. Fossil otoliths of Plagioscion and Pachypops were described from Pliocene and Miocene deposits in Trinidad (Nolf 1976). This suggests that the freshwater sciaenids were widely distributed and may have originated in marine or estuarine environments. The modern freshwater sciaenids are endemic to the At- lantic drainages of the New World. Knowledge of their taxonomy and biology is rather sparse. Further studies on these freshwater sciaenids and on eastern Pacific sciaenids (Chao, see footnote 4) should supplement the present study, and only when this is done, shall we have adequate evidence for phylogenetic groupings. Based on the morphology of otoliths (Schmidt 1968; Nolf 1976), the family Sciaenidae is most similar to Lut- janidae, Pomadasyidae, Sparidae, and other closely related perciform fish families. KEY TO THE GENERA AND SPECIES OF WESTERN ATLANTIC SCIAENIDAE (WITH FRESHWATER AMERICAN GENERA) la. Lower jaw with one or more barbels (Fig. 32E-H) 2 2a. Only one barbel at tip of lower jaw 3 3a. Anal fin with one spine; body elongate fusiform, and rounded in cross section; swim blad- der absent or rudimentary in adult Menticirrhus 3b. Anal fin with two spines; body compressed, not fusiform; swim bladder well developed in adult 4 4a. Barbel stout and rigid, its tip rounded and perforated with an apical pore (Fig. 36C); mouth distinctly inferior; body with longitudinal stripes and vertical bars, without a well-defined black spot at pertoral fin base Umbrina 4b. Barbel slender and flexible, its tip tapering and without an apical pore (Fig. 36A); mouth slightly inferior, subterminal; body uniform in color, a very distinct and well-defined black spot at pectoral fin base. [D. X+I, 20-24; A. II, 7-9; gill rakers short, (7-9) + (13-17) = 21-25] Ctenosciaena gracilicirrhus (Central and South America from Honduras to Brazil) 2b. Two or more barbels on lower jaw 5 5a. Barbels only present on tip of lower jaw 6 6a. Lower jaw with two barbels only (Fig. 32F); eye small, 9 to 12 times in head length; pectoral fins very long, jet black, upper rays filamentous, reaching to caudal pe- duncle; swim bladder atrophied in adult (Fig. 14A); estuarine and marine. [D. X-XI+I, 37-39; A. II, 7-9; gill rakers short, (4-6) + (ll-13) = 15-18] . . . Lonchurus lanceolatus (Venezuela to Brazil) 6b. Lower jaw with three to many barbels; eye large, 3 to 5 times in head length; pec- 47 toral fins pale, upper rays not reaching beyond vent; swim bladder well developed in adult; freshwater and sometimes estuarine Pachypops (Freshwater South America) 5b. Barbels present at tip and a series of small barbels along rami (medial edges) of lower jaw 7 7a. Pairs of small barbels along rami of lower jaw, not in tuft at tip of lower jaw (Fig. 32G); eye diameter about 5 times in head length, second anal fin spine strong and long, more than two-thirds the length of first anal fin ray 8 8a. Preopercular margin strongly serrate; body covered with relatively small scales, 50 to 56 pored scales along lateral line; body silvery with pinkish cast, with many oblique stripes along the scale rows, swim bladder with a pair of tubelike diverticula (Fig. 9) Micropogonias 8b. Preopercular margin smooth; body covered with relatively large scales, 41 to 46 pored scales along lateral line; body gray to dark, juveniles with 4 to 5 broad vertical bars; swim bladder very complex with many interconnected divertic- ula (Fig. 10). [D. X+I, 19-23; A. II, 5-7; gill rakers short, (4-6) + ( 12-16) = 16- 21] Pogonias cromis (Gulf of Maine to Gulf of Mexico and southern Brazil to Argentina) 7b. Three pairs of barbels in a tuft at tip and more than 10 pairs barbels along rami of lower jaw (Fig. 32H); eye diameter about 9 times in head length, second anal fin spine weak and short, less than half the length of first anal fin ray Paralonchurus lb. Lower jaw without barbel (Fig. 32A-D) 9 9a. Mouth strongly oblique, tip of lower jaw projecting beyond upper jaw, preopercular margin smooth without strong spines (Fig. 37A, B) 10 10a. Eye very small, 8 to 11 times in head length (Fig. 37B); body rounded in cross section. [D. Vn+I, 31-32; A. II, 9-10; gill rakers long and slender, (5-9) + (14-15) = 20-24] . Nebris microps (Costa Rica to Brazil) 10b. Eye moderate to large, 3 to 6 times in head length (Fig. 37A); body compressed in cross section 11 11a. Body elongate and fusiform; teeth conical, set in narrow ridges on both jaws, usually with a pair of enlarged caninelike teeth at the tip of upper jaw; anal fin spines small, inconspicuous; swim bladder with a pair of large hornlike divertic- ula anteriorly (Fig. 12) 12 12a. Anterior (spinous) and posterior (soft) portions of dorsal fin well separated, with a gap in between; anal fin base about equal to the length of soft dorsal fin base; anal fin rays 16 to 20. [D. VII+I, 18-20; gill rakers moderately long, (2-3) + (7-9) = 9-12] Isopisthus parvipinnis (Panama to southern Brazil) 12b. Anterior (spinous) and posterior (soft) portion of dorsal fin, with a deep notch in between, but not separated by a gap; soft dorsal fin base much longer than anal fin base; anal fin rays 7 to 12 13 13a. Large caninelike teeth with arrow-headed tips present on both upper and lower jaws (Fig. 37A); lower jaw canines exposed externally when mouth closed. [D. X + I, 27-29; A. II, 8-9; gill rakers slender, (2-3) + (7-9) = 9-12J Macrodon ancylodon (Venezuela to southern Brazil) 48 13b. Large canines, if present, only at tip of upper jaw, none in lower jaw and tips never arrow-headed Cynoscion lib. Body short and robust; teeth minute, set in one to two rows on both jaws; without caninelike teeth; anal fin spines large, conspicuous; swim bladder simple or with a pair of small projections at the anterior end (Fig. 5E, F) Larimus 9b. Mouth not strongly oblique, tip of lower jaw usually not projecting beyond upper jaw, if so the predpercular margin with strong spines (Figs. 37C, 38) 14 14a. Preopercular margin usually serrate, sometimes strongly with one or more spines at the angle 15 15a. Scales along lateral line considerably enlarged but almost entirely concealed by small scales (Fig. 40); swim bladder with one chamber and a pair of anterior horns (Fig. 12D); lapillus (small earstone) small, less than 10% the size of sagitta (large earstone); freshwater and estuarine Plagioscion (Freshwater and estuarine South America) 15b. Scales along lateral line about same size as adjacent rows; swim bladder with two chambers, anterior one yoke-shaped and separated by a constriction from main posterior chamber (Fig. 15); lapillus (small earstone) large, more than 50% the size of sagitta (larger earstone) (Fig. 27); marine and estuarine 16 16a. Interorbital width usually more than 1.5 times of eye diameter, tip of an- terior swim bladder chamber terminating subcutaneously, frequently visible on superficial inspection in adult and juvenile under the upper end of operculum. Mouth inferior to terminal; snout usually projecting; dorsal view of head blunt, cavernous 17 17a. Anterior chamber of swim bladder with a pair of variably developed posterior diverticula (Fig. 16); bony interorbital width 3.5 or less in head Stellifer 17b. Anterior chamber of swim bladder without posterior diverticula; bony interorbital width 3.5 or more in head Ophioscion 16b. Interorbital width usually less than 1.2 times of eye diameter, tip of an- terior swim bladder chamber visible externally only in juvenile under the upper end of operculum. Mouth terminal; snout not projecting; dorsal view of head tapered, not cavernous Bairdiella 14b. Preopercular margin without obvious serrations, sometimes ciliated but never with spines at the angle 18 18a. Mouth small, horizontal, inferior; snout projecting in front of upper jaw; lower jaw included 19 19a. Body short, deep; dorsal profile strongly elevated at nape 20 20a. Body silvery with faint oblique stripes along the oblique scale rows dorsally; a dark spot above upper angle of gill slit; gill rakers 30 or more on first gill arch. [D. IX-XI+I, 29-35; A. II, 12-13; gill rakers (8-12) + (20-24) = 30-36] Leiostomus xanthurus (United States Atlantic and Gulf of Mexico coasts to Bay of Campeche, Mexico) 20b. Body pale to dark brown; with dark longitudinal stripes and/or oblique bars on sides, gill rakers less than 20 on first gill arch 21 21a. Height of anterior (spinous) portion of dorsal fin much greater 49 than head length; body pale brownish with three oblique bars on