Order Beloniformes: Needlefishes, Sauries, Halfbeaks, and Flyingfishes: Part 10 9781933789347

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Fishes of the Western North Atlantic

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MEMOIR SEARS FOUNDATION FOR MARINE RESEARCH

Number I

Fishes of the Western North Atlantic

PART TEN

Order Beloniformes

Needlefishes, Sauries, Halfbeaks, and Flyingfishes BELONIDAE SCOMBERESOCIDAE HEMIRAMPHIDAE EXOCOETIDAE

PEABODY MUSEUM OF NATURAL HISTORY, YALE UNIVERSITY Distributed by Yale University Press New Haven and London

Osprey (Pandion haliaetus) with Redfin Needlefish (Strongylura notata) at the J. N. “Ding” Darling National Wildlife Refuge, Florida, U.S.A. Photograph by Gene Helfman. Used with permission.

Fishes of the Western North Atlantic

Series Editors BRUCE B. COLLETTE National Systematics Laboratory National Marine Fisheries Service

THOMAS J. NEAR Yale University

Authors BRUCE B. COLLETTE KATHERINE E. BEMIS National Systematics Laboratory National Marine Fisheries Service

NIKOLAY V. PARIN ILIA B. SHAKHOVSKOY P. P. Shirshov Institute of Oceanology Russian Academy of Sciences

PEABODY MUSEUM OF NATURAL HISTORY, YALE UNIVERSITY Distributed by Yale University Press New Haven and London

Published with assistance from the National Systematics Laboratory of the National Marine Fisheries Service, NOAA Fisheries, and the Peabody Museum of Natural History, Yale University. on the cover: Atlantic Blackwing Flyingfish, Hirundichthys volador, western Atlantic Ocean. Illustrated by Howard E. Hamman.

ISBN 978-1-933789-21-7 (pbk.) ISBN 978-1-933789-34-7 (e-book) © 2019 Peabody Museum of Natural History, Yale University. All rights reserved. This book may not be reproduced, in whole or in part, including illustrations, in any form (beyond that copying permitted by Sections 107 and 108 of the U.S. Copyright Law and except by reviewers for the public press), without the written permission of the publisher. Published by Peabody Museum of Natural History, Yale University P.O. Box 208118, New Haven CT 06520-8118 USA peabody.yale.edu Distributed by Yale University Press NEW HAVEN AND LONDON

Printed in the United States of America Printed on acid-free paper Library of Congress Control Number: 49000120

Dedicated to the memories of three beloved ichthyological colleagues who began this research on beloniforms and helped me understand these fascinating fishes — Fred Berry, Bob Gibbs, and Nik Parin

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 | 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, Squirrelfishes 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, Halfbeaks, and Flyingfishes MEMOIR II The Elementary Chemical Composition of Marine Organisms by A. P. Vinogradov

Table of Contents Foreword xi Introduction xiii Acknowledgments xiii Order Beloniformes. By Bruce B. Collette and Katherine E. Bemis 1 Key to the Western Atlantic Families of Beloniformes 4 Literature Cited 4 Family Belonidae. Needlefishes. By Bruce B. Collette and Katherine E. Bemis 5 Introduction 5 Materials and Methods 6 Key to the Belonidae of the Western Atlantic 7 Genus Ablennes 8 Ablennes hians 8 Genus Platybelone 12 Platybelone argalus argalus 12 Genus Strongylura 18 Strongylura hubbsi 18 Strongylura marina 21 Strongylura notata 30 Strongylura notata notata 31 Strongylura notata forsythia 34 Strongylura timucu 37 Genus Tylosurus 42 Tylosurus acus acus 42 Tylosurus crocodilus crocodilus 48 Literature Cited 54 Family Scomberesocidae. Sauries. By Katherine E. Bemis and Bruce B. Collette 79 Introduction 79 Materials and Methods 80 Key to the North Atlantic Sauries 80 Genus Scomberesox 80 Scomberesox saurus saurus 81 Scomberesox simulans 83 Literature Cited 86 Family Hemiramphidae. Halfbeaks. By Bruce B. Collette and Katherine E. Bemis 89 Introduction 89 Materials and Methods 91 Key to the Hemiramphidae of the Western Atlantic 92 Genus Chriodorus 93 Chriodorus atherinoides 93 Genus Euleptorhamphus 96 Euleptorhamphus velox 96

Table of Contents

Genus Hemiramphus 98 Hemiramphus balao 99 Hemiramphus bermudensis 103 Hemiramphus brasiliensis 105 Genus Hyporhamphus 109 Hyporhamphus brederi 110 Hyporhamphus collettei 112 Hyporhamphus meeki 113 Hyporhamphus mexicanus 117 Hyporhamphus roberti 119 Hyporhamphus roberti roberti 119 Hyporhamphus roberti hildebrandi 122 Hyporhamphus unifasciatus 124 Genus Oxyporhamphus 127 Oxyporhamphus micropterus similis 127 Literature Cited 130 Family Exocoetidae. Flyingfishes. By Nikolay V. Parin, Ilia B. Shakhovskoy, Katherine E. Bemis, and Bruce B. Collette 149 Introduction 149 Materials and Methods 150 Key to the Species of Exocoetidae in the Western Atlantic 151 Genus Cheilopogon 152 Cheilopogon cyanopterus 153 Cheilopogon exsiliens 160 Cheilopogon furcatus furcatus 165 Cheilopogon heterurus 170 Cheilopogon melanurus 176 Cheilopogon pinnatibarbatus 180 Genus Cypselurus 186 Cypselurus comatus 186 Genus Exocoetus 191 Exocoetus obtusirostris 191 Exocoetus volitans 196 Genus Hirundichthys 199 Hirundichthys affinis 199 Hirundichthys rufipinnis 206 Hirundichthys speculiger 209 Hirundichthys volador 214 Genus Parexocoetus 219 Parexocoetus hillianus 219 Genus Prognichthys 223 Prognichthys glaphyrae 224 Prognichthys occidentalis 228 Literature Cited 232 Index

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Foreword The publication of this tenth and final title of the Fishes of the Western North Atlantic is the culmination of an eighty-year effort conceived in New Haven in 1937, when Albert E. Parr, with a gift from Henry Sears, established the Sears Foundation for Marine Research at Yale University to promote research and publication in marine sciences. The Fishes of the Western North Atlantic series, first published in 1948, was from its beginnings intended to synthesize and make accessible the wealth of information in widely scattered published accounts of the fish fauna of the region for both the layman and the specialist. The series has always been collaborative and international in scope, a history reflected in its who’s who of contributors. Along with Parr, the first editorial board consisted of prominent ichthyologists of the day—Charles M. Breder, Jr., Samuel F. Hildebrand, William C. Schroeder, and John Tee-Van—assisted by an advisory board of other renown students of the sea, among them William Beebe, Rolf L. Bolin, William K. Gregory, Carl L. Hubbs, George S. Myers, John T. Nichols, Luis Howell-Rivero, Leonard P. Schultz, and Daniel Merriman, the latter then director of Yale’s Bingham Oceanographic Laboratory, which sponsored basic oceanographic expeditions for research, some with the Woods Hole Oceanographic Institution. Part 10 of the series, Order Beloniformes, follows in this rich tradition of scientific cooperation in the service of excellence. In his preface, Henry Bigelow wrote that the first book “has been written on the premise that it should be useful to those in many walks of life—to those casually or vitally interested in the general phenomena of life in our waters, to the sportsman whose interests are closely associated with pleasure and relaxation, to the fisherman whose livelihood depends upon knowledge of where fishes are gathered together, as well as to the amateur ichthyologist and the professional scientist.” That this and the works that followed remain authoritative studies still valuable and of interest today is a testament to the legacy of the scientific literature even in this age of electronic publishing. The exceptional black and white drawings of the earlier works especially harken back to the heyday of scientific illustration and the unsung artists, many of them women, that provided vital support to this research. With the submission of the Beloniformes manuscript came the opportunity to reissue the already published nine parts of the Sears Memoir I, Fishes of the Western North Atlantic. The Yale Peabody Museum of Natural History is pleased to again make these titles available, many long out of print, through Yale University Press, and to include Memoir II, A. P. Vinogradov’s The Elementary Chemical Composition of Marine Organisms, an historically important work greeted in 1953 as a significant contribution to the science of biogeochemistry. The Yale Peabody Museum is grateful to Tim White, Director of Collections and Operations, and Rosemary Volpe, Publications Manager, for coordinating the project; to Yuki Hoshino, Sloan Wilson, Sally Pallatto, and Nate Uthrup for their technical and production assistance; and to Jean E. Thomson Black, Senior Executive Editor, Jenya Weinreb, Director of Editorial, Design, and Production, and Orna Johnston, Reprints Manager, at Yale University Press, for their efforts in the reissue of these works. The assistance of the National Systematics Laboratory of the National Marine Fisheries Service at the United States National Oceanic and Atmospheric Administration for the publication of this volume is deeply appreciated. Thomas J. Near Yale University July 2018 New Haven, Connecticut

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Introduction Shortly after I was hired by the U.S. Bureau of Commercial Fisheries in 1960, Giles Mead, former director of the National Systematics Laboratory and then chair of the editorial board for the Sears Foundation’s monographic series, asked me to take over preparation of the section on the Hemiramphidae, the halfbeaks. Research began shortly thereafter when Frederick H. Berry (Collette and Anderson 2002) transferred his data and material on halfbeaks to me. We found that there were many problems with inshore halfbeaks identified as Hyporhamphus unifasciatus within the United States and elsewhere in the world. Description of a new species of Hemiramphus from Bermuda was the first product (Collette 1962) of this research. Long delays in publishing the sections of the Fishes of the Western North Atlantic led me to turn my attention to other regions, resulting in publications on the halfbeak fauna of the eastern Atlantic (Collette 1965), Australia and New Zealand (Collette 1974a), the Western Indo-Pacific (Parin, Collette and Shcherbachev 1980), and the Far East (Collette and Su 1986). In 1964, Berry also transferred his material on needlefishes to me and research for the needlefish account began. We collaborated on an evaluation of previous papers on needlefishes (Collette and Berry 1965, 1966). For the western Atlantic species, two problems were uncovered and published: separation of Strongylura timucu from S. marina (Collette 1968) and a description of S. hubbsi from freshwaters of Guatemala and Mexico (Collette 1974d). Research with my graduate student Heidi Banford solved the taxonomic problems with what had been considered a single western hemisphere species, Hyporhamphus unifasciatus, by describing new species from the east coast of the United States, the tropical eastern Pacific, and Bermuda (Banford and Collette 1993, 2001; Banford 2010). Additional delays in the publication of the series allowed me to concentrate on the needlefishes of the eastern Atlantic (Collette and Parin 1970) and South America (Collette 1966, 1974a, 1974b, 1982). Robert H. Gibbs, Jr. began the study of Atlantic flyingfishes (Gibbs and Staiger 1970; Gibbs 1978) and commissioned many illustrations (which we have used in the flyingfishes section here) before directing his research to deep-sea fishes. Nikolay Parin agreed to take on the task and completed several species accounts, publishing on the west-central Atlantic (Parin 2003) and east-central Atlantic flyingfishes for the Food and Agriculture Organization of the United Nations (Parin and Shakhovskoy 2016). The premature deaths of both Gibbs in 1988 (Springer and Collette 1989) and Nikolay Parin in 2012 (Collette 2013) challenged completion of this book. I enlisted the help of Katherine E. Bemis to put together all the material on western Atlantic beloniform fishes as a project for her 10-week internship in the summer of 2014, under the NOAA Ernest F. Hollings Scholarship from the U.S. National Oceanic and Atmospheric Administration, and she has continued to work with me to finish all the species accounts. Ilia B. Shakhovskoy joined us to finish the flyingfishes section; without his help, we could not have completed the volume. Acknowledgments. It is rewarding to complete this book, nearly 60 years in the making, and I am grateful to the many who have been involved in the process. Much of the material for this study was collected in the 1950s and 1960s on cruises of the old Bureau of Commercial Fisheries (now the National Marine Fisheries Service) exploratory fishing vessels Combat, Gill, Oregon, and Delaware. I was privileged to participate in several cruises of the Oregon and Delaware and spent many nights leaning over railings dipnetting halfbeaks, needlefishes, and flyingfishes that were attracted to our underwater light. In particular, I thank our former scientific illustrator at the National Marine Fisheries Service, Mildred H. Carrington, who carefully drew all but one of the figures of halfbeaks and

Introduction needlefishes included here, and Howard E. Hamman, who drew most of the flyingfish illustrations for Bob Gibbs. Heidi Banford volunteered to complete the halfbeak illustrations by drawing the halfbeak that she had named for me, Hyporhamphus collettei. The distribution maps were skillfully made by William E. Bemis from the locality data of specimens examined, which were summarized by Janet Nestlerode, Katherine Bemis, and Ilia Shakhovskoy, or based on published maps by Nikolay Parin and Hubbs and Wisner (1980). Janet also prepared preliminary tables of meristic data. Martin Wiley measured gonad lengths and made egg counts. Ilia Shakhovskoy’s work was partially supported by the Federal Agency for Scientific Organizations (FASO Russia), theme no. 0149-2018-0008. A long series of interns and collection support personnel, especially including Ruth Gibbons, made thousands of radiographs that have been transferred to the radiograph collection in the Division of Fishes at the Smithsonian, under the care of Lisa Palmer. The original handwritten data sheets for western Atlantic needlefishes and halfbeaks have been scanned for permanent deposition in the Division of Fishes (USNM ARC 300,000). All original illustrations are also available through the Fish Division. I thank the curators and collection managers of the many collections of fishes that I have worked in or borrowed from over the last half century. These are too numerous to list here, but that does not diminish my profound gratitude for their cooperation. We are deeply appreciative for the detailed reviews of our manuscript by Lynne Parenti, Michael Oliver, and Greg Watkins-Colwell, each of whom provided useful and insightful comments and suggestions that greatly improved our manuscript. And, finally, we have been guided to successful completion of this project by Rosemary Volpe, publications manager at the Yale Peabody Museum of Natural History. Bruce B. Collette Systematics Laboratory National Marine Fisheries Service Washington, D.C. Literature Cited

———. 1974a. The garfishes (Hemiramphidae) of Australia and New Zealand. Records of the Australian Museum 29(2):11–105. ———. 1974b. Potamorrhaphis petersi, a new species of freshwater needlefish (Belonidae) from the upper Orinoco and Rio Negro. Proceedings of the Biological Society of Washington 87(5):31–40. ——— . 1974c. Geographic variation in the central Pacific halfbeak, Hyporhamphus acutus (Günther). Pacific Science 28(2):111–122. ——— . 1974d. Strongylura hubbsi, a new species of freshwater needlefish from the Usumacinta Province of Guatemala and Mexico. Copeia 1974(3):611–619. ——— . 1982. South American freshwater needlefishes of the genus Potamorrhaphis (Beloniformes: Belonidae). Proceedings of the Biological Society of Washington 95(4):714–747. ——— . 2013. Obituary: Nikolai Vasil’evich Parin (1932– 2012). Copeia 2013(4):768–780. Collette, B. B. and W. Anderson, Jr. 2002. Obituary: Frederick H. Berry, 1927–2001. Copeia 2002(2): 558–567. Collette, B. B. and F. H. Berry. 1965. Recent studies on the needlefishes (Belonidae): an evaluation. Copeia 1965(3):386–392. ——— .1966. Proposed suppression of three nomina oblita in the family Belonidae (Pisces). Z.N.(S.) 1723. Bulletin of Zoological Nomenclature 22(5–6):325–329. Collette, B. B. and N. V. Parin. 1970. Needlefishes (Belonidae) of the eastern Atlantic Ocean. Atlantide Report 11:7–60.

