Order Gasterosteiformes: Part 8 9781933789316

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Fishes of the Western North Atlantic

PUBLICATIONS OF THE SEARS FOUNDATION FOR MARINE RESEARCH, YALE UNIVERSITY The Sears Foundation for Marine Research at Yale University was established in 1937 by Albert E. Parr, director of Yale's Bingham Oceanographic Laboratory, through a gift from Henry Sears, to promote research and publication in marine sciences. The Foundation's Memoirs, inaugurated in 1948, remain important references. In 1959 the Bingham Oceanographic Collection was incorporated into the Yale Peabody Museum of Natural History. Distributed by Yale University Press www.yalebooks.com I yalebooks.co.uk MEMOIR I FISHES OF THE WESTERN NORTH ATLANTIC Part One Lancelets, Cyclostomes, Sharks Part Two Sawfishes, Guitarfishes, Skates and Rays, Chimaeroids Part Three Soft-rayed Bony Fishes: Orders Acipenseroidei, Lepisostei, and Isospondyli Sturgeons, Gars, Tarpon, Ladyfish, Bonefish, Salmon, Charrs, Anchovies, Herring, Shads, Smelt, Capelin, et al. Part Four Soft-rayed Bony Fishes: Orders Isospondyli and Giganturoidei Argentinoids, Stomiatoids, Pickerels, Bathylaconids, Giganturids Part Five Orders Iniomi and Lyomeri Lizardfishes, Other Iniomi, Deepsea Gulpers Part Six Orders Heteromi (Notacanthiformes), Berycomorphi (Beryciformes), Xenoberyces (Stephanoberyciformes), Anacanthini (Gadiformes) Halosauriforms, Killifishes, SquirreIrishes and Other Beryciforms, Stephanoberyciforms, Grenadiers Part Seven Order Iniomi (Myctophiformes) Neoscopelids, Lanternfishes, and Atlantic Mesopelagic Zoogeography Part Eight Order Gasterosteiformes Pipefishes and Seahorses Part Nine, Volume One Orders Anguilliformes and Saccopharyngiformes Part Nine, Volume Two Leptocephali Part Ten Order Beloniformes Needlefishes, Sauries, Half beaks, and Flyingfishes MEMOIR II THE ELEMENTARY CHEMICAL COMPOSITION OF MARINE ORGANISMS by A. P Vinogradov

MEMOIR SEARS FOUNDATION FOR MARINE RESEARCH Number I

Fishes of the Western North Atlantic

PART EIGHT

Order Gasterosteiformes, Suborder Syngnathoidei SYNGNATHIDAE (Doryrhamphinae, Syngnathinae, Hippocampinae)

NEW HAVEN SEARS FOUNDATION FOR MARINE RESEARCH, YALE UNIVERSITY

Yale ISBN 978-1-933789-18-7 (pbk.) ISBN 978-1-933789-31-6 (e-book) Issued in paperback by the Peabody Museum of Natural History, Yale University, New Haven, Connecticut 06511 USA Part Eight first published in hardcover in 1982 by the Sears Foundation for Marine Research, Yale University ISBN 978-0-912532-89-9 (cloth) Library of Congress Control Number: 49000120 Distributed by Yale University Press NEW HAVEN AND LONDON Printed in the United States of America Printed on acid-free paper

Fishes of the Western North Atlantic

Authors C. E. DAWSON Gulf Coast Research Laboratory Museum

RICHARD P. VARI Smithsonian Institution

NEW HAVEN SEARS FOUNDATION FOR MARINE RESEARCH, YALE UNIVERSITY

Editorial Board Editor-in-Chief

JAMES E. BOHLKE Academy of Natural Sciences Philadelphia, Pennsylvania

THEODORE W. PIETSCH

DANIEL M. COHEN

University of Washington Seattle, Washington

National Marine fisheries Service Seattle, Washington

BRUCE B. COLLETTE

WILLIAM J. RICHARDS

National Marine fisheries Service Miami, Florida

National Marine fisheries Service Washington, D.C.

WILLIAM N. ESCHMEYER

C. LAVETT SMITH

California Academy of Sciences San Francisco, California

American Museum of Natural History New York, New York

ROBERT H. GIBBS, JR.

KEITH S. THOMSON

Smithsonian Institution Washington, D.C.

- Ex Officio

Yale University New Haven, Connecticut

Editor Emeritus ALBERT E. PARR Hamden, Connecticut VI

Table of Contents INTRODUCTION Family Syngnathidae.

ix The Pipefishes. By C. E. DAWSON

General Characters Skeletal Characters Development Diversity and Range Economic Value Systematics Western Atlantic Syngnathidae Subfamilies Doryrhamphinae and Syngnathinae.

1 1 1 2 2 2 2 By C. E. DAWSON

Acknowledgments Western Atlantic Genera and Species Rejected Names Extralimital Species Methods Key to the Western Atlantic Genera Genus Amphelikturus A. dendriticus Genus Penetopteryx P. nanus Genus Oostethus O. brachyurus lineatus Genus Micrognathus Key to Western Atlantic Species of Micrognathus M. crinitus M. erugatus Genus Anarchopterus Key to the Species of Anarchopterus A. criniger A. tectus Genus Leptonotus L. blainvilleanus Genus Pseudophallus Key to the Species of Pseudophallus P. starksii P. mindii P. elcapitanensis vii

4 4 5 5 6 10 10 11 13 14 16 17 25 26 26 31 33 34 34 38 40 41 44 45 45 49 53

Table of Contents

viii

Genus Syngnathus Key to Western Atlantic Species of Syngnathus S. floridae S. springeri S. louisianae S. fuscus S. scovelli S. affinis S. makaxi S. folletti S. dawsoni S. pelagicus S. caribbaeus Genus Bryjc Key to Western Atlantic Species of Bryx B. dunckeri B. randalli Genus Cosmocampus Key to Western Atlantic Species of Cosmocampus C. albirostris C. profundus C. brachycephalus C. elucens C. hildebrandi Literature Cited Subfamily Hippocampinae.

The Seahorses.

55 56 57 67 70 76 83 90 93 95 101 103 108 HI 112 112 117 120 121 121 125 128 132 139 141

By RICHARD P. VARI

Acknowledgments Western Atlantic Genera and Species Terminology and Methods Genus Hippocampus Key to Western Atlantic Species of Hippocampus H. zosterae H. reidi H. erectus Literature Cited Index

173 173 173 175 175 176 178 181 185 190

Introduction ""PHIS eighth part of FISHES OF THE WESTERN NORTH ATLANTIC treats the family Syng-*• nathidae of the Order Gasterosteiformes. The general account of the family and the accounts of the species comprising the pipefish subfamilies Doryrhamphinae and Syngnathinae were prepared by C. E. Dawson. These were based upon a large amount of material, a considerable portion collected by the author. Included are four species not known from the area defined as western North Atlantic in this series. Two of the species of the genus Pseudophallus treated are eastern Pacific forms; these, however, live mostly in streams and rivers and the author thinks that there is a reasonable chance that either or both eventually will enter the Atlantic. Two other species included, Leptonotus blainvilleanus and Syngnathus folletti, are as yet known only from the southern West Atlantic. These complete the monographic treatment for all western Atlantic pipefishes, and at least Syngnathus folletti may be expected in the region we define as western North Atlantic. At any rate, there is John Tee-Van's statement in Part 1 of this series (p. xiv): "... this range [Hudson Bay to the Amazon Delta] is not strictly adhered to in all instances; a number of species living close to the outer borders of the region covered by this publication are included, particularly when their inclusion assists in a more adequate understanding of the group under consideration"; also, his comment in Part 2 of the series (p. xi): "... occasional species from adjoining seas are described." The account of the seahorse subfamily Hippocampinae with its three western Atlantic species was prepared by Richard P. Vari. These curious little fishes with prehensile tails are popular in the aquarium trade and also are widely sold as curios in a dried state. The following abbreviations have been used throughout in order to avoid repetition of names of natural history collections housing the preserved specimens upon which the accounts have been based. AMNH — American Museum of Natural History, New York. ANSP — Academy of Natural Sciences, Philadelphia. ASM — Alabama State Museum, Tuskaloosa. BMNH — British Museum (Natural History), London. BOC — Bingham Oceanographic Collection, Yale University, New Haven. CAS — California Academy of Sciences, San Francisco. CAS(SU) — Stanford University collections, now at California Academy of Sciences. CIMAR — Centro de Investigaciones del Mar, Universidad Catolica de Valparaiso, Chile. FDNR — Florida Department of Natural Resources, St. Petersburg. FLSM — Fundacion La Salle de Ciencias Naturales, Isla Margarita, Venezuela. FMNH — Field Museum of Natural History, Chicago. ix

Introduction FSM FSU GCRL GMBL HBF HMBL IBMC IRSNB LACM LCM LS MACN MBUCV MCZ MHNM MNHN MNMH MZUSP NMC NMW NYZS RMNH SHML SIO STRI UCLA UCR UDONECI UFP UMML UMMZ UNAM UNC UOIO UP UPR USA

— Florida State Museum, Gainesville. — Florida State University, Tallahassee. — Gulf Coast Research Laboratory Museum, Ocean Springs, Mississippi. — Grice Marine Biological Laboratory, Charleston, South Carolina. — Harbor Branch Foundation, Fort Pierce, Florida. — Huntsman Marine Biological Laboratory, St. Andrews, New Brunswick. — Institute de Biologia Marina, Mar del Plata, Argentina. — Institut Royal des Sciences Naturelles de Belgique, Brussels. — Los Angeles County Museum of Natural History. — Laboratorio de Ciencias do Mar, Universidade Federal do Ceara, Fortaleza, Brazil. — Linnaean Society, London. — Museo Argentina de Ciencias Naturales x/Bernardino Rivadavia," Buenos Aires. — Museo de Biologia, Universidad Central de Venezuela, Caracas. — Museum of Comparative Zoology, Harvard University. — Museo de Historia Natural, Montevideo, Uruguay. — Museum National d'Histoire Naturelle, Paris. — Museo Nacional de Historia Natural, Santiago, Chile. — Museu de Zoologia, Universidade de Sao Paulo, Brazil. — National Museum of Canada, Ottawa. — Naturhistorisches Museum Fischsammlung, Wien. — New York Zoological Society. — Rijksmuseum van Natuurlijke Historic, Leiden. — Sandy Hook Marine Laboratory, Highlands, New Jersey. — Scripps Institution of Oceanography, La Jolla, California. — Smithsonian Tropical Research Institute, Panama. — University of California at Los Angeles. — Universidad de Costa Rica, San Jose. — Centro de Investigaciones Cientificas del Niicleo de Nueva Esparta de la Universidad de Oriente, Boca de Rio, Isla Margarita, Venezuela. — Universidade Federal de Pernambuco, Brazil. — University of Miami, Marine Laboratory, Rosenstiel School of Marine and Atmospheric Sciences. — University of Michigan Museum of Zoology, Ann Arbor. — Universidad Nacional Autonoma, Mexico, D.F. — University of North Carolina, Institute of Marine Sciences, Morehead City. — Universidad de Oriente, Institute Oceanografico, Cumana, Venezuela. — Universidad de Panama, Panama. — University of Puerto Rico, Mayagiiez. — University of South Alabama, Mobile.

Introduction USC USNM UWF ZFMK ZMA ZMB ZML ZMUC

XI

— University of South Carolina, Baruch Coastal Research Institute, Columbia. — National Museum of Natural History, Washington, D.C. — University of West Florida, Pensacola. — Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn. — Zoologisch Museum, Amsterdam. — Zoologisches Museum, Museum fur Naturkunde der HumboldtUniversitat, Berlin. — Zoological Museum, Lund, Sweden. — Zoologisk Museum, University of Copenhagen.

The editors are grateful to the two authors for their contributions and to the home institutions of the authors for supporting their research. While the authors have acknowledged those who assisted them, it is appropriate here to reiterate the importance of the inter-institutional cooperation of curators of collections to make possible comprehensive studies as those in FISHES OF THE WESTERN NORTH ATLANTIC. We are indebted to the Sears Foundation for the publication of this volume. The Editor-in-Chief acknowledges the Academy of Natural Sciences for supporting his work with the volume and thanks Dr. Alfred E. Schuyler, also of the Academy, for preparing the manuscript for the printer. April, 1981 Philadelphia, Pennsylvania

JAMES E. BOHLKE Editor-in-Chief

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Family Syngnathidae C. E. DAWSON Gulf Coast Research Laboratory Museum Ocean Springs, Mississippi 39564

Acknowledgments. See those for the Pipefishes.

General Characters. Typically slender and elongate gasterosteiforms, with or without a prehensile tail, with body armored by dermal plates arranged in a more or less regular series of rings and with or without an external fleshy integument. Mouth small, terminal on protruding snout (except in Enchelyocampus). Jaws edentate but odontoid processes (Dawson and Fritzsche, 1975) are present on dentaries and premaxillae in some genera. Gill opening reduced to a pore in the opercular membrane and located above upper posterior angle of opercle, gill membranes otherwise fused to isthmus or body. Spinous dorsal and pelvic fins absent, other fins variously present or absent. Without pyloric sphincter or distinct stomach. Kidney aglomerular (except in Halicampus macrorhynchus; see Khalil, 1980), only present on the right side. Sexually dimorphic with males incubating eggs under trunk or tail. Eggs carried exposed on ventral surface of male or partially to completely concealed within a well-defined brood pouch. Pouch with or without ventrolateral extensions of lateral dermal plates (pouch plates), with or without bilateral membranous folds which

overlap the eggs and meet or nearly meet on ventral midline, or pouch developed as a sealed sac-like structure which opens to the exterior through an anterior pore. Skeletal Characters. Four complete gill arches, gill lobate, pseudobranchiae present, two infrapharyngobranchials present and without basisphenoid or metapterygoid. Posttemporals coossified with cranium, supracleithra and postcleithra absent. The three anterior vertebrae fused to form a compressed median longitudinal partition. Mobile vertebrae without articular processes (zygopophyses); ribs absent. Development. Larvae and early postlarvae usually replicate adults in gross morphology but there are significant differences between young and adults in some genera. Pectoral fins are present in larvae but absent in adults of the eastern Atlantic genus Nerophis, and both dorsal and pectoral fins are similarly lost during transition from larva to adult in both Penetopteryx and Apterygocampus (Dawson and Allen, 1978). The caudal fin is present in young of the prehensiletailed Indo-Pacific genus Haliichthys but this fin is absent in adults. It has also been shown (Dawson et al., 1979) that larvae and/or postlarvae of

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Memoir 1, Part 8—Sears Foundation for Marine Research

the Indo-Pacific genera Yozia and Trachyrhamphus have elongate dermal appendages, absent in adults, which presumably function as buoyant processes during the term of planktonic life. Diversity and Range. The Syngnathidae are distributed from boreal regions of the Atlantic and Pacific Oceans to the southern cold-temperate region. The family, including some 55 genera and ca. 200 species, is predominantly marine but freshwater and estuarine forms are not uncommon. A few are oceanic but the Syngnathidae are most common in coastal waters and within the 50 m depth range. Species abundance is perhaps highest in temperate-subtropical areas and greatest diversity (ca. 68% of recognized species) is found in the subtropical-tropical Indo-Pacific region. Economic Value. Syngnathids have limited economic value and are not presently considered an exploitable source of animal protein. Seahorses and pipefishes are sometimes dried or embedded in plastic for sale as curios and Syngnathidae are also utilized to some extent by the aquaria trade. The seahorse has long been a popular aquarium fish but increased demand for estuarine or freshwater pipefishes (e.g. species of Oostethus, Doryichthys, Pseudophallus) is to be expected. Syngnathids are recorded as food items for some sea birds and predatory fishes (e.g. tunas) but they are not known to constitute the principal or preferred diet of any species. Systematics. Since the first significant treatment of the family (Kaup, 1856), classification of the Syngnathidae has been based largely on differences in location and development of the eggdeposition area or brood pouch of mature males. This has been supplemented by differences in other major characters such as the presence or absence of certain fins or head ridges, as well as by differences in configuration of the principal body ridges. Duncker (1912, 1915) divided the Syngnathidae into two groups; those wherein eggs are deposited under the abdomen of males (Gastrophori) and those wherein eggs are deposited under the tail (Urophori). He further divided each group into three subfamilies on the basis

of differences in types of egg-deposition or brood-pouch morphology. Following Duncker, Herald (1943, 1959) included the subfamilies Nerophinae, Syngnathoidinae, and Doryrhamphinae in the Gastrophori and the Solegnathinae, Syngnathinae and Hippocampinae in the Urophori. The Nerophinae are characterized by having the eggs deposited loosely on the trunk and eggs are not individually isolated. The Syngnathoidinae and Solegnathinae have eggs individually isolated within a spongy mass or in membranous compartments. Eggs are similarly deposited in the Doryrhamphinae and Syngnathinae, but they are additionally protected with bilateral pouch plates and/or covering membranes which are not completely sealed along the ventral midline of the pouch. The Hippocampinae have eggs deposited within a sealed sac-like pouch which opens only through an anteromesial pore. Herald (1959), utilizing variations in types of pouch closure and lateral ridge configurations within the Syngnathinae, also postulated an evolutionary sequence leading from the presumably primitive open or semi-open pouch-closure of Corythoichthys (Syngnathinae) to the sealed pouch of seahorses (Hippocampinae). Present studies showing that type of pouch closure varies within some syngnathine species and description of a sealed hippocampine-like brood pouch in the Indo-Pacific syngnathine Apterygocampus (Dawson and Allen, 1978) show that Herald's (1959) phylogenetic concept and Duncker's (1912, 1915) subfamily designations require reexamination. Fritzsche's (1976, 1980) studies on branchial arches and those of Dawson and Allen (1978) and Dawson et al. (1979) on syngnathine larvae tend to further demonstrate the uncertain status of generally accepted concepts of relationships within the Syngnathidae. Western Atlantic Syngnathidae. Following Duncker (1912,1915) and Herald (1959), western Atlantic Syngnathids include one genus and species referable to the gastrophorine subfamily Doryrhamphinae, whereas urophorine forms include 9 genera and 26 species in the Syngnathinae and one genus and three species in the

Fishes of the Western North Atlantic Hippocampinae. In more practical terms, there are 10 genera and 27 species of pipefishes and one genus and three species of seahorses in the western Atlantic Ocean. The remainder of this

3

treatment has, for convenience, been divided between pipefishes and seahorses. These may be distinguished in the western Atlantic by the character combinations in the couplet below.

la. Caudal fin present; tail usually straight, not prehensile (except in Amphelikturus); longitudinal axes of head and trunk essentially parallel (head angled ca. 30° ventrad in Amphelikturus); brood pouch under trunk or tail, but not sac-like or opening only through an anteromesial pore. Pipefishes, p. 4 Ib. Caudal fin absent; tail coiled ventrad, prehensile; longitudinal axis of head angled ca. 70-90° from axis of body; brood pouch under tail, sac-like and opening only through an anteromesial pore. Seahorses, p. 173

THE PIPEFISHES (Subfamilies Doryrhamphinae and Syngnathinae) C. E. DAWSON Gulf Coast Research Laboratory Museum Ocean Springs, Mississippi 39564

Acknowledgments. For permission to examine specimens in their care, for prompt response to inquiries or loan requests and for other courtesies, I thank the following: D. E. Rosen and C. L. Smith (AMNH); J. E. Bohlke and W. F. Smith-Vaniz (ANSP); A. C. Wheeler and P. J. P. Whitehead (BMNH); K. Thompson (BOC); W. N. Eschmeyer and Pearl Sonoda (CAS); D. Prinz (FLSM); R. K. Johnson (FMNH); C. R. Gilbert (FSM); R. W. Yerger (FSU); W. D. Anderson, Jr. (GMBL); R. G. Gilmore, Jr. and R. S. Jones (HBF); E. E. Boschi and Maria Berta Cousseau (IBMC); J. P. Gosse (IRSNB); R. J. Lavenberg and C. C. Swift (LACM); R. B. Lopez (MACN); F. Mago, L. (MBUCV); W. Fink and K. E. Hartel (MCZ); MarieLouise Bauchot (MNHN); N. Bahamonde, N. (MNMH); N. Menezes (MZUSP); D. E. McAllister (NMC); R. Hacker and P. Kahsbauer (NMW); M. Boeseman (RMNH); R. H. Rosenblatt (SIO); I. Rubinoff (STRI); B. Walker (UCLA); W. A. Bussing (UCR); F. Cervigon, M. and A. G. Caspar (UDONECI); Aida M. Eskinazi (UFP); C R. Robins (UMML); R. M. Bailey and R. R. Miller (UMMZ); A. Resendez, M. (UNAM); F. J. Schwartz (UNC); G. Reyes-Vasquez (UOIO); L. Howell Rivero, formerly (UP); P. L. Colin (UPR); R. L. Shipp (USA); E. A. Lachner and associates (USNM); S. Bortone (UWF); K. H. Liiling (ZFMF); H. Nijssen (ZMA); H. J. Paepke (ZMB); and J. Nielsen (ZMUC). Appreciation is expressed for gift or exchange specimens received from many of these colleagues as well as from other students and professional biologists throughout the study area. Especially important material was made available by L. N. Chao and L. Pereira (Universidade do Rio Grande, Brazil), D. W. Greenfield (Northern Illinois Univeristy), L. W. Knapp (Smithsonian Oceanographic Sorting Center), P. S. Lobel (MCZ), D. W. Moore (Ichthyological Associates) and J. Villa (University of Missouri). Special acknowledgment is due B. Fernholm (Roskilde University) and J. Nielsen (ZMUC) for assistance with problems concerning Scandinavian literature and specimens. I thank W. I. Follett and Lillian Dempster (CAS) for advice on nomenclatural matters and A. C. Wheeler (BMNH) for arranging for my examination of Linnaean material. Pertinent records and unpublished notes of the late E. S. Herald were kindly made available

by W. N. Eschmeyer (CAS) and Mrs. E. S. Herald. Computer programming and related services were provided by D. Boyes and Mrs. Lois Coquet, GCRL Computer Section. Since this work has been complemented by information on the worldwide pipefish fauna, I also thank the many individuals and institutions that have otherwise contributed to my studies. This project would have suffered severely without the personal interest and cooperation of my associates in the GCRL Museum. Mrs. Elizabeth Heal has provided expert technical secretarial assistance and has cheerfully coped with a demanding work-load and many manuscript changes. Mr. F. N. Jackson has assisted in the photographing of specimens and has efficiently handled the receipt, curation and dispatch of innumerable specimen loans. The majority of drawings are by Mrs. Nancy Gordon but some were prepared by Mrs, Anne Langenfeld and Mrs. Dianne Yandell. For permission to reproduce Figures 50, 51, and 53,1 thank the Director of the Western Australian Museum. This study was supported in part by National Science Foundation Grant BMX 75-19502.

Western Atlantic Genera and Species. Twentyseven species of pipefishes are recognized in the western Atlantic Ocean. These belong in ten genera. One genus (Oostethus) is a trunk-pouch form (Gastrophori: Doryrhamphinae), whereas the remainder are tail-pouch genera (Urophori: Syngnathinae). Five genera (Amphelikturus, Leptonotus, Oostethus, Penetopteryx, Pseudophallus) are each represented by single species or subspecies. Three genera (Anarchopterus, Bryx, Micrognathus) are each represented by two species. There are five species referred to Cosmocampus and eleven to Syngnathus. One genus, Anarchopterus, is restricted to the western Atlantic Ocean. Another, Pseudophallus, is restricted to the Western Hemisphere. The majority are low-temperate to tropical

Fishes of the Western North Atlantic species. Only one (Leptonotus blainvilleanus) is known from below 40° S and only one (Syngnathus fuscus) occurs commonly above 40° N. Three species not known from the western North Atlantic are included here. Leptonotus blainvilleanus (South Atlantic) has been added to complete the coverage of western Atlantic species. Two species of Pseudophallus (starksii, elcapitanensis), occurring in the geologically unstable Pacific drainage of Central America, are not treated adequately in the literature and may, through stream capture, eventually occur in the Atlantic drainage. A number of other nominal genera and species have been recorded from western Atlantic waters but these represent rejected names or unconfirmed records of extralimital species. Rejected Names. Poey (1876) reported four varieties or undescribed species of Cuban pipefishes merely as Syngnathus with his personal species number appended to each. He also provided brief descriptions, without figures, of five species: Syngnathus flavirostris, S. picturatus, S. linea, S. marmoreus and S. ascendens. Jordan and Evermann (1896a), without examining the types (evidently lost), referred these names to the synonymy of Siphostoma elucens (Poey), and this action has generally been accepted by subsequent authors. Poey's descriptions consist mainly of total length, some proportional data, notes on coloration and some counts (except for picturatus). Color descriptions and given proportions are not diagnostic. Counts for four species (rings 16-17 + 33, dorsal-fin rays 24-25, subdorsal rings 1.5-0.5 + 4-4.5) could apply to Syngnathus pelagicus, S. caribbaeus and several species of Cosmocampus (including elucens). Furthermore, the description of Syngnathus ascendens states that the lateral trunk ridge is continuous with lateral tail ridge and confluent with superior tail ridge. This is assumed to be anomalous, since Poey's counts of 16 trunk rings and 1.5 + 4 subdorsal rings do not occur in Pseudophallus mindii, the only tropical western Atlantic pipefish with this lateral ridge configuration. Since these five nominal species cannot with certainty be referred to

5

Syngnathus elucens Poey or any western Atlantic pipefish, all are nomina dubia. Jordan (1884) introduced the name Siphostoma miurum Swain and Meek in his list of fishes from Key West, Florida. Later, Jordan (1886a) states that this was a manuscript name, that the holotype had been lost or destroyed and that the planned description had been suppressed. In the absence of description or illustration, Siphostoma miurum is a nomen nudum. Jordan and Evermann (1896a) introduced Siphostoma poeyi as a replacement name for the preoccupied Syngnathus tenuis (non Blyth 1859) Poey 1868. Duncker (1915) doubtfully referred this species to the synonymy of Microphis lineatus (=Oostethus brachyurus lineatus) and it has since been treated as a junior synonym by most authors. The holotype of Syngnathus tenuis Poey is lost, the species was not figured, but Poey did give counts of 30 dorsal-fin rays and 1 + 6 subdorsal rings for his single specimen. The dorsalfin ray count is 3 less than that of any known specimen of Oostethus and 8 less than that of any O. b. lineatus examined from the Antilles (Table VI). Furthermore, subdorsal trunk rings are 1.5 or more in all examined western Atlantic Oostethus (Table III). Since no clearly diagnositc characters were given in the original description and since these counts show, beyond a reasonable doubt, that the holotype was not conspecific with O. b. lineatus, Syngnathus tenuis Poey and Siphostoma poeyi Jordan and Evermann are nomina dubia. Extralimital Species. Peters (1869) described Syngnathus fistulatus from "Puerto Cabello, gesammelt von Appun." In addition to the holotype (ZMB 4348), there are evidently only seven other specimens referred to this species. All were cataloged prior to 1900, detailed collection data are lacking, and all are now in poor condition. Six are labeled "Jamaica" [CAS 24768 (1, 249); USNM 12688 (5, ca. 230-254)] and one, ca. 250 mm SL (MNHN 93-97), is labeled "Martinique" with notation of its receipt from "Musee de Macon." These fish have long snouts (snout length 1.6-1.7 in hi), lateral trunk and tail ridges are, variably, interrupted or confluent below

6

Memoir I , Part 8—Sears Foundation for Marine Research

dorsal-fin base, and principal meristic features are as follows: rings 17-20 + 39-43, dorsal-fin rays 35-37, subdorsal rings 1-0 + 6.5-8, pectoralfin rays 13-15, anal rays 3, brood pouch below 22-25 tail rings. This combination of characters does not occur in any western Atlantic pipefish but it is essentially diagnostic of the eastern Atlantic Syngnathus acus Linnaeus. There is some confusion as to the site of Peters' type-locality. Jordan and Evermann (1896b), without discussion and evidently without supporting evidence, state that Puerto Cabello was "near Aspinwall," an early name for Colon, Panama. This was accepted by Herald (1942) but he later questioned the matter (Herald, 1943), and Schultz (1949) stated that the type-locality was undoubtedly Puerto Cabello, Venezuela. The internal jar label of the holotype and the original catalog entry both show "Porto Cabello" but this has been crossed out in the catalog and remains uncorrected (H. J, Paepke, in litt.). Nothing can be here added toward identification of Peters' locality but it may well represent an error in labeling. In any event, Syngnathus fistulatus Peters is here considered conspecific with S. acus and, in the absence of confirmed records, this species is omitted from the western Atlantic pipefish fauna. The eastern North Atlantic snake pipefish, Entelurus aequoreus Linnaeus, has occasionally been included in the American pipefish fauna, but, as noted by Herald (1942), all such records are highly questionable. This species, as Osphyolax pellucidus, has been taken in mid-Atlantic waters (Cope, 1875) and has been reported, as Nerophis martinicensis from Martinique (Kaup, 1856; Dumeril, 1870), as Nerophis heckeli from Bogota (Kaup, 1856; Dumeril, 1870) and as Nerophis aequoreus from Brazil (Castelnau, 1855; Dumeril, 1870). In the absence of recent or confirmed records, Entelurus aequoreus is also omitted from the western Atlantic pipefish fauna. METHODS Fish Characters. The identification of western Atlantic pipefishes (best aided by X30 or higher

magnification) depends strongly on the accurate determination of the following: 1. Differences in the configuration of the principal body ridges (Fig. 1: STRR, ITRR, LTRR, STAR, ITAR) separate treated forms into four groups of 1-4 genera (Fig. 2). The ridge pattern is obscure in some genera (e.g. Anarchopterus) and may be difficult to determine in early juveniles of many species. In problem specimens, slight drying usually enhances the ridge pattern. The ridge configuration is a conservative character in present populations wherein unilateral anomalies occur in less than 5% and bilateral anomalies in less than 1% of examined material. 2. Presence or absence of dorsal, pectoral or anal fins. The dorsal and pectoral fins are variously present or absent in Penetopteryx and the anal fin is absent in Anarchopterus, Bryx, Penetopteryx and Pseudophallus. The anal fin is often minute and difficult to see. It may be concealed within the anterior portion of the brood pouch in mature males. 3. Presence, degree of development or absence of certain head ridges and dermal flaps (Fig. 1) are important factors in some determinations. Dermal flaps are absent in some genera (e.g. Syngnathus) but typically present in others (e.g. Anarchopterus). In the latter case, however, they may be sparse or, in poorly preserved material, completely lacking. 4. Counts of body rings (Fig. 1) are required for species determination. Trunk ring counts begin with the ring bearing the pectoral-fin base (pectoral cover plate of authors) and end with the ring bearing the anus (anal ring). Tail ring counts begin with the 1st ring behind the anus (usually bearing anal fin) and end with the penultimate ring, excluding the terminal element bearing the caudal fin. In the few cases where anus and anal fin occur on the same ring, the minimum value is recorded (e.g. 16 rather than 16.5). Counts of brood-pouch rings include all rings bearing any trace of pouch development; the pouch usually begins on the 1st tail ring in syngnathine species. The number of subdorsal rings is estimated to the nearest fourth of ring length and is indicated by the formula: trunk

Fishes of the Western North Atlantic MDHR.

7

PF

STRR

LTRR ! ITRR !—MVTRR

DF

DFO

STRR_: LTRR_ ITRR_

ITAR LTAR

FIGURE 1. Sections of generalized pipefish delineating morphological features referenced in text: AF, anal fin; DF, dorsal fin; DFL, dermal flap; DFO, dorsal-fin origin; GO, gill opening; ITAR, inferior tail ridge; ITRR, inferior trunk ridge; LSR, lateral snout ridge; LTAR, lateral tail ridge; LTRR, lateral trunk ridge; MDHR, median dorsal head ridges; MDSR, median dorsal snout ridge; MVTRR, median ventral trunk ridge; N, naris; OPR, opercular ridge; PF, pectoral fin; PFB, pectoral-fin base; S, scutellum; SOPR, supraopercular ridge; SOR, supraorbital ridge; STAR, superior tail ridge; STRR, superior trunk ridge; TARI, tail ring; TRRI, trunk ring.

rings covered by dorsal fin + tail rings covered = total subdorsal rings. Point of dorsal-fin origin is recorded in V4 ring intervals before (+) and behind (—) the anterior margin of the 1st tail ring (0-point). 5. All fin rays are counted separately. Present counts were usually made on both pectoral fins.

Data are tabulated as total frequencies of pectoral-fin rays and as paired or equivalent counts, wherein equal counts occurred in both left and right fins. Caudal-fin rays are typically 10 in syngnathine genera and 9 in the doryrhamphine genus Oostethus. Frequency of caudal-fin rays is a highly conservative character and

Memoir 1, Part 8—Sears Foundation for Marine Research

8

'/ // // * '' ' //'/'/'S''/' / / / / / / / / / / ' / / ' / , ' '','. *, / / / / / / / / / / / X / V X x>PENETOPTERYX

LEPTONOTUS PSEUDOPHALLUS

B

/ *

*

/

/

/

*S

OOSTETHUS AMPHELIKTURUS ANARCHOPTERUS MICROGNATHUS

BRYX COSMOCAMPUS SYNGNATHUS FIGURE 2. Lateral aspect of posterior trunk and anterior tail rings illustrating the four configurations of principal body ridges occurring in western Atlantic pipefishes.

counts differing from these indicate regeneration and concomitant high probability of an atypical tail-ring count.

6. Proportional data are based on the following measurements (mm): standard length (SL)— straight line distance from tip of lower jaw

Fishes of the Western North Atlantic (mouth closed) to articular base of median caudal-fin rays; head length (hi)—tip of lower jaw to rear margin of opercle; snout length—tip of lower jaw to inner anterior margin of bony orbit; snout depth—least vertical dimension of snout, posteriad of gape; pectoral-fin length— length of longest fin ray from articular base to undamaged tip; length of pectoral-fin base— straight line distance between articular bases of upper and lowermost rays; length of dorsal-fin base—distance between anterior and posterior angles of insertion; anal ring depth—least distance between outer margins of superior and inferior trunk ridges at anal ring; trunk depth— maximum depth of trunk, usually measured between outer margins of superior and median ventral trunk ridges. Although some species exhibit sexual dimorphism in certain dimensions (e.g. trunk depth), proportional data are here combined for both sexes. 7. Persistent color markings in well-preserved material are often useful aids in the identification of some species (e.g. Syngnathus floridae, S. pelagicus). However, coloration alone should not be used for the identification of any western Atlantic pipefish. 8. Type of brood-pouch closure (Fig. 3) may also aid in the assignment of adult males to genera. However, this character is of limited value, since the type of closure can usually be determined only in specimens bearing a full pouch of recently deposited eggs. Synonymies and References. Synonymies are intended to be complete and have, where possible, been verified by examination of pertinent typematerial. Considerable effort has been expended to include all literature references to treated species. There is little doubt however, that some have been omitted. Hopefully, these are not sufficiently numerous to constitute a serious deficiency in this report. Distribution Maps. The maps delineate general distribution of material examined and may not show all collection sites within the immediate

INVERTED

OVERLAPPING

9

SEMI

EVERTED

FIGURE 3. Types of brood-pouch closure occurring in western Atlantic syngnathine (tail pouch) pipefishes; terminology from Herald, 1959.

vicinity of symbols. In most cases, the maps indicate the extent of known distribution but this may be supplemented with additional information under Distribution in species treatment. Keys. Keys to genera and species are designed to minimize the use of sexually dimorphic characters and preserved coloration. Proportional characters are required for some identifications but the need for measurements has been kept to a minimum. Keys should work equally for wellpreserved early juveniles and adults of most species.

Memoir I, Part 8—Sears Foundation for Marine Research

10

Key to the western Atlantic Genera1 la. Superior trunk and tail ridges confluent (Fig. 2a), not interrupted behind anal ring or near rear of dorsal fin. Penetopteryx Lunel 1881, p. 13 Ib. Superior trunk and tail ridges interrupted near rear of dorsal fin 2 2a. Inferior trunk and tail ridges interrupted near anal ring (Fig. 2c) 3 2b. Inferior trunk and tail ridges confluent 6 3a. Tail curled (prehensile); dorsal-fin base elevated. Amphelikturus Parr 1930, p. 10 3b, Tail not curled; dorsal-fin base not elevated 4 4a. Caudal-fin rays typically 9; pectoral-fin rays 17-23; dorsal-fin rays more than 30; brood pouch abdominal. Oostethus Hubbs 1929, p. 16 4b. Caudal-fin rays typically 10; pectoral-fin rays fewer than 16; dorsal-fin rays fewer than 30; brood pouch caudal 5 5a. Anal fin present; pectoral-fin rays usually (99%) 12-14. Micrognathus Duncker 1912, p. 25 5b. Anal fin absent; pectoral-fin rays usually (99%) 6-11. Anarchopterus Hubbs 1935, p. 33 6a. Lateral trunk and tail ridges confluent (Fig. 2b) 7 6b. Lateral trunk and tail ridges interrupted near anal ring (Fig. 2d) 8 7a. Anal fin present; trunk rings 17-20. Leptonotus Kaup 1853, p. 40 7b. Anal fin absent; trunk rings 13-15. Pseudophallus Herald 1940, p. 44 8a. Without supraopercular ridge, lateral snout ridge or dermal flaps; anal fin present; dorsal-fin rays 22-49, usually 28 or more (except 22-27 in S. makaxi); subdorsal trunk rings 0.25-6.5; inverted pouch closure (Fig. 3). Syngnathus Linnaeus 1758, p. 55 8b. With supraopercular ridge, lateral snout ridge or dermal flaps (all may be present); anal fin present or absent; dorsal-fin rays 19-27; subdorsal trunk rings 0-2.75; pouch closure not inverted 9 9a. Anal fin absent. Bryx Herald 1940, p. Ill 9b. Anal fin present. Cosmocampus Dawson 1979, p. 120

Genus Amphelikturus Parr 1930 Amphelikturus Parr, 1930:31 (type-species Amphelikturus brachyrhynchus Parr 1930 (—Siphostoma dendriticum Barbour 1905), by original designation). Characters. Superior trunk and tail ridges discontinuous near rear of dorsal fin, inferior trunk ridge ends at anal ring, lateral tail ridge ends below dorsal-fin base, lateral trunk ridge confluent with inferior tail ridge near anal ring (Fig. 2c). Head usually angled somewhat ventrad from principal body axis; snout short; median 1

After this report was in proof, I received a western Caribbean specimen of a species of Minyichthys Herald and Randall 1972, taken in a mid water trawl. This genus, previously unknown from the western Atlantic Ocean, is in part characterized by the Fig. 2c ridge configuration, absence of prehensile tail and presence of anal fin and an 8-rayed caudal fin. A report on this fish will be published elsewhere.

dorsal snout ridge entire or with 1-3 irregular emarginations; snout ridge originates near middle of snout length, not confluent behind with orbital ridges. Median dorsal head ridges prominent, somewhat conical; complete or nearly complete ridge curves dorsad across opercle with several rather distinct radiating striae below; pectoral-fin base protrudes strongly laterad, superior and inferior ridges present in subadults. Principal body ridges prominent; posterior angles of rings produced as points on trunk and anterior tail rings, ridges and ring angles becoming indistinct caudad; scutella well-defined on trunk, reduced or obsolete on posterior %-% of tail. Simple to abundantly branched dermal flaps on eye, remainder of head and body. Trunk rings 13-15, total rings 50-55. Dorsal-fin rays 15-18; dorsal-fin origin between penultimate trunk ring and 2nd tail ring, the fin-base

Fishes of the Western North Atlantic elevated. Pectoral-fin rays 12-15; anal fin usually with two rays. Posterior % or more of tail curled ventrad (prehensile) in preservative; caudal fin small, with 9-10 rays. Brood pouch below 10-11 tail rings; pouch plates little enlarged. Pouchclosure uncertain, arrangement of pouch eggs unknown. Discussion. Herald (1943, 1959) considerd Amphelikturus to be the genus of pipefishes (Syngnathinae) most closely related to seahorses (Hippocampinae). This opinion was derived principally from the shared configuration of principal body ridges and the prehensile tail, as well as from examination of two empty brood pouches of Amphelikturus. On the basis of present material, I cannot confirm Herald's observation of everted pouch closure nor his (1959) statement that "the pouch is closed for the two most posterior rings." Herald considered the partially fused pouch folds to be intermediate between the open (unsealed) pouch of typical pipefishes and the sealed pouch of seahorses, and that the condition in Amphelikturus was unique among the Syngnathinae. Herald was evidently unaware that the Australian syngnathine Kaupus costatus (Waite and Hale) also has pouch folds fused over the distal 2-3 rings and everted pouch closure, although differing from Amphelikturus in body ridge configuration (Fig. 2d versus c). The Amphelikturus ridge configuration is shared with several genera of doryrhamphine and syngnathine genera. Furthermore, the sealed brood pouch of the Indo-Pacific syngnathine genus Apterygocampus Weber has recently been shown (Dawson and Allen, 1978) to bridge previously accepted subfamilial lines between the pipefishes and seahorses. Further study is required, but morphological similarities of Amphelikturus and seahorses may well reflect parallelism rather than close phylogenetic relationship. Size. Small fishes not known to exceed 81 mm. Range. Western Atlantic between about 42° N and 09 °S; eastern tropical Atlantic and Indo-Pacific. Species. The genus includes one species, Amphelikturus dendriticus, in the western Atlantic. Material under study suggests the presence of

11

one species in the eastern Atlantic and two or more Indo-Pacific species. Amphelikturus dendriticus (Barbour) 1905 Seahorse pipefish Figures 4, 5, 48, 49 Siphostoma dendriticum Barbour, 1905:115, PL 1 (orig. descr.; Ireland Island, Bermuda; holotype, MCZ 29057); Bean, 1906:39 (compiled); Mellen, 1919:134, fig. (prehensile tail); Collette, 1962:440 (compiled). Syngnathus dendriticus. Jordan et al., 1930:242 (n. comb.; compiled). Amphelikturus dendriticus. Parr, 1930:32 (n. comb.; in key); Beebe and Tee-Van, 1933:79, 284, fig. (descr.; in key; prehensile-tailed pipefish; Bermuda); Herald, 1942:132 (in key; Bermuda and Crooked Island, Bahamas); Herald, 1943:41, 72, 291-295, Fig. 31 (descr.; phylogenetic position); Herald, 1959:470, Figs. 1, 2 (characters; relationships); Herald, 1961:150 (descr.); Herald, 1965:364 (compiled; seahorse pipefish); Bohlke and Chaplin, 1968:181, fig. (descr.; behavior; pipehorse; Bermuda and Bahamas); Dawson, 1972:846 (nekton, off Mississippi); Fahay, 1975:20 (plankton; 29°05'N, 80°19'W); Walls, 1975:156, fig. (descr.); Guitart, 1975:278, Fig. 212 (in key; diagn.; Cuba); Wheeler, 1975:109 (descr. notes; range, in part); Shiino, 1976:107 (compiled); Markle et al., 1980:56, Fig. 7c (surface neuston; 42°19'N, 65°30'W). Amphelikturus brachyrhynchus Parr, 1930:32, 144, Fig. 7 (orig. descr.; type-species of Amphelikturus', Crooked Island, Bahamas; holotype, BOC 2532); Herald, 1942:131 (=A. dendriticus); Fowler, 1944a:442 (compiled).

Distinctive Characters. The combination of ventrally coiled tail, together with presence of caudal fin, elevated dorsal-fin base and essentially unsealed brood pouch readily separates this species from other Atlantic syngnathids. Description. Proportional data from measurements of holotype. Rings: 13-15 + 37-41, modally 14 + 39. Subdorsal rings: 1.0-0.0 + 3.25-4.5 = 3.75-5.0, modally 0.5 + 3.75. Fin rays: dorsal 15-18 (x = 16.2), pectoral 12-15 (x = 14.3); anal 2-3; caudal 9 (in 4), 10 (in 9). Head. Head length 7.2 in SL, snout length 3.1 in hi, snout depth 3.0 in snout length. Lateral snout ridge usually a blunt protrusion; opercular ridge essentially complete in small fish (*3&&*^ ^&x..v^fj£*

sffi///fffis*$$$$&^, fmizFte^^?*'*?*** ^-JL^tt^t

|t

M^g^WA^^gj^^^y **ff^T"*g^1fl-n*mn

^**'^^

^j9

FIGURE 9. Micrognathus crinitus: holotype (BMNH 1917.17.7.14.29), female, 84.5 mm.

1942:131 (=Micrognathus vittatus); Duarte-Bello and Buesa, 1973:91 (not recorded, Cuba); Dawson, 1978a:13, Fig. 1 (sr. synon. of M. vittatus; descr. and fig. of holotype; type-locality: Brazil). Micrognathus vittatus. Duncker, 1915:77 (n. comb.; descr.; doubtful species); Hubbs, 1935:3 (lateral ridge configuration); Herald, 1942:131, 134 (synon., in part; in key); Herald, 1943:99, 275, Fig. 25 (descr.; comparisons; distr.); Bertin and Esteve, 1950:46 (listed); Kahsbauer, 1950:271 (meristic characters, Brazil); Herald, 1953:257 (color note); Springer and Bullis, 1956:67 (listed); Briggs, 1958:267 (compiled; distr., in part);

Duarte-Bello, 1959:51 (probably, Cuba); Herald, 1959:469, Fig. 2 (lateral ridge configuration); Bailey et al., 1960:23 (banded pipefish); Collette 1962:440 (compiled); Herald, 1965:372 (only Atlantic member of subgenus Micrognathus; comparisons; primarily insular); Caldwell, 1966:36 (compiled); Cervigon, 1966a:932 (descr.; "ensenadae" coloration; Los Roques, Venezuela); Bohlke and Chaplin, 1968:157, fig. (descr.; comparisons; ecol. note; distr., in part); Starck, 1968:18 (Alligator Reef, Florida); Bailey et al., 1970:33 (compiled); Mago, 1970:92 (compiled); Resendez, 1971:25 (compiled); Shiino, 1972:61 (compiled); Duarte-Bello

28

Memoir I , Part 8—Sears Foundation for Marine Research

i-H

LT>

ITS -*

LT> \O

«M in in

o

ri . • ' ; .'V

^;^>^^f^7^^H^'x^:- ^ -^.'^ /"•' .(!' --' k":^*.^ j^K'i'^vl',/,.•.-. ; :,y • , ..-

FIGURE 21. Syngnathus floridae: Top three, female (GCRL 7396) 175 mm. Bottom, delineation of characteristic markings on posterior tail rings.

ISyngnathus floridae. Gunter and Hall, 1963:225, 233 (possibly mixed with 5. louisianae). Syngnathus floridae hubbsi Herald, 1965:364, 368 (Chesapeake dusky pipefish; orig. descr.; York River, Virginia; holotype, USNM 91321); Dahlberg, 1975:53 (characters, distr.); Shiino, 1976:109 (compiled). Syngnathus floridae mckayi Herald, 1965:364, 367, 370 (Key dusky pipefish; Miami to Tortugas, Florida; leetotype selection); Dahlberg, 1975:53 (characters; distr.).

Syngnathus floridae nesiotes. Herald, 1965:364, 368 (n. comb.; Bermuda dusky pipefish; notes); Herald and Dawson, 1972:782 (Bermudan dwarf); Shiino, 1976:109 (compiled).

Distinctive Characters. The following combination of characters differentiates S. floridae from western Atlantic congeners: broad preorbital (Fig. A), moderately slender snout (depth aver-

Memoir I, Part 8—Sears Foundation for Marine Research

60

TABLE XXIII. Geographic variation in frequencies of trunk and tail rings in Syngnathus floridae. Tail rings

Trunk rings Locale Bermuda Bahamas United States Maryland Virginia N. Carolina E. Florida W. Florida Alabama Mississippi Louisiana Texas Mexico Belize Nicaragua Panama

16

3 2 8 1

17

18

19

43 4

6 2

1

2 29 31 86 83*

30 4 27 17 2

2

46

4

4 6 53 289 1 5 2 5 4 2 3

29

1 15

30

32

33

34

35

36

37

2

4

13 2

16

11 4

3 1

2 16 14 41 187

15 57 86

2 29 11 1

1 3

1 1 14

1 2

31

1

5

1

3 15

14 7 15 88*

2

18

16

2 1

17 2

9 13

1 3 1 5 2

3

38

39

1

1

1 1 1

1

10

29

* Primary type.

ages ca. 7.0 in length), low pectoral-fin base in pectoral-fin length ratio (averages 1.3), usually 17-18 trunk rings and frequent presence of characteristic tail markings (Fig. 21). Description. Proportional data based on 397 specimens from various localities, 49.5-243.0 (x = 146.5) mm. Rings: 16-19 + 29-39 = 46-58; trunk rings modally 18 in West Florida population, modally 17 elsewhere (Tables XXIII, XXVII, XL, XLI). Subdorsal rings: 3.0-0.25-h4.0-7.0 = 5.75-8.5

(Tables XXV, XXVI, XXIX, XXX, XLIV, XLV, XL VI). Fin rays: dorsal 26-35, 28-31 in 84% (Tables XXIV, XXVIII, XLII); pectoral 12-16, modally 14 (Table XLIII). Head. Head length 5.3-7.9 (x = 6.5) in SL, snout length 1.6-2.1 (1.7) in hi, snout depth 4.8-10.4 (7.2) in snout length. Preorbital area between nares and anterior margin of orbit (lateral ethmoid) is typically wide and appears externally as a broad flat bony process (Figs. A in

TABLE XXIV. Geographic variation in frequencies of dorsal-fin rays in Syngnathus floridae. Dorsal-fin rays Locale Bermuda Bahamas United States Maryland Virginia N. Carolina E. Florida W. Florida Alabama Mississippi Louisiana Texas Mexico Belize Nicaragua Panama * Primary type.

26

1 7

1

27

28

29

30

31

32

1

5 1

16

13 1

18 2

1 3

1 17 13 30 134*

1 8 15 41 71

5 9 41 24

1

17

8 6 3 1 2

1 1 50

9 2 11 105

1

1 2

8 1

2

4 1

7 1

3 9 3

12

20

17

6

19 2 1 2 1 11 2

33

34

35

3

2

1

7

2 1

Fishes of the Western North Atlantic

61

TABLE XXV. Geographic variation in frequencies of subdorsal trunk rings in Syngnathus floridae. Subdorsal trunk rings Locale Bermuda Bahamas United States Maryland Virginia N. Carolina E. Florida W. Florida Alabama Mississippi Louisiana Texas Mexico Belize Nicaragua Panama

3.00

2.75

2.50

2.25

2.00

1.75

1.50

1

1 2

3 2

7

25 1

11 1

10

2

2

3

1 1 1

5

6 1 1 5

7

1 41 7 1 2 2

25 13 2 2 4 1

9 1 2 12

9

1.25

1.00

11 6 16 63

1 7 19 16 149

4 5 1 48*

2 3 4 31

17

1

5

12 1

1

3

1 9 6 39 83 2 1 5 4 1 1 19

7 1 6 1 7

0.75

0.50

0.25

1

1

4

* Primary type.

key, 21); width varies somewhat among individuals of similar size and is proportionately narrower in young, but S. floridae typically has a broader preorbital than western Atlantic congeners. Body. Depth at anal ring 3.2-9.5 (5.1) in hi, trunk depth 2,2-6.1 (3.8) in hi. Mature females often with deep trunk (Fig. 74), particularly in Florida populations. Fins. Length of dorsal-fin base 1.0-2.0 (1.4) in hi, pectoral-fin length 5.8-10.5 (7.8) in hi, length

of pectoral-fin base 0.9-1.8 (1,3) in pectoral-fin length. Color. Ground color near white to brownish, markings mostly tan to near black. Snout usually with diffuse dusky lateral stripe, suborbital and lower third of opercle often blotched with iridescent silver, opercle usually dusky elsewhere. Markings variable on trunk and anterior half or more of tail (Figs. 73-76), usually darker on dorsum and upper half of sides, mottled, blotched, with irregular vertical rows of small

TABLE XXVI. Geographic variation in frequencies of total subdorsal rings in Syngnathus floridae. Total subdorsal rings Locale Bermuda Bahamas United States Maryland Virginia N. Carolina E. Florida W. Florida Alabama Mississippi Louisiana Texas Mexico Belize Nicaragua Panama * Primary type.

5.75

6.00

6.25

1 4

21 2 2

1

14

16

6.50

3 3 5 77 1 3 1 1 1

21

6.75

7.00

7.25

7.50

775

8.00

8.25

1

4

11 1

13 1

15 2

10 1

2 2

1

9 8 35 58

3 12 46 15

2 2 21 2

12

1

1

11

5

2

13 5

3 4 1

2 3

1

1

9 3 7 96* 6 4 3 2 1 3

1 13 13 27 121

10 1 6 7 1 1 1

1

2

8.50

Memoir 1, Part 8—Sears Foundation for Marine Research

62

TABLE XXVII. Geographic variation in frequencies of trunk, tail and total rings in Florida populations of Syngnathus floridae. Locale

19

16

17

18

8

1 1 26 58

1 14 38

1

15 36 40 222 7 21

14 1

East Florida St. Lucie Co. Martin Co. Dade Co. Florida Keys West Florida Cape Sable area Lee-Citrus Co. Taylor-Franklin Co. Escambia-Santa Rosa Co.