sides; third bar running from nape to caudal fin; dorsal fin with more than 45 soft rays Equetus 21b. Height of anterior (spinous) portion of dorsal fin less than head length; body pale brown to dark brown with longitudinal stripes on sides; dorsal fin with less than 40 soft rays Pareques 19b. Body elongate, dorsal profile not strongly elevated at nape 22 22a. Snout with peculiar conical appearance from the preorbital region being swollen and enlarged; maxillary almost entirely concealed under snout; freshwater Pachyurus (Freshwater South America) 22b. Snout rounded, not swollen, maxillary not concealed under snout 23 23a. Body elongate; one or more large dark spots (larger than eye) on the base of caudal peduncle and/or side of body in adult. [D. X + I, 23-25; A. II, 7-9; gill rakers (4-5) + (7-9) = 12-14] Sciaenops ocellata (United States Atlantic and Gulf of Mexico coasts to northern Mexico) 23b. Body robust; color uniform, no black spot on caudal pe- duncle nor on side of body. [D. IX-X+I, 27-33; A. II, 7; gill rakers 6+ (10- 12) = 16-18]; freshwater Aplodinotus grunniens (Freshwater, Canada to Guatemala) 18b. Mouth large, moderately oblique, terminal or subterminal; lower jaw not in- cluded 24 24a. A row of enlarged caninelike teeth present on both jaws; a distinct black blotch at pectoral fin base. [D. XI-XII+I, 23-26; A. II, 8-9; gill rakers long, (5-9) + (14-17) = 19-25] Odontoscion dentex (Florida, Antilles, along South American coast to Brazil) 24b. Without enlarged caninelike teeth on jaws; no distinct blotch at pectoral fin base Sciaena (Colombia and adjacent Caribbean Sea) Key to the Species of Bairdiella la. Preopercular margin with distinct strong spines at the angle, vertebrae 11 + 14 = 25; length of second anal fin spine less than 2.2 times in head 2 2a. Second anal fin spine very strong, about the length of first soft ray; preopercular margin with strong spinelike serrations. [D. X+I, 21-26 (usually 23-25); A. II, 7-9 (usually 8); gill rakers slender, (6-10) + (15-18) = 21-27 (usually 24-25)] B. ronchus (Antilles and from Hondurus to Brazil) 2b. Second anal fin spine not as strong, about two-thirds the length of first soft ray; preopercular margin with distinct strong spines. [D. X-XI+I, 19-23; A. II, 8-10 (usually 9); gill rakers slen- der, (7-8) + (14-16) = 22-24] B. chrysoura (U.S. Atlantic and Gulf of Mexico coasts) lb. Preopercular margin with only weak serrations, no spines at the angle, vertebrae 12+13 = 25; length of second anal fin spine more than 2.5 times in head 3 3a. Mouth slightly inferior; side with distinct stripes on scale rows; length of second anal fin spine 50 2.5-3.0 times in head; D. X+I, 25-29; A. II, 8; gill rakers rather short, (5-6) + (13-16) = 18-22 B. batabana (Southern Florida to Antilles) 3b. Mouth terminal; side without stripes; length of second anal fin spine 3.2-3.6 in head; D. X-XI +1, 22-24; A. II, 9; gill rakers moderately long, (7-8) + (16- 18) = 23-26 B. sanctaeluciae (Antilles. Honduras to Guyana, and occasionally in Florida) Key to the Species of Cynoscion la. Scales on body cycloid 2 2a. Posterior (soft) portion of dorsal fin covered with small scales beyond basal half of fin. [Lower jaw with a row of enlarged and widely spaced teeth; D. X+I, 22-25 (usually 23-24); A. II, 8-10 (mostly 9); gill rakers (2-3) + (6-8) = 8-ll; 150 to 160 scales in transverse row above lateral line; anterior horns of swim bladder long and straight (Fig. 13A) ] C. microlepidotus (Colombia to Brazil) 2b. Posterior (soft) portion of dorsal fin with only one or two rows of small scales at base 3 3a. Dorsal fin with 20 to 24 soft rays; lower jaw teeth uniform in size, closely set; A. II, 10-12 (mostly ID; gill rakers (2-3) + (5-8) = 7-ll; 115 to 125 scales in transverse row above lateral line; anterior horns of swim bladder strongly curved (Fig. 13D, D') sagitta without a notch on ventral margin C. leiarchus (Venezuela to Brazil) 3b. Dorsal fin with 27-31 soft rays; lower jaw teeth with a row slightly enlarged, widely set; A. II, 7-9 (mostly 8); gill rakers (l-3) + (6-8) = 7-ll; 120-130 transverse scale rows above lateral line; anterior horns of swim bladder slightly curved; sagitta with a notch on ven- tral margin (Fig. 23C) C. virescens (Honduras to Argentina) lb. Scales on body ctenoid 4 4a. More than 20 gill rakers on first gill arch. [D. X+I, 18-21; A. II, 8-9; gill rakers long, (7-9) + (14-17) = 21-26] C. striatus (Brazil to Argentina) 4b. Less than 20 gill rakers on first gill arch 5 5a. Dorsal fin usually with 22 or fewer soft rays; anal fin usually with 8 soft rays. [D. X+I, 17-23; A. II, 7-9; gill rakers (2-6) + (8-10) = 10-16; posterior (soft) portion of dorsal fin with only one to two rows small scales at base; caudal fin rhomboidal in adult] C. acoupa (Nicaragua to Argentina) 5b. Dorsal fin usually with more than 22 soft rays; anal fin usually with 9 or more soft rays 6 6a. Posterior (soft) portion of dorsal fin covered with small scales up to or beyond basal third of fin, especially on membranes between anterior rays 7 7a. Caninelike teeth absent at the tip of upper jaw; pectoral fins usually shorter than pelvic fins, 2 times or more in head length. [D. X+I, 21-24; A. II, 10-12; gill rakers (3-5) + (8-10) = 11-15] C. steindachneri (Guyana to Brazil) 7b. A pair of large caninelike teeth present at the tip of upper jaw; pectoral fins longer than pelvic fins, less than 2 times in head length 8 51 8a. Side of body with small dark spots forming distinct oblique streaks above lateral line; D. X+I, 24-29; A. II, 10-13 (usually 12); gill rakers (4-5) + (10-12) = 14-17 C. regalis (Gulf of Maine to Florida, U.S.A) 8b. Side of body pale without distinct spots, sometimes with faint streaks above lateral line 9 9a. Anal fin with 10 to 12 soft rays (usually 11). [D. IX-X+I, 25-29; gill rakers (3-4) + (9-10) = 12-14 (or more); soft portion of dorsal fin covered with small scales up to basal half of fin] C. arenarius (Gulf of Mexico coast, U.S.A to Bay of Campeche, Mexico) 9b. Anal fin with 8 to 10 soft rays (usually 9) 10 10a. Soft portion of dorsal fin covered with small scales up to (or beyond) three-fourths of fin height; total vertebrae 25; D. X + I, 23-27; gill rakers (2-3) + (7-10) = 9-13 C. jamaicensis (Lesser Antilles and Panama to Argentina) 10b. Soft portion of dorsal fin covered with small scales less than two-thirds of soft dorsal fin height; total vertebrae 27; D. X+I, 26-31; gill rakers (3-4) + (8- 10) = 11-14 C. nothus (Chesapeake Bay to Texas, U.S.A to Bay of Campeche, Mexico) 6b. Posterior (soft) portion of dorsal fin with only one to two rows small scales at base [caudal fin truncate in adults] 11 11a. Sides of body scattered with large dark spots; D. IX-X+I, 24-28; A. II, 9-12; gill rakers (2-3) + (7-9) = 9-12 C. nebulosus (New York to Florida and Gulf of Mexico coast, U.S.A. to northern Mexico) lib. Side of body without dark spots; D. IX-X+I, 24-29; A. II, 8-10; gill rakers (3-4) + (6-9) = 9-12 C. similis (Venezuela to Brazil) Key to the Species of Equetus la. Body with two narrow longitudinal stripes above and below the third oblique bar; dorsal, anal, and caudal fins dark brown with white spots; pectoral fin dark brown. [D. XI-XII+I, 45-47; A. II, 6-8; gill rakers short, 5+(10-13) = 15-18] E. punctatus (Bermudas, Florida, Antilles; and Panama to Brazil) lb. Body without narrow longitudinal stripes; broad oblique bars with distinct white margins; pectoral, dorsal, and anal fins pale. [D. XII-XIII+I, 47-55; A. II, 6; gill rakers short, (5-6) + (10-13) = 14- 18] E. lanceolatus (Bermudas, North Carolina to Florida, western Gulf of Mexico, Antilles; and Venezuela to Brazil) Key to the Species of Larimus la. Body dark gray above and silvery below without vertical bars on the sides; D. X+I, 26-28; gill rakers slender, longer than eye diameter, (9-ll) + (19-22) = 28-33 L. breviceps (Caribbean Islands to Brazil) lb. Body dark with 7 to 9 vertical bars on the sides; D. X+I, 24-27; gill rakers slender, about equal to eye diameter, (11- 13) + (22-25) = 34-36 L. fasciatus (U.S. Atlantic and Gulf of Mexico coasts) 52 Key to the Species of Menticirrhus la. Breast scales not uniform in size, those towards head notably smaller than scales along lateral line; molariform teeth present on phryngeal plates; pectoral fins short, not reaching beyond tip of pelvic fins; gill rakers in adult usually three or more on the lower limb of first branchial arch; color plain silvery