Banford, H. M. 2010. Hyporhamphus collettei, a new species of halfbeak (Hemiramphidae) endemic to Bermuda, with comments on the biogeography of the Hyporhamphus unifasciatus species group. Proceedings of the Biological Society of Washington 123:345–358. Banford, H. M. and B. B. Collette. 1993. Hyporhamphus meeki, a new species of halfbeak (Teleostei: Hemiramphidae) from the Atlantic and Gulf coasts of the United States. Proceedings of the Biological Society of Washington 106:369–384. ——— . 2001. A new species of halfbeak, Hyporhamphus naos (Beloniformes: Hemiramphidae), from the tropical eastern Pacific Ocean. Revista de Biología Tropical 49(suppl 1):39–49. 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. ——— . 1965. Hemiramphidae (Pisces, Synentognathi) from tropical West Africa. Atlantide Report 8: 217–235. ——— . 1966. Belonion, a new genus of freshwater needlefishes from South America. American Museum of Natural History Novitates 2274. 22 pp. ——— . 1968. Strongylura timucu (Walbaum): A valid species of western Atlantic needlefish. Copeia 1968(1):189–192.

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Introduction ———. 1978. Five new species of halfbeaks (Hemiramphidae) from the Indo-West Pacific. Proceedings of the Biological Society of Washington 91(3):731–747. Collette, B. B. and J. Su. 1986. The halfbeaks (Pisces, Beloniformes, Hemiramphidae) of the Far East. Proceedings of the Academy of Natural Sciences of Philadelphia 138(1):250–301. Gibbs, R. H., Jr. 1978. Exocoetidae. In W. Fischer, ed. FAO Species Identification Sheets for Fishery Purposes, Western Central Atlantic (Fishing Area 31), Volume 2. Rome: Food and Agriculture Organization of the United Nations. 32 pp. Gibbs, R. H., Jr. and J. C. Staiger. 1970. Eastern tropical Atlantic flyingfishes of the genus Cypselurus (Exocoetidae). Studies in Tropical Oceanography 4:432–466. Hubbs, C. L. and R. L. Wisner. 1980. Revision of the sauries (Pisces, Scomberesocidae) with descriptions of two new genera and one new species. Fishery Bulletin 77:521–566. Parin, N. V. 2003. Family Exocoetidae, flyingfishes. In K. E. Carpenter, ed. The Living Marine Resources of

the Western Central Atlantic. Volume 2, Bony Fishes. Rome: Food and Agriculture Organization of the United Nations. FAO Species Identification Sheets for Fishery Purposes. American Society of Ichthyologists and Herpetologists Special Publication 5. pp. 1116–1134. Parin, N. V., B. B. Collette, and Y. N. Shcherbachev. 1980. Preliminary review of the marine halfbeaks (Hemiramphidae, Beloniformes) of the tropical IndoWest Pacific. Trudy Instituta Okeanologii 97:7–173. [In Russian with English abstract]. Parin, N. V. and I. Shakhovskoy. 2016. Exocoetidae, flyingfishes. In K.E. Carpenter and N. de Angelis, eds. The Living Marine Resources of the Eastern Central Atlantic. Volume 3, Bony fishes, part 1 (Elopiformes to Scorpaeniformes). Rome: Food and Agriculture Organization of the United Nations. FAO Species Identification Guide for Fishery Purposes. pp. 2132–2156. Springer, V. G. and B. B. Collette. 1989. Obituary: Robert H. Gibbs, Jr. 1929–1988. Copeia 1989(1):245–251.

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Order Beloniformes BRUCE B. COLLETTE and KATHERINE E. BEMIS

The Beloniformes (or Synentognathi) is an order of atherinomorph fishes comprising 2 suborders, 6 families, 37 genera, and at least 230 species (Rosen and Parenti 1981; Collette et al. 1984; Collette 2004). Genetic and morphological data all support the Beloniformes as a monophyletic order (Mirande 2017). All these fishes lack true stomachs (Manjakasy et al. 2009; Iwamatsu 2012). The suborder Adrianichthyoidei contains only the family Adrianichthyidae, which lives in brackish and freshwaters of East and South Asia, and the Indo-Australian Archipelago as far east as Timor (Parenti 2008). This suborder differs from the Exocoetoidei in lacking a lateral line on the body, having the lower pharyngeal bones separate, and in having only four to seven branchiostegal rays instead of nine or more (Collette and Parenti, in press). The suborder Exocoetoidei is divided into two superfamilies with five families (Aschliman et al. 2005): Superfamily Scomberesocoidea (Belonidae and Scomberesocidae) and superfamily Exocoetoidea (Hemiramphidae, Zenarchopteridae, and Exocoetidae), which live in tropical and subtropical marine, fresh, and brackish water habitats. Features common to Exocoetoidei include body elongate, subcylindrical or slightly compressed, dorsal and anal fins on the rear half of the body, pelvic fins abdominal with six soft rays, no fin spines, lateral line running along the ventral edge of the body, nasal organ an open pit with a protruding tentacle and lower pharyngeal bones fused into a triangular plate (leading to the name Synentognathi). Historically (Schlesinger 1909; Nichols and Breder 1928), and before inclusion of the Adrianichthyoidei (Rosen and Parenti 1981; Collette et al. 1984), the Scomberesocidae and Belonidae formed the superfamily Scomberesocoidea or Microsquamati, and the Exocoetidae and Hemiramphidae were grouped together as the Exocoetoidea or Macrosquamati (Fig. 1.1). Three derived characters diagnose the superfamily Scomberesocoidea: presence of a pre-

maxillary lateral line canal; interdigitation of bony processes from each side of the lower jaw (Collette et al. 1984; also E. M. G. Kokkelmans, “The functional morphology and phylogenetic implications of lower jaw structure of the Scomberesocoidea,” unpublished manuscript), unique among teleosts; and upper jaw at least slightly elongate. Other diagnostic characters include a third pair of upper pharyngeal bones separate, fourth upper pharyngeal bone usually present, and scales on the body small (Collette et al. 1984). The superfamily Exocoetoidea is diagnosed by one derived character and three inferred losses: third pair of upper pharyngeal bones united (but not fused) into a plate; fourth upper pharyngeal bone lost; and postorbital branch of canalis frontalis and secondary bony canals on the head lost (Parin and Astakhov 1982). Other diagnostic characters include scales on body large, lower jaw frequently elongate, but upper jaw never significantly elongate; premaxillary canal lost (Collette et al. 1984; Fig. 1.2) and no interdigitation between left and right halves of the lower jaw (Kokkelmans, unpublished manuscript). Analysis of molecular data (Lovejoy et al. 2004) indicates a different grouping of genera than the classic four families; e.g., Scomberesocidae with Belone and Petalichthys of the Belonidae and Zenarchopteridae closer to the Belonidae rather than the Hemiramphidae (Lovejoy et al. 2004; Aschliman et al. 2005; Fig. 1.3). Development has long been of interest in Beloniformes (Schlesinger 1909; Nichols and Breder 1928; Collette et al. 1984; Lovejoy 2000; Lovejoy et al. 2004; Collette 2005). Most beloniform fishes produce large spherical eggs with attaching filaments, characters they share with other atherinomorph fishes (Rosen and Parenti 1981). Freshwater and estuarine genera of halfbeaks in the Asian family Zenarchopteridae practice internal fertilization and three genera are viviparous (Meisner and Collette 1999). Adrianichthyid eggs are the smallest (1.0–1.5 mm in diameter),

Fishes of the Western North Atlantic

Figure 1.1. Early ideas on the phylogeny of Beloniformes. From Nichols and Breder (1928: fig. 171). Microsquamati = Scomberesocoidei; Macrosquamati = Exocoetoidei.

Figure 1.2. Cladogram of the Beloniformes. From Collette et al. (1984: fig. 184).

followed by exocoetids (generally 1.5–2.0 mm), hemiramphids (typically 1.5–2.5 mm), scomberesocids (slightly elliptical, 1.5–2.5 mm), and belonid eggs that are generally the largest (most 3.4 mm). The eggs typically have a homogeneous yolk and a relatively small perivitelline space. The incubation period is relatively long in exocoetoids (Kovalevskaya 1982). With a pattern similar to that reported for egg size, belonids hatch at the largest sizes (6.8–14.4 mm) followed by hemiramphids (4.8–11 mm), scomberesocids (at least as small as 6.0–8.5 mm), exocoetids (3.5–6.1 mm), and adrianichthyids (3.5–4.5 mm) (Collette et al. 1984). Fins generally begin to form during the embryonic stages or soon after hatching. Caudal, dorsal, and anal fins generally form first, followed by the pectorals, and lastly the pelvics (except in exocoetids, in which the pectorals form last). During postembryonic development, exocoetoids undergo complex changes in barbel development, beak length, melanistic dorsal fin lobe, body bars, and pelvic fin pigmentation, features important in both phyloge-

ny reconstruction and identification of species. Jaw evolution has been of particular interest to several investigators, most recently Boughton et al. (1991), Lovejoy (2000), and Lovejoy et al. (2004). Representatives of four of the five families of Exocoetoidei occur in the western North Atlantic: Belonidae (4 genera, 8 species); Exocoetidae (6 genera, 16 species); Hemiramphidae (5 genera, 12 species), and Scomberesocidae (1 genus, 2 species). Most beloniform species are tropical epipelagic marine fishes, but several genera of Belonidae and Hemiramphidae are restricted to freshwaters and a few genera contain estuarine and freshwater as well as marine species. We include a freshwater species of Strongylura (S. hubbsi) and two freshwater species of Hyporhamphus (H. mexicanus and H. brederi), because they are freshwater relatives of largely marine genera. We exclude the three freshwater endemic South American genera (Belonion, Potamorrhaphis, and Pseudotylosurus), which have been reviewed elsewhere (Collette 1966, 1974, 1982). 2

Order Beloniformes

Figure 1.3. Phylogeny of Beloniformes, based on total molecular evidence (Rag2, Tmo, 16S, cyt b). From Lovejoy et al. (2004:369, fig. 1). Numbers above branches are decay indices; numbers below branches are bootstrap proportions.

3

Fishes of the Western North Atlantic Key to the Western Atlantic Families of Beloniformes 1a. Both upper and lower jaws elongate; scales small. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Neither jaw or only lower jaw elongate; scales large. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2a. Jaws slightly elongate with small, weak teeth; several finlets present behind dorsal and anal fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sauries, Scomberesocidae 2b. Both jaws very long with larger, sharp teeth; finlets absent. . . . . . . . . . . . . . . . . . . . . . . Needlefishes, Belonidae 3a. Lower jaw usually elongate; pectoral and pelvic fins not enlarged (except in Oxyporhamphus in which adults have short lower jaws and long pectoral fins). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halfbeaks, Hemiramphidae 3b. Neither jaw elongate; pectoral and pelvic fins usually enlarged (except in Exocoetus). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Flyingfishes, Exocoetidae

Literature Cited

Kovalevskaya, N. V. 1982. Reproduction and development of flying fishes of the family Exocoetidae. Voprosy Ikhtiologii 22(4):582–587. [In Russian, English translation in Journal of Ichthyology 22(4):48–54.] Lovejoy, N. R. 2000. Reinterpreting recapitulation: systematics of needlefishes and their allies (Teleostei: Beloniformes). Evolution 54:1349–1362. Lovejoy, N. R., M. Iranpour, and B. B. Collette. 2004. Phylogeny and jaw ontogeny of beloniform fishes. Integrative and Comparative Biology 44(5):366–377. Manjakasy, J. M., R. D. Day, A. Kemp, and I. R. Tibbetts. 2009. Functional morphology of digestion in the stomachless piscivorous needlefishes Tylosurus gavialoides and Strongylura leiura ferox (Teleostei: Beloniformes). Journal of Morphology 270(10):1155–1165. Meisner, A. D. and B. B. Collette. 1999. Generic relationships of the internally-fertilized southeast Asian halfbeaks (Hemiramphidae: Zenarchopterinae). In B. Séret and J.-Y. Sire, eds. Proceedings of the Fifth Indo-Pacific Fish Conference; Nouméa, New Caledonia; 3–8 November 1997. Paris: Société Française d’Ichtyologie and Institut de Recherche pour le Développement. pp. 69–76. Mirande, J. M. 2017. Combined phylogeny of ray-finned fishes (Actinopterygii) and the use of morphological characters in large-scale analyses. Cladistics 33(4):333–350. Nichols, J. T. and C. M. Breder, Jr. 1928. An annotated list of the Synentognathi with remarks on their development and relationships; collected by the Arcturus. Zoologica: Scientific Contributions of the New York Zoological Society 8(7):423– 448. Parenti, L. R. 2008. A phylogenetic analysis and taxonomic revision of ricefishes, Oryzias and relatives (Beloniformes, Adrianichthyidae). Zoological Journal of the Linnean Society 154(3):494–610. Parin, N. V. and D. A. Astakhov. 1982. Studies on the acoustico-lateralis system of beloniform fishes in connection with their systematics. Copeia 1982(2):276–291. Rosen, D. E. and L. R. Parenti. 1981. Relationships of Oryzias, and the groups of atherinomorph fishes. American Museum of Natural History Novitates 2719. 22 pp. Schlesinger, G. 1909. Zur Phylogenie und Ethologie der Scombresociden. Verhandlungen der Zoologisch-Botanischen Gesellschaft Wien 59:302–339.

Aschliman, N. C., I. R. Tibbetts, and B. B. Collette. 2005. Relationships of sauries and needlefishes (Teleostei: Scomberesocoidea) to the internally fertilizing halfbeaks (Zenarchopteridae) based on the pharyngeal jaw apparatus. Proceedings of the Biological Society of Washington 118(2):416–427. Boughton, D. A., B. B. Collette, and A. R. McCune. 1991. Heterochrony in jaw morphology of needlefishes (Teleostei: Belonidae). Systematic Zoology 40(3):329–354. Collette, B. B. 1966. Belonion, a new genus of freshwater needlefishes from South America. American Museum of Natural History Novitates 2274. 22 pp. ——— . 1974. South American freshwater needlefishes (Belonidae) of the genus Pseudotylosurus. Zoologische Mededelingen Leiden 48(16):169–186. ——— .1982. South American freshwater needlefishes of the genus Potamorrhaphis (Beloniformes: Belonidae). Proceedings of the Biological Society of Washington 95(4):714–747. ——— . 2004. Family Hemiramphidae Gill 1859—halfbeaks. California Academy of Sciences Annotated Checklists of Fishes 22. 35 pp. ——— . 2005. Chapter 75, Order Beloniformes. In W. J. Richards, ed. Early Stages of Atlantic Fishes: An Identification Guide for the Western Central North Atlantic, Volume 1. Boca Raton, FL: CRC Press. p. 903. Collette, B. B., G. E. McGowen, N. V. Parin, and S. Mito. 1984. Beloniformes: development and relationships. In National Marine Fisheries Service, National Oceanic and Atmospheric Administration; H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Kendall, Jr., and S. L. Richardson, eds. Ontogeny and Systematics of Fishes: based on an international symposium dedicated to the memory of Elbert Halvor Ahlstrom. [Lawrence, KS]: American Society of Ichthyologists and Herpetologists. Special Publication 1. pp. 335–354. Collette, B. B. and L. R. Parenti. Order Beloniformes. In P. C. Heemstra, E. Heemstra, and D. A. Ebert, eds. Coastal Fishes of the Western Indian Ocean. In press. Iwamatsu, T. 2012. Growth of the Medaka (I)—formation of vertebrae, changes in blood circulation, and changes in digestive organs. Bulletin of Aichi University of Education 61:55–63.