Total rings

Tail rings

Trunk rings 29

30

1 1

1

7 8

34

32

33

13 28

1 1 1 3 16 40 24

1 16 24 9 47 137 2 10 13

8 73 4

15* 13

1

3

1 21* 10

31

35

3

2 9

50

51

52

53

12 45

1 2 8 20

1 11

1

1

5 19 21 14 65 119 4 11 11

4 66 3

2 10

1

2

11* 15

47

48

1

2 7

49

16 21

4

* Primary type.

pale spots or with diffuse bars; venter of trunk pale to dusky. Sides and dorsum of posterior tail rings usually with characteristic pattern of irregular narrow brownish stripes and pale interspaces (Fig. 21); stripes sometimes obscured by ground color, interrupted or confined to last 3-4 rings but some evidence persists in most wellpreserved subadults and adults. Dorsal fin usually pale, occasionally with diffuse dusky bars or blotches on basal half; pectoral fins mainly pale; caudal fin brownish, often margined with pale. Longley (1916, 1917) and Behre (1933) report adaptive color change to changing substrate. Specimens from Thalassia beds or sand flats are usually green, brown-olive or green with gray

dorsum (Longley, 1917); specimens from the northern Gulf of Mexico are usually olive-green to brownish in life. Size. The largest examined specimen is 258 mm. Development and Growth. Fertilized eggs are 0.6-1.2 mm in diameter and average 0.8-0.9 mm (Brown, 1972). Embryology, originally described by Gudger (1905), has been summarized by Hardy (1978). Incubation requires about 10 days (Smith, 1907), pouch-larvae reach ca. 11.5 mm TL (Lippson and Moran, 1974) and 14.25 mm TL larvae were described by Hardy (1978). Brown (1972) found 40-50 mm juveniles during April, May, August and December at Cedar Key, Florida, and his length-frequency data indicate that

TABLE XXVIII. Geographic variation in frequencies of dorsal-fin rays in Florida populations of Syngnathus floridae. Dorsal-fin rays Locale East Florida St. Lucie Co. Martin Co. Dade Co. Florida Keys West Florida Cape Sable area Lee-Citrus Co. Taylor-Franklin Co. Escambia-Santa Rosa Co. * Primary type.

26

27

28

29

30

31

32

33

34

35

1

5 6

11 19

12 29

3 13 25

3 16

3

2

1

7 78 15 5

26 93 5 10*

12 46 1 12

6 14

1

4

1

1

6 1

1 41 7 1

Fishes of the Western North Atlantic

63

TABLE XXIX. Geographic variation in frequencies of subdorsal trunk rings in Florida populations of Syngnathus floridae. Subdorsal trunk rings Locale East Florida St. Lucie Co. Martin Co. Dade Co. Florida Keys West Florida Cape Sable area Lee-Citrus Co. Taylor-Franklin Co. Es cambia- Santa Rosa Co.

3.00

2.75

1 1

2

2.50

2.25

2 1

2.00

2 4

1.75

1.50

1.25

7 34

11 14

2 11 26

1 4 11

1 6

3 9 1

12 64 6 1

7 46 4 6

1.00

0.75

0.50

5 10

1

2 2

21 100 11 17

6 35 3 4*

2 21 4 4

1

* Primary type.

life-span is less than one year, that breeding is essentially year-round and maximum size is 200-210 mm in this population. Mature females were generally larger and outnumbered mature males by as much as 2.9:1 in Cedar Key samples. The smallest examined female with ripe ova was 121.5 mm, immature ova were common in 120-140 mm fish and most females longer than 140 mm were sexually mature. Reid (1954) found ovarian eggs to average ca. 519 in examined Cedar Key females; a 192mm fish contained ca. 1100 eggs. Brown (1972) found most males to be sexually mature at 140-150 mm and the smallest mature male in Cedar Key samples was 102 mm. Broodpouch eggs are usually in 1-2 layers and in 2-8 transverse rows. Brown (1972) showed the num-

ber of pouch eggs to generally increase with increasing SL; the average number counted in 177 males (ca. 125-180 mm) was 358, the maximum 886. Herald (1943) noted incipient pouch development at 46 mm but there is usually no indication of pouch folds or plates in fish less than 80-85 mm. Among material examined here, the smallest male with eggs in situ is 91 mm, brood pouch extends below 11-22 rings and minute papillae may be present or absent on opposing inverted margins of pouch folds. Habits and Behavior. Autumn-winter migration to deeper or offshore waters has been reported for the Chesapeake Bay population (Hildebrand and Schroeder, 1928; Schwartz, 1964 and others), and Mercer (1973) found the species present in inshore waters June-October, with

TABLE XXX. Geographic variation in frequencies of total subdorsal rings in Florida populations of Syngnathus floridae. Total subdorsal rings Locale

6.00

6.25

East Florida St. Lucie Co. Martin Co. Dade Co. Florida Keys West Florida Cape Sable area Lee-Citrus Co. Taylor-Franklin Co. Escambia-Santa Rosa Co. * Primary type.

2 1 1

18 3

6.75

7.00

7.25

7.50

7.75

8.00

8.25

5

2 5

1 1 9 16

1 11 23

1 15 30

4 17

3 9

1

6 56 8 7

14 62 6 14*

15 89 8 9

10 45 2 1

6 8 1

6.50

1 1

8.50

1

64

Memoir 1, Part 8—Sears Foundation for Marine Research

peak abundance during July-August. Seasonal variations in inshore abundance of Florida populations may also be temperature related (Springer and Woodburn, 1960; Springer and McErlean, 1962; Brown, 1972 and others). Courtship or liebspiel of mating pairs has been described by Gudger (1905), Norman (1948) and Herald (1961). Food. The food of S. floridae consists almost entirely of microcrustaceans (Reid, 1954; Brown, 1972) but larval Syngnathus sp. are occasionally ingested by subadults or adults. Brown (1972) found copepods to account for 69% of the food items in 36 young fish (51-90 mm), whereas small shrimp or mysids accounted for more than 80% of the food items in 92.5-181 mm specimens; other food items included amphipods, isopods, ostracods and larval decapods. Brown also noted evidence of seasonal variations in food items, with some increase in variety during summer and fall coupled with increased consumption of small shrimp and decreased utilization of copepods. Brook (1977) reported only daytime feeding in a South Florida population. Enemies. Aside from Strawn's (1958) note that S. floridae were eaten by the blue crab (Callinectes sapidus) in pushnet collections, there appear to be no published records of predators. It is problematical whether the crab is an effective predator on free-living pipefish. Parasites. Linton (1905) reported cestodes, trematodes and gregarines from a sample of 90 "Siphostoma fuscum" from North Carolina and it is probable that at least some Syngnathus floridae were included therein. Overstreet (1969) failed to find digenetic trematodes in six specimens examined. Sawyer et al. (1975) record infection by a leech, Myzobdella lugubris Leidy. Variation and Relationships. This species exhibits considerable meristic and color variation and there are both population and sex-linked differences in some body proportions (Herald, 1942, 1943; Brown, 1972). The original description was based on material from the northern Gulf of Mexico and North Carolina, but the Florida Keys population was described shortly thereafter as Siphostoma mckayi. Subsequently,

Herald (1942) erected a subspecies (Syngnathus mackayi nesiotes) for the Bermuda population and retained species rank for S. floridae and S. mckayi. Herald (1942, 1943) differentiated these nominal taxa mainly on the presence or absence of papillae on pouch folds, meristic differences (including number of pouch rings), relative trunk depth of mature females and maximum size attained. Herald finally (1965) considered the "floridae complex77 to include four subspecies: S. /. floridae (Gulf of Mexico: Cape Sable, Fla. to Corpus Christi, Tex.), S. /. mckayi (E Florida: Miami to Dry Tortugas), S. /. hubbsi (Chesapeake Bay to South Carolina) and S. /. nesiotes (Bermuda). Herald also suggested that Panamanian material probably represented an additional subspecies. Since Herald7s subspecies were never compared or diagnosed adequately, recent workers have generally failed to accept his proposed treatment of S. floridae. Present data delineate observed geographic variation in counts of rings, dorsal-fin rays and subdorsal rings. Where adequate material is available, trunk rings are modally 17 except in W Florida samples where rings are modally 18, and high tail-ring counts occur in Bermuda, Bahamas and Caribbean samples (Table XXIII). Dorsal-fin ray counts (Table XXIV) are most variable in E Florida samples and counts from other areas fall within this range. Counts of subdorsal trunk rings (Table XXV) indicate a somewhat more anterior dorsal-fin insertion in Bermuda, Bahamas and Caribbean samples and in some E Florida specimens, whereas counts of total subdorsal rings (Table XXVI) are lowest in Panamanian material and generally higher in E Florida, Bermuda and Bahamas samples. Herald considered the modal shift from 17 to 18 trunk rings to be one of the principal characters differentiating S. /. floridae from S. /. mckayi, but this appears to be an essentially local variation resulting from presently undetermined ecological influences. Present frequency data on rings, dorsal-fin rays and subdorsal rings of Florida samples have been tabulated north to south on the east coast and northward on the west coast from Cape Sable to the Florida-Ala-

Fishes of the Western North Atlantic bama boundary. Trunk rings (Table XXVII) are 17 in 55% and 18 in 36% of counts from the Florida Keys, whereas counts in material from Cape Sable (ca. 100-150 km distant) show reversal of frequencies to 30% with 17 and 70% with 18 trunk rings. Furthermore, 18 trunk rings occur in 70-80% of examined specimens between Cape Sable and the vicinity of Apalachicola Bay, but this count reverts to 17 in 70% of specimens examined from the Pensacola area. Compared to the Florida Keys population, W Florida material tends to have somewhat lower counts of dorsalfin rays (Table XXVIII), subdorsal trunk rings (Table XXIX) and total subdorsal rings (Table XXX). Among examined material, there are 12-17 (usually 13-15) pouch rings in Bermuda and Bahamas samples, 11-15 (12-14) in Caribbean, 17-22 (19-21) in Chesapeake Bay, 13-17 (13-15) in E Florida and 13-19 (15-18) in Gulf of Mexico samples. Pouch papillae are most common in Chesapeake Bay material but they also occur in specimens from E Florida, the Gulf of Mexico and the Caribbean. Herald (1943) noted that the percentage of deep-bodied females was variable between populations and this character is not considered diagnostic. Most variations noted here occur to greater or lesser degree in western Atlantic congeners (e.g. S. scovelli, S. fuscus), characters exhibit considerable overlap between nominal subspecies, at least two of these (S. /. mckayi and S, /. floridae) are sympatric in South Florida and there appears to be no valid basis for subspecific treatment of S. floridae. There is little doubt that gene flow between mainland and insular populations (Bermuda, Bahamas) is maintained by planktonic juveniles and apparent gaps in mainland distribution may reflect inadequate sampling. Range and Abundance. This is one of the more common pipefishes in inshore waters of Atlantic and Gulf of Mexico coasts from Chesapeake Bay (Plum Point, Md.) to the Laguna Madre at Port Isabel, Texas and, apparently smaller, populations occur at Bermuda, the Bahamas and on Caribbean mainland coasts from Mexico to Panama (Fig. 28). There are no acceptable or confirmed records from Caribbean islands and a

65

highly questionable record from the Azores (Dooley, 1972) has not been verified Furthermore, S.floridaeis presently unreported between Seabrook's Beach, S.C. (Fowler, 1945a) and Fort Pierce, Fla. and from the Mexican coast north of Quintana Roo. Although occurring in a wide range of coastal or estuarine habitats, S. floridae is most commonly collected on or near "grass beds'7 or bottoms vegetated with Agardhiella, Halodule, Thalassia, Ulva, Zostera, etc. The apparent absence of Syngnathus floridae from portions of the Atlantic seaboard and western Gulf of Mexico probably reflects the absence of extensive vegetated bottoms and inadequate sampling. Juveniles (37-54 mm) have been taken in offshore surface nekton samples over depths of 18-55 m (Dawson, 1972) and specimens have been trawled in offshore depths of 18-22 m (Springer and Bullis, 1956), but most collections are from depths of 5 m or less. Wang and Raney (1971) report a 211-mm fish from freshwater and maximum reported salinity is 40.0%o (Roessler, 1970). Most other collections apparently fall within 10-39%o and Mercer (1973) found greatest abundance at 17-22%o. Recorded temperature extremes are 10.0°C (Reid, 1954) and 32.5° C (Springer and Woodburn, 1960). Tabb and Manning (1961) found S. floridae to be the most abundant pipefish in Florida Bay and it is often recorded as "common" in the literature. Reid (1954), Brown (1972) and others have found this species to be second in abundance after S, scovelli in W Florida seine or trawl collections. Few specimens have been examined here from Alabama and Louisiana but S. floridae occurs with some frequency in these areas and it is not, as reported by Whatley (1969), rare in Mississippi waters. Resendez (1975) found this species to be common on the Caribbean coast of Mexico, Weinstein and Heck (1979) ranked S. floridae 23rd in abundance among 106 species taken in shallow-water trawls in Panama, but there are no useful abundance data for intervening Caribbean coasts. Study Material. 873 specimens, 37-258 mm SL, including lectotype and five paralectotypes.

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Memoir 1, Part 8—Sears Foundation for Marine Research

Lectotype. MCZ 35958, orig. USNM 30826 (149 mm, male), Laguna Grande, Pensacola Bay, Fla., March 1882, D. S. Jordan and S. Stearns. Paralectotypes. USNM 30826 (5, 121.5-156), taken with lectotype. Other Material. BERMUDA: AMNH 10135 (1, 106.5); AMNH 35898 (3,ca. 72-139); ANSP133435 (1,101); BOC 7789 (1, 46.5); BOC 7790 (1, 85); BOC 7791 (2, 60-73); BOC 7792 (1, 109); BOC 7793 (1, 125); CAS(SU) 36484 and 36485 (2, 108-152, paratypes of S. mackayi nesiotes); FMNH 5340, 5341, 5342, 5343, 5347, 5348, 5349 and 5351 (10, 74-156.5, paratypes of S. m. nesiotes); FMNH 48956 (5, 107-118); UMMZ 172340 (9, 75-149.5); USNM 170915, orig. NYZS 8919 (ca. 145, holotype of S. m. nesiotes); USNM 170916, 170917, 170918, 170919 and 170920 (8, 108-150, paratypes of S. m. nesiotes); USNM 178531 (8, 54.5-154); USNM 178679 (2,123-160); USNM 178784 (1, 156); USNM 178841 (6, 109-153.5); USNM 178854 (1, 98.5); USNM 217816 (1, 128). BAHAMAS: AMNH 28255 (1, 128); ANSP 94583 (1, 165); BOC 86 (1, 141.5); UMML 14139 (2, 227.5-232). UNITED STATES, Maryland: UMMZ 118119 (2, 59-80). Virginia: CAS 24295 (2, 137-177.5, paratypes of S. floridae hubbsi); MCZ 45171 (2,131-133); UMMZ 197416 (1,126); USNM 91321 (177, holotype of S. /. hubbsi); USNM 133053 (31, 44-177.5, paratypes of S. /. hubbsi); USNM 218463 (3, 92-98). North Carolina: ASM 86 (1, 110); GCRL 12232 (1, 84); GCRL 12233 (2, 69-108); GCRL 12235 (1, 100); GCRL 12237 (1, 110); GCRL 12242 (1, 56); GCRL 12243 (1, 60); GCRL 12244 (4, 71-147); GCRL 14090 (1, 105); GCRL 14824 (4, 98-104); USNM 198233 (23, 84-160). East Florida: AMNH 2800 (4, 173-209); AMNH 2823 (3, 137-161); AMNH 2829 (1, 130); ANSP 30617 (1, 92.5); ANSP 30623-24 (2, 114.5-157); ANSP 30625-29 (5, 80.5-157); ANSP 78646 (1, 183); ANSP 78650 (3, ca. 182.5-235); ANSP 78659 (4, 114.5-195); ANSP 98159 (1, 124); ANSP 98162 (3, 129-160); ANSP 134206 (4, 94-173.5); CAS 33314 (5, 130-161); CAS(SU) 1894 (138, male lectotype and 3 (169.5-210.5) paralectotypes of Siphostoma mckayi); FDNR 1659 (8, 68-179); FDNR 9020 (1, 74); GCRL 14249 (4, 110-159); GCRL 15235 (7, 119.5-172); GCRL 15240 (13, 71.5-176); GCRL 15241 (4, 147-186); GCRL 15242 (2, 142.5-171.5); GCRL 15244 (4, 139-146); GCRL 15245 (7, 101-163); GCRL 15246 (2, 113-142); GCRL 15257 (2, 127-134); HBF 1205 (1, 69.5); HBF 1207 (1, 136); HBF 1208 (1, 94); HBF 1209 (1, 143); UMML 905 (5, 54.5-174); UMML 1688 (4, 95.5-200); UMML 8173 (1, ca. 102); UMML 8186 (7, 85-137); UMML 8327 (8, 75-189); UMML 8844 (5, 121-195); UMML 13719 (4, 208-223); UMML 13953 (4, 135-190); UMML 13954 (3, 57-196.5); UMML 14100 (2, 171-195); UMML 33191 (2, 164-183.5); UMML 33192 (1, 121); USNM 34989 (2, 159-164, paralectotypes of S. mckayi); USNM 57388 (2, 163.5-191); USNM 57389 (1, 156.5); USNM 57399 (2, 120-162); USNM 217833 (3, 115-144.5).

West Florida: ASM 65 (1, 123); CAS(SU) 407 (2, 121.5-139.5); FSU 18286 (16, 76.5-177); FSU 18507 (2, 141.5-193); FSU 23194 (4, 168-200); FSU 24097 (3, 143-173); FSU 25139 (4, 124.5-135); GCRL 2761 (1, 185); GCRL 6968 (1, 126); GCRL 7053 (8,100-211); GCRL 7388 (1, 186); GCRL 7396 (11, 108-194); GCRL 7416 (1, 188); GCRL 7417 (1, 225); GCRL 7425 (1, 190); GCRL 7697 (12, 142-208); GCRL 7881 (1, 89); GCRL 8361 (1, 141); GCRL 8638 (1, 133); GCRL 8641 (3, 74-188); GCRL 8645 (3, 190-213); GCRL 8646 (1, 214); GCRL 8647 (2, 150-183); GCRL 8650 (2, 157-257); GCRL 8652 (4, 181-207); GCRL 8903 (2, 190-202); GCRL 9302 (9, 123-211); GCRL 10006 (1, 165); GCRL 10009 (1, 177); GCRL 10012 (1, 159); GRCL 10025 (2, 162-176); GCRL 10042 (2, 159-184); GCRL 10068 (1, 182); GCRL 10073 (4, 159-191); GCRL 10076 (1, 171); GCRL 10080 (4, 143-181); GCRL 10081 (1, 184); GCRL 10083 (6, 111-191); GCRL 10468 (6, 187-216); GCRL 11536 (7, 117-202); GCRL 11539 (2, 182-218); GCRL 11740 (2, 120-258); GCRL 11744 (2, 129-251); GCRL 12197 (8, 138-204); GCRL 12199 (2, 156-171); GCRL 12766 (2,115-136); GCRL 12770 (1,135); GCRL 13186 (2, 128-150); GCRL 13194 (2, 128-137); GCRL 13200 (2, 101-188); GCRL 13201 (5, 81-171); GCRL 13205 (1, 161); GCRL 13207 (1, 174); GCRL 13208 (1, 154); GCRL 13226 (1, 145); GCRL 13230 (1, 228); GCRL 13253 (1, 69); GCRL 13265 (13, 95-187); GCRL 13795 (5, ca. 95.5-208); GCRL 13807 (6, 179-216); GCRL 13815 (1, 178); GCRL 13831 (2, 145-170); GCRL 13852 (2, 142-152); GCRL 13856 (1, 204.5); GCRL 13865 (11, 125.5-202); GCRL 13867 (5, 103.5-177); GCRL 13931 (4, 144-183.5); GCRL 13934 (1, 232.5); GCRL 13936 (3, 206.5-227); GCRL 13937 (10, 153-243); GCRL 13938 (5, 159-185); GCRL 13948 (2,204-206); GCRL 13952 (1,135); GCRL 14099 (2, 143.5-144); GCRL 14908 (11, 95.5-227); GCRL 15231 (15, 113-226); GCRL 15491 (2, 169-202); GCRL 15492 (15, 86-216); MCZ 46450 (1, 165); UMML 8720 (17, 114.5-249); UMML 8925 (9, 110-205); UMML 10292 (6, 125-209); UMML 10295 (3, 158-197); UMML 31685 (1, 150); UMMZ 139182 (4, 130-197); UMMZ 153572 (5, 84-112.5); UMMZ 153617 (10, 130-168.5); UMMZ 153651 (10, 142-181); UMMZ 153870 (32, 89.5-191); UMMZ 197415 (2, 130-138); USNM 30773 (9, 90.5-181); USNM 73225 (1, 93.5); USNM 119832 (1, 136). Alabama: USNM 185813 (1, 98). Mississippi: GCRL 1817 (1, 186); GCRL 1820 (3, 94-136); GCRL 4256 (1, 54); GCRL 4265 (1, 37); GCRL 4635 (4,166-183); GCRL 14134 (27, 69-169). Louisiana: USNM 132723 (4, 71-158). Texas: GCRL 15233 (12, 156.5-215); GCRL 15243 (1, 92); GCRL 15250 (14,103.5-219); USNM 213523 (2,120-127), MEXICO, Quintana Roo: GCRL 3640 (3, 90-140); GCRL 8256 (3, 42-73); GCRL 13881 (4, 58.5-135.5); GCRL 14539 (1, 120.5); UNAM IB/CML P.475 (11, 75.5-172); USNM 192285 (1, 142.5); USNM 192286 (1, 174); USNM 192362 (1, 41.5). BELIZE: AMNH 24631 (1, 138.5); ANSP 123835 (1, 193.5); FMNH 83845 (1, 105); FMNH 83846 (1, 127);

Fishes of the Western North Atlantic

67

FIGURE 22. Syngnathus springeri: female (GCRL 3642), 271 mm. GCRL 16079 (1, 171.5); USNM 218066 (1, 102). NICARAGUA: GCRL 16129 (1, 288); UCR 367-12 (3, 61.5-157); UCR 387-28 (1,286). PANAMA: AMNH 11173 (1,133.5); FMNH 8295 (2, 159); FMNH 8296 (2, ca. 150-153); FMNH 8297 (1, ca. 182); FMNH 8298 (1, ca. 155); FMNH 8299 (3,150-175); FSU 25392 (1,172); FSU 25428 (1,155); FSU 25507 (1, 146); FSU 25542 (1, 161); FSU 25789 (1, 200); FSU 25812 (2, 170.5-200); FSU 25831 (3, 143-177); FSU 25888 (1, 129); FSU 25927 (1, 171); FSU 26020 (1, 168); FSU 26073 (1, 160); FSU 26128 (2, 117-180); FSU 26141 (2, 153-161); FSU 26203 (1, 122); FSU 26360 (1, 140); FSU 26469 (1, 176); FSU 26476 (1, 154); FSU 26551 (2,136-159); FSU 26582 (1, 177); FSU 26585 (1, 147); FSU 26587 (1, 154.5); GCRL 14542 (1, 145); USNM 79700 (1, 65); USNM 79704 (1,150); USNM 79705 (1, 168); USNM 79706 (3, 151-209); USNM 79707 (1, 167); USNM 79708 (1, 164); USNM 79709 (1, 193); USNM 133593 (2, 136-166); USNM 218464 (3, 90-156); USNM 218465 (1, 75.5). LOCALITY UNCERTAIN: AMNH 16846 (1, 156), Florida, no other data. LOCALITY QUESTIONABLE: BOC 87 (1, ca. 119), Cuba, Isle of Pines.

Syngnathus springeri Herald 1942 Bull pipefish Figures 22, 28, 77-79

comparisons); Fowler, 1945a:285 (compiled); Bohlke, 1953:62 (compiled); Anderson and Gehringer, 1958:48, 49 (ref. to predator); Briggs, 1958:267 (Springer's pipefish; distr.); Herald, 1959:473 (inverted pouch closure); Bailey et al., 1960:23 (bull pipefish); Springer and Woodburn, 1960:33, 93 (off Tampa Bay, Florida); Bullis and Thompson, 1965:36 (listed); Herald, 1965: 364, 370 (biol. notes; distr.); Moe et al., 1966:27 (listed); Bohlke and Chaplin, 1968:188, fig. (descr., coloration; Bahamas and Florida coast); Starck, 1968:19 (Alligator Reef, Florida); Dragovich, 1969:16 (ref.); Bailey et al., 1970:34 (compiled); Wang and Raney, 1971:29 (seasonal distr.; Placida Harbor and Pine Island Sound, Florida); Dawson, 1972:844-847 (off Mississippi); Herald, 1972:139 (biol. note; distr.); Miller and Jorgenson, 1973:310 (meristic characters); Herrema, 1974:155 (distr. note); Dahlberg, 1975:52, 54 (in key); Fahay, 1975:4, 20 (in part; North Carolina to Cape Canaveral, Florida); Kerr, 1975:52 (listed); Walls, 1975:162, fig. (descr.; west coast of Florida); Shiino, 1976:110 (compiled); Gilmore, 1977a:134 (listed); Schwartz and Porter, 1977:436 (records; North Carolina). Syngnathus elucens (not of Poey). Fahay, 1975:20 (misident.; 117.4 mm spec, from Sta. FF-3; 31°49.5'N, 80°21.5'W).

Syngnathus louisianae (not of Giinther 1870). Fowler, 1922a:444 (in part, only 23 ring spec, subsequently designated paratype [ANSP 15753] of S. springeri); Longley and Hildebrand, 1941:64 (in part, only 23 ring spec, noted by Herald, 1942). Syngnathus springeri Herald, 1942:125, 132 (orig. descr.; in key; Lemon Bay, Englewood, Florida; holotype, CAS(SU) 36460); Herald, 1943:96, 222 (in key; descr.;

Distinctive Characters. Syngnathus springeri is best differentiated from western Atlantic congeners by the high number of trunk rings (22-24 against 21 or fewer). Compared with the only two species which may have as many as 21 trunk rings (S. louisianae, S. fuscus), S. springeri has a higher average hi in SL ratio than S. lou-

68

Memoir 1, Part 8—Sears Foundation for Marine Research

isianae (8.4 against 7.0) and a more slender snout than S. fuscus (snout depth in snout length averages 7.8 against 5.1). Description. Proportional data based on 71 specimens from various localities, 45.0-345 ( x = 186.1) mm. Rings: 22-24 + 34-37 = 57-61, modally 23 + 36 (Tables XXXI, XL, XLI). Subdorsal rings: 4.75-3.0 + 3.5-5.0 = 7.5-9.25 (Tables XLIV, XLV, XLVI). Fin rays: dorsal 32-38, 34-37 in 88% (Tables XXXI, XLII): pectoral 12-14, modally 13 (Table XLIII). Head. Head length 7.6-9.5 (x = 8.4) in SL, snout length 1.7-2.0 (1.9) in hi, snout depth 4.2-11.8 (7.8) in snout length. Preorbital moderate to narrow (Fig. 22); opercular ridge complete or nearly so in young (ca. 55 mm), often crosses half or more of opercle in adults. Body. Depth at anal ring 3.1-7.9 (5.0) in hi, trunk depth 2.2-6.4 (5.3) in hi. Median ventral trunk ridge often prominent in subadults and adults; trunk not exceptionally deep in mature females; principal ridges with prominent denticulations or minute spines in planktonic juveniles (ca. 45-100 mm), ridges essentially smooth to minutely denticulate in subadults and adults. Fins. Length of dorsal-fin base 0.9-1.3 (1.1) in hi, pectoral-fin length 4.2-7.9 (5.3) in hi, length of pectoral-fin base 1.3-2.6 (1.9) in pectoral-fin length. Color. Preserved coloration variably plain to heavily pigmented (Figs. 77-79), ground color pale to brown, markings tan to dark brown. Subadults and adults often with brownish lateral stripe on snout; dorsum of snout, dorsum and upper half of sides of head blotched or shaded with brown; remainder of head mainly pale; dorsum and upper half of sides of trunk and tail brownish with indications of narrow pale bars; venter and lower part of sides usually plain, dusky or variably blotched with brown but some adult females retain indications of dark-margined pale bars on lower half of each trunk ring; dorsal fin of mature males plain or with traces of 10-11 faint diagonal brownish

bars; adult females typically with prominent dusky to near-black blotch on anterior 4-6 dorsal-fin rays and membranes, the fin elsewhere hyaline or with faint bars; pectoral fins hyaline; caudal fin mainly brown, usually margined with pale. Planktonic young (ca. 60-100 mm) usually with ca. 13-15 brown bands (4-5 rings wide), separated by 0.5-1.0 ring diffuse pale interspaces, crossing sides and dorsum of trunk and tail; venter of trunk and tail speckled with brown; caudal fin brown with broad pale margin above and below; other fins hyaline. One juvenile (Fig. 78) with persistent narrow brown lateral stripe on trunk and anterior third of tail; one specimen examined was generally brown throughout, whereas many others lack persistent markings. Size. Herald (1965) reported a maximum length of 355 mm but this is evidently a misprint as the specimen on which this record was based (CAS 39807) is now 345 mm. Development. Among material examined, there are 17 brooding males taken in all months except February, August, September and November; 13 were taken in the December-April period. Brood pouch is developed below 16-19 rings in 25 males (208-343 mm) and it appears that males mature at about 200 mm. Pouch eggs are in 1-2 layers and in up to 10 transverse rows; Herald (1965) counted 1390 pouch eggs in a 274 mm fish. Enemies. Anderson and Gehringer (1958) report S. springeri as a food item of tuna, Euthynnus alletteratus. Parasites. Two argulid copepods from the gill chamber of a 164-mm specimen (CAS 24878) have been identified as Argulus flavescens Wilson by Dr. Roger Cressey (USNM). Variation and Relationships. Based on material examined, S. springeri has a modal count of 23 trunk rings throughout the geographic range. There is no clear indication of geographic variation in frequencies of total ring counts but the data (Table XXXI) suggest a tendency toward reduced counts of dorsal-fin rays in Florida and Mississippi material. Among western Atlantic congeners, only S. fuscus shares the modal count

Fishes of the Western North Atlantic

69

TABLE XXXI. Geographic variation in frequencies of total rings and dorsal-fin rays in Syngnathus springeri. Dorsal-fin rays

Total rings Locale Bahamas United States N. Carolina S. Carolina Georgia E. Florida1 W. Florida Mississippi

57

58

1

2

1 2

1

2

12

7* 3

59

60

61

32

33

34

35

36

37

1 9 3 27 13 4

7 4 5 9 1 2

2

1 2

2

1 1* 2

8 4 2

2 2 3 10 6 3

4 7 13 6 2

38 1

5 2 4 10 2 2

4 1 2

1

Including Fla. Keys and Tortugas. * Holotype.

of 13 pectoral-fin rays with S. springeri (Table XLIII). Frequencies of pectoral-fin rays are more variable in S. fuscus where only 43% of the counts are 13; this value occurs in 74.5% of counts in S. springeri. Range and Abundance. Syngnathus springeri is known from the western Bahamas (ca. 23°27'N, 76°38'-78°W), from the offings of Massachusetts (41°06'N, 66°31'W) south to the Dry Tortugas, and westward to about 30°13'N, SS^SW in the northern Gulf of Mexico (Fig. 28). Herald (1965) noted that S. springeri appears to be most common in depths of 18.3-128 m and that it may be found as far as 185 km offshore. This species evidently occurs throughout the water column and has been taken at the surface, in midwater trawls, in bottom trawls and dredges; individual samples include a maximum of 4 specimens. Juveniles (45-117 mm) are found in surface plankton or nekton samples (Dawson, 1972; Fahay, 1975), and specimens as large as 120 mm are recorded from floating Sargassum. Other young fish (51-144 mm) and at least two adults (268-288 mm) are recorded from surface nightlight and dipnet collections. The majority of adults were taken in bottom trawls or scallop dredges over a depth range of 11-91.4 m. The 128 m depth (Herald, 1965) has not been confirmed, but the specimen was taken in a midwater trawl and this may actually be the total depth rather than capture depth. Although most commonly collected in offshore waters, S. springeri is occasionally taken in protected shallow

inshore habitats along the W Florida coast. The holotype was collected in Lemon Bay, two specimens (121 mm) were trawled in Johnson Bay (Collier Co.), Wang and Raney (1971) reported 123-132 mm fish from Placida Harbor and Pine Island Sound, and there are four specimens (142-195 mm) in the UMMZ collections from Boca Ciega Bay near St. Petersburg. There are few useful data on substrate preference but sand and shell bottom was noted for a number of dredge samples; Starck (1968) reported the species as "rare" in a coral reef study. Recorded salinity range is 19.9-37.6%o (Dawson, 1972; Fahay, 1975); temperature range is 11.6-27.5°C (Fahay, 1975 [data for Sta. FFS, D-68-1]; Wang and Raney, 1971). Study Material. 128 specimens, 45-345 mm SL, including holotype and three paratypes. Holotype. CAS(SU) 36460 (125.5 mm, juvenile), Lemon Bay, Englewood, Fla.; Apr. 1939, S. Springer. Paratypes. ANSP 15753 (ca. 266 mm, female), Pensacola, Fla., 1942. USNM 25628 (88 mm, juvenile), Charleston, S.C., 9 June 1880. USNM 100453 (75.5 mm, juvenile), Boca Grande Light, Fla., FISH HAWK Sta. 7796, 2 Jan. 1913. Other Material. BAHAMAS: ANSP 94586 (1, 225); CAS 24736 (4, 51.5-62.5); UMML 1479 (1, 144); UMML 2759 (1,136). UNITED STATES, Massachusetts: HMBL H7566 (1, 101.5). North Carolina: ANSP 121862 (1, 157.5); ANSP 122686 (1, 186.5); CAS 39652 (1, 97); CAS 39667 (1, 267); CAS 39671 (1, 286); CAS 39672 (1, 325); CAS 39803 (1, ca. 226); CAS 39804 (1, 321); FSM 23893 (2, 306-323); FSM 23895 (1, 296); FSM 23897 (1, 222.5); GCRL 14347 (1, 66); GMBL uncat. (1,310); USNM 214276 (3, 281-296). South Carolina: FMNH 83837 (2, 273-339);

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Memoir 1, Part 8—Sears Foundation for Marine Research

FSM 23898 (1, 86); GCRL 14346 (1, 45). Georgia: CAS 24758 (2, 282-333); CAS 39816 (1, 278); FSM 23091 (1, 118); FSM 23900 (1, 75); FSM 23902 (1, 219); GCRL 14356 (1, 56.5); GCRL 14357 (1, 61.5); SHML 196 (1, 113); SHML 251 (2,75-77.5). East Florida: CAS 39521 (1, 342); CAS 39668 (1, 280); CAS 39665 (4, 255-305); CAS 39666 (1, 378); CAS 39669 (1, 323); CAS 39670 (1, 286); CAS 39673 (1, 345); CAS 39805 (1, 315); CAS 39806 (1, 333); CAS 39807 (1, 345); CAS 39808 (1, 320); CAS 39809 (2, 192-225); CAS 39813 (1, 322); CAS 39814 (1, 302); CAS 39815 (1, 315); FMNH 70547 (1, 288); FSM 2543 (2, 246-282); FSM 11777 (2, 215-239); FSM 23891 (1, 290); FSM 23892 (1, ca. 160); FSM 23894 (1, 220); FSM 23896 (3, 71.5-73); GCRL 3642 (1, 271); GCRL 13553 (1, 290); GCRL 14358 (1, 60.5); GCRL 14359 (2, 62.5-66.5); GCRL 14360 (1, 69); GCRL 14361 (1, 57); HBF 1210 (1, 232); HBF 1211 (1, 268.5); HBF 1212 (1, 243); MCZ 26251 (1, 92.5); UMML 1812 (1, 284); UMML 4133 (1, 308); UMML 8211 (1, 309); UMML 8378 (2, 208-268); UMML 15579 (1, 237); UMML 30492 (1, 227); UMML 31636 (1, 300); UMML 33203 (1, 119.5); USNM 132741 (1, 269). West Florida: ANSP 112286 (1, 174.5); CAS 24877 (1, 148); CAS 24878 (1, 164); CAS 39656 (1, 96.5); CAS 39810 (1, 268); CAS 39811 (1, ca. 191); CAS 39812 (1, 298); FSM 23899 (1, 228); GCRL 12451 (2,121); GCRL 13850 (1,135); GCRL 15720 (1, 284); UMML 16375 (1, 251.5); UMMZ 153435 (1, 143); UMMZ 154083 (1, 195); UMMZ 154893 (2, 142-176.5); USA 2914 (1, 122.5); USA 3165 (1, 280); USNM 73227 (1, 81); USNM 155534 (1,322). Mississippi: CAS 24749 (1, 86); CAS 39817 (1, 239); GCRL 4257 (1, 100); GCRL 4258 (1, 104); GCRL 4259 (1, 87); GCRL 4260 (1, 61); GCRL 4262 (1, 82); GCRL 4264 (2, 69-71); GCRL 4266 (1, 66); GCRL 4392 (1, 67). LOCALITY UNCERTAIN: AMNH 28629 (1, 280), no data.

Syngnathus louisianae Gunther 1870 Chain pipefish Figures 23, 28, 80-82 Syngnathus pelagicus, var. Linnaeus, 1766:416 (Charleston, South Carolina). Syngnathus louisianae Gunther, 1870:160 (orig. descr.; New Orleans, Louisiana; purchased; holotype, BMNH 1852.8.16.25); Fowler, 1906a:94, Fig. 10 (color note; Florida Keys); Weymouth, 1910:137 (Cameron or Chandeleur Islands, Louisiana); Fowler, 1915c:248 (Palm Beach, Florida); Nichols, 1920b:63 (Bermuda); Fowler, 1922a:444 (in part; Florida); Hildebrand and Schroeder, 1928:182, 184 (in key; Chesapeake Bay); Breder, 1929:102, 103 (in key); Truitt et al., 1929:59 (St. Mary's County, Maryland); Burkenroad, 1931:21 (sound production); Beebe and Tee-Van, 1933:81, 284, fig. (characters, in part; in key); Behre, 1933:51 (coloration); Gunter, 1935:39 (Barataria Bay, Louisiana);

Fowler, 1940a:ll (Boca Grande, Florida); Fowler, 1941c:84 (Sanibel Island, Florida); Longley and Hildebrand, 1941:64 (in part; Dry Tortugas, Florida); Herald, 1942: 125, 126, 129, errata sheet (compared with S. springeri; in key; range); Herald 1943:12, 72, 97, 213 (in key; descr.; comparisons); Fowler, 1945a:182, 285, 374 (synon., in part; range); Gunter, 1945:48 (Aransas Bay, Texas); Reid, 1954:25 (Cedar Key, Florida); Hildebrand, 1954:297 (Sabine Bank, Texas; Obregon, Mexico); Hildebrand 1955:205 (Punta Morros, Mexico); Kilby, 1955:228-234 (Cedar Key, Florida); McLane, 1955:143, 144 (sound production; St. Johns River, Florida); Reid, 1955:436 (East Bay, Texas); Springer and Bullis, 1956:68 (listed); Joseph and Yerger, 1956:129 (Alligator Harbor, Florida); Boschung, 1957:216, 219 (in key; descr.; Mobile Bay, Alabama); Joseph, 1957:76 (feeding behavior); Simmons, 1957: 183, 191 (Upper Laguna Madre, Texas); Briggs, 1958:257 (compiled); Hoese, 1958:328 (compiled); Duarte-Bello, 1959:52 (probably, Cuba); Herald, 1959:473 (inverted pouch closure); Bailey et al., 1960:23 (chain pipefish); Phillips and Springer, 1960:27 (Caloosahatchee estuary, Florida); Springer and Woodburn, 1960:32, 93 (seasonal distr.; ecol.; Tampa Bay, Florida); Richmond, 1962:93 (Horn Island, Mississippi); Springer and McErlean, 1962:54 (seasonal distr.; Lower Matecumbe Key, Florida); Powell and Strawn, 1963:115 (West Florida); Bullis and Thompson, 1965:36 (listed); Christensen, 1965:84 (ecol.; Jupiter, Florida); Gunter and Hall, 1965:27, 49 (ecol.; Caloosahatchee estuary, Florida); Herald, 1965:364, 371 (compared with S. springeri); Parker, 1965:210 (Galveston Bay, Texas); Breder and Rosen, 1966:296 (ref.); Caldwell, 1966:36 (listed; Jamaica); Moe et al., 1966:26 (listed); Norden, 1966:127 (Vermillion Bay, Louisiana); Starck, 1968:19 (Alligator Reef, Florida); Jared and Wallace, 1968:328 (chromatography of yolk protein); Tagatz, 1968:38 (ecol.; St. Johns River, Florida); Overstreet, 1969:172 (ref.); Whatley, 1969:459, 462 (Mississippi Sound); Bailey et al., 1970:34 (compiled); Dahlberg and Odum, 1970:386 (Georgia); Resendez, 1970:106, Fig. 21 (descr.: ecol.; 3.9 m/1 O2; Laguna de Tamiahua, Veracruz, Mexico); Roessler, 1970:863, 884 (ecol.; Everglades National Park, Florida); Swingle, 1971:31, Tabs. 21, 24,25, 27, 29 (seasonal distr.; Alabama estuaries); Wang and Raney, 1971:29, Tab. 23 (seasonal distr.; Charlotte Harbor, Florida); Brown, 1972:1 (biol.; food; breeding, etc.; Cedar Key, Florida); Dahlberg, 1972:844 (estuarine; Georgia); Dawson, 1972:844 (nekton; off Mississippi); Herald and Dawson, 1972:783 (ref.); Mountain, 1972:6 (Crystal River, Florida); Parker et al., 1972:70 (in key); Powell et al., 1972:60 (listed); Christmas and Waller, 1973:349, 396 (listed); Duarte-Bello and Buesa, 1973:91 (not recorded, Cuba); Miller and Jorgenson, 1973:310

Fishes of the Western North Atlantic

71

FIGURE 23. Syngnathus louisianae: juvenile (GCRL 14136), 151.5 mm. (meristic characters); Sabins, 1973:52 (Caminada Pass, Louisiana); Herrema, 1974:54 (Palm Beach and Broward Counties, Florida); Swingle and Bland, 1974:34, Tabs. 4, 5, 8, 18-20 (uncommon in tidal rivers; Alabama); Dahlberg, 1975:53, Fig. Ill (characters; common in Georgia estuaries; rare in oligohaline creek); Jones et al., 1975:74 (listed); Kerr, 1975:52 (in part; Indian River, Florida); Walls, 1975:160, fig. (descr.; uncommon; usually in deeper water; Gulf of Mexico); Hastings and Bortone, 1976:124 (off Destin, Florida); Livingston, 1976:382, 383 (Apalachicola, Florida); Shiino, 1976:110 (compiled); Gilmore, 1977a:133 (listed); Hoese and Moore, 1977:161, 162 (in key; most common pipefish; rare offshore; Gulf of Mexico); Livingston et al., 1977:90 (listed); Hardy, 1978:407, Fig. 244 (characters, ecoL, deveL); Weinstein and Heck, 1979:105 (Cape Romano and Marco Island, Florida). Siphonostoma fuscum (not of Storer). Jordan and Gilbert, 1879:368 (in part, misident.; spec, referred to S. louisianae and S. floridae by Jordon and Gilbert, 1882; Beaufort Harbor, North Carolina). ^Syngnathus fuscus. Goode, 1879:110 (listed, St. Johns River, Florida; questionable ident.; may refer to louisianae, scovelli or floridae). Siphonostoma louisianae. Jordan, 1880:22 [n. comb.; compared with S. fuscum and S. sp. (=scavelli)]. Siphostoma louisianae. Jordan and Gilbert, 1882a:264 (n. comb.; refer in part to Beaufort Harbor spec, reported as S. fuscum by Jordan and Gilbert, 1879); Jordan and Gilbert, 1882b:383 (descr.; distr.); Swain, 1882:308,313 (in key; descr.; Atlantic coast of United States); Jordan, 1884:114 (common; Key West, Florida); Swain and Meek, 1884:238, 239 (in key; Key West, Florida); Jordan, 1886b:30 (compiled); Jordan, 1887b:850 (compiled); Bean, 1891:84 (Cape Charles City, Virginia and St. Jerome's, Maryland; rare, Chesapeake Bay); Henshall, 1891:375 (Egmont Key, Florida); Evermann and Kendall, 1894:92, 109 (Corpus Christi, Galveston and

Dickinson Bayou, Texas); Jordan and Evermann, 1896a:762, 770 (in key; descr.; range); Jordan and Evermann, 1896b:327 (compiled); Evermann and Bean, 1897:242 (Titusville and Cocoa, Florida); Smith and Kendall, 1898:176 (Kings Creek, Cape Charles City, Virginia); Evermann and Kendall, 1899:63 (Florida rec, in part); Smith, 1907:171 (diagn.; biol. notes; range coincides with S. floridae). ^Syngnathus louisianae. Jordan and Gilbert, 1883:588 (inadequate descr.; probably mixed sample including S. fuscus and S.floridae;abundant; Charleston, South Carolina); Gunter and Hall, 1963:225, 230, 233, 252 (evidently mixed sample with S. floridae). ?Siphostoma fuscum. Linton, 1905:359, (records parasites in 90 pipefish from Beaufort, North Carolina; sample probably included mostly Syngnathus floridae, S. louisianae and some S. fuscus). Syrictes louisianae. Jordon et al., 1930:242 (n. comb.; range); Gunter, 1935:39 (Barataria Bay, Louisiana). Oostethus lineatus (not of Kaup). Powell et al., 1972:59 (misident.; Gulf of Mexico). Syngnathus springeri (not of Herald). Fahay, 1975:20 (misident.; 61 mm spec, from Sta. JJ-3, 29°43'20"N, 80°02'W). Syngnathus elucens (not of Poey). Kerr, 1975:52 (misident.; Indian River, Florida).

Distinctive Characters. The presence of 19-21 trunk rings differentiates S. louisianae from all western Atlantic congeners except S. floridae, S. fuscus and S. affinis, and from these it differs in the more slender snout (snout depth in snout length averages ca. 10 against 7 or less). This species usually has more dorsal-fin rays than S. floridae (33 or more in 96% against 32 or fewer in 98%) and it lacks the broad preorbital of S. floridae. Furthermore, S. louisianae has a longer

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Memoir I, Part 8—Sears Foundation for Marine Research

snout (snout length in hi averages 1.7 against 2.1 or more) and generally fewer subdorsal trunk rings (3.25 or fewer in 94% against 3.5 or more in 97-100%) than either S. fuscus or S. affinis. Description. Proportional data based on 188 specimens from various localities, 52.5-344.0 ( x = 151.2) mm. Rings: 19-21+33-38 = 52-59, usually 20 + 35-36 (Tables XXXII, XL, XLI). Subdorsal rings: 4.0-1.5 + 375-6.0 = 6.5-9.0, modally 3.0 + 5.0 (Tables XLIV, XLV, XLVI). Fin rays: dorsal 29-42 (Tables XXXIII, XLII); pectoral 12-16, modally 14 (Table XLIII). Head. Head length 5.7-8.7 (x = 7.0) in SL, snout length 1.5-1.9 (1.7) in hi, snout depth 6.9-13.2 (10.2) in snout length. Preorbital moderate to narrow; snout long and slender; opercular ridge complete in planktonic juveniles (to ca. 60-70 mm), usually reduced to anterior third of opercle in subadults and adults (Fig. 23). Body. Depth at anal ring 4.2-9.1 (6.3) in hi, trunk depth 3.4-9.1 (5.6) in hi. Mature females without enlarged or deepened trunk. Fins. Length of dorsal-fin base 1.0-1,7 (1.3) in hi, pectoral-fin length 3,4-8.2 (6.5) in hi, length of pectoral-fin base 1.2-2.3 (1.7) in pectoral-fin length. Color. Ground color usually near white to light tan; markings tan to brown, variable (Figs. 80-82), infrequently dark brown throughout. Most inshore or estuarine specimens (ca. 65-200 mm) with diffuse brownish lateral stripe on posterior half of snout and postorbital area above opercle, snout elsewhere hyaline or shaded lightly with brown; opercle and dorsum of head brownish; trunk with upper half of sides tan to brown, the dorsum often somewhat lighter; usually with a moderate to narrow brown stripe on or just above the lateral ridge and often with narrow brown bars extending dorsad from stripe on margin of each ring; some fish with lateral and dorsal scutella margined with brown (hence the name "chain pipefish"), young (ca. 70-80 mm) infrequently with narrow brown stripes on dorsum; markings on sides and dorsum of tail may replicate those on trunk, or tail

mainly brownish, mottled or blotched with pale; venter of trunk usually near white to tan, venter of tail shading to brown caudad; dorsal fin hyaline in small specimens, often with 7-10 dusky bars in larger fish; pectoral fins usually hyaline; caudal fin brownish. Large fish may have silvery iridescence on opercle; dorsum and upper half of sides mainly brown; lateral scutella of tail with rear margin edged broadly with brown; venter of tail largely plain but often with indications of dusky bars posteriad; pectoral fins shaded with microchromatophores; occasional specimens with body barred or banded throughout (Fig. 82). Planktonic juveniles (ca. 50-75 mm) shaded with brown on snout and dorsum of head; body with ca. 14-15 brown bands (2-4 rings wide) and 1-2 ring pale interspaces, bands usually confined to sides and dorsum of trunk but completely encircle tail; dorsal and pectoral fins hyaline; caudal fin brownish. Brown (1972) noted life coloration as pale brown. Some color change in response to variations in artificial illumination was reported by Behre (1933). Size. Brown (1972) reported maximum length as 355 mm and Walls (1975) stated that S. louisianae reaches a total length of 381 mm (15 inches). Development and Growth. Brooding males have been reported for all months except March, May, October and November (Smith, 1907; Hildebrand, 1954, 1955; Joseph and Yerger, 1956; Brown, 1972) and ripe females occur during April, July, August and September in Florida (Brown, 1972). Planktonic young (28-100 mm) occurred off Mississippi during all months except February and December, with maximum abundance during April-July (Dawson, 1972). Other young fish (ca. 45-115 mm) have been taken in all months except December, and it appears likely that breeding is essentially yearround in southern habitats. Brown (1972) suggested that sexual maturity is reached within the 150-200 mm size range. The smallest recorded male with developing pouch is apparently 116 mm (Springer and

Fishes of the Western North Atlantic Woodburn, 1960), and Joseph and Yerger (1956) noted brooding males within the 125-185 mm size range. Mature males are evidently uncommon in inshore collections and there are few useful data on their occurrence. Among material examined, the smallest specimen with developing pouch is 145.5 mm; 25 brooding males (155.5-344 mm) have pouch extending below 15-20 rings; an occasional specimen was found to have papillae on pouch folds as described for S. floridae. Brood-pouch eggs of S. louisianae are 0.6-1.4 mm in diameter (Herald, 1943; Brown, 1972), arranged in 1-2 layers and in 2-10 transverse rows (Herald, 1943). Brown (1972) found number of pouch eggs to range from 454 to 898 in six fish (202-294.5 mm). In addition to the complete opercular ridge, planktonic young (ca. 20-70 mm) may have elevated median dorsal head ridges and laterally protruding spine-like projections on posterior end of principal ridges of all rings. Best developed in smallest specimens, these features are reduced and eventually lost in larger fish. Estuarine young (ca. 50-70 mm) retain little or no trace of these common modifications of planktonic specimens. Habits and Behavior. Burkenroad (1931) described a clicking sound produced by rapid head movement in this species and Yerger (1957) reported a similar sound accompanying feeding in aquarium specimens. McLane (1955) also noted high frequency vibrations in freshly caught material. Food. Small shrimp, mysids and amphipods accounted for the bulk of food items in 18 examined S. louisianae, but small quantities of copepods, isopods and ostracods were also included (Brown, 1972). Enemies. There appear to be no confirmed records of predators. Fowler (1945a) notes that a specimen was regurgitated by a Royal Tern (Thalasseus maximus) but this could have been taken while feeding on trash fish. Parasites. Pipefishes reported as Siphostoma fuscum by Linton (1905) probably included some Syngnathus louisianae but reported parasites cannot now be definitely related to this species. Di-

73

TABLE XXXII. Geographic variation in frequencies of trunk and tail rings in Syngnathus louisianae. Trunk rings Locale United States New Jersey Virginia N. Carolina S. Carolina Georgia E. Florida W. Florida Alabama Mississippi Louisiana Texas Mexico

19

3 1 7 16 16 1 49 1 8 3

Tail rings

20 21 33 34 35 36 37 38 1 4 53 4 21 48 34 8 102 2* 8 1

2 8 1 1 3

1

11 1 10 2 10 3 3

1 3 2 9 35 22 4 61 1 10 1

1 2 2 1 11 38 12 2 13 5 2 1 1 17 15 1 1 2 1 70 13 2* 3

* Primary type.

genetic trematodes were not found in 3 specimens examined by Overstreet (1969). Of four S. louisianae from Horn Island, Mississippi, one had an encysted cystacanth of an acanthocephalan (Paleoacanthocephala) and about 25 plerocercoids of a tetraphyllidean cestode in the intestine (R. M. Overstreet, pers. comm.). Variation. Brown (1972) found some differences in body proportions and counts in compared samples of S. louisianae from the Miami area and Cedar Key (West Florida) and material examined here exhibits clinal variation in several meristic features. Nineteen trunk rings occur in only 15% of Atlantic seaboard material (reach 24% in East Florida), whereas this count occurs in 32% of Gulf of Mexico specimens and reaches or approaches 50% in Texas and Mississippi samples (Table XXXII). Similarly, 7% of Atlantic coast fish have 21 trunk rings but this count occurred in only 3 (1.3%) of 234 Gulf of Mexico specimens. Tail rings are modally 37 in North Carolina, usually 35-36 in East Florida and the Gulf of Mexico and a north-south reduction occurs in counts of dorsal-fin rays (Table XXXIII). Similar variation is found in counts of subdorsal rings and dorsal-fin origin is usually somewhat more advanced in Atlantic coast specimens (78% on 2.75 or more trunk rings against 73% on 2.75 or less).