gray. [Young fish (less than 100 mm SL) with only a vestige of swim bladder. D. IX-X+I, 19-26; gill rakers short, (3-5) + (0-8) = 3-12] M. littoralis (U.S. Atlantic coast, Gulf of Mexico and Caribbean coast to Brazil) lb. Breast scales uniform in size; about as large as those along lateral line; no molariform teeth on pharyngeal plates; pectoral fin reaching to or beyond tip of pelvic fin; gill rakers in adult tuberculate or absent on the lower limb of first branchial arch; side with dark oblique bars 2 2a. Anal fin rays usually 7(6-8); depressed spinous portion of dorsal fin seldom extends past base of second soft ray in dorsal fin; longest dorsal spine 16.2-24.1% of SL; young fish (less than 100 mm SL) with only a vestige of swim bladder. [D. IX-X+I, 20-26; gill rakers short, (2-3) + (0- 7) = 2-10] M. americanus (U.S. Atlantic coast, Gulf of Mexico and Antilles, Caribbean coast to Argentina) 2b. Anal fin rays usually 8(7-9); depressed spinous portion of dorsal fin often extends past base of second soft ray in dorsal fin; longest dorsal spine 24.6-38.9% of SL; young fish (less than 100 mm SL) with well-developed swim bladder. [D. IX-X+I, 22-27; gill rakers short, (3-5) + (0- 7) = 3-12; a longitudinal dark stripe usually present below lateral line posteriorly M. saxatilis (U.S. Atlantic and Gulf of Mexico coasts to Yucatan, Mexico) Key to the Species of Micropogonias* la. Dark spots on scales above lateral line forming distinct continuous streaks particularly below the posterior (soft) portion of dorsal fin; 6 or 7 scales between dorsal fin origin and lateral line in vertical series. [D. X+I, 26-30 (usually 26-28); A. II, 7-8; gill rakers short, (7-9) + (12-15) = 21-25] .... M. furnieri (Antilles, Venezuela to Argentina) lb. Dark spots on scales above lateral line not forming distinct continuous streaks below the posterior (soft) portion of dorsal fin; 8 or 9 scales between dorsal fin origin and lateral line in vertical series. [D. X+I, 26-30 (usually 28-29); A. II, 7-9 (usually 8); gill rakers short, (8-10) + (14-18) = 22-29] . M. undulatus (Cape Code, Mass., to Bay of Campeche and maybe from southern Brazil to Argentina) Key to the Species of Ophioscion 1 la. Dorsal fin with 20 to 21 soft rays; pored lateral line scales 47-49; gill rakers (7-9) + 14= 21-23 0. panamensis (Only known from types, Panama) lb. Dorsal fin with 23 to 24 soft rays; pored lateral line scales 52-53; gill rakers (7-8) + (13-16) = 20-24 O. punctatissimus (Panama to Brazil) Key to the Species of Paralonchurus la. Pectoral fin short and pale, not extending beyond the tip of pelvic fin; eye diameter moderate (about 5 times in head); soft dorsal with 28-31 rays; body with vertical dark stripes and dark humeral spot above pectoral fin origin; vertebrae 11 + 18=29; gill rakers short, (3-5) + (6-9) = 10-14 P. brasiliensis (Venezuela to Argentina) 'This key is based on types, specimens examined, and literature descriptions of Jordan and Eigenmann (1889), Jordan and Evermann (1898), Meek and Hildebrand (1925), MacDonagh (1934), Cervigon (1966a), and Roux (1973). Scale counts failed to key out species in several cases, but color patterns were useful in all cases. The morphology of sagittae (Figs. 26, 41) is useful within limits (also see Remarks on Micropogonias in the "Synopsis section"). 'Also see Remarks on Ophioscion in the "Synopsis section." 53 lb. Pectoral fin very long and jet black, reaching to anal fin origin; eye diameter small (about 9 times in head); soft dorsal 31-33 rays; body uniform brown; vertebrae 10+15=25; gill rakers very short, (3-4) +(4-8) =7-11 P. elegans (Surinam to Brazil) Key to the Species of Pareques la. Body with a broad oblique bar running from the base of anterior (spinous) portion of dorsal fin to pelvic fins; only one longitudinal stripe present at the midline of sides, reaching almost to the tip of caudal fin. [D. X+I, 38-39; A. II, 7; gill rakers short, (5-6) + (12-13) = 17-19] Pareques sp. 8 (Gulf of Mexico) lb. Body without oblique bar; usually several distinct longitudinal stripes present on sides 2 2a. Side with 7 to 10 narrow longitudinal stripes narrower than pupil; young with an anteriorly directed, V-shaped dark bar connecting the orbits across top of the head; depressed anterior (spinous) portion of dorsal fin usually not reaching base of the fourth soft ray. [D. IX-X+I, 38-40; A. II, 7; gill rakers short (4-6) + (10-12) = 15-18] P. umbrosus (Chesapeake Bay to Florida and Gulf of Mexico coast, U.S.A.). 2b. Side with 3 to 5 broad longitudinal stripes wider than pupil; and narrow stripes in between them; young with a straight dark bar connecting the orbits across the top of the head; de- pressed anterior (spinous) portion of dorsal fin usually reaching base of sixth soft ray. [D. VIII-IX+I, 37-41; A. II, 7-8; gill rakers short, (5-6) + (9-14) = 14-20] P. acuminatum (Chesapeake Bay to Gulf of Mexico, U.S.A., south to Bay of Campeche, Mexico; Antilles; and Honduras to Brazil). Key to the Species of Sciaena la. Dorsal fin with 24-26soft rays; gill rakers (6-7) + (13-15) = 19-21 S. trewavasae (Colombia and adjacent Caribbean Sea) lb. Dorsal fin with 21-23 soft rays; gill rakers (7-9) + (14-19) = 22-27 S. bathytatos (Colombia and adjacent Caribbean Sea) Key to the Species of Stellifer la. Preopercular margin with two or three strongly developed spines at the angle 2 2a. Propercular margin with three strong spines (occasionally four); gill rakers (12-14) + (20- 25) = 32-38; D. XI+I, 18-20; A. II, 8. [Pectoral fin about 28.5 to 31% of SL (72-106 mm SL specimens)] S. stellifer (Venezuela to Brazil, one record from Puerto Rico) 2b. Preopercular margin with two strong spines; gill rakers on first arch 40 or more. [D. XI-XH+I, 21-24; A. II, 8-9] 3 3a. Gill rakers (16-21) + (23-30) = 40-49 (rarely more than 50); tip of lower jaw with a dark band inside dentary teeth; interorbital width usually less than 2.8 in head; pectoral fin length usually less than 30% of SL S. rastrifer (Venezuela to Brazil) 3b. Gill rakers (20-23) + (32-36)=52-59; tip of lower jaw without a dark band inside dentary teeth; interorbital width usually more than 2.8 in head; pectoral fin length usually more more than 30% of SL S. griseus (Venezuela and Trinidad) 'Manuscript species of L. Woods (FMNH) and G. Miller (NMFS, Miami). 54 lb. Preopercular margin with four or more strongly developed spines at the angle 4 4a. Gill rakers 37 to 41; snout 4.1 to 4.5 in head length; roof of mouth black, [D. XI+I, 23-24; A. II, 8-9] Stellifer sp. A (Venezuela, Wintersteen and Chao, see footnote 3) 4b. Gill rakers 36 or fewer; snout 3.2 or less in head length; roof of mouth pale 5 5a. Gill rakers 28 or fewer; upper jaw gape length 9 2.8 or more in head length, mouth more or less ventral in position; when mouth closed, snout projecting in front of premaxillae (upper jaw) 6 6a. Scales on top of head mostly cycloid; pelvic fins moderately long, its tip reaching (or almost) to vent; with a medial row of slightly enlarged teeth on lower jaw; swim bladder with tubular diverticula (Fig. 16C, D) 7 7a. Gill rakers (7-9) + (13-16) = 20-24; D. X-XI+I, 19-21; A. II, 8-10 (usually 9); diverticula on swim bladder short, digitlike (Fig. 16C) S. microps (Colombia to Brazil, and West Indies) 7b. Gill rakers 8+ ( 14-16) = 22-24; D. X+I, 22; A. II, 9; diverticula on swim bladder very long, U-shaped (Fig. 16D) S. brasiliensis (Brazil) 6b. Scales on top of head ctenoid; pelvic fins relatively short, tip falling far short of vent; without an enlarged medial row of teeth on lower jaw; swim bladder with bulb or bean-shaped diverticula (Fig. 16A) 8 8a. Pelvic fin 5.3 to 6.0 in standard length, its tip ends slightly anterior to the tip of pectoral fin; eye 3.7 to 4.1 in head; gill rakers (8-9) + (15-16) = 23-25 [D. XI+I, 20-22; A. II, 8] S. naso (Venezuela to Brazil) 8b. Pelvic fin 6.8 in standard length, its tip ends much anterior to tip of pectoral fin; eye 4.6 to 5.3 in head; gill rakers (9-10) + (16-19) = 26-28 [D. XI-XH+I, 21-22; A. II, 8 (rarely 9)] S. venezuelae (Venezuela) 5b. Gill rakers 29 or more; upper jaw gape length (see footnote 9) 2.6 or less in head length, mouth more or less oblique and terminal in position; when mouth closed, snout equal to or projecting slightly in front of premaxillae (upper jaw) 9 9a. Underside of lower jaw with 4 pores; gill arch dark; gill rakers at the corner distinc- tively longer than the filaments; posterior diverticula of the anterior swim bladder chamber short, pear-shaped [ head cavernous, but not spongy to touch, tip of upper lip on horizontal with or above ventral margin of eye; snout usually not projecting beyond upper lip; D.XI + II, 20; gill raker (ll-12)+(18-21)=30-33;eye 5.5-6.3 times in head] Stellifer sp. B. (French Guiana and Northern Brazil; Wintersteen and Chao, see footnote 3) 9b. Underside of lower jaw with 6 pores; gill arch pale; gill rakers at the corner about equal or short of the filaments; posterior diverticula of the anterior swim bladder chamber small, knoblike 10 10a. Head extremely cavernous, spongy to touch, tip of upper lip on horizontal with or above ventral margin of eye; snout usually not projecting beyond upper lip; D. XI-XII+I, 20-24; gill rakers (10-13) + (22-23) = 32-36; eye 4.7-5.5 "Upper jaw gape length is measured from the tip of premaxillary to the corner of mouth (junction of premaxillary and dentary bones). 