4

Family Belonidae NEEDLEFISHES

BRUCE B. COLLETTE and KATHERINE E. BEMIS

Introduction The Belonidae, the needlefishes, is one of the six families of the order Beloniformes. Along with its sister group, the Scomberesocidae, the sauries, it forms the superfamily Scomberesocoidea (Collette et al. 1984). Characters of the pharyngeal jaw apparatus support this sister group relationship (Aschliman et al. 2005). The belonids are diagnosed by one derived reductive character (Collette et al. 1984): interruptions in the cephalic lateralis system. Other diagnostic characters include no finlets following dorsal and anal fins; both upper and lower jaws usually elongate and studded with relatively large sharp teeth; pectoral branch of lateral line present; and post-temporal forked. Their long jaws give most needlefishes (and the distantly related gars, Lepisosteidae) the fastest opening and closing mandibles yet measured in fishes (Westneat 2004). Fish prey is usually swallowed and digested head first. Needlefishes are unusual in that they are carnivorous but lack a stomach and have a straight short gut (Manjakasy et al. 2009). Needlefishes show significantly lower α-amylase activity than halfbeaks, but other digestive enzymes were not substantially different (Day et al. 2011). The dorsal and anal fins are posterior on the body near the caudal fin, yet needlefishes swim with anguilliform motion (Liao 2002). The bones, scales, and fin rays are green due to the presence of the bile pigment biliverdin (Çaglar 1945, 1950). The Belonidae contains 10 genera and 35 species (Collette 2003b). Three genera are monotypic: the southern African Petalichthys and the broadly distributed Ablennes and Platybelone. The Asian freshwater Xenentodon contains two species, as does the eastern Atlantic Belone. Three genera are restricted to freshwaters of South America (Lovejoy and Collette 2003): Pseudotylosurus (two species), Potamorrhaphis (four), and Belonion (two). Tylosurus contains six strictly marine species; Strongylura comprises 14 species, some marine, some estuarine, and three strictly freshwater species. The genera Belone and Petalichthys are the most

generalized (Collette and Berry 1965; Parin 1967), having well-developed gill rakers, large scales, comparatively weak canine teeth, and other generalized characters. Belone also is characterized by the most completely developed cranial lateralis system (Parin and Astakhov 1982). Of the other genera, the greatest number of primitive characters are found in Platybelone, which is also characterized by several specialized features (in particular, a depressed caudal peduncle). The remaining three genera of marine needlefishes (Strongylura, Ablennes, and Tylosurus) are more advanced, but Collette and Berry (1965), Parin (1967), and Astakhov (1980) have interpreted their relationships differently. The three South American freshwater genera of needlefishes are secondarily freshwater fishes that were presumably derived from marine needlefishes. The first review of American needlefishes was by Jordan and Fordice (1887). They recognized only two genera: Tylosurus, for the marine species, and Potamorrhaphis, for one South American freshwater species. They recognized 16 species, 12 in the western Atlantic (although one of the western Tylosurus was another freshwater species, Pseudo­ tylosurus microps) and four in the eastern Pacific. Today the needlefish fauna of the western Atlantic (Collette 2003a) includes seven marine species in four genera: Platybelone (one species), Ablennes (one species), Tylosurus (two species), and Strongy­ lura (three species). There is also a freshwater species of Strongylura, Strongylura hubbsi, which lives in rivers draining into the Gulf of Mexico in Mexico and Guatemala and is included in this account. The three endemic South American freshwater genera have been covered adequately in a series of earlier publications (Belonion by Collette [1966]; Pseudotylo­ surus by Collette [1974a]; Potamorrhaphis by Collette [1974b, 1982], Lovejoy and Collette [2003], Sant'Anna et al. [2012]). There are some valid published fossil records from the Americas. These include dentaries, premaxillae, and a vertebra from Florida, U.S.A., referred to Belonidae (Hulbert 2001), which is perhaps closely related

Fishes of the Western North Atlantic and T. acus (see Collette and Parin 1970: fig. 12) and Ablennes hians have bars on the posterior part of the body. These bars are retained in adult Ablennes, as is the posterior dorsal fin lobe. Interestingly, analysis of molecular data supports Ablennes as the sister taxon of Tylosurus (Lovejoy et al. 2004). Relationships to Humans. Needlefishes are valued food fishes in many parts of the world, including Australia, India, the West Indies, and South America, but are not currently considered an important food fish resource in the United States. All needlefishes that have been evaluated by the IUCN (International Union for the Conservation of Nature) have been considered Least Concern. Needlefishes are known to have caused injuries, particularly to fishermen in tropical regions (Barss 1982). A recent study described five cases of injuries caused by needlefishes penetrating the eyes, ears, face, or legs of people in the Arabian Gulf, two of which resulted in fatalities (Jawad 2016).

to Strongylura, dated at ca. 2.5 million years old from the late Pliocene, from the Richardson Road Shell Pit 6A and the Macasphalt Shell Pit 1A sites in Sarasota County, and a partial skull and several vertebrae referred to S. marina, late Pleistocene, probably less than 500,000 years old, from the Fort Thompson Rock Pit, Palm County (R. C. Hulbert, pers. comm.). †Belone countermani Sant’Anna et al., 2013 was described from the Miocene St. Marys Formation of Calvert Cliffs, MD, and is the only record of this genus for the western Atlantic. This species was described from a well-preserved three-dimensional pair of upper and lower jaws and is characterized by a unique dental pattern on the dentary (Sant’Anna et al. 2013). Needlefish eggs are generally larger (2.3–4.3 mm in diameter) than other beloniform eggs (Collette et al. 1984: table 90) and the filaments on belonid eggs are typically long, numerous, and uniformly spaced over the chorion. In at least one species, Strongylu­ ra strongylura, some filaments are markedly longer than others, as in the adrianichthyids. During post-embryonic development, needlefishes, like other beloniform fishes, undergo complex changes in the beak length, melanistic dorsal fin lobe, and body bars. Most species of Belonidae pass through a “halfbeak” stage in ontogeny in which the lower jaw, but not the upper jaw, is greatly elongate (Boughton et al. 1991). Juveniles of Belone belone remain in the halfbeak stage for a longer time than do other needlefishes. Petalichthys and Platybelone also remain in the halfbeak stage for a long time. The duration of this stage differs among species of Strongylura. Comparative development of Platybelone (as Strongylura longleyi), Strongylura marina, S. notata, and two species of Tylosurus (T. acus and T. crocodilus) was illustrated by Breder (1932a: figs. 7, 10, pl. 1, 2). Tylosurus crocodilus lacks a halfbeak stage: upper and lower jaws grow at the same rate from larval to adult stages of development (Breder 1932a: pl. 2, fig. 2, as T. raphidoma). The South American freshwater genus Belonion (maximum observed body length size of 42 mm) matures while still in the halfbeak stage and was considered paedomorphic by Collette (1966). Ablennes and Tylosurus share a prominent enlarged melanistic lobe in the posterior part of the dorsal fin (Figs. 2.1, 2.12, 2.14). Other genera of needlefishes have no trace of this posterior dorsal lobe. Breder (1932a: pl. 3–5) illustrated the development of this posterior lobe in Tylosurus acus and T. crocodilus and noted its absence in Strongylura and Platybelone. The lobe is apparently sloughed off in T. crocodilus (Breder and Rasquin 1952), resorbed in T. acus (Breder and Rasquin 1954), and retained in adult Ablennes. Juveniles of two species of Tylosurus, T. gavialoides

Materials and Methods Counts and measurements follow papers by Collette (e.g., Collette and Parin 1970) and are summarized here. All elements in the dorsal, anal, and pectoral fins were counted. Vertebrae were counted from radiographs and recorded as “precaudal + caudal = total” where available (the total is given for all species). Predorsal scales were counted from the origin of the dorsal fin anteriorly to the posterior end of the head. Meristic data is summarized by species in Table 2.1. Morphometric data recorded to distinguish species include the following: total length (TL); standard length (SL); body length (BL), end of opercle to caudal base (jaws of belonids are frequently broken, so standard length is not always measurable); head length, from the tip of upper jaw to posterior end of opercular membrane; lower jaw extension, from the tip of upper jaw to tip of lower jaw; P1 length, from the base of uppermost pectoral ray to tip of longest ray; P1–P2, distance from base of uppermost pectoral ray to base of anteriormost pelvic ray; P2–C, distance from base of anteriormost pelvic ray to caudal base (C); length of dorsal fin base; and length of anal fin base. Ratios of several of these measurements have proved particularly useful: P1–P2:P2–C; head length in standard length; lower jaw length in head length; pectoral and pelvic fin length in standard length; length of anal fin base in dorsal fin base. Standard two-letter postal codes are used for U.S. locations. Standard symbolic codes for institutions housing museum specimens at the time they were examined follow Sabaj (2016). 6

Family Belonidae Key to the Belonidae of the Western Atlantic 1a. Gill-rakers present, 7–14; caudal peduncle strongly depressed, with well-developed lateral keel . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platybelone argalus

1b. Gill-rakers absent, caudal peduncle rounded, not strongly depressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Dorsal and anal fins very long, almost confluent with very small caudal fin; dorsal rays 27–43; anal rays

24–39 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Potamorrhaphis (Freshwater, South America)

2b. Dorsal and anal fins much shorter, well separated from a much larger caudal fin; dorsal rays 11–27; anal

rays 12–29. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Upper jaw short, as in halfbeaks; pectoral rays 5 or 6. . . . . . . . . . . . Belonion (Freshwater, South America) 3b. Upper jaw elongate in adults, although it may not be as long as lower jaw; pectoral rays 8–14. . . . . . . . . 4 4a. Body strongly compressed and marked with a series of 12–14 dark vertical bars; pectoral fins falcate

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ablennes hians

4b. Body rounded in cross section; vertical bars absent; pectoral fins rounded or pointed, but not falcate. . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5a. Caudal fin distinctly forked, with lower lobe longer than upper lobe; narrow raised dark lateral keel on

caudal peduncle; juveniles with an expanded black lobe in posterior part of dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tylosurus … 6

5b. Caudal fin rounded or truncate; no keel on caudal peduncle; no posterior black dorsal lobe at any size

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

6a. Vertebrae 90–95; dorsal rays 22–26, usually 23 or 24; anal rays 22–26; dorsal and anal fin lobes relatively

low compared to body length (10.5–13.3 and 9.7–11.7 times in BL, respectively); pectoral and pelvic fins relatively short (8.0–12.4 and 10.0–14.1 times in BL, respectively); upper and lower jaw teeth straight at all sizes; left gonad absent or greatly reduced. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tylosurus acus

6b. Vertebrae 79–86; dorsal rays 20–24; anal rays usually 19 or more; dorsal and anal fin lobes relatively

high compared to body length (5.4–10.6 and 5.5–8.0 times in BL, respectively); pectoral and pelvic fins relatively long (6.6–8.3 and 7.3–10.6 times in BL, respectively); upper and lower jaw teeth point distinctly anterior in juveniles; left gonad distinct, although shorter than right. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tylosurus crocodilus

7a. Body lateral line with long dorsal and ventral side branches; caudal peduncle strongly depressed;

caudal fin divided into two lobes, with upper lobe equal to or longer than lower lobe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudotylosurus (Freshwater, South America)

7b. Lateral line without side branches; caudal peduncle rounded; caudal fin rounded or truncate . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strongylura … 8

8a. Maxilla completely covered by preorbital; anal fin rays 12–15; predorsal scales 76–110; vertebrae 53–61

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strongylura notata … 9

8b. Maxilla exposed posteriorly; anal fin rays 16–20; predorsal scales 120–305; vertebrae 64–77 . . . . . . . . 10 9a. Predorsal scales usually 90 or less (x̅ 84.2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strongylura notata notata 9b. Predorsal scales usually more than 90 (x̅ 95.3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strongylura notata forsythia 10a. Predorsal scales 120–185. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strongylura timucu 10b. Predorsal scales 205–305 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11a. Vertebrae 69–77; only right gonad present in males and females. . . . . . . . . . . . . . . . . . . Strongylura marina 11b. Vertebrae 64–68; only right ovary present in females, males with both testes. . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strongylura hubbsi (Freshwater, Guatemala and Mexico)

7

360–430 No data 103–128 116.5 216–293 240.0 205–305 256.0 76–96 84.2 76–110 95.3 120–185 156.0 320–389 347.3 240–290 260.5

Genus Ablennes Jordan and Fordice, 1887 Athlennes Jordan and Fordice, 1887:324, 358–359 (original description as subgenus of Tylosurus, type by monotypy Belone hians Valenciennes in Cuvier and Valenciennes 1846). Ablennes Jordan and Fordice, orthography corrected by decision of the International Commission on Zoological Nomenclature (ICZN 1912).

93–97 70–74 64–68 69–77 56–61 53–59 69–75 90–95 80–84

94.8 71.6 66.4 72.1 58.5 56.4 71.7 91.8 82.3

Diagnosis. Ablennes differs from all other genera of Belonidae in its greatly compressed body, presence of prominent dark vertical bars on the body (Fig. 2.1), and its porous pre-orbital bone. Ablennes lacks the branch of the lateral line ascending to the base of the pectoral fin that is present in the other genera (except the South American freshwater Pseudotylosurus and Belonion). The pectoral fin is falcate in Ablennes, whereas it is pointed or rounded in other needlefishes. Ablennes is similar to Tylosurus in having a deeply forked caudal fin and many pectoral and dorsal fin rays and vertebrae. Both genera have a melanistic posterior lobe in the dorsal fin in juveniles; it is lost in adult Tylosurus and retained in Ablennes. Ablennes resembles Strongylura in lacking a narrow keel on the caudal peduncle, in having more anal than dorsal fin rays, and in having a distinct halfbeak stage in juveniles. Are these similarities of Ablennes to Tylosurus due to phylogenetic relationship or convergence (Collette and Parin 1970:40)? With molecular data (Lovejoy et al. 2004), Ablennes clusters with species of Tylosurus. Species. Ablennes contains only one species, Ablennes hians (Collette 2003b). Range. Marine, widespread in tropical and subtropical waters (Collette 2003b).

26.4 18.2 18.0 18.0 14.3 13.0 17.8 21.5 20.5

13–15 10–12 10–11 10–12 10–12 10–12 10–12 13–14 13–15

13.8 10.9 10.9 11.4 10.1 9.8 11.0 13.2 14.2

60–63 44–47 40–43 41–50 35–40 34–38 44–49 60–64 53–57

56.5 45.0 41.3 45.1 37.2 36.0 46.2 61.2 54.3

30–35 25–28 23–28 23–29 20–23 19–22 24–27 28–32 25–29 23–26 12–15 14–17 13–17 13–15 12–15 14–17 22–26 20–24 Ablennes hians Platybelone argalus argalus Strongylura hubbsi Strongylura marina Strongylura notata forsythia Strongylura notata notata Strongylura timucu Tylosurus acus acus Tylosurus crocodilus crocodilus

32.2 26.9 25.1 26.6 21.5 20.9 25.7 30.8 28.3

Mean Range Range Mean Range Mean Range Mean Range Mean Range Mean Species

24–28 17–20 16–19 16–21 13–15 12–15 16–20 20–24 17–22

Ablennes hians (Valenciennes in Cuvier and Valenciennes, 1846) Flat Needlefish Figures 2.1, 2.2 Tables 2.1, 2.2, 2.3

24.6 13.3 15.8 15.2 13.7 13.3 15.7 23.9 22.3

Range Mean

Caudal Pectoral Anal Dorsal

Table 2.1. Summary data for western Atlantic Belonidae.