Memoir I , Part 8—Sears Foundation for Marine Research

74

TABLE XXXIII. Geographic variation in frequencies of dorsal-fin rays in Syngnathus louisianae. Dorsal-fin rays Locale United States New Jersey Virginia N. Carolina S. Carolina Georgia E. Florida W. Florida Alabama Mississippi Louisiana Texas Mexico

29

30

31

32

33

34

1

1

1

1

1 5 7 1

2 9 2 2 3 1

6 10 15 1 36 1 6 1

35

3 2 6 15 12 2 42 1* 5 3

36

1 3 3 12 24 5 4 20 3

37 1 2 9

38

39

40 1

19

1 1 16

10 2

2

5

1

7

41

42

3

1

3

1

* Primary type.

Distribution and Abundance. Syngnathus louisianae is known from the Forked River, Barnegat Bay, New Jersey to the Dry Tortugas, westward along Gulf of Mexico shores to Port Izabel, Texas and from the Mexican states of Veracruz (Resendez, 1970), Tabasco and Campeche (Fig. 28). The New Jersey specimens were recently collected from cooling water screens at a power plant. Extralimital records for Bermuda (Nichols, 1920a) and Jamaica (Caldwell, 1966) are highly questionable. The former is evidently based on a single adult (AMNH 9059) received from L. L. Mowbray in a collection of Bermuda fishes. Separate collection data were not provided for this pipefish and it is known that Mowbray collected in other areas and received material from workers outside of Bermuda. In the absence of subsequent collections from Bermuda, it appears likely that this specimen originated elsewhere and was inadvertently mixed with the shipment of Bermuda fishes. The sole Jamaican record is based on a specimen (orig. USNM 6922, now USNM 117400) labeled "Jamaica—C. B. Adams/' The original catalog entry for USNM 6922 shows that "Syngnathus" has been crossed out and replaced by "Harengula" in different handwriting. There is no apparent reason for recataloging and the entry for USNM 117400 repeats the original data on locale and collector. Absence of subsequent collections of Syngnathus louisianae from

Jamaica and evident cataloging problems cast considerable doubt on the occurrence of this species at Jamaica. The majority of collections are from inshore or estuarine habitats at depths of 10 m or less. Herald (1965) indicated a maximum depth of 128 m but, as noted under S. springeri, this was from a midwater trawl sample and actual collection depth may well have been less. Among material examined, maximum recorded depth is from a tumbler dredge sample in 38.4 m (SILVER BAY Sta. 3621). Juveniles may occur well offshore in surface nekton or plankton collections (Dawson, 1972; Fahay, 1975) and one (MCZ 43241) was taken with nightlight and dipnet over a depth of 1463 m. Although an associate of floating Sargassum (Herald, 1943; Hastings and Bortone, 1976), Syngnathus louisianae is most commonly found among or near such aquatic vegetation as eelgrass, Halodule or Thalassia (Smith, 1907; Herald, 1943; Hildebrand, 1954). Juvenile S. louisianae may be the most abundant pipefishes in offshore Gulf of Mexico surface collections (Dawson, 1972), but only one specimen occurred in similar samples off the Atlantic seaboard (Fahay, 1975; see synonymy). Present data and the literature (Springer and Woodburn, 1960; Powell and Strawn, 1963; Brown, 1972, etc.) show that adults are uncom-

Fishes of the Western North Atlantic mon in shallow inshore habitats. There is less agreement in reports of species abundance and S. louisianae is variously recorded as rare, uncommon, abundant or most common. The species is reported infrequently from Chesapeake Bay, and its apparent absence from the Mexican coast between northern Verzcruz and Tabasco probably reflects reduction of vegetated bottom areas and inadequate sampling. In Georgia estuaries, this species has been described both as uncommon (Dahlberg and Odum, 1970; Dahlberg, 1972) and common (Dahlberg, 1975). Florida data of Reid (1954), Springer and Woodburn (1960) and Wang and Raney (1971) show S. louisianae to be less abundant in inshore samples than either S. floridae or S. scovelli, and it accounted for roughly 11% of the total annual catch of the three species. Although rare in Mississippi Sound during summers of 1964 and 1965 (Whatley, 1969), a single collection during July 1975 produced 63 S. louisianae, 27 S. floridae and 17 S. scovelli. Syngnathus louisianae is recorded from fresh water (Tagatz, 1968) to a maximum salinity of 45%o (Simmons, 1957) but most collections appear to be from salinities of 13-37%o. Reported temperature extremes are apparently 10°C (Reid, 1954) and 34.9°C (Christmas and Waller, 1973). Study Material. 426 specimens, 19-344 mm SL, including holotype. Holotype. BMNH 1852.8.16.25 (200 mm, female), New Orleans, purchased. Other Material. UNITED STATES, New Jersey: GCRL 16089 (1, 301); GCRL 16255 (1, 251); GCRL 16256 (1,299). Virginia: USNM 43206 (2, 276-295); USNM 72316 (2, 176-181.5). North Carolina: AMNH 3036 (12, 82.5-155); AMNH 3301 (3, 280-323); AMNH 4372 (18, 58-327); BOC 5204 (12, 58.5-125); FMNH 50344 (1, 232); GCRL 12231 (1,245); GCRL 12238 (1,100); GCRL 12240 (1,110); GCRL 12241 (1, 107); GCRL 14091 (1, 96); GCRL 14831 (3, 255-344); GCRL 14833 (3, 252-306); USC uncat. (1, 53); USNM 117454 (1, 291); USNM 117455 (1, 273); USNM 117456 (1, 263). South Carolina: BOC 7794 (1, 205); FMNH 38261 (1, 152); GMBL uncat. (2, 47-69); GMBL uncat. (1,157); LS (1, ca. 230). Georgia: FSM 23884 (1, 295); GCRL 13788 (1, 221); GCRL 13789 (1, 153); GCRL 13796 (1, 165.5); GCRL 13798 (1, 159.5); GCRL 13799 (1, 109); GCRL 13818 (2, 79.5); GCRL 13820 (1, 126.5); GCRL 13836 (1, 75.5); GCRL 13845 (1, 94.5);

75

GCRL 13846 (1, 165); GCRL 13893 (2, 98-146.5); GCRL 13895 (2, 63-127.5); GCRL 13897 (1, 208); GCRL 13908 (1, 73); GCRL 13910 (1, 71.5); GCRL 13913 (1, 81.5); GCRL 13915 (1, 67.5); GCRL 13917 (1, 114.5); GCRL 14082 (1, 80); GCRL 14125 (1, 67.5); GCRL 14127 (1, 77); GCRL 14130 (1, 93.5); GCRL 14133 (1, 80); MCZ 45364 (1, 261); USNM 213527 (1, 92). East Florida: ANSP 30618 (1, 253); ANSP 30620-22 (3, 155.5-222); ANSP 98154 (3, 93.5-135); GCRL 13791 (1, 157.5); GCRL 13886 (1, 248); GCRL 13887 (3, 231.5-256); GCRL 14354 (1, 58.5); GCRL 15238 (1, 141); GCRL 15239 (7, 69-176); GCRL 16278 (1, 254); HBF 477 (1, 53.5); HBF 1213 (7, 86.5-120.5); HBF 1214 (2, 183.5-233); HBF 1215 (6, 66-141); HBF 1216 (3, 89-128); HBF 1217 (5, 69.5-109.5); HBF 1218 (4,127-139); UMML 33200 (1, 128); UMML 33201 (2, 186-217); UMML 33222 (10, 117.5-173); Uncat. (1, 265); USNM 68577 (4, 78-135); USNM 117247 (1, 189.5); USNM 117248 (1, ca. 184); USNM 214745 (1, 157); USNM 214746 (2, 97.5-107); USNM 217834 (1, 113). West Florida: ANSP 89708 (1, 284); CAS(SU) 67482 (1, 90.5); FDNR 6507 (1, 27.9); GCRL 12767 (1, 124); GCRL 12771 (1, 136); GCRL 13227 (1, 91); GCRL 13229 (1, 129); GCRL 13255 (1,247); GCRL 13256 (1, 265); GCRL 13257 (1,210); GCRL 13301 (2, 103-105); GCRL 13458 (4, 52.5-108); GCRL 13749 (2, 111-118); GCRL 13752 (1, 143.5); GCRL 13754 (1, 147); GCRL 13755 (1, 177); GCRL 13756 (1, 120.5); GCRL 13757 (1, 120); GCRL 13759 (2, 82.5-150); GCRL 13762 (1, 84); GCRL 13763 (2, 114-129); GCRL 13766 (7,105.5-132); GCRL 13855 (1,155.5); GCRL 13863 (1, 274); GCRL 13864 (1, 191); GCRL 13929 (1, 216); GCRL 13935 (1, 184); GCRL 15230 (1, 94); GCRL 15236 (5, 112-159); UMMZ 153590 (1, 267); USNM 26606 (1, ca. 80); USNM 214747 (1, 141); USNM 214748 (3, 57-73.5); uncat. (2, 62-71). Alabama: GCRL 14901 (8, 78-149); MCZ 43241 (1, ca. 78); USA 4053 (1, 205). Mississippi: GCRL 4214 (1, 60); GCRL 4215 (2, 60-101); GCRL 4216 (2, 44-70); GCRL 4217 (1, 73); GCRL 4218 (1, 64); GCRL 4219 (1, 47); GCRL 4220 (3, 57-60); GCRL 4221 (1, 65); GCRL 4222 (1, 58); GCRL 4223 (1,42); GCRL 4224 (1, 50); GCRL 4225 (3,55-90); GCRL 4226 (3,28-88); GCRL 4227 (20, 58-80); GCRL 4228 (3, 54-71); GCRL 4230 (3, 54-65); GCRL 4231 (1, 48); GCRL 4232 (1, 50); GCRL 4233 (1, 55); GCRL 4234 (1, 54); GCRL 4247 (23, 55-80); GCRL 4248 (2, 51-54); GCRL 4249 (1, 50); GCRL 4250 (1, 35); GCRL 4340 (5, 55-86); GCRL 4341 (1, 19); GCRL 13822 (2, 60-70.5); GCRL 14135 (3, 146.5-190); GCRL 14136 (59, 102.5-186); GCRL 14444 (1, 252.5); GCRL 15166 (2, 109.5-176); USNM 218466 (2, 80-91). Louisiana: USNM 131164 (2, 261-270). Texas: ANSP 129702 (2, 87-145.5); GCRL 15234 (7, 61-146); GCRL 15237 (2, 110-127); GCRL 15251 (2, 167.5-213); GCRL 15253 (1, 61.5); USNM 118657 (7, 52-60); USNM 213524 (2, 90.5-124). MEXICO, Tabasco: CAS 13921 (1, 145.5). Campeche: CAS 13920 (1, 304); UNAM IB/CML P.468

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Memoir I , Part 8—Sears Foundation for Marine Research

(2,124-217). LOG QUESTIONABLE: AMNH 9029 (1, ca. 263), Bermuda; USNM 117400 (1, 170.5), Jamaica.

Syngnathus fuscus Storer 1839 Northern pipefish Figures 24, 28, 83, 84 Syngnathus typhle (not of Linnaeus, 1758). Mitchill, 1814:475 (misident.); Dumeril, 1870:569 (ref.). Syngnathus peckii Storer, 1839:3,163 (nomen nudum, nomenclatural note). Syngnathus fuscus Storer, 1839:162 (orig. descr.; Nahant (?), Massachusetts); DeKay, 1842:321 (characters); Linsley, 1884: 74 (compiled; Connecticut); Dumeril, 1870:568, 574 (in key; descr. based on Savannah spec, reported as S. fasciatus by Kaup, 1856); Bean, 1881:77 (Woods Hole); Schreiner and Miranda Ribeiro, 1903:88 (Woods Hole); Fowler, 1906b:231 (descr.; New Jersey); Kendall, 1908:65 (compiled); Fowler, 1909:407 (Corson's Inlet, New Jersey); Fowler, 1911:12 (Delaware); Fowler, 1912:37, 38 (Virginia); Fowler, 1913:64 (Chincoteague, Virginia); Sumner et al, 1913:746 (Woods Hole); Fowler, 1914a:2 (Sea Isle City, New Jersey); Fowler, 1914b:3 (Ocean City, Maryland); Fowler, 1916:41 (Manasquan, New Jersey); Fowler, 1917:116 (listed); Fowler, 1919a:294 (New Jersey); Kindred, 1921:425 (descr. of chondrocranium in 8-15 mm spec.); Fowler, 1922a:443 (counts); Fowler, 1922b:4 (New Jersey); Huntsman, 1922:62 (Bay of Fundy); Breder, 1924:26 (Sandy Hook, New Jersey); Kindred, 1924:421 (descr. of chondrocranium in 45 mm spec.); Fish, 1925:165, 166, 170 (seasonal distr.; plankton; Woods Hole); Jordan, 1925:451 (listed); Fowler, 1926b:147 (Longport, New Jersey); Fowler 1927:89 (Ocracoke, North Carolina); Jordan and Evermann, 1927:504 (type-species of Syrictes); Nichols and Breder, 1927:66, fig. (biol.; distr.); Hildebrand and Schroeder, 1928:182, Fig. 99 (descr.; in key; biol. notes; seasonal distr.; banded pipefish; Chesapeake Bay); Breder, 1929:102, fig. (in key, notes); Fowler, 1929:609 (New Jersey); Truitt et al., 1929:59 (descr.; Maryland); Breder, 1932:180 (Sandy Hook Bay, New Jersey); Dawson, 1932:492 (crystallization of haemoglobin in erythrocytes); Dawson, 1933:36 (data on erythrocytes); Breder and Nigrelli, 1934:193 (Sandy Hook, New Jersey); Vladykov and McKenzie, 1935:80, Fig. 75 (Nova Scotia); Bigelow and Schroeder, 1936:329 (Maine); Fowler, 1937:306 (Ventnor, New Jersey); Hunninen and Cable, 1940:373 (trematode host); Gunter, 1942:312 (euryhaline); Herald, 1942:129, 133 (in key; distr.); Herald, 1943:11, 15, 69, 97, 143, 149, PL 1, Figs. 17a, b (descr.; in key; comparisons); Warfel and Merriman, 1944:23 (seasonal distr.; rel. abundance; growth; breeding; Connecticut); Fowler, 1945a:62, 181 (Mary-

land and South Carolina); Kahsbauer, 1950:267 (meristic characters; New Jersey and Boston); Fowler, 1952:116 (New Jersey); Bigelow and Schroeder, 1953:312, Fig. 172 (in key; descr.; biol. notes; distr.); Herald, 1953:232 (sr. synon. of S. martini); Mansueti and Scheltema, 1953:5, 14 (Chesapeake Bay); Gordon, 1956:243, 244, 276, figs, (descr.; Rhode Island); Gunter, 1956:350 (compiled); Ray and Ciampi, 1956:201, Col. PL 1 (descr.); Mansueti, 1957:4 (compiled); Kadam, 1958:562 (ref.); Herald, 1959:472 (inverted pouch closure); Bailey et al., 1960:23 (northern pipefish); Bousfield and Leim, 1960:12 (Minas Basin, Nova Scotia); Gordon, 1960:45, PI. 33 (Rhode Island); Leim, 1960:732 (Cape Sable, Nova Scotia); Williams, 1960:342, 346, Figs. 6-8 (rel. abundance; Woods Hole); Bearden, 1961:40, 43 (South Carolina); Kadam, 1961:257, 290, 292 (ref.); Perlmutter, 1961:278, 339, fig. (in key); Schwartz, 1961:394 (seasonal distr.; Chincoteague and Sinepuxtent Bays); de Sylva et al., 1962:27, 99, 139 (seasonal distr.; Delaware River estuary); Pearcy and Richards, 1962:250, 253, 254 (distr. of larvae; Mystic River, Connecticut); Herman, 1963:106 (seasonal distr.; Narraganset Bay); Alperin and Schaefer, 1964:10 (Great South Bay, New York); Anderson, 1964:4A (cytol. of oocyte); Schwartz, 1964:182 (ecol. notes; near Ocean City, Maryland); Bullis and Thompson, 1965:36 (listed); Christensen, 1965:84 (near Jupiter Inlet, Florida); Herald, 1965:370 (compared with S. affinis); Branch, 1966:69 (name); Breder and Rosen, 1966:294, 296 (summary on breeding and reproduction); Leim and Scott, 1966:177, fig. (descr.; biol. and ecol. notes; Canadian distr.); Schaefer, 1967:11 (Fire Island, New York); Anderson, 1968:24 (cytol. of cortical alveoli and yolk); Boyle, 1968:32 (Croton on Hudson, New York); McAllister, 1968:114 (two filiform branchiostegals); Squires and Gorham, 1968:280 (Public Landing, New Brunswick); Tagatz, 1968:38 (St. Johns River, Florida); Whitworth et al., 1968:96, fig. (descr.; possibly enters fresh water; Connecticut); Wicklund et al., 1968:26 (off Jones Inlet, New York); Clark et al., 1969:50 (off New York and Cape Hatteras); Doss and Farr, 1969:634 (trematode host); Haefner, 1969:201 (13.4 ppm dissolved O2; Penobscot River estuary, Maine); Bailey et al., 1970:34 (compiled); Dahlberg and Odum, 1970:386 (seasonal distr.; Georgia); Richards and Castagna, 1970:238, 239, 244 (seasonal distr.; Eastern Shore, Virginia); Dovel, 1971:10 (Chesapeake Bay); Kindred, 1971:420 (type C acidophils present); Smith, 1971:85 (ecol.; distr.; Delaware River); Dahlberg, 1972:329, 339 (seasonal distr.; Georgia); Herald, 1972:139 (Nova Scotia to Florida); Herald and Dawson, 1972:783 (ref.); Shiino, 1972:61 (banded pipefish); Bleakney and McAllister, 1973:372 (Minas Basin, Nova Scotia); Chenoweth, 1973:107, 110 (occurrence of "larvae," near Boothbay Harbor, Maine); Derickson

Fishes of the Western North Atlantic and Price, 1973:555, 556, 558 (Delaware); Maurer and Watling, 1973:25, 62 (ecol; Delaware Bay); Mercer, 1973:7,13 (ecol.; York River, Virginia); Miller and Jorgenson, 1973:310 (meristic characters); Hasse, 1974:282 (early growth rate compared with Syngnathoides biaculeatus); Lippson and Moran, 1974:161, fig. (summary; characters; spawning; devel.; comparisons); Targett and McCleave, 1974:327 (rel. abundance; Maine); Dahlberg, 1975:52, Fig. 110 (in key; characters; Georgia); Fahay, 1975:20 (off Cape Canaveral, Florida); Kerr, 1975:52 (listed); Rounsefell, 1975:75 (note); Shiino, 1976:109 (compiled); Gilmore, 1977a:133 (listed); Hardy, 1978:402, Figs. 240-243 (characters; ecol.; devel.). Syngnathus Peckianus Storer, 1839:163, PI. 1, Fig. 2 (orig. descr.; Peck's pipefish; Holmes Hole (Martha's Vineyard), Massachusetts; holotype, not located); Linsley, 1844:74 (Connecticut); Storer, 1846:490 (descr.); Gill, 1862:58 (compiled); Storer, 1863:412, PI. 33, Fig. 3 (color notes; synon.; Boston harbor); Gill, 1865:263 (Bay of Fundy); Storer, 1867a:218, PI. 33, Fig. 3 (reprint of Storer, 1863); Dumeril, 1870:571 (descr. compiled); Gill, 1873:15 (Newfoundland to Cape Hatteras); Jones, 1882:94 (Nova Scotia). Syngnathus fasciatus (not of Risso, 1818; not of Gray, 1842) DeKay, 1842:319, PL 54, Fig. 174 (orig. descr.; banded pipefish; New York; holotype, not located); Ayres, 1843:283 (=S. peckianus); Kaup, 1853:232 (name); Kaup, 1856:45 (descr.; notes spec, in MNHN); Dumeril, 1870:569 (compiled in synon. of S. DeKayi); Giinther, 1870:157 (compiled in synon. of S. peckianus)', Jordan and Gilbert, 1882b:383 (=Siphostoma fuscum). Syngnathus viridescens DeKay, 1842:321, PI. 54, Fig. 176 (orig. descr.; New York; holotype, not located); Ayres, 1843:283 (=S. peckianus)', Dumeril, 1870:568, 570 (descr. of spec, from Cape Cod); Giinther, 1870:156 (compiled in synon. of S. peckianus); Swain, 1882:315 (=Siphostoma fuscum). Syngnathus pechianus. DeKay, 1842:321 (misspelling; descr. notes). Syngnathus (sp.?). Storer, 1867a:256 (compared with S. peckianus); Storer, 1867b:280 (ref.). Syngnathus DeKayi Dumeril, 1870:568, 569 (orig. descr.; replacement name for S. fasciatus DeKay, preoccupied; holotype, MNHN 4842). Syngnathus Milbertianus Dumeril, 1870:568, 573 (orig. descr.; replacement name for New York spec, of "S. fasciatus'' of Kaup, 1856:45; holotype, MNHN 6109). Syngnathus peckianus. Ayres, 1843:282 (behavior; color notes; Long Island Sound); Giinther, 1870:157 (descr.); Uhler and Lugger, 1876:76 (St. Mary's River, Maryland); Jordan and Gilbert, 1882b:383 (=Siphostoma fuscum); Ryder, 1882:194 (ref. to early devel.); Ryder, 1887:508 (early devel.); Jordan, 1929:114 (in key; New-

77

foundland to Virginia; states that Storer preferred peckianus although fuscus has priority); Greeley, 1939:84 (Long Island Sound); Merriman, 1947:281 (notes on tidal distr. of young; Connecticut); Merriman and Sclar, 1952:179 (notes on pelagic larvae); Herald, 1953:232 (sr. synon. of S. martini); Wheatland, 1956:262, 300 (occurrence of postlarvae; Long Island Sound). ISyngnathus peckianus. Yarrow, 1877:204 (Bird Shoal, North Carolina); Jordan and Gilbert, 1879:368 (comment on Yarrow's material); McMurrich, 1883:623 (osteol. and devel.; Beaufort, North Carolina); Jungerson, 1910:348, 352 (comment on McMurrich, 1883). ISyngnathus fuscus. Goode, 1879:110 (St. Johns River, Florida; questionable ident.; probably scovelli, louisianae or floridae). Siphonostoma fuscum. Jordan and Gilbert, 1879:368 (n. comb.; in part; Beaufort, North Carolina); Goode and Bean, 1879:4 (Massachusetts); Jordan, 1880:22 (comparisons); Jordan and Gilbert, 1882a:264 (refer 1879 record chiefly to louisianae and floridae). Siphonostoma Peckianum. Goode and Bean, 1879:4 (n. comb.; Salem, Massachusetts). Siphonostoma fasciatus (of DeKay). Jordan, 1880:22 (n. comb.; seems to = S. fuscum). Siphonostoma peckianus. Jordan, 1880:22 (n. comb.; seems to = S. fuscum). Siphonostoma viridescens. Jordan, 1880:22 (n. comb.; seems to = S. fuscum). Siphostoma fuscum. Jordan and Gilbert, 1882b:383 (n. comb.; synon., in part; descr.); Swain, 1882:302, 314 (in key; synon.; descr.; range); Bean, 1883:495 (ref.); Jordan, 1887b:850 (listed); Bean, 1888:134 (New Jersey); Bean, 1891:84 (Chesapeake Bay); Kendall, 1896:623 (sympatric with Gasterosteus; Maine); Bean, 1897:356 (food; aquarium spec.); Slonaker, 1897:471, 482, PI. 30, Fig. 59 (descr. of eye foveae); Smith, 1898:94 (seasonal distr.; Woods Hole); Smith and Bean, 1899:185 (Virginia); Gorham, 1901:33 (gas-bubble disease in aquarium fish); Linton, 1901:443 (cestode host; Woods Hole); Bean, 1903:347, 472 (descr.; common pipefish or billed eel; New York); Sharp and Fowler, 1904:507 (Nantucket); Smith, 1907:170 (in key; diagn.; North Carolina); Evermann and Hildebrand, 1910:160 (Riverside, Maryland and Hampton Creek, Virginia); Tracy, 1910:92 (growth rate; seasonal distr.; Rhode Island); Kendall, 1914:45 (range); Wiegmann and Nichols, 1915:44 (Coney Island, New York); Latham, 1916:39 (Long Island); Cox, 1921:111 (Cape Breton Island); Bigelow and Welsh, 1925:175, Fig. 81 (descr.; biol. notes; range); Fish and Johnson, 1937:258, 269 (distr. of young; Bay of Fundy and Gulf of Maine); Needier, 1940:38 (ecol. note; Malpeque Bay); Munk, 1975:111 (ref.). Syngnathus dekayi. Swain, 1882:315 (^Siphostoma fuscum); Bertin and Esteve, 1950:48 (holotype listed).

Memoir 1, Part 8—Sears Foundation for Marine Research

78

1 1 fo^-^^-^-'-'^^:' --;; if^.v^'jn.-.: - j-.^.j'.j^ ,H. y;.^.7-I-:.:: . . . . i -- :•/

.-\ f£v :?tt -o- 1, •.-:•,-.'..*'.--

FIGURE 24. Syngnathus fuscus: female (GCRL 16087), 177 mm. Syngnathus milbertianus. Swain, 1882:315 (=Siphostoma fuscum); Bertin and Esteve, 1950:48 (holotype listed). Siphostoma (Siphostoma) fuscum. Jordan and Evermann, 1896a:762, 770 (n. comb.; descr.; range); Jordan and Evermann, 1896b:327 (compiled). ISiphostoma fuscum. Linton, 1905:359 (records cestodes, trematodes and gregarines in 90 spec, from Beaufort, North Carolina; probably mostly Syngnathus floridae and 5. louisianae but some fuscus may have been included). Syngnathus pellegrini Fowler, 1919b:265, Fig. 11 (orig. descr.; "Gabun, West Africa," locality error; holotype, ANSP 975); Fowler, 1922a:444 (type material listed); Fowler, 1936:556, 558 (in key, descr.).3 3

See also Dawson, C. E., 1981, Proceedings of the Biological Society of Washington, 94(2):464-478.

Syrictes fuscus. Jordan et al., 1930:242 (n. comb.; compiled); Breder, 1938:26, 28 (lower Hudson River); Linton, 1940:76 (trematode host); Pearson, 1941:85 (seasonal distr.; Chesapeake Bay). Syngnathus martini Herre, 1936:358, PL 1, Fig. 2 (orig. descr.; Manila Bay, Philippine Islands; holotype, CAS(SU) 30973); Bohlke, 1953:62 (holotype listed); Herre, 1953:214 (questionable sp.; unique holotype indistinguishable from S. fuscus); Herald, 1953:232 [=S. peckianus (or S, fuscus)]. Syngnathus pekianus. Greeley, 1937:98 (misspelling; lower Hudson River and Long Island Sound). Syngnathus fuscus fuscus. Herald, 1965:363 (n. comb.; name only); Herald and Dawson, 1972:783 (range).

Distinctive Characters. The combination of modally 19 trunk rings, 53-58 tail rings, short

Fishes of the Western North Atlantic head (hi averages ca. 8 in SL), moderate snout depth (averages ca. 5 in snout length) and low ratios of pectoral-fin length in hi (averages 4.4) and dorsal-fin base in hi (averages 0.9) differentiates S. fuscus from western Atlantic congeners. Description. Proportional data based on 128 specimens from various localities, 62.5-278 (x= 124.1) mm. Rings: 18-21 + 34-39, usually 19 + 35-38 (Tables XL, XLl). Subdorsal rings: 6.5-3.0 + 3.75-7.0 = 7.75-12.75 (Tables XXXV, XLIV-XLVI). Fin rays: dorsal 33-49 (Tables XXXIV, XLII); pectoral 12-15, usually 13-14 (Table XLIII). Head. Head length 7.1-9.5 (x = 8.3) in SL, snout length 1.7-2.4 (2.1) in hi, snout depth 2.3-7.9 (5.1) in snout length. Preorbital moderate to narrow (Fig. 24); opercular ridge usually restricted to anterior third of opercle in adults, complete or nearly so in young (ca. 40 mm). Body. Depth at anal ring 3.1-10.1 (5.0) in hi, trunk depth 1.7-5.4 (3.8) in hi. Median ventral trunk ridge often prominent in subadults and adults, sometimes with a fleshy keel in large females; mature females often with deep trunk, particularly in northern populations. Fins. Length of dorsal-fin base 0.6-1.2 (0.9) in hi, pectoral-fin length 3.3-6.3 (4.4) in hi, length of pectoral-fin base 1.4-2.7 (2.0) in pectoral-fin length. Color. Some young and adults are essentially brown throughout, but ground color is usually light tan to brown and the highly variable markings are in darker shades of tan or brown. Young fish (ca. 70-120 mm) usually with dusky lateral stripe on snout and postorbital area above opercle, and with a diagonal bar from posteroventral margin of orbit to anteroventral part of opercle; dorsum of head and snout brownish, the venter pale or shaded lightly with brown; body brownish, often somewhat lighter on dorsum, often mottled or blotched with pale; some fish with dorsum and part or all of sides crossed by 12-13 brown bands (3-4 rings wide) and 0.5-1.0 ring diffuse pale interspaces; venter of

79

trunk and tail tan to brownish, shading to darker caudad. Large specimens (ca. 200 mm) usually with similar markings on head (Fig. 84); dorsum and upper half of sides brown, remainder of sides tan to near white; body bands, when present, usually composed of a close-set series of irregular brown lines and pale interspaces; bands reduced to scattered dark blotches on lower half of sides on trunk; each tail ring often with diffuse brown bar extending ventrad from lateral scutellum; venter of trunk and most of tail unmarked but some fish with irregular dusky blotches on posterior third of tail. Dorsal fin banded, hyaline or shaded with chromatophores; caudal fin dusky to brownish with narrow pale margin. Color in life variably dark greenish, olive, brownish or (infrequently) red on dorsum and sides; venter pale to golden yellow; trunk and tail mottled or with 12-13 brown bands, sometimes with pale bars on lower half of sides of each trunk ring; dorsal fin pale or with diagonal bands; caudal fin brownish (Smith, 1907; Bigelow and Schroeder, 1953; Hardy, 1978). Bigelow and Welsh (1925) note that color may change with changing habitats. Size. Nichols and Breder (1927) record a 305mm TL specimen but examples larger than ca. 280 mm are uncommon. Development and Growth. Fertilized eggs are 0.75-1.0 mm in diameter (Ryder, 1887; Herald, 1943), incubation time is estimated at ca. 10 days (Bigelow and Schroeder, 1953) and pouch larvae reach ca. 10-12.2 mm TL before dispersal (Lippson and Moran, 1974). Available information on larval development has been summarized and illustrated (3.0-14.7 mm TL) by Hardy (1978). Growth to 70 mm in two months is reported for aquarium hatchlings (Tracy, 1910) and length frequency data (de Sylva et al, 1962) indicate average length increment from ca. 60 to 120 mm TL during the June-October period in the Delaware River estuary. Sexual maturity is apparently reached in about 12 mo (Bigelow and Welsh, 1925) but a period of two years or more was estimated by Warfel and Merriman (1944). A 190-mm female contained 860 nearly ripe

80

Memoir 1, Part 8—Sears Foundation for Marine Research

ovarian eggs (Hildebrand and Schroeder, 1928) but minimum size at maturity is unknown for females. Brood-pouch eggs of similar or varying stages of development are usually arranged in 1-3 layers and in 2-8 transverse rows; 104 pouch eggs were counted in a 120-mm male and a 190mm fish contained 570 (Hildebrand and Schroeder, 1928). Males may be brooding at 83 mm (Herald, 1943) but the pouch is seldom developed in fish smaller than 90-100 mm; brood pouch is reported to occupy a minimum of 13 rings (Lippson and Moran, 1974). Among 19 brooding males (94-278 mm) examined here, the pouch was developed below 15-20 rings. Specimens from Nova Scotia and Massachusetts have 17-20 (usually 18-19) pouch rings, whereas fish from Virginia-Florida have 15-18 (usually 16-17). Papillae are not uncommon on contacting margins of brood-pouch folds, but they are seemingly less frequent in males from southern areas. Breeding apparently occurs from early March to October and peaks during May-June in the Chesapeake Bay-New England region (Lippson and Moran, 1974; Hardy, 1978); present material includes brooding males collected in Georgia during January, February and March and in Florida during April and June. Habits and Behavior. Nichols and Breder (1927) noted that S. fuscus had a "rank odor" and suggested that this serves a protective function; this observation has not been repeated by recent workers. Variable behavior of young (over 30 mm) is suggested by data from subsurface plankton samples (Williams, 1960) wherein S. fuscus were more abundant at night and in flood currents than during the day and ebb tides. Hildebrand and Schroeder (1928) and others have noted seasonal migratory patterns in the Chesapeake Bay area with movement toward deep water in autumn and return to shallows in spring. Wicklund et al. (1968) observed torpid individuals lying motionless on or partially buried in sand bottom or curled around sand dollars in depths of 14-17 m at 10.6-11.0°C It is not known whether this is the typical winter be-

havior pattern in northern populations or merely a response to low temperature. Food. Hildebrand and Schroeder (1928) list copepods, amphipods, fish fry, insects and algae as food items of S. fuscus. Bean (1897) fed aquarium specimens "shrimp with eggs" and small Gammarus sp. Enemies. Aside from a report of entanglement in tentacles of Physalia pelagica (Gordon, 1956), there are apparently no records of enemies. One specimen, ca. 150 mm (USNM 154775), is labeled "removed from stomach of cod" (Gadus morhua?) from the Grand Banks. Parasites. Infection by the cestode Rhynchobothrium heterospine [=Lacistorhynchus tenuis (van Benedin)] was reported by Linton (1901). Digenetic trematodes reported by Hunninen and Cable (1940), Linton (1940) and Doss and Farr (1969) include: Cymbephallus vitellosus (Linton) and Opecoebides manteri (Hunninen and Cable). Variation and Relationships. As noted for some congeners, planktonic young of S. fuscus may have a complete opercular ridge and have posterior angles of principal ridges produced to spine-like points on each ring. Herald (1943) found that large specimens may have superior body ridges elevated above the intervening dorsum, that some mature females develop a rather deep trunk and that there is clinal variation in frequency of dorsal-fin rays. Although there is no evidence of geographical variation in counts of trunk or tail rings, present data clearly demonstrate clinal variation in frequencies of dorsalfin rays (Table XXXIV) and subdorsal trunk rings (Table XXXV). Dorsal-fin rays and subdorsal trunk rings are most numerous and dorsalfin origin most advanced in northern samples (Nova Scotia-Massachusetts), whereas counts are lowest and dorsal fin originates somewhat farther caudad in southern material (North Carolina-Florida). Similar variation is reflected in counts of subdorsal trunk rings and total subdorsal rings, and variation in frequency of brood-pouch rings has been noted above. Pectoral-fin rays are modally 13 in most material (Nova Scotia-South Carolina) but modally 14 in

Fishes of the Western North Atlantic

81

TABLE XXXIV. Geographic variation in frequency of dorsal-fin rays in Syngnathus fuscus. Dorsal-fin rays Locale

33

34

35

36

37

38

1

1

1 1

6 2 1

Canada Nova Scotia United States Maine Massachusetts Connecticut New York New Jersey Delaware Virginia N. Carolina S. Carolina Georgia E. Florida

1

2 1

1 1 5

1

3 5 7 25 2

1 8 10 29 7

5

39

4 3 2 4 3

3 21 8

40

41

42

43

44

45

46

1

4

6

6

6

11

1

3 2

6 6

9 1

8 1

2

1 2

2

4 1 2 1

4

1

1

1

2

47

48

49 1

2 1

samples from Georgia and Florida. Present material fails to indicate geographical trends in preserved coloration, Syngnathus affinis Giinther from the Gulf of Mexico, considered a subspecies of S, fuscus by Herald (1965), is here treated as a separate species (see Key and description). Primary Type. Storer's type-material is evidently lost. Distribution and Abundance. This species is known (Fig. 28) from Prince Edward and Cape Breton Islands in the Gulf of St. Lawrence (Needier, 1940; Cox, 1921) south to Jupiter Inlet and lower reaches of the Loxahatchee River,

Florida (Christensen, 1965). A reference to Newfoundland (Gill, 1873) has not been confirmed and records from the Gulf of Mexico refer to S. affinis rather than S, fuscus. Gorham (1901) reported this species to a depth of 182.9 m but this remains unconfirmed and the usually accepted maximum depth is 49.4 m (Hildebrand and Schroeder, 1928). Specimens have been stranded by receding tides (Bleakney and McAllister, 1973), found in surf (Fowler, 1922b; Schaefer, 1967) and young may occur in plankton samples to ca. 178 km offshore (Fish and Johnson, 1937; Merriman and Sclar, 1952; Fahay, 1975).

TABLE XXXV. Geographic variation in frequency of subdorsal trunk rings in Syngnathus fuscus. Subdorsal trunk rings Locale Canada Nova Scotia United States Maine Massachusetts Connecticut New York New Jersey Delaware Virginia N. Carolina S. Carolina Georgia E. Florida

6.50

6.25

6.00

1 1 2

4.25

5.75

5.50

5.25

5.00

4.75

4.50

5

4

8

12

4

5

1

1 1

2 1

12 2

6 4

11 4

2

1 1 5

4

3

1

1

4

1 2 1

1 6 3

4.00

3.75

3.50

3.25

3.00

6 6 17 4

2

1

5

1

1

1 3 1

4 2 1 2

2

7 7 8 30 8

1 2 8 25 3

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Memoir 1, Part 8—Sears Foundation for Marine Research

Although reported from "floating rockweed" (Kendall, 1896) and a broad range of substrates, S. fuscus is a common associate of aquatic vegetation (e.g. eelgrass, Zoster a, Ulva\ Bigelow and Welsh (1925) considered it a year-round resident of Zoster a beds and Needier (1940) mentions that Syngnathus fuscus was not seen in Malpeque Bay for about six years following the 1932 Zostera mortality. This species may be an associate of Thalassia in southern Florida. Syngnathus fuscus is the only pipefish occurring commonly in coastal and estuarine habitats from Long Island Sound to the Gulf of St. Lawrence. Local abundance is variable seasonally but it has been reported among the most abundant fishes in spring-summer collections near New Haven, Connecticut (Williams, 1960) and Woods Hole, Massachusetts (Warfel and Merriman, 1944); it is evidently the most common pipefish in the Chesapeake Bay area (Hildebrand and Schroeder, 1928). This species is evidently uncommon in Florida but it was usually represented by 1-28 specimens in a series of shallow seine collections at St. Simons Island, Georgia. Breder and Nigrelli (1934) found S. fuscus to be "rare" in collections at Sandy Hook, New Jersey during 1933 but "fairly common" during 1934 and similar annual variations probably occur throughout the species range. The species was listed as euryhaline by Gunter (1942, 1956) and, although recorded from a salinity range of 0.0-38.8%o (Christensen, 1965; Dahlberg, 1972), it is apparently most common at 13.0-20.0%o (Mercer, 1973). Maximum recorded temperature range is 3-35°C (de Sylva et al., 1962). Study Material. 545 specimens, 37-278 mm SL. CANADA, Nova Scotia: MNHN 4842 (250 mm, holotype of S. DeKayi); NMC 58-296 (1, 108.5); NMC 61171 (1, 151); NMC 62-64 (7, 69.5-200.5); NMC 72-108 (1, 176); NMC 64-528 (1,181.5); NMC 64-879 (1,130); NMC 66-610 (2, 64.5-83); NMC 72-42 (1, 158.5); NMC 72-110 (6, 88-178); USNM 132856 (18, 114.5-179.5). UNITED STATES, Maine: USNM 82121 (1, 262). Massachusetts: BOC 7786 (30, 77.5-198); MCZ 36046 (2, 208-218); MCZ 50758 (1, 117.5); MCZ 50946 (2, 108.5-128.5); NMC 67477 (1, 101); NMC 67-478 (2, 129-171); RMNH 11545 (1, 166.5); USNM 109525 (1, ca. 164); USNM 109533 (1,205);

USNM 154775 (1, ca. 150); USNM 218821 (1, 146). Connecticut: BOC 5757 (5, 111-149); BOC 5790 (5, 113-138); BOC 5887 (3, 124-159.5); BOC 7541 (1, 181). New York: MNHN 6109 (260, holotype of S. Milbertianus). New Jersey: USNM 104888 (11, 119-175). Delaware: BOC 429 (1, 186). Virginia: USNM 132732 (17, 73-172); USNM 218351 (1, 68). North Carolina: GCRL 6015 (1, 121); GCRL 12229 (1, 251); GCRL 12234 (1, 130); GCRL 12236 (1, 112); GCRL 12239(1, 144); GCRL 14825(11,71.5-110); USNM 51868 (3,155-167). South Carolina: GMBL uncat. (24 [10 lots], 57-260.5); MCZ 36049 (1, 103.5). Georgia: FSM 5441 (1, 66); FSM 23882 (1, 218); FSM 23883 (1, 278); FSM 23885 (1, 203); GCRL 12424 (1, 222); GCRL 13792 (2, 93.5-98); GCRL 13797 (1,116); GCRL 13803 (2, 66-67); GCRL 13806 (1, 115.5); GCRL 13813 (1, 84.5); GCRL 13819 (2, 64.5-66); GCRL 13821 (2, 62.5-69); GCRL 13829 (5, 95-133.5); GCRL 13832 (3, 118-134); GCRL 13835 (1, 77); GCRL 13838 (3, 100-120.5); GCRL 13843 (3, 87-113); GCRL 13844 (3, 85.5-115); GCRL 13889 (2, 72.5-73.5); GCRL 13890 (2, 88-130); GCRL 13891 (3, 71-80); GCRL 13892 (1, 125); GCRL 13894 (5, 83-128); GCRL 13896 (2, 131-134); GCRL 13900 (3, 96-109); GCRL 13901 (6, 76-130); GCRL 13902 (17, 51.5-108); GCRL 13903 (5, 89-142); GCRL 13904 (28, 47.5-96); GCRL 13905 (4, 67-112); GCRL 13906 (2, 71-85); GCRL 13907 (4, 74.5-122.5); GCRL 13909 (2, 60-86); GCRL 13911 (7, 43-136); GCRL 13912 (1, 72.5); GCRL 13914 (7, 66-76.5); GCRL 13916 (6, 64.5-88.5); GCRL 13918 (1, 85.5); GCRL 13919 (2, 58-105.5); GCRL 13920 (5, 67-98.5); GCRL 13921 (10, 67.5-91.5); GCRL 13922 (9, 59-113); GCRL 13953 (5, 41-108); GCRL 13954 (1, 154.5); GCRL 13955 (4, 79.5-158); GCRL 13956 (5, 92-120.5); GCRL 13957 (18, 43-88.5); GCRL 14080 (24, 37-105.5); GCRL 14081 (5, 74-92); GCRL 14083 (10, 46-82); GCRL 14084 (9, 66.5-93); GCRL 14085 (6, 67-122.5); GCRL 14086 (15, 65-90.5); GCRL 14087 (14, 51-83); GCRL 14088 (5, 61-99); GCRL 14089 (9, 58-95); GCRL 14124 (5,68-73); GCRL 14126 (1, 69); GCRL 14128 (3, 59-90); GCRL 14129 (2, 65-66); GCRL 14131 (11, 56-80.5); GCRL 14132 (8, 50-86); GCRL 16087 (5, 148-180); MNHN 6121 (1, 132); USNM 131256 (1, 233). East Florida: CAS 41147 (1, 243); FSM 12022 (6, 78-156); FSU 11901 (8,67.5-173.5); FSU 12614 (1,117);FSU 12916 (1, 97); FSU 13202 (1, 51.5); FSU 13234 (1, 68); SHML 253 (1, 63.5). LOCALITY UNCERTAIN: ANSP 975 (107.0, holotype of S. pellegrini), originally described in error from Gabon, West Africa); ANSP 976-978 (3, 83-99; paratypes of S. pellegrini); CAS(SU) 30973 (168.5, holotype of S. martini), originally reported from Philippine Is., but locality questioned by Herald (1953); MCZ 11711 (10, 106.5-166), "Rio Grande de Sol, Albuquerque," no other data; USNM 133051 (1, ca. 183), originally in lot of S. louisianae (USNM 117247) reportedly collected by Longley at Tortugas on 15 Apr. 1921 (Longley did not arrive in Tortugas until June 1921).

Fishes of the Western North Atlantic Syngnathus scovelli (Evermann and Kendall) 1896 Gulf pipefish Figures 25, 26, 85-87 ?Syngnathus fuscus (not of Storer). Goode, 1879:110 (probable misident; may refer to S. louisianae or S. scovelli). Siphonostoma sp. Jordan, 1880:22 (comparisons; St. Johns River, Florida); Jordan and Gilbert 1882b:383 (descr.). ?Syngnathus louisianae (not of Giinther). Goode and Bean, 1880:333 (misident.; San Marco Island, Florida). Siphostoma affine (not of Giinther). Jordan and Gilbert, 1882a:264 (descr.; Pensacola Bay, Florida); Jordan and Gilbert, 1882b:904, 906 (misident. of sp. det. as Siphonostoma sp. by Jordan, 1880; in key); Swain, 1882:308, 313 (in key; descr.; Pensacola, Florida); Bean, 1883:420 (compiled); Jordan, 1884:114 (Key West, Florida); Jordan and Swain, 1884:231 (color note; Cedar Keys, Florida); Swain and Meek, 1884:238 (in part; distr., except doubtful rec. from Abrolhos Reef, Brazil); Jordan, 1887a:569 (listed, West Indies; from Swain and Meek, 1884); Henshall, 1891:375 (ecol. notes; Florida distr., except doubtful rec. from Garden Key). Siphostoma fuscum (not of Storer). Evermann and Kendall, 1894:109 (misident.; meristic characters; Corpus Christi, Texas); Evermann and Kendall, 1899:63 (compiled). Siphostoma scovelli Evermann and Kendall, 1896:113 (orig. descr.; Shamrock Point, Corpus Christi, Texas; lectotype, USNM 47300); Evermann and Bean, 1897:242 (Indian River, Florida); Evermann, 1899:309 (Grand Plains Bayou and Lake Lapourde (=Palourde), Louisiana); Evermann and Kendall, 1899:63 (compiled); Bohlke, 1953:61 (SU 5470 and 8613 listed incorrectly as syntypes). Siphostoma (Siphostoma) scovelli. Jordan and Evermann, 1896a:762, 769 (n. comb.; in key; descr.; comparisons; range); Jordan and Evermann, 1896b:327 (compiled). Siphostoma louisianae (not of Giinther). Evermann and Kendall, 1899:63 (in part; compiled). Syngnathus scovelli. Jordan and Thompson, 1905:235 (n. comb.; compiled); Weymouth, 1910:137 (Cameron or Chandeleur Islands, Louisiana); Breder, 1929:102 (in key; Scovell's pipefish; Gulf of Mexico); Fowler, 1933:60 (Lake Charles, Louisiana); Hubbs, 1936:281 (Campeche, Mexico); Fowler, 1940a:ll (Boca Grande, Florida); Fowler, 1941b:238, Fig. 4 (St. Johns River, Florida); Gunter, 1942:314 (euryhaline); Hamilton, 1942:188 (fecundity, etc.; Bonita Springs, Florida); Herald, 1942:129,133 (descr. note; in key; range); Herald, 1943:11, 14, 69, 161, Figs. 18a, b (in key; descr.; comparisons; range); Fowler, 1945a:285 (Florida); Gunter, 1945:48 (seasonal distr.; ecol.; Copano and Aransas Bays, Texas); Baughman, 1950:138 (compiled);

83

Behre, 1950:43 (ref.); Bailey et al., 1954:146 (Escambia River, Florida); Hildebrand, 1954:297 (Texas); Reid, 1954:24 (seasonal distr.; ecol.; biol.; Cedar Key, Florida); Kilby, 1955:227 (ecol.; Cedar Key and Bayport, Florida); McLane, 1955:136 (seasonal distr.; biol.; ecol.; St. Johns River, Florida); Gunter, 1956:350 (compiled); Joseph and Yerger, 1956:129 (Alligator Harbor, Florida); Boschung, 1957:216 (in key; descr.; Mobile Bay, Alabama); Caldwell, 1957:145 (ref.); Joseph, 1957:1 (syst.; life hist.; Florida); Simmons, 1957: 183 (Laguna Madre, Texas); Briggs, 1958:267 (range, in part); Hoese, 1959:328 (compiled); Hubbs, 1958:8 (compiled); Strawn, 1958:21 (note on predator); Can and Coin, 1959:32, 109 (in key; descr.; invades St. Johns River to Deland, Florida, also in N Florida springs); Duarte-Bello, 1959:52 (probably, Cuba); Herald, 1959:473 (inverted pouch closure); Bailey et al., 1960:23 (Gulf pipefish); Hoese, 1960:332 (ecol.; Mesquite Bay, Texas); Phillips and Springer, 1960:27 (Caloosahatchee River, Florida); Renfro, 1960:86 (ecol.; Aransas River, Texas); Springer and Woodburn, 1960:31, 93 (seasonal distr.; ecol.; biol.; Tampa Bay and Stuart, Florida); Springer and McErlean, 1962:54 (Matecumbe Key, Florida); Tabb et al., 1962:51 (Coot Bay, Florida); Whatley, 1962:220 (breeding; Lake St. John, Louisiana); Gunter and Hall, 1963:225, 230, 252 (ecol.; St. Lucie estuary, Florida); Hoese and Jones, 1963:42 (Redfish Bay, Texas); Holliman, 1963:85 (trematode host); Powell and Strawn, 1963:115 (breeding; fecundity; W Florida); Turnage, 1964:9 (Lake Palourde, Louisiana); Christensen, 1965:86 (ecol.; biol.; near Jupiter Inlet, Florida); Gunter and Hall, 1965:27, 51 (ecol.; behavior note; Caloosahatchee Estuary, Florida); Herald, 1965:364 (listed); Parker, 1965:210 (Galveston Bay, Texas); Roessler, 1965:314 (trawl data; Biscayne Bay, Florida); Cervigon, 1966b:240 (descr.; Laguna de Arestinga, Venezuela); Miller, 1966:796 (peripheral freshwater species); Moe et al., 1966:27 (listed); Norden, 1966:127 (Vermilion Bay, Louisiana); Hellier, 1967:21 (breeding; Santa Fe River, Florida); Fox and Mock, 1968:46, 52 (Barataria Bay, Louisiana); Richmond, 1968:238 (Horn Island, Mississippi); Smith-Vaniz, 1968:81 (ref.); Tagatz, 1968:38 (St. Johns River, Florida); Whitaker, 1968:10 (in key); Doss and Fair, 1969:635 (compiled); Overstreet, 1969:172 (ref.); Whatley, 1969:441 (descr.; ecol.; biol.; Lake St. John, Louisiana and Mississippi Sound); Bailey et al., 1970:34 (compiled); Hudson et al., 1970:11 (Porpoise Lake, Florida Bay); Mago, 1970:92 (compiled); Resendez, 1970:106, Fig. 2 (descr.; Laguna Tamiahua, Mexico); Roessler, 1970:864, 884 (ecol.; Everglades National Park, Florida); Swingle, 1971:31 (Mobile area, Alabama); Wang and Raney, 1971:28, Tab. 22 (in part only sp. under 200 mm; seasonal distr.; Charlotte Harbor, Florida); Brown, 1972:1 (life hist; biol.; variation;

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Memoir 1, Part 8—Sears Foundation for Marine Research

FIGURE 25. Syngnathus scovelli: female (GCRL 7394), 128 mm. Florida); Herald, 1972:139 (range); Hubbs, 1972:6 (Texas); Mountain, 1972:6 (Crystal River, Florida); Parker et al., 1972:70 (in key); Powell et al., 1972:61 (listed); Christmas and Waller, 1973:350, 396 (ecol.; Mississippi); Duarte-Bello and Buesa, 1973:92 (expected, Cuba); Miller and Jorgenson, 1973:310 (meristic characters); Resendez, 1973:218 (descr.; Laguna de Alvarado, Mexico); Sabins, 1973:52 (Caminada Pass, Louisiana); Clark, 1974:780 (trawl data; Whitewater Bay, Florida); Douglas, 1974:276, 400, fig., map (descr.; Louisiana); Herrema, 1974:54 (Palm Beach and Broward counties, Florida); Swingle and Bland, 1974:34 (ecol. notes; Mobile area, Alabama); Cervigon, 1975:13, 16, Tab. 1 (meristic characters; descr.; comparisons; in key; Laguna la Restinga, Venezuela); Dahlberg, 1975:53 (in key; characters; range); Jones et al., 1975:68, 80, 86 (listed); Kerr, 1975:52 (listed); Moffett, 1975:22, 66 (Chocolate Bayou, Texas); Subrahmanyam and Drake, 1975:453, 455 (St. Marks and Wakula, Florida); Walls, 1975:161, fig. (descr; Gulf of Mexico); Livingston, 1976:384 (Apalachicola Bay, Florida); Shiino, 1976:110 (compiled); Stevenson, 1976:29, 196 (in key; Florida); Brook, 1977:226 (food and feeding; Card Sound, Florida); Gilmore, 1977a:134 (listed); Hoese and Moore, 1977:161 (in key; descr.; range); Livingston et al.,

1977:84, 90, 92 (listed); Swift et al., 1977:40 (Ochlockonee River, Florida); Cooley, 1978:104 (Pensacola area, Florida); Gunter, 1979:270 (ref.); Weinstein and Heck, 1979:105 (Cape Romano and Marco Island, Florida); Lee et al., 1980:580, fig. (notes on biol.; distr.; habitat). Syngnathus elucens (not of Poey). Fowler, 1906a:94, Fig. 9 (misident.; Marquesas and Big Pine Key, Florida); Fowler, 1922a:443 (compiled). Corythoichthys albirostris (not of Kaup). Fowler, 1920:395 (misident.; Marco, Florida); Fowler, 1922a:442 (compiled). Syngnathus pelagicus (not of Linnaeus). Fowler, 1922a:443 (in part; misident. of sp. from Lake George, Florida); Schultz, 1949:102 (Point Macolla, Venezuela); Cervigon, 1966b:239 (compiled); Mago, 1970:92 (compiled). Syrictes scovelli. Jordan et al., 1930:242 (n. comb.; range); Carr, 1936:86 (freshwater; Florida). Syngnathus affinis (not of Giinther). Fowler, 1933:60 (Prien Lake, Louisiana); Fowler, 1941a:147 (compiled, from Swain and Meek, 1884). Syngnathus scovilli. Hubbs, 1936:159 (misspelling; range). ?Syngnathus rousseau (not of Kaup). Fowler, 1940b:761, Fig. 34 (probable misident.; Rio de Janeiro, Brazil); Figueiredo and Menezes, 1980:11, 13, Fig. 23 (in key; characters).