55 times in head S. lanceolatus (Chesapeake Bay to Gulf of Mexico coasts, U.S.A.) 10b. Head cavernous, but not spongy to touch; tip of upper lip usually on horizon- tal line with or well below ventral margin of eye; snout projecting slightly beyond upper lip; D. XI+I, 23-24; gill rakers (10-12) + (19-22) = 29-33; eye 5.5- 6.2 times in head S. colonensis (Panama, Puerto Rico and Haiti) Key to the Species of Umbrina la. Anal fin rays 7 or 8; mental (chin) barbel short; total gill rakers 19 to 20 2 2a. Eye larger, about 9.8 to 10.7% of SL; caudal peduncle circumferential scales 22; mental barbel with an apical pore; D. X+I, 24-25; A. II, 7; gill rakers (8-9) + (12-13) = 20-22 U. canosai (Southern Brazil to Argentina) 2b. Eye smaller, about 5.9 to 6.2% of SL; caudal peduncle circumferential scales 18 or 19; mental barbel with a pore on the middle anterior surface; D. X+I, 22-23; A. II, 8; gill rakers (7-8) + (ll- 13) = 19-20 U. milliae (Offshore Colombia and adjacent Caribbean seas) lb. Anal fin rays 6; mental (chin) barbel relatively long and slender; total gill rakers 13 to 15 3 3a. Scale rows beneath spinous dorsal fin and longitudinal stripes approximately parallel to lateral line; scales in diagonal series from dorsal fin origin to lateral line 5 or 6; stripes on body less distinct; D. X+I, 23-26; gill rakers (5-7) + (7-10) = 13-15 U. broussonetii (Antilles) 3b. Scale rows beneath spinous dorsal fin and longitudinal stripes situated at an angle of about 30° to lateral line; scales in diagonal series from dorsal fin origin to lateral line usually 8, some- times 7; stripes on body distinct; D. X+I, 26-31 (except for specimens from western Gulf of Mexico, which have slightly lower counts, 24-26); gill rakers (5-7) + (7-10) = 13-15 U. coroides (Gulf of Mexico to Brazil) ACKNOWLEDGMENTS I thank my graduate committee chairman, J. A. Musick, and members, D. F. Boesch, B. B. Collette, G. C. Grant, P. A. Haefner, Jr., and J. V. Merriner, for their help and guidance. Many people helped me to obtain museum specimens and fresh material, especially those who worked as museum assistants and crew members of various research vessels. To those who remain anony- mous, I express my appreciation. For permission to examine specimens in their institu- tions or for making specimens available, I thank: J. W. Atz (AMNH), M.-L. Bauchot (MNHN), M. Boeseman (RMNH), J. E. and E. B. Bohlke (ANSP), B. B. Collette (NMFS Systematics Laboratory, Washington, D.C.), W. E. Eschmeyer (CAS), C. R. Gilbert (UF), G. S. Glodek (FMNH), K. E. Hartel (MCZ), L. M. Hirsch (AMNH), T. Iwamoto (CAS), R. K. Johnson (FMNH), S. Kar- nella (USNM), P. Kahsbauer (NHMV), L. W. Knapp (SOSC), E. A. Lachner (USNM), K. F. Liem (MCZ), J. D. McEachran (Texas A&M University), R. V. Miller (NMFS, Washington, D.C.), J. G. Nielsen (ZMK), H. Nyssen (ZMA), T. W. Pietsch (formerly at MCZ), C. R. Robins (UMML), D. E. Rosen (AMNH), C. Roux (MNHN), R. Schoknecht (MCZ), C. L. Smith (AMNH), W. G. Saul (ANSP), W. Smith-Vaniz (ANSP), V. G. Springer (USNM), C. Swiff (LACM), W. R. Taylor (USNM), E. Trewavas (BMNH), S. H. Weitzman (USNM), and A. Wheeler (BMNH). Field sampling was essential to this study. I thank D. Flescher, M. Grosslein, H. Jansen, and H. Boyer of NMFS, Woods Hole, Mass., and T. Azarovitz, A. Pachaeco, and M. Silverman of NMFS, Sandy Hook, N.J., for inviting me to participate on various oceano- graphic cruises. I also thank the crew members of RV Al- batross IV, Woods Hole; Atlantic Twin, Sandy Hook; RV Eastward, Beaufott, N.C.; MV Oregon I and Oregon II Miami, Fla., and Pascagoula, Miss. I also express my appreciation to the following persons at VIMS: R. Bradley, J. Gilley, and M. Williams for preparation of graphs; W. H. Kriete, Jr., J. Colvocores- ses, D. F. Markle, J. E. Illowsky, and J. Green for help with field work; D. A. Sprinkle and J. F. Millen for typ- ing drafts of this manuscript; my fellow graduate students, K. Able, J. Colvocoresses, E. Lawler, D. Markle, J. McEachran, L. Mercer, G. Sedberry, and C. 56 Wenner for allowing me to discuss with them about sciaenids and for their helpful suggestions; and my major professor, J. Musick, for his courage in bailing me out of trouble, for his good judgment in allowing me freedom to do this work, and for his friendship. Large specimens of Sciaenops ocellata, which were very important to this study, were caught by C. A. Wenner. Early or obscure references vital to this study, were ob- tained through much appreciated efforts of: B. B. Collette and D. M. Cohen (NMFS Systematics Laboratory, Washington, D.C.); J. Marquardt and his staff at the Smithsonian library; S. Barrick and her staff at VIMS library; E. B. and J. E. Bohlke (ANSP); and L. P. Woods (FMNH). J. E. Fitch, California Department of Fish and Game, provided valuable comments and materials on the otolith section. E. Trewavas (BMNH) offered advice in all stages of this study. J. Wintersteen kindly offered in- formation on the genus Stellifer. A. Wheeler (BMNH), J. Nielsen (ZMK), and M. Bauchot (MNHN) helped by ex- amining type-specimens in their care. G. Clipper and J. L. Russo (NMFS Systematics Laboratory, Washington, D.C.) helped in taking photos and radiographs of type- specimens at USNM. B. B. Collette, C. R. Gilbert, D. F. Markle, J. A. Musick, and E. Trewavas have reviewed drafts of this manuscript and made very helpful comments. D. E. McAllister, National Museums of Canada, reviewed the final draft. J. McConnell and her staff at the Word Processing Centre, National Museums of Canada, Government of Canada, typed the final draft. I also ap- preciate a postdoctoral fellowship from the National Research Council, Canada (1976-78) which gave me an opportunity to finish this study. LITERATURE CITED AGASSIZ, L. 1829-1831. In J. B. Spix and L. Agassiz, Selecta genera et piscium quos in itinere per Brasiliam peracto colleget et pingendos curavit J. B. de Spix. Monachii. 138 p., 101 pis. (Part II, p. 83-138, pis. 46-76, published in 1821.) 1854. Notice of a collection of fishes from the southern bend of the Tennessee River, in the State of Alabama. Am. J. Sci. Arts, Ser. 2, 17:297-308. ARTEDI, P. 1738. In C. Linnaeus (editor), Iehthyologia sive Opera Omnia de Piscibus. [Five papers in one.] Lungduni Batavorum, apud conradum wishoff. BEARDEN, C. M. 1963. A contribution to the biology of the king whitings, genus Menticirrhus, of South Carolina. Contrib. Bears Bluff Lab. 38, 27 p. BERG, C. 1895. Enumeracion sistematica y sinonimica de los peces de las costas Argentina y Uruguaya. Ann. Mus. Nac. Buenos Aires 4:1-102. BLEEKER, P. 1865. Poissons du Japon, du Cap de Bonne Esperance et de Suri- name, conserves au Musee de liede. Ned. Tijdschr. Dierkd. 2:250-269. 1873. Description de deux especes nouvelles de Sciaenoidea de Surinam. Arch. Neerl. Sci. Nat. 5:456-461. 1874. Memoire sur les Scienoides et les Sillaginoides de 1'Inde achi- pelagique. Verh. Akad. Amst. 14:1-76. 1876. Systema Percarum revisum, Part II. Arch. Neerl. Sci. Nat. 11:289-340. BLOCH, M. E. 1788. Beskrivning over tvende nve Aborrer (Perca) fra Indien. K. Dan. Selsk. Skr. N. Saml. Copenh. 3:383-385. 1785-1795. Naturgeschichte der auslandischen Fishe. 9 pts. Berlin. [Also French translation, in Laveau, 1797, Ichthyologie, ou histoire naturelle, generale et paticuliere des poissons. Avec des figures enluminees, dessines d'apres nature. 12 pts. in 6 vols., 452 pis. Berlin.] BLOCH, M. E., and J. G. SCHNEIDER. 