Precaudal

Vertebrae Fin rays

Total

Predorsal scales

Fishes of the Western North Atlantic

Belone hians Valenciennes in Cuvier and Valenciennes, 1846: 432–436 (original description, Bahia, Brazil), pl. 548. Günther 1866:248 (description, after Valenciennes in Cuvier and Valenciennes, 1846). Jones 1876:134 (Bermuda). Goode 1877:292 (Bermuda). Goode and Bean 1879:46 (range, first U.S. record, from North Carolina). Jordan and Gilbert 1879: 383 (Beaufort, NC). Bean 1880:103 (New York market). Günther 1880:9 (Bermuda). Jordan and Gilbert 1881 (gill rakers absent). Cockerell 1892 (Jamaica). Posada Arango 1909:297 (Colombia). Mees 1962:29–31 (description, synonymy, range). Mees 1964:318, 325 (listed). Briggs 1964:707 (widespread species). Belone maculata Poey, 1860:290–291 (original description, Havana, Cuba). Cope 1871:481 (St. Kitts, West Indies). Tylosurus hians. Jordan and Gilbert 1883a:373 (description). Jordan 1883:283 (Belone maculata Poey not different from

8

Family Belonidae B. hians). Jordan 1886a:26 (Beaufort, NC). Jordan 1886b:35 (Havana, Cuba). Eigenmann and Eigenmann 1891:65 (Bahia, Brazil). Tylosurus (Athlennes) hians. Jordan and Fordice 1887:342, 357– 359 (original description of Athlennes as subgenus, description, synonymy). Jordan 1887b:568 (West Indies). Belone (Athlennes) hians. Metzelaar 1919:33 (Curaçao). Athlennes hians. Jordan and Evermann 1896a:321 (listed). Jordan and Evermann 1896b:717–718 (description, synonymy). Smith 1898:93 (2.5-foot specimen from Woods Hole, MA, summer, 1895). Evermann and Kendall 1899:61 (Key West, FL). Bean 1906:35 (Bermuda). Smith 1907:159–160 (Beaufort, NC). Kendall 1908:57 (Woods Hole, MA). Sumner et al. 1913:745 (Buzzards Bay, MA, trap). Gudger 1913:11 (North Carolina). Longley 1929:289 (found at Tortugas bird rookery). Norman and Fraser 1949:104 (Brazil). Miranda Ribero 1961:7 (Rio de Janeiro, Brazil). Ablennes hians. Miranda Ribeiro 1915:4–5 (description, Brazil). Fowler 1919b:6, 1919c:145 (St. Christophers = St. Kitts, West Indies). Meek and Hildebrand 1923:231–232 (description based on specimens from Beaufort, NC). Breder 1927:18 (Misteriosa Bank). Nichols and Breder 1927:58 (Woods Hole, MA), figure. Beebe and Tee-Van 1928:65–66 (description, Port-au-Prince Bay, Haiti). Hildebrand and Schroeder 1928:150–151 (description, Buckroe Beach and Cape Charles, mouth of Chesapeake Bay). Nichols and Breder 1928:435 (fig. 169, gill arch of juvenile with vestigial gill rakers). Jordan et al. 1930:197 (range, synonymy). Beebe and Tee-Van 1933:61 (Bermuda). Fowler 1940a:12 (New Jersey). Longley and Hildebrand 1941:29–30 (Tortugas). Fowler 1941d:141 (Bahia and São Salvador, Brazil). Breder 1948:89–90 (description, Florida straying north to Massachussetts). Fowler 1951b:74 (New Jersey). Fowler 1952a:112 (Atlantic and Ocean counties, New Jersey). Fowler 1952b:87 (Hispaniola, after Beebe and Tee-Van 1928). Bigelow and Schroeder 1953:168–169 (description, range, specimen from North Truro, MA). Fowler 1953:52 (Colombia, after Medellin 1909). Voss 1953:229–230 (eaten by Istiophorus americanus). Baughman 1955:55 (Port Aransas, TX). Erdman 1956:324 (Aguadilla, Puerto Rico, locally called tijerilla). Springer and Bullis 1956:57 (Oregon stations, Gulf of Mexico, TU and USNM). Boschung 1957:23 (Alabama). Springer and Hoese 1958:343 (Port Aransas and Heald Bank, TX). Hoese 1959:325 (Texas, after Baughman 1955, Springer and Hoese 1958). Gordon 1960:84 (Rhode Island). Springer and Woodburn 1960:24 (plaster cast of specimen from Clearwater, FL, at Sea Orama, Clearwater Beach). Berry and Rivas 1962 (description, western Atlantic). Erdman 1962:226 (in stomach of Makaira nigricans). Lowe (McConnell) 1962:693 (listed, British Guiana). Roux 1963:398 (types of Belone hians Valenciennes). Cervigón 1965:26 (Venezuela). Collette and Berry 1965:389 (validity of genus). Collette 1966:4–7 (meristic data). Cervigón 1966:216–217 (description, Venezuela). Caldwell 1966a:31 (Jamaica, Oregon, Sta. 3558, USNM 198026). Moe et al. 1966:16 (FSBC collection). Randall 1967:693 (food of Sphyraena barracuda), 754 (food of Scomberomorus regalis). Böhlke and Chaplin 1968:115 (description, Bahamas). Starck 1968:17 (Alligator Reef, FL). Collette and Parin 1970:37–41 (description, comparison of eastern Atlantic with western Atlantic specimens), fig. 9. Cressey and Collette 1970:396 (five species of parasitic copepods in western Atlantic), 411 (fig. 175, widespread distribution). Mago Leccia 1970:89 (listed, Venezuela). Dahl 1971:180–181 (description, outline figure, Colombia). Swingle 1971:27 (Alabama, after Boschung 1957). Powell et al.

1972:48 (three collections at FSBC). Duarte-Bello and Buesa 1973:78 (synonymy, range). Turner and Johnson 1973:7, 19 (lower reaches Newport River estuary, North Carolina). Palacio 1974:34 (Caribbean Colombia, UMML collections). Belyanina 1975:140 (Gulf of Mexico). Guitart 1975:244–245 (Cuba, synonymy, description, fig. 178). Shiino 1976:92 (common names). Gilmore 1977:132 (Indian River area, FL, after Anderson and Gehringer 1965). Hoese and Moore 1977:153 (description, range), 49 (pl. 131). Pequegnat et al. 1977:11 (Flower Gardens, Gulf of Mexico). Collette 1978 (description, range, figure). Figueiredo and Menezes 1978:63 (Brazil), fig. 108. Hardy 1978:87–89 (development, description, range, figures). Lima and Oliveira 1978:15 (Brazilian common name zambaiataba). Román 1979:81 (Venezuela, in key). Gilmore et al. 1981:10 (Indian River Lagoon, FL). Fahay 1983:212 (counts). Murdy 1983:88 (Texas, in key). Collette et al. 1984:336 (eggs), 344 (fig. 179D, 36 mm juvenile), 352 (counts). Rodríguez et al. 1984:15 (agujón de golfo, Cuba). Acero P. and Garzón F. 1987:90 (Colombia). Ditty et al. 1988:813 (larva, northern Gulf of Mexico). Zavala-Camin 1988a:11, 1988b:108 (in stomachs of Coryphae­ na hippurus, Brazil). Schwartz 1989:340 (North Carolina). Boughton et al. 1991 (ontogeny of upper jaw). Cervigón 1991:224–225 (description, Venezuela, fig. 168). Able 1992:4 (New Jersey, rare). Boschung 1992:83 (Alabama). Cervigón et al. 1993:269 (northern coast of South America). Humann 1994:56–57 (description, range, color photograph). Hensley and Hensley 1995:814 (eaten by Sooty Terns and Brown Noddies, Dry Tortugas, FL). Collette et al. 1997:7 (types). Debelius 1997:56–57 (description, color underwater photograph). Murdy et al. 1997:117–118 (description, Chesapeake Bay, fig. 96). Smith 1997:392 (description, range, not color pl. 109, which is a Tylosurus). Ho 1998:251 (parasitic copepod, Brazil). McEachran and Fechhelm 1998:935, figure (Gulf of Mexico). Carvalho Filho 1999:81 (description, Brazil). Smith-Vaniz et al. 1999:165 (Bermuda records). Lovejoy 2000:1352 (fig. 2, molecular phylogeny). Claro and Parenti 2001:43 (Cuba). Collette 2002:287–288 (Gulf of Maine), fig. 154. Briggs and Waldman 2002:58 (New York after 1964 manuscript “Long Island fish records by Roy Latham” cited in Briggs and Waldman [2002]). Humann and DeLoach 2002:60–61 (description, range, color photograph). Collette 2003a:1107 (figure, description, range in west-central Atlantic). Collette 2003b:1 (synonymy). Feiroza et al. 2003:68 (St. Paul’s Rocks, Brazil). McBride and Styer 2002:23 (common year-round in south Florida lampara net fishery). Menezes and Figueiredo 2003:67 (Brazil). Smith et al. 2003:15 (Belize). Banford et al. 2004 (molecular phylogeny). Lovejoy et al. 2004:369, 376 (molecular phylogeny). Reckel and Melzer 2004:15 (curtain-like falciform process in eye). Collette 2005:918–919 (early life history, figs. A–E). Schwartz 2006:64 (North Carolina records). Fahay 2007:776, 780–781 (early stages, figs. A–E). Luque and Tavares 2007:3, 9, 14–15 (parasitic copepods, Brazil). Nóbrega and Lessa 2007:68 (artisanal fishery, northeast Brazil). Casazza and Ross 2008:351 (associated with Sargassum, Gulf Stream off North Carolina). Setiamarga et al. 2008:603 (relationships to other Atherinomorpha). McEachran 2009:1285 (Gulf of Mexico). Collette 2010:32 (reproduction and development). Pepperell 2010:232 (color figure). Kells and Carpenter 2011:152– 153 (description, color painting). Schwartz 2012:44 (rare in North Carolina). Nolf 2013:75, pl. 156 (otoliths of Australian specimens). Schwartz and Purifoy 2013:69 (Newport River, Carteret County, North Carolina). Barbanti et al. 2013:279 (fig. 1p), 282 (Canal de Bertioga, São Paulo, Brazil). Humann

9

Fishes of the Western North Atlantic

Figure 2.1. Flat Needlefish, Ablennes hians. A. USNM 202583, BBC 904, 418 mm BL. Collected by B. B. Collette in the Gulf of Guinea, 16 November 1964. B. USNM 198035, 187 mm BL. Collected off Honduras from M/V Oregon, Sta. 3630, 7 June 1962. C. USNM 188843, 36.1 mm BL. Collected by B. B. Collette off Honduras, 9 June 1962. Illustrations by M. H. Carrington.

Brazilian common name is agulha (Figueiredo and Menezes 1978) with a local name of zambaia-taba in the state of Ceará (Lima and Oliveira 1978). Several other names have been used in other parts of the world (Shiino 1976). Diagnosis. Ablennes is monotypic; the species diagnosis is the same as the generic diagnosis. Description. Body elongate and greatly compressed laterally. Upper and lower jaws greatly elongate and studded with small sharp teeth. Gill rakers absent. Anterior parts of dorsal and anal fins with high falcate lobes, pectoral fins falcate. Caudal peduncle without lateral keels; caudal fin deeply forked, lower lobe longer than upper lobe. Females lack the right gonad and males either lack or have the right gonad greatly reduced in length. Dorsal fin rays numerous, 23–26, usually 24 or 25; anal fin rays numerous, 24–28, usually 26 or 27; pectoral fin rays 13–15 (Table 2.2). Vertebrae 60–63 precaudal + 30–35 caudal = 93–97 total (Table 2.3). Predorsal scales 360–430.

and DeLoach 2014:62–63 (description, range, color photograph). Smith-Vaniz and Jelks 2014:32 (St. Croix, U.S. Virgin Islands records). Snyder and Burgess 2016:102 (listed, Florida). Starck et al. 2017:91 (Florida Keys). Ablennes hians hians. Briggs 1958:264 (range). Duarte-Bello 1959:42 (range). Albennes [sic] hians. Schwartz 1962 (Maryland). Rees 1969: 539–540 (Bermuda, cestode Ptychobothrium belones in intestine).

Types of Atlantic Nominal Species. Belone hians Valenciennes in Cuvier and Valenciennes, 1846. Lectotype, designated by Collette and Parin (1970:40), MNHN B.1125 (451 mm BL); Bahia, Brazil. Dorsal fin rays 26; anal fin rays 27; pectoral fin rays 14–14. Paralectotype MNHN B.1204 (359 mm BL); Bahia, Brazil. Belone maculata Poey, 1860. No types known. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Flat Needlefish (English) and agujón sable (Spanish, Mexico). The common name in Cuba is agujón or agujón de golfo (Duarte-Bello 1959; Rodríguez et al. 1984; Claro and Parenti 2001); the 10

Family Belonidae Color. Bluish green above, silvery white below. A broad dark blue stripe along sides and about 12–14 prominent dark vertical bars on body; tip of lower jaw red. Juveniles and adults have an elevated black lobe in the posterior part of the dorsal fin. Size. The maximum observed size for Ablennes hians is 82.5 cm SL and 63 cm BL, commonly to 70 cm SL. The all-tackle gamefish record is 4.8 kg, for a fish caught in Mozambique in 1997 (IGFA 2018). Biology Predators. There are records of several species of fishes in the western Atlantic that eat Ablennes: Great Barracuda, Sphyraena barracuda (Randall 1967); Cero, Scomberomorus regalis (Randall 1967); Sailfish, Istio­ phorus platypterus (Voss 1953); Blue Marlin, Makaira nigricans (Erdman 1962), and Dolphinfish, Coryphaena hippurus (Zavala-Camin 1988a). They have also been picked up at the Bird Key Rookery, Tortugas, where they are eaten by Brown Noddies and Sooty Terns (Longley 1929; Longley and Hildebrand 1941; Hensley and Hensley 1995). Parasites. Nine species of parasites have been reported from the western Atlantic Ablennes hians: six copepods, two isopods, and a cestode. copepods: A total of nine species of parasitic copepods were reported from Ablennes by Cressey and Collette (1970), but only six of these were found in the western Atlantic: Bomolochidae, Bomolochus bel­ lones Burmeister (genus following Ho and Lin 2009), 12 males and 96 females from 31 specimens; Caligidae, Caligodes laciniatus (Krøyer), 8 females from 6 specimens; Anthosomatidae, Lernanthropus belones Krøyer, 3 males and 1 female from a single specimen from Puerto Rico, and L. tylosuri Richardi, 6 females and 3 females from 4 specimens; and Philichthyidae, Colobomatus goodingi Cressy and Collette, 2 females from interorbital canals of 2 specimens, from Cuba and Haiti. isopods: Two species of the parasitic isopod genus Mothocya were reported by Bruce (1986) but only M. longicopa Bruce was found in the Atlantic, in the Gulf of Guinea, first reported as Myothocya sp. by Collette and Parin (1970). cestodes: The single cestode record is of a postlarval Ptychobothrium belones (Dujardin) in the intestine of an Ablennes from Bermuda (Rees 1969). Food. No published information was located, but Ablennes undoubtedly feeds on small fishes like other marine needlefishes. Reproduction. Ablennes probably spawns offshore (Berry and Rivas 1962), presumably in the spring as a specimen collected in May from North Carolina

Figure 2.2. Atlantic distribution of Flat Needlefish, Ablennes hians, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

appeared ripe (Hildebrand and Schroeder 1928). Only the right gonad is developed. A 278 mm female had 660 eggs (3.0–3.15 mm in diameter) in the ovary. Ovarian eggs are 3.0–3.5 mm in diameter (Hildebrand and Schroeder 1928; Collette et al. 1984) and are covered with uniformly spaced tufts of filaments longer than the diameter of the egg (Collette et al. 1984). Development. Like most needlefishes, Ablennes passes through a halfbeak stage in which the lower jaw is much longer than the upper jaw (Fig. 2.1). Juvenile Ablennes, like juvenile Tylosurus, have a prominent melanistic posterior lobe in the dorsal fin. This lobe is either sloughed off or resorbed in Tylo­ surus species as they mature, but is retained in adult Ablennes. Juvenile Ablennes and some species of Tylo­ surus have vertical dark bars on their bodies that are lost in Tylosurus, but retained in Ablennes (Collette et al. 1984). Relationship to Humans. Not commonly used as food in the western Atlantic although Posada Arango (1909:297) wrote that in Colombia it “es bueno para comer.” Large Ablennes were seen for sale in the fish market in Cochin, India (B.B. Collette, pers. comm.). These were caught with drift nets (floating gill nets). Range. Widespread in tropical and subtropical seas within the 23.9 °C isothere (Cressey and Collette 1970: fig. 175). In the western Atlantic, from 11

Fishes of the Western North Atlantic Table 2.2. Fin ray counts for Ablennes hians, based on Collette and Parin (1970). Dashes indicate a value of zero. Fin rays Dorsal

Anal

Locality

22

23

24

25

26

24

25

26

27

28

29

Western Atlantic Gulf of Guinea Indian Ocean Persian Gulf Red Sea–Gulf of Aden West Central Pacific Eastern Pacific Species total

— — 1 2 — 3 — 6

11 3 11 10 12 15 3 65

40 33 14 3 19 52 7 168

51 29 2 — 5 27 3 117

13 4 — — — 3 — 20

3 — — 3 1 1 — 8

7 4 12 5 5 25 6 64

48 34 14 5 22 51 5 179

52 28 2 2 8 19 3 114

4 3 — — — 3 — 10

— — — — — 1 — 1

Table 2.3. Vertebral counts of Ablennes hians, based on Collette and Parin (1970). Dashes indicate a value of zero. Vertebrae Locality

82

83

84

85

86

87

88

89

90

91

92

93

94

95

96

97

Western Atlantic Gulf of Guinea Indian Ocean Persian Gulf Red Sea–Gulf of Aden West Central Pacific Eastern Pacific Species total

— — — 1 — — — 1

— — — 3 — — — 3

— — — 3 1 2 — 6

— — 2 3 2 6 — 13

— — — 4 2 8 1 15

— 1 3 — 5 3 4 16

— 1 1 — 2 2 2 8

— 3 2 — 1 6 — 12

— 14 5 — — 1 — 20

— 23 2 — 1 — — 26

— 17 1 — — 1 — 19

8 5 2 — 1 — — 16

23 — — — 1 — — 24

23 — — — — — — 23

17 — — — — — — 17

4 — — — — — — 4

North Truro, MA, on the north shore of Cape Cod (Bigelow and Schroeder 1953; MCZ 37040) and Woods Hole, MA, south past Bermuda through the Gulf of Mexico and Caribbean Sea to Rio de Janeiro, Brazil (Fig. 2.2). Study Material. From the western Atlantic, 183 specimens (57.5–610 mm BL), including the types of Belone hians, from 90 collections at CU, MCZ, MNHN, MNRJ, MZUSP, NMC, TU, UMML, USNM, YPM, and ZMK. From the eastern Atlantic, 69 specimens (76.6– 511 mm BL) from 26 collections from the Cape Verde Islands and Gulf of Guinea from Dakar, Senegal, south to MoÇamedes, Angola, as listed by Collette and Parin (1970).

there is no wide keel. Gill rakers are present in Platy­ belone as in Belone and Petalichthys. The lower jaw extension is greater than in most other genera of needlefishes, with the tip of the upper jaw falling short of the tip of the lower jaw even in adults. Analysis of molecular data (Lovejoy et al. 2004:369) does not show Platybelone as closely related to any other clade of needlefishes. Species. Platybelone contains only one species, Platybelone argalus, but worldwide seven subspecies are recognized (Collette 2003b). Range. Marine, widespread in tropical and subtropical waters, particulary around islands (Collette 2003b). In the western Alantic Platybelone is present from Virginia south, at Bermuda, throughout the Gulf of Mexico and the Caribbean Sea to Trinidad (Fig. 2.4).