Fishes of the Western North Atlantic Syngnathus folleti (not of Herald). Tommasi, 1964:34 (misident., in part; descr. of female; Brazil). Syngnathus scovelli scovelli. Herald and Dawson, 1972:782 (n. comb.; meristic characters; comparisons); Lee et al., 1980:568 (ref.).

Distinctive Characters. The combination of modally 16 trunk rings, moderate snout length (averages 2.3 in hi) and depth (averages 3.7 in snout length), relatively low dorsal-fin base in hi ratio (averages 1.0) and narrow preorbital distinguishes S. scovelli from western Atlantic congeners. Description. Proportional data based on 339 specimens from various localities, 42.5-169.0 (x = 90.3) mm. Rings: 15-18 + 28-34, usually (85%) 16 + 30-32 (Tables XXXVI, XL, XLI). Subdorsal rings: 5.0-1.0 + 3.25-5.5 = 5.5-9.5 (Tables XXXVIII, XXXIX, XLIV, XLV, XLVI). Fin rays: dorsal 25-37, 29-34 in 98% from U.S.A. and Mexico, 26-31 in 97% from Caribbean and South America (Tables XXXVII, XLII); pectoral 11-17,13-15 in 93% except 12-13 in Brazilian material (Tables XXXVII, XLIII). Head. Head length 6.3-8.9 (x = 7.6) in SL, snout length 1.9-2.7 (2.3) in hi, snout depth 2.4-5.4 (3.7) in snout length. Preorbital narrow (Fig. 25); opercular ridge often curved somewhat dorsad, usually restricted to anterior third of opercle in adults but complete in young to ca. 46 mm. Body. Depth at anal ring 2.5-5.6 (3.8) in hi, trunk depth 1.5-4.7 (2.7) in hi. Superior ridges somewhat elevated in large specimens; mature females with deep trunk, the venter clearly V-shaped, often with a low but distinct median longitudinal ridge, sometimes with a well-developed fleshy keel. Fins. Length of dorsal-fin base 0.9-1.5 (1.0) in hi, pectoral-fin length 3.6-7.7 (5.1) in hi, length of pectoral-fin base 1.1-2.1 (1.5) in pectoral-fin length. Color. Ground color light tan to dark olive brown, markings mainly various shades of brown but some fish are generally plain without distinct secondary markings. Snout usually with a diffuse dusky lateral stripe, opercles and dor-

85

sum of head shaded or mottled with brown, suborbital area and venter of head plain, blotched or shaded with brown. Dorsum of body often lighter than sides, plain, streaked with narrow brown lines, with broad brown bars (3-5 rings wide) and 0.5-1 ring pale interspaces or with broad pale bars and narrow brown interspaces. Upper half of sides of trunk streaked with fine brown lines, faintly barred or with rather well-defined brown stripe; lower half variously mottled or streaked in young and males. Mature or nearly mature females usually have one or more dark margined narrow pale bars on lower half of each trunk ring (Figs. 86, 87); bars may be in parallel series of 2-3 or there may be a single bar which is branched at one or both ends; most prominent on side of trunk, bars may continue on venter and on anterior tail rings. Sides of tail otherwise variously streaked, barred or blotched with brown; median ventral trunk ridge dark brown in some adult females, venter of trunk otherwise mainly shaded with brown, whereas venter of tail is plain, brownish or with indications of brown bars between each ring. Dorsal fin hyaline, shaded with chromatophores or with indications of dusky blotches or bands; pectoral fins mainly hyaline; caudal fin brownish, mottled with pale. In life, ground color varies from dark green to olive brown; snout mottled or with dark lateral stripe; trunk and tail encircled with dark bands and lighter interspaces; adult females with black ventral keel and silvery marks on side of trunk rings; dorsal fin generally translucent but dark diagonal bars present in some females; caudal fin dark, margined with white spots (Brown, 1972). Size. Reaches 183 mm but examples larger than ca. 165 mm are uncommon. Development and Growth. Fertilized eggs are 0.9-1.3 mm in diameter (Brown, 1972), incubation time is 11-15 days and pouch larvae reach 12-15 mm (TL?) before dispersal (Joseph, 1957; Whatley, 1969). Aquarium hatchlings can reach 28-30 mm in a month (Joseph, 1957), and 12mm fish may reach 60 mm in 83 days (Whatley,

86

Memoir 1, Part 8—Sears Foundation for Marine Research

1969). Length frequency studies (Joseph, 1957; Brown, 1972) indicate that sexual maturity is reached in about 6 months in Florida and Louisiana populations. Hamilton (1942) noted ovarian eggs in a 52-mm female and counted 94 welldeveloped eggs in an 87-mm female; Reid (1954) found 143 mature eggs in a 121-mm female. Brown (1972) did not find ovarian development below 84 mm and most mature females exceed 100 mm in the Cedar Key area. Males may be brooding at 47 mm (Christensen, 1965) in the East Florida population, and Brown (1972) found 90% of examined Cedar Key males to be mature at 87 mm. Brood-pouch eggs are arranged in 1-3 layers and in 2-8 (usually 2-4) transverse rows; number of pouch eggs generally increases with size of male. The average number of pouch eggs is 56-58 in Cedar Key males (Joseph, 1957; Brown, 1972), whereas average counts are 29.5 in the St. Johns River population (McLane, 1955) and 12.2 in the Santa Fe River (Hellier, 1967); ca. 180 pouch eggs were counted in a 110-mm fish (Brown, 1972). Among 148 brooding males (51-119 mm) examined here, brood pouch extended below 8-16 rings. Specimens from East Florida have 10-13 (usually 11-12) pouch rings, those from the Gulf of Mexico have 8-16 (11-13) and there are 11-14 (12-13) in Venezuelan material. Breeding is continuous in some populations (Reid, 1954; McLane, 1955; Hellier, 1967), but Brown (1972) found reduced breeding frequency during December-March at Cedar Key and reproduction peaks in both spring and early fall. Breeding may be continuous in northern Gulf of Mexico populations during warm winters but is reduced or halted during November-January in cold years (Joseph, 1957). Habits and Behavior. Courtship and mating behavior have been described for aquarium specimens of S. scovelli by Joseph (1957) and Whatley (1969), and apparent differences from behavior described for S. floridae are discussed by Joseph (1957). Brook (1977) found S. scovelli an almost exclusive day feeder in the S Florida Thalassia community. Feeding, described by Joseph (1957), is usually accompanied by a clicking sound. Food. The diet of Syngnathus scovelli consists

largely of microcrustaceans but insect larvae and pupae, gastropods and hatchling pipefish are also included (Reid, 1954; McLane, 1955; Joseph, 1957; Brook, 1977). Aquarium specimens will also accept mosquito larvae, newly hatched brine shrimp, freshwater cladocera and hatchling pipefish (Joseph, 1957; Whatley, 1969). Fish under 70 mm feed mainly on copepods, whereas amphipods are the most common food at lengths over 100 mm (Brown, 1972). There is some increase in variety in the summer and fall diet of the Cedar Key population, with increased consumption of shrimp and copepods coupled with decreased utilization of ostracods (Brown, 1972). Enemies. Strawn (1958) noted predation by the blue crab (Callinectes sapidus) in pushnet collections but references to other enemies of Syngnathus scovelli have not been located. Parasites. Overstreet (1969) failed to find digenetic trematodes in two fish examined from Biscayne Bay, but a monogene, Gyrodactylus shorti was originally described from the pouch of a male Syngnathus scovelli collected in Tampa Bay, Florida (Holliman, 1963). The copepod Lernaea cyprinacea Linnaeus was found on a pipefish from Charlotte Harbor, Florida (R. Cressey, in litt.), and a planarian (Dugesia tigrina) has been reported to infest aquarium specimens (Whatley, 1969). Variation and Relationships. In addition to differences in coloration (Figs. 85-87), mature females typically have a deeper and slightly longer trunk than mature males (Herald, 1943; Joseph, 1957; Brown, 1972). There are some differences in proportional characters between populations (Joseph, 1957; Brown, 1972) and fish from low salinity habitats have relatively longer snouts than those from high salinity areas (Joseph, 1957). Herald (1943) and Joseph (1957) have shown that samples from Alabama, Mississippi and Louisiana often have modal counts of 17 trunk rings (usually 16 elsewhere) and clinal variation in frequencies of dorsal-fin rays has been reported by Joseph (1957), Herald and Dawson (1972) and others. Although few specimens are included from low salinity habitats

Fishes of the Western North Atlantic

87

TABLE XXXVI. Geographic variation in frequencies of trunk and tail rings in Syngnathus scovelli. Trunk rings Locale United States E. Florida W. Florida Alabama Mississippi Louisiana Texas Mexico Belize Guatemala Honduras Panama Venezuela Brazil

15

16

17

6 20

65 165 11 21 2 45* 11

3 9 1 14 4 15

1

3 1 6 9 4

1

14 2 1 1 1 18 2

93 49

Tail rings 18

28

29

30

31

32

2

10 2

23 22

26 97 7 19 5 24* 10

13 68 5 13

3 1 19 6

1

19

2 37 3

60

34

5 1

15 8 2 3 1 3 2 12

1

1

33

2 1

1 1

1

2 1 12

28

* Primary type.

along the northern Gulf of Mexico, present data show a trend toward higher trunk-ring counts in samples from Mississippi, Louisiana and Texas (Table XXXVI); trunk rings are most frequently 15 in Mexican specimens but modally 16 in fish from other areas. Venezuelan samples show the greatest range in tail-ring counts and there is a tendency toward high counts in Caribbean material. Frequencies of dorsal-fin rays (Table XXXVII) are somewhat higher in Gulf of Mexico samples and lowest values occur in some Caribbean and South American specimens. Concomitant with lower counts of dorsal-fin rays,

Caribbean and South American samples have reduced frequencies of subdorsal rings and dorsal-fin origin is somewhat more caudad than that of most other material (Tables XXXVIII, XXXIX). Pectoral-fin rays are modally 14 in East Florida, Gulf of Mexico and Venezuelan samples but this count is often 13 in Caribbean specimens and 12-13 in all examined Brazilian material (Table XXXVII). Although Syngnathus scovelli exhibits considerable geographic and ecological variation in some meristic and morphometric characters, available data do not indicate subspecific treatment for any population.

TABLE XXXVII. Geographic variation in frequencies of dorsal- and pectoral-fin rays in Syngnathus scovelli. Pectoral-fin rays

Dorsal-fin rays Locale

25

United States E. Florida W. Florida Alabama Mississippi Louisiana Texas Mexico Belize Guatemala Honduras Panama Venezuela Brazil * Primary type.

26

27

28

29

30

31

32

33

2 4

19 15

15 48

19 61 1 3

12 41 2 13 2 12 3

4 18 4 7 1 18 3

2 1

1

1 1

2

2

1 4 11 23 20 20

1 40 9

1

1

1 3 3

2

3 4

2

1 4 1

5 8

30 2

11

34

35

36

5 3 5

1 1 4

1

12

7*

1

2

3

37

11

12

1

14

15

32 51 10 53 172 58 4 3 17 3 3 20 1 5 2 10 56* 21 1 14 3 7 3 2 1 1 4 7 1 2 52 120 30 37 36 5

1

13

16

17

5

1

1 2

2

Memoir 1, Part 8—Sears Foundation for Marine Research

88

TABLE XXXVIII. Geographic variation in frequency of subdorsal trunk rings in Syngnathus scovelli. Subdorsal trunk rings Locale

3.75 3.50 3.25 3.00 2.75 2.50 2.25 2.00

5.00 4.75 4.50 4.25 4.00

United States E. Florida W. Florida Alabama Mississippi Louisiana Texas Mexico Belize Guatemala Honduras Panama Venezuela Brazil

1 1

1 1

1

2 1 6 4

3

1 6 2 4 2 5

11 1 1 9 1 23 6

8 8 2 7 1 9* 2

18 31 4 7 1 10 8

9 35 3 3 3 4

17 67

6 14

4 24

5

1.75

1.50 1.25

1.00

1

1 1

2

2 1 2 3

16 7

2 16 3

1 62 32

1 13 5

1 2 1 2 9 10

1

1

1 1 1

* Primary type.

The nominal S. scovelli makaxi (Herald and Dawson, 1972) is here considered a separate species (see Key and description). Primary Type. Although Evermann and Kendall (1896) provided data on some 42 specimens in the original species description, type-material was restricted to single male and female syntypes in USNM 47300. The female, now 113.5 mm, is here selected as the lectotype of Siphostoma scovelli Evermann and Kendall. This specimen has a deep trunk (depth ca. 2 in hi) and retains evidence of pale bars on trunk rings; see Study Material and Tables XL-XLVI for addi-

tional data. The male paralectotype has been recataloged as USNM 218366. Distribution and Abundance. Syngnathus scovelli is known (Fig. 26) from the St. Johns River drainage south to Key West in East Florida, along Gulf of Mexico shores to Campeche, Mexico, from various Caribbean localities from Belize to Panama, from Venezuela and from as far south as Ilha Sao Sebastao, state of Sao Paulo, Brazil. The species also occurs in a number of isolated or semi-isolated freshwater habitats in Florida and Louisiana (McLane, 1955; Hellier, 1967; Whatley, 1962, 1969). Henshall's (1891)

TABLE XXXIX. Geographic variation in frequency of total subdorsal rings in Syngnathus scovelli. Total subdorsal rings Locale

5.50 5.75 6.00 6.25 6.50

United States E. Florida W. Florida Alabama Mississippi Louisiana Texas Mexico Belize Guatemala Honduras Panama Venezuela Brazil * Primary type.

6.75 7.00 7.25 7.50 7.75 8.00 8.25 8.50 2 14

1 1 1 1

2

2

2

2 1 7

1 5 9

2 3 25 22

1

1

10 54 1 3

1

2 7

3 7

38 5

5 1

1

5 27

2

47 10

5 50 3 4 2 5 3

9 55 7 1 16 4

5 18 6 11 1 10*

5 5 1 7 1 17 1

8.75 9.00 1 1 1 2 1 3 1

9.25 9.50

1

2 1

2

Fishes of the Western North Atlantic record from Garden Key has not been verified by subsequent collections from the Dry Tortugas. Similarly, the record from Abrolhos, Brazil (Swain and Meek, 1884) has not been verified. This species is not recorded from offshore trawl or plankton collections and available data indicate that collections are typically from depths of less than 6.1 m. Syngnathus scovelli is a common associate of algal or grass (Thalassia, Halodule) bottoms; it occurs in the Spartina marsh (Joseph, 1957) and is found with Najas, Vallisneria, Ruppia, etc., in freshwater areas (McLane, 1955; Whatley, 1969). This is usually the most abundant pipefish in East Florida and Gulf of Mexico shore collections (Reid, 1954; Springer and Woodburn, 1960; Christensen, 1965, etc.). Although infrequent in Caribbean collections, S. scovelli is reasonably common from the Brazilian states of Pernambuco and Bahia and it is the only pipefish taken in a number of samples from Laguna la Restinga, Isla Margarita, Venezuela. Syngnathus scovelli is the only western Atlantic species of Syngnathus known to have breeding populations in freshwater and it occurs to salinities of 45%o in the Laguna Madre of Texas (Simmons, 1957) and 56%o in Laguna la Restinga, Venezuela (Cervigon, 1975). Recorded temperature extremes are apparently 10°C (Reid, 1954) and 36°C (Christensen, 1965). Study Material. 1325 specimens, 24-183 mm SL, including lectotype and paralectotype. Lectotype. USNM 47300 (113.5 mm, female), Shamrock Point, Corpus Christi, Texas, 29 November 1891, B. W. Evermann and party. Paralectotype. USNM 218366 (100 mm, male), taken with lectotype. Other Material UNITED STATES, East Florida: ANSP 30630-30872 (2, 95-96.5); ANSP 78773 (6, 61-99); ANSP 98148 (1, 91); ANSP 98173 (1, 104); ANSP 134493 (5, 54-106); FDNR 3264 (7,81-115.5); FSM 8664 (2,57-57.5); GCRL 13837 (1, 48.5); GCRL 15248 (1, 53); GCRL 15258 (1, 61); GCRL 15259 (1, 124); HBF 1200 (29, 46-79); HBF 1201 (25, 42-86.5); HBF 1202 (18, 52-87); HBF 1203 (15, 39-76); HBF 1204 (15, 36-82.5); UMML 1062 (6, 47.5-92); UMML 2731 (2, 108-113); UMML 33199 (15, 66-120.5); USNM 214738 (1, 123); USNM 214739 (9, 88-119); USNM 214743 (20, 42.5-125). West Florida: ANSP 50635 (1, 74.5); ANSP 78636 (1, 138.5); ANSP 78658 (7, 46-102);

89

45° N.

30°

15°

- 0°

15°

95° W.

60°

65°

50°

35°

30'S. 20°

FIGURE 26. Distribution of Syngnathus scovelli based on material examined. ANSP 83851 (2, 56-86); FSU 18285 (3, 109-128); GCRL 7394 (3, 119-133); GCRL 12198 (2, 85-117); GCRL 12200 (1, 96); GCRL 12201 (1, 109); GCRL 12202 (1, 78); GCRL 12765 (4, 92-100); GCRL 12768 (2, 83-114); GCRL 12769 (1,113); GCRL 12772 (1, 73); GCRL 12774 (1,116); GCRL 12780 (1, 113); GCRL 12781 (2, 73-94); GCRL 12782 (1, 119); GCRL 13183 (3, 67-115); GCRL 13185 (1, 114); GCRL 13187 (1, 96); GCRL 13188 (1, 75); GCRL 13189 (1, 132); GCRL 13193 (2, 84-101); GCRL 13204 (2, 65-77); GCRL 13206 (1, 100); GCRL 13209 (2, 47-91); GCRL 13210 (2, 98-100); GCRL 13224 (5, 91-109); GCRL 13225 (2, 97-111); GCRL 13228 (2, 60-68); GCRL 13231 (1, 72); GCRL 13249 (3, 75-82); GCRL 13250 (2, 88-116); GCRL 13251 (2, 67-69); GCRL 13252 (3, 79-98); GCRL 13254 (2, 82-90); GCRL 13258 (1,109); GCRL 13263 (5, 78-108); GCRL 13264 (3, 78-100); GCRL 13266 (2, 75-90); GCRL 13300 (3, 64-94); GCRL 13343 (11, 62-102); GCRL 13459 (17, 60-108); GCRL 13750 (2, 96-102.5); GCRL 13751 (6, 76.5-113.5); GCRL 13753 (4,96-114); GCRL 13758 (1, 87); GCRL 13760 (1, 105); GCRL 13761 (1, 89); GCRL 13764 (3, 81-85); GCRL 13765 (4, 99-11L5); GCRL 13790 (2, 99.5-107); GCRL 13793 (1, 112.5); GCRL 13805 (1, 74.5); GCRL 13814 (4,84-89.5); GCRL 13830 (5, 68-107); GCRL 13851 (1, 61); GCRL 13853 (2, 108-114); GCRL 13854 (2, 96-105); GCRL 13866 (5, 50-118.5); GCRL 13930 (2, 87-89.5); GCRL 13932 (2, 76-97.5); GCRL 13939 (13, 96.5-118); GCRL 13949 (1, 87.5); GCRL 13950 (3, 67-91); GCRL 13951 (4, 62-78); GCRL 13958 (5,92.5-109); GCRL 14100 (10, 110-169); GCRL 14101 (2, 118-138.5); GCRL 15232 (2, 120-126.5); GCRL 15247 (8, 57.5-116); MNHN 87-512/513 (4, 85-99); UMMZ 60558 (5, 46-76); UMMZ

90

Memoir I , Part 8—Sears Foundation for Marine Research

153641 (96, 59.5-128.5); UMMZ 153649 (112, 70-152.5); UMMZ 153681 (119, 77-133); UMMZ 153697 (92, 73-149.5); USNM 214740 (56, 62-91.5); USNM 214741 (13, 24.3-103); USNM 214742 (62, 61.5-105). Alabama: ASM 254 (2, 82-98); GCRL 14902 (31, 70-102). Mississippi: GCRL 427 (1, 75); GCRL 428 (1, 102); GCRL 847 (4, 74-115); GCRL 1813 (1, 98); GCRL 1814 (1, 88); GCRL 1821 (2, 87-104); GCRL 14139 (17, 82-105); GCRL 14445 (1, 100); GCRL 15165 (2, 46-55); GCRL 15167 (6, 65-105.5). Louisiana: UMMZ 105655 (6, 61-80.5). Texas: CAS(SU) 5470 (4, 70-112); CAS(SU) 8613 (3, 66.5-84.5); FMNH 42911 (1, 90); GCRL 12826 (1, 55); GCRL 12827 (1, 66); GCRL 12828 (1, 94); GCRL 15249 (14, 57.5-92); GCRL 15254 (4, 105-117); GCRL 15502 (1, 60); USNM 69367 (11, 63.5-114); USNM 125617 (14, 69-94); USNM 213525 (6, 72.5-92); USNM 214526 (3, 43-88.5). MEXICO, Veracruz: UNAMIB/CML P.381 (8, 51-95); UNAM uncat. (1, 95). Campeche: UNAM IB/CML P.469 (8, 43-87); UNAM uncat. (8, 42-86). BELIZE: FMNH 83844 (1, 64); USNM 218067 (2, 49-52); USNM 218068 (1, 51). GUATEMALA: CAS 24160 (1, 108); GCRL 4427 (1, 103); UMMZ 202746 (3, 51.5-112.5). HONDURAS: FMNH 84369 (1, 63); FMNH 84374 (1, 65). PANAMA: FMNH 42910 (1, 145.5); GCRL 7974 (2, 50-95.5); USNM 128529 (1, 97.5); USNM 218845 (1, 98); USNM 218846 (1, 104); USNM 218848 (1, 84). VENEZUELA, Falcon: USNM 123162 (4, 115-147); USNM 123163 (1, 102); USNM 123164 (1, 183). Nueva Esparta: FLSM 137 (5, 70-112); FLSM 141 (6,66-101); FLSM 751 (1, 101); FLSM 1202 (1, 87); GCRL 15526 (28, 53-119.5); GCRL 15527 (28, 42.5-116); UDONECI 353 (4, 84-110); UDONECI 853 (18, 52-96); UDONECI 855 (3, 80-113); UDONECI 856 (2, 94-107); UDONECI 857 (2, 68-84); UDONECI 1040 (3, 77-104); UDONECI 1041 (3, 69-76); UDONECI 1042 (6, 69-100); UDONECI 1043 (10, 68-86); UDONECI 1046 (1, 109). BRAZIL, Pernambuco: GCRL 15956 (21, 41-80); GCRL 15957 (1, 87); GCRL 15958 (7,63-82); GCRL 15959 (1, 97); GCRL 15960 (2, 62-79); UFP uncat. (10, 49-91). Bahia: GCRL 10700 (1,106); GCRL 15219 (1, 61); MZUSP uncat. (7, 35-61); MCZ 11705 (2, ca. 118-118.5). Sao Paulo: USNM 144052 (2, 124-129); USNM 144055 (2, 73-90). Loc. uncertain: CAS 6794 (1, ca. 107), "Abrolhos"; MNHN 9243 (2, 112-127.5), "Brazil."

Syngnathus affinis Giinther 1870 Figures 27, 28, 88, 89 Syngnathus affinis Giinther, 1870:163 (orig. descr.; Louisiana; holotype, BMNH 1854.7.3.2); Herald, 1942:131 (=S. fuscus); Herald, 1943: 158(=S. fuscus); Joseph, 1957:4 (ref.). Siphostoma affine. Jordan and Gilbert, 1882b:383 (n. comb.; descr. compiled); Jordan, 1887a:569 (listed); Jordan, 1887b:850 (compiled; distr, in part).

Siphostoma (Siphostoma) affine. Jordan and Evermann, 1896a:769 (n. comb.; descr. compiled); Jordan and Evermann, 1896b:327 (compiled). Syngnathus affine. Breder, 1929:102 (in key; Gulf pipeEsh). Syrictes affinis. Jordan et al., 1930:242 (n. comb.; compiled). Syngnathus fuscus (not of Storer). Hoese, 1958:328 (in part; Texas population referred to S. /. affinis)', Parker et al., 1972:70 (in key; characters, in part; Corpus Christi, Texas); Walls, 1975:159, 162, Eg. (descr.; Louisiana and Texas); Hoese and Moore, 1977:161 (in key; descr., in part; Corpus Christi Bay). Syngnathus fuscus affinis. Hoese, 1958:328 (n. comb.; evidently based on E. S. Herald, in litt.); Herald, 1965:363, 370 (descr., in part; relict northern pipefish; Corpus Christi area, Tex.); Herald and Dawson, 1972:783 (ref.; distr.); Walls, 1975:160 (distr.); Shiino, 1976:109 (compiled); Hoese and Moore, 1977:161 (ref.; distr., in part).

Distinctive Characters. The combination of modally 18 trunk rings, usually 33-34 tail rings and 37-41 dorsal-fin rays, moderate snout length (averages 2.4 in hi) and depth (averages 3.0 in snout length) and narrow preorbital distinguishes S. affinis from western Atlantic congeners. Description. Proportional data based on 26 specimens from Texas, 55.0-218.0 (x = 94.8) mm. Rings: 17-19 + 32-35, modally 18 + 33 (Tables XL, XLI). Subdorsal rings: 5.75-4.0 + 3.5-5.5 = 8.75-10.0 (Tables XLIV, XLV, XLVI). Fin rays: dorsal 35-41, usually (85%) 38-41 (Table XLII); pectoral 13-16, usually (90%) 14-15 (Table XLIII). Head. Head length 7.2-10.0 (x = 8.1) in SL, snout length 2.1-2.6 (2.4) in hi, snout depth 2.4-5.0 (3.0) in snout length. Preorbital narrow (Fig. 27); interorbital somewhat depressed between supraorbital ridges; opercular ridge crosses half of opercle in small fish (55 mm), restricted to anterior third of opercle in largest specimens examined. Body. Depth at anal ring 2.7-5.0 (4.2) in hi, trunk depth 2.1-4.4 (3.3) in hi. Superior ridges somewhat elevated in large specimens; mature females with moderately deep trunk (depth 2.1-2.3 in hi), the venter V-shaped but without prominent median ridge or fleshy keel.

Fishes of the Western North Atlantic

91

' / / : : ^t*-^

%r

l^ffiW

FIGURE 27.

Syngnathus affinis: female (GCRL 15255), 125 mm.

Fins. Length of dorsal-fin base 0.7-1.0 (0.9) in hi; pectoral-fin length 3.0-5.5 (4.4) in hi, length of pectoral-fin base 1.3-2.2 (1.6) in pectoral-fin length. Color. Ground color highly variable and may range from almost white to dark brown within a single collection; markings tan to brown. Among recently preserved specimens (GCRL 15252, 15255), about half are essentially without distinctive markings, whereas the remainder exhibit some degree of body banding. Well marked fish (Fig. 89) usually have a brown snout, a whitish blotch below eye, a pale diag-

onal bar across opercle and remainder of sides and dorsum of head mottled or plain brownish. Body with dorsum and part or all of sides crossed by 11-12 (usually 11) brown bands (2-4 rings wide) and 1-2 ring pale interspaces; venter of trunk and tail variously plain, shaded with brown or with faint continuations of body bands caudad. Dorsal and pectoral fins hyaline in most material but traces of diagonal brownish bands persist on dorsal fins of two large females (FMNH 40309); caudal fin pale, shaded lightly with chromatophores or dark brown with pale distal margin.

Memoir 1, Part 8—Sears Foundation for Marine Research

92

o rM co fM M3 CT\

CO f-«

«-i ON vO ^«

co

co m co CM

in co ^< co 60

C

in vo in

ON CM CO ^

VD 00

O O CO

ON vO

CO

ON

O

8 G o, u d

00

CM

CM

CO

CO

C

'C T3 C rt C H

C O

CM 00 CO

O (N rt ^*

* *

ON r^ vO

vO

CM t-n

Tf ON

QJ

tu X CD

H

*

O ON in O r^ ON CO

H

*+ f-n

Size. Reaches at least 218 mm. Development. Brood pouch extends below 12 and 16 rings in two males (144, 174 mm), the larger fish has pouch eggs arranged in two layers and six transverse rows. Relationships. Among western Atlantic congeners, S. afftnis is most similar to S. fuscus and it was treated as a subspecies of the latter by Herald (1965). Present data (Tables XL, XLI, XLIII) show S. afftnis to differ in counts of trunk rings (modally 18 against 19 in fuscus), tail rings (96% with 34 or fewer against 99% with 35 or more in fuscus), total rings (96% with 52 or fewer against 53 or more in all examined fuscus) and pectoral-fin rays (modally 14 against 13 in fuscus). Although S. fuscus exhibits clinal variation in frequencies of subdorsal trunk rings (Table XXXV), southern samples (North CarolinaFlorida) have lower average counts than examined S. afftnis (3.84 against 4.75). Furthermore, the snout is usually shorter (averages 2.4 in hi against 2.1) and deeper (depth averages 3.0 in snout length against 5.1) in S. afftnis and well marked specimens have fewer body bands (11-12 against 12-13 in fuscus). These differences, together with allopatric distribution (Fig. 28) are considered sufficient basis for retaining species rank for S. afftnis. Primary Type. The holotype (Fig. 88) has a more slender snout than most other study material and the exact type-locality is unknown. The specimen was purchased (Lot 233) at Steven's sales rooms (London) and registered simply "Louisiana." Another miscellaneous lot of fishes and reptiles, registered two days earlier and purchased from the same vendor, was also labeled "Louisiana" with Mr. W. P. Smith as collector (A. C. Wheeler, in litt.). Syngnathus afftnis has not been recognized in subsequent collections from Louisiana and the questionable nature of the type-locality was indicated by Giinther's (1870) failure to discuss the source of the holotype. Should the holotype of S. afftnis prove to be of extralimital origin, a replacement name will be required for the western Gulf of Mexico species described above. Distribution and Abundance. This species is presently known only from the Corpus Christi

Fishes of the Western North Atlantic

93

TABLE XLI. Frequency distributions of total rings in western Atlantic species of Syngnathus. Species floridae springeri louisianae fuscus scovelli affinis makaxi folletti dawsoni pelagicus caribbaeus

43

1

44

45

46

47

48

49

50

1

15

96* 212

251

6

42 122

8

38* 29 1

207* 152

97 11

54

55

56

37

6

2

3

70 149 37 113

103* 95

136

66

1 10 4

49

15 4

5 13*

1 140 48

3 63 75

17 29 28 13* 3 5* 1 14 2 31* 4

4

39

Total rings 51 52 53

8

57

58

59

4 53 22

1 27* 56 20 2 6

60

61

28

3

1 4

* Primary type.

Bay area of Texas and from Laguna de Terminos, Campeche, Mexico. The latter record, omitted from Fig. 28 and Study Material, is based on a 35.5-mm specimen (subsequently lost) collected in a depth of 0-1.2 m on 4 May 1970 by C. E. Dawson and A. Resendez. Counts, from unpublished notes of E. S. Herald, follow: rings 18 + 34, dorsal-fin rays 36, subdorsal rings 4.5 + 5. Although little material is available, the species may be locally abundant. Twenty of the 26 known specimens of S, affinis were taken (with S. louisianae and S. scovelli) in two seine collections at Fish Pass, Corpus Christi Bay, on 27 May 1976. Study Material. 26 specimens, 55-218 mm SL, including holotype. Holotype. BMNH 1854.7.3.2 (144 mm, male), Louisiana, purchased. Other Material Texas, Corpus Christi Bay and vie.: CAS 39674 (1, 79.5); FMNH 40309 (2, 206-218); GCRL 15252 (10, 55-122.5); GCRL 15255 (10, 63-125); USNM 132675(2, 117-174).

Syngnathus makaxi Herald and Dawson 1972 Figures 28, 29, 90, 91 Syngnathus scovelli makaxi Herald and Dawson, 1972:781 (orig. descr.; Laguna Makax, Isla Mujeres, Mexico; holotype, USNM 205235); Resendez, 1975:92,98 (Laguna de Nichupte, Mexico). Syngnathus scovelli makayi. Lee et al., 1980:568 (ref.; lapsus calami).

Distinctive Characters. Syngnathus makaxi is distinguished from western Atlantic congeners by

the combination of modally 14 trunk rings, 22-27 dorsal-fin rays, modally 12 pectoral-fin rays, rather short pectoral fins (length averages 6.7 in hi) and high dorsal-fin base in hi ratio (averages 1.5). Description. Proportional data based on holotype and 15 paratypes, 37.8-62.5 (x = 50.9) mm. Rings: 14-15 + 30-32, modally 14 + 31 (Tables XL, XLI). Subdorsal rings: 3.0-1.5 + 3.5-5.5 = 5.5-6.75 (Tables XLIV, XLV, XLVI). Fin rays: dorsal 22-27, usually (99%) 26 or fewer (Table XLII); pectoral 10-13, 12 or fewer in 94% (Table XLIII). Head. Head length 5.9-6.9 (x = 6.5) in SL, snout length 2.1-2.4 (2.2) in hi, snout depth 3.1-4.7 (3.8) in snout length. Preorbital narrow (Fig. 29); interorbital depressed between supraorbital ridges; opercular ridge complete in small fish (ca. 24-30 mm), restricted to anterior half of opercle in most large specimens. Body. Depth at anal ring 3.8-5.8 (5,0) in hi, trunk depth 2.8-4.5 (3.5) in hi. Dorsum somewhat depressed between superior ridges; trunk moderately deep in mature females (Fig. 91), clearly V-shaped and usually with a low median ventral bony keel. Fins. Length of dorsal-fin base 1.4-1.7 (1.5) in hi, pectoral-fin length 5.4-8.3 (6.7) in hi, length of pectoral-fin base 1.3-1.9 (1.7) in pectoral-fin length. Color. Most specimens tan to light brown, plain or with pale or brownish markings; occa-

Memoir 1, Part 8—Sears Foundation for Marine Research

94

V}

.fi

I

O O^

en -*

O -* oo i-t

S"1 rt ^J CO

CM

O

00 ON

co

cN o.

ON

i-* 4T) -^

ON

O "**

in

sC ^

CM CM CM i-
i^iiiid^^ FIGURE 36. Bryx dunckeri: adult male (GCRL 15743), 74 mm. Arrow indicates anal ring. Islas de San Bernado, Colombia); Cervigon, 1975:15, Tab. 1 (descr.; in key; Venezuela); Kerr, 1975:52 (listed); Fritzsche, 1976:43 (referred to Bryx); Kahsbauer, 1976:385 (Virgin Islands); Shiino, 1976:109 (compiled); Gilmore, 1977a:133 (listed); Figueiredo and Menezes, 1980:11, Fig. 20 (in key; characters). Siphostoma albirostro (not of Kaup). Luederwaldt, 1929:41, 45 (misident., in part; Ilha de Sao Sebastao, Brazil). Syngnathus crinigerum (not of Bean and Dresel). Breder, 1927a:23 (misident., in part; Bahamas). Syngnathus elucens (not of Poey). Beebe and Tee-Van,

1928:85, fig. (misident.; Haiti); Nichols, 1929:217, Fig. 60 (in part; fig. and Haitian record are dunckeri); Beebe and Hollister, 1935:212 (misident.; Union Island, Grenadines). Corythoichthys bermudensis Beebe and Tee-Van, 1932:113 (orig. descr.; Nonsuch Island, Bermuda; holotype, USNM 170913); Beebe and Tee-Van, 1933:80, Fig. 284 (diagn.; Bermuda pipefish; in key); Longley, 1934:271 (=Syngnathus dunckeri); Mead, 1958:132 (holotype listed); Collette, 1962:440 (compiled); Shiino, 1976:107 (compiled).

Memoir I , Part 8—Sears Foundation for Marine Research

114

TABLE XLVII. Geographic variation in frequencies of trunk rings and tail rings in Bryx dunckeri. Tail rings

Trunk rings Locale Bermuda Bahamas Greater Antilles Lesser Antilles United States N. Carolina E. Florida Mexico Belize Honduras Providencia Is. Costa Rica Panama Venezuela Brazil

15

16 1 2 4 4

1 2 1 1 3 1

17

31

32

33

1 18 8 9

2 40 17 43*

12 13 11

3 2 1

1 3

3 3

1

2 4

1

2

1 1 1 9 4

17

18

30

3 67 34 59*

3 2

1

1

1

7 8 5 3 6 1 22 6 1

34

35

1

1

2

5

2

1 1 9 2 2

4

36

10

3

4

* Primary type. Syngnathus jaumei Howell Rivero, 1936:58 (orig. descr.; Guanabo, Cuba; holotype, MCZ 34038); Herald, 1942:131 (=S. dunckeri); Duarte-Bello and Buesa, 1973:91 (compiled). Syngnathus (Microsyngnathus) dunckeri. Herald, 1943:15, 29, 71, 239-246, Fig. 22 (n. comb.; descr.; Duncker's pipefish; comparisons; distr.). Syngnthus dunckeri. Bohlke, 1953:62 (misspelling; Stanford syntypes listed). Syngnathus dunkeri. Springer and McErlean, 1962:54 (misspelling; Matecumbe Key, Florida). ?Synganthus dunckeri. Austin, 1971:32 (misspelling; Puerto Rico), Syngnathus duncheri. Eskinazi, 1972b:293 (misspelling; Pernambuco, Brazil).

Syngnathus rousseaui (not of Kaup), Kahsbauer, 1976:286 (misident; Virgin Islands). Bryx dunckeri. Fritzsche, 1976:58 (n. comb.; name only), Bryx (Bryx) dunckeri. Fritzsche, 1980:192, 197 (n. comb.)

Distinctive Characters. Characters in key (p. 112) distinguish B. dunckeri from its only Atlantic congener, B. randalli. Description. Proportional data based on 49 specimens from various localities, 37.0-95.0 (x =62.2) mm. Rings: 15-18 + 30-36 = 47-52; usually 17 + 31-33, except usually 16 + 34-35 in Brazil (Tables XLVII, LII, LIII).

TABLE XLVIII. Geographic variation in frequencies of dorsal- and pectoral-fin rays in Bryx dunckeri. Dorsal-fin rays Locale Bermuda Bahamas Greater Antilles Lesser Antilles United States N. Carolina E. Florida Mexico Belize Honduras Providencia Is. Costa Rica Panama Venezuela Brazil * Primary type.

21

22

23

24

25

26

27

2 23 12 12

3 32 18 25

9 2 10

2

3

1 5 3 10*

4 2 1 3

1 2

2

4 1 2 1 6

1 2 1 3

3 2 1 3

9

Pectoral-fin rays

3

4 1 4

1 1 8 4 2

2

1 2

10

11

12

2 5 6 10

6 39 25 49

2 13 20 12

6

2 8 2 7 2 6 2 8 3 1

4 2

1 5

3 2 5 2 2 1

13 2

Fishes of the Western North Atlantic

115

TABLE XLIX. Geographic variation in frequencies for dorsal-fin origin in relation to anterior margin of 1st tail ring (0-point) in Bryx dunckeri. + = portion of ring before 0-point; - = portion of ring behind 0-point. Point of dorsal-fin origin Locale

Bermuda Bahamas Greater Antilles Lesser Antilles United States N. Carolina E. Florida Mexico Belize Honduras Providencia Is. Costa Rica Panama Venezuela Brazil

-hl.OO

+0.75

1 1

+0.50

+0.25

5 2 1

1

1

0.00

-0.25

8 2 3

2 28 14 41*

2 13 7 9

1 2

5 2

1

1 3 6 3 12 2 9

1

-0.50

-0.75

1 16 11 3

1 2 1 5

4 1 4

1

1 1

1

1 2 2 2

-1.00

8 1 7

1 1 1

2

* Primary type.

Subdorsal rings: 1.0-0.0 + 5.0-7.5 = 5.0-7.5 (Tables LV, VLI, LVII). Fin rays: dorsal 21-27; pectoral 9-13, modally 10-11 (Tables XLVIII, LIV). Head. Head length 9.0-12.7 (x = 10.9) in SL, snout length 2.8-3.7 (3.3) in hi, snout depth 0.7-2.2 (1.7) in snout length. Snout short and deep; interorbital a little concave between supraorbital ridges; opercular ridge restricted to anterior third of opercle in subadults and adults, usually angled somewhat dorsad, may cross opercle to upper posterior angle in young (ca. 35 mm), opercle elsewhere with low radiating striae; supraopercular ridge inconspicuous or obsolete, other ridges usually low; with or without short simple dermal flaps on eye as well as on dorsum and sides of head (Fig. 36). Body. Depth at anal ring 2.0-6.2 (2.7) in hi, trunk depth 1.6-2.4 (2.0) in hi. Principal ridges usually distinct but not elevated strongly, a little indented between rings, the margins entire to minutely denticulate; dorsum flat to somewhat concave between superior ridges; venter of trunk V-shaped; trunk not exceptionally deep in adult females (Fig. 105). Fins. Length of dorsal-fin base 0.7-0.9 (0.8) in hi, pectoral-fin length 3.6-5.2 (4.5) in hi, length of pectoral-fin base 1.5-2.4 (1.9) in pectoral-fin

length. Dorsal-fin origin usually (90%) at or behind anterior margin of 1st tail ring (Table XLIX); pectoral-fin base usually without clearly defined ridges. Color. Highly variable; ground color ranges from pale to nearly black. Adult females often mainly pale; usually with short radiating brown bars or streaks on eye; body mottled faintly with brownish, usually without bars on dorsum and sides; fins mainly pale or hyaline. Adult males and juveniles more often heavily marked with brown (Figs. 103, 104); usually with dark stripe on side of snout, radiating brown bars or streaks on eye and a dark blotch on postorbital area and upper anterior part of opercle which may be margined by a pale blotch below; dorsum of body often crossed by irregular pale or brown bars or blotches which may continue ventrad on sides, sometimes body merely mottled or blotched; venter and lower half of sides of trunk occasionally with a row of 6-8 small brown spots near middle of each ring; fins plain or the rays spotted with brown. Size. This is a small species which probably seldom reaches 100 mm. Austin's (1971) length of 165 mm (TL?) is undoubtedly based on a misidentification. Development. Among material examined, the

116

Memoir 1, Part 8—Sears Foundation for Marine Research

- 45°N.

30°S. 95°W. 80° 65° 50° 35° 20° FIGURE 37. Distribution of Bryjc dunckeri based on material examined.

smallest male with developing brood pouch was 42 mm. Brood pouch was developed below 14-19 rings in 35 brooding fish, 49.5-95 mm. Brood-pouch eggs are deposited in 2-7 or 8 transverse rows and in 1-3 layers. The smallest brooding male contained 8 eggs in a 14-ring pouch. An 80 mm fish had a single layer of 6 transverse rows through 17 of 18 pouch rings with 45 eggs in the outer right row. An 87 mm specimen had about three layers of eggs in 6 transverse rows through 18 of 19 pouch rings with 50 eggs in the ventralmost right row. Brooding males are here recorded for all months except January, February and September and Bohlke and Chaplin (1968) note brooding fish taken during February in the Bahamas. Enemies. Anderson and Gehringer (1958) list this species among the food items of Thunnus atlanticus. Variation. Brazilian specimens usually have one less trunk ring, one or two more tail rings and somewhat lower counts of dorsal and pectoral-fin rays than other examined material (Tables XLVII, XLVIII). This is not unusual in Brazilian populations of pipefishes and similar meristic variations have been noted here for

Oostethus b. lineatus, Pseudophallus mindii and Syngnathus scovelli. Despite the modal shift from 17 to 16 trunk rings, Brazilian Bryx dunckeri are considered to be conspecific with other Atlantic and Caribbean material Primary Type. Metzelaar (1919:28, Fig. 9) illustrated a brooding male (ca. 50 mm SL) and mentioned specimens from Curasao, Bonaire, St. Eustatius and St. Martin. However, no holotype was designated and no information was given on the length range or number of syntypes. Metzelaar's Bonaire collection has not been located, but there are 22 specimens (39.5-95 mm), identified as Syngnathus dunckeri by Metzelaar and collected by J. Boeke during 1904-1905, which are here considered to be syntypes. Of these, only two brooding males (51.5 and 53.5 mm) approach the calculated length of the illustrated specimen and only the larger of these has the illustrated 15-ring brood pouch. This fish (CAS(SU) 23319) is here designated as the lectotype of Syngnathus dunckeri and the remaining 21 specimens are paralectotypes. The jar containing the lectotype includes a label of unknown origin stating that the specimen was drawn 4X natural size; an enlargement of 3X is indicated in Metzelaar's figure legend. The pectoral and caudal fins are damaged in the lectotype; see Tables XLVII-XLIX and LII-LVII for counts. Distribution and Abundance. Bryx dunckeri is known from the offings of Cape Hatteras, North Carolina (35°23'N, 75°24.5'W), Bermuda and vicinity, the Bahamas, Antilles, SE Florida and Caribbean mainland and Brazilian coasts to 23°50'S (Fig. 37). Herald (1942, 1943) considered this to be an insular or "semi-pelagic" species but present data show that B. dunckeri occupies a variety of habitats from mainland estuaries to coral reefs. Young and subadults (35-51.5 mm) occur in surface plankton and nightlight collections, often at some distance offshore, but there are no confirmed records of brooding males in surface samples. A 45.5-mm fish was taken in an IsaacsKidd trawl off Bermuda (34°27'N, 69°31'W) and a 45-mm specimen was found among floating

Fishes of the Western North Atlantic Sargassum some 9.3-18.5 km E of Alligator Light, Monroe County, Florida. In shore collections, Bryx dunckeri is most commonly taken in depths to 11 m but maximum recorded depth is 72 m (Eskinazi and Holanda Lima, 1968). The species is recorded from Thalassia beds (Bohlke and Chaplin, 1968; Cervigon, 1975), from Diplanthera (Tabb and Manning, 1961), and from calcareous algae (Eskinazi and Holanda Lima, 1968). Tabb et al. (1962) indicate Florida Bay samples from salinities of 20-34%o, Eskinazi (1972b) notes Brazilian collections at 35%o, and four recent samples from Panama were taken in 28-32%o and at temperatures of 23-28.2°C Bohlke and Chaplin (1968) report B. dunckeri as the most common pipefish in their Bahamas collections, whereas it was rare in Herrema's (1974) collections from SE Florida. Among material examined, samples of several or more specimens are not uncommon. Study Material. 346 specimens, 28-95 mm SL, including lectotype and 21 paralectotypes. Lectotype. CAS(SU) 23319 (53.5 mm, male), Curasao, 1904-05, J. Boeke colL Paralectotypes. BMNH 1902.1.8.18 (1, 81.5); CAS(SU) 68582 (1, 51.5); RMNH 9322 (8, 44-69); ZMA 113.100 (2, 91.5-95); data as for lectotype. ZMA 113.101 (9, 39.5-48.5), harbor at St. Eustatius (5.5-11.0 m) and St. Martin, 1904-05, J. Boeke coll. Other Material. BERMUDA: AMNH 16236 (2, 64.5-82.5); ANSP 133402 (1, 71.5); BOC 7783 (1, 87); FMNH 48572 (1, 54.5); FMNH 48785 (1, 48); USNM 170913 (60.3, holotype of Corythoichthys bermudensis). BAHAMAS: AMNH 25850 (1, 42); ANSP 72700 (1, 43.5); ANSP 94549 (25, 34.5-70.5); ANSP 94550 (2, 60.5-66); ANSP 94551 (1, 73); ANSP 94552 (1, 39.5); ANSP 94553 (4, 44.5-70); ANSP 94554 (13, 35.5-49.5); ANSP 94555 (2, 35.5-42); ANSP 94557 (1, 39); ANSP 94558 (15, 39-56.5); ANSP 94559 (8, 35-62.5); ANSP 94560 (3, 34.5-52); ANSP 94561 (6, 38-58.5); ANSP 94562 (1, 39); ANSP 95556 (1, 37); ANSP 101228 (4, 71.5-75.5); ANSP 101229 (6, 35.5-47.5); ANSP 115118 (4, 35.5-44); ANSP 119811 (5, 40.5-47); BOC 91 (1, 40); BOC 92 (1, 49); CAS 30762 (1, 36); UMML 1733 (1, 74.5); UMML 7181 (1, 64.5); UMML 12824 (1, 38.5); UMML 27343 (1, 37); USNM 92727 (1, 77.5). GREATER ANTILLES, Cuba: MCZ 34038 (49, holotype of Syngnathus jaumei); MCZ 34039 (4, 43.5-46.5, paratypes of S. jaumei); USNM 82382 (1, ca. 74); USNM 82383 (1, 47); USNM 107440 (15, 34.5-46). Cayman Is.: BMNH 1939.5.12.66 (1, 49); FSM 17749 (1, 40); FSM 17827 (1, 40.5); FSM 18040 (1, 55.5).

117

Puerto Rico: ANSP 134471 (6, 38.5-50.5); MCZ 51915 (2, 47.5-50); UMML 1965 (2, 56.5-58); UPR 1382 (1, ca. 71); UPR 2619 (2, 45.5-47); UPR 2651 (1, 55); UPR 2789 (2, 45-49); UPR 3261 (13, 41.5-49.5); UPR uncat. (1, 51.5); USNM 130650 (5, 39.5-43). LESSER ANTILLES, Virgin Is.: ANSP 102025 (8, 41.5-50.5); BMNH 1976.7.14.69 (1, 57.5); FMNH 83225 (1, 48.5); GCRL 1726 (2, 45-47); GCRL 15743 (4, 74-83); GCRL 15744 (1, 78.5); ZMUC P.3996 (1, 35); ZMUC P.3997-8 (2, 39-42.5); ZMUC P.3999 (1, 39). Anguilla: ANSP 121881 (3, 40-47.5). St. Barthelemy: ANSP 121868 (2, 49.5-54.5). Barbuda: USNM 190488 (1, 56); USNM 223851 (2, 56.5-63.5). Martinique: ANSP 121866 (1, 70). Grenadine Is.: AMNH 16228 (4, 42.5-50). Grenada: GCRL 13800 (1, 43.5). Klein Bonaire: RMNH 18740 (1, 41). Curasao: RMNH 23228 (1, 62); ZMA 114.118 (7, 41.5-74). Aruba: ZMA 114.168 (1, 48.5). UNITED STATES, North Carolina: AMNH 29637 (1, 45.5); FSM 23886 (8, 39-48); USNM 92668 (1, 40.5); USNM 101559 (5, 35.5-49); USNM 177942 (1, 44.5). East Florida: ANSP 75758 (1, 58); ANSP 134470 (1, 51.5); HBF 1221 (1, 56.5); UMML 3557 (1, 59.5); UMML 4422 (1, 45); USNM 117249 (3, 80-86). MEXICO, Quintana Roo: USNM 192404 (1, 57). BELIZE: GCRL 15866 (1, 45); USNM 218063 (1, 65); USNM 218065 (5, 64-79). HONDURAS, Roatan Is.: FMNH 84367 (2, 39.5-71.5); FMNH 84376 (1, 46). PROVIDENCIA IS.: FSM 18864 (4, 38-55.5); FSM 23551 (1, 43); FSM 24468 (1, 49.5); FSM 24479 (1, 50); FSM 24482 (1, 44). COSTA RICA: UCR 1016-40 (1,38.5); UCR 1128-42 (2, 43.5-45.5). PANAMA: CAS 30758 (6, 37-50); CAS 30760 (7, 39-45); GCRL 4679 (2, 41-44); GCRL 6564 (1, 40); GCRL 9241 (2, 36-59); GCRL 10128 (1, 50); GCRL 10623 (1, 49); GCRL 14042 (2, 44.5-55.5); GCRL 14060 (1, 58); MCZ 50753 (1, 54); MCZ 50754 (1, 43). VENEZUELA: FLSM 1725 (1, 42.5); MBUCV V.1053 (1, 61.5); UDONECI 833 (1, 42); UDONECI 863 (1, 40); UDONECI 877 (1, 47); UDONECI 1051 (1, 42). BRAZIL, Pernambuco: GCRL 12447 (1, 57); UFP uncat. (5, 38-59). Bahia: GCRL 10717 (1, 34); GCRL 15218 (1, 42); MZUSP uncat. (7, 28-59.5). Sao Paulo: MZUSP 9414 (1, 63.5); MZUSP 9416 (1, 55); MZUSP 9417 (2, 44-49); MZUSP 9991 (1, 65). LOC. UNCERTAIN: BOC 7784 (1, 33), 26 Feb. 1933, plankton, sta. 1479, no other data. USNM 217842 (1, 45.9), between Miami and Trinidad, 12-26 June 1968.