1801. Systema ichthyologiae. Berlin, 584 p. BOCOURT, M. 1869. Description de quelques reptiles et poissons nouveaux ap- partenant a la faune tropicale de l'Amerique. Nouv. Arch. Mus. Hist. Nat. Paris, Bull. 5:19-24. BOESEMAN, M. 1948. On new, little known, or otherwise interesting species of Suri- nam fishes. Proc. K. Ned. Akad. Wet., Amst. 51(3):3-11. BOIE, B. F. 1826. Oder encyclopaedische Zeitung. Isis (Jena) 19:970-982. BOLIN, R. L. 1947. The evolution of the marine Cottidae of California, with a discussion of the genus as a systematic category. Stanford Ich- thyol. Bull. 3:153-168. BONAPARTE, C. 1831. Saggio d'una distribuzione metodica degli animali vertebrati a sangue freddo. G. Acad. Sci. Lett. Art., Rome 53:129-209. BOSC, M. 1816. Ancylodon. In Nouveau dictionnaire d'histoire naturelle. 2nd ed. Vol. 1. Paris. [See p. 497.] BOULENGER, G. A. 1895. On a collection of fishes formed by Dr. C. Ternetz at various localities in Matto Grosso and Paraguay, with descriptions of new species. Proc. Zool. Soc. Lond. 1895:523-529. CALDWELL. D. K. 1958. A new fish of the genus Ophioscion, family Sciaenidae, from Caribbean Costa Rica. Q. J. Fla. Acad. Sci. 21:117-124. CAMPOS, A. A. 1942. Sciaenidae da agua doce: estudo das especies que habita os rios de Brasil. Arq. Mus. Parana. 2(ll):9-22. CASTELNAU, F. L. 1855. Animaux nouveaux ou rares recueillis pendant l'expedition dans les parties centrales de l'Amerique du Sud, de Rio de Janeiro a Lima, et de Lima au Para. Tome 2, Poissons, 112 p. Bertrand, ed., Paris. CATESBY, M. 1743. The natural history of Carolina, Florida, and the Bahama Is- lands, S.C.. Revised ed. Vol. II, 100 p., 20-p. appendix. Pri- vately published, London. CERVIGONM..F. 1966a. Los peces marinos de Venezuela. Fondo Cult. Cient. Cara- cas 2:449-951. 1966b. Una nueva especie de Stellifer de las costal de Venezuela (Pisces: Sciaenidae). Contrib. Ocas. Mus. Hist. Nat. La Salle, ser. zool. No. 24, 4 p. CERVIGON M., F., and H. DE HOLANDA LIMA. 1972. Las especies del genero Cynoscion (Pisces: Sciaenidae). Contrib. Cient. Univ. Oriente., Porlamar, Venez., No. 3, 44 p. CHAO L. N. 1976. Aspects of systematics, morphology, life history and feeding of western Atlantic Sciaenidae (Pisces: Perciformes). Ph.D. Thesis, College of William and Mary, Williamsburg, Va., 342 p. [Diss. Abstr. 37(5):2105-B (order no. 76-25, 354).] CHAO, L. N., and R. V. MILLER. 1975. Two new species of sciaenid fishes (tribe: Sciaenini) from the Caribbean Sea and adjacent waters. Bull. Mar. Sci. 25:259-271. CHAO, L. N., and J. A. MUSICK. 1977. Life history, feeding habits and functional morphology of juvenile scianid fishes in the York River estuary, Virginia. Fish. Bull., U.S. 75:657-702. 57 CHI". YUAN-TING, LO YUN-LING, and WU HAN-LING. 1963. Monographs of fishes of China. A study on the classification of the sciaenoid fishes of China, with description of new genera and species. Sci. Tech. Press. Shanghai, 100 p. (Engl, sum m. p. 83- 94). [Engl, transl. by L. N. Chao (J. A. Musick and L. P. Mer- cer, editors). 1975. Transl. Ser. 27, Virginia Institute of Marine Science. 91 p.] COPE. E. D. 1S67. On some new species of American and African fishes. Trans. Am. Philos. Soc. Phila. 13:400-407. CUVIER, G. 1S17. Le regne animal. Tome II. 532 p. Paris. 1829. Le regne animal. Nouvelle ed. Tome II, 406 p. Paris. 1830. In G. Cuvier and A. Valenciennes, Histoire naturelle des poissons. Vol. 5. 499 p. F. G. Levrault, Paris. DAHL. G. 1971. Los peces del norte de Colombia. INDERENA, Bogota, 391 p. DANERI. C. A. 1954. Una nueva especie de corvina de aqua dulce Plagioscion mac- donaghi n. sp. (Pisces, Sciaenidae). Comun. Inst. Invest. Mus. Argent. Cienc. Nat. Buenos Aires. Zool. 2:179-188. 1956. Pachyurus paranensis n.sp.. de corvina de agua dulce. Com. Jornadas Icticas 1956. Dir. Princ. Biol. Anim. Zootec. Santa Fe, 16 p. (Not seen.) DELSMAN. H. C. 1941. Resultats scientifiques des croisieres du Navire-ecole Beige "Mercator". Vol. II, No. 3, Pisces. Mem. Mus. R. Hist. Nat. Belg.. Ser. 2. Fasc. 21:47-82. DESMAREST. A. G. 1823. Premiere decade ichthyologique, ou description complete de dix especes de poissons nouvelles, ou imparfaitement connues, habitant la mer qui baigne les cotes de File cuba. Mem. Soc. Linn.. Paris 2:163-210. DEVINCENZI. G. J. 1924. Peces del Uruguay. II. Nuestra fauna ictiologica segun neustras coloecciones (continuacion). Ann. Mus. Nac. Montevi- deo. Ser. 2. Pt. 5:139-290. DIJKGRAAF. S. 1947. Ein Tone erzeugender Fisch im Neapler Aquarium. Experi- entia (Basel) 3:493-494. DRUZHININ. A. D. 1974. The distribution, biology and fisheries of croakers. [In Rus- sian.] Akad. SSSR Moscow, Izd. Pishch. Prom-st', 120 p. EDWARDS. G. 1751. A natural history of birds. Pt. 4, 248 p. [Fishes in the appendix. | EIGENMANN. C. H. 1912. The freshwater fishes of British Guiana, including a study of the ecological grouping of species, and the relation of the fauna of the plateau to that of the lowlands. Mem. Carnegie Mus. 5:1- 554. FISH. M. P.. and W. H. MOWBRAY. 1970. Sounds of western north Atlantic fishes: a reference file of biological underwater sounds. John Hopkins Press, Baltimore and Lond., 207 p. FOWLER, H. 1916. Cold-blooded vertebrates from Costa Rica and the Canal Zone. Proc. Acad. Nat. Sci. Phila. 68:389-439. 1937. A collection of Haytian fishes obtained by Mr. Stanley Wood- ward. Proc. Acad. Nat. Sci. Phila. 89:309-315. 1942. A list of the fishes known from the coast of Brazil. Arq. Zool. Estado. Sao Paulo 3:115-184. 1944. The fishes. In Results of the fifth George Vanderbilt expedi- tion (1941) (Bahamas, Carribean Sea, Panama, Galapagos Archi- pelago and Mexican Pacific Islands), p. 57-529. Acad. Nat. Sci. Phila., Monogr. No. 6. 1954. Os peixes de agua doce do Brasil. Arq. Zool. Estado. Sao Paulo 9:1-400. FOWLER. H. W.. and B. A. BEAN. 1924. Descriptions of eighteen new species of fishes from the Wilkes exploring expedition preserved in the United States National Museum. Proc. U.S. Natl. Mus. 63:1-27. FRIZZELL, D. L„ and J. H. DANTE. 1965. Otoliths of some early Cenozoic fishes of the Gulf coast. J. Paleontol. 39:687-718. GILBERT. C. R. 1966. Western Atlantic sciaenid fishes of the genus Umbrina Bull. Mar. Sci. 16:230-258. GILL, T. 1862a. Catalogue of the fishes of the eastern coast of North Amer- ica, from Greenland to Georgia. Proc. Acad. Nat. Sci. Phila. 13: 1-63. 1862b. Revision of the genera of North American Sciaeninae. Proc. Acad. Nat. Sci. Phila. 13:79-89. 1862c. On the Liostominae. Proc. Acad. Nat. Sci. Phila. 13:89-93. 1862d. On the Haploidonotinae. Proc. Acad. Nat. Sci. Phila. 13: 100-105. 1863. Notes on the sciaenoids of California. Proc. Acad. Nat. Sci. Phila. 14:16-18. 1864a. Catalogue of the North American sciaenoid fishes. Proc. Acad. Nat. Sci. Phila. 15:28-32. 1864b. Descriptive enumeration of a collection of Fishes from the western coast of Central America, presented to the Smithsonian Institution by Capt. John M. Dow. Proc. Acad. Nat. Sci. Phila. 15:162-174. 1873. Catalogue of the fishes of the east coast of North America. Smithson. Misc. Collect. 283, 50 p. GINSBURG, I. 1929. Review of the weakfishes (Cynoscion) of the Atlantic and Gulf coasts of the United States, with a description of a new spe- cies. Bull. U.S. Bur. Fish. 45:71-85. 1952. Eight new fishes from the Gulf Coast of the United States, with two new genera and notes on the geographic distribution. J. Wash. Acad. Sci. 42:84-101. GIRARD, C. 1859. Notes upon various new genera and new species of fishes, in the museum of the Smithsonian Institution, and collected in con- nection with the United States and Mexican Boundary Survey. Proc. Acad. Nat. Sci. Phila. 10:167-171. GISTEL, J. 1848. Naturgeschichte des Thierreichs. Fur hohere Schulen. Scheitlin and Krais, Stuttgart, 216 p. GOODE, G. B. 1876. Catalogue of the fishes of the Bermudas. Based chiefly upon the collections of the United States National Museum. Bull. U.S. Natl. Mus. No. 5, 82 p. GRONOVIUS, L. T. 1854. Catalogue of fish collected and described by Laurence Theo- dore Grownow, now in the British Museum. J. E. Gray (edited), Lond., 196 p. GUNTHER, A. 1860. Catalogue of the acanthopterygian fishes in the collection of the British Museum. Vol. 2, 548 p. 1867. On the fishes of the states of Central America; founded upon specimens collected in the fresh and marine waters of various parts of that country by Messrs. Salvin, Godman and Capt. J. M. Dow. Proc. Zool. Soc. Lond., 1866, Pt. 3:600-604. 