Genus Platybelone Fowler, 1919 Platybelone Fowler, 1919b:2 (original description as subgenus of Belone, type species Belone platyura Bennett 1832 by original designation).

Platybelone argalus argalus (Lesueur, 1821) Western Alantic Keeltail Needlefish Figures 2.3, 2.4 Tables 2.1, 2.4, 2.5, 2.6

Diagnosis. Platybelone differs from all other needlefishes in having a greatly depressed caudal peduncle with a wide, flattened keel. The South American freshwater Pseudotylosurus is slightly similar, but

Belona argalus Lesueur, 1821:125–126 (original description, Guadaloupe), figure. Mees 1966:151–152 (validity of name).

12

Family Belonidae Belone argalus. Valenciennes in Cuvier and Valenciennes 1846:439–440 (description, after Lesueur). Cockerell 1892:14 (Jamaica). Fowler 1919b:135–136 (description, St. Martins, West Indies). Fowler 1930:270 (Grenada). Fowler 1941d:141 (Fernando de Noronha, Brazil, after Schreiner and Miranda Ribeiro 1903, MNRJ 1337). Fernández-Yépez 1948:3 (B. depressa Poey considered a junior synonym). Santos 1952:93 (Brazil, B. trachura Valenciennes considered a junior synonym). Nomura and Menezes 1964:354 (Brazil, reaches length of 35 cm). Tylosurus ardeolus (not of Valenciennes in Cuvier and Valenciennes, 1846). Jordan and Fordice 1887:349 (synonymy, in part). Jordan 1887:568 (listed, West Indies). Tylosurus ardeola (not of Valenciennes in Cuvier and Valenciennes, 1846). Jordan and Evermann 1896a:320 (listed). Jordan and Evermann 1896b:713 (description, synonymy). Bean 1906:36 (Bermuda, FMNH 5475–8). Evermann and Marsh 1902:99 (Puerto Rico). Parr 1930:18 (description, Bahamas). Belone trachura (not of Valenciennes in Cuvier and Valenciennes, 1846). Miranda Ribeiro 1915:5–7 (Fernando de Noronha, Brazil). Carvalho 1943:44 (description, Estado São Paulo, Brazil). Belone (Tylosurus) ardeola (not of Valenciennes in Cuvier and Valenciennes, 1846). Metzelaar 1919:29 (description, Curaçao). Strongylura ardeola (not of Valenciennes in Cuvier and Valenciennes, 1846). Breder 1927:18 (description, Bahamas). Nichols and Breder 1928:423 (17°56′N, 63°12′W). Breder 1929:279 (mature at 250 mm SL). Breder 1932a:3–19 (comparative development). Beebe and Tee-Van 1933:60 (Bermuda, USNM 164894, 164776, 164899). Anderson and Gehringer 1959:44, 48 (off Cape Canaveral, FL). Nahhas and Cable 1964:199 (trematodes Steganoderma nitens and Schikobalotrema acutum in intestines, Curaçao and Jamaica). Woods 1952:40 and Springer and Bullis 1956:57 (Alacran Reef, Gulf of Mexico). Strongylura argalus. Jordan et al. 1930:196 (range, synonymy). Strongylura longleyi Breder, 1932a:12–13 (original description, Tortugas). Longley and Hildebrand 1941:27–28 (description, Tortugas). Springer and Bullis 1956:57 (Gulf of Mexico, TU 5993). Briggs 1958:264 (listed, Florida). Duarte-Bello 1959:43 (Cuba). Hoese 1959:325 (listed after Springer and Bullis 1956).

Belone ardeola ardeola (not of Valenciennes in Cuvier and Valenciennes, 1846). Norman 1935:4–5 (subspecies from West Indies, Azores, and Cape Verde Islands). Belone argalus argalus. Fowler 1943:43 (Mona Island, West Indies). Fowler 1944:71 (Serranilla Bank, Caribbean Colombia, description, synonymy), 94 (Courtown Keys, description). Fowler 1953:52 (Courtown Keys, Serranilla Bank, after Fowler [1944]). Belone platyura. Mees 1962:58–63 (synonymy, description, in part). Mees 1964:325 (listed). Belone argala. Berry and Rivas 1962 (characters, synonymy, distribution). Hildebrand et al. 1964:114 (Alacran Reef, Mexico). Reséndez Medina 1971:24 (Alacran Reef, after Hildebrand et al. 1964). Duarte-Bello and Buesa 1973:287 (synonymy, range). Obregón-Barboza et al. 1994:84 (Veracruz). Platybelone argalus. Collette and Berry 1965:388 (validity of genus). Collette 1966:4–7 (counts). Caldwell 1966a:31 (Jamaica, USNM 198043, LACM 5865). Caldwell 1966b:340 (Miocene jaw fragments, Jamaica). Collette 1967:197 (validity of name Belone argalus). Randall 1967:683 (stomach contents 97% Jenkinsia, Virgin Islands). Birdsong and Emery 1968:192 (Albuquerque and Courtown keys off Nicaragua). Randall 1968:40 (description), fig. 41 (photograph). Starck 1968:17 (Alligator Reef, FL). Cressey and Collette 1970:399–400 (parasitic copepods), 412 (fig. 176, world distribution). de Sylva 1970:51 (Biscayne Bay, FL). Mago Leccia 1970:89 (Venezuela, listed). Bright and Cashman 1974:342, 348– 349 (description, West Flower Garden Bank, Gulf of Mexico), 378 (fig. 14, photograph). Fischthal and Nasir 1974:80 (digenetic trematode Multitestis blennii from intestine, La Tortuga Island, Venezuela). Palacio 1974:34 (Caribbean Colombia, UMML). Guitart 1975:246–247 (Cuba, synonymy, description, fig. 181). Erdman 1976:23 (ripe in June, northeastern Caribbean). Shiino 1976:92 (common name Keeltail Needlefish). Gilmore 1977:132 (Indian River Lagoon, FL). Hoese and Moore 1977:152 (description, range, Gulf of Mexico), 48 (pl. 129, photograph). Pequegnat et al. 1977:11 (Flower Gardens, Gulf of Mexico). Terrero and Bonnelly de Calventi 1978:7 (Dominican Republic). Vitvitsky 1978:662 (collagen thermostability). Román 1979:82 (Venezuela, figure). Gilmore et al. 1981:10 (Indian River Lagoon area, FL). Pezold and Edwards 1982:103 (Laguna Madre, TX). Amato

Figure 2.3. Western Atlantic Keeltail Needlefish, Platybelone argalus argalus. A. USNM 198043, 168 mm BL. Collected by B. B. Collette off Honduras from M/V Oregon, 17 May 1962. B. USNM 198102, 96 mm BL. Collected at 39°28′N, 69°30′W, 30 October 1956. Illustrations by M. H. Carrington.

13

Fishes of the Western North Atlantic species because: (1) it is the earliest name for the species, (2) the figure in the original description is clearly of this species and could not be of any other species, and (3) Fowler (1919b, 1930) clearly pointed out its applicability to the species. The name argalus (sometimes incorrectly emended to argala) has been extensively used in the Atlantic and, starting with Parin (1967), in the Pacific. Mees (1962, 1966) argued that the name argalus was a composite, but these arguments have been countered (Collette and Berry 1965; Berry 1967; Collette 1967; ICZN 1970). Strongylura longleyi Breder, 1932. Holotype AMNH 9686; Tortugas. Dorsal fin rays 13; anal fin rays 18. Belone ardeola Valenciennes in Cuvier and Valenciennes, 1846 from Martinique, West Indies. The whereabouts of the holotype are unknown (Collette et al. 1997; Collette 2003b). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Keeltail Needlefish (English) and agujón de quilla (Spanish, Mexico). The common name in Cuba is agujón aquillado (Rodríguez et al. 1984; Claro and Parenti 2001) and marao in Venezuela (Cervigón 1965, 1966; Mago Leccia 1970; Román 1979). Diagnosis. Meristically, Platybelone argalus argalus is most similar to the eastern Atlantic P. a. lovii. Both subspecies have moderately high numbers of fin rays (Table 2.4) and vertebrae (Table 2.5). Morphometrically, P. a. argalus is also most similar to P. a. lovii but differs significantly in the slopes of four of eight characters: interorbital width, head depth, head width, and orbit diameter (Collette and Parin 1970: table 10). Description. Body elongate, rounded in cross section. Upper and especially lower jaws greatly elongate and studded with fine teeth. Gill rakers present. Anterior parts of dorsal and anal fins not forming prominent lobes; pectoral fin not falcate. Caudal peduncle greatly depressed, least caudal peduncle depth about half the width, with very large lateral keels; caudal fin forked, upper and lower lobes of about equal length. Both right and left gonads present, right longer than left. Dorsal fin rays few, 12–15, usually 13; anal fin rays 17–20, usually 18; pectoral fin rays 10–12, usually 11 (Table 2.4). Vertebrae 44–47 precaudal + 25–28 caudal = 70–76 total (Table 2.5). Predorsal scales comparatively few and large, 103–128 (Table 2.6). Color. Bluish green above, silvery below. A dark blue stripe along sides. Fins clear, without pigment. Scales and bones green. Size. The maximum observed size for Platybelone argalus argalus is 38.2 cm SL and 25.6 cm BL in the west-central Atlantic, commonly to 30 cm SL.

1983:93 (digenetic trematode, Multitestis brasiliensis, from Venezuela). Murdy 1983:88 (figure), 89 (in key, Texas). Acero P. et al. 1984:41 (Caribbean Colombia). Rodríguez et al. 1984:15 (listed, Cuba, common name). Dyer et al. 1985:86 (digenetic trematode Lecithostaphylus nitens, Puerto Rico). Bruce 1986:1178 (parasitic isopod Mothocya sp., Bermuda). Acero P. and Garzón F. 1987:90 (Colombia). ValdésMuñoz and Garrido 1987:10 (Cuba). Schwartz 1989:340 (North Carolina). Boughton et al. 1991 (ontogeny of upper jaw). Cervigón 1991:228–229 (description, Venezuela). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Humann 1994:56–7 (description, range, color photograph). Debelius 1997:56–57 (description, color underwater photograph). Ho 1998:251 (parasitic copepod, Brazil). McEachran and Fechhelm 1998:935, figure (Gulf of Mexico). Mejía Aristizábal et al. 1998:30 (Colombian Caribbean oceanic atolls, listed). Allué et al. 2000:83 (Bahamas). Lovejoy 2000:1352 (fig. 2, molecular phylogeny). Claro and Parenti 2001:43 (agujón aquillado, Cuba). Gasparini and Floeter 2001:1643 (Trindade Island, South Atlantic). González-Gándara and Arias-González 2001:249 (Alacranes Reef, Gulf of Mexico). Humann and DeLoach 2002:58–59 (description, range, color photograph). McBride and Styer 2002:23 (common yearround in south Florida lampara net fishery). Menezes and Figueiredo 2003:67 (Brazil). Casazza and Ross 2008:351 (associated with Sargassum, Gulf Stream off North Carolina). Headley et al. 2009:672 (table 1, prey of Blackfin Tuna). Collette 2010:32 (reproduction and development). Pepperell 2010:232 (color figure). Kells and Carpenter 2011:152–153 (description, color painting). Nolf 2013:75, pl. 155 (otoliths of western Atlantic specimens). De Angelo et al. 2014:1043 (seagrass beds in eastern Gulf of Mexico). Humann and DeLoach 2014:62–63 (description, range, color photograph). Pinheíro et al. 2015:10 (underwater photograph, Trindade Island, Brazil). Snyder and Burgess 2016:102 (listed, Florida). Platybelone argala. Cervigón 1965:26 (Venezuela). Cervigón 1966:212–213 (description, Venezuela, fig. 83). Böhlke and Chaplin 1968:116 (description, figure, Bahamas). Belyanina 1975:140 (larvae, Gulf of Mexico and Caribbean Sea). Fedoryako 1980:581 (Sargasso Sea). Platybelone argala argala. Parin 1967:13 (western Atlantic subspecies). Platybelone argalus argalus. Collette and Parin 1970:28 (fig. 5, range in Atlantic), 1970:29–31 (7 subspecies recognized, eye size, fin ray and vertebral counts). Collette 1978 (description, range, figure). Collette et al. 1984:344 (fig. 179E, 96 mm juvenile in halfbeak stage), 352 (counts). Alvarez-Guillen et al. 1988:320 (Quintano Roo, Mexico). Cervigón et al. 1993:269 (northern coast of South America). Smith-Vaniz et al. 1999:166 (Bermuda records), color pl. 2–26. Collette 2003a:1108 (figure, description, range in west-central Atlantic). Collette 2003b:3–4 (synonymy). Smith et al. 2003:14 (Belize). Lovejoy et al. 2004:369, 376 (molecular phylogeny). Collette 2005:920–921 (early life history, figs. a–b). McEachran 2009:1285 (Gulf of Mexico). Smith-Vaniz and Jelks 2014:32 (St. Croix, U.S. Virgin Islands records). Starck et al. 2017:91 (Florida Keys).