Bryx randalli (Herald) 1965 Ocellated pipefish Figures 38, 39, 106 Syngnathus randalli Herald, 1965:364, 373 (orig. descr.; ocellated pipefish; Isla Venados, Venezuela; holotype, USNM 198903); Cervigon, 1966b:239 (descr. notes; Golfo de Cariaco); Mago, 1970:92 (compiled); Cervigon, 1975:17 (in key); Fritzsche, 1976:43 (referred to Bryx); Shiino, 1976:110 (compiled).

118

Memoir 1, Part 8—Sears Foundation for Marine Research

Bryx randalli. Fritzsche, 1976:43 (n. comb.; name only). Bryx (Bryx) randalli. Fritzsche, 1980:197 (n. comb.).

Distinctive Characters. Characters in key (p. 112) best distinguish B. randalli from its only western Atlantic congener, B. dunckeri. Description. Proportional data based on 12 specimens from various localities, 46-93 (x = 64.2) mm. Rings: 17-18 + 30-33 = 47-51 (Tables LII, LIII). Subdorsal rings: 1.0-0.0 + 4.5-6.5 = 5.0-6.5 (Tables LV, LVI, LVII). Fin rays: dorsal 22-27; pectoral 12-14, modally 13 (Table LIV). Head. Head length 8.4-9.7 (x = 9.0) in SL, snout length 2.4-2.8 (2.6) in hi, snout depth 2.8-3.7 (3.3) in hi. Ridges defined clearly; opercular ridge angled dorsad and essentially complete in juveniles (ca. 32 mm), restricted to anterior portion of opercle in adults (Fig. 38), opercle ornamented elsewhere with network of low radiating striae; supraopercular ridge usually distinct; minute simple dermal flaps may be present on opercular, supraopercular and median dorsal ridges. Body. Depth at anal ring 2.4-4.9 (3.3) in hi, trunk depth 2.0-2.9 (2.5) in hi. Principal ridges somewhat elevated, indented to distinctly notched between rings, margins entire to finely denticulate; posterior angles of last 10 or more tail rings sometimes produced to short spinelike points; body surfaces somewhat depressed between principal ridges, ornamented with minute striae; scutella prominent; venter of trunk clearly V-shaped in adults; trunk not exceptionally deep in mature females (Fig. 106); with or without minute simple flaps on superior ridges. Fins. Length of dorsal-fin base 0.9-1.2 (1.1) in hi, pectoral-fin length 5.4-6.9 (5.9) in hi, length of pectoral-fin base 1.2-1.8 (1.4) in pectoral-fin length. Pectoral-fin base with prominent inferior ridge, superior ridge usually smaller, occasionally obsolete. Color. Study material largely faded or bleached in preservative. One specimen (UPR 3266) with a persistent, dark-margined, narrow pale diagonal bar directed posteroventrad on suborbital and with four similar, more or less equally

spaced, bars crossing opercle; dorsum of body mainly light tan; sides mottled brown above lateral trunk ridge and on tail, remainder of trunk with indications of brown bars crossing venter and lower half of sides on anterior V4-% of each ring. The bars on head were described by J. E. Randall as light yellow in the fresh holotype, ground color was light yellowish gray, body markings were various shades of brown, dorsal fin pale, caudal fin dusky yellow and eye with radiating pale yellow lines and a red iris; for additional color notes, see Herald (1965). Size. Reaches at least 93 mm, but doubtless a small pipefish which seldom exceeds 100 mm. Development. Among examined material, the brood pouch is developed below 10-14 rings in five males, 60-76 mm. A 73 mm fish has eggs deposited in single layer of two transverse rows and there are about 60 eggs in the 14-ring pouch. Variation. There are 17 trunk rings in eight specimens from Venezuela and Haiti but this count is 18 in other material. Herald (1965), noting differences from the holotype in head profile and hi in SL ratio, questionably referred two young (31-32 mm) Haitian fish (USNM 164831) to B. randalli. These differences reflect allometry and the Haitian specimens agree with B. randalli in all other features. Despite differences in trunk-ring count and depth of capture (see Distribution), other Caribbean specimens show no basis for separate treatment from the Venezuelan holotype. Distribution. Bryx randalli is known from Haiti, the Lesser Antilles, Belize, Providencia Islands and eastern Venezuela to Golfo de Cariaco (Fig. 39). The holotype was taken over coral bottom, off a rocky shore with sparse growth of mangrove, at a depth of 0.3-0.6 m and a temperature of 22.5° C. Data from six SCUBA samples from Belize and the Antilles indicate collections over rock or coral bottoms with some growth of gorgonians and sponge. Four specimens were taken in 6.1-9.4 m, and single fish were taken in depths of 15.2, 19.8-22.9 and 25.9-27.4 m. Although mentioned in the description of the type-locality (Herald, 1965), mangroves were ev-

Fishes of the Western North Atlantic

119

''

'•

r

'

FIGURE 38. Bryx randalli: holotype (USNM 198903), female, 98 mm. Arrow indicates anal ring.

idently somewhat distant from the collection site. Cervigon (1966b) also notes collections in shallow water near mangrove but it is questionable whether B. randalli is truly a mangrove associate.

Study Material. 16 specimens, 31.5-93 mm SL, including holotype. Holotype. USNM 198903 (93 mm, female), S side of Isla Venados, Venezuela, coral (Porites furcata) bottom, 0.3-0.6 m, 27 January 1965, J. E. Randall. Other Material. GREATER ANTILLES, Haiti: USNM

Memoir I, Part 8—Sears Foundation for Marine Research

120

45°N.

95°W.

65'

50'

FIGURE 39. Distribution of Bryx randalli based on material examined. 164831 (2, 31.5-32). LESSER ANTILLES, Virgin Is., St. Croix: UPR 3266 (1, 65.5). St. Barthelemy: ANSP 121879 (1, 46). Dominica: ANSP 121867 (1, 54.5). St. Vincent: ANSP 121878 (1, 62). Grenadines: ANSP 121876 (1, 60.5); ANSP 121877 (1, 57.5). BELIZE: FMNH 93579 (1, 52). PROVIDENCIA IS.: FSM 24481 (1, 76). VENEZUELA, Sucre: CAS 23945 (1, 73); FLSM 143 (1, 61); FLSM 1802 (2, 60-60.5); UMML 21864 (1, 76).

Genus Cosmocampus Dawson 1979 Cosmocampus Dawson, 1979b:674 (type-species Corythoichthys albirostris Kaup 1856, by original designation). Simocampus Fritzsche, 1980:193 (type-species Siphostoma arctum Jenkins and Evermann 1888, by original designation; as subgenus of Bryx).

Characters. Superior trunk and tail ridges discontinuous near rear of dorsal fin, inferior trunk and tail ridges continuous, lateral trunk and tail ridges discontinuous below dorsal-fin base (Fig. 2d). Snout length 1.7-3.3 in hi; median dorsal snout ridge low, entire to denticulate; median lateral snout ridge present or absent; median dorsal head ridges distinct to strongly elevated; supraopercular ridge present; opercle with complete or incomplete median ridge, usually prominent, often angled dorsad and margined with radiating striae; pectoral-fin base low or protruding somewhat laterad, usually with two dis-

tinct ridges. Principal body ridges prominent, occasionally strongly elevated; ridge margins entire, denticulate to finely serrate in adults, indented to deeply notched between rings; scutella usually well defined; juveniles and adults often with posterior angles of tail rings produced as short spines; dermal flaps usually present in juveniles and/or adults. Trunk rings 15-18, total rings 40-57; dorsal-fin rays 19-27; dorsal-fin origin usually on trunk, total subdorsal rings 4.25-6.25, dorsal-fin base not elevated. Pectoral-fin rays 10-14; anal fin with 2-4 rays; caudal-fin rays typically 10. Brood pouch below 12-20 tail rings; pouch plates usually well developed; pouch folds present with everted, overlapping or semi closure (Fig. 3). Discussion. Among genera of western Atlantic pipefishes, the principal body ridge configuration of Cosmocampus (Fig. 2d) is shared only with Syngnathus Linnaeus and Bryx Herald. This genus differs from Syngnathus sensu stricto in the presence of a supraopercular ridge (Fig. 1) and dermal flaps (absent in Syngnathus) and lacks the inverted pouch closure (Fig. 3) characteristic of Syngnathus. Cosmocampus differs from Bryx in possessing an anal fin and in generally lower frequencies of subdorsal tail rings (mostly 2.25-5.25 against mostly 5.5-7.5 in Bryx). Dawson (1977a) provisionally referred three western Atlantic species (albirostris Kaup, brachycephalus Poey, profundus Herald), previously included in the Indo-Pacific genus Corythoichthys Kaup, to Syngnathus. These species were subsequently referred to Cosmocampus (Dawson, 1979b) and two other western Atlantic species (elucens Poey, hildebrandi Herald) are here included in this genus. In addition, several subtropical-tropical Indo-Pacific forms, now incorrectly included in Syngnathus, may be referable to Cosmocampus. Western Atlantic species of Cosmocampus share a number of common characters but vary considerably in general morphology. More than one phyletic lineage may be represented but, pending further study, introduction of monotypic genera for the accommodation of morphologically atypical species (e.g. elucens) serves no useful purpose. Diagnos-

Fishes of the Western North Atlantic tic characters of Fritzsche's (1980) subgenus Simocampus coincide with those of Cosmocampus. Size. Small to medium-sized pipefishes, seldom exceeding 210 mm. Range. In western Atlantic from about 39° N

121

to 05° S; probably also in subtropical-tropical Indo-Pacific. Species. The number of species referable to Cosmocampus is uncertain but probably approaches 12-15. Five western Atlantic species are herein recognized,

Key to Western Atlantic Species of Cosmocampus la. Snout short and deep; snout length 2.2-3.8 (x = 2.4-3.3) in hi; snout depth 1.4-4.0 in snout length Ib. Snout longer and more slender; snout length 1.7-2.5 (x = 1.7-2.0) in hi; snout depth 3.9-10.2 in snout length 2a. Tail rings 31-35 (usually 32 or more); pectoral-fin rays 10-13 (modally 11). hildebrandi (Herald) 1965, p. 2b. Tail rings 24-31 (usually 30 or fewer); pectoral-fin rays 11-15 (modally 13) 3a. Snout length 2.2-2.8 (x = 2.4) in hi; snout depth 2.6-4.0 (x = 3,4) in snout length, albirostris (Kaup) 1856, p, 3b. Snout length 2.9-3.8 (x = 3.3) in hi; snout depth 1.4-2.2 (x = 1.9) in snout length. brachycephalus (Poey) 1868, p. 4a. Tail rings 29-34; dorsal-fin rays 21-25; snout depth 3.9-9.3 (x = 6.4) in snout length. elucens (Poey) 1868, p. 4b. Tail rings 38-39; dorsal-fin rays 27; snout depth 9.1-10.2 in snout length. profundus (Herald) 1965, p, Cosmocampus albirostris (Kaup) 1856 Whitenose pipefish Figures 40, 41, 43, 107-109 Corythoichthys albirostris Kaup, 1853:231 (nomen nudum; name only, based on Heckel MS); Kaup, 1856:25 (orig. descr.: Mexico and Bahia); Duncker, 1915:74 (descr.); Jordan, 1919:253 (type-species of Corythoichthys Kaup); Parr, 1930:28-30 (generic relationships; in key); Herald, 1940:53 (generic relationships); Herald, 1942:131, 132 (sr. synon. of C. brederi; in key; distr.); Herald, 1943:96,119, 125, 126, Figs. 28a, b (in key; descr.; comparisons; white-nosed pipefish); Whitley, 1948:268 (unavailable as type-species of Corythoichthys); Briggs, 1958:266 (distr.); Duarte-Bello, 1959:50 (probably, Cuba); Herald, 1959:468 (semi pouch closure); Bailey et al., 1960:23 (whitenose pipefish); Springer and Woodburn, 1960:33 (ref.); Springer and McErlean, 1962:44 (Matecumbe Key, Florida); Hildebrand et al., 1964:115 (Alacran Reef, Mexico); Bullis and Thompson, 1965:36 (listed); Herald, 1965:363 (listed); Roessler, 1965:314 (Biscayne Bay, Florida); Moe et al., 1966:24 (listed); Bohlke and Chaplin, 1968:187, fig. (characters; comparisons; Bahamas); Starck, 1968:18 (rare; Alligator Reef, Florida); Overstreet, 1969:169

2 4 139 3 121 128 132 125

(ref.); Bailey et al., 1970:33 (compiled); Resendez, 1971:24 (compiled); Powell et al., 1972:56 (listed); Duarte-Bello and Buesa, 1973:89 (expected, Cuba); Dahlberg, 1975:52, 54 (in key; characters; distr., in part); Kerr, 1975:52 (listed); Resendez, 1975:92 (Laguna de Nichupte, Mexico); Shiino, 1976:107 (listed); Dawson, 1977a:295, 297 (provisionally referred to Syngnathus); Gilmore, 1977a:133 (listed); Dawson, 1979b:674 (type-species of Cosmocampus). Syngnathus albirostris. Giinther, 1870:170 (n. comb.; descr.); Jordan and Gilbert, 1882a:256 (identical with Siphostoma zatropis); Fowler, 1941a:147 (listed, Brazil). Syngnathus (Corythoichthys) albirostris. Dumeril, 1870:550, 563 (in key; n. comb.; descr.). Siphostoma zatropis Jordan and Gilbert, 1882a:264 (orig. descr.; from mouth of "Red Snapper"; conspecific with Mexican spec, descr. by Giinther (1870) as Syngnathus albirostris; holotype, USNM 30856); Swain, 1882:307, 308 (in key; descr; type-locality: "Snapper Banks" near Pensacola, Florida); Jordan and Gilbert, 1882b:905, 906 (in key; descr.; Gulf of Mexico, N to Pensacola); Jordan, 1884:115 (Key West, Florida); Swain and Meek, 1884:237, 238 (in key; Key West); Jordan, 1887a:570 (listed, West Indies); Jordan,

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FIGURE 40. Cosmocampus albirostris: female (GCRL 15535), 102 mm. 1887b:849 (compiled); Jordan and Evermann, 1896a:772 (=S. albirostre). Corythroichthys albirostris. Jordan and Gilbert, 1882a:265 (misspelling; ref.); Jordan and Evermann, 1898:2838 (name); Evermann and Kendall, 1899:63 (Tampa Bay, Florida); Jordan and Thompson, 1905:235 (Tortugas, Florida). Siphostoma (Corythroichthys) albirostre. Jordan and Evermann, 1896a:763, 772 (n. comb.; in key; descr.; distr.); Jordan and Evermann, 1896b:328 (compiled). Siphostoma albirostre. Bean, 1905:321 (listed, Bahamas); Rosen, 1911:50 (compiled); Miranda Ribeiro, 1915:12 (descr. copied).

Syngnathus albirostre. Breder, 1929:102, 104 (in key; ecol. note). Corythoichthys brederi Parr, 1930:30, Fig. 6 (orig. descr.; Cat Island, Bahamas; holotype, BOC 2528); Herald, 1942:131 (=C. albirostris). Hippichthys albirostris. Jordan et al., 1930:243 (n. comb.; compiled); Fowler, 1941c:84 (Howe Key, Florida); Longley and Hildebrand, 1941:56, 57 (in key; descr.; Tortugas, Florida). Syngnathus brederi. Longley, 1934:271 (n. comb.; incorrectly given as sr. synon. of Syngnathus pipulus Beebe and Tee-Van).

Fishes of the Western North Atlantic

123

FIGURE 41. Cosmocampus albirostris: juvenile (UPR 3267), 72 mm.

Distinctive Characters. Cosmocampus albirostris is best distinguished from western Atlantic congeners by characters in key (p. 121). Description. Proportional data based on 64 specimens from various localities, 51-208 (x = 134.1) mm. Rings: 16-18 + 25-31 = 42-49, usually 17-18 + 26-30 (Tables LII, LIII). Subdorsal rings: 2.0-0.0 + 3.5-5.75 = 4.5-6.0, usually 1.5-0.5 + 3.5-4.75 (Tables LV, LVI, LVII). Fin rays: dorsal 21-25; pectoral 11-15, modally 13 (Table LIV). Head. Head length 7.2-10.5 (x = 8.8) in SL, snout length 2.2-2.8 (2.4) in hi, snout depth 2.6-4.0 (3.4) in snout length. Lateral snout ridge present in smaller fish (to ca. 110 mm), often indistinct or obsolete in largest specimens; opercular ridge, straight or angled slightly dorsad, crosses more than half of opercle in both young and adults; opercle elsewhere ornamented with fine radiating striae; supraopercular ridge prominent; supraorbital ridges elevated and flared outward with interorbital rather deeply concave between; other head ridges prominent in both adults and young (Figs. 40, 41); ridge margins

granular to denticulate; usually with minute simple dermal flaps on eye, with or without longer simple or branched flaps on principal ridges. Body. Depth at anal ring 1.9-4.5 (2.7) in hi, trunk depth 1.5-3.1 (2.2) in hi. Principal ridges indented to clearly notched between rings, somewhat elevated, with surfaces of sides and dorsum concave or depressed between; ridge margins granular to denticulate, occasionally somewhat spiny in young (ca. 70 mm); superior posterior angles of most tail rings usually with an enlarged denticule or spine-like point; breadth across superior tail ridges less than that across inferior ridges; venter slightly V-shaped; trunk not exceptionally deepened in mature females; simple or branching dermal flaps present or absent on principal lateral ridges. Fins. Length of dorsal-fin base 1.0-1.5 (1.2) in hi, pectoral-fin length 3.8-6.1 (4.6) in hi, length of pectoral-fin base 1.1-1.8 (1.4) in pectoral-fin length. Pectoral-fin base convex, usually protruding somewhat laterad; ridges distinct to elevated; with or without dermal flaps. Color. Ground color nearly white to brownish.

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Memoir 1, Part 8—Sears Foundation for Marine Research

Snout usually pale, occasionally shaded with brown on dorsum and upper half of sides; upper half of opercle and dorsum of head pale to brown; suborbital and lower half of opercle usually with irregular series of alternating brown and pale bars or blotches directed posteroventrad to isthmus. Sides and dorsum of body (Figs. 107, 108) crossed by 9-11 dark bars (1-3 rings wide) separated by 2-5 ring pale interspaces, bars diffuse or well-defined and mottled with pale; sides and dorsum often pale elsewhere or with irregular mottling above inferior ridge in specimens with dark ground color; venter mainly pale, occasionally with dusky continuations of dark lateral bars; dermal flaps pale; dorsal and pectoral fins hyaline; caudal fin pale or shaded with brown. Size. Reaches at least 208 mm. Development. The smallest examined male with developing brood pouch was 92 mm. Brood pouch extends below 16-20 rings in 12 brooding males, 102-164.5 mm; pouch closure, the semi type (Fig. 3). Eggs are deposited in up to 8-9 transverse rows and usually in a single layer. A 155-mm fish had eggs in a single layer of 7-8 rows, with ca. 73 in the outer right row, through 19 of 20 pouch rings. Present material includes brooding males taken in all months except May and September through December. Enemies. With the exception of a reported occurrence in the mouth of a "red snapper" (Jordan and Gilbert, 1882a), there are apparently no references to predators. Parasites. Digenetic trematodes were not found in two specimens examined by Overstreet, 1969. Variation and Relationships. Available data indicate some meristic differences between samples from East and West Florida. Trunk rings are usually (76%) 17 in Atlantic specimens and total ring counts average 43.9. The 28 available West Florida (Gulf of Mexico) fish have 17 (53.6%) or 18 (46.4%) trunk rings and total rings average 45.7. No significant differences are found in frequencies of dorsal-fin rays but pectoral-fin rays are usually (57%) 11-12 in East Florida fish and 13-15 in 91% of West Florida specimens. Sam-

ples from other areas are few but all fall within the range of Florida material. There is considerable variation in the occurrence of dermal flaps on the body and they are either lost or obsolete in some fish. When best developed, flaps occur just above or on the inferior ridge, on the lateral trunk and tail ridges and on or just below the superior ridge. Flaps are somewhat flattened, sometimes broad, and branching is largely peripheral. The largest and most profusely branched flaps occur on every 4th-5th ring and there may be similar but smaller flaps on the intervening rings. Young fish (Fig. 41) have head and body ridges strongly developed and ridge margins are clearly denticulate to somewhat serrate. In larger specimens (Fig. 40), ridges are less prominent and there is much less ornamentation on ridge margins. Among western Atlantic species of Cosmocampus, this species is most similar to C. elucens. Tail rings are usually fewer in C. albirostris (87% with 25-29 against 30-34 in 99% of elucens) and the snout is shorter (length averages 2.4 in hi against 2.0 in elucens) and deeper in C. albirostris (depth averages 3.4 in snout length against 6.4 in elucens). Primary Type. Kaup's (1856) original description was based on three specimens, supposedly from Mexico and Bahia (Brazil), including a female in the Vienna Museum and a male and female in the Stuttgart Museum. Kaup's trunk ring count of 12 is obviously in error and he included only two pertinent measurements, "length of the body 2.67 inches, of the tail 3.30 inches, ...." Kaup's original material is evidently no longer in these collections. There is a lot, including two females and a male, in Vienna (NMW 59127) labeled "Mexico, 1848. III.l," but the females (162-173.5 mm SL) are longer than the 152 mm TL indicated by Kaup. Search of the Stuttgart collection by Drs. G. von Wahlert and H. Wermuth has failed to locate Kaup's specimens. The British Museum, however, does have a specimen from Mexico, donated by Kaup and bearing a metal tag numbered 13/164, which

Fishes of the Western North Atlantic approximates the length of the original material. This female specimen has 24 dorsal-fin rays and 28 tail rings rather than counts of 27 dorsal rays and 29 rings as given by Kaup. This is presumably also the fish described by Giinther (1870), but trunk and tail ring counts are both one less than those given by Giinther, and it may or not be one of Kaup's syntypes. Distribution. This species is known from Bermuda, the Bahamas, Antilles, Atlantic seaboard and Gulf of Mexico from off South Carolina (32°52'N, 79°09'W) to Texas and the Caribbean coast of Mexico (Fig. 43). The Brazilian record of Luederwaldt (1929) is based on misidentifications of Syngnathus folletti and Bryx dunckeri and there are no confirmed records from areas south of Curagao. There appears little doubt that Kaup's (1856) reference to Brazil is erroneous. Jordan (1884) and Roessler (1965) report occurrence with Thalassia, whereas Jordan and Evermann (1896a) reported occurrence in deeper waters, especially around coral. Present data add nothing useful regarding preferred substrate but collections range from beach shallows to a depth of 49.4 m. There are no records of C. albirostris from areas of low salinity. The maximum number of fish in any examined collection is four and most samples include only single specimens. Present data suggest that the species is most abundant along the South Atlantic seaboard and in the eastern Gulf of Mexico. Collection effort has doubtless been higher in these areas, but it is questionable whether increased effort alone accounts for the fact that 85% of the known specimens have been taken off United States coasts from South Carolina to Alabama. Study Material. 92 specimens, ca. 20-208 mm SL BERMUDA: FMNH 48841 (1, 127); FMNH 48848 (1, 131). BAHAMAS: ANSP 94587 (1, 164.5); BOC 2528 (115, holotype of Corythoichthys brederi); FSM 23889 (1, 96.5). GREATER ANTILLES, Puerto Rico: UPR 3267 (1, 72). LESSER ANTILLES, Grenadines: ANSP 121857 (1, 73). Curasao: CAS 23509 (1, 151). UNITED STATES, South Carolina: CAS 31195 (1, 126); CAS 31197 (1, 103). Georgia: USNM 155578 (1, 87.5). East Florida: AMNH 2801 (2, ca. 123-137); AMNH 8065 (1, 167); AMSP 70583-85

125

(3, 122-155); CAS 339 (1, 142); CAS 31174 (1, 160); CAS 31175 (1, 152); CAS 31176 (1, 120); CAS 31198 (1, 144); FDNR 1916 (1, 106); FMNH 46847 (1, 139.5); FSM 7899 (2, 52-60); FSM 8668 (1, 66); GCRL 15503 (1, 156); GCRL 15535 (1, 102); UMML 2415 (2, 137-155); UMML 2459 (1, 130); UMML 2871 (3, 147-150); UMML 5635 (1, 129.5); UMML 6813 (2, 146-166); UMML 8058 (1, 51); UMML 33194 (1, 107); UMML 33195 (1, 130); UMML 33228 (1, 156); UMMZ 55586 (1, ca. 20); UMMZ 87976 (1, 94.5); UMMZ 201043 (1, 150); USNM 44999 (4, 97-123); USNM 67595 (1, 163.5); USNM 72311 (1, 92); USNM 107195 (1, 135); USNM 131977 (1, 114); USNM 143448 (1, 139); USNM 217820 (1, 136). West Florida: CAS 31172 (2, 166-181); CAS 31173 (1, 184); CAS 31192 (1, 199); CAS 31194 (1, 170); CAS 31199 (1, 179); FDNR 298 (1, 208); FDNR 1890 (1, 205); FDNR 3690 (1, 196); FDNR 4460 (1, 102.5); FDNR 7439 (1, 163); FDNR 7440 (1, 183); FDNR 5123 (1, 171); FDNR 5179 (1, 204); FSM 9237 (1, 102); GCRL 15967 (1, 109); USA 1457 (1, 130); USA 2913 (1, 113.5); USNM 30856 (134, holotype of Siphostoma zatropis); USNM 43869 (1, 131); USNM 72310 (1, 140.5); USNM 73234 (1, 66); USNM 73236 (1, 86); USNM 73237 (1, 104); USNM 73238 (1, 147); USNM 73243 (1, 92); USNM 109531 (1, 140); USNM 132740 (1, 89); USNM 142869 (1, 81.5); UWF uncat. (1, 57.5). Alabama: GCRL 16073 (1, 155). Texas: CAS 31196 (1, 160). MEXICO, Quintana Roo: UNAM IBCML P.474 (1, 76). Loc. unknown: BMNH 1978.9.12.4 (1, 146); NMW 59127 (3, 149.5-173.5).

Cosmocampus profundus (Herald) 1965 Deepwater pipefish Figures 42, 43, 110 Corythoichthys profundus Herald, 1965:363, 364 (orig. descr.; E of Melbourne, Florida; holotype, USNM 198096); Bailey et al., 1970:33 (compiled); Robins, 1971:319, Fig. 1 (descr.; Arrowsmith Bank); Herrema, 1974:164 (name); Shiino, 1976:107 (compiled); Dawson, 1977a:295, 297 (provisionally referred to Syngnathus); Dawson, 1979b:675 (referred to Cosmocampus).

Distinctive Characters. Cosmocampus profundus is distinguished from western Atlantic congeners by higher counts of tail rings (38-39 against 24-35), total rings (56-57 against 40-52) and dorsal-fin rays (27 against 19-25). Description. Proportional data are based on two specimens, 191-197 mm. Rings: 18 + 38-39 (Tables LII, LIII). Subdorsal rings: 0.75-0.5 + 5.25-5.5 = 5.75-6.0 (Tables LV-LVII).

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Memoir I , Part 8—Sears Foundation for Marine Research

% ^^y^r^f^

ly-i-^irt^PlST"

.

..

'j^^^t^ FIGURE 42. Cosmocampus profundus: adult male (UMML 26978), 191 mm.

rays: dorsal 27, pectoral 14-15 (Table LIV). . Head length 8.6-9.0 in SL, snout length 1.7 in hi, snout depth 9.1-10.2 in snout length. Median dorsal snout ridge denticulate or irregularly emarginate over posterior third of snout; lateral snout ridge apparently obsolete; opercular ridge complete or nearly so in young (ca. 100 mm), angled slightly dorsad and restricted to anterior third of opercle in adults (Fig. 42), opercle elsewhere ornamented with radiating striae; supraorbital ridges elevated and flared laterad with interorbital deeply concave between; supraopercular ridge distinct; other head ridges prominent and somewhat elevated. Body. Depth at anal ring 4.4-5.1 in hi; trunk depth 4.0-4.4 in hi. Principal ridges largely notched between rings, elevated and with surfaces of sides and dorsum concave between; ridge margins denticulate on trunk rings of adults but essentially entire on tail rings, den-

ticulate throughout in young; posterior angles of last few tail rings produced to short points; simple dermal flaps present on principal ridges, at least in adults. Fins. Length of dorsal-fin base 1.3-1.4 in hi, pectoral-fin length 6.1-6.7 in hi, length of pectoral-fin base 1.4-1.6 in pectoral-fin length. Pectoral-fin base somewhat convex and protruding laterad, with two prominent ridges. Color. Herald (1965) described the preserved holotype as light tan with small black flecks; dorsal fin with "pigment spots" along base on proximal portion of rays; pectoral and anal fins colorless; caudal-fin membranes with small blackish spots. Robins (1971) described life coloration of a second specimen as pale rosy red with seven dark brown spots, equal to or larger than eye diameter, on side of trunk and anterior part of tail; all fins without pigment. This specimen (UMML 26978) is now largely whitish (Fig.

Fishes of the Western North Atlantic

-

45°N.

127

— 45°N.

—

• hildebrondi O profundus

95°W.

80°

65°

— 45°N.

- 45°N.

95°W.

50°

95°W.

80

65°

FIGURE 43. Distribution of western Atlantic species of Cosmocampus based on material examined.

110); persistent brownish blotches on lower half of side overlie junctures of trunk rings 3-4, 5-6, 8-9, 10-11, 13-14, 15-16 and that of 1st two tail rings; fins hyaline. Size. Reaches at least 197 mm. Development. Brood pouch is developed below 18-21 rings in the two available males. Herald (1965) reported a single layer of six transverse

egg rows in the holotype and counted 131 eggs deposited along 9 pouch rings; pouch closure is the semi type (Fig. 3). The only available young fish is in poor condition and no evidence of dermal flaps persists. Moderately long, slender, simple flaps are present on principal ridges of a number of trunk and tail rings in one adult (UMML 26978), but

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Memoir 1, Part 8—Sears Foundation for Marine Research

flaps are absent (presumably lost) from head ridges; the bases of one or two flaps may be present on eyes. Distribution. Cosmocampus profundus is known from the offings of East Florida (27°56'N), the Virgin Islands and the Yucatan Peninsula, Mexico (Fig. 43). The holotype was taken over coral and sand bottom (Herald, 1965). Two of the three known specimens were trawled in depths of 146-265 m. The third was found in the stomach of a "red snapper" caught in 219.5-274.3 m. This is apparently a benthic species and, if so, it occupies a habitat deeper than that of any other known pipefish. Study Material. Three specimens, ca. 100-197 mm SL, including holotype. Holotype. USNM 198096 (197 mm, male), ca. 167 km ESE of Melbourne, Florida, 27°56'N, 79°05'W, 182.9 m, tumbler dredge; SILVER BAY Sta. 3466; 25 October 1961. Other Material. MEXICO: UMML 26978 (1,191), Banco Arrowsmith, 21°05'N, 86°23'W, 146-265 m, trawl, PILLSBURY Sta. P.581. LESSER ANTILLES, Virgin Is.: USNM 220007 (1, ca. 100), edge of shelf N of St. Thomas, 219.5-274.3 m.

Cosmocampus brachycephalus (Poey) 1868 Crested pipefish Figures 43, 44, 111, 112 Syngnathus brachycephalus Poey, 1868:444 (orig. descr.; Cuba; lectotype, MCZ 11726); Giinther, 1870:156 (name); Poey, 1876:375 (annotation of orig. descr.); Jordan and Thompson, 1905:235 (Tortugas, Florida); Jordan et al., 1930:241 (compiled); Howell Rivero, 1938:184 (syntypes listed). Siphostoma brachycephalum. Jordan, 1887a:570 (n. comb.; compiled). Siphostoma (Siphostoma) brachycephalum. Jordan and Evermann, 1896a:762, 769 (n. comb.; in key; diagn.); Jordan and Evermann, 1896b:327 (listed, Cuba). Corythroichthys cayorum Evermann and Kendall, 1898:128, PI. 7, Fig. 7 (orig. descr.; Key West, Florida; holotype, USNM 48784); Jordan and Evermann, 1898:2838 (descr. compiled); Evermann and Kendall, 1899:63 (Key West); Evermann and Marsh, 1900:107, 108, Fig. 21 (in key; descr.; Puerto Rico); Bean, 1905:321 (listed, Bahamas); Jordan and Thompson, 1905:235 (compiled); Rosen, 1911:50 (listed); Longley, 1917:580 (Tor-

tugas, Florida); Nichols, 1921:22 (Turk Island, Bahamas); Longley, 1932:299 (^Hippichthys brachycephalus); Bohlke, 1953:60 (type-material listed, in part). Syngnathus (Corythroichthys) cayorum. Metzelaar, 1919:27 (n. comb.; descr.; Curasao). Hippichthys cayorum. Nichols, 1929:218, Fig. 61 (n. comb.; diagn.; distr.; Puerto Rico); Jordan et al., 1930:243 (compiled). Corythoichthys cayorum. Parr, 1930:30 (in key); Herald, 1942:131 (=C. brachycephalus). Hippichthys brachycephalus. Longley, 1932:299 (sr. synon. of Corythoichthys cayorum); Fowler, 1941c:84 (Tortugas, Florida); Longley and Hildebrand, 1941:56,57 (in key; descr.; comparisons; Tortugas, Florida); Fowler, 1944a:442 (listed, Bahamas). Corythoichthys brachycephalus. Herald, 1942:132 (n. comb.; in key; distr.); Herald, 1943:96, 133, Figs. 29a, b (in key; descr.; comparisons; distr.); Briggs, 1958:266 (distr.); Duarte-Bello, 1959:50 (compiled); Herald, 1959:468 (semi pouch closure); Bailey et al., 1960:23 (crested pipefish); Boeseman, 1960b:12 (St. Martin and Curasao); Springer and McErlean, 1962:45 (Matecumbe Key, Florida); Caldwell and Caldwell, 1964:15 (Caledonia Bay, Panama); Hildebrand et al., 1964:114 (Alacran Reef, Mexico); Herald, 1965:363 (name); Mago, 1965b:372 (descr.; Los Roques, Venezuela); Caldwell, 1966:35 (listed); Moe et al., 1966:24 (listed); Bohlke and Chaplin, 1968:186, fig. (comparisons; Bahamas); Starck, 1968:18 (Alligator Reef, Florida); Bailey et al., 1970:33 (compiled); Mago, 1970:92 (compiled); Resendez, 1971:24 (compiled); Powell et al., 1972:56 (listed); Shiino, 1972:62 (compiled); DuarteBello and Buesa, 1973:89 (compiled); Guitart, 1975:282, Fig. 215 (diagn.; Cuba); Kerr, 1975:52 (listed); Shiino, 1976:107 (compiled); Dawson, 1977a:295, 297 (provisionally referred to Syngnathus); Gilmore, 1977a:133 (listed); Dawson, 1979b:675 (referred to Cosmocampus). Corythoichthys brachicephalus. Cervigon, 1966b:241 (misspelling; descr.; Los Roques, Venezuela). Corythoichthys branquicephalus. Guitart, 1975:279 (misspelling in key). Cosmocampus brachycephalus. Fritzsche, 1980:Fig. 17c (branchial skeleton).

Distinctive Characters. The combination of short snout (length averages 3.3 in hi), low snout depth/snout length ratio (averages 1.9) and modally 25-26 tail rings and 22 dorsal-fin rays distinguishes Cosmocampus brachycephalus from western Atlantic congeners. Description. Proportional data based on 101 specimens from various localities, 36-93 (x = 70.4) mm.

Fishes of the Western North Atlantic

FIGURE 44. Cosmocampus brachycephalus: adult female (GCRL 14545), 69 mm.

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Memoir 1, Part 8—Sears Foundation for Marine Research

130

TABLE L. Geographic variation in frequencies of trunk, tail and total rings in Cosmocampus brachycephalus. Locale Bahamas Greater Antilles Lesser Antilles United States E. Florida Mexico Belize Guatemala Honduras Panama Colombia Venezuela

15

17

18

59 8* 13

2

103 1 14 7 15 85 11 9

3

16

1 1

3 2 3 1 1

Total rings

Tail rings

Trunk rings 24

25

2 2

4

25

26

27

9 1

38 6* 5

14 1 8

72 1 2 2 4 29 7

13

28

40

43

44

45

9 1

36 6* 6

16 1 7

1

69 1 3 2 4 30 8 1

14 4

1

42

1 1

8 5 7 58 5 7

41

26

1

5

2 3

2 3 3

8 5 7 52 4 6

2 7 3

* Primary type.

Rings: 15-18 + 24-28 = 40-45, modally 17 + 26 (Tables L, LII, LHI). Subdorsal rings: 2.75-0.5 + 2.25-4.75 = 4.25-6.25 (Tables LI, LV-LVII). Fin rays: dorsal 19-25, modally 22; pectoral 11-14, modally 13 (Tables LI, LIV). Head. Head length 7.0-10.6 (x = 9.0) in SL, snout length 2.9-3.8 (3.3) in hi, snout depth 1.4-2.2 (1.9) in snout length. Lateral snout ridge infrequently vestigial, usually obsolete; supraorbital ridges elevated strongly, with interorbital deeply concave between; opercular ridge angled slightly dorsad, crosses half or more of opercle in young and adults; opercle elsewhere finely striate; suborbital crossed by elevated ridge ex-

tending posteroventrad from anteroventral margin of orbit; other ridges prominent, mostly elevated (Fig. 44); eye usually with circlet of minute simple flaps; short; simple or branching flaps usually present on anteroventral margin of snout, on most ridges and on opercular membranes. Body. Depth at anal ring 1.4-2.7 (1.8) in hi, trunk depth 1.4-2.3 (1.6) in hi. Principal ridges elevated, with sides and dorsum concave or depressed between; ridges angled somewhat laterad on posterior half or more of tail; ridges clearly notched or indented between rings, the margins more or less denticulate; angles of posterior tail rings usually pointed; with or without

TABLE LI. Geographic variation in frequencies of dorsal-fin rays and total subdorsal rings in Cosmocampus brachycephalus. Dorsal-fin rays Locale Bahamas Greater Antilles Lesser Antilles United States E. Florida Mexico Belize Guatemala Honduras Panama Colombia Venezuela * Primary type.

19

20

21

22

23

24

22 3 8

17 2* 2

7

1

14 2 4 30

41

28

8

4 2 6 31 4 1

10 2 7 21 3 4

1

6 1

Total subdorsal rings

5 1 4 3 2 28 5

4 4

25

4.25

4.75

5.00

5.25

5.50

5.75

18 1 2

16 2 4

13 2* 6

6

2

4 1 1

1

6

30

30

13

2

3 1

6

13 4 1

1 4 5 33 4 2

8 2 3 29 3 4

31 1 2 6 7 2 2

1 6

1

1

4.50

2

1

6.00 5

1 4

6.25 1

Fishes of the Western North Atlantic short, usually simple, flaps on principal ridges of sides and dorsum of each ring. Fins. Length of dorsal-fin base 0.9-1.4 (1.1) in hi, pectoral-fin length 3.4-5.5 (4.3) in hi, length of pectoral-fin base 1.1-1.7 (1.3) in pectoral-fin length. Pectoral-fin base with two prominent ridges; caudal fin short and rounded (Figs. Ill, 112). Color. Ground color variable, may range from light tan to nearly black within a single collection; markings mainly shades of brown. Side of snout usually blotched in front but crossed posteriad by alternating brown and pale bars which continue on suborbital and ventrad toward isthmus; opercle often with 4-6 more or less equally spaced, narrow, dark-margined diagonal pale lines or bars; head otherwise mottled or blotched in most specimens; some fish with irregular interrupted rows of narrow brown streaks on pale ground color and, in these, the snout and opercular bars may be reduced or absent. Body most commonly mottled or blotched, often with diffuse bars crossing sides and dorsum (Fig. Ill); some fish pale, with irregular interrupted rows of narrow brown streaks on sides and dorsum; venter plain or blotched with brown. Dorsal and pectoral-fin rays narrowly edged with brown, the membranes hyaline; caudal fin variously shaded or blotched, sometimes with pale distal margin. The streaked pattern occurs in both sexes and appears to have no clear relationship to differences in habitat. In life, the posterior third or more of tail is often abruptly pale in contrast to remainder of body and head. Size. Reaches at least 97 mm (Herald, 1943); most specimens are smaller and the species probably seldom exceeds 100 mm. Development. Among examined material, the smallest male with developing pouch was 57 mm; 22 brooding fish (64-92.5 mm) had pouch developed below 12-20 tail rings. Pouch closure is the semi type (Fig. 3) and margins of pouch folds are often minutely scalloped. Eggs are usually deposited in single layer of up to 5 transverse rows; an 85-mm fish had a single layer of 4 rows with 24 eggs in the outer right row through 13 rings of a 17-ring pouch. Herald

131

(1943) noted pouch-egg diameters of 0.8 X 1.8 mm. Present material and the August record of Longley and Hildebrand (1941) indicate that brooding males occur throughout the year. Variation. Trunk-ring counts show little geographic variation, tail rings are modally one less in East Florida samples than those from other areas (Table L). Similarly, specimens from Mexico to Venezuela (Table LI) tend to have slightly lower frequencies of dorsal-fin rays and total subdorsal rings than those from other examined populations. Perhaps the most striking feature is the within-sample variation in coloration of specimens from a limited collection site. This often ranges from nearly black, without obvious markings, to diffusely banded or mainly tan with interrupted brown streaks. Primary Type. The syntypes consist of a male and a female from Cuba and the latter (MCZ 11726) is here designated the lectotype of Syngnathus brachycephalus Poey. This fish retains traces of the streaked color pattern and there are 3 anal-fin rays (see Tables L-LVII for counts). Distribution and Abundance. Cosmocampus brachycephalus is known from the Bahamas, Antilles, lower East Florida coast (Martin County southward), Alacran Reef (Hildebrand et al., 1964) and Quintana Roo, Mexico and western Caribbean localities, south to Bahia de Mochima, Venezuela (Fig, 43). There is one adult male (FMNH 80460) labeled: Whale Bay, Bermuda, 26 June 1910. However, the species has evidently never again been taken at Bermuda and, pending collection of additional material, this is a questionable record. Longley and Hildebrand (1941) found the species to be common on "grass flats" and it appears to be associated with Thalassia or algal beds throughout its range. As many as 6-10 specimens have often been taken with ichthyocide over a 10-15 m long Thalassia patch in Panama. Available records indicate collections in 0-1,8 m. Similarly, salinity and temperature ranges are 33.5-37.8%o and 23.5-34°C, respectively. Study Material. 363 specimens, 19-93 mm SL, including lectotype and paralectotype.

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Memoir 1, Part 8—Sears Foundation for Marine Research

Lectotype. MCZ 11726 (85.5 mm, female), Cuba, 1857, P. Poey. Paralectotype. MCZ 54085 (73.5 mm, male), data as for lectotype. Other Material BAHAMAS: ANSP 72204 (2, 61-67); ANSP 94564 (1, 87); ANSP 94565 (1, 20); ANSP 94566 (1, 27.5); ANSP 94567 (4, 25.5-37.5); ANSP 94568 (2, 62.5-65); ANSP 94569 (8, 25.5-49); ANSP 94570 (1, 36); ANSP 94571 (5, 23.5-43); ANSP 94572 (1, 85); ANSP 94573 (1, 80); ANSP 94582 (1, 19.2); ANSP 101240 (34, 32.5-72.5); ANSP 116456 (1,45.5). GREATER ANTILLES, Cuba: MCZ 36025 (1, 59). Cayman Is.: FSM 13251 (1, 30). Hispaniola: ANSP 120523 (2,73-75.5). Puerto Rico: UPR 3262 (1,60); UPR 3264 (1,39); UPRuncat. (1, 22). LESSER ANTILLES, Virgin Is.: ANSP 102023 (1, 80); ANSP 102024 (1, 77); ANSP 130818 (1, 90); GCRL 1727 (3, 23-36); GCRL 1728 (1, 52); GCRL 1844 (1, 46); UPR 3010 (1, ca. 56). St. Martin: ANSP 105759 (2, 34-50); RMNH 23227 (1, 88.5). Grenada: ANSP 105722 (1, 56.5). Curacao: CAS 39645 (1, 82); RMNH 23226 (1, 81); ZMA 114.120 (1, 93). UNITED STATES, East Florida: ANSP 74887 (1, 76); ANSP 78489 (1, 69.5); ANSP 124223 (11, 52-84); CAS(SU) 6313 (1, 75.5); FDNR 1807 (3, 40-84.5); FDNR 1858 (1, 61); FDNR 1893 (1, 39); FDNR 1947 (1, 72.5); FDNR 5767 (1, 79.5); FDNR 8396 (1, 60); FMNH 46848 (1, 80); FMNH 50691 (89, 51-86.5); HBF 1206 (1, 39); USNM 48784 (87.9, holotype of Corythroichthys cayorum); USNM 127138 (89, paratype of C. cayorum). MEXICO, Quintana Roo: ANSP 114893 (1, 54). BELIZE: AMNH 24626 (1, 50.5); FMNH 83842 (2, 49.5-53); FMNH 83843 (1, 38); USNM 218059 (6, 71-85); USNM 218060 (1, 83); USNM 218061 (1, 82); USNM 218062 (5, 65-85). GUATEMALA: GCRL 4402 (3, 59-67); GCRL 4755 (3, 46-58); USA 702 (1, 73). HONDURAS, Roatan Is.: FMNH 84366 (6, 40.5-85); FMNH 84368 (5, 39-69); FMNH 84370 (1, 29); FMNH 84373 (2, 76.5-84); FMNH 84375 (1, 43). PROVIDENCIA IS.: FSM 24323 (2, 47-51). PANAMA, Canal Zone: GCRL 1891 (6, 55-75); GCRL 3202 (7,41-77); GCRL 3285 (1, 59); GCRL 3319 (6, 53-71); GCRL 6566 (2,56-68); GCRL 7804 (2, 69-70); GCRL 7972 (10, 49-80); GCRL 9719 (1, 73); GCRL 10174 (8, 46-80); GCRL 10304 (1, 66); GCRL 11330 (1, 68); GCRL 11364 (5, 36-71); GCRL 12021 (6, 51-75); GCRL 12658 (1, 77); GCRL 13092 (2, 65-82); GCRL 14544 (5, 48-77); GCRL 14545 (9, 52-73.5); GCRL 15509 (2,71.5-84); GCRL 15510 (4, 65-78); MCZ 50757 (1, 92). Colon: GCRL 4678 (1, 76); UP 477 (1, 56). San Bias: CAS 30761 (1, 27); GCRL 9690 (1, 52); GCRL 10126 (3, 72-86); GCRL 10129 (3, 55-66); GCRL 11025 (1, 67); GCRL 11064 (1, 53); GCRL 11164 (2, 63-66). COLOMBIA, Cartegena: GCRL uncat. (2, 84-88); USNM 213599 (1, 30); USNM 213600 (3, 67-74); USNM 213801 (1, 57); USNM 213802 (5, 59-70); USNM 215333 (1, 32.5). VENEZUELA, Los Roques: MBUCV V.I 168 (1,90.5); UDONECI528 (1, 85); UDONECI uncat.

(5, 38-89). Sucre: UOIO 69.103 (3, 72-85). LOG QUESTIONABLE: FMNH 80460 (1, 92.5), Bermuda.

Cosmocampus elucens (Poey) 1868 Shortfin pipefish Figures 43, 45, 46, 113-116 Syngnathus elucens Poey, 1868:443 (orig. descr.; Cuba; holotype missing); Giinther, 1870:165 (=S, pelagicus); Poey, 1876:373 (redescr.; compared with S. pelagicus); Jordan, 1890b:647 (=Siphostoma rousseau); Jordan and Thompson, 1905:235 (Garden Key, Florida); Breder, 1929:102, fig. (in key; Poey's pipefish; distr.); Nichols, 1929:217 (in part; Havana and Puerto Rico only); Jordan et al, 1930:241 (compiled); Beebe and Tee-Van, 1933:82, Fig. 285 (descr.; Bermuda); Longley, 1933:294 (note); Nichols, 1937:1 (compared with Syngnathus walcotti); Fowler, 1941c:84 (Tortugas, Florida); Longley and Hildebrand, 1941:56, 61 (in part; in key; compared with S. robertsi); Herald, 1942:131, 132 (synon.; in key; distr.); Herald, 1943:229 (referred to subgen. Parasyngnathus Duncker); Springer and Bullis, 1956:68 (listed); Briggs, 1958:267 (distr.); Duarte-Bello, 1959:52 (compiled); Herald, 1959:468 (everted pouch closure; in subgen. Parasyngnathus); Bailey et al., 1960:23 (shortfin pipefish); Springer and Woodburn, 1960:33, 93 (Bella Vista Beach, Florida); Collette, 1962:440 (compiled); Hildebrand et al., 1964:115 (Alacran Reef, Mexico); Nahhas and Cable, 1964:224 (Jamaica); Bullis and Thompson, 1965:36 (listed); Herald, 1965:363, 366 (comparisons): Moe et al., 1966:25 (listed); Bohlke and Chaplin, 1968:192, fig. (descr.; comparisons; Bahamas); Starck, 1968:16 (Alligator Reef, Florida); Bailey et al., 1970: 34 (compiled); Mago, 1970:92 (listed, Venezuela); Resendez, 1971:25 (compiled); Powell et al., 1972:59 (listed); Dawson, 1972:847 (nekton; off Mississippi); Eskinazi, 1972a:37 (estuaries; Ceara, Brazil); Duarte-Bello and Buesa, 1973:91 (compiled); Miller and Jorgenson, 1973:310 (meristic characters); Cervigon, 1975:12, 16, Tab. 1, Fig. 3 (descr.; ecol. notes; in key; incorrect data in table; Isla Cubagua and Bahia de Mochima, Venezuela); Guitart, 1975:279, 282, Fig. 216 (in key; diagn.; Cuba); Walls, 1975:159, 162, fig. (descr.; Gulf of Mexico); Shiino, 1976:109 (listed); Hoese and Moore, 1977:160 (in key); Figueiredo and Menezes, 1980:11, Fig. 21 (in key; Rio Grande do Sul, Brazil). Syngnathus modestus Giinther, 1870:166 (orig. descr.; "1 New Hebrides"; "South America"; syntypes, BMNH 1847.5.14.39 and BMNH 1860.7.18.63); Giinther, 1910:429 (descr.; probably Aneitum); Fowler, 1928:113 (compiled); Herre, 1931:11 (compiled); Fowler, 1944b:189 (compiled); Fowler, 1949:66 (compiled). Siphostoma elucens. Jordan, 1887a:569 (n. comb.; West In-

Fishes of the Western North Atlantic

133

x.:...;^.t;,;^^.i.,.,;:,.,fc.-,.j^ FIGURE 45. Cosmocampus elucens: adult male (USNM 117241), 126 mm.

dies); Carman, 1896:93 (listed); Evermann and Kendall, 1899:63 (compiled); Evermann and Marsh, 1900:107 (in key; Puerto Rico); Bean, 1906:38 (Duck Fish; Bermuda); Metzelaar, 1919:27 (compiled in synon. of Syngnathus rousseau). Siphostoma (Siphostoma) elucens. Jordan and Evermann, 1896a:762, 768 (descr.; prob. ident. with Syngnathus rousseau, closely related to 5. pelagicus); Jordan and Evermann, 1896b:327 (compiled). Siphostoma (Siphostoma) robertsi Jordan and Evermann, 1896b:327 (nomen nudum; incorrectly cite 1896 descr. by Jordan and Rutter). Siphostoma robertsi Jordan and Rutter, 1897:97 (orig. descr.; Kingston, Jamaica; holotype, CAS(SU) 4988);

Jordan and Evermann, 1898:2837 (descr.; comparisons); Bohlke, 1953:61 (holotype listed). Syngnathus robertsi. Metzelaar, 1919:27 (n. comb.; counts; Curacao, Aruba, St. Martin); Metzelaar, 1922:134 (Curacao); Jordan et al., 1930:241 (compiled); Longley, 1932:299 (=S. elucens)', Longley, 1933:294 (prob. not conspecific with S. elucens Poey); Longley and Hildebrand, 1941:56, 60 (in key; descr.; Tortugas, Florida); Herald, 1942:131 (=S. elucens); Fowler, 1944a:461 (compiled); Manter, 1947:295, 375 (trematode host); Doss and Farr, 1969:635 (compiled). Syngnathus rousseau (not of Kaup). Breder, 1927a:23 (misident., in part; Saddle Rock, Bahamas). Syngnathus pipulus Beebe and Tee-Van, 1932:115 (orig.