1880. Report on the shore fishes produced during the voyage of H.M.S. Challenger in the years 1873-1876. Zool. Challenger Ex- ped. Pt. 6, 82 p. HECK EL, J. J. 1840. Johann Natterer's neue Fliissfische Brasilien's, nach den Beobachtungen und Mittheilungen des Entdeckers brescrieben. Ann. Mus. Wien 2:327-470. HOLBROOK, J. E. 1855. Ichthyology of South Carolina. Charleston, S.C., 182 p. [Second edition, 1860, 205 p.] HUBBS, C. L., and K. F. LAGLER. 1964. Fishes of the Great Lakes region. Rev. ed. University Michigan Press, 213 p. INTERNATIONAL COMMISSION FOR ZOOLOGICAL NOMEN- CLATURE. 1972. Opinion 988: Sciaena Linnaeus, 1758 (Pisces): Designation of a type-species under the plenary powers. Bull. Zool. Nomencl. 29:123-124. 58 IRWIN, R. J. 1970. Geographical variation, systematics, and general biology of shore fishes of the genus Menticirrhus, family Sciaenidae. Ph.D. Diss., Tulane Univ., 295 p. JARDIM, L. F. A. 1973. Sciaenideas marinhos da costa do Brasil Sul (Acanthoptery- gii-Perciformes-Sciaenidae). Comun. Mus. Cienc. PUCRGS, Ser. Zool. No. 3, 64 p. JENYNS, L. 1842. The zoology of the voyage of H.M.S. Beagle, Part 4, Fishes. London, 172 p. JOHNSON, M. W. 1948. Sound as a tool in marine ecology, from data on biological noises and the deep scattering layer. J. Mar. Res. 7:443-458. JONES, F. R. H., and N. B. MARSHALL. 195.3. The structure and functions of the teleostean swimbladder. Biol. Rev. 28:16-83. JORDAN, D. S. 1876. Manual of the vertebrates of the northern United States, in- cluding the district east of the Mississippi River, and north of North Carolina and Tennessee, exclusive of marine species. 1st ed. Chicago, 342 p. 1889. Scientific results of explorations by the U.S. Fish Commis- sion Steamer Albatross. No. IX — Catalogue of fishes collected at Port Castries, St. Lucia, by the steamer ALBATROSS, November, 1888. Proc. U.S. Natl. Mus. 12:645-652. 1917-1923. [reprinted 19631 The genera of fishes and a classifica- tion of fishes. Stanford LIniv. Press, Stanford, 800 p. JORDAN, D. S., and C. H. BOLLMAN. 1889. Scientific results of explorations by the LI.S. Fish Commission Steamer Albatross. No. IV. — Descriptions of new species of fishes collected at the Galapagos Islands and along the coast of the United States of Colombia, 1887-'88. Proc. U.S. Natl. Mus. 12: 149-183. JORDAN, D. S., and M. C. DICKERSON. 1908. Notes on a collection of fishes from the Gulf of Mexico at Vera Cruz and Tampico. Proc. U.S. Natl. Mus. 34:11-22. JORDAN, D. S„ and C. H. EIGENMANN. 1889. A review of the Sciaenidae of America and Europe. Rep. Comm., U.S. Coram, Fish Fish. 1886:343-451. JORDAN, D. S., and B. W. EVERMANN. 1896. A check-list of the fishes and fish-like vertebrates of North and Middle America. Rep. Comm. U.S. Comm. Fish Fish. 1895, 21:207-584. 1898. The fishes of North and Middle America. Bull. U.S. Natl. Mus. No. 47, Pt. 2:1241-2183. 1927. New genera and species of North America fishes. Proc. Calif. Acad. Sci. 16:501-507. KLEIN, J. T. 1749. Historia pescium naturalis. Vol. 5, 102 p. Leipzig. KNUDSEN, V. 0., R. S. ALFORD, and J. W. EMLING. 1948. Underwater ambient noise. J. Mar. Res. 7:410-429. LACEPEDE, B. G. E. V. 1798-1803. Histoire naturelle des poissons. 5 vols., Paris. [Vol. 1 (1798), 532 p.; vol. 2 (1800), 632 p.; vol. 3 (1802), 558 p.; vol. 4 (1803), 728 p.; vol. 5 (1803), 803 p.j LAL MOHAN, R. S. 1969. A synopsis to the Indian genera of the fishes of the family Sciaenidae. Indian J. Fish. 16:82-98. LeSUEUR, C. A. 1822. Description of three new species of the genus Sciaena. J. Acad. Nat. Sci. Phila., New Ser. 2:251-256. LINNAEUS, C. 1758. Systema naturae. Tome I. Regnum animale. 10th ed. Lip- siae, 824 p. 1766. Systema naturae. Tome I. Regnum animale. 12th ed. Vindobonae, 523 p. LOWE (McCONNELL), R. H. 1966. The sciaenid fishes of British Guiana. Bull. Mar. Sci. 16: 20-57. LUTKEN. C. F. 1875. Vehas-flodens fiske et bidrag til Brasiliens ichthyologi. Vidensk. Selsk. Skr., Copenh. 13:123-254. MACDONAGH, E. J. 1931. Sobre la corvina de San Bias: Macropogon patagonensis. Physis (Buenos Aires) 10:409-410. 1934. Nuevos conceptos sobre la distribucion geografica de los peces Argentinos basados en expediciones del Museo de La Plata. Rev. Mus. La Plata 34:21-170. McClelland, j. 1842. Remarks on a new genus of thoracic percoid fishes, Cestreus. J. Nat. Hist. (Calcutta) 2:150-152. McFARLAND, W. N. 1963. Seasonal changes in the number and the biomass of fishes from the surf at Mustang Island, Texas. Publ. Inst. Mar. Sci. Univ. Tex. 9:91-105. McPHAIL, J. D. 1961. A review of the tropical eastern Pacific species of Pareques (Sciaenidae). Copeia 1961:27-32. MARCGRAVE, G. 1648. Historia naturalis Brasilae. 2nd sect., 303 p. Amsterdam. [Translation by A. De E. Taunay, 1941, Sao Paulo, 293 p.] MEEK, S. E., and S. F. HILDEBRAND. 1925. The marine fishes of Panama. Field Mus. Nat. Hist. Publ. Zool. Ser. No. 226, 15(Pt. II):331-707. METZELAAR, J. 1919. Report on the fishes collected by Dr. J. Boeke in the Dutch West Indies 1904-1905. With comparative notes on marine fishes of tropical West Africa. In J. Boeke, Rapport betreffende een voorloopig onderzoek naar den toestand van de Visscherij en de Industrie van Zeeproducten in de Kolonie Curasao, etc. Pt. I, 314 p. Leiden. MILLER, R. V. 1971. A new sciaenid fish (Pisces: Umbrinini) with a single mental barbel, from the Southern Caribbean. Copeia 1971:300-306. MIRANDA RIBEIRO, A. de. 1915. Fauna Brasiliense. Peixes, Tomo V. (Eleutherobranchios aspirophros) — physoclisti. Arq. Mus. Nac. Rio de Janeiro, Vol. 17, 600 p. [Without continuous pagination, Sciaenidae, 46 P-] MIRANDA RIBEIRO, A. de. 1955. Tipos das especies e subespecies do prof. Alipio de Miranda Ribeiro depositados no Museu Nacional. Arq. Mus. Nac. Rio de Janeiro 42:389-417. MITCHILL, S. L. 1814. Fishes of New York. Report in part, reprinted with new introduction by T. Gill, 1898, Washington, 29 p. 1815. The fishes of New York. Trans. Lit. Phila. Soc, New York 1:355-492. MOHSIN, A. K. M. 1973. Comparative osteology of the weakfishes (Cynoscion) of the Atlantic and Gulf Coasts of the United States (Pisces-Sciaenidae). Ph.D. Diss., Texas A&M Univ., 158 p. MULLER, J., and F. H. TROSCHEL. 1848. Fische. In R. Schomburgk (edited by), Reisen in British- Guiana in den Jaren 1840-1844, p. 618-644. Leipzig. NAVAL ORDINANCE LABORATORY. 1942. Measurement of background noise in the water at Cape Henry, Va. due to surf and marine life. Nav. Ord. Lab. Rep., 594 p. 1944. Investigation of biological underwater background noises in vicinity of Beaufort, N.C. Nav. Ord. Lab. Rep., 880 p. NEAVE, S. A. (editor). 1939. Nomenclator zoologicus. Zool. Soc. Lond., 4 vols., 3805 p. [Vols. 1 and 2 published in 1939; vols. 3 and 4 in 1940.] NOLF, D. 1976. Les otolithes des teleosteens neogenes de Trinidad. Ecol- ogae Geol. Helv. 69:703-742. NORMAN, J. R. 1966. A draft synopsis of the orders, families and genera of recent fishes and fish-like vertebrates. Br. Mus., Lond., 649 p. OKEN, L. 1817. Order encyclopaedische Zeitung. Isis (Jena.) 8:1181-1183. PARKER, G. H. 1908. Structure and functions of the ear of the squeteague. Bull. [U.S.] Bur. Fish. 28:1211-1224. 59 PARRA. D. A. 1TS7. Descripcion de diferentes piezas de historia natural, las mas del ramo maritime, representadas en setenta y cinco laminas. Havana. Fishes. 66 p. POEV. F 1860-1861. Poissonsde Cuba, espees nouvelles. Vol. 2. In Memoria sobre la Historia natural de la Isla de Cuba. p. 115-356. Havana. 1875. Poissons de l'ile de Cuba. Especes nouvelles decrites. Ann. Lyceum Nat. Hist. New York 11:58-70. 1881. Peces. In J. Gudlach, Apuntes para la fauna puerto-riquena. Vol. 5, p. 317-350. An. Soc. Esp. Hist. Nac, Madrid. PUYO, J. 1949. Poissons de la Ouyane Francaise, Faune de l'Empire Fran- chise. Vol. 12. Off. Rech. Sci. Outre-Mer, Paris, 280 p. QUOY. J. R. C. and P. GAIMARD. 1824. Voyage autour du monde, . . . execute sur les corvettes de L.M. l'Uranie et la Physicienne, pendant les annees 1817-1820. Zoologie, Poissons. Paris, p. 183-401. RAFINESQUE-SCHMALTZ, C. S. 1815. Analyse de la nature ou tableau de l'univers et des corps or- ganises. Barravecchia, Palermo, 224 p. 1819. Prodrome de 70 nouveaux genres d'animaux decouverts dans 1'interieur des Etats-Unis d'Amerique durant l'annee 1818. J. Phys. Chim. Hist. Nat., Paris 88:417-429. 1820. Ichthyologia Ohiensis. Lexington, Ky., 90 p. RANDALL, J., and F. CERVIGON M. 1968. L>n neuvo pez Sciaenidae del genero Cynoscion de Venezuela y notas sobre C. jamaicensis y C. obliquatus. Mem. Soc. Cienc. Nat.. La Salle 27:176-192. REGAN, C. T. 1903. Descriptions of new South-American fishes in the collection of the British Museum. Ann. Mag. Nat. Hist., Ser. 7, 12:621-630. 