Types of Western Atlantic Nominal Species. Belone argalus Lesueur, 1821. The whereabouts of Lesueur’s type are unknown. The name argalus is used for this 14

Family Belonidae Biology

in the eastern Atlantic (Collette and Parin 1970), and throughout Oceania. In the western Atlantic, known from off the Atlantic coast of the United States, from Bermuda, and from throughout the Gulf of Mexico and the Caribbean Sea (Fig. 2.4). Also known from the islands of Fernando de Norohna and Trindade off Brazil (Pinheiro et al. 2015). Geographic Variation. There is considerable geographic variation in the number of fin rays (Table 2.4), vertebrae (Table 2.5), and predorsal scales, and in body proportions among populations of Platybe­ lone. Previous authors have recorded the differences in number of fin rays among populations. Norman (1935) concluded that there were two Atlantic subspecies: Belone ardeola ardeola from the West Indies, Azores, and Cape Verde Islands, and B. ardeola tra­ chura from Ascension and St. Helena. Mees (1962) noted variation in the species (as Belone platyura Bennett) worldwide, but refrained from recognizing any subspecies. In his study of Indo-West Pacific needlefishes, Parin (1967:13) provisionally recognized five subspecies: Platybelone argala argala from the West Indies and West Africa; P. a. trachura (Valenciennes in Cuvier and Valenciennes, 1846) from St. Helena, Ascension, and Brazil; P. a. platyura (Bennett 1832) from the tropical Indo-West Pacific; P. a. platura (Rüppell 1837) from the Red Sea, Gulf of Aden, and

Predators. Platybelone are eaten by Blackfin Tuna, Thunnus atlanticus (Headley et al. 2009), probably by other epipelagic tunas and billfishes, and by seabirds. Parasites. Seven species of parasites have been reported from western Atlantic Platybelone argalus ar­ galus: three copepods, an isopod, and three digenetic trematodes. copepods: Cressey and Collette (1970) reported a total of seven species of parasitic copepods from Platybelone argalus, but only three of these were found on the western Atlantic subspecies: Bomolochidae, Bomolochus bellones Burmeister (genus following Ho and Lin 2009), 1 male and 46 females from 15 specimens; Caligidae, Caligus belones Krøyer, 1 male and 1 female from 2 specimens; and Anthosomatidae, Ler­ nanthropus belones Krøyer, 2 males and 1 female from 3 specimens from Bermuda. isopods: An unidentified parasitic isopod of the genus Mothocya was found on Bermudian specimens (Bruce 1986). digenetic trematodes: Three species of digenetic trematodes have been reported from Platybelone argalus argalus: Zoogonidae, Lecithostaphylus nitens (Linton, 1898) (Nahhas and Cable 1964; Dyer et al. 1985); Haplosplanchnidae, Schikhobalotrema acutum (Linton, 1910) (Nahhas and Cable 1964); and Lepocreadidae, Multitestis blennii Manter from Venezuela (Fischthal and Nasir 1974). Food. Feeds primarily on small fishes (Carvalho 1943) such as Jenkinsia, 97% by volume in Virgin Islands specimens (Randall 1967). Reproduction. Females were reported to be ripe in June in the northeastern Caribbean (Erdman 1976). The right ovary is longer than the left (ratio of left to right is 1.0–2.0). A 266 mm female contained 944 eggs (1.5–1.8 mm in diameter) in the left ovary, 1,136 eggs (1.5–1.6 mm) in the right ovary (Collette 2005). Development. Platybelone remains in the halfbeak stage, in which the upper jaw is much shorter than the lower jaw, to a larger size than most other needlefishes (Collette 2005), to beyond 100 mm BL (Fig. 2.3). Breder illustrated the development of the posterior part of the body for 33 mm SL, 71 mm SL, and 397 mm SL (Breder 1932a: pl. 3, fig 5; Collette 2005:912, figs. a–c). Range. Widespread in tropical and subtropical seas within the 23.9 °C isothere (Cressey and Collette 1970: fig. 17). Particularly abundant around islands such as the West Indies, and the other subspecies are primarily known from around islands such as Ascension, St. Helena, Annobón, and Fernando Po

Figure 2.4. Atlantic distribution of Keeltail Needlefish, Platybelone argalus, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

15

Fishes of the Western North Atlantic Table 2.4. Fin ray counts for Platybelone argalus. Dashes indicate a value of zero. Fin rays Dorsal Species and locality

Anal

11

12

13

14

15

16

15

16

17

18

19

20

Platybelone argalus argalus Western Atlantic Platybelone argalus lovii Cape Verde Islands

—

18

127

73

7

—

—

—

26

142

55

4

—

1

4

12

3

—

—

—

1

16

3

—

Platybelone argalus spp. Azores

—

—

—

4

—

1

—

—

—

—

5

—

Platybelone argalus annobonensis Gulf of Guinea

—

7

38

22

—

—

—

18

44

5

—

—

— —

— —

— —

4 2

33 15

9 2

— —

— —

— —

4 —

31 5

11 14

4 — —

37 1 2

26 11 45

1 23 64

— 5 15

— — —

7 — —

37 — —

25 9 24

— 21 76

— 9 25

— 1 1

3 7

38 104

66 317

9 214

— 78

— 12

2 9

46 101

54 183

14 278

— 133

— 31

Platybelone argalus trachura Ascension Saint Helena Platybelone argalus platyura Persian Gulf–Red Sea Indian Ocean West Central Pacific Platybelone argalus pterura Eastern Pacific Species total

Table 2.5. Vertebral counts of Platybelone argalus. Dashes indicate a value of zero. Vertebrae Species and locality Platybelone argalus argalus Western Atlantic Platybelone argalus lovii Cape Verde Islands Platybelone argalus spp. Azores Platybelone argalus annobonensis Gulf of Guinea Platybelone argalus trachura Ascension Saint Helena Platybelone argalus platyura Persian Gulf–Red Sea Indian Ocean West Central Pacific Platybelone argalus pterura Eastern Pacific Species total

62

63

64

65

66

67

68

69

70

71

72

73

74

75

76

—

—

—

—

—

—

—

—

8

38

48

11

2

—

1

—

—

—

—

—

—

—

1

4

10

3

2

—

—

—

—

—

—

—

—

—

—

—

—

—

—

—

2

2

1

—

—

—

—

2

15

27

9

1

—

—

—

—

—

—

— —

— —

— —

— —

— —

— —

— —

— —

— —

— —

5 —

13 —

17 3

6 10

— 3

6 — —

17 — —

13 — —

10 — —

5 — —

— — —

— — —

— 10 6

— 34 29

— 36 35

— 8 7

— 5 6

— 1 —

— 1 —

— — —

— 6

2 19

23 36

36 46

6 13

— 15

— 27

— 26

— 76

— 119

— 71

— 37

— 25

— 19

— 5

16

Family Belonidae Table 2.6. Number of predorsal scales of Platybelone argalus. Dashes indicate a value of zero. Predorsal scales Species and locality

101–102

103–104

105–106

107–108

109–110

111–112

113–114

115–116

117–118

119–120

P. argalus argalus Western Atlantic

—

2

1

6

6

4

6

17

8

9

P. argalus lovii Cape Verde Islands

—

—

—

2

2

3

5

1

1

4

P. argalus spp. Azores

—

—

—

—

—

—

—

—

—

—

P. argalus annobonensis Gulf of Guinea

—

—

—

—

1

1

1

2

7

3

P. argalus trachura Ascension Saint Helena

— —

— —

— —

— —

— —

— —

— —

2 —

1 —

3 —

P. argalus platyura Persian Gulf–Red Sea Indian Ocean West Central Pacific

1 — —

1 — —

2 — 1

2 2 3

5 2 11

10 2 10

4 1 7

7 4 6

1 1 10

6 1 8

P. argalus pterura Eastern Pacific

2

4

8

5

5

6

7

5

7

—

Species total

3

7

12

20

32

36

31

44

36

34

Predorsal scales 121–122

123–124

125–126

127–128

129–130

131–132

133–134

135–136

137–138

P. argalus argalus Western Atlantic

5

2

2

1

—

—

—

—

—

P. argalus lovii Cape Verde Islands

2

—

—

—

—

—

—

—

—

P. argalus spp. Azores

—

—

2

—

2

—

—

—

1

P. argalus annobonensis Gulf of Guinea

12

8

9

7

2

—

3

1

—

P. argalus trachura Ascension Saint Helena

8 —

3 —

5 2

8 2

3 3

5 2

2 2

1 1

— 2

1 1 3

1 1 2

1 — —

— — —

— 1 2

— — —

— — —

— — —

— — —

— 32

— 17

— 21

— 18

— 13

— 7

— 7

— 3

— 3

P. argalus platyura Persian Gulf–Red Sea Indian Ocean West Central Pacific P. argalus pterura Eastern Pacific Species total

Persian Gulf; and P. a. pterura (Osburn and Nichols 1916) from the eastern tropical Pacific. Seven subspecies are recognized worldwide, as there is variation among populations at groups of islands associated with the eastern Atlantic, compared with the relative lack of variation found from Mauritius and Tanganyika to Hawaii (Collette and Parin 1970). Within the

Atlantic, there are five distinct populations: P. a. ar­ galus (Lesueur, 1821) from the western Atlantic; P. a. lovii Günther, 1866 from the Cape Verde Islands; P. a. trachura Valenciennes in Cuvier and Valenciennes, 1846 from Ascension and St. Helena; P. a. annobon­ ensis Collette and Parin, 1970 from Annobón and Fernando Po (the most distinct subspecies meris17

Fishes of the Western North Atlantic spread in tropical and subtropical marine waters. Strongylura also has freshwater representatives, including S. fluviatilis, S. hubbsi, and S. krefftii. Stron­ gylura fluviatilis is known from the Pacific slope of Ecuador, S. hubbsi from Guatemala and Mexico, and S. krefftii from rivers of northern Australia and southern Papua New Guinea (Collette 2003b).

tically and morphometrically); and five specimens examined from the Azores, for which no subspecific name was applied. Study Material. From the western Atlantic, 393 specimens (23.7–276 mm BL) from 116 collections in AMNH, ANSP, BMNH, CAS, GCRL, LACM, MCZ, MNRJ, ROM, SIO, TU, UMML, UMMZ, USNM, YPM, ZMH, and ZMK. From the eastern Atlantic, 19 specimens (144–242 mm BL) from 9 collections, Cape Verde Islands; 60 specimens (211–320 mm BL) from 13 collections, Ascension and St. Helena; 57 specimens (149–233 mm BL) from 4 collections, Annobón and Fernando Po; and 5 specimens (252–274 mm BL), Azores, listed by Collette and Parin (1970).

Strongylura hubbsi Collette, 1974 Maya Needlefish Figures 2.5, 2.6 Table 2.1 Tylosurus timucu (not of Walbaum). Evermann and Goldsborough 1902 (USNM 50477 from Teapa River at Teapa, Tabasco, Mexico). Tylosurus marinus (not of Walbaum). Meek 1904 (FMNH 4690 from Tesechoacan River at Perez, Veracruz, Mexico). Strongylura sp. Miller and Nelson 1961 (UMMZ 184751 from Almoloya River, on Trans-Isthmian Highway, Oaxaca, Mexico, about 32 km north of its southern terminus). Salgado-Maldonado 2006:211, 291 (acanthocephalan parasite, Tonalá River basin, Mexico). Salgado-Maldonado 2008:42– 43 (parasites). Strongylura marina (not of Walbaum). Cressey and Collette 1970:360 (ergasilid copepod Acusicola tenax (Roberts) on gill filaments of “a freshwater population now referred to S. marina but possibly representing an undescribed form” from the Usumacinta River, Guatemala). Strongylura hubbsi Collette, 1974c (original description). Miller 1974:470 (Almoloya River, Coatzacoalcos Basin, Oaxaca, Mexico). Miller 1976a:155 (Atlantic coast streams from Papaloapam River to Usumacinta River). Miller 1976b:18 (Tesechoacan River, Perez, Veracruz, Mexico). Collette et al. 1992:4 (types). Espinoza Pérez et al. 1993:32 (after Collette 1974c). Lara-Domínguez et al. 1993:364 (Mexico). Lovejoy 2000:1352 (fig. 2, molecular phylogeny). Rodiles-Hernández et al. 1999 (Lacanja River, Chiapas, Mexico). Lovejoy and Collette 2003:587 (after Collette 1974c). Banford et al. 2004 (molecular phylogeny). Collette 2003b:6 (distribution). Lovejoy et al. 2004:369, 377 (molecular phylogeny). Miller et al. 2005:202–203 (range, habitat, fig. 6.233, map 6.199). Lozano-Vilano et al. 2007:46, 48 (Lacantún River, Chiapas, Mexico). González-Díaz et al. 2008:50 (La Venta River, Chiapas, Mexico). Miller et al. 2009:225–226 (range, habitat, fig. 6.233, map 6.199).

Genus Strongylura van Hasselt, 1824 Strongylura van Hasselt, 1824:374. Type species Strongylura caudimaculata van Hasslet, 1824. Type by monotypy.

Diagnosis. Body elongate, rounded in cross section. Upper and lower jaws greatly elongate and studded with sharp teeth; maxilla exposed posteriorly. Gill rakers absent. Anterior parts of dorsal and anal fins not forming prominent lobes; pectoral fins not falcate. Caudal peduncle without lateral keels, deeper than wide; caudal fin emarginate, not deeply forked. Strongylura is more advanced than basal belonids such as Belone, Petalichthys, and Platybelone, but lacks the characters of the more advanced Ablennes and Tylosurus. The genus has no known synapomorphies and species currently assigned to Strongylura may not be monophyletic, but are placed together because they lack the characters defining other genera (Collette 2003b). Analysis of molecular data (Lovejoy et al. 2004) recovers species of Strongylura as polyphyletic, spread among several clades of needlefishes (see Fig. 1.3). Species. Strongylura is the most speciose genus in the family, with 14 species recognized (Collette 2003b): S. anastomella, S. exilis, S. fluviatilis, S. hubb­ si, S. incisa, S. krefftii, S. leiura, S. marina, S. notata, S. scapularis, S. senegalensis, S. strongylura, S. timucu, and S. urvillii. Four of these species (S. hubbsi, S. marina, S. notata, and S. timucu) occur in the western North Atlantic. Strongylura marina is most closely related to the freshwater S. hubbsi and the eastern Pacific S. exilis, S. timucu is closest to the eastern Atlantic S. senegalensis, and S. notata is not closely related to any of the other American species of the genus (Banford et al. 2004; Lovejoy et al. 2004). Range. Most species are marine and are wide-

Types. Holotype UMMZ 194998 out of UMMZ 143506; male (183 mm BL, 304 mm SL); Guatemala, El Petén, Pasión River, immediately above mouth of Arroyo de Petexbatúm at Sayaxché. Dorsal fin rays 16; anal fin rays 17; pectoral fin rays 11–11. Vertebrae 42 precaudal + 25 caudal = 67 total. Predorsal scales 230. Testis lengths (left–right) 19–40 mm. See Study Material below for paratype information. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Maya Needlefish (English) and agujón maya (Spanish). 18

Family Belonidae Diagnosis. Strongylura hubbsi has both left and right gonads in the males (ratio of left to right is 1.1–4.0) and only the right gonad in females, whereas S. marina has only one gonad in both sexes, and S. notata and S. timucu have two gonads in both sexes. Pigment extends ventrally nearly to the lateral line, giving S. hubbsi a very dark appearance, like S. timu­ cu. As would be expected with a species of needlefish that is shorter (has fewer vertebrae) than its relatives, several proportional measurements (head length, snout length, and preopercle length) average distinctly longer than in S. marina and S. timucu (Collette 1974c: table 2). Counts of fin rays are very similar in all three species (Table 2.1). Vertebrae are fewer, 64– 68 total, than in either S. marina or S. timucu (68–77), but more than in S. notata (53–61). Predorsal scales are numerous, 216–293, (x̅ 245 for 82 specimens) as in S. marina (205–305, x̅ 256 for 145 specimens) and more than in either S. timucu (120–185) or S. notata (76–117). Color. Color descriptions are taken from the unpublished field notes of C. L. Hubbs. UMMZ 143513, field no. M35-100: “Olive above with blackish freckling, a wide, deep, sooty lateral band with silvery overcast separated from back by a bright metallic stripe which is broadest and brightest posteriorly. Fins amber green except anal which is mostly whitish. Eye golden just outside pupil. A few others seen before this one was caught look different from those seen in Río San Pedro largely because of the sooty stripe. Others speared, however, did not show this stripe so distinctly. Tip of mandible green in one seined.” For specimen UMMZ 143512, field no. M3568: “Yellow-green above on caudal and on dorsal rays. White below with metallic blue luster.” UMMZ 143511, field no. M35-86: “…olive above, silvery blue on sides, some have a wash of rose on the sides and on the paired fins and also dusky rose vertical fins and tip of mandible pink.” During his field work in Guatemala, R. M. Bailey (pers. comm.) noted that “males have the caudal fin and the distal halves of the dorsal and anal fins red (checked on seven specimens). Nine females with large eggs have these fins yellow-olive with no trace of red. None of these specimens has red on the snout.” Size. The maximum observed size for male Stron­ gylura hubbsi is 275 mm BL; females are known to 307 mm BL.

copepod, an acanthocephalan, and a nematode. copepods: An ergasilid copepod Acusicola tenax (Roberts) was collected from the gill filaments of Strongylura hubbsi (as S. marina) from the Usumacinta River, Guatemala (Cressey and Collette 1970). acanthocephalans: An acanthocephalan, Neo­ echinorhynchus golvani Salgado-Maldonado, family Neoechinorhynchidae, was reported from Strongylu­ ra, presumably S. hubbsi, from Lago El Rosario in the Tonalá River basin (Salgado-Maldonado 2006, 2008). nematodes: Procamallanus rebecae Addrade-Salas, Pineda-López, and Garcia-Magaña was reported from Strongylura, presumably S. hubbsi, from Lago El Rosario in the Tonalá River basin (Salgado-Maldonado 2006, 2008). Reproduction. Fifty males ranged in size from 97.8–275 mm BL and 45 females from 93.9–307 mm BL. Of the 50 males, 48 had the left and right gonads developed; 2 apparently lacked the left gonad. The right testis is longer than the left, with the ratio of the left to right 1.1–4.0 (x̅ 2.27). Seven of the 14 smallest males (97.8–165 mm BL) had poorly developed testes. The seven largest females (320, 307, 294, 288, 285, 284, and 280 mm BL) were all larger than the largest male. Of the 45 females, 2 had a tiny left ovary, 10–11 mm long; all others lacked any trace of the left ovary. Two of the four smallest females (93.9–136 mm BL) had a poorly developed ovary. Most of the other females had the ovary filled with developing eggs. The ovaries of two large females were removed and, through volumetric displacement, the egg count for one (285 mm BL, Usumacinta River, UMMZ 14360) was estimated at 367 eggs and for the other (294 mm BL, Grijalva River, USNM 50477) at 1,000 eggs. The eggs in specimen USNM 50477 were very large and seemed to be mature; 10 eggs ranged from 2.50 to 3.14 mm in diameter (x̅ 2.75). The eggs in the other specimen (UMMZ 14360) were smaller and variable in size, with 20 eggs ranging from 1.29–1.89 mm in diameter (x̅ 1.58). Development. Four collections, taken in March and April, contain juveniles. UMMZ 194743, 6 March 1948, Jaltepec River, tributary to Coatzocoalcos River, juvenile (24.6 mm BL). AMNH 24530, 15 March 1963, upper Pasión River, juvenile (52.1 mm BL). UMMZ 143504: Pasión River; 10 juveniles (24.4–68.4 mm BL) and 2 subadults (121–165 mm BL). UMMZ 153506, Arroyo de Petexbatúm, El Petén, juvenile (37.3 mm BL) and 11 adults (130–285 mm BL). Even the two smallest specimens (24.4 and 24.6 mm BL) are beyond the halfbeak stage of development, with head lengths of 12.9 and 14.0 mm, snout lengths of 8.1 and 8.5 mm, and distances between the tips of the upper and lower jaws of 3.0 and 1.8 mm.