Memoir I, Part 8—Sears Foundation for Marine Research

134

TABLE LII. Frequency distributions of trunk rings and tail rings in western Atlantic species of Bryx and Cosmocampus. Genus Species

Tail rings

Trunk rings 15

16

17

18 24

25

Bryx dunckeri rand alii

1 36 222* 8*

Cosmocampus albirostris profundus brachycephalus elucens hildebrandi

4 2 56 27 _ 2* 1 10 326* 9 27 127 4 28 150 11 3 32*

26

33 34 35 36 37 38 39

32

27 28 29 30 31

4 50 122* 62 19 1 5 5* 3

6 6

21

151*

19 16 14 37

2

5

3

3

8

1

17 3 15* 7

62 9

1 28 82 2

1*

2

* Primary type. descr.; The Reach, Bermuda; holotype, USNM 170914); Beebe and Tee-Van, 1933:81,285, fig. (characters; little pipefish; in key); Longley, 1934:271 (=S. brederi); Herald, 1942:131 (=S. elucens)', Herald, 1943:232 (tail regenerated in holotype); Mead, 1958:134 (holotype listed); Collette, 1962:440 (compiled); Shiino, 1976:110 (listed). Syngnathus walcotti Nichols, 1937:1 (orig. descr.; St. John, Virgin Islands; holotype, AMNH 13685); Herald, 1942:131 (=S. elucens). Syngnathus (Parasyngnathus) elucens. Herald, 1943:96,229, Fig. 21 (n. comb.; in key; descr.; comparisons; distr.). Syngnathus ? elucens. Eskinazi, 1972b:293 (listed; Pernambuco, Brazil). Parasyngnathus elucens. Fritzsche, 1980:223, Fig. 16 (n. comb.; branchial skeleton).

Distinctive Characters. The combination of long and slender snout (snout depth in snout length ratio averages 6.4), usually (98%) 30-33 tail rings and absence of strongly elevated head ridges (Fig. 45) distinguishes C. elucens from western Atlantic congeners.

Description. Proportional data based on 104 specimens from various localities, 32-164 (x = 108.8) mm. Rings: 15-18 + 29-34 = 45-52, modally 17 + 31 (Tables LII, LIII). Subdorsal rings: 1.5-0.0 + 3.5-5.25 = 4.25-6.0 (Tables LV, LVI, LVH). Fin rays: dorsal 19-25, modally 23; pectoral 11-15, usually (94%) 13-14 (Table LIV). Head. Head length 6.5-9.1 (x = 7.6) in SL, snout length 1.2-2.5 (2.0) in hi, snout depth 3.9-9.3 (6.4) in hi. Lateral snout ridge obsolete; median dorsal snout ridge with granular or finely denticulate margin in adults; supraorbital ridges somewhat elevated and flared laterad with interorbital concave between; opercular ridge usually angled slightly dorsad, crosses %-% of opercle in adults, often reaches to or near upper posterior angle in small fish (ca. 20-30 mm); supraopercular ridge present; other ridges

TABLE LIII. Frequency distributions of total rings in western Atlantic species of Bryx and Cosmocampus. Genus Species

40

41

42

43

44

45

46

Bryx dunckeri rand alii Cosmocampus albirostris profundus brachycephalus elucens hildebrandi

* Primary type.

Total rings 47 48 49 4 1

5 1

31 127

21

14

9

144* 41

2 1

21

7

50

51

52

57 124* 59 3 5* 3

16 2

5

5

53

54

55

56

3 1

2

39

78 1

57

56 9

13 18*

3 5

1 2

1*

Fishes of the Western North Atlantic

135

TABLE LIV. Frequency distributions of dorsal-fin rays, pectoral-fin rays and equivalent (paired) counts of pectoral-fin rays in western Atlantic species of Bryx and Cosmocampus. Genus Species

Dorsal-fin rays

19 20 21

Bryx dunckeri randalli Cosmocampus albirostris profundus brachycephalus elucens hildebrandi

9

10

22

Pectoral-fin rays

23 24 25 26 27

6 37* 75 95 30 1 3 2* 3 5

35

24

12

2

8 49 98 121 58* 16 12 38 94 47 11 11 10* 2

2 9

5 1

Equivalent counts

10 11

12

13

14

2 35 160

68 7

2 13*

8

52

72

9

15 9

1 12 56 25 3

20 1 3* 1, 1 111 197 71* 2 14 161 152 4 6 28 7* 1 2

2*

10 11 12

13 14 1 5*

3

8 1* 38 67 14* 3 58 52

1 20 30

2 9 2 *

* Primary type.

distinct but not elevated strongly; eye usually with one or more minute simple flaps; dermal flaps usually present on principal ridges behind eye (Fig. 45), often narrow and simple but sometimes spatulate with entire or frilled margins, occasionally long and branched on median dorsal ridges. Body. Depth at anal ring 3.0-5.7 (3.7) in hi, trunk depth 2.8-4.2 (3.4) in hi. Principal ridges somewhat elevated with surfaces of sides and dorsum depressed or concave between; ridges indented between rings, the margins usually entire or granular, seldom denticulate in adults; posterior angles of tail rings sometimes produced as short points; venter of trunk V-shaped but trunk not exceptionally deep in mature females (Fig. 115); dorsum of tail rings slightly narrower than breadth across inferior ridges; dermal flaps often present on principal ridges of

most trunk and tail rings, additional flaps sometimes occur between ridges; flaps usually slender and simple, sometimes spatulate, frilled or with a few branches. Fins. Length of dorsal-fin base 1.2-1.9 (1.6) in hi, pectoral-fin length 5.3-9.0 (7.1) in hi, length of pectoral-fin base 1.2-1.8 (1.4) in pectoral-fin length. Pectoral-fin base not protruding strongly laterad, with two distinct ridges and usually with one or more dermal flaps. Color. Without distinctive markings (Figs. 113-116), usually tan to greenish brown. Side and dorsum of snout plain brownish, mottled or spotted, sometimes with a distinct brown stripe to orbit; dorsum and sides of head usually mottled, darkest above and shading to pale ventrad; eye usually with irregular radiating blotches of pale and brown. Dorsum and sides of body usually mottled, sometimes with indications of dif-

TABLE LV. Frequency distributions of subdorsal trunk rings in western Atlantic species of Bryx and Cosmocampus. *: Genus Species

Subdorsal trunk rings

2.75

2.50

2.25

2.00

1.75

1.50

1.25

Bryx dunckeri randalli Cosmocampus albirostris profundus brachycephalus elucens hildebrandi * Primary type.

1 3

8

12

113*

1.00

0.75

0.50

0.25

0.00

1 2

2 3

9 6*

16

128* 4

6 1 1 56 2

7 2* 1 31 8*

3

2

27 12

5 6

2

7

21

36

66

126 2

25 7

3 75 4

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Memoir I, Part 8—Sears Foundation for Marine Research

TABLE LVI. Frequency distributions of subdorsal tail rings in western Atlantic species of Bryx and Cosmocampus. Genus Species

Subdorsal tail rings

2.25

2.50

2.75

3.00

3.25

3.50

3.75

4.00

4.25

Bryx dunckeri randalli Cosmocampus albirostris profundus brachycephalus elucens hildebrandi

2

9

24

63

61

4.50

4.75

2*

1

8

4

25

17

24

5

125* 3

44 8

22 32

4 67

2 52

2 18

1

3

10

4

* Primary type.

TABLE LVI, Continued. Subdorsal tail rings

Genus Species

5.00

5.25

5.50

Bryx dunckeri randalli

1 3

1

8 4

Cosmocampus albirostris profundus brachycephalus elucens hildebrandi

1

16 8

2

1*

7 8*

1

5.75

6.00

6.25

6.50

6.75

7.00

7.25

7.50

6

30 2

37*

69 2

60

44

11

4

1

* Primary type.

fuse bars; some fish pale on lower half of sides of trunk, some are mottled, but others may have rather distinct brownish bars encircling scutella; venter usually pale or brownish, sometimes with indications of a brown bar on each ring. Dorsal and pectoral fins typically hyaline, caudal fin sometimes shaded with brown. Planktonic young (ca. 21-46 mm) encircled by about 10 broad brown bands separated by diffuse pale interspaces (ca. 0.5-1 ring wide); caudal fin brown, other fins hyaline. Size. Largest specimen examined is 164 mm; probably fails to exceed 175 mm. Development. Present material includes brooding males collected during all months except February, March, June and November. The smallest male with developing brood pouch was 68.5 mm; the pouch was developed below 15-20 rings in 37 brooding fish (102-155.5 mm). Pouch eggs are usually deposited in a single layer and in up to ca. 10 transverse rows. A 120-mm male

with ca. 9 rows had 58 eggs in the outer right row through 17 rings of a 19-ring pouch, and a 155-mm fish with ca. 10 rows had 66 eggs in the outer row. Pouch closure apparently variable, everted in most but clearly the overlapping or semi closure in others (Fig. 3). Planktonic juveniles (ca. 21-45 mm) have minute denticulations on the principal ridges but the protruding spine-like processes occurring in juveniles of some species of Syngnathus (e.g. louisianae) are evidently absent in Cosmocampus elucens. Parasites. Manter (1947) reported this species as host for the trematode Genitocotyl atlantica Manter. Nahhas and Cable (1964) failed to find trematodes in eleven specimens examined from Jamaica. Variation. Materials examined show no evidence of significant geographic variation in meristic characters or coloration. Nine specimens (87.5-115 mm) trawled off

Fishes of the Western North Atlantic

137

FIGURE 46. Cosmocampus elucens: female (UMML 23933), 107 mm.

Panama in 47-79 m at R/V PILLSBURY Stas. 392, 425, 433 and 434 (09°14'36"-09°45'06"N, 79°09'06"-80°13'30"W) exhibit atypical development of dermal flaps. In other material, flaps are commonly short and there is usually a single flap on each principal ridge of most rings (Fig. 45). In these Panamanian specimens, flaps are exceptionally long and slender (Fig. 46), they often occur between the principal ridges and

there may be as many as five on the inferior ridge of a single ring. These specimens are all bleached in preservative (Fig. 116) but they agree with C. elucens in all other characters and there is no ready explanation for this atypical development of dermal flaps. Herald (1943, 1959) referred C. elucens to Duncker's (1915) nominal subgenus Parasyngnathus due to apparent sharing of the everted

TABLE LVII. Frequency distributions of total subdorsal rings in western Atlantic species of Bryx and Cosmocampus. Genus Species

Total subdorsal rings

4.25

4.50

4.75

Bryx dunckeri rand alii Cosmocampus albirostris profundus brachycephalus elucens hildebrandi * Primary type.

9 3

5.00

5.25

5.50

1 1*

2

9 2

3 1

29 4

6 1 20 4 2*

1 2* 9 1

3

7

19

30

19

25 4 1

61 22 4

100 68 9

73 75 9

60* 26 10

5.75

6.00

6.25 6.50

6.75

7.00

66 3

63

48

35* 2

1

7.25 7.50 12

4

138

Memoir 1, Part 8—Sears Foundation for Marine Research

FIGURE 47. Cosmocampus hildebrandi: holotype (USNM 117251), female, 86 mm.

pouch closure with the type-species P. argyrostictus (Kaup). Present observations indicate that pouch closure may be variable in Cosmocampus elucens (see Development) and that this species cannot be referred to Duncker's Indo-Pacific taxon. This Atlantic species further differs from Parasyngnathus sensu stricto in lacking a complete opercular ridge in adults (present in argyrostictus) and in the presence of dermal flaps (absent in argyrostictus).

Distribution. The species is known (Fig. 43) from the offings of New Jersey (39°04'N), Bermuda, Bahamas, the Antilles, SE Florida (Monroe Co.)/ the eastern Gulf of Mexico and from western Caribbean and Atlantic localities south to Cabo do Norte (ca. 01°40'N) and the state of Rio Grande do Sul, Brazil (Figueiredo and Menezes, 1980). Literature and present data indicate that Cosmocampus elucens occurs over a broad spectrum

Fishes of the Western North Atlantic of habitats. Longley and Hildebrand (1941) found it common in turtle grass (Thalassia). Cervigon (1975) reported it from coral (Acropora) and Thalassia bottom and noted that the species did not occur in areas of mud bottoms or turbid water. Eskinazi (1972a) records C. elucens from estuaries and there are collections from areas of mangrove, "mud and grass" and sandy beaches. There appears little doubt that the most common inshore habitat is among algae or Thalassia. Juveniles and subadults (ca. 21-120 mm) are recorded from surface nightlight, nekton or neuston collections at distances to 100 km or more offshore but there are evidently no records of direct association with floating Sargassum. Fahay's (1975) records of this species are based on misidentifications of Syngnathus springeri and S. pelagicus. Present materials include several dredge collections in depths of 51-171.9 m but most specimens are from depths of less than 5 m. Dawson (1972) reported collections at a temperature of 26.4° C; recorded salinities range from 29,4-45%o (Dawson, 1972; Eskinazi, 1972a). Study Material. 214 specimens, 21-164 mm SL. BERMUDA: AMNH 16239 (1, 48); BOC 7777 (3, 53-86); FMNH 5338 (1, ca. 105); FMNH 5344 (1, 152); FMNH 48697 (1, 68); UMMZ 175977 (1, 108.5); UMMZ 176371 (1, 98); USNM 170087 (1, 32.5); USNM 170914 (112, holotype of Syngnathus pipulus); USNM 217338 (1, 132). BAHAMAS: AMNH 23988 (1, ca. 27); AMNH 28687 (1, 73); AMNH 31290 (3, 102.5-132); ANSP 94574 (1, 41.5); ANSP 94575 (1, 128.5); ANSP 94576 (1, 34); ANSP 94577 (10, 49.5-102.5); ANSP 94578 (1, 39.5); ANSP 94579 (2, 30.5-36.5); ANSP 94580 (1, 31.5); ANSP 97986 (1, 99.5); ANSP 124059 (2, 100-103.5); BOC 89 (1, 104); CAS 31178 (1, 39); CAS 31179 (1, 38); FSM 14002 (1, 73.5); UMML 33194 (1, 84); UMML 33195 (1, 115.5); UMML 33196 (1, 39.5). GREATER ANTILLES, Cayman Is.: FSM 24575 (1, 54). Jamaica: CAS(SU) 4988 (113.5, holotype of Siphostoma robertsi); FMNH 83836 (1, 147). Hispaniola: AMNH 15056 (1, 71); CAS 31180 (1, 74). Puerto Rico: FMNH 74790 (1, 83); FSM 12052 (1, 148.5); LACM 677M (1, 122); UMML 1964 (1, 85); UPR 3260 (1, 145); UPR 3263 (1, 115); UPR 3265 (1, 92); USNM 127137 (1, ca. 155.5). LESSER ANTILLES, Virgin Is.: AMNH 13685 (144, holotype of Syngnathus walcotti); BMNH 1976.7.14.68 (1, 95); GCRL 15745 (3, 103-120); GCRL 15746 (1, 101); GCRL 15747 (2, 102-107.5); GCRL 15748 (1, 99); GCRL 15749 (1, 114.5); GCRL 15750 (1, 122); USNM 178047 (1, 141.5); USNM 217337 (1, 95). St.

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Barthelemy: ANSP 121860 (2, 62-74); ANSP 121880 (2, 32-66). Dominica: CAS 31181 (1, 120); USNM 217339 (1, 164). Martinique: ANSP 113001 (1, 71.5); ANSP 130819 (1, 82). Barbados: MCZ 28044 (1, ca. 115). Curasao: USNM 85475 (1, ca. 122); ZMA 114.117 (1, 112). Aruba: ZMA 114.116 (3, 87.5-99.5). UNITED STATES, off New Jersey (ca. 39°04'N): GCRL 14446 (1, 46.5). East Florida: FSM 4154 (2, 117-136.5); FSM 11807 (1, 130.5); FSM 11826 (1, 56.5); MCZ 26202 (1, 156.5); UMMZ 87982 (2, 128-132); USNM 68575 (1, 131.5); USNM 117241 (47, ca. 92-149); USNM 117250 (7, 117-145); USNM 117252 (2, ca. 129-132); USNM 119118 (1, ca. 138); USNM 217336 (2, 73.5-104). West Florida: CAS 31200 (1, 120); FDNR 6521 (1, 36.5); FDNR 5422 (1, ca. 61); FDNR 6524 (1, 68.5). Mississippi: GCRL 4269 (1, 21). BELIZE: FMNH 83847 (1, 146); FMNH 83848 (1, 130). NICARAGUA: BMNH 1969.6.26.535 (1, 93.5); CAS 31191 (1, 98). COSTA RICA: UCR 387-30 (1, 157). PROVIDENCIA IS.: FSM 24379 (1, 134); FSM 24435 (1, 85.5); FSM 24471 (1, 129.5). PANAMA: GCRL 10127 (1, 68); GCRL 11163 (1, 121); GCRL 15673 (1, 103); GCRL 15764 (1, 107); UMML 23933 (3,103-115); UMML 26594 (1,87.5); UMML 26662 (1, 100); UMML 26735 (2, 94-100). COLOMBIA: CAS 31190 (1, 116); GCRL uncat. (1, 121); UMML 29173 (1, 95.5); USNM 213596 (1, 119). VENEZUELA: FLSM 138 (1,123); MBUCV V.4956 (1,155); UDONECI687 (2,150); UDONECI 834 (2, 142-145); UDONECI 878 (1, 108); UDONECI 922 (3, 103-120); UDONECI 976 (8, 96-145); UDONECI 977 (1, 137); UOIO 67-060 (2, 54-120). GUYANA: RMNH 26982 (3, 92-106). SURINAM: CAS 31201 (1,150); CAS 31202 (1,107). BRAZIL, Amapa: UFP uncat. (1, 88). LOG UNCERTAIN: BMNH 1847.5.14.39 (1, 147.5), "S. America, purchased" and BMNH 1860.7.18.63 (1, ca. 138), "New Hebrides, Aneitum," syntypes of Syngnathus modestus; BMNH uncat (1, 145), "purchased from Mr. Damon"; CAS 39651 (1, 90), GILL Cr. 3, 25 July 1953; RMNH 9320-21 (3, 100-120), Curacao and St. Martin.

Cosmocampus hildebrandi (Herald) 1965 Dwarf pipefish Figures 43, 47, 117, 118 Syngnathus elucens (not of Poey). Longley and Hildebrand, 1941:61 (misident, in part; Tortugas, Florida). Syngnathus sp. Briggs, 1958:257 (Hildebrand's pipefish; eastern Gulf of Mexico). Syngnathus hildebrandi Herald, 1965:364,366 (orig. descr; dwarf pipefish; Tortugas, Florida; holotype, USNM 117251); Bailey et aL, 1970:34 (compiled); Powell et al., 1972:60 (listed); Walls, 1975:160, 162, fig. (descr.; West Florida); Fritzsche, 1976:43, 46 (referred to MS subgenus of Bryx); Shiino, 1976:110 (compiled). Bryx (Simocampus) hildebrandi. Fritzsche, 1980:192 (n. comb.).

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Distinctive Characters. The combination of short snout (snout length averages 2.8 in hi), 19-22 dorsal-fin rays and modal counts of 11 pectoral-fin rays and 33 tail rings distinguishes C. hildebrandi from western Atlantic congeners. Description. Proportional data based on 20 specimens from Florida, 29-86 (x = 59.0) mm. Rings: 16-17 + 31-35 = 48-52, modally 17 + 33 (Tables LII, LIII). Subdorsal rings: 1.0-0.0 + 4.0-5.5 = 4.5-5.75 (Tables LV, LVI, LVII). Fin rays: dorsal 19-22, usually (94%) 19-21; pectoral 10-13, modally 11 (Table LIV). Head. Head length 7.0-11.0 (x = 9.6) in SL, snout length 2.6-3.2 (2.8) in hi, snout depth 2.2-3.6 (2.8) in snout length. Median dorsal snout ridge somewhat elevated along posterior l 2 /2- /3 of snout; supraorbital ridges elevated, flared laterad, with interorbital concave between; opercular ridge angled somewhat dorsad, crosses %-Vi of opercle in adults, reaches to or near upper posterior angle in 20-mm juvenile; opercle ornamented elsewhere with radiating low striae; supraopercular ridge present, often prominent; simple, slender to spatulate, dermal flaps present or absent on eye, anteroventral part of snout, and most head ridges (Fig. 47). Body. Depth at anal ring 2.5-3.2 (2.9) in hi, trunk depth 2.0-2.6 (2.4) in hi. Principal ridges somewhat elevated, with sides and dorsum depressed or concave between; ridge margins entire to minutely denticulate, indented to somewhat notched between rings; posterior tail rings often with posterior angles produced to points; venter of trunk V-shaped; trunk not greatly deepened in mature females (Fig. 117); simple dermal flaps present or absent on principal ridges of most rings. Fins. Length of dorsal-fin base 0.9-1.2 (1.1) in hi, pectoral-fin length 4.2-7.1 (4.8) in hi, length of pectoral-fin base 1.3-2.8 (1.9) in hi. Pectoralfin base protruding slightly laterad, with two ridges, often with one or more dermal flaps. Color. Most specimens completely faded or bleached in preservative. Two fish (43-50.5 mm)

retain a light tan ground color, sides and dorsum are irregularly shaded with faint brownish microchromatophores, the venter is mainly pale with fine dark stripe on midline and all fins are hyaline. A 20-mm juvenile is largely brown with 9-10 diffuse pale bars (0.5-1.0 ring wide) crossing dorsum and upper part of sides, the fins are hyaline. Life color is largely red (R. L. Shipp, pers. comm.) and the head and body of one fish (GCRL 15966) retained some shading of pink for some days following preservation. Size. This is a small species which probably fails to reach 100 mm; largest known specimen is 86 mm. Development. Among examined material, the smallest male with developing pouch is 44 mm; the pouch extends below 13-18 rings in five brooding fish (50.5-74 mm); pouch plates are angled somewhat laterad; pouch closure is the semi type (Fig. 3). Brood-pouch eggs are arranged in a single layer of 2-4 transverse rows. A 50.5-mm fish had a total of ca. 38 eggs in two rows throughout a 16-ring pouch, whereas another (65 mm) had four rows of eggs through 12 rings of a 16-ring pouch and there were 20 eggs in the outer right row. Variation. Available material shows no significant geographic variation in examined characters. There are differences in the development or persistence of dermal flaps but these are considered to reflect individual variation or loss in the course of collection and preservation. The anal fin may be concealed within the anterior portion of the brood pouch in mature males and careful examination under X30 or greater magnification may be required to verify its presence. Distribution. The species is presently known from North Carolina (34°19.5'N, 76°53.5'W) and SE Florida (Monroe Co.) on the Atlantic seaboard and from 26°24' to ca. 30°N off W Florida (Fig. 43). Among examined material, there are two SCUBA collections in 4.6-7.6 and 27.4 m and 24 lots from dredge or trawl samples in depths of

Fishes of the Western North Atlantic 13.3-73.2 m. Available data indicate collections over substrates of coral, rock and sand. Study Material. 42 specimens, 20-86 mm SL, including holotype and 5 paratypes. Holotype. USNM 117251 (86 mm, female), Dry Tortugas, Monroe Co., Florida, W. H. Longley. Paratypes. West Florida: USNM 73235 (73.5 mm, female), off Pepperfish Key, 29°13'15"N, 83°32'3(TW, rocky bottom, 13.3 m, dredge, 21 Nov. 1901, FISH HAWK Sta. 7165. USNM 134312 (59.5 mm, female), St. Martin's Key, 28°34'30"N, 83°15'45"W, 13.7 m, 15 Jan. 1902, FISH HAWK Sta. 7220. USNM 109826 (ca. 67 mm, female), St. Martin's Reef, 28°27WN, 83°13'00"W, rock and sand bottom, 20.1 m, 15 Jan. 1902, FISH HAWK Sta. 7217. CAS 24294 (1, 61.5 mm, female) and USNM 73239 (1, 54 mm, female), St. Martin's Reef, 28°26'30"N, 83°08WW, 18.3 m, dredge, 15 Jan. 1902, FISH HAWK Sta. 7216. Other Material UNITED STATES, North Carolina: USNM 219600 (1, 57). East Florida: UMML 19678 (1, ca. 74.5). West Florida: FDNR 6508 (2, 52-60); FDNR 6509 (1, 71); FDNR 6510 (1, 34); FDNR 6511 (1, ca. 52); FDNR 6512 (1, 57); FDNR 6513 (1, 52); FDNR 6514 (1, 31.5); FDNR 6515 (1, 63); FDNR 6516 (2, 51-55); FDNR 6517 (1, 61.5); FDNR 6518 (2, 24.9-28.9); FDNR 6519 (3, 22.2-56); FDNR 6525 (1, 60); GCRL 13552 (1, 74); GCRL 15966 (1, 50.5); GCRL 16071 (1, 20); GCRL 16072 (1, 43); USA 4726 (5, 37-44); USA 4796 (3, 51-65); USA 5376 (1, 32); USA 5377 (2, 23.5-36); USA 5396 (1, 26).

LITERATURE CITED ABBOTT, J. F. 1899. The marine fishes of Peru. Proceedings of the Academy of Natural Sciences of Philadelphia, 51:324-364. ADAMS, J. A. 1960. A contribution to the biology and postlarval development of the Sargassum fish, Histrio histrio (Linnaeus), with a discussion of the Sargassum complex. Bulletin of Marine Science of the Gulf and Caribbean, 10(l):55-82. ALPERIN, I. M., AND R. H. SCHAEFER. 1964. Juvenile bonefish (Albula vulpes) in Great South Bay, New York. New York Fish and Game Journal, 11(1):1-12. ALVAREZ DEL VILLAR, J. 1970. Peces mexicanos (claves). Secretaria de Industria y Comercio, Mexico. Serie Investigacion Pesquera, Estudio, 1:1-166. ANDERSON, E. 1964. Cytologic changes during oocyte differentiation and formation of the vitelline envelope in certain teleost fish. Journal of Cell Biology, 23:4A. -. 1968. Cortical alveoli formation and vitellogenesis during oocyte differentiation in the pipefish, Syngnathus fuscus, and the killifish, Fundulus heteroclitus. Journal of Morphology, 125(1):23-31.

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ANDERSON,W.W.,ANDJ.W.GEHRINGER. 1958. Physical oceanographic, biological, and chemical data . . . South Atlantic coast of the United States . . . GILL Cruise 5. United States Fish and Wildlife Service, Special Scientific Report (Fisheries), 248:1-220. ANONYMOUS. 1976. Catalogo de peces marinos Mexicanos. Institute Nacional de Pesca, Mexico, D.F.: xvi + 462 pp. AUSTIN, H. M. 1971. A survey of the ichthyofauna of the mangroves of western Puerto Rico during December, 1967-August, 1968. Caribbean Journal of Science, ll(l-2):27-39. AYRES, W. O. 1843. Enumeration of the fishes of Brookhaven, Long Island, with remarks upon the species observed. Boston Journal of Natural History, 4(3):265-292. BACKUS, R. H., J. E. CRADDOCK, R. L. HAEDRICH, AND D. L. SHORES. 1969. Mesopelagic fishes and thermal fronts in the western Sargasso Sea. Marine Biology, 3(2):87-106. BAHAMONDE N., N., AND G. PEQUENO R. 1975. Peces de Chile—Lista sistematica. Publicacion Ocasional, Museo Nacional de Historia Natural, Santiago, 21:1-20. BAILEY, R. M., H. W. WINN, AND C. L. SMITH. 1954. Fishes from the Escambia River, Alabama and Florida, with ecological and taxonomic notes. Proceedings of the Academy of Natural Sciences of Philadelphia, 106:109-164. BAILEY, R. M., E. A. LACHNER, C. C. LINDSEY, C. R. ROBINS, P. M. ROEDEL, W. B. SCOTT, AND L. P. WOODS. 1960. A list of common and scientific names of fishes from the United States and Canada. 2nd edn. American Fisheries Society Special Publication, 2:1-102. BAILEY, R. M., J. E. FITCH, E. S. HERALD, E. A. LACHNER, C. C. LINDSEY, C. R. ROBINS, AND W. B. SCOTT. 1970. A list of common and scientific names of fishes from the United States and Canada. 3rd edn. American Fisheries Society Special Publication, 6:1-150. BAISSAC, J. DE B. 1956. Contribution a 1'etude des poissons de Tile Maurice. VI. Proceedings of the Royal Society of Arts & Sciences of Mauritius, l(4):319-365. BANKS, J. 1768-1771. The ENDEAVOUR journal of Joseph Banks. J. C. Beaglehole, ed. Trustees of the Public Library of New South Wales. 2nd edn., 1963. Angus and Robertson, Sydney, 2:xvi + 406 pp. BARBOUR, T. 1905. Notes on Bermudian fishes. Bulletin of the Museum of Comparative Zoology, Harvard College, 46(7):109-134. BARNARD, K. H. 1927. A monograph of the marine fishes of South Africa. Annals of the South African Museum, 21:1-418.

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BAUGHMAN, J. L. 1950. Random notes on Texas fishes. Part 1. Texas Journal of Science, 2(1):117-138. BEAN, B. A. 1891. Fishes collected by William P. Seal in Chesapeake Bay, at Cape Charles City, Virginia, September 16 to October 3,1890. Proceedings of the United States National Museum, 14:83-94. . 1905. Fishes of the Bahama Islands (pp. 291-325), in The Bahama Islands, G. B. Shattuck, ed. Geographical Society of Baltimore, Johns Hopkins Press: xxxii -I- 630 pp. BEAN, T. H. 1880. A list of European fishes in the collection of the United States National Museum. Proceedings of the United States National Museum, 2:10-44. . 1881. Check-list of duplicates of North American fishes distributed by the Smithsonian Institution in behalf of the United States National Museum, 1877-1880. Proceedings of the United States National Museum, 3:75-116. . 1883. Catalogue of the collection of fishes exhibited by the United States National Museum. Bulletin of the United States National Museum, 27:387-510. . 1888. Report on the fishes observed in Great Egg Harbor Bay, New Jersey, during the summer of 1887. Bulletin of the United States Fish Commission, 7:129-152. . 1890. Notes on fishes collected at Cozumel, Yucatan, by the U.S. Fish Commission, with descriptions of new species. Bulletin of the United States Fish Commission, 8:193-206. . 1897. Notes upon New York fishes received at the New York Aquarium, 1895-1897. Bulletin of the American Museum of Natural History, 9(24):327-375. . 1903. Catalogue of the fishes of New York. Bulletin of the New York State Museum, 60:1-784. . 1906. A catalogue of the fishes of Bermuda, with notes on a collection made in 1905 for the Field Museum. Field Columbian Museum Publication 108, Zoological Series, 7(2):21-89. BEAN, T. H., AND H. G. DRESEL. 1884. Diagnoses of three new species of fishes from the Gulf of Mexico. Proceedings of the Biological Society of Washington, 2:99-100. BEARDEN, C. M. 1961. Common marine fishes of South Carolina. Contributions from Bears Bluff Laboratories, 34:1-47. BEEBE, W., AND G. HOLLISTER. 1935. The fishes of Union Island, Grenadines, British West Indies, with the description of a new species of star-gazer. Zoologica, New York, 19(6):209-224. BEEBE, W., AND J. TEE-VAN. 1928. The fishes of Port-auPrince Bay, Haiti, with a summary of the known

species of marine fish of the island of Haiti and Santo Domingo. Zoologica, New York, 10(l):l-279. . 1932. New Bermuda fish, including six new species and forty-three species hitherto unrecorded from Bermuda. Zoologica, New York, 13(5):109-120. . 1933. Field book of the shore fishes of Bermuda. G. P. Putnam's Sons, New York: xiv + 337 PPBEHRE, E. H. 1933. Color recognition and color changes in certain species of fishes. Copeia, 1933(2):49-58. . 1950. Annotated list of the fauna of the Grand Isle region—1928-1946. Occasional Papers of the Marine Laboratory, Louisiana State University, 6:1-66. BERG, C. 1895. Enumeracion sistematica y sinonimica de los peces de las costas Argentina y Uruguaya. Anales del Museo Nacional de Buenos Aires, Serie 2, 4:1-120. BERTIN, L., AND R. ESTEVE. 1950. Catalogue des types de poissons du Museum National d'Histoire Naturelle. 6e partie. Haplomes, Heteromes, Catosteomes. Imprimerie Nationale, Paris, 1-60. BIGELOW, H. B., AND W. C. ScHROEDER. 1936. Supplemental notes on fishes of the Gulf of Maine. Bulletin of the United States Bureau of Fisheries, 48:319-343. . 1953. Fishes of the Gulf of Maine. Fishery Bulletin, 53(74):vii + 577 pp. BIGELOW, H. B., AND W. W. WELSH. 1925. Fishes of the Gulf of Maine. Bulletin of the United States Bureau of Fisheries, 40(1): 1-567. BINI, G. 1970. Atlante dei pesci delle coste Italiane. 3 Osteitti. Mondo Sommerso Editrice: 232 pp. BLACHE, J., J. CADENAT, AND A. STAUCH. 1970. Cles de determination des poissons de mer signales dans 1'Atlantique oriental (entre le 20° parallele N. et le 15° parallele S.). Faune Tropicale, O. R. S. T. O. M., 18:1-479. BLEAKNEY, J. S., AND D. E. MCALLISTER. 1973. Fishes stranded during extreme low tides in Minas Basin, Nova Scotia. Canadian Field-Naturalist, 87:371-376. BLEEKER, P. 1849. Bijdrage tot de kennis der ichthyologische fauna van het eiland Madura, met beschrijving van eenige nieuwe soorten. Verhandelingen van het Bataviaasch genootschap van kunsten en wetenschappen, 22(8):1-16. . 1853. Bijdrage tot de kennis der Troskieuwige visschen van den Indischen Archipel. Verhandelingen van het Bataviaasch genootschap van kunsten en wetenschappen, 25(6):1-30. . 1854-57. Index specierum piscium in voluminibus XXI, XXII, XXIII, XXIV, XXV et XXVI Actorum Societatis Artium et Scientiarum Bataviensis descriptarum adjectis citacionibus ubi descriptiones

Fishes of the Western North Atlantic Bleekerianae recentiores emendataeque reperiuntur. Verhandelingen van het Bataviaasch genootschap van kunsten en wetenschappen, 26(6):l-24. -. 1860a. Over eenige vischoorten van de Kaap de Goede Hoop. Natuurkundig Tijdschrift voor Nederlandsch Indie, 21:49-80. --. 1860b. Vischsoorten. . . . in de zogenaamde Krooszee gevangen. Natuurkundig Tijdschrift voor Nederlandsch Indie, 21:331. --. 1860c. Achtste bijdrage totde kennis der vischfauna van Sumatra (Visschen van Benkoelen, Priaman, Tandjong, Palembang en Djambi). Actorum Societatis Scientiarum Indo-Neerlandicae, 8:1-88. --. 1873. Addition au memoire sur la faune ichthyologique de Chine. Nederlandsch Tijdschrift voor de Dierkunde, 4:233-234. -. 1878. Contribution a la faune ichthyologique de Tile Maurice. Wis — en Natuurkundige Verhandelingen der Koninkijke Akademie van Wetenschappen te Amsterdam, 18:1-23. BLOCK, M. E. 1785. Naturgeschichte der auslandischen fische. Erster theil. Berlin: 136 pp. BOESEMAN, M. 1960a. The fresh-water fishes of the island of Trinidad. Studies on the Fauna of Curasao and other Caribbean Islands, 10(48):72-153. --. 1960b. Notes on marine fishes from the Netherlands Antilles, with the description of a new species, Eutyx tumidifrons (Brotulidae). Zoologische Mededelingen, 37(2):11-16. BOHLKE, J. E. 1953. A catalogue of the type specimens of recent fishes in the Natural History Museum of Stanford University. Stanford Ichthyological Bulletin, 5(1):1-168. BOHLKE, J. E., AND C. C. G. CHAPLIN. 1968. Fishes of the Bahamas and adjacent tropical waters. Livingston, Wynnewood, Pa.: xxiii + 771 pp. BONNATERRE, M. L. 1788. Tableau encyclopedique et methodique des trois regnes de la nature . . . Ichthyologie. Paris: Ivi + 215 pp. BORODIN, N. A. 1928. Scientific results of the yacht ARA Expedition during the years 1926 to 1928, while in command of William K. Vanderbilt. Fishes. Bulletin of the Vanderbilt Marine Museum, BOSCHI, E. E., AND J. L. FENUCCI. 1972. Contribucion al conocimiento de la fauna marina del Golf o San Jose. Physis, Buenos Aires, 31(82):155-167. BOSCHUNG, H. T. 1957. The fishes of Mobile Bay and the Gulf coast of Alabama. Ph.D. thesis, University of Alabama, Montgomery. BOULENGER, G. A. 1904. Fishes, systematic account of Teleostei. The Cambridge Natural History, London, 7:539-727. --. 1912. Poissons recueillis dans la region du Bas-

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Congo par M. le Dr. W. J. Ansorge. Annales du Musee du Congo Beige. Zoologie. Serie 1, 2(3):l-27. BOUSFIELD, E. L., AND A. H. LEiM. 1960. The fauna of Minas Basin and Minas Channel. Bulletin. National Museum of Canada, 166:1-30. BOYLE, R. H. 1968. Notes on fishes of the lower Hudson River. Underwater Naturalist, 5(2):32-33. BRANCH, G. M. 1966. Contributions to the functional morphology of fishes. Part III. The feeding mechanism of Syngnathus acus Linnaeus. Zoologica Africana, 2(l):69-89. BRAVO-NUNEZ, E., AND A. YANEZ-ARANCIBIA. 1979. Ecologia en la boca de Puerto Real, Laguna de Terminos I. Description del area y analisis estructural de las comunidades de peces. Anales Centre Ciencias del Mar y Limnologia. Universidad Nacional Autonoma de Mexico, 6(1):125-182. BREDER, C. M., JR. 1924. Nineteen twenty three fish notes from Sandy Hook Bay. Copeia, 1924(127): 25-29. -. 1927a. Scientific results of the first oceanographic expedition of the "PAWNEE," 1925. Fishes. Bulletin of the Bingham Oceanographic Collection, ---

. 1927b. The fishes of the Rio Chacunaque drainage, eastern Panama. Bulletin of the American Museum of Natural History, 57(3):91-176. --. 1929. Field book of marine fishes of the Atlantic coast from Labrador to Texas. G. P. Putnam's Sons, New York: xxxviii + 332 pp. --. 1932. Fish notes for 1931 and 1932 from Sandy Hook Bay. Copeia, 1932(4): 180. --. 1933. The fishes of Barro Colorado, Gatun Lake, Panama. Zoologica, New York, 9(16):565-572. . 1934. The oceanographic vessel ATLANTIS in the West Indies. Bulletin of the New York Zoological Society, 37(2):31-42. . 1938. The species of fish in New York harbor. Bulletin of the New York Zoological Society, 41(l):23-29. . 1946. An analysis of the deceptive resemblances of fishes to plant parts, with critical remarks on protective coloration, mimicry and adaptation. Bulletin of the Bingham Oceanographic Collection, 10(2):l-49. BREDER, C. M., JR., AND R. F. NIGRELLI. 1934. Fish notes for 1933 and 1934 from Sandy Hook Bay and other New York localities. Copeia, 1934(4): 193-194. BREDER, C. M., JR., AND D. E. ROSEN. 1966. Modes of reproduction in fishes. Natural History Press, Garden City, New York: xv + 941 pp. BRIGGS, J. C. 1958. A list of Florida fishes and their distribution. Bulletin of the Florida State Museum, Biological Sciences, 2(8):224-318.

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. 1974. Marine zoogeography. McGraw-Hill, New York: xi -I- 475 pp. BROOK, I. M. 1977. Trophic relationships in a seagrass community (Thalassia testudinum), in Card Sound, Florida. Fish diets in relation to macrobenthic and cryptic faunal abundance. Transactions of the American Fisheries Society, 106(3):219-229. BROWN, J. D. 1972. A comparative life history study of four species of pipefishes (Family Syngnathidae) in Florida. Ph.D. thesis, University of Florida, Gainesville. BROWNELL, R. L., JR., J. OLAZARRI, AND F. ACHAVAL. 1973. Marine mammal and bird observations and trawling off Uruguay: R/V HERO Cruise 72-3a. Antarctic Journal of the United States, 8(1):9-10. BULLIS, H. R., JR., AND J. R. THOMPSON. 1965. Collections by the exploratory fishing vessels OREGON, SILVER BAY, COMBAT and PELICAN made during 1956-1960 in the southwestern North Atlantic. United States Fish and Wildlife Service, Special Scientific Report (Fisheries), 510:1-130. BURGESS, G. H., G. W. LINK, JR., AND S. W. Ross. 1980. Additional marine fishes new or rare to Carolina waters. Northeast Gulf Science, 3(2):74-87. BURKENROAD, M. D. 1931. Notes on the sound-producing marine fishes of Louisiana. Copeia, 1931(l):20-28. BUSSING, W. A. 1967. New species and new records of Costa Rican freshwater fishes with a tentative list of species. Revista de Biologia Tropicale, San Jose, 14(2):205-249. C ADENAT, J. 1950. Initiations Africaines. III. Poissons de mer du Senegal. Institut Francais d'Afrique Noire, Dakar: 345 pp. CALDWELL, D. K. 1957. The biology and systematics of the pinfish, La god on rhomboides (Linnaeus). Bulletin of the Florida State Museum, Biological Sciences, 2(6):77-173. . 1966. Marine and freshwater fishes of Jamaica. Bulletin of the Institute of Jamaica. Science Series, 17:1-120. CALDWELL, D. K., AND M. C. CALDWELL. 1964. Fishes from the southern Caribbean collected by VELERO III in 1939. Allan Hancock Atlantic Expedition Report, 10:1-61. CALDWELL, D. K., L. H. OGREN, AND L. GIOVANNOLI. 1959. Systematic and ecological notes on some fishes collected in the vicinity of Tortuguero, Caribbean coast of Costa Rica. Revista Biologia Tropicale, San Jose, 7(l):7-33. CARCASSON, R. H. 1977. A field guide to the coral reef fishes of the Indian and West Pacific Oceans. Wm. Collins Sons, London: 320 pp. CARR, A. F., JR. 1936. A key to the fresh-water fishes of

Florida. Proceedings of the Florida Academy of Sciences, 1:72-86. CARR, A. F., JR., AND C. J. COIN. 1959. Guide to the reptiles, amphibians and fresh-water fishes of Florida. University of Florida Press, Gainesville: 341 pp. CARUS, J. V. 1893. Prodromus faunae Mediterraneae, . . . Vertebrata. 1. Class. Pisces. Stuttgart: 2:ix -I- 854 PPCASTELNAU, F. L. 1855. Animaux nouveaux ou rares requeillis pendant 1'expedition dans les parties centrales de 1'Amerique du Sud, de Rio de Janeiro a Lima, et de Lima au Para; executee par ordre du Gouvernement Francais pendant les annees 1843 a 1847 ____ Poissons. C. P. Bertrand, Paris: 3:xii -I- 112. CERVIGON, M. F. 1966a. Los peces marines de Venezuela. Tomo II. Monografia de la Fundacion La Salle de Ciencias Naturales, Caracas, 12:449-951. --. 1966b. Los peces marines de Venezuela. Tomo I. Monografia de la Fundacion La Salle de Ciencias Naturales, Caracas, 11:1-436. --. 1975. Los peces marinos de Venezuela. Complemento IV. Contribuciones Cientificas, Universidad de Oriente, Porlamar, 5:1-45. CHABANAUD, P., AND T. MONOD. 1926. Les Poissons de Port-Etienne. Bulletin du Comite d'Etudes Historiques et Scientifiques de 1'Afrique Occidentale Francaise, 9(2):225-287. CHENOWETH, S. B. 1973. Fish larvae of the estuaries and coast of central Maine. Fishery Bulletin, CHICKERING, A. M. 1930. An Atlantic pipefish caught in transit through the Panama Canal. Copeia, 1930(173):85-86. CHIRICHIGNO F., N. 1974. Clave para identificar los peces marinos del Peru. Informe del Institute del Mar del Peru, Callao, 44:1-387. CHRISTENSEN, F. R. 1965. An ichthyological survey of Jupiter Inlet and Loxahatchee River, Florida. M.Sc. thesis, Florida State University, Tallahassee. CHRISTMAS, J. Y., AND R. S. WALLER. 1973. Estuarine vertebrates, Mississippi. In Cooperative Gulf of Mexico estuarine inventory and study, Mississippi (pp. 320-406). Gulf Coast Research Laboratory: 434 PPCHU, Y. T. 1931. Index piscium Sinensium. Biological Bulletin of St. John's University, Shanghai, l:iv + 290pp. CLARK, J. E., W. G. SMITH, A. W. KENDALL, JR., AND M. FAHAY. 1969. Studies of estuarine dependence of Atlantic coastal fishes. Data report I: Northern section, Cape Cod to Cape Lookout, R. V. DOLPHIN cruises 1965-66: Zooplankton volumes, midwater trawl collections, temperatures and salinities. United States Fish and Wildlife Service Technical Paper, 28:1-132.

Fishes of the Western North Atlantic CLARK, S. H. 1974. A study of variation in trawl data collected in Everglades National Park, Florida. Transactions of the American Fisheries Society, 103(4):777-785. COLLETTE, B. B. 1962. Hemiramphus bermudensis, a new halfbeak from Bermuda, with a survey of endemism in Bermudian shore fishes. Bulletin of Marine Science of the Gulf and Caribbean, 12(3):432-449. COOLEY, N. R. 1978. An inventory of the estuarine fauna in the vicinity of Pensacola, Florida. Florida Marine Research Publications, 31:1-119. COPE, E. D. 1875. On a new genus of lophobranchiate fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 27:450. Cox, P. 1921. List of the fishes collected in 1917 of f the Cape Breton coast and the Magdalen Islands. Contributions to Canadian Biology, 1918-1920(11): 109-114. DAHL, G. 1971. Los peces del norte de Colombia. Instituto de Desarrollo de los Recursos Naturales Renovables, Bogota: xvii H- 391 pp. DAHLBERG, M. D. 1972. An ecological study of Georgia coastal fishes. Fishery Bulletin, 70(2):323-353. . 1975. Guide to coastal fishes of Georgia and nearby states. University of Georgia Press, Athens: xvii -I- 186 pp. DAHLBERG, M. D., AND E. P. ODUM. 1970. Annual cycles of species occurrence, abundance, and diversity in Georgia estuarine fish populations. American Midland Naturalist, 83(20):382-392. D'ANCONA, U. 1934. Le specie mediterranee del genere Syngnathus. Memorie. R. Comitato Talassografico Italiano, 210:1-79. DAWSON, A. B. 1932. Intracellular crystallization of haemoglobin in the erythrocytes of the northern pipefish Syngnathus fuscus. Biological Bulletin, Woods Hole, 63:492-495. . 1933. The relative numbers of immature erythrocytes in the circulating blood of several species of marine fishes. Biological Bulletin, Woods Hole, 64:33-43. DAWSON, C. E. 1970. A Mississippi population of the opossum pipefish, Oostethus line at us (Syngnathidae). Copeia, 1970(4):772-773. . 1972. Nektonic pipefishes (Syngnathidae) from the Gulf of Mexico off Mississippi. Copeia, 1972(4):844-848. . 1974. Pseudophallus brasiliensis (Pisces: Syngnathidae), a new freshwater pipefish from Brazil. Proceedings of the Biological Society of Washington, 87(36):405-410. —. 1977a. Review of the pipefish genus Corythoichthys with description of three new species. Copeia, 1977(2):295-338. . 1977b. Synopsis of syngnathine pipefishes

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usually referred to the genus Ichthyocampus Kaup, with description of new genera and species. Bulletin of Marine Science, 27(4):595-650. . 1978a. Micrognathus vittatus (Kaup), a junior synonym of M. crinitus (Jenyns), with description of the insular pipefish, M. tectus, new sp. Copeia, 1978(1):13-16. . 1978b. Syngnathus parvicarinatus, a new Australian pipefish, with notes on S. sauvagei (Whitley) and Leptonotus caretta (Klunzinger). Copeia, 1978(2):288-293. . 1978c. Review of the Indo-Pacific pipefish genus Hippichthys (Syngnathidae). Proceedings of the Biological Society of Washington, 91(1):132-157. . 1979a. Review of the polytypic doryrhamphine pipefish Oostethus brachyurus (Bleeker). Bulletin of Marine Science, 29(4):465-480. . 1979b. Notes on western Atlantic pipefishes with description of Syngnathus caribbaeus n. sp. and Cosmocampus n. gen. Proceedings of the Biological Society of Washington, 94(4):671-676. . 1980. Synopsis of the pipefishes (Syngnathidae) of New Zealand. National Museum of New Zealand Records, 1(17):281-291. DAWSON, C. E., AND G. R. ALLEN. 1978. Synopsis of the 'finless' pipefish genera (Penetopteryx, Apterygocampus and Enchelyocampus, gen. nov.). Records of the Western Australian Museum, 6(4):391-411. DAWSON, C. E., AND R. A. FRITZSCHE. 1975. Odontoid processes in pipefish jaws. Nature, 257(5525):390. DAWSON, C. E., F. YASUDA, AND C. IMAI. 1979. Elongate dermal appendages in species of Yozia (Syngnathidae) with remarks on Trachyrhamphus. Japanese Journal of Ichthyology, 25(4):244-250. DE BUEN, F. 1940. Lista de peces de agua dulce de Mexico. En preparacion de su catalogo. Trabajos de la Estacion Limnologica de Patzcuaro, 2:1-66. . 1959. Lampreas, tiburones, rayas y peces en la Estacion de Biologia Marina de Montemar, Chile. Revista de Biologia Marina, Valparaiso, 9(1-3):1-200. . 1963. Peces Chilenos. Beloniformes, Syngnathiformes y Gobiidae. Boletin de la Sociedad de Biologia de Concepcion, Chile, 35-36:81-101. DEKAY, J. E. 1842. Zoology of New York; or, the New York fauna. Part IV. Fishes. In Natural History of New York. New York State Geological Survey, Albany: 415 pp. DE LEON P., L. 1975. The freshwater pipefishes of Pacific Panama, aspects of their biology and ecology contributing to niche separation. Thesis, Universidad de Panama. DELFIN, F. T. 1899. Lista metodida de los peces de la Bahia de Concepcion i sus alrededores. Revista Chilena de Historia Natural, 3(12):176-178.