1905. Description de six poissons nouveaux faisant partie de la col- lection du Musee d'histoire naturelle de Geneve. Rev. Suisse Zool. 13:389-393. REINHART. ■}. T. 1851. Nye sydamerikanske Ferskvandsfisk. Vidensk. Meddr. Dan. Naturhist. Foren. Kbh. 1849:29-57. 1855. Notits om Slaegten Pachyurus Agass. og de dertil h^rende Arter. Vidensk. Meddr. Dan. Naturhist. Foren. Kbh. 1854:108- 112. RICHARDSON, J. 1836. Fauna Boreali-Americana; or the zoology of the northern parts of British America., Part 3, The fish. London,_ 327 p. RINQUELET, R. A., R. H. ARAMBURU, and A. A. De ARAMBURU. 1967. Los peces Agentinos de aqua dulce. Com. Invest. Cienc, La Plata, 602 p. ROBINS, C. R. 1964. Sciaena bedoti and Corvina subaequalis, junior synonyms of Bairdiella ronchus, a western Atlantic sciaenid fish. Ann. Mag. Nat. Hist.. Ser. 13, 7:705-706. ROBINS, C. R., and D. C. TABB. 1965. Biological and taxonomic notes on the blue croaker, Bair- diella batabana. Bull. Mar. Sci. 15:495-511. ROUX, C. 1973. Resultats scientifiques des campagnes de la Calypso. Fax. X. Poissons teleosteens du plateau continental bresilien. An. Inst. Oceanogr. 49(suppl.):23-207. SAUVAGE, H. E. 1879. Description de quelques poisson9 d'especes nouvelles de la collection du Museum d'Histoire Naturelle. Bull. Soc. Philo- math. Paris 3:204-210. SCHINZ, H. R. 1822. Das Thierreich. Vol. 2. Fische. Stuttgart and Tubingen, 835 p. SCHMIDT, W. 1968. Vergleichend morphologische studie liber die Otolithen mar- iner Knochenfische. Arch. Fischereiwiss. 19(suppl.):l-95. SCHOMBURGK, R. H. 1860. Ichthyology. Fishes of British Guiana. Part II. In The Naturalist's Library. Vol. 40, p. 129-214. London. SCHULTZ, L. P. 1945. Three new sciaenid fishes of the genus Ophioscion from the Atlantic Coasts of Central and South America. Proc. U.S. Natl. Mus. 96:123-137. 1949. A further contribution to the ichthyology of Venezuela. Proc. U.S. Natl. Mus. 99:1-211. SEBA. A. 1761. Lncupletissimi rerum naturalium thesauri accurata de- scriptio. et iconibus artificiosissimus expressio, per universam physices historiam opus, cui ex tuto terrarum orbe collegit, diges- sit, descripsit et depingendum curavit A. Seba. Amstelaedami, vol. 3, 212 p. SMITH. H. M. 1905. The drumming of the drum-fishes (Sciaenidae). Science, New Ser. 22:376-378. STARK. J. 1828. Elements of natural history. Edinb. and Lond., 527 p. STARKS. E. C. 1913. The fishes of the Stanford expedition to Brazil. Stanford Univ. Publ., Univ. Ser., 77 p. STEINDACHNER, F. 1863. Beitrage zur Kenntniss der Sciaenoiden Brasiliens und der Cyprinodonten Mejikos. Sitzungsber. Akad. Wiss. Wien 48:162- 185. 1864. Ichthyologische Notizen I. Sitzungsber. Akad. Wiss. Wien 49:200-214. 1867. Ichthyologische Notizen VI. vii. Uber einige neue order sel- tene Fischarten von Westindien und Surinam. Sitzungsber. Akad. Wiss. Wien 56:347-357. 1870. Ichthyologische Notizen VIII. Sitzungsber. Akad. Wiss. Wien 60:120-191. 1875. Ichthyologische Bietrage II. (ii) Uber einige neue Fischarten von der Ost-und West-Kiiste Sud-Amerikas. Sitzungsber. Akad. Wiss. Wien 71:468-480. 1877. Ichthyologische Bietrage V. (iii) Uber einige Meeresfische von den Kusten Brasiliens. Sitzungsber. Akad. Wiss. Wien 74: 167-176. 1878. Zur FischFauna des Magdalena-Stromes. Denksch. Akad. Wiss. Wien 39:19-78. 1879. Uber einige neue und seltene Fisch-Arten aus den k.k. zoolo- gischen Museen zu Wien, Stuttgart und Warschau. Denkschr. Akad. Wiss. Wien 41:1-52. 1880. Ichthyologische Bietrage VIII. (i) Beschreibungen von 48 Fischarten. Sitzungsber. Akad. Wiss. Wien 80:119-191. 1917. Ichthyologische Beitrage XIX. Uber die Arten der Gattung Plagioscion Gill (= Diplolepis Steind.). Stizungsber. Akad. Wiss. Wien 126:657-676. STINTON, F. C. 1975. Fish otoliths from the English Eocene. Palaeontogr. Soc. (Monogr.), Lond., p. 1-56. SULLIVAN, B.. J. BONAVENTURA, and C. BONAVENTURA. 1975. The structure and evolution of parvalbumins. I. Amino acid compositional studies of parvalbumins from four perciform spe- cies. J. Mol. Evol. 5:103-116. TA VOLGA, W. N. (editor). 1964. Marine bio-acoustics. N.Y., vol. I, 413 p.; vol. 2, 353 p. TAYLOR, W. R. 1967. An enzyme method of clearing and staining small vertebrates. Proc. U.S. Natl. Mus. 122(3596), 17 p. TOPP, R. W., and C. F. COLE. 1968. An osteological study of the sciaenid genus, Sciaenops Gill (Teleostei, Sciaenidae). Bull. Mar. Sci. 18:902-945. TOWER, R. W. 1908. The production of sound in the drumfishes, the sea-robin and the toadfish. Ann. N.Y. Acad. Sci. 18:149-180. TRAVASSOS, H. 1973. Pescadas e corvinas de aguos marinhos Brasileiras. Bol. Mer. Pesq. 5(718):233-235. TRAVASSOS, H., and M. P. PAIVA. 1957. Lista dos Sciaenidae marinhos barsileiros, contendo chave de identificacao e proposta de "nomes vulgares oficias". Bol. Inst. Oceanogr. Sao Paulo 8:139-169. TRAVASSOS, H., and R. REGO-BARROS. 1971. Lista das especies Brasileiras de familia Sciaenidae Owen, 1846 (Perciformes, Percoide). Arq. Cienc. Mar 11(2):59-71. 60 TRRWAVAS, E. 1950. The status of the American mullets, Mugil brasiliensis and M. curema. Copeia 1950:149. 1962. A hasis for classifying the sciaenid fishes of tropical West Africa. Ann. Mag. Nat. Hist., Ser. 13, 5:167-176. 1963. Sciaena Linnaeus, 1758 (Pisces); proposed validation under the plenary powers of the ruling given in Opinion 93 concerning the type-species. Z.N. (S.) 850. Bull. Zool. Nomencl. 20:349-360. 1964. The sciaenid fishes with a single mental barbel. Copeia 1964:107-117. 1966. Comments on type-species of Sciaena Linnaeus Z.N. (S.) 850. Bull. Zool. Nomencl. 23:4-5. 1977. The sciaenid fishes (croakers or drums) of Indo- West-Pacific. Trans. Zool. Soc. Lond. 33:253-541. VAILLANT, L. L., and F. BOCOURT. 1915. Mission scientifique au Mexique et dans l'Amerique Centrale, recherches zoologiques. Etudes sur les Poissons Pt. 4:201-265. Paris. VALENCIENNES, A. 1833. In G, Cuvier and A. Valenciennes, Histoire naturelle des poissons. Vol. 9, p. 475-481. F. G, Levrault, Paris. VAZZOLER, A. E. A. de M. 1969. Ictiofauna de al Bahia de Santos. I. Sciaenidae (Percoidea. Percomorphi). CARPAS, Doc. Tech., No. 15, 19 p. WALBAUM, J. J. 1793. J.T. Kleinii ichthyologia enodata, sive index rerum ad his- torian! piscium naturalem synonymis recentissimorum systemati- corum explicatus. Lipsiae, 114 p. WHEELER, A. C. 1958. The Gronovius Fish Collection: A catalogue and historical account. Bull. Br. Mus. (Nat. Hist.), Hist. Ser. 1:185-249. 61 SYSTEMATIC INDEX Amblodon 41 concinnus 42 lineatus 42 neglectus 42 Amblyscion 31 argenteus 31 Ancylodon 36, 37 atricauda 37 jaculidens 37 parvipinnis 37 Aplodinotus 8, 14, 22, 25, 41, 47 grunniens 41, 43, 44, 45, 50 Apseudobranchus 34 Archoscion 34 petranus 36 Argyrosomus regium 6 Atractoscion aequidens 10, 14 Attilus 29 Bahaba falvolabiata 14 Bairdiella .... 10, 13, 20, 23, 24, 25, 26, 38, 39, 40, 45, 49, 50 batabana 38, 39, 51 chrysoleuca 40 chrysoura 25, 38, 39, 50 ronchus 25, 38, 39, 50 sanctaeluciae 25, 38, 39, 51 uerae-crucis 39 Bodianus argyroleucus 38, 39 costatus 32 exiguus 39 fulvus punctatus 39 pallidus 39 stellifer 40, 41 Buccone 34 Cestreus 34 carolinensis 34, 36 predatorius 34 steindachneri 36 Chaetodon lanceolatus 27 Cheilodipterus acoupa 34, 35 Cheironebris 31 Cirrimens 30 Corvina 43 adusta 40 biloba 42 crawfordi 32, 33 dentex 39 fulgen 39 gilli 40 grunniens 43 macrops 39 microps 41 monacantha 44 oscula 42 richardsoni 41, 42 ronchus 39 subaequalis 39 trispinosa 41 vermicularis 39 Corvula 38, 39 macrops 38 sanctaeluciae 39 sialis 39 Ctenosciaena 8, 14, 23, 25, 26, 29, 44, 45 gracilicirrhus 29, 44, 47 Cynoscion .... 10, 11, 13, 15, 20, 21, 22, 24, 25, 26 27, 34, 37, 43, 45, 46, 49, 51 acoupa 35, 51 arenarius 35, 36, 52 jamaicensis 35, 36, 52 leiarchus 35, 36, 51 maracaiboensis 35 microlepidotus 35, 36, 51 nebulosus 34, 35, 36, 52 nothus 35, 36, 52 regalis 13, 35, 36, 52 similis 35, 36, 52 steindachneri 34, 35, 36, 51 striatus 35, 36, 51 virescens 35, 36, 51 Cyprinus americanus 30 Daysciaena albida 27 Diplolepis 43 squamosissimus 43 Diplolepsis 43 Dipterodon chrysourus 38, 39 Eques 27 acuminatum 28 acuminatus umbrosus 28 americanus 27 balteatus 28 lineatus 28 pulches 28 punctatus 28 umbrosus 28 Equetus 8, 14, 24, 25, 26, 27, 44, 50, 52 lanceolatus 27, 52 punctatus 27, 28, 52 Eriscion 34 Eutychelithus 41 Genyanemus brasiliensis 38 Grammistes acuminatus 28 Haploidonotus 41 Homoprion 40 lanceolatus 40, 41 xanthurus 39 Isopisthus 10, 18, 20, 21, 24, 25, 34, 37, 45 af finis 37 harroweri 37 parvipinnis 37, 45, 48 Johnius 46 amazonicus 44 auratus 43 batabanus 38, 39 courvina 44 62 heterolepis 43 regalis 34, 36 saxatilis 31 Labrus cromis 34 maculatus 36 squeteague 36 squeteague var. maculatus 36 Larimus ..... 