Biology Parasites. Three species of parasites have been reported from western Atlantic Stongylura hubbsi: a 19

Fishes of the Western North Atlantic

Figure 2.5. Maya Needlefish, Strongylura hubbsi. USNM 214022, 187 mm BL. Collected by D. E. Rosen and R. M. Bailey from Guatemala, 10 April 1966. Illustration by M. H. Carrington.

Ecology. A summary of the ecological information in field notes for 18 UMMZ collections by C. L. Hubbs in 1935 and five by R. M. Bailey in 1966–1968 indicates that there was virtually no aquatic vegetation at most localities, only some water hyacinth and Myriophyllum at one. The water was usually clear, sometimes chalky blue. Bottom visibility ranged from 0.1–3 m, and was usually 0.3–0.7 m. The current was none to slight at most localities. Riffles occurred in a few, but there is no indication that any of the Strongylura hubbsi were taken in other than slow waters. The bottom consisted, at least partly, of mud at 21 of the 23 stations. The character of the shoreline varied widely: brushy; high bush; steep, heavily wooded hillside; sloping banks; tropical bush; claymud slopes; thick, high tropical bush; wooded; dense woods; open mud flat; and rainforest. Miller and Nelson (1961:6) described the habitat in the Almoloya River, a tributary to the Coatzacoalcos River, where one specimen (UMMZ 184751) of Stron­ gylura hubbsi was collected: “A clear winding stream of moderate gradient; average 20 ft. wide; maximum depth 8–9 ft.; swift, rocky riffles alternating with long pools; partly shaded by marginal vegetation; bottom included much sand and silt, alternating with rocks and boulders; steep banks; current swift to slight.” Range. Strongylura hubbsi is restricted to fresh water in the western part of the Usumacinta Province as defined by Miller (1966: fig. 2), centered in the basin that is drained by the upper Usumacinta River in Guatemala (Fig. 2.6). However, one specimen (FMNH 4690) was taken in the Tesechoacan River, tributary to the Papaloapan River; four in the Coatzocoalcos River; and one (USNM 50477) in the Teapa River, tributary to the Grijalva River. The Papaloapan enters the Bay of Campeche some 350 km west of the Usumacinta; the Coatzacoalcos 190 km west. Thus, S. hubbsi occupies Miller’s two major divisions of Usumacinta Province, the Usumacinta–Grijalva and the Papaloapan–Coatzacoalcos, and is absent from the two lesser divisions, the Yucatán Peninsula and the

Polochic–Lake Izabal. The species of Strongylura in Lake Izabal is S. marina. This distribution pattern is strikingly similar to that of the other Central American freshwater beloniform, Hyporhamphus mexicanus Alvarez, which was originally described from one specimen from the upper Coatzacoalcos River. Most of the additional 36 specimens now available are from the upper Usumacinta River in Guatemala. However, there are also two specimens from the Nuevo Gonzalez River, Tabasco, near the coast between the Coatzacoalcos and the Usumacinta.

Figure 2.6. General distribution of three freshwater Beloniformes: Maya Needlefish, Strongylura hubbsi, El Petén Halfbeak, Hyporhamphus mexicanus, and Breder’s Halfbeak, Hyporhamphus brederi; based on specimens examined by B. B. Collette. Map by W. E. Bemis.

20

Family Belonidae The distribution of Strongylura hubbsi is coincident with that of the poeciliid genus Priapella, as shown by Rosen and Bailey (1963: map 3; Miller 1966: fig. 3), an example of an autochthonous endemic genus in Usumacinta Province. In a series of papers (Rosen 1967, 1970; Rosen and Kallman 1969) four endemic upland characins and poeciliids have been described from the basin drained by the Usumancinta River, but all of these are restricted to the upper Usumacinta. Geographic Variation. Specimens from the Grijalva (one), Coatzaccalcos (three), and Papaloapan (one) rivers average higher predorsal scale counts (255; 285, 290, 293; and 276, respectively) than the Usumacinta population and were not designated paratypes. Vertebral counts (41 precaudal + 24 caudal; 40 precaudal + 25 caudal, 41 precaudal + 25 caudal, 40 precaudal + 25 caudal; and 43 precaudal + 25 caudal, respectively) average about the same as in the Usumacinta. Dorsal and anal fin rays for these specimens (15–16; 15–18, 15–17, 16–18; and 15–18, respectively) are also about the same as in the Usumacinta. Four of the five specimens are females with only the right ovary present; the fifth is a male with a ratio of left to right testis length of 2.2. Study Material. In addition to the holotype, there are 90 paratypes (24.4–307 mm BL) from 39 collections, plus 6 non-type specimens (24.6–320 mm BL) from 6 collections from the Papaloapan, Coatzacoalcos, and Grijalva rivers in Mexico (Collette 1974c).

Gaimard specimens MNHN 4508, 4511, Lesson and Garnot specimen MNHN 4510, Gay specimens MNHN 944; Cayenne, French Guiana, Poiteau specimen MNHN 721). Jordan 1887a:529–530 (Cayenne specimen MNHN 721). Belone scrutator Girard, 1858:170 (original description, Brazos and St. Joseph’s Island, TX; USNM 834). Girard 1858b:30 (description after Girard 1858a), pl. 13. Belone longirostris. Gill 1873:30 (silver gar; Cape Cod to Florida). Yarrow 1877:214 (description, behavior, Beaufort, NC). Goode 1879:116 (St. John’s River, FL). Jordan and Gilbert 1879:383 (abundant, Beaufort, NC). Bean 1880:103 (Noank, CT, and Woods Hole, MA). Ryder 1882 (development, single gonad). Tylosurus longirostris. Jordan and Gilbert 1881 (gill rakers absent, placed in Tylosurus). Jordan and Gilbert 1883a:374 (description, range, synonymy). Abbott 1888 (breeding in fresh water, Delaware and Raritan Canal, NJ). Evermann and Kendall 1894:123 (Galveston, TX, Belone scrutator a junior synonym). Tylosurus marinus. Jordan and Gilbert 1883b:587 (Charleston, SC). Bean 1883:450 (common names, range). Bean 1884:342 (Woods Hole, MA). Jordan 1886a:26 (Beaufort, NC). Jordan and Fordice 1887:351–352 (synonymy, range, description). Bean 1890a:39–40 (silver gar, ecology, Great South Bay, Long Island, NY). Bean 1891:92 (Chesapeake Bay). Jordan and Evermann 1896a:320 (common names, range). Jordan and Evermann 1896b:714 (description, range, synonymy). Evermann and Bean 1898:242 (Indian River, FL). Smith 1898:93 (Woods Hole, MA). Evermann and Kendall 1899:61 (Florida). Smith and Bean 1899:185 (ascends Potomac River). Linton 1901a:268, 277, 298, 1901b:442 (stomach contents, parasites, Woods Hole, MA). Bean 1900:99–100, 1902:405 (Long Island, NY). Linton 1905:356–357 (stomach contents, parasites, Beaufort, NC). Wilson 1905:631 (parasitic copepod, Woods Hole, MA). Tennent 1906:640, 674 (trematode, Beaufort, NC). Smith 1907:157 (common names, food, North Carolina), fig. 61. Kendall 1908:56–57 (New England localities). Wilson 1908:126, 623, 626 (parasitic copepods). Tracy 1910:87 (food, Rhode Island). Weymouth 1910:137 (Cameron, LA). Fowler 1911:11 (Delaware). Fowler 1913:63 (Franklin City, VA). Nichols 1913:96 (near New York City). Sumner et al. 1913:744–745 (food, parasites, Woods Hole, MA). Fowler 1914:343 (Ocean City, NJ). Kendall 1914:41 (Maine). Miranda Ribeiro 1915:19 (Brazil). Fowler 1916:41 (Sandy Hook, NJ). Starks 1916:23 (sesamoid articular). Fowler 1917a:111 (Somers Point, NJ). Fowler 1917b:121 (Woods Hole, MA). MacCallum 1917:55–58 (parasitic worm, Diplectanum tylosuri). Latham 1918:54, 1919:55–56 (Orient, Long Island, NY). Fowler 1918:14 (Lewes, DE). Nichols 1918:107 (New York City area). Wilson 1922:48–50 (parasitic copepod Lernanthropus chlam­ ydotes, Beaufort, NC). Latham 1923:61 (Orient, Long Island, NY). Breder 1926:122–123 (growth of jaws, Sandy Hook Bay, NJ). Nichols and Breder 1927:57–58 (description, New York and southern New England). Hildebrand and Schroeder 1928:148–149 (description, biology, Chesapeake Bay). Pearson 1929:136 (young, creek off Copano Bay, TX). Chandler 1935:126–127 (parasites, Galveston Bay, TX). Nagaty 1937:36 (trematodes, after Tennant 1906). Manter 1940:534 (trematodes). Sproston 1946:229–230, 541 (monogenes). Norman and Fraser 1949:102 (Atlantic coast of the United States). Bigelow and Schroeder 1953:167–168 (common name, description), fig. 79. Hargis 1955:188 (trematodes). Gordon 1960:31 (common name, Rhode Island records). Miranda Ribeiro 1961:6 (Brazil, MNRJ 1331, 1332). Pil-

Strongylura marina (Walbaum, 1792) Atlantic Needlefish Figures 2.7, 2.8 Tables 2.1, 2.7, 2.8, 2.9 Sea pike. Schöpf 1788:177. Esox marinus Walbaum, 1792:88 (original description, after Schöpff 1788; Long Island, NY). Esox belone var. marinus. Bloch and Schneider, 1801:391 (after Schöpff 1788). Esox belone var. Houttuyni Bloch and Schneider, 1801:391 (original description). Esox longirostris Mitchill, 1818:322 (original description, Hudson River near Albany, NY). Belona truncata Lesueur, 1821:126–127 (original description; New York, NY, Philadelphia, PA, and Newport, RI, MNHN 4505–6). Belone almeida Quoy and Gaimard, 1824:226–227 (original description, Rio de Janeiro, Brazil, MNHN 4508, 4511). Roux 1963:398 (reference to Quoy and Gaimard). Belone truncata. Valenciennes in Cuvier and Valenciennes 1846:402–405 (description, after Lesueur 1821). Günther 1866:244–245 (synonymy, in part, description). Belone galeata Valenciennes in Cuvier and Valenciennes 1846: 429–430 (original description, Cayenne, French Guiana, MNHN A.8896). Belone timucu (not of Walbaum). Valenciennes in Cuvier and Valenciennes 1846:426–428 (description, Brazil, Quoy and

21

Fishes of the Western North Atlantic

Figure 2.7. Atlantic Needlefish, Strongylura marina. A. TU 6950, 272 mm BL. Collected just south of mouth of Tangipahoa River in Lake Pontchartrain, Louisiana, U.S.A., 1953. B. USNM 189006, 23.5 mm BL. Dipnetted at night by B. B. Collette off Bluefields, Nicaragua, from M/V Oregon, 4 June 1962. Illustrations by M. H. Carrington.

lai 1967:262 (parasitic copepod Parabomolochus concinnus, Beaufort, NC). Tylosaurus [sic] marinus. Wilson 1911:365, 374 (parasitic copepods). Tylosurus almeida. Starks 1913:40 (Lake Papary, Natal, Brazil, in part, four specimens of Strongylura marina and four of S. timucu). Tylosorus [sic] marinus. Latham 1917:19 (Orient, Long Island, NY). Tylosurus marianus [sic]. Kudo 1919:36 (myxosporidian, Chloro­ myxum granulosum). Strongylura marinus. Gudger 1912:168 (Beaufort, NC). Breder 1948:89 (description, range). Pearse and Williams 1951:145 (Beaufort, NC). Thomson et al. 1971:80–81 (description, range), fig. 53. Hinegardner and Rosen 1972:639 (cellular DNA). Santos et al. 2008:6, 10 (acanthocephalan parasite, Rio de Janeiro, Brazil). Strongylura marina. Fowler 1919a:5 (Massachusetts to FL). Fowler 1920:155 (New Jersey localities). Fowler 1922:6 (Atlantic City, NJ). Fowler 1926:250 (Captiva Pass, FL). Fowler 1928b:609 (Longport, NJ). Jordan et al. 1930:196 (range, synonymy). Breder 1932a:6–7 (New York, New Jersey, Louisiana, Massachusetts, growth), pl. 1, fig. 1 (growth of upper and lower jaws), pl. 3, fig. 1 (caudal fin), pl. 11, fig. 2 (photograph in the New York Aquarium). Fowler 1935:161 (Island Heights, NJ). Carr 1936:161, 163 (Lake Eustis, St. Johns system, FL). Price 1937:127 (monogene Ancyrocephalus tylosu­ ri). Fowler 1938:60 (Barnegat Bay, NJ). Greeley 1939:84 (Long Island, NY). Buen 1940:45 (synonymy, range). Linton 1940:3, 23, 30–31 (trematodes, Woods Hole, MA). Fowler 1941a:238 (Lake Apopka, FL). Fowler 1941b:59 (Wildwood, NJ). Gunter 1942:314 (fresh water records). Hubbs and Allen 1943:122 (Silver Springs, FL). Fowler 1945:59–60 (description, synonymy, MD), figs. 172–174 (South Carolina records). Gunter 1945:46 (salinity 3.0‰–36.9‰, Texas). Fowler 1946:59 (Ocean City, NJ). Sproston 1946:541 (monogenes). Herald and Strickland 1949:104 (Homosassa Springs, FL). Baughman 1950:135 (Texas records). Alvarez