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—. 1900. Catalogo de los peces de Chile. Revista Chilena de Historia Natural, 4(2):21-25. . 1901. Catalogo de los peces de Chile. Ictiolojia Chilena. Imprenta Gillet, Valparaiso: 133 pp. DERICKSON, W. K., AND K. S. PRICE, JR. 1973. The fishes of the shore zone of Rehoboth and Indian River Bays, Delaware. Transactions of the American Fisheries Society, 102(3):552-562. DE SYLVA, D. P. 1954. The live bait fishery of the northeast coast of Florida. Florida State Board of Conservation, Technical Series, 11:1-35. DE SYLVA, D. P., F. A. KALBER, JR., AND C. N. SHUSTER, JR. 1962. Fishes and ecological conditions in the shore zone of the Delaware River estuary, with notes on other species collected in deeper water. University of Delaware Marine Laboratory, Information Series, 5:1-164. DEVINCENZI, C. J. 1939. Notas ictiologicas. Sobre peces de la region Patagonica. Analisis de la obra de J. R. Norman: Coast Fishes II. The Patagonian Region (Discovery Reports, Vol. XVI, 1937). Anales del Museo de Historia Natural de Montevideo, 2nd Series, 4(14):3-21. DOLLO, L. 1904. Poissons. In Resultats du voyage du S. Y. BELGICA en 1897-1898-1899. J. E. Buschmann, Anvers: 240 pp. DOOLEY, J. K. 1972. Fishes associated with the pelagic Sargassum complex, with a discussion of the Sargassum community. Contributions in Marine Science, University of Texas, 16:1-32. Doss, M. A., AND M. M. FARR. 1969. Index-Catalogue of Medical and Veterinary Zoology. Subjects: Trematoda and trematode diseases. Parts 11 and 12. United States Department of Agriculture: 694 pp. DOUGLAS, N. H. 1974. Freshwater fishes of Louisiana. Claitor's Publishing Division, Baton Rouge: xi -I- 443 PPDOVEL, W. L. 1971. Fish eggs and larvae of the Chesapeake Bay. Special Report, Natural Resources Institute, University of Maryland, 4:1-71. Not seen. DRAGOVICH, A. 1969. Review of studies of tuna food in the Atlantic Ocean. United States Fish and Wildlife Service, Special Scientific Report (Fisheries), 593:1-21. DUARTE-BELLO, P. P. 1959. Catalogo de peces Cubanos. Monografia de la Universidad de Villanueva, Marianao, 6:1-208. DUARTE-BELLO, P. P., AND R. J. BUESA. 1973. Catalogo de peces Cubanos (Primera revision). I. Indice taxonomico. Ciencias, Investigaciones Marinas, Universidad de la Habana, Serie 8, 3:1-255. DUKE, J. A., AND A. W. RUDOLPH. 1969. Bioenvironmental and radiological-safety feasibility studies. Atlantic-Pacific interoceanic canal. Darien fish di-

etary. Battelle Memorial Institute, Columbus, Ohio: 150 pp. DUMERIL 1870. Histoire naturelle des poissons, ou ichthyologie generale. II. Ganoi'des, Dipnes, Lophobranches. Librairie Encyclopedique de Roret, Paris: 624 pp. DUNCKER,G. 1899. Preliminary report on the results of statistical and ichthyological investigations made at the Plymouth laboratory. Journal of the Marine Biological Association of the United Kingdom, 5(2):172-175. . 1905. Ober regeneration des Schwanzendes bei Syngnathiden. Archiv fur Entwicklungsmechanik der Organismen, 20(l):30-37. . 1907. Ober regeneration des Schwanzendes bei Syngnathiden (Zweite Mitteilung). Archiv fur Entwicklungsmechanik der Organismen, 24:656-662. . 1910. On some syngnathids ("Pipe Fish") from Ceylon. Spolia Zeylanica, 7(25):25-34. . 1912. Die Gattungen der Syngnathidae. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 29:219-240. . 1915. Revision der Syngnathidae. Erster Teil. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 32:9-120. EATON, T. H., JR. 1942. Leaves out of a jungle. The Chicago Naturalist, 5(3):47-52. EIGENMANN, C. H. 1903. The fresh-water fishes of western Cuba. Bulletin of the United States Fish Commission, 22:211-236. . 1909. The fresh water fishes of Patagonia and an examination of the Archiplata-Archhelenis theory. Reports of the Princeton University Expeditions to Patagonia, 1896-1899, 3(3):211-374. . 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. Memoirs of the Carnegie Museum, 5:xxii + 1-578. EKMAN, S. 1953. Zoogeography of the Sea. Sidgwick and Jackson, London: xiv -1-417 pp. ERDMAN, D. S. 1956. Recent fish records from Puerto Rico. Bulletin of Marine Science of the Gulf and Caribbean, 6(4):315-340. ESKINAZI, A. M., DE OLIVEIRA. 1972a. Peixes estuarinos do nordeste oriental Brasileiro. Arquivos de Ciencias do Mar, Fortaleza, 12(1):35-41. . 1972b. Peixes do Canal de Santa Cruz-Pernambuco-BrasiL Travalhos de Oceanografia do Universidade Federal de Pernambuco, 13:283-302. . 1976. Composicao e distribuicao da ictiofauna, nas aguas estuarinas do Rio Jaguaribe (Ceara-Brasil). Arquivos de Ciencias do Mar, Fortaleza, 16(1):9-18.

Fishes of the Western North Atlantic ESKINAZI, A. M., DE OLIVEIRA, AND H. DE HOLANDA LIMA. 1968. Peixes marinhos do norte e nordeste do Brasil, coletados pelo Akaroa, Canopus et Noc. Almirante Saldanha. Arquivos de Ciencias do Mar, Fortaleza, 8(2):163-172. EVERMANN, B. W. 1899. Report on investigations by the U.S. Fish Commission in Mississippi, Louisiana, and Texas, in 1897. Report of the United States Fish Commission, 24:287-310. EVERMANN, B. W., AND B. A. BEAN. 1897. Indian River and its fishes. Report of the United States Commissioner of Fish and Fisheries, 22(Appendix 5):223-262. EVERMANN, B. W., AND S. F. HILDEBRAND. 1910. On a collection of fishes from the lower Potomac, the entrance of Chesapeake Bay and from streams flowing into these waters. Proceedings of the Biological Society of Washington, 23:157-164. EVERMANN, B. W., AND W. C. KENDALL. 1894. The fishes of Texas and the Rio Grande Basin, considered chiefly with reference to their geographic distribution. Bulletin of the United States Fish Commission, 12:57-126. . 1896. Description of a new species of pipefish (Siphostoma scovelli) from Corpus Christi, Texas. Proceedings of the United States National Museum, 18:113-115. . 1898. Descriptions of new or little-known genera and species of fishes from the United States. Bulletin of the United States Fish Commission, 17:125-133. . 1899. Check-list of the fishes of Florida. Report of the United States Commission of Fish and Fisheries, 25:35-103. EVERMANN, B. W., AND M. C. MARSH. 1900. The fishes of Puerto Rico. United States Commission of Fish and Fisheries, Washington: vi + 1-350. Also, 1902, Bulletin of the United States Fish Commission, 20(1):1-350. EVERMANN, B. W., AND L. RADCLIFFE. 1917. The fishes of the west coast of Peru and the Titicaca Basin. Bulletin of the United States National Museum, 95:1-166. EYDOUX, J. F. T., AND F. L. P. GERVAIS. 1837a. Voyage de "La Favorite." Poissons. Magasin de Zoologie, Paris, 7(4):l-4. . 1837b. Voyage autour du monde de la corvette de S. M. "La Favorite," execute pendant les annees 1830, 1831, 1832 sous le commandement de M. Laplace. Zoologie, Poissons. Paris, 5:1-200. FAHAY, M. P. 1975. An annotated list of larval and juvenile fishes captured with surface towed meter net in the South Atlantic Bight during four R. V. DOLPHIN cruises between May 1967 and February

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1968. NOAA Technical Report, National Marine Fisheries Service, SSRF-685, 39 pp. FERNANDEZ- YEPEZ, A. 1972. Analysis ictiologico del complejo hidrografico "Rio Yaracuy." Direccion de Obras Hidaulicas, Caracas: 25 pp. FIGUEIREDO, J. L., AND N. A. MENEZES. 1980. Manual de peixes marinhos do sudeste do Brasil. III. Teleostei (2). Museu de Zoologia, Universidade de Sao Paulo: 90pp. FINUCANE, J. H. 1969. Antimycin as a toxicant in a marine habitat. Transactions of the American Fisheries Society, 98(2):288-292. FISCHER, W. 1963. Die fische des brackwassergebietes Lenga bei Concepcion (Chile). Internationale Revue der gesamten Hydrobiologie, 48(3):419-511. FISH, C. J. 1925. Seasonal distribution of the plankton of the Woods Hole region. Bulletin of the United States Fish Commission, 41:91-179. FISH, C. J., AND M. W. JOHNSON. 1937. The biology of the zooplankton population in the Bay of Fundy and Gulf of Maine with special reference to production and distribution. Journal of the Biological Board of Canada, 3(3)189-322. FOWLER, H. W. 1906a. Some cold-blooded vertebrates of the Florida Keys. Proceedings of the Academy of Natural Sciences of Philadelphia, 58:77-113. --. 1906b. Fishes of New Jersey. Annual Report of the New Jersey State Museum, 1905:35-477. --. 1909. A new species of fish of the genus Atopichthys, with notes on New Jersey fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 61:406-407. --. 1911. The fishes of Delaware. Proceedings of the Academy of Natural Sciences of Philadelphia, 63:3-16. -. 1912. Records of fishes for the middle Atlantic states and Virginia. Proceedings of the Academy of Natural Sciences of Philadelphia, 64:34-59. --. 1913. Notes on the fishes of the Chincoteague region of Virginia. Proceedings of the Academy of Natural Sciences of Philadelphia, 65:61-65. --. 1914a. Notes on New Jersey fishes. Copeia, . 1914b. Notes on the fishes at Ocean City, Maryland. Copeia, 1914(2):! -2. . 1915a. Fishes of Trinidad, Grenada and St. Lucia, British West Indies. Proceedings of the Academy of Natural Sciences of Philadelphia, 67:520-546. . 1915b. A list of Santo Domingo fishes. Copeia, 1915(24):50-51. . 1915c. Cold-blooded vertebrates from Florida, the West Indies, Costa Rica and eastern Brazil. Proceedings of the Academy of Natural Sciences of Philadelphia, 67:244-269.

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Memoir 1, Part 8—Sears Foundation for Marine Research . 1916. Records of northern New Jersey fishes. Copeia, 1916(31):41-42. . 1917. Notes on New England fishes. Proceedings of the Boston Society of Natural History, 35(4): 109-138. . 1919a. Notes on New Jersey, Pennsylvania, and Virginia fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 71:292-300. . 1919b. The fishes of the United States Eclipse Expedition to West Africa. Proceedings of the United States National Museum, 56:195-292. . 1919c. Notes on tropical American fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 71:128-155. . 1920. Description of a new cyprinoid fish (Notropis stonei), with notes on other fishes obtained in the United States. Proceedings of the Academy of Natural Sciences of Philadelphia, 72:385-402. . 1922a. Notes on hemibranchiate and lophobranchiate fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 73:437-448. . 1922b. Records of fishes for the eastern and southern United States. Proceedings of the Academy of Natural Sciences of Philadelphia, 74:1-27. . 1926a. Fishes from Florida, Brazil, Bolivia, Argentina, and Chile. Proceedings of the Academy of Natural Sciences of Philadelphia, 78:249-285. . 1926b. Records of fishes in New Jersey, 1925. Copeia, 1926(156):146-150. . 1927. Fishes from Ocracoke, North Carolina. Copeia, 1927(165):89. . 1928. The fishes of Oceania. Memoirs of the Bernice P. Bishop Museum, 10:iii -I- 540 pp. . 1929. Notes on New Jersey fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 80:607-614. . 1931. A collection of fishes from the Texas coast. Copeia, 1931(2):46-50. . 1933. Notes on Louisiana fishes. Proceedings of the Biological Society of Washington, 46:57-64. . 1934. A synopsis of the fishes of China. Part V continued. The cods, opahs, flounders, soles, John Dories, berycoids, pipe fishes, silversides, mullets, barracudas and thread fishes. The Hong Kong Naturalist, 5(4):304-319. . 1936. The marine fishes of West Africa. Bulletin of the American Museum of Natural History, 70(1):1-605. . 1937. Notes on fishes from the Gulf Stream and the New Jersey coast. Proceedings of the Academy of Natural Sciences of Philadelphia, 89:297-308. . 1940a. A collection of fishes obtained on the west coast of Florida by Mr. and Mrs. C. G. Chaplin. Proceedings of the Academy of Natural Sciences of Philadelphia, 92:1-22.

. 1940b. The fishes obtained by the Wilkes Expedition, 1838-1842. Proceedings of the American Philosophical Society, 82(5):733-800. . 1941a. A list of the fishes known from the coast of Brazil. Arquivos de Zoologia do Estado de Sao Paulo, 3(6):115-184. . 1941b. A collection of fresh-water fishes obtained in Florida, 1939-1940, by Francis Harper. Proceedings of the Academy of Natural Sciences of Philadelphia, 92:227-244. . 1941c. Notes on Florida fishes with descriptions of seven new species. Proceedings of the Academy of Natural Sciences of Philadelphia, 93:81-106. . 1942. Notes on marine fishes from Honduras. Fish Culturist, 22(2):9-12. . 1944a. The fishes. In Results of the fifth George Vanderbilt Expedition (1941). The Academy of Natural Sciences of Philadelphia, Monographs, 6:57-529. . 1944b. Fishes obtained in the New Hebrides by Dr. Edward L. Jackson. Proceedings of the Academy of Natural Sciences of Philadelphia, 96:155-199. . 1945a. A study of the fishes of the Southern Piedmont and Coastal Plain. The Academy of Natural Sciences of Philadelphia, Monographs, 7:vi -I- 408 pp. . 1945b. Los peces del Peru. Catalogo Sistematico de los peces que habitan en aguas peruanas. Museo de Historia Natural "Javier Prado" Universidad Nacional Mayor de San Marcos, Lima: 298 pp. . 1945c. Fishes of Chile. Systematic catalog. Revista Chilena de Historia Natural, 45-47:1-171. . 1949. The fishes of Oceania. Supplement 3. Memoirs of the Bernice P. Bishop Museum, 12(2):37-186. . 1952. A list of the fishes of New Jersey, with off-shore species. Proceedings of the Academy of Natural Sciences of Philadelphia, 104:89-151. Fox, L. S., AND W. R. MOCK, JR. 1968. Seasonal occurrence of fishes in two shore habitats in Barataria Bay, Louisiana. Proceedings of the Louisiana Academy of Sciences, 31:43-53. FRITZSCHE, R. A. 1976. A revision of the eastern Pacific Syngnathidae (Pisces: Syngnathiformes). Ph.D, thesis, University of California, San Diego. . 1980. Revision of the eastern Pacific Syngnathidae (Pisces: Syngnathiformes), including both recent and fossil forms. Proceedings of the California Academy of Sciences, 42(6):181-227. CARMAN, S. 1896. Report on the fishes collected by the Bahama Expedition, of the State University of Iowa, under Professor C C Nutting in 1893. Bulletin from the Laboratories of Natural History of the State University of Iowa, 4(l):76-93.

Fishes of the Western North Atlantic GILBERT, C. H., AND E. C. STARKS, 1904. The fishes of Panama Bay. Memoir of the California Academy of Sciences, 4:1-304. GILBERT, C. R., AND D. P. KELSO. 1971. Fishes of the Tortuguero area, Caribbean Costa Rica. Bulletin of the Florida State Museum, Biological Sciences, GILL, T. N. 1862. Catalogue of the fishes of the eastern coast of North America from Greenland to Georgia. Proceedings of the Academy of Natural Sciences of Philadelphia, 1861 (Supplement): 1-63. --. 1863. Synopsis of the lophobranchiate fishes of western North America. Proceedings of the Academy of Natural Sciences of Philadelphia, 1862:282-284. --. 1865. Synopsis of the fishes of the Gulf of St. Lawrence and Bay of Fundy. The Canadian Naturalist and Geologist (new series), 2:244-266. --. 1873. Catalogue of the fishes of the east coast of North America. Report of the United States Commissioner of Fish and Fisheries, 1871-72:779-822. Also, Smithsonian Miscellaneous Collections, 283:1-50. --. 1893. A comparison of antipodal faunas. Memoirs of the National Academy of Sciences, 6:91-124. --. 1909. Angler fishes: their kinds and ways. Annual Report of the Smithsonian Institution, 1908:565-615. GILMORE, R. G., JR. 1977a. Fishes of the Indian River Lagoon and adjacent waters, Florida. Bulletin of the Florida State Museum, Biological Sciences, 22(3):101-147. --. 1977b. Notes on the opossum pipefish Oostethus line at us from the Indian River Lagoon and vicinity, Florida. Copeia, 1977(4):781-783. GOODE, G. B. 1876. Catalogue of the fishes of the Bermudas. Bulletin of the United States National Museum, 5:1-82. --. 1879. A preliminary catalogue of the fishes of the St. John's River and the east coast of Florida, with description of a new genus and three new species. Proceedings of the United States National Museum, 2:108-121. GOODE, G. B., AND T.H. BEAN. 1879. A list of the fishes of Essex County, including those of Massachusetts Bay, according to the latest results of the work of the U.S. Fish Commission. Bulletin of the Essex Institute, Salem, 11:1-38. --. 1880. Catalogue of a collection of fishes obtained in the Gulf of Mexico, by Dr. J. W. Velie, with descriptions of seven new species. Proceedings of the United States National Museum, 2:333-345. GORDON, B. L. 1956. The paternal pipefish. Nature Magazine, 49(5):243-44, 276.

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. 1960. The marine fishes of Rhode Island. The Book & Tackle Shop, Watch Hill: 136 pp. GORHAM, F. P. 1901. The gas-bubble disease of fish and its causes. Bulletin of the United States Fish Commission, 19:33-37. GOTSCHLICH, B. 1913. Llanquihue i Valdivia. Boletin del Museo Nacional (Santiago, Chile), 6(l):7-626. GREELEY, J. R. 1937. A biological survey of the lower Hudson watershed. II. Fishes of the area with annotated list. Report of the New York State Conservation Department, 26(Supplement 11):45-103. . 1939. A biological survey of the salt waters of Long Island, 1938. Section II. Fishes and habitat conditions of the shore zone based upon July and August seining investigations. Report of the New York State Conservation Department, 28(Supplement 2):72-91. GREY, M. 1947. Catalogue of type specimens in Chicago Natural History Museum. Fieldiana, Zoology, 32(3):109-205. GUDGER, E. W. 1905. The breeding habits and the segmentation of the eggs of the pipefish, Siphostoma floridae. Proceedings of the United States National Museum, 29:447-500. GUICHENOT, A. 1848. Peces de Chile. In Gay, C., Historia fisica y politica de Chile. Zoologia. Paris, 2:137-372. GUITART, D. J. 1975. Sinopsis de los peces marines de Cuba. Academia de Ciencias de Cuba, Institute de Oceanologia, Habana, 2:141-323. GUNTER, G. 1935. Records of fishes rarely caught in shrimp trawls in Louisiana. Copeia, 1935(1 ):39-40. . 1942. A list of the fishes of the mainland of North and Middle America recorded from both freshwater and sea water. American Midland Naturalist, 28(2):305-326. . 1945. Studies on marine fishes of Texas. Publications of the Institute of Marine Science, University of Texas, 1(1):1-190. . 1956. A revised list of euryhaline fishes of North and Middle America. American Midland Naturalist, 56(2):345-354. . 1979. Marine fishes of Panama as related to the canal. Gulf Research Reports, 6(3):267-273. GUNTER, G., AND G. E. HALL. 1963. Biological investigations of the St. Lucie estuary (Florida) in connection with Lake Okeechobee discharges through the St. Lucie Canal. Gulf Research Reports, 1(5): 189-307. --. 1965. A biological investigation of the Caloosahatchee estuary of Florida. Gulf Research Reports, GUNTHER, A. 1870. Catalogue of fishes in the British Museum (Natural History), 8:xxv + 549 pp. London. --. 1874. Descriptions of new species of fishes in

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the British Museum. Annals and Magazine of Natural History, 4th series, 14:368-371, 453-455. - . 1880a. Report on the shorefishesprocured during the voyage of H.M.S. CHALLENGER in the years 1873-1876. Report on the Scientific Results of the voyage of H.M.S. CHALLENGER, Zoology, . 1880b. An introduction to the study of fishes. Adams & Black, Edinburgh: 720 pp. . 1910. Andrew Garrett's Fische der Siidsee, beschrieben und redigirt von A. C. L. G. Giinther. Journal des Museum Godeffroy, Hamburg, 6(17): 389-514. HAEFNER, P. A., JR. 1969. Occurrence of a larval alligator fish (Agonidae) in brackish water. Copeia, 1969(1):201-202. HAMILTON, W. J., JR. 1942. Habits of Syngnathus scovelli (Evermann and Kendall). Copeia, 1942(3):188. HARDY, J. D., JR. 1978. Anguillidae through Syngnathidae. In Development of fishes of the mid-Atlantic bight. An atlas of egg, larval and juvenile states. United States Fish and Wildlife Service, Washington, 2:1-458. HASSE, J. J. 1974. Observations on egg hatching of Syngnathoides biaculeatus (Bloch) (Pisces: Syngnathidae). Micronesica, 10(2):279-283. HASTINGS, P. A., AND S. A. BORTONE. 1976. Additional notes on tropical marine fishes in the northern Gulf of Mexico. Florida Scientist, 39(2): 122-125. HELLIER, T. R., JR. 1967. The fishes of the Santa Fe River system. Bulletin of the Florida State Museum, Biological Sciences, 2(l):l-46. HENSHALL, J. A. 1891. Report upon a collection of fishes made in southern Florida in 1889. Bulletin of the United States Fish Commission, 9:371-389. HERALD, E. S. 1940. A key to the pipefishes of the Pacific American coasts with descriptions of new genera and species. Allan Hancock Pacific Expeditions, 9(3):51-64. . 1941. A systematic analysis of variation in the western American pipefish, Syngnathus californiensis. Stanford Ichthyological Bulletin, 2(3):49-73. . 1942. Three new pipefishes from the Atlantic coast of North and South America, with a key to the Atlantic American species. Stanford Ichthyological Bulletin, 2(4):125-134. . 1943. Studies on the classification and interrelationships of the American pipefishes. Ph.D. thesis, Stanford University. . 1950. Ichthyocampus pawned, a new pipefish from the Bahamas. Journal of the Washington Academy of Sciences, 40(8):269. . 1953. Family Syngnathidae: Pipefishes. In L. P. Schultz et al., Fishes of the Marshall and Mari-

anas Islands. Bulletin of the United States National Museum, 202(1 ):231-278. . 1959. From pipefish to seahorse . . . a study of phylogenetic relationships. Proceedings of the California Academy of Sciences, Fourth Series, 29(13):465-473. . 1961. Living fishes of the world. Doubleday, New York: 304 pp. . 1965. Studies on the Atlantic American pipefishes with descriptions of new species. Proceedings of the California Academy of Sciences, Fourth Series, 32(12):363-375. . 1969. A new pipefish from the Virgin Islands, Micrognathus dawsoni. Occasional Papers of the California Academy of Sciences, 73(4):l-3. . 1972. Fishes of North America. Doubleday, New York: 254 pp. HERALD, E. S., AND C. E. DAWSON. 1972. A new subspecies of the Gulf pipefish, Syngnathus scovelli makaxi (Pisces: Syngnathidae). Copeia, 1972(4):781-784. . 1974. Micrognathus erugatus, a new marine pipefish from Brazil (Pisces: Syngnathidae). Proceedings of the Biological Society of Washington, 87(4):27-30. HERALD, E.S., AND J.E.RANDALL. 1972. Five new IndoPacific pipefishes. Proceedings of the California Academy of Sciences, Fourth Series, 39(11):121-140. HERMAN, S. S. 1963. Planktonic fish eggs and larvae of Narragansett Bay. Limnology and Oceanography, 8(1):103-109. HERRE, A. W. (C. T.) 1931. A check list of the fishes recorded from the New Hebrides. Journal of the Pan-Pacific Research Institute, 6(4):11-14. . 1936. Notes on fishes in the Zoological Museum of Stanford University, V. New or rare Philippine fishes from the Herre 1933 Philippine Expedition. Philippine Journal of Science, 59(3):357-373. . 1942. Notes on a collection of fishes from Antigua and Barbados, British West Indies. Stanford University Publications, Biological Sciences, 7(2):287-305. . 1953. Check list of Philippine fishes. United States Fish and Wildlife Service Research Report, 20:1-970. HERREMA, D. J. 1974. Marine and brackish water fishes of southern Palm Beach and northern Broward counties, Florida. M.Sc. thesis, Florida Atlantic University. HILDEBRAND, H. H. 1954. A study of the fauna of the brown shrimp (Penaeus aztecus Ives) grounds in the western Gulf of Mexico. Publications of the Institute of Marine Science, University of Texas, 3(2):233-366.

Fishes of the Western North Atlantic . 1955. A study of the fauna of the pink shrimp (Penaeus duorarum Burkenroad) grounds in the Gulf of Campeche. Publications of the Institute of Marine Science, University of Texas, 4(1): 169-232. HILDEBRAND, H. H., H. CHAVEZ, AND H. COMPTON.

1964. Aporte al conocimiento de los peces del Arrecife Alacranes, Yucatan (Mexico). Ciencia, Mexico, 23(3):107-134. HILDEBRAND, S. F. 1946. A descriptive catalog of the shore fishes of Peru. Bulletin of the United States National Museum, 189:xi -I- 530 pp. HILDEBRAND, S. F., AND W. C. SCHROEDER. 1928. Fishes of Chesapeake Bay. Bulletin of the United States Bureau of Fisheries, 43(1 ):1-366. HOESE, H. D. 1958. A partially annotated checklist of the marine fishes of Texas. Publications of the Institute of Marine Science, University of Texas, 5:312-352. . 1960. Biotic changes in a bay associated with the end of a drought. Limnology and Oceanography, 5(3):326-336. HOESE, H. D., AND R. S. JONES. 1963. Seasonally of larger animals in a Texas turtle grass community. Publications of the Institute of Marine Science, University of Texas, 9:37-47. HOESE, H. D., AND R. H. MOORE. 1977. Fishes of the Gulf of Mexico, Texas, Louisiana and adjacent waters. Texas A.&M. University Press: xv -I- 327 pp. HOLLIMAN, R. B. 1963. Gyrodactylus shorti, a new species of monogenetic trematode from the brood pouch of the southern pipefish, Syngnathus scovelli (Evermann and Kendall). Tulane Studies in Zoology, 10(2):83-86. HOWELL RIVERO, L. 1936. Some new, rare and little known fishes from Cuba. Proceedings of the Boston Society of Natural History, 41(4):41-76. . 1938. List of fishes, types of Poey, in the Museum of Comparative Zoology. Bulletin of the Museum of Comparative Zoology, Harvard University, 82(3): 169-227. HUBBS, C. 1958. A checklist of Texas fresh-water fishes. Texas Game and Fish Commission IF Series, 3:1-14. . 1972. A checklist of Texas freshwater fishes. Texas Parks and Wildlife Department Technical Series, 11:1-11. HUBBS, C. L. 1929. Oostethus: A new generic name for a doryrhamphine pipefish. Occasional Papers of the Museum of Zoology, University of Michigan, 199:1-4. . 1935. Anarchopterus, a new genus of syngnathid fishes from the western Atlantic. Occasional Papers of the Museum of Zoology, University of Michigan, 320:1-3. . 1936. Fishes of the Yucatan Peninsula. Publi-

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cations of the Carnegie Institution of Washington, 457:157-287. HUBRECHT, A. A. W. 1879. Catalogue des collections formees et laissees par M. P. Bleeker. De Breuk & Smits, Leiden: 71 pp. HUDSON, J. H., D. M. ALLEN, AND T. J. COSTELLO. 1970. The flora and fauna of a basin in central Florida Bay. United States Fish and Wildlife Service Special Scientific Report (Fisheries), 604:1-14. HUNNINEN, A. V., AND R. M. CABLE. 1940. Studies on the life history of Anisoporus manteri sp. nov. (Trematoda: Allocreadiidae). Biological Bulletin, Woods Hole, 79:373-374. HUNTSMAN, A. G. 1922. The fishes of the Bay of Fundy. Contributions to Canadian Biology, 1921(3):49-72. HUTTON, F. W. 1872. Fishes of New Zealand. Catalogue with diagnoses of the species. Colonial Museum and Geological Survey Department, Wellington: xvi -I- 133 pp. . 1904. Index faunae Novae Zelandiae. Dulau, London: 372 pp. JARED, D. W., AND R. A. WALLACE. 1968. Comparative chromatograhy of the yolk proteins of teleosts. Comparative Biochemistry and Physiology, 24(2):437-443. JENKINS, O. P. 1887. A list of the fishes of Beaufort harbor, North Carolina. Studies from the Biological Laboratory, Johns Hopkins University, 4:83-94. JENYNS, L. 1835. A systematic catalogue of British vertebrate animals. Cambridge: 22 pp. . 1842. Fish. In The zoology of the voyage of H.M.S. BEAGLE, under the command of Captain Fitzroy, R.N., during the years 1832 to 1836. Smith, Elder & Co., London, 4:1-172. JONES, J. M. 1882. List of the fishes of Nova Scotia. Proceedings and Transactions of the Nova Scotian Institute for Natural Science, 5(Appendix):87-97. JONES, R. S., R. G. GILMORE, JR., G. R. KULCZYCKI, W. C. MAGLEY, AND B. GRAUNKE. 1975. Studies of the fishes of the Indian River Coastal Zone. Annual Report of the Indian River Coastal Zone Study. Harbor Branch Consortium 1974-1975, 1:57-88. JORDAN, D. S. 1880. Notes on a collection of fishes from Saint John's River, Florida, obtained by Mr. A. H. Curtiss. Proceedings of the United States National Museum, 3:22. . 1884. List of fishes collected at Key West, Florida, with notes and descriptions. Proceedings of the United States National Museum, 7:103-150. . 1886a. Notes on some fishes collected at Pensacola, by Mr. Silas Stearns, with descriptions of one new species (Chaetodon aya). Proceedings of the United States National Museum, 9:225-229. . 1886b. Notes on fishes collected at Beaufort,

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North Carolina, with a revised list of the species known from that locality. Proceedings of the United States National Museum, 9:25-30. . 1887a. A preliminary list of the fishes of the West Indies. Proceedings of the United States National Museum, 9:554-608. . 1887b. A catalogue of the fishes known to inhabit the waters of North America, north of the Tropic of Cancer, with notes on the species discovered in 1883 and 1884. Report of the United States Commission of Fish and Fisheries, 12:789-973. . 1890a. Scientific results of explorations by the U.S. Fish Commission steamer ALBATROSS. XVIII. List of fishes obtained in the harbor of Bahia, Brazil, and in adjacent waters. Proceedings of the United States National Museum, 13:313-336. . 1890b. Scientific results of explorations by the U.S. Fish Commission steamer ALBATROSS. IX. Catalogue of fishes collected at Port Castries, St. Lucia, by the steamer ALBATROSS, November, 1888. Proceedings of the United States National Museum, 12:645-652. . 1895. The fishes of Sinaloa. Proceedings of the California Academy of Sciences, Series 2,5:377-513. . 1917. The genera of fishes, from Linnaeus to Cuvier, 1758-1833, seventy-five years, with the accepted type of each. A contribution to the stability of scientific nomenclature. Leland Stanford Junior University Publications, University Series: 161 pp. . 1919. The genera of fishes, part II, from Agassiz to Bleeker, 1833-1858, twenty-six years, with the accepted type of each. A contribution to the stability of scientific nomenclature. Leland Stanford Junior University Publications, University Series; ix -I- 163-284 -I- xiii pp. . 1925. Fishes. D. Appleton, New York: 773 pp. . 1929. Manual of the vertebrate animals of the northeastern United States inclusive of marine species. 13th edn. World Book Co., Yonkers-onHudson: 446 pp. JORDAN, D. S., AND M. C. DICKERSON. 1908. Notes on a collection of fishes from the Gulf of Mexico at Vera Cruz and Tampico. Proceedings of the United States National Museum, 34:11-22. JORDAN, D. S., AND B. W. EVERMANN. 1896a. The fishes of North and Middle America. Bulletin of the United States National Museum, 47(1 ):lx + 1240 pp. . 1896b. A check-list of the fishes and fish-like vertebrates of North and Middle America. Report of the United States Commission of Fish and Fisheries, 21:207-584. . 1898. The fishes of North and Middle America. Bulletin of the United States National Museum, 47(3):xxiv -I- 2183-3136.

. 1927. New genera and species of North American fishes. Proceedings of the California Academy of Sciences, Fourth Series, 16(15):501-507. JORDAN, D. S., B. W. EVERMANN, AND H. W. CLARK. 1930. Check list of the fishes and fishlike vertebrates of North and Middle America north of the northern boundary of Venezuela and Colombia. Report of the United States Commissioner of Fisheries for 1928, Part II: 670 pp. JORDAN, D. S., AND C. H. GILBERT. 1879. Notes on the fishes of Beaufort Harbor, North Carolina. Proceedings of the United States National Museum, 1:365-388. . 1882a. Notes on fishes observed about Pensacola, Florida, and Galveston, Texas, with description of new species. Proceedings of the United States National Museum, 5:241-307. . 1882b. Synopsis of the fishes of North America. Bulletin of the United States National Museum, 16: Iv -I- 1016 pp. . 1883. Notes on a collectiorf of fishes from Charleston, South Carolina, with descriptions of three new species. Proceedings of the United States National Museum, 5:580-620. JORDAN, D. S., AND C. RUTTER. 1897. A collection of fishes made by Joseph Seed Roberts in Kingston, Jamaica. Proceedings of the Academy of Natural Sciences of Philadelphia, 49:91-133. JORDAN, D. S., AND J. SWAIN. 1884. Notes on fishes collected by David S. Jordan at Cedar Keys, Florida. Proceedings of the United States National Museum, 7:230-234. JORDAN, D. S., AND J. C. THOMPSON. 1905. Fish fauna of the Tortugas Archipelago. Bulletin of the United States Bureau of Fisheries, 24:229-256. JOSEPH, E. B. 1957. A study of the systematics and life history of the Gulf pipefish, Syngnathus scovelli (Evermann and Kendall). Ph.D. thesis, Florida State University. JOSEPH, E. B., AND R. W. YERGER. 1956. The fishes of Alligator Harbor, Florida, with notes on their natural history. Florida State University Studies, 22:111-156. JUNGERSON, H. F. E. 1910. Ichthyotomical contributions. II. The structure of the Aulostomidae, Syngnathidae and Solenostomidae. Det kongelige danske videnskabernes selskabs skrifter, 7. raekke, naturvidenskabelig og matematisk afdeling, 8(5):268-364. KADAM, K. M. 1958. The development of the chondrocranium in the sea-horse, Hippocampus (Lophobranchii). Journal of the Linnean Society of London. Zoology, 43(293):557-573. . 1961. The development of the skull in Nerophis (Lophobranchii). Acta Zoologica, 42:257-298.

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LONGLEY, W. H. 1916. The significance of the colors of tropical reef fishes. Carnegie Institution of Washington Yearbook, 15:209-212. . 1917. Studies upon the biological significance of animal coloration. I. The color and color changes of West Indian reef fishes. Journal of Experimental Zoology, 23:533-601. . 1932. Preparation of a monograph o n the Tortugas fishes. Carnegie Institution of Washington Yearbook, 31:299-301. . 1933. Preparation of a monograph o n the Tortugas fishes. Carnegie Institution of Washington Yearbook, 32:293-295. . 1934. Preparation of a monograph of Tortugas fishes. Carnegie Institution of Washington Yearbook, 33:270-272. LONGLEY, W. H., AND S. F. HiLDEBRAND. 1941. Systematic catalogue of the fishes of Tortugas, Florida, with observations on color, habits, and local distribution. Carnegie Institution of Washington Publication, 535:xiii + 1-331. LOZANO KEY, L. 1947. Peces Ganoideos y Fisostomos. Memorias de la Real Academia de Ciencias Exactas, Fisicas y Naturales de Madrid, Serie de Ciencias Naturales, 11:1-839. LUEDERWALDT, H. 1929. Resultados de uma excursao scientifica a Ilha de Sao Sebastiao no littoral do Estado de Sao Paulo e'em 1925. Revista do Museu Paulista, 16:21-46. LULING, K. H. 1974. Wissenschaftliche Ergebnisse der Argentinien-Siidbrasilien-Expedition Dr. K. H. Liiling 1970/71. Der Rio Mage-Roncador und seine charakteristischen Fische von der Einmiindung in der Bucht von Rio de Janeiro nebst einer kurzen Notiz iiber einen Bach im Orgelgebirge (Siidostbrasilien). Zoologischer Anzeiger, 193(3/4):193-205. LUNEL, G. 1881. Melanges ichthyologiques. Liste de quelques especes de poissons, nouvelles pour la faune de Tile Maurice. Memoires de la Societe de Physique et d'Histoire Naturelle de Geneve, 27(2):267-303. MACLEAY, W. 1882. Descriptive catalogue of the fishes of Australia. Proceedings of the Linnean Society of New South Wales, 6:202-387. MAGO L., F. 1965a. Contribucion a la sistematica y ecologia de los peces de la Laguna de Unare, Venezuela. Bulletin of Marine Science, 15(2):274-330. . 1965b. Nuevas adiciones a la ictiofauna de Venezuela. I. Acta Biologica Venezuela, Caracas, 4(13):366-420. . 1970. Lista de los peces de Venezuela incluyendo un estudio preliminar sobre la ictiogeografia del pais. Ministerio de Agricultura y Cria, Caracas: 283 pp.

MANN F., G. 1954. Vida de los peces en aguas chilenas. Institute de Investigaciones Veterinarias, Santiago: 342 pp. MANSUETI, R. 1957. Revised key to Maryland freshwater fishes. Chesapeake Biological Laboratory Report, 57-22:1-26. Not seen. . 1962. Checklist of fishes of the Patuxent River drainage and of Chesapeake Bay off Calvert County, Maryland. Chesapeake Biological Laboratory Report, 62-36:1-5. Not seen. MANSUETI, R., AND R. S. SCHELTEMA. 1953. Summary of fish collections made in the Chesapeake Bay area of Maryland and Virginia during October 1953. Chesapeake Biological Laboratory Field Summary, 1:1-18. Not seen. MANTER, H. W. 1947. The digenetic trematodes of marine fishes of Tortugas, Florida. American Midland Naturalist, 38(2):257-416. MARKLE, D. F., W. B. SCOTT, AND A. C. KOHLER. 1980. New and rare records of Canadian fishes and the influence of hydrography on the resident and nonresident Scotian Shelf ichthyofauna. Canadian Journal of Fisheries and Aquatic Sciences, 37(l):49-65. MAURER, D., AND L. WATLING. 1973. The biology of the oyster community and its associated fauna in Delaware Bay. University of Delaware, Delaware Bay Report Series, 6:3-97. MCALLISTER, D. E. 1968. The evolution of branchiostegals and associated opercular, gular, and hyoid bones and the classification of teleostome fishes, living and fossil. Bulletin of the National Museum of Canada, 221:xiv + 239 pp. McCosKER, J. E., AND C. E. DAWSON. 1975. Biotic passage through the Panama Canal, with particular reference to fishes. Marine Biology, 30(4):343-351. McCuLLOCH, A. R. 1929. A check-list of the fishes recorded from Australia. Memoirs of the Australian Museum, Sydney, 5(1):1-144. McLANE, W. M. 1955. The fishes of the St. Johns River System. Ph.D. thesis, University of Florida. MCMURRICH, J. P. 1883. On the osteology and development of Syngnathus peckianus. Quarterly Journal of Microscopical Science, 23:623-652. MEAD, G. W. 1958. A catalog of the type specimens of fishes formerly in the collections of the Department of Tropical Research, New York Zoological Society. Zoologica, New York, 43(4):131-134. MEEK, S. E. 1904. The fresh-water fishes of Mexico north of the Isthmus of Tehuantepec. Publication of the Field Columbian Museum, Zoological Series, 5:lxiii + 252 pp. . 1914. An annotated list of fishes known to occur in the fresh waters of Costa Rica. Field Museum

Fishes of the Western North Atlantic of Natural History Publication 174, Zoological Series, 10(10):101-134. MEEK, S. E., AND S. F. HILDEBRAND. 1923. The marine fishes of Panama. Field Museum of Natural History Publication 215, Zoological Series, 15(l):xi -I- 330 PPMELLEN, I. M. 1919. Prehensile-tailed pipefishes. Bulletin of the New York Zoological Society, 22(6):133-135. MERCER, L. P. 1973. The comparative ecology of two species of pipefish (Syngnathidae) in the York River, Virginia. M. A. thesis, College of William and Mary. MERRIMAN, D. 1947, Notes on the midsummer ichthyofauna of a Connecticut beach at different tide levels. Copeia, 1947(4):281-286. MERRIMAN, D., AND R. C. SCLAR. 1952. The pelagic fish eggs and larvae of Block Island Sound. Bulletin of the Bingham Oceanographic Collection, 13:165-219. METZEIJ\AR, ]. 1919. Report on the fishes collected by Dr. ]. Boeke in the Dutch West Indies 1904-1905, with comparative notes on marine fishes of tropical West Africa. A. H. Kruyt, Amsterdam: 315 pp. --. 1922. On a collection of marine fishes from the Lesser Antilles. Bijdragen tot de Dierkunde, 22:133-141. MILLER, G. L., AND S. C. JORGENSON. 1973. Meristic characters of some marine fishes of the western Atlantic Ocean. Fishery Bulletin, 71(1):301-312. MILLER, R. R. 1966. Geographical distribution of Central American freshwater fishes. Copeia, 1966(4):773-802. --. 1976. An evaluation of Seth E. Meek's contribution to Mexican ichthyology. Fieldiana: Zoology, MIRANDA RIBEIRO, A. 1915. Fauna Brasiliense, Peixes. Archivos do Museu Nacional do Rio de Janeiro, 17:1-227. --. 1918a. Fauna Brasiliense (Peixes) Tomo V. Eleutherobranchios, Aspirophoros, Physoclisti. Archivos do Museu Nacional do Rio de Janeiro, 21:1-227. --. 1918b. Lista dos peixes Brazileiros do Museu Paulista. Revista do Museu Paulista, 10:763-783. MIRANDA RIBEIRO, P. 1962. Catalogo dos peixes do Museu Nacional, Part X. Publica9oes Avulsas do Museu Nacional, 44:1-13. MITCHILL, S. L. 1814. The fishes of New York, described and arranged. Transactions of the Literary and Philosophical Society of New York, 1:355-492. MOBius, K. 1894. Ueber Eiernester Pelagischer Fische aus dem mittelatlantischen Ocean. Sitzungsberichten der Koniglich Preussischen Akademie der Wis-

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. 1921. A list of Turk Island fishes, with a description of a new flatfish. Bulletin of the American Museum of Natural History, 44(3):21-24. . 1929. The fishes of Porto Rico and the Virgin Islands. Branchiostomidae to Sciaenidae. Scientific Survey of Porto Rico and the Virgin Islands, New York Academy of Sciences, 10(2):161-399. . 1937. Syngnathus walcotti, a new West Indian pipefish. American Museum Novitates, 934:1. . 1946. On young western Atlantic seriolas. Copeia, 1946(4):259-260. NICHOLS, J. T., AND C. M. BREDER, JR. 1927. The marine fishes of New York and southern New England. Zoologica, New York, 9(1):1-192. NOBRE, A. 1935. Fauna marinha de Portugal. I. Vertebrados (Mamiferos, Reptis e Peixes). Publica^oes do Institute de Zoologia, Universidade do Porto: 577 PPNOMURA, H., AND N. MENEZES. 1964. Peixes marinhos. In Historia Natural dos Organismos aquaticos do Brasil (pp. 343-385), P. E. Vanzolini ed. Sao Paulo: 452 pp. NORDEN, C. R. 1966. The seasonal distribution of fishes in Vermilion Bay, Louisiana. Transactions of the Wisconsin Academy of Science, Arts & Letters, 55:119-137. NORMAN, J. R. 1937. Coast fishes. Part II. The Patagonian region. Discovery Reports, 16:1-150. . 1948. A history of fishes. A. A. Wyn, New York: 463 pp. NORMAN, J. R., AND P. H. GREENWOOD. 1975. A history of fishes. Third (revised) edn. J. Wiley, New York: 467 pp. OSBECK, P. 1757. Dagbok ofver en Ostindisk resa aren 1750-52, med anmarkningar uti naturkundigheten, frommande folkslags sprak, etc. L. Grefing, Stockholm: 376 pp. . 1765, Reise nach Ostindien und China aus dem Schwedischen iibersetzt von J. G. Georgi. J. C. Koppe, Rostock: xxvi -I- 552 pp. . 1771. A voyage to China and the East Indies. Translation by J. R. Forster. Benjamin White, London: 2:1-147. OVERSTREET, R. M. 1969. Digenetic trematodes of marine teleost fishes from Biscayne Bay, Florida. Tulane Studies in Zoology, 15(4): 119-175. PAIVA CARVALHO, J. DE. 1941. Nota preliminar sobre a fauna ictiologica do littoral sul do Estado de Sao Paulo. Boletim de Industria Animal, 150:27-81. PALACIO, F. J. 1974. Peces colectados en el Caribe Colombiano por la Universidad de Miami. Boletin Museo del Mar, Bogota, 6:1-137. PARKER, J. C. 1965. An annotated checklist of the fishes of the Galveston Bay system, Texas. Publications of

the Institute of Marine Science, University of Texas, 10:201-220. PARKER, J. C., B. J. GALLAWAY, AND D. MOORE. 1972. Key to the estuarine and marine fishes of Texas. Texas A.&M. University: 177 pp. PARR, A. E. 1930. Teleostean shore and shallow-water fishes from the Bahamas and Turks Island. Bulletin of the Bingham Oceanographic Collection, 3(4):1-148. PEARCY, W. G., AND S. W. RICHARDS. 1962. Distribution and ecology of fishes of the Mystic River estuary, Connecticut. Ecology, 43(2):248-259. PEARSON, J. C. 1941. The young of some marine fishes taken in lower Chesapeake Bay, Virginia, with special reference to the gray sea trout Cynoscion regalis (Bloch). Fishery Bulletin, 50:79-102. PELLEGRIN, J. 1914. Missions Gruvel sur la cote occidentale d'Afrique (1905-1912). Poissons. Annales de 1'Institut Oceanographique, 6(4):l-99. PENRITH, M. J. 1976. Distribution of shallow water marine fishes around southern Africa. Cimbebasia, Windhoek, Series A, 4(7): 137-154. PEQUENO R., G. 1975. Vision del estudio biologico de los peces marinos chilenos. Revista de Estudios del Pacifico, Valparaiso, 9:37-56. PERLMUTTER, A. 1961. Guide to marine fishes. New York University Press: 431 pp. PETERS, W. 1869. Ueber eine neue Nagergattung, sowie iiber einige neue oder weniger bekannte Amphibien und Fische. Monatsbericht der K. Preussichen Akademie der Wissenschaften zu Berlin, 1868:448-460. PHILIPPI, R. A. 1896. Peces nuevos de Chile. Anales de la Universidad, Memorias Cientificas i Literarias, Santiago, 93-95:375-390. PHILLIPS, R. C., AND V. G. SPRINGER. 1960. A report on the hydrography, marine plants and fishes of the Caloosahatchee River area, Lee County, Florida. Florida State Board of Conservation, Special Scientific Report, 5:1-34. POEY, F. 1868. Synopsis piscium Cubensium. Repertorio fisico-natural de la isla de Cuba, Habana, 2:279-468. . 1876. Enumeratio piscium Cubensium. Parte tercera. Anales de la Sociedad Espanola de Historia Natural, Madrid, 5:373-404. POLL, M. 1949. Poissons. Resultats scientifiques des Croisieres du Navire-Ecole Beige "MERCATOR," 4(4):173-269. POWELL, C. R., AND K. STRAWN. 1963. Notes on the fringed pipefish, Micrognathus crinigerus, from the west coast of Florida. Publications of the Institute of Marine Science, University of Texas, 9:112-116. POWELL, D., L. M. SWINELL, AND S. E. SWINELL. 1972.

Fishes of the Western North Atlantic An annotated listing of the fish reference collection at the Florida Department of Natural Resources Marine Research Laboratory. Florida Department of Natural Resources, Special Scientific Report, 36:1-179. Pozzi, A. J., AND L. F. BORDALE. 1935. Cuadro sistematico de los peces marinos de la Republica Argentina. Anales de la Sociedad Cientifica Argentina, Buenos Aires, 120:145-189. Pozzi, A. J., AND E. M. SICCARDI. 1948. Descripcion del alotipo de Syngnathus folletti Herald, 1942 (Pisces, Syngnath.). Comunicaciones del Institute Nacional de Investigation de las Ciencias Naturales y Museo Argentine de Ciencias Naturales 'Bernardino Rivadavia/ Ciencias Zoologicas, 8:1-8. QUIJADA, B. B. 1912. Catalogo de la coleccion de los peces Chilenos i estranjeros del Museo Nacional. Boletin del Museo Nacional de Chile, 4:69-109. . 1913. Catalogo ilustrado i descriptive de los peces Chilenos i estranjeros conservados en el Museo Nacional. Boletin del Museo Nacional de Chile, 5(1):7-130. RAFINESQUE, C. S. 1810. Caratteri di alcuni nuovi generi e nuove specie di animali (principalmente de pesci) e piante della Sicilia, con varie osservazioni spora i medisimi. Palermo: 105 pp. . 1815. Analyse de la nature, ou tableau de 1'univers et des corps organises. Palermo: 224 pp. RANDALL, J. E. 1968. Caribbean reef fishes. T. F. H. Publications, Jersey City: 318 pp. RAY, C., AND E. CIAMPI. 1956. The underwater guide to marine life. A. S. Barnes, New York: 338 pp. REEVES, C. D. 1927. A catalogue of the fishes of northeastern China and Korea. Journal of the Pan-Pacific Research Institution, 2(3):3-16. REGAN, C. T. 1906. On the freshwater fishes of the island of Trinidad based on the collection, notes and sketches made by Mr. Lechmere Guppy, Jr. Proceedings of the Zoological Society of London, 1906(l):378-393. . 1908. Pisces. In Godman, F. Due., and O. Salvin, Biologia Centrali-Americana. London. Part 193, 203 pp. REID, G. K., JR. 1954. An ecological study of the Gulf of Mexico fishes in the vicinity of Cedar Key, Florida. Bulletin of Marine Science of the Gulf and Caribbean, 4(1): 1-94. . 1955. A summer study of the biology and ecology of East Bay, Texas. Part II. The fish fauna of East Bay, the Gulf beach and summary. Texas Journal of Science, 7(4):430-453. RENFRO, W. C. 1960. Salinity relations of some fishes in the Aransas River, Texas. Tulane Studies in Zoology, 8(3):83-91.

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California Academy of Sciences, Second series, 6:245-267. RYDER, J. A. 1887. On the development of osseous fishes, including marine and freshwater forms. Report of the United States Fish Commission, 13:489-605. SABINS, D. S. 1973. Diel studies of larval and juvenile fishes of the Caminada Pass area, Louisiana. M.S. thesis, Texas A.&M. University. SAUVAGE, H. E. 1882. Description de quelques poissons de la collection du Museum d'histoire naturelle. Bulletin de la Societe Philomathique de Paris, Septieme serie, 6:168-176. . 1891. Histoire naturelle des poissons. In A. Grandidier, Histoire physique, naturelle et politique de Madagascar. Imprimerie National, Paris, 16:1-543. SAWYER, R. T., A. R. LAWLER, AND R. M. OVERSTREET. 1975. Marine leeches of the eastern United States with a key to the species. Journal of Natural History, 9:633-667. SCHAEFER, R. H. 1967. Species composition, size and seasonal abundance of fish in the surf waters of Long Island. New York Fish and Game Journal, 14(1): 1-46. SCHNEIDER, J. B. 1801. Systema Ichthyologiae iconibus ex illustratum. S. Berolina: Ix + 584 pp. SCHREINER, C, AND A. MIRANDA RlBEIRO. 1903. A COl-

leccao de peixes do Museu Nacional do Rio de Janeiro. Archives do Museu Nacional, Rio de Janeiro, 12:67-109. SCHULTZ, L. P. 1949. A further contribution to the ichthyology of Venezuela. Proceedings of the United States National Museum, 99:1-211. SCHWARTZ, F. J. 1961. Fishes of Chincoteague and Sinepuxent bays. American Midland Naturalist, 65(2):384-408. . 1964. Fishes of Isle of Wight and Assawoman bays near Ocean City, Maryland. Chesapeake Science, 5(4):172-193. SCHWARTZ, F. J., AND H. J. PORTER. 1977. Fishes, macroinvertebrates, and their ecological interrelationships with a calico scallop bed off North Carolina. Fishery Bulletin, 75(2):427-446. SCOTT, E. O. G. 1939. Observations on some Tasmanian fishes. Part IV. Papers and Proceedings of the Royal Society of Tasmania, 1938:139-159. . 1942. Syngnathus tuckeri sp. nov.: a new Tasmanian pipefish. Records of the Queen Victoria Museum, Launceston, 1(1): 17-20. . 1955. Observations on some Tasmanian fishes. Part VII. Papers and Proceedings of the Royal Society of Tasmania, 89:131-146. SHARP, B., AND H. W. FOWLER. 1904. The fishes of Nantucket. Proceedings of the Academy of Natural Sciences of Philadelphia, 56:504-512.

SHAW, G. 1804. General zoology or systematic natural history. Pisces. London, 5(2):451-457. SHIINO, S. M. 1972. List of English names of Japanese fishes with proposition of new names. Science Report of Shima Marineland, 1:1-210. . 1976. List of common names of fishes of the world, those prevailing among English-speaking nations. Science Report of Shima Marineland, 4:1-262. SICCARDI, E. M. 1954. Consideraciones sobre el modo de reproduccion de "Leptonotus blaenvillanus" Eydoux & Gervais, 1837 (Pisces; Syngnath.). Comunicaciones del Institute Nacional de Investigacion de las Ciencias Naturales y Museo Argentine de Ciencias Naturales "Bernardino Rivadavia." Ciencias Zoologicas, 2(14):211-242. SILVESTER, C. F. 1916. Fishes new to the fauna of Porto Rico, Carnegie Institution of Washington Yearbook, 14:214-217. . 1918. Fishes new to the fauna of Porto Rico, with descriptions of eight new species. Papers from the Department of Marine Biology of the Carnegie Institution of Washington, 12:19-24. SIMMONS, E. G. 1957. An ecological survey of the upper Laguna Madre of Texas. Publications of the Institute of Marine Science, University of Texas, 4(2):156-200. SLONAKER, J. R. 1897. A comparative study of the area of acute vision in vertebrates. Journal of Morphology, 13:455-502. SMITH, B. A. 1971. The fishes of four low-salinity tidal tributaries of the Delaware River estuary, M.S. thesis, Cornell University. Not seen. SMITH, H. M. 1898. The fishes found in the vicinity of Woods Hole. Bulletin of the United States Fish Commission, 17:85-111. . 1907. The fishes of North Carolina. North Carolina Geological and Economic Survey, Raleigh, 2:1-453. SMITH, H. M., AND B. A. BEAN. 1899. List of fishes known to inhabit the waters of the District of Columbia and vicinity. Bulletin of the United States Fish Commission, 18:179-187. SMITH, H. M., AND W. C. KENDALL. 1898. Notes on the extension of the recorded range of certain fishes of the United States coasts. Report of the United States Fish Commission, 22:169-176. SMITH, J. L. B. 1949. The sea fishes of southern Africa. Central News Agency, Cape Town: 550 pp. (also, 1953, 1961 and 1965 editions). SMITH, J. L. B., AND M. SMITH. 1963. The fishes of Seychelles. Department of Ichthyology, Rhodes University: 215 pp. SMITH-VANIZ, W. F. 1968. Freshwater fishes of Alabama. Auburn University: vii + 211 pp.