13, 14, 20, 21, 22, 24, 26, 27, 31, 33, 45, 46, 49, 52 breviceps 8, 13, 31, 45, 52 fasciatus 8, 13, 31, 52 Leiostomus 8, 14, 24, 25, 26, 27, 29, 44 humeralis 29 xanthurus 14, 21, 29, 32, 36, 39, 49 Lepipterus 42 corvina 43 francisci 42 Liostomus 29 Lonchiurus 37 Lonchurus ... 8, 10, 11, 13, 14, 18, 20, 21, 24, 25, 26 27, 37, 42, 45, 46 ancylodon 36, 37 barbatus 37 depressus 37 lanceolatus 18, 37, 38, 45, 46, 47 Lutjanus cayennensis 35 triangulum 34 Macrodon . . 10, 19, 20, 21, 22, 24, 25, 34, 36, 37, 45 ancylodon 37, 45, 48 Menticirrhus . .6, 8, 13, 15, 18, 20, 21, 23, 25, 26, 27 30, 45, 46, 47, 48, 53 alburnus 30 americanus 9, 30, 31, 53 atlanticus 31 forcaliger 31 littoralis 9, 30, 31, 53 saxatilis 9, 30, 31, 53 Micropogon 3, 32 argenteus 32, 33 barretoi 32, 33 lineatus 32 ornatus 38 patagonensis 32, 33 trifilis 42 Micropogonias . . .9, 13, 19, 20, 21, 24, 25, 26, 32, 33, 40, 46, 48, 53 furnieri 19, 32, 33, 40, 53 undulatus 19, 32, 33, 53 Monosira 31 stahli 31 Mugil cepnalus 35 gigus 34 grunniens 34 obliquus 29 Nebris 9, 13, 15, 20, 21, 22, 24, 26, 31, 46 microps 31, 46, 48 occidentalis 31 Nector 40 Nomalus 37 Odontoscion ... 10, 13, 20, 21, 23, 24, 26, 38, 39, 45 dentex 39, 50 Ophioscion ... 10, 13, 20, 21, 23, 25, 26, 38, 39, 40 45, 49, 53 adusta 40 adustus 40 brasiliensis 41 costaricensis 41 panamensis 40, 53 punctatissimus 23, 40, 53 typicus 39 venezuelae 41 woodwardi 32, 40 Otolithinae 4, 5, 14 Otolithus analis 34 bairdi 34, 36 carolinensis 36 drummondi 36 guatucupa 36 jamaicensis 36 leiarchus 36 microlepidotus 36 microps 36 nebulosus 36 nothus 36 obliquatus 36 regalis 34 rhomboidalis 35 striatus 36 thalassinus 36 toe-roe 34, 35 virescens 36 Pachypops 14, 27, 41, 42, 45, 47, 48 adspersus 42 camposi 42 fourcroi 10, 42, 45 steindachneri 42 trifilis 42 Pachyurus 8, 14, 24, 25, 27, 41, 42, 47, 50 adsperus 42 biloba 42 bonariensis 8, 9, 13, 14, 42 francisci 42 grunniens 42, 43 lundii 42, 43 nattereri 43 paranensis 42, 43 schomburgkii 8, 14, 42, 43 squamipennis 42, 43 squamipinnis 43 Paralarimus 34 patagonicus 34, 35 Paralonchurus .... 10, 13, 14, 18, 23, 24, 26, 37, 38, 45, 48, 53 brasiliensis 18, 38, 45, 53 elegans 18, 38, 45, 54 goodei 38 63 petersi 38 rathbuni 38 Paraplesichthys 37 Pareques 8, 14, 24, 25, 26, 27, 28, 44, 50, 54 acuminatus 28, 54 umbrosus 28, 54 Paraques species 54 Perca alburnus 30, 31 fourcroi 42 lanceolata 37 ocellata 34 punctata 39 punctatus 39 undulata 32 undulatus 32 Plagioscion ... 10, 14, 22, 24, 34, 41, 43, 45, 47, 49 auratus 43 heterolepis 43 macdonaghi 43, 44 microps 43, 44 monacantha 43, 44 pauciradiatus 43, 44 squamosissimus 43, 44 surinamensis 14, 43, 44, 45 ternetzi 43, 44 Pogonathus 34 courbina 34 Pogonias .... 6, 9, 13, 14, 20, 21, 24, 25, 26, 27, 34, 44, 45, 46 cromis 34, 44, 45, 46, 48 fasciatta 34 Polycirrhus 38 dumerili 38 fasciatus 34 ruthbuni 38 Polyclemus 14, 38 Pseudosciaena surinamensis 44 Roccus comes 36 Roncador stearnsii 14 Sagenichthys 37 Sciaena . . 6, 8, 13, 14, 20, 21, 24, 25, 26, 27, 28, 29, 31, 35, 40, 44, 46, 50, 54 alburnus 30, 31 bathytatos 28, 54 bedoti 39 cirrosa 29 croker 32 dubia 29 edwardi 28 fusca 34 grisea 42 im berbis 34 magdalenae 44 multifasciata 29 nebulosa 31 operculars 32, 33 oscula 42 rubella 44 squamosissima 43, 44 trewavasae 28, 44, 54 umbra 28, 29 Sciaenidae 3, 4, 5, 6, 14, 21, 41, 47 western Atlantic 3, 6, 22, 26, 27, 37 Sciaeninae 4, 5, 6 Sciaenini 6, 28 Sciaenops . . .9, 10, 11, 13, 14, 21, 24, 25, 26, 27, 33, 34, 45, 46 ocellata 34, 46, 50 Sigmurus 39 Stellicarens 40 Stellifer . . . 10, 11, 13, 20, 21, 23, 24, 26, 27, 38, 40, 41, 43, 45, 46, 49, 54 brasiliensis 40, 41, 55 capensis 40 colonensis 25, 40, 41, 56 griseus 13, 25, 40, 41, 54 illecebrosus 40 lanceolatus 23, 26, 40, 41, 56 microps 13, 25, 26, 40, 41, 55 mecatoris 37 mindii 41 naso 40, 41, 55 oscitans 40 rastrifer 25, 26, 40, 41, 54 stellifer 25, 26, 40, 41, 54 venenzuelae 25, 40, 41, 55 zestocarus 40 Stellifer species 26, 40, 41, 55 Stelliferus 40 naso 41 rastrifer 41 Symphysoglyphus 34 Umbrina . . .6, 8, 14, 24, 25, 26, 29, 44, 45, 46, 47, 56 arenata 30 broussonetii 29, 56 canosai 29, 56 coroides 29, 30, 56 furnieri 32 gracilicirrhus 29 gracilis 30 januaria 30 littoralis 30, 31 martinicensis 30 milliae 29, 30, 44, 56 ophiocephalus 30 phalaena 30 Umbrula 30 Vacuoqua 38, 39 Wak 46 Zaclemus 38 Zestidium 40 Zestis 40 Zonoxcion 38 64 tr U S GOVERNMENT PRINTING OFFICE: 1976-797-733/21 REGION .188. Proceedings of the first U.S. -Japan meeting on aquaculture at Tokyo, -Japan, October 18-19, 1971. William N. Shaw (editor). (18 papers, 14 authors.) February 1974, iii 4- 133 p. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. .'!89. Marine flora and fauna of the northeastern United States. Crustacea: Decapoda. By Austin B. Williams. April 1974, iii + SO p.. Ill figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 390. Fishery publications, calendar year 1973: Lists and indexes. By Mary Ellen Engett and Lee 0. Thorson. September 1974, iv + 14 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 391. Calanoid copepods of the genera Spinncalanus and Mimncalanus from the central Arctic Ocean, with a review of the Spinocalanidae. By David M. Damkaer. .lune 1975, x + 88 p., 22.S figs., 4 tables. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington. D.C. 20402. 392. Fishery publications, calendar year 1974: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. -June 1975, iv + 27 p.. 1 fig. 393. Cooperative Gulf of Mexico estuarine inventory and study — Texas: Area description. Bv Richard A. Diener. September 1975, vi + 129 p., 55 figs., 2H tables. 394. Marine Flora and Fauna of the Northeastern United States. Tar- digrada. Bv Leland W. Pollock. May 1976, iii + 25 p., figs. For sale In i In 1 Superintendent of Documents, U.S. Government Printing Office. Washington, D.C. 20402. 395. Report of a colloquium on larval fish mortality studies and their relation to fishery research. January 1975. By John R. Hunter. May 1976. iii + 5 p. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. UNITED STATES DEPARTMENT OF COMMERCE C^-4. CCEANlC AND ATMOSPHERIC ADMINISTRATION n»TiOn», marine FISHERIES SERVICE SCIENTIFIC PUBLICATIONS STAFF ROOM 450 1107 N E 45TH ST SEATTLE. WA 98105 OFFICIAL BUSINESS PENN STATE UNIVERSITY LIBRARIES A000Q7201AT3LI NOAA SCIENTIFIC AND TECHNICAL PUBLICATIONS NOAA, the National Oceanic and Atmospheric Administration, was established as part of the Department of Commerce on October 3, 1970. The mission responsibilities of NOAA are to monitor and predict the state of the solid Earth, the oceans and their living resources, the atmosphere, and the space environment of the Earth, and to assess the socioeconomic impact of natural and technological changes in the environment. The six Major Line Components of NOAA regularly produce various types of scientific and technical infor- mation in the following kinds of publications: PROFESSIONAL PAPERS— Important definitive research results, major techniques, and special in- vestigations. TECHNICAL REPORTS— Journal quality with extensive details, mathematical developments, or data listings. TECHNICAL MEMORANDUMS— Reports of preliminary, partial, or negative research or tech- nology results, interim instructions, and the like. CONTRACT AND GRANT REPORTS— Reports prepared by contractors or grantees under NOAA sponsorship. TECHNICAL SERVICE PUBLICATIONS— These are publications containing data, observations, instructions, etc. A partial listing: Data serials; Pre- diction and outlook periodicals; Technical manuals, training papers, planning reports, and information serials: and Miscellaneous technical publications. ATLAS — Analysed data generally presented in the form of maps showing distribution of rainfall, chemical and physical conditions of oceans and at- mosphere, distribution of fishes and marine mam- mals, ionospheric conditions, etc. Information on availability of NOAA publications can be obtained from: ENVIRONMENTAL SCIENCE INFORMATION CENTER ENVIRONMENTAL DATA SERVICE NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION U.S. DEPARTMENT OF COMMERCE 3300 Whitehaven Street, N.W. Washington, D.C. 20235