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on the animal communities in two north Florida salt marshes. Part 1. Fish communities. Bulletin of Marine Science, 25(4):445-465. SUMNER, F. B., R. C. OSBURN, AND L. J. COLE. 1913.

A

biological survey of the waters of Woods Hole and vicinity. Part II. Section III. A catalogue of the marine fauna. Bulletin of the United States Bureau of Fisheries, 31:549-782. SWAIN, J. 1882. A review of the Syngnathinae of the United States, with a description of one new species. Proceedings of the United States National Museum, 5:307-315. SWAIN, J., AND S. E. MEEK. 1884. Notes on the pipefishes of Key West, Florida, with description of Siphostoma McKayi, a new species. Proceedings of the United States National Museum, 7:237-239. SWIFT, C., R. W. YERGER, AND P. R. PARRISH. 1977. Distribution and natural history of the fresh and brackish water hshes of the Ochlockonee River, Florida and Georgia. Bulletin of Tall Timbers Research Station, Tallahassee, 30:1-111. SWINGLE, H. A. 1971. Biology of Alabama estuarine areas—Cooperative Gulf of Mexico estuarine inventory. Alabama Marine Research Bulletin, 5:1-123. SWINGLE, H. A., AND D. G. BLAND. 1974. A study of the hshes of the coastal watercourses of Alabama. Alabama Marine Research Bulletin, 10:17-102. TABB, D. C., D. L. DUBROW, AND R. B. MANNING. 1962. The ecology of northern Florida Bay and adjacent estuaries. Florida State Board of Conservation, Technical Series, 39:1-81. TABB, D. C., AND R. B. MANNING. 1961. A checklist of the flora and fauna of northern Florida Bay and adjacent brackish waters of the Florida mainland collected during the period July, 1957 through September, 1960. Bulletin of Marine Science of the Gulf and Caribbean, ll(4):552-649. TAGATZ, M. E. 1968. Fishes of the St. Johns River, Florida. Quarterly Journal of the Florida Academy of Sciences, 30(1):25-50. TARGETT, T. E., AND J. D. MCCLEAVE. 1974. Summer abundance of hshes in a Maine tidal cove with special reference to temperature. Transactions of the American Fisheries Society, 103(2):325-330. THOMPSON, W. F. 1916. Fishes collected by the United States Bureau of Fisheries steamer "ALBATROSS" during 1888, between Montevideo, Uruguay and Tome, Chile, on the voyage through the Straits of Magellan. Proceedings of the United States National Museum, 50:401-416. TOMMASI, L. R. 1964. Sobre tres especies de peces marinos del litoral de San Pablo (Brasil). Neotropica, Buenos Aires, 10(31):29-35. TORTONESE, E. 1970. Osteichthyes (Pesci Ossei). Parte Prima. Fauna d'ltalia, Bologna, 10:xiii -I- 565 pp.

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TRACY, H. C. 1910. Annotated list of fishes known to inhabit the waters of Rhode Island. Annual Report of the Commissioners of Inland Fisheries (Rhode Island), 40:35-176. TRUITT, R. V., B. A. BEAN, AND H. W. FOWLER. 1929. The fishes of Maryland. State of Maryland Conservation Bulletin, 3:1-120. TURNAGE, D. 1964. A survey of fishes in Lake Palourde, Louisiana. Proceedings of the Louisiana Academy of Sciences, 27:8-11. UHLER, P. R., AND O. LUGGER. 1876. List of fishes of Maryland. Report of the Commissioner of Fisheries of Maryland, 1876:67-176. VAILLANT, L. 1888. Poissons. In Mission scientifique du Cap Horn, 1882-1883. Zoologie. Paris, 6:1-35. VERGARA RODRIGUEZ, R. 1974. Nuevos registros para la ictiofauna Cubana, II. Poeyana, 134:1-7. VLADYKOV, V. D., AND R. A. MCKENZIE. 1935. The marine fishes of Nova Scotia. Proceedings of the Nova Scotian Institute of Science, 19(1):17-113. WAITE, E. R. 1907. A basic list of the fishes of New Zealand. Records of the Canterbury Museum, ---

. 1911. Scientific results of the New Zealand Government trawling expedition of 1907. Pisces, Part II. Records of the Canterbury Museum, l(3):157-272. WALLS, J. G. 1975. Fishes of the northern Gulf of Mexico. T. F. H. Publications, Neptune City: 432 pp. WANG, J. S. C., AND E. C. RANEY. 1971. Distribution and fluctuations in the fish fauna of the Charlotte Harbor estuary, Florida. Charlotte Harbor Estuarine Studies, Mote Marine Laboratory: 56 pp. WARFEL, H. E., AND D. MERRIMAN. 1944. Studies on the marine resources of southern New England. 1. An analysis of the fish population of the shore zone. Bulletin of the Bingham Oceanographic Collection, WEBER, M., AND L. F. DE BEAUFORT. 1922. The fishes of the Indo- Australian Archipelago. IV. Heteromi, Solenichthyes, Synentognathi, Percesoces, Labyrinthici, Microcyprini. E. J. Brill, Leiden, 4:1-410. WEINSTEIN, M. P., AND K. L. HECK. 1979. Ichthyofauna of seagrass meadows along the Caribbean coast of Panama and in the Gulf of Mexico: Composition, structure and community ecology. Marine Biology, 50:97-107. WEYMOUTH, F. W. 1910. Notes on a collection of fishes from Cameron, Louisiana. Proceedings of the United States National Museum, 38:135-145. WHATLEY, E. C. 1962. Occurrence of breeding Gulf pipefish, Syngnathus scovelli, in the inland fresh waters of Louisiana. Copeia, 1962(1):220. - . 1969. A study of Syngnathus scovelli in fresh waters of Louisiana and salt waters of Mississippi. Gulf Research Reports, 2(4):437-474.

WHEATLAND, S. B. 1956. Oceanography of Long Island Sound, 1952-54. VII. Pelagic fish eggs and larvae. Bulletin of the Bingham Oceanographic Collection, 15:234-314. WHEELER, A. C. 1975. Fishes of the world. MacMillan, New York: 366 pp. WHITAKER, J. O. 1968. Keys to the vertebrates of the eastern United States, excluding birds. Burgess, Minneapolis: iii -I- 225 pp. WHITLEY, G. P. 1948. New sharks and fishes from Western Australia. The Australian Zoologist, ll(3):259-276. . 1951. Studies in ichthyology No. 15. Records of the Australian Museum, 22(4):389-408. . 1955. Sidelights on New Zealand ichthyology. The Australian Zoologist, 12(2):110-119. WHITWORTH, W. R., P. L. BERRIEN, AND W. T. KELLER. 1968. Freshwater fishes of Connecticut. Bulletin of the State Geological and Natural History Survey of Connecticut, 101:vi-t- 134 pp. WlCKLUND, R. L, S. J. WlLK, AND L. OGREN. 1968.

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servations on wintering locations of the northern pipefish and spotted seahorse. Underwater Naturalist, 5(2):26-28. WIEGMANN, W. H., AND J. T. NICHOLS. 1915. Notes on fishes near New York. Copeia, 1915(23):43-45. WILLIAMS, G. C. 1960. Dispersal of young marine fishes near Woods Hole, Massachusetts. Publications of Michigan State University, Biological Series, l(10):329-368. WILSON, C. 1916. Some marine fishes from Colombia and Ecuador. Annals of the Carnegie Museum, 10:57-70. WILSON, H. V. 1900. Marine biology at Beaufort. The American Naturalist, 34:339-360. YARROW, H. C. 1877. Notes on the natural history of Fort Macon, N.C. and vicinity (No. 3). Proceedings of the Academy of Natural Sciences of Philadelphia, 29:203-218. ZARET, T. M., AND R. T. PAINE. 1973. Species introduction in a tropical lake. Science, 182(4lll):449-455. ZHUDOVA, A. M. 1969. Materials on the study of the eggs and larvae of some species of fish from the Gulf of Guinea and the adjacent waters of the open ocean. Trudy Atlanticheskii Nauchino-Issledovatelskii Institut Rybnogo Khoziaistva i Okeanografii, 22:135-163. (Translated from Russian by W. L. Klawe, Inter-American Tropical Tuna Commission, La Jolla). ZUGMAYER, E. 1911. Poissons provenant des campagnes du yacht "Princesse Alice" (1901-1910). Resultats des Campagnes Scientifiques accomplies sur son yacht par Albert 1", Prince souverain de Monaco, 35:1-174.

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FIGURES 48-51. 48. Amphelikturus dendriticus: adult male (BOC 2532), holotype of A. brachyrhynchus, Bahamas. 49. Amphelikturus dendriticus: juvenile (GCRL 4235), 33 mm, Gulf of Mexico. 50. Penetopteryx nanus: lectotype (ZML 5a), female, 31.7 mm, Bahamas. 51. Penetopteryx nanus: adult male (AMNH 26032), 22 mm, Bahamas.

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53

FIGURES 52-54. 52. Penetopteryx nanus: planktonic "pawneei" form (BOC 3327), 22 mm, holotype of Ichthyocampus pawneei, Bahamas. 53. Penetopteryx nanus: photomicrograph of body section of pouch larva (ca. 3.8 mm) showing dorsal-fin rays, pterygiophores and ventral fin-fold. 54. Oostethus brachyurus lineatus: female, 119 mm (GCRL 3892), Mississippi.

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FIGURES 55-61. 55. Oostethus brachyurus lineatus: adult male, 140 mm (GCRL 3892), Mississippi. 56. Oostethus brachyurus lineatus: adult male (GCRL 13359), 120 mm, Brazil. 57. Oostethus brachyurus lineatus: adult male (GCRL 13396), 121 mm, ventral aspect of partially filled brood pouch, Brazil. 58. Micrognathus crinitus: female (GCRL 10979), 85 mm, Brazil. 59. Same as 58. 60. Micrognathus crinitus: female (FSM 18678), 89 mm, "ensenadae" color pattern, East Florida. 61. Micrognathus erugatus: holotype (MZUSP 9408), female, 64 mm, Brazil.

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FIGURES 62-67. 62. Anarchopterus criniger: female (GCRL 2800), 88 mm, Louisiana. 63. Anarchopterus criniger: adult male (GCRL 15256), 88.5 mm, West Florida. 64. Anarchopterus tectus: holotype (USNM 216592), female, 92 mm, East Florida. 65. Leptonotus blainvilleanus: adult male, 159 mm (GCRL 16257), Argentina. 66. Same as 65. 67. Leptonotus blainvilleanus: mature female, 174.5 mm (GCRL 16257), Argentina.

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FIGURES 68-74. 68. Pseudophallus starksii: adult male (GCRL 11873), 106 mm, Panama, Pacific drainage. 69. Pseudophallus starksii: adult male (GCRL 15646), 133 mm, Nicaragua, Pacific drainage. 70. Pseudophallus starksii: mature female (GCRL 12461), 98 mm, Panama, Pacific drainage. 71. Pseudophallus mindii: male (GCRL 12775), 114 mm, Atlantic drainage of Panama. 72. Pseudophallus elcapitanensis: male (GCRL 11874), 120 mm, Pacific drainage of Panama. 73. Syngnathus floridae: male (GCRL 14824), 103.5 mm, North Carolina. 74. Syngnathus floridae: mature female (GCRL 15242), 172 mm, East Florida.

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FIGURES 75-81. 75. Syngnathus floridae: adult male (GCRL 14908), 186 mm, West Florida. 76. Syngnathus floridae: mature female (GCRL 15250), 201 mm, Texas. 77. Syngnathus springeri: adult male (GCRL 15720), 284 mm, West Florida. 78. Syngnathus springeri: juvenile (GCRL 13850), 135 mm, West Florida. 79. Syngnathus springeri: juvenile (GCRL 4258), 104 mm, Mississippi. 80. Syngnathus louisianae: adult male (GCRL 13256), 265 mm, West Florida. 81. Syngnathus louisianae: juvenile (GCRL 14136), 170 mm, Mississippi.

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FIGURES 82-87. 82. Syngnathus louisianae: adult female (GCRL 13887), 225 mm, East Florida. 83. Syngnathus fuscus: juvenile (GCRL 14825), 110 mm, North Carolina. 84. Syngnathus fuscus: adult male (GCRL 12424), 222 mm, Georgia. 85. Syngnathus scovelli: adult male (GCRL 14100), 114 mm, West Florida. 86. Syngnathus scovelli: mature female (GCRL 14902), 102 mm, Alabama. 87. Syngnathus scovelli: mature female (GCRL 15527), 101 mm, Venezuela.

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FIGURES 88-93. 88. Syngnathus affinis: holotype (BMNH 1854.7.3.2), male, 144 mm, Louisiana. 89. Syngnathus affinis: female (GCRL 15252), 125 mm, Texas. 90. Syngnathus makaxi: adult male, 45 mm, paratype (CAS 13617), Mexico. 91. Syngnathus makaxi: mature female, 41.5 mm, paratype (CAS 13617), Mexico. 92. Syngnathus folletti: adult male, 158 mm (GCRL 16355), inshore benthic, Brazil. 93. Syngnathus folletti: mature female, 133mm (GCRL 16355), inshore benthic, Brazil.

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FIGURES 94-100. 94. Syngnathus folletti: adult female (MHNM 638), 179.5 mm, offshore benthic, Uruguay. 95. Syngnathus folletti: adult male (GCRL 16259), 74 mm, offshore planktonic, Brazil. 96. Syngnathus dawsoni: holotype (USNM 203148), adult female, 135.5 mm, Virgin Islands. 97. Syngnathus dawsoni: adult male (AMNH 26294), 131 mm, Virgin Islands. 98. Syngnathus dawsoni: juvenile or female (ANSP 113002), 91.5 mm, Martinique. 99. Syngnathus pelagicus: adult male (GCRL 16073), 144 mm, Northwestern Atlantic. 100. Syngnathus pelagicus.: mature female (USNM 219901), 139.5 mm, Northwestern Atlantic.

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FIGURES 101-107. 101. Syngnathus canbbaeus: paratype (GCRL 15525), adult male, ca. 190 mm, Venezuela. 102. Syngnathus caribbaeus: paratype (GCRL 1956), female, 170 mm, Panama. 103. Bryx dunckeri: adult male, 78 mm. 104. Same as 103. 105. Bryx dunckeri: adult female, 82.5 mm (GCRL 15743), Virgin Islands. 106. Bryx randalli: holotype (USNM 198903), female, 98 mm, Venezuela. 107. Cosmocampus albirostris: female (GCRL 15967), 109 mm, West Florida.

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FIGURES 108-113. 108. Cosmocampus albirostris: adult male (GCRL 16073), 155 mm, Alabama. 109. Cosmocampus albirostris: juvenile (UPR 3267), 74 mm, Puerto Rico. 110. Cosmocampus profundus: adult male (UMML 26978), 191 mm, off Yucatan, Mexico. 111. Cosmocampus brachycephalus: adult male (GCRL 10126), 68 mm, Panama. 112. Cosmocampus brachycephalus: adult female (GCRL 14545), 68 mm, Panama. 113. Cosmocampus elucens: adult male (GCRL 15745), 120 mm, Virgin Islands.

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FIGURES 114-118'. 114. Cosmocampus elucens: adult male (GCRL 15750), 122 mm, Virgin Islands. 115. Cosmocampus elucens: mature female (GCRL 11163), 121 mm, Panama. 116. Cosmocampus elucens: female (UMML 23933), 103 mm, Panama. 117. Cosmocampus hildebrandi: holotype (USNM 117251), female, 86 mm, West Florida. 118. Cosmocampus hildebrandi: adult male (GCRL 15966), 50.5 mm, West Florida.

THE SEAHORSES (Subfamily Hippocampinae) RICHARD P. VARI Department of Vertebrate Zoology, National Museum of Natural History Smithsonian Institution, Washington, D.C. 20560

Acknowledgments. I thank the following for information, permission to examine specimens in their care, and prompt response to loan requests: C. L. Smith and D. E. Rosen (AMNH); J. E. Bohlke and W. F. Smith-Vaniz (ANSP); P. H. Greenwood and G. J. Howes (BMNH); W. N. Eschmeyer and P. M. Sonoda (CAS); D. J. Stewart and R. K. Johnson (FMNH); W. L. Fink and K. E. Hartel (MCZ); M. L. Bauchot (MNHN); and R. Hacker (NMW). R. H. Kanazawa and E. N. Gramblin provided technical assistance in the study and Ms. P. Whorl, secretarial assistance.

commencing immediately posterior to the terminal trunk ring. In the posterior portion of the tail, however, these bony processes become increasingly less distinct. The rings in this region of the tail are, however, demarcated by deep transverse grooves along the ventral surface of the tail, with each segment bounded anteriorly and posteriorly by a groove. Additional shallower intermediate grooves are often present midway along some rings. However, the shallower depth of the latter depressions makes their discrimination from the transverse grooves marking the ring boundaries relatively simple. In some individuals the groove between the ultimate and penultimate tail rings is lacking. Nonetheless, the increased length of the resultant, apparently single, element relative to the preceding ring, and the mobility between its parts allows the discrimination of these two rings. The failure to recognize two elements in such cases is not critical for distinguishing western Atlantic species of seahorses. 3. Ring form. Each ring is composed or four or more segments, with each segment usually, although not always, bearing a prominent central tubercle which forms part of one of the longitudinal ridges extending along the trunk and tail. The majority of trunk segments are septangular with the body planes bounded by the paired superior, lateral and inferior trunk ridges (STRR, LTRR and ITRR) and a median ventral trunk ridge (MVTRR) (see Fig. 1). The latter element is lacking, however, on the ultimate trunk ring. Tail rings are nearly all quadrangular, bearing paired superior and inferior tail ridges (STAR and ITAR). In the region ventral to the dorsal-fin base, the superior tail ridge extends anteriorly two or three segments to intercalate between the superior trunk ridge (STRR) and

Western Atlantic Genera and Species. Three species of seahorses are recognized from the western Atlantic ocean, all in the genus Hippocampus. All three species are primarily tropical and temperate with one species (H. erect us) sporadically occurring to beyond 40°N and 35°S. Records of other nominal Hippocampus species from western Atlantic waters appear to be erroneous or are unconfirmed. TERMINOLOGY AND METHODS Fish Characters. The various morphometric and meristic characters used to distinguish species of seahorses differ notably from those of most other fishes. The terms utilized and their definitions are as follows: 1. Number of trunk rings. The anteriormost trunk ring is that bearing the pectoral fin and the anteriormost tubercle on the lateral trunk ridge (LTRR). The terminal trunk ring is located immediately anterior to the anal fin and is most readily distinguished externally by the posteriormost tubercle of the inferior trunk ridge (ITRR) (see Fig. 1). 2. Number of tail rings. The number of rings along the anterior portion of the tail can be accurately determined by counting the bony spurs or tubercles along the inferior tail ring (ITAR) 173

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LTRR

MVTRR

STAR

FIGURE 1. Exoskeleton of Hippocampus showing morphological features referred to in the text: DFB, dorsalfin base; ITAR, inferior tail ridge; ITRR, inferior trunk ridge; LTRR, lateral trunk ridge; MVTRR, median ventral trunk ridge; STAR, superior tail ridge; STRR, superior trunk ridge; TARI, tail ring; TRRI, trunk ring (after Ginsburg, 1937).

lateral trunk ridge (LTRR) (see Fig. 1). The variation in the anterior extent of the lateral trunk ridge and differences in the posterior limit of the superior trunk ridge result in variable numbers of segments on three rings. The penultimate trunk ring ranges from septangular to novemangular, the ultimate from sexangular to octangular and the first tail ring from quadrangular to sexangular. In some individuals the segment of the ultimate trunk ring forming part of the superior tail ridge lacks a distinct tubercle.

Although this condition results in an apparently single body plane between the superior and lateral trunk ridges, careful examination will reveal the intermediate ossification. 4. Proportional measurements. To determine the total length (TL) of a specimen, the tail is straightened and the head oriented perpendicular to the longitudinal axis of the tail. As a consequence of the relative inflexibility of the trunk it is impossible, however, to eliminate the trunk curvature. Contrary to Ginsburg (1937), the total length is measured from the tip of the tail to the median groove of the coronet at the tip of the head. Measuring to the latter point rather than to the terminal point of the coronet reduces the variation introduced by differing degrees of ontogenetic development of the dorsolateral coronet processes. Trunk length is the distance between the median groove of the coronet and the midlateral point intermediate to the boundaries of the terminal trunk and first tail rings. Tail length is the distance between the latter point and the tail tip. Greatest body depth is measured between the two deepest trunk rings. The measurement is the minimum distance at that point between the superior trunk ridge (STRR) and the median ventral trunk ridge (MVTRR). Head length (HL) is measured from the tip of the snout to the posterodorsal fleshy margin of the gill opening. Snout length is the distance from the snout tip to the anterior margin of the bony orbit. Orbital diameter is measured between the anterior and posterior bony ridges bordering the orbit. The postorbital length is the distance from the bony ridge along the posterior margin of the eye to the posterodorsal margin of the gill opening. In contrast to Ginsburg (1937:508), subunits of the head are expressed as percentages of head length rather than total length. This method reduces the variation introduced by difficulties in straightening specimens and by differing degrees of contractability in body parts. Length of the dorsal-fin base is measured between the anterior and posterior angles of insertion of the fin. Greatest body depth, trunk length and tail length are sexually dimorphic and are consequently segre-

Fishes of the Western North Atlantic gated by sex in the species descriptions. The ranges presented for all other measurements are inclusive of both sexes. 5. Fin rays. All fin rays are counted separately even when closely approximated basally. Pectoral-fin ray counts were taken from the right fin except when it was damaged. Synonyms and References. Synonymies are as complete as possible with all available type series examined. Where feasible, the specimens that served as the basis for literature citations were examined. Distribution Maps. The distribution maps show the distribution of all material examined although not necessarily all collections from the immediate vicinity of each symbol. Literature records of a species from beyond the geographic range of the material examined are noted in the Range section for the species. Genus Hippocampus Rafinesque 1810 Hippocampus Rafinesque, 1810:18 (type-species Syngnathus hippocampus Linnaeus 1758, by original designation). Macleayina Fowler, 1907:426 (type-species Hippocampus bleekeri Fowler 1907 [=Hippocampus abdominalis Lesson 1827], by original designation). Parlapiscus Whitley, 1931:313 (type-species Hippocampus breviceps Peters 1870, by original designation). Hippohystrix Whitley, 1940:414 (type-species Hippocampus spinosissimus Weber 1913, by original designation). Jamsus Ginsburg, 1937:584 (type-species Hippocampus regulus Ginsburg 1933, by original designation).

Characters. Superior trunk and tail ridges discontinuous and overlapping under dorsal-fin base, lateral trunk ridge confluent with inferior tail ridge, inferior trunk ridge ends at ultimate trunk ring, median ventral trunk ridge terminates at penultimate trunk ring. Points of inter-

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section of longitudinal trunk and tail ridges with transverse rings usually bearing tubercles. Tubercles usually more prominent in juveniles, often elongate and spinous, becoming varyingly reduced in adults. Ridges on tail and associated tubercles less prominent posteriorly. Dermal flaps often extending from body tubercles and head ridges and spines, usually more prominent in juveniles, sometimes branching. Head angled distinctly ventrad relative to principal body axis, moveable in median plane over a considerable angle. Snout moderate to long, no prominent mid-dorsal snout ridge; internasal and supraopercular spines variously developed. Spination on coronet usually more prominent in juveniles, reduced or absent in some species. Trunk rings 9 to 13, penultimate trunk ring septangular to novemangular, ultimate trunk ring hexangular to septangular, remaining trunk rings septangular. Tail prehensile, tail rings 28 to 49, first ring quadrangular to hexangular, remaining rings quadrangular. Dorsal-fin rays 10 to 31, dorsal-fin origin on 2 to 5 trunk and 1 or 2 tail rings, fin base elevated. Anal-fin rays 3 to 5. Pectoralfin rays 10 to 19. Caudal fin absent. Brood pouch located entirely under tail, pouch a sealed saclike structure with an anteroventral pore-like opening, pouch lacks ventrolateral extensions of lateral dermal plates. Size. Small fish, reaching 300 mm TL. Range. Atlantic, Indian and Pacific oceans and associated seas, primarily between 30°N and 30°S. Species. As a consequence of our poor understanding of seahorse taxonomy, it is not possible to accurately estimate the number of species worldwide. Approximately two dozen species are currently recognized.

Key to Western Atlantic Species of Hippocampus la. Dorsal-fin rays 10-13; pectoral fin rays 10-13 zosterae Jordan and Gilbert 1882, p. 176 Ib. Dorsal-fin rays 16-20; pectoral fin rays 14-17 2 2a. Head and body pigmentation in the form of distinct dark round spots of varying sizes, snout length .41-.49 of head length reidi Ginsburg 1933, p. 178

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2b. Head and body pigmentation in the form of blotches, streaks and lines, snout length .3S-.46 of head length erectus Perry 1810, p. 181 Hippocampus zosterae Jordan and Gilbert 1882 Dwarf seahorse Figures 2-4 Hippocampus zosterae Jordan and Gilbert, 1882:265 (orig. descr.; Laguna Grande, Pensacola, Florida; holotype USNM 30852 [missing]); Bean, 1883:44 (Pensacola, Florida); Jordan, 1887:850 (North America); Jordan and Evermann, 1896:778, PL 121, Fig. 328 (Pensacola Bay, Florida); Evermann and Kendall, 1900:63 (Florida: Pensacola, Cape Florida and Key West); Weymouth, 1910:137 (Cameron, Lousiana); Breder, 1929:105 (Pensacola, Florida); Jordan et al., 1930:244 (Pensacola Bay, Florida); Ginsburg, 1937:589 (Florida: Cape Florida, Biscayne Bay, Newfound Harbor Key, Key West, Boca Chica, Captiva Pass, Pepperfish Key, Apalachicola Bay, St. Martins, Pensacola); Breder, 1940:137 (birth of young); Kanazawa, 1952:78 (Bermuda); Strawn, 1954:215 (breeding); Hoese, 1958:238 (Texas); Strawn, 1958:16 (life history); Starck, 1958:18 (Alligator Reef, Florida); Briggs, 1958:267 (compiled); Springer and Woodburn, 1960:30 (Boca Ciega Bay, Florida); Tabb and Manning, 1961:617 (ecology); Breder and Rosen, 1966:298 (compiled; reproduction); Bohlke and Chaplin, 1968:182 (Hog Island, Bahamas); Duarte-Bello and Buesa, 1973:90 (Cuba); Guitart, 1975:280 (Cuba); Hoese and Moore, 1977:162 (Gulf of Mexico). Hippocampus regulus Ginsburg, 1933:563 (orig. descr.; Harbor Island, Texas; holotype USNM 92950); Hubbs, 1936:282 (Campeche, Mexico); Ginsburg, 1937:585, Figs. 70, 71 (Texas: Harbor Island, Hog Island; Campeche, Mexico); Reid, 1954:27 (Cedar Key, Florida); Briggs, 1958:267 (compiled); Hoese, 1958:328 (=H. zosterae); Strawn, 1958:16 (=H. zosterae). Hippocampus rosamondae Borodin, 1928:16. PL 1, Fig. 3 (orig. descr.; Cuba, syntypes VMM, not examined); Ginsburg, 1937:586 (=H. zosterae).

Distinctive Characters. The possession of 10 to 13 dorsal-fin rays and 10 to 13 pectoral-fin rays distinguishes this species from all other western Atlantic Hippocampus species. Description. Proportional and meristic data from 146 specimens, 15 to 42 mm TL. Rings: 9-11 (11 rare) + 28-34 (34 rare). Subdorsal rings: 1.5-2.5 + 0.0-1.5. Fin rays: Dorsal 10-13 (modally 11, see Table I), pectoral 10-13, anal 4, rarely 3.

Head. Head length .19-.25 of TL, snout length .26-.3S of HL, orbital diameter .18-.23 of HL, postorbital length .42-.51 of HL, dermal flaps variously present, less prominent in adults. Body. Depth between deepest trunk rings .12-. 19 of TL in females, .14-.21 of TL in males; trunk length .S7-.47 of TL in females, .S4-.44 of TL in males; tail length .S3-.63 of TL in females, .S6-.66 of TL in males; superior trunk ridge and superior tail ridge overlapping on 0-2 rings (modally 1); penultimate trunk ring septangular (octangular in 1 specimen), ultimate trunk ring sexangular or octangular, rarely septangular; first caudal ring quadrangular or sexangular; dermal flaps variously developed; length of dorsal-fin base .06-.08 of TL; brood pouch of eggcarrying males extending along 5-7 caudal rings. Color. Coloration variable, mottling varies from distinct to absent, without definite pattern when present, masked in some nearly black individuals; life coloration green to black (Hoese and Moore, 1977:162). Dorsal fin with a submarginal band, slightly wider anteriorly, intensity of pigmentation in band highly variable. Size. The largest specimen examined measured 42 mm TL; males with a distinct developing brood pouch at 16 mm TL and with brood-pouch eggs or embryos at 24 mm TL. Development and Growth. Number of eggs in ovaries of females and number of developing young in brood pouch of males increasing with adult size, females with up to 69 large eggs in ovaries, males with up to 55 young in brood pouch. All eggs in brood pouch of a male evidently from a single female. Breeding season mid-February to late October, evidently primarily correlated with day length, partially with water temperature. Developing young carried in brood pouch approximately 10 days. Average male carrying two broods per month. Young expelled from brood pouch by muscular contractions of body and pouch. Juveniles 7-9 mm TL at hatching, growing in captivity to 18 mm TL in 17 days. Growth most rapid in summer when

Fishes of the Western North Atlantic

FIGURE 2. Hippocampus zosterae, male, 28.1 mm TL; Harbor Island, Texas; USNM 92950, holotype of H. regulus (from Ginsburg, 1937).

individuals attain breeding size in three months. Approximately three generations per breeding season. Few individuals surviving to one year, none to two years. Females outnumbering males in population (Strawn, 1958). Habits and Behavior. Inhabits grass flats in shallow waters, periodically found on floating vegetation, most common in bays during periods of high salinity (Tabb and Manning, 1961:617). Food. Small crustaceans. Variation. Ginsburg (1937:585) in his discussion of the nominal forms Hippocampus zosterae and H. regulus noted that no trenchant character separated the species, but recognized them as distinct on the basis of a modal divergence in dorsal-fin ray and tail-ring counts, although

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FIGURE 3. Hippocampus zosterae, female, 28.3 mm TL; Harbor Island, Texas; USNM 124854 (from Ginsburg, 1937).

these intergraded significantly between the species. Examination of a larger series of specimens than available to Ginsburg fails to reveal any sharp break in the distribution of the modal counts of dorsal-fin rays (see Table I). Similarly, although the number of tail rings demonstrates some geographic variation, the differences form a relatively smooth cline rather than demonstrating a bimodal distribution (see Table II). Thus, available data fail to provide a basis for the continued recognition of H. regulus as distinct from H. zosterae. Hippocampus rosamondae of Borodin (1934) is tentatively placed as a synonym of H. zosterae following Ginsburg (1937:589). In the original

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178

TABLE I. Frequency distribution of the number of dorsal-fin rays of Hippocampus zosterae across its range. 10

Locality Bermuda, Bahamas, East coast of Florida West coast of Florida Northern Gulf Coast, Florida to Louisiana Texas Coast

Dorsal-fin rays 12 11

3 19

22 25

12 12

1 9

9 28

1 2

13 1

20"

description the species is described as having 14 dorsal-fin rays, a count closer to the range of the H. zosterae material examined (dorsal-fin rays 10-13) than of H. erectus and H. reidi (16-20). Furthermore, the stated count of 11 H- 30 body rings falls within the range for H. zosterae but not of the other two species. Relationships to Man. Hippocampus zosterae is relatively common in the aquarium trade. Range. This species is known from Bermuda, the Bahamas, Cuba (Guitart, 1975), the east coast of Florida and the coasts of the Gulf of Mexico from the Florida Keys to the Gulf of Campeche. Study Material. 146 specimens, 15 to 42 mm TL. BERMUDA: FMNH 49019 (1, 36.4); FMNH 49020 (4, 31.8-36.0). BAHAMAS: ANSP 88677 (5, 26.5-36.0); ANSP 88676 (1,32.0). FLORIDA: USNM143070 (1, 24.2); USNM 176006 (1, 26.2); USNM 26595 (1, 29.7); USNM 31920 (4, 35.5-41.0); SU (CAS) 1671 (1, 28.5, paratype of H. zosterae)', USNM 176166 (8, 19.8-26.8); USNM 57438 (1, 28.5); USNM 57453 (2, 29.5-30.0); USNM 57236 (2, 33.0); USNM 176167 (5, 33.3-35.7); AMNH 39169 (2, 39.0-42.5); AMNH 39171 (1, 42.2); USNM 132633 (1,

TABLE II. Frequency distribution of the number of tail rings of Hippocampus zosterae across its range. Locality Bermuda, Bahamas, East coast of Florida West coast of Florida Northern Gulf Coast, Florida to Louisiana Texas coast

Tail rings 28 29 30 31 32 33 34

1

1 6 10 11 10 15 20 9 3 4 3 8 17 11

100

8 1

FIGURE 4. Hippocampus zosterae, geographic distribution based on material examined. 31.7); USNM 132631 (1, 30.2); USNM 132630 (1, 35.8); USNM 132638 (1, 30.0); USNM 132635 (2, 33.5-34.3); USNM 176171 (5,28.3-34.4); USNM 176187 (5,34.4-40.4); USNM 176189 (5, 29.8-35.4); USNM 176185 (5,30.3-37.1); USNM 184201 (4,30.3-37.5); USNM 176172 (4,26.0-31.3); USNM 176173 (5, 27.7-33.0); USNM 107014 (1, 29.0); AMNH 39170 (4,25.5-26.3); USNM 176170 (5,28.1-30.0); USNM 176183 (5, 25.4-30.5). MISSISSIPPI: USNM 132629 (4, 19.2-20.5); USNM 218799 (1, 20.0). TEXAS: USNM 92950 (1, 28.1, holotype of H. regulus); USNM 176179 (6, 21.5-25.7); USNM 176182 (8, 23.7-31.7); USNM 124854 (1, 28.3); USNM 124857 (1, 27.8); USNM 124858 (1, 21.7); USNM 124855 (1, 27.3); USNM 132627 (1, 30.2); USNM 176176 (9, 25.5-34.7); USNM 176175 (8, 23.8-30.5); ANSP 129761 (2, —). MEXICO: USNM 176169 (4, 20.7-25.2).

Hippocampus reidi Ginsburg 1933 Longsnout seahorse Figures 5-7 Hippocampus longirostris. Kaup, 1856:12, PI. 3, Figs. 2, 2a (misidentification, Martinique and St. Lucia). Hippocampus guttulatus. Goode, 1877:291 (misidentification, Bermuda). Hippocampus punctulatus. Starks, 1913:40 (misidentification; Natal, Brazil); Meek and Hildebrand, 1923:255 (misidentification in part, specimens from Porto Bello, Panama). Hippocampus reidi Ginsburg, 1933:561 (orig. descr.; St. George, Grenada, British West Indies; Porto Bello; Panama; Jamaica; Haiti; holotype USNM 86596);

Fishes of the Western North Atlantic

179

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FIGURE 5. Hippocampus reidi, male, holotype, 116.1 mm TL; St. George, Grenada, British West Indies; USNM 86590 (from Ginsburg, 1937).

Beebe and Tee-Van, 1935:318 (Haiti); Ginsburg, 1937:572, Figs. 65, 66 (Porto Bello, Panama; Grenada, British West Indies; Port-au-Spain, Haiti; Jamaica; Bermuda); Fowler, 1939:10 (Jamaica); Caldwell and Caldwell, 1964:15 (Caledonia Bay, Panama); Caldwell, 1966:36 (Jamaica: Kingston, Palisadoes); Cervigon, 1966:236 (Punta de Piedras, Venezuela); Bohlke and Chaplin, 1968:184 (Bahamas); Starck, 1968:18 (Alligator Reef, Florida); Mago, 1970:92 (compiled; Venezuela); Duarte-Bello and Buesa, 1973:90 (Cuba). Hippocampus obtusus Ginsburg, 1933:562 (orig. descr.; off Cape Hatteras, North Carolina; holotype USNM 84527); Ginsburg, 1937:576, Fig. 67 (after Ginsburg, 1933); Kanazawa, 1952:77 (Bermuda); Hardy, 1978:397 (compiled); Robins et al., 1980. Hippocampus poeyi Howell Rivero, 1934:32, fig. (orig. descr.; Havana, Cuba; holotype MCZ 33913); Ginsburg, 1937:561 (=H. hudsonius punctulatus).

FIGURE 6. Hippocampus rtiJftrfeiihhle, 126.8 mm TL; St. George, Grenada, British West Indies; USNM 223673 (from Ginsburg, 1937).

Hippocampus hudsonius punctulatus. Ginsburg, 1937:561 (in part).

Distinctive Characters. The spotted pigmentation, 16-19 dorsal-fin rays, 15-17 pectoral-fin rays, and a snout length (.41-.49 of HL) always longer than postorbital length, in combination distinguish this species from all other western Atlantic Hippocampus species. Description. Proportional and meristic data from 69 specimens, 30 to 175 mm TL. Rings: 11+33-37. Subdorsal rings: 2.0-3.0 + 1.0-2.5. Fin rays: Dorsal 16-19 (modally 17); pectoral 15-17, anal 4, rarely 3. Head. Head length .2S-.27 of TL, snout length .41-.49 of HL, orbital diameter .13-.18 of HL,

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Memoir I, Part 8—Sears Foundation for Marine Research

FIGURE 7. Hippocampus reidi, geographic distribution based on material examined.

postorbital length .35-.41 of HL; tuberculation poorly developed; coronet knob-like, with no prominent dorsolateral projections; no dermal flaps. Body. Depth between deepest trunk rings .11-.15 of TL in females, .13-.17 of TL in males; trunk length .39-.4S of TL in females, .35-.41 in males; tail length .55-.61 of TL in females, .59-.6S of TL in males. Superior trunk ridge and superior tail ridge overlapping on 1 to 3 rings (2 modal, 1 in only 1 specimen); penultimate trunk ring septangular (modally) or novemangular, ultimate trunk ring octangular, first caudal ring sexangular, rarely quadrangular or pentangular (segment of superior tail ridge present only unilaterally in the latter). Tuberculation weakly developed, no strong spination or dermal flaps. Length of dorsal-fin base .07-.10 of TL. Brood pouch in egg-carrying males extending 5-7 tail rings. Color. Head and body covered relatively densely with dark rounded spots of varying sizes against a lighter background, ground col-

oration in larger specimens sprinkled with very small whitish dots, lighter transverse bars often present on trunk and tail. Some specimens nearly black and spotted pattern not evident. Dorsal clear with a series of chromatophores forming a submarginal band and an irregular pattern of spotting at fin base. Size. The largest specimen examined measured 175 mm TL; males with a distinct developing brood pouch at 70 mm TL and with brood-pouch eggs or embryos at 80 mm TL. Discussion. No single character easily separates all specimens of Hippocampus reidi and H. erectus. The markedly different pigmentation patterns readily separate those specimens in which the body coloration is evident. In those cases where the pigmentation is either very faint or masked by a dark overall body coloration, specimens of these two forms are usually easily separable by snout length. In H. reidi the snout is always longer than the postorbital length. In H. erectus, in contrast, snout length is shorter than postorbital length in all but some of the largest specimens. These latter individuals can, however, be identified by their considerably deeper bodies compared to those of adult H. reidi (compare Figs. 6 and 10). Range. Bermuda, Atlantic coasts of North America from Cape Hatteras south, the Bahamas, west coast of Florida, Cuba, Haiti, Jamaica, Caribbean coast of Panama and Colombia, Venezuela, Lesser Antilles, Atlantic coast of South America south to Rio de Janeiro (see Fig. 7); occurring to depths of 55 m. Study Material 68 specimens, 30.5-180 mm TL, including holotype. Holotype. USNM 86590 (116.1 mm, male), St. George, Grenada, British West Indies. Other Material. CAPE HATTERAS: USNM 84527 (1, 65.3, holotype of H. obtusus). BERMUDA: USNM 21933 (1, 130.5); USNM 169958 (1, 30.5). FLORIDA: USNM 222670 (1, 66.5); USNM 142897 (1, 83.0); USNM 93699 (1, —). BAHAMAS: ANSP 88672 (1, —); ANSP 116291 (1, -); AMNH 26116 (1, 53.0). ST. LUCIA: USNM 223667 (1, 98.9); USNM 170162 (1, 103.1). DOMINICA: USNM 170221 (1, 64.5). GUADELOUPE: USNM 132619 (1, 55.2). GRENADA: USNM 223673 (1, 126.8). HAITI: AMNH 19206 (1,180.0); USNM 164770 (1,122.5); USNM

Fishes of the Western North Atlantic 85958 (3, -). CUBA: USNM 171747 (1,95.9); MCZ 33913 (1, 48.7, holotype of H. poeyi); USNM 108446 (15, 68.0-129.5); USNM 108447 (7, 121.3-132.4). PUERTO RICO: ANSP 144650 (3, 97.5-143.0); ANSP 142993 (1, —). JAMAICA: USNM 130481 (1, —); USNM 92684 (3, 48.3-79.8); USNM 93732 (1, 41.7); ANSP 68638 (1, —). DOMINICA: AMNH 36625 (1, -). BRITISH VIRGIN ISLANDS: ANSP 117441 (1, —). "CARIBBEAN SEA": USNM 223638 (1, 94.7). PANAMA: USNM 79685 (1, 86.0). COLOMBIA: USNM 223636 (1, 88.0). TRINIDAD: USNM 170352 (1, 144.0). VENEZUELA: ANSP 104627 (1, 140.0); ANSP 114576 (2, —); ANSP 114575 (2, —): SURINAM: AMNH 26275 (1, 175.0). BRAZIL: AMNH 3860 (2, 160.0-178.0); USNM 131966 (1, 78.5).

Hippocampus erectus Perry 1810 Lined seahorse Figures 8-11 Hippocampus erectus Perry, 1810: PL XLV (orig. descr.; American seas, coasts adjacent to Mexico and the West Indies; no type specimen); Bailey et al., 1960 (common name); Tabb and Manning, 1961:616 (ecology); Linton and Soloff, 1965:45 (brood pouch physiology); Croker, 1965:92 (eggs and larvae, Sandy Hook, New Jersey); Breder and Rosen, 1966:297 (reproduction, sonic ability; compiled); Leim and Scott, 1966:76 (Nova Scotia); Starck, 1968:18 (Alligator Key, Florida); Bohlke and Chaplin, 1968:183 (Great Bahama Bank, Bahamas); Miller and Jorgenson, 1973:310 (meristics); DuarteBello and Buesa, 1973:90 (Cuba); Palacio, 1974:37 (Gulf of Uraba, Colombia); Fahay, 1975:20 (continental shelf, North Carolina to Florida); Hoese and Moore, 1977:162 (Gulf of Mexico); Hardy, 1978:391 (development and life history; compiled); Hastings, 1979:7 (Okaloosa County, Florida); Wang and Kernehan, 1979:157 (Delaware estuary). Hippocampus hudsonius De Kay, 1842:322, PI. 53, Fig. 171 (orig. descr.; Hudson River, New York; no type specimen); Lockwood, 1867:225 (development); Dumeril, 1870:523 (after De Kay); Jordan, 1887:850 (North America); Henshall, 1891:375 (southern Florida); Henshall, 1895:212 (Tampa, Florida); Jordan and Evermann, 1896:777, PL 121, Fig. 327 (Cape Cod to Charleston); Smith, 1898:94 (Vineyard Bay, Massachusetts); Evermann and Kendall, 1900:63 (Florida: Pensacola, St. John's River, Key West, Gulf Stream); Bean, 1902:411 (New York: Gravesend Bay, Southampton, Fire Island); Sherwood and Edwards, 1902:27 (Gayhead, Massachusetts); Bean, 1903:349 (New York); Jordan and Thompson, 1905:235 (Middle Key, Florida); Bean, 1906:40 (Bermuda: Flatts, Cable Bay); Fowler, 1906:233 (New Jersey); Smith, 1907:172 (North Carolina); Fowler, 1907:291 (Cape May, New Jersey); Ken-

181

dall, 1908:65 (Rhode Island, Massachusetts); Fowler, 1908:165 (Delaware Bay); Tracy, 1910:94 (Rhode Island); Evermann and Hildebrand, 1910:160 (Cape Charles); Fowler, 1911:12 (Delaware); Halkett, 1913:74 (Nova Scotia); Sumner, Osburn and Cole, 1913:166 (compiled); Fowler, 1918:18 (Smith Island, Virginia); Fowler, 1920:154 (New Jersey: Cape May, Cumberland and Monmouth Counties); Fowler, 1921:445 (Rhode Island, New Jersey, Delaware); Bigelow and Welsh, 1925:177 (Gulf of Maine); Nichols and Breder, 1927:67 (Long Island, New York); Hildebrand and Schroeder, 1928:185 (Chesapeake Bay); Truitt et al., 1929:59 (Chesapeake Bay); Breder, 1929:104 (Atlantic coast, Charleston to Nova Scotia); Jordan et al., 1930:244 (Cape Cod to Tortugas); Vladykov, 1935:5 (Nova Scotia); Vladykov and McKenzie, 1935:81 (Nova Scotia); Fowler, 1937:306 (New Jersey); Pearson, 1941:85 (Chesapeake Bay); Fish, 1952:98 (sound production); Merriman and Sclar, 1952:165 (Rhode Island); Bigelow and Schroeder, 1953:315 (Gulf of Maine, George's Bank, Nova Scotia); Hildebrand, 1955:205 (Gulf of Campeche); Hoese, 1958:328 (Texas); Springer and Woodburn, 1960:30 (Tampa Bay, Florida); Bergman, 1967:751 (motor nerves); Fish and Mowbray, 1970:19 (sound production). Hippocampus punctulatus Guichenot, 1853:174, PL 5, Fig. 2 (orig. descr.; Cuba; syntypes MNHN 6092); Kner, 1867:390 (Rio de Janeiro, Brazil); Dumeril, 1870:40 (redescription of syntypes); Jordan, 1887:850 (North America); Jordan and Eigenmann, 1888:269 (Charleston, South Carolina); Bean, 1888:134 (Great Egg Harbor, New Jersey); Jordan and Evermann, 1896:777 (in part, North Carolina); Carman, 1896:93 (Key West, Florida); Jordan and Rutter, 1898:98 (Jamaica); Evermann and Kendall, 1906 (Florida: Key West and Tarpon Springs); Evermann and Marsh, 1902:109 (Ponce, Puerto Rico); Jordan and Thompson, 1905:235 (Bush Key, Florida); Bean, 1906:40 (Flatts, Bermuda); Fowler, 1906:234 (New Jersey); Smith, 1907:173 (North Carolina); Miranda Ribeiro, 1915:11 (West Indies to Rio de Janeiro); Wilson, 1916:60 (Cartagena, Colombia); Miranda Ribeiro, 1918:45 (compiled); Metzelaar, 1919:28 (Windward Islands: St. Martin, St. Eustatius). DeVincenzi, 1924:194 (Uruguay: Montevideo and Maldonado); Meek and Hildebrand, 1923:255 (in part, Panama); Devincenzi, 1925:318 (Maldonado, Uruguay); Devincenzi and Baratini, 1928, PL 19, Figs. 5, 6 (Uruguay); Beebe and Tee-Van, 1928:83 (Port-auPrince, Haiti); Nichols, 1929:220 (San Juan, Puerto Rico); Breder, 1929:105 (West Indies to New Jersey); Pozzi and Bordale, 1935:19 (Argentina to 43°30'S). Jordan et al., 1930:244 (in part, West Indies, Brazil, Gulf Stream, North Carolina); Longley and Hildebrand, 1941:65 (Tortugas); Fowler, 1941:147 (compiled, Brazil: Bahia, Natal, Sao Sebastiao, Rio de Ja-

182

Memoir 1, Part 8—Sears Foundation for Marine Research

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^^tv

FIGURE 8. Hippocampus erectus, male, 130 mm TL; Horn Island, Mississippi, FMNH16191 (from Ginsburg, 1937). neiro); Breder, 1949:89 (diurnal coloration changes); Guitart, 1955:280 (Cuba); Ringuelet and Aramburu, 1960:56 (compiled; Argentina). Hippocampus marginalis Kaup, 1856:15 (orig. descr.; Mexico; holotype NMW [missing]); Dumeril, 1870:522 (after Kaup, 1856); Jordan and Evermann, 1896:778 (=H. punctulatus); Ginsburg, 1937:561 (=H. hudsonius punctulatus). Hippocampus fascicularis Kaup, 1856:15 (orig. descr.; Mexico; holotype NMW [missing]); Dumeril, 1870:526 (after Kaup, 1856); Jordan and Evermann, 1896:778 (=H. punctulatus); Ginsburg, 1937:561 (=H. hudsonius punctulatus). Hippocampus laevicaudatus Kaup, 1856:16 (orig. descr.; North America; holotype NMW 75535); Dumeril, 1870:513 (redescription of type); Giinther, 1876:205 (after Kaup, 1856); Jordan and Evermann, 1896:777 (=H. hudsonius hudsonius). Hippocampus brevirostris. Knight, 1866:9 (Nova Scotia). Hippocampus guttulatus. Giinther, 1876:202 (in part, Caribbean). Hippocampus antiquorum. Goode, 1877:291 (Bermuda); Goode, 1878:45 (St. George's Bank); Goode and Bean,

1879:4 (George's Bank); Jones, 1882:95 (Nova Scotia); Ryder, 1882:191 (development). Hippocampus villosus Giinther, 1880:8, PL 1, Fig. D (orig; descr.; off Bahia, Brazil; holotype BMNH 1879.5.14; 464); Miranda Ribeiro, 1915:10 (after Giinther, 1880); Ginsburg, 1937:583 (?Panama); Fowler, 1941:148 (after Giinther, 1880). Hippocampus stylifer Jordan and Gilbert, 1882:265 (orig. descr.; Pensacola Florida; holotype USNM 30876). Jordan and Evermann, 1896:778 (coasts of Florida); Jordan, 1887:850 (North America); Jordan and Thompson, 1905:235 (West Channel, Tortugas, Florida); Bean, 1906:40 (Flatts, Bermuda); Jordan et al., 1930:244 (coasts of Florida); Breder, 1929:105 (Florida); Longley, 1933:293 (=H. punctulatus)', Ginsburg, 1937:561 (=H. hudsonius punctulatus). Hippocampus kincaidi Townsend and Barbour, 1906:305, fig. (orig. descr.; Bermuda; holotype MCZ 29127); Beebe and Tee-Van, 1933:140 (?=H. punctulatus)', Longley, 1933:293 (=H. punctulatus). Hippocampus brunneus Bean, 1907:32 (orig. descr.; Long Bird Island, Bermuda; holotype FMMH 5494); Fowler, 1921:446 (?=H. punctulatus)', Jordan et al., 1930:244 (Bermudas); Longley, 1933:293 (=H. punctulatus)', Ginsburg, 1937:568 (=H. hudsonius kincaidi). Hippocampus hudsonius hudsonius. Ginsburg, 1937:551, Figs. 59-62 (Atlantic coast: Florida to St. George's Bank; Gulf of Mexico: Alabama to Rio Grande). Hippocampus hudsonius punctulatus. Ginsburg, 1937:561, Figs. 63,64 (southern and western Florida, Cuba, Florida Keys); Breder and Edgerton, 1942:145 (locomotion); Herald and Rakowicz, 1951:234 (rearing, life history); Reid, 1954:27 (Cedar Key, Florida); Martin, 1956:96 (Los Roques, Venezuela); Caldwell and Caldwell, 1964:15 (Cubagua Island, Venezuela); Cervigon, 1966:236 (Los Roques, Venezuela); Mago, 1970:92 (compiled, Venezuela); Dahl, 1971:172 (Colombia: Islas del Rosario, Islas de San Bernardo, las Bahias). Hippocampus hudsonius kincaidi. Ginsburg, 1937:568 (Bermuda). Hippocampus erectus hudsonius. Briggs, 1958:267 (compiled); Schwartz, 1961:393 (Maryland: Chincoteague and Sinepuxent Bays). Hippocampus erectus erectus. Briggs, 1958:267 (compiled).

Distinctive Characters. The mottled or striped coloration which lacks a distinctive spotted pattern, 16-20 dorsal-fin rays, 14-17 pectoral-fin rays, snout length .3S-.46 of HL and shorter than the postorbital length in all but the largest specimens, in combination distinguish this species from all other western Atlantic Hippocampus species.

Fishes of the Western North Atlantic

183

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