1950:59 (description, enters rivers, Mexico). Gunter 1950:93 (Aransas National Wildlife Refuge, TX). Gunter and Hildebrand 1951:732 (hypothermal mortality, Mustang Island, TX). Fowler 1952a:111 (New Jersey records). Mansueti and Scheltema 1953:4 (Chesapeake Bay). Raney and Massman 1953:426, 429, 431 (Pamunkey River, VA). Hopkins 1954:356–360 (trematodes, Virginia, Louisiana, Texas). Massman 1954:76–77 (freshwater portions of five Virginia rivers). Mansueti 1955 (Potomac River). Gunter 1956:350 (euryhaline species). Simmons 1957:183, 191 (salinity, upper Laguna Madre, TX). Briggs 1958:264 (range). Sparks 1958:72–73, 78 (trematodes, Grand Isle, LA). Yamaguti 1958:13 (digene Rhipidocotyle transversalis). Chambers and Sparks 1959:243 (Crystal Lake, TX). Darnell 1959:367 (food, Lake Pontchartrain, LA). Hoese 1959:325 (Texas). Arnold et al. 1960:11 (ecology, Galveston Island, TX). Boschung and Hemphill 1960:73 (Black Warrier River at Tuscaloosa, AL). Sparks 1960:293 (trematodes, Texas). Darnell 1961:558– 560 (food, Lake Ponchartrain, LA). Schwartz 1961a:392 (Chincoteague Bay, MD). Schwartz 1961b:5 (Patuxent River, MD). Breuer 1962:174 (lower Laguna Madre, TX). Hellier 1962:16 (Laguna Madre, TX). Schwartz 1962 (description, food, outline figure, Chesapeake Bay and lower Susquehanna River). Hoese and Jones 1963:42 (Redfish Bay, TX). McFarland 1963:96 (Mustang Island, TX). Schwartz 1964:180 (Ocean City, MD). Collette and Berry 1965:390 (validity of name S. marina). Gooding and Collette 1965 (parasitic copepod Colobomatus). Parker 1965:208 (Galveston, TX). Mago Leccia 1965:293–294 (description, ecology, Laguna de Unare, Venezuela, MIOUDO 00357– 60). Schwartz and Elser 1965 (20.9-inch specimen, Patuxent River, MD). Breder and Rosen 1966:302 (reproduction, references to Ryder [1882], Abott [1888]). Collette and Berry 1966:325 (validity of name). Miller 1966:796 (range, in part). Collette 1967:196–197 (validity of name). Dovel 1967:127 (larva, Magothy River, MD). Fiske et al. 1967:17, 19, 23, 25 (found at three of seven stations, Pleasant Bay, Cape Cod, MA, salinities 27.5‰–31.4‰). Goodyear

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Family Belonidae 1967:298 (eaten by Lepisosteus osseus along the coast of Mississippi). Hellier 1967:18–19 (Sante Fe River, FL, UF 8157, 8293). Norden 1966:126 (Vermillion Bay, LA). Patrick et al. 1967:401 (mile 134, Savannah River, GA). Perlmutter et al. 1967:52 (lower Hudson River, NY). Schaefer 1967:11, 18–19 (Fire Island, Long Island, NY). Woodwell et al. 1967:822 (DDT residue, Carmans River estuary, Long Island, NY). Collette 1968:190 (comparisons with S. notata and S. timu­ cu). Corkum 1968:346–347 (trematodes, Barataria Bay, LA). Fox and Mock 1968:46, 51 (salinity, Barataria Bay, LA). Jorgenson and Miller 1968:3, 11 (SL–TL conversions). Starck 1968:17 (Alligator Reef, FL). Whitworth et al. 1968:82 (previous Connecticut records, figure of juvenile). Dragovich 1969:15 (eaten by tunas). Kingston et al. 1969:545–546 (monogenes, Chesapeake Bay). Laird and Bullock 1969:1080 (no haematozoa found, Woods Hole, MA). Cressey and Collette 1970:401 (parasitic copepods). Collette and Parin 1970:23, 25–27 (comparison with S. timucu and S. senegalen­ sis). de Sylva 1970:51 (Biscayne Bay, FL). Lunz and Schwartz 1970:22 (South Carolina). Mago Leccia 1970:89 (listed, Venezuela). Richards and Castagna 1970:240, 243 (ecology, eastern shore, Virginia). Roessler 1970:864, 883 (ecology, Buttonwood Canal, FL). Schwartz 1970:33 (Mullet Pond, Shackleford Banks, NC). Schwartz and Tyler 1970:8 (outline figure), 30 (listed, North Carolina). Wiley 1970:156 (lower Potomac River). Briggs 1970:55–56 (parasitic isopod Lironeca ovalis, Great South Bay, NY). Ager 1971:60 (ripe adults, juveniles, Lake Okeechobee, FL). Briggs and O’Connor 1971:21, 24–27, 30–33 (ecology, Great South Bay, NY). Dahlberg and Scott 1971:32 (fresh water records, Georgia). Damon 1971:8 (photograph of specimen from British Honduras). Gilbert and Kelso 1971:27 (Tortuguero, Costa Rica). Grimes 1971:42 (Crystal River, FL). Perret et al. 1971:45 (Louisiana estuaries, salinities 4.0‰–31.2‰). Swingle 1971:27 (Alabama records). Buen 1972:159 (300 predorsal scales, Mexico), 160 (photograph). Carvajal Rojas 1972:16 (description, Gulf of Cariaco drainage, Venezuela). Dahlberg 1972:338 (Georgia estuaries). Oliveira 1972:36 (salinity, northeastern Brazil estuaries). Mountain 1972:54 (Crystal River, FL). Odum and Heald 1972:674 (North River, Everglades National Park, FL). Williams and Rogers 1972 (monogenes). Derickson and Price 1973:555–556, 558 (Delaware). Eskinazi 1972:286, 292 (ecology, north of Recife, Brazil). Kilgen and Harris 1973:100 (two individuals, 467– 601 mm TL, brackish-water pipeline canals, coastal marsh, Louisiana). Kroger and Guthrie 1973:115–116 (vertebral anomaly, North Carolina). Mahoney et al. 1973:599 (fin rot disease, New York Bight). Moss 1973:30 (Westport River estuary, MA). Reséndez Medina 1973:197–199 (description, ecology, Laguna de Alvaredo, Veracruz, Mexico). Reisman and Nicol 1973:27 (ecology, Gardiner’s Island, NY). Tagatz and Wilkens 1973:8–9 (ecology, Pensacola Bay, FL). Turner and Johnson 1973:7 (Newport River, NC). Collette 1974c:612 (comparison of the three western Atlantic species of Strongylura), 612 (drawing). Dineen 1974:382 (The Everglades, Florida). Foster 1974:125–126 (development, Potomac River, figure of 14.4 mm TL larva). Kushlan and Lodge 1974:119 (southern Florida canals into the Everglades). Livingston et al. 1974:254 (Apalachicola Bay, FL). Menhinick et al. 1974:37 (salt and brackish waters, occasionally enters fresh waters, North Carolina). McNulty et al. 1974:108 (western Florida localities). Rose and Hassler 1974:96 (eaten by Coryphaena hippurus off Cape Hatteras, NC). Weaver and Holloway 1974:62 (Louisiana tidal marsh). Williams and Gaines 1974:139 (no Acanthocephala, Mobile

Bay region). Grimes 1975:174 (Crystal River, FL). Sawyer et al. 1975:641 (marine leech Calliobdella vivida). Subrahmanyam and Drake 1975:451–456 (northern Florida salt marshes). Hubbs 1976:5 (listed, fresh waters, Texas). Miller 1976b:18 (no Mexican fresh water records). Seehorn 1975:19 (national forests, southeastern United States). Shiino 1976:92 (common name). Stunkard 1976 (life cycle trematode parasite). Beckham 1977:4 (Escatawpa River, AL, MS). Beecher et al. 1977:144 (oxbow lake, Escambia River, FL). Gilmore 1977:132 (previous records, Indian River, FL). Hoese and Moore 1977:152–153 (description, range, in part), 49 (color pl. 130). Hoff and Ibara 1977:671 (Slocum River, Buzzards Bay, MA). Hillman et al. 1977:361–362, 366 (abundance, Long Island Sound, NY). Livingston et al. 1977:92 (Apalachicola Bay, FL). Schauss 1977:278 (Lynnhaven Bay, VA). Stevenson 1977:182 (description, enters rivers, FL). Swift et al. 1977:34–35 (Ochlockonee River, FL). Yerger 1977:26–30 (Appalachicola River, FL). Castro-Aguirre 1978:60–61 (Atlantic Mexican fresh water records). Collette 1978 (description, range, figure). Cressey 1978:8 (branchiuran Argulus alosae). Cooley 1978:102 (Santa Rosa Sound, Pensacola, FL). De Vane 1978:584 (eaten by Scomb­ eromorus cavalla, Onslow Bay, NC). Figueiredo and Menezes 1978:64 (description, Brazil), fig. 109. Hardy 1978:90–95 (description, development, figures). Naughton and Saloman 1978:46, 50 (abundance, St. Andrew Bay, FL). Snelson and Bradley 1978:6 (present in area but not killed by cold, Mosquito Lagoon, FL). Hastings 1979:7 (jetties, St. Andrew Bay, FL). Perry and Carter 1979:29, 36 (southwest Louisiana). Román 1979:82 (in key, figure, Venezuela). Burgess 1980 (range, map of freshwater distribution in the United States). Love 1980:811 (reference to Hinegardner and Rosen 1972). McEwan and Hirth 1980:230 (remains found near Bald Eagle nests in north-central Florida). Modde and Ross 1980:914 (Horn Island, MS). Subrahmanyam 1980:132, Subrahmanyam and Coultas 1980:800 (northern Florida salt marshes). Weinstein and Davis 1980:101 (Cape Fear River, NC). Álvarez Cadena and Flores-Coto 1981:7 (larva, Laguna de Términos, Campeche, Mexico), fig. 6 (14 mm juvenile). Dovel 1981:27 (larvae, Patuxent River, MD). Gilmore et al. 1981:10 (Indian River Lagoon, FL). Hackney and de la Cruz 1981:659 (tidal creek, Bay St. Louis, MS). Lucena and Lucena 1981:41 (records from Florianópolis, Torres, Tramandí, Brazil). Reséndez Medina 1981b:486 (Carmen-Machona-Redanda lagoon system, Tabasco, Mexico). Crabtree and Dean 1982:8 (estuarine tide pools, South Carolina). Flores-Coto and Álvarez Cadena 1982:74 (Laguna de Términos, Campeche, Mexico). Guillory 1982:5 (Grand Isle, LA). Reséndez Medina 1982:394 (Laguna de Zontecompanan, Veracruz, Mexico). Schwartz et al. 1982:27 (Cape Fear estuary, NC). Wenner et al. 1982:4 (Santee River estuary, SC). Fahay 1983:212–213 (development), figure (14.4 mm larva from Foster 1974). Flores-Coto et al. 1983:252 (spawn in Laguna de Tamiahua, Mexico). Hickey and Lester 1983:105 (Fort Pond Bay, Long Island, NY). Holt and Holt 1983:465 (hypothermal mortality, Port Aransas, TX). Murdy 1983:88 (outline figure), 89 (in key, Texas). Snelson 1983:192 (Indian River, FL). Bireley 1984:244 (Long Island Sound, CT). Collette et al. 1984:336 (eggs), 344 (fig. 179C, 23.5 mm juvenile). Koltes 1984:120 (collected with Menidia menidia, Pettaquamscutt River, RI). Manooch and Raver 1984:134–135 (description, excellent baitfish, color figure). McKeown 1984:190 (lower Susquehanna River). Millican et al. 1984:32 (Lake Maurepas, LA). Rozas and Hackney 1984:216 (oligohaline marshes, Cape Fear River, NC). Wil-

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Fishes of the Western North Atlantic liams and Rogers 1984:582 (acanthocephalan Pomphorhyn­ chus lucyi). Kobelkowsky Díaz 1985:155 (Laguna de Tampamachoco, Veracruz, Mexico). Manooch et al. 1985:1212 (eaten by Euthynnus alletteratus). Perez and Rylander 1985:643 (hemoglobin heterogeneity, Venezuela). Rosen 1985:39 (fig. 34, occipital region and first vertebra). Bruce 1986:1116, 1181 (isopod Mothocya xenobranchia). Castro-​ Aguirre et al. 1986:164 (Veracruz). Gilmore 1987:129 (seagrass communities). Hastings 1987:24 (Manchac Wildlife Area, LA). Hastings et al. 1987:92 (Lake Maureps, tributary of Lake Pontchartrain, LA). Loftus and Kushlan 1987:202 (fresh water records, southern Florida), 203 (fig. 24, distribution records). Martins-Juras et al. 1987:109 (Ilha da São Luis, Maranhão, Brazil). Paiva Filho and Toscano 1987:157 (São Paulo, Brazil). Ross et al. 1987 (Horn Island, MS). Thayer et al. 1987:30 (few found in seagrasses near red mangrove prop root habitat, southern Florida). Gomez-​ Soto and Contreras-Balderas 1988:12, 14 (Tamaulipas, Mexico). Boschung 1989:74 (Mobile River basin, AL). Garzón F. 1989:155 (Colombia). Schwartz 1989:340 (North Carolina). Sogard et al. 1989a:58, 1989b:195 (Florida Bay). Norcross and Hata 1990:448 (Virginia Barrier Islands). Brown-Peterson and Eames 1990:230 (associated with spoil islands, Indian River Lagoon, FL). Peterson and Ross 1991:473 (Old Fort Bayou, MS). Schwartz et al. 1990:42 (Shackleford Banks, NC). Smith and Lake 1990:9 (Hudson River, NY). Boughton et al. 1991 (ontogeny of upper jaw). Cervigón 1991:229–230 (description, Venezuela, fig. 170). Hubbard et al. 1991:10 (Sucarnoochee River, Tombigbee drainage, AL). Szedlmayer 1991:78 (juveniles, Anclote River estuary, Florida). Able 1992:4 (New Jersey, common). Boschung 1992:84 (Alabama). Monteleone 1992:232 (larvae, Great South Bay, NY). Brown-Peterson et al. 1993:181 (Indian River Lagoon, FL). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Rodenas and López-Rojas 1993:75 (Laguna de Tacarigua, Venezuela). Zeckua-Ramos and Martínez-Perez 1993 (development in the estuarine system of Tecolutla, Veracruz, Mexico). Obregón-Barboza et al. 1994:84–85 (Veracruz, Mexico). Ross and Doherty 1994:62 (Horn Island, MS). Bull et al. 1995:336 (Lake Okeechobee, FL). Giannini and Paivo Filho 1995:144 (São Paulo, Brazil). Mettee et al. 1996:438–441 (distribution, biology, Mobile River basin, AL). González Bencomo 1997:39–40 (juveniles, Strait of Lake Maracaibo, Venezuela). González Bencomo et al. 1997:159 (Sector San Carlos del Lago de Maracaibo, Venezuela). Greenfield and Thomerson 1997:94 (not yet taken in Belize), fig. 100. Murdy et al. 1997:118–119 (description, Chesapeake Bay, fig. 97). Schwartz 1997:96 (found in all eight of North Carolina’s coastal watersheds). Smith 1997:392–393 (description, range, color pl. 110). Able and Fahay 1998:100–101 (early life history, Middle Atlantic Bight). Duffy and Baltz 1998:205 (Lake Pontchartrain, LA). Fuentes Zambrano 1998:69–72 (new monogene from northeast Venezuela). McEachran and Fechhelm 1998:936, figure (Gulf of Mexico). Schmitter-Soto 1998:74–76 (Quintana Roo, Mexico). Shute and Etnier 1998:16 (Tennessee River). Carvalho Filho 1999:81 (description, Brazil, figure). Fuller et al. 1999:287 (entered the Tennessee drainage through the Tennessee–Tombigbee connection). Suttkus et al. 1999:53 (lower St. Louis Bay, MS). Wagner and Austin 1999:201 (low salinity reaches of Chesapeake Bay). Allué et al. 2000:83 (western North Atlantic). M. E. de Araújo et al. 2000:140 (estuaries, Ceará, Brazil). Gordon et al. 2000:470 (comprise 54% of diet of Black Skimmers). Loftus 2000:34 (Everglades National Park, FL). Lovejoy 2000:1352 (fig. 2,

24

molecular phylogeny). Pérez-Hernández and Torres-Orozco 2000:434 (Mexican coastal lagoons). Vendel et al. 2000:180 (tidal flat, Paraná, Brazil). Gonzáles-Gándara and Arias-González 2001:249 (Alacranes Reef, Gulf of Mexico). Lovejoy and Collette 2001 (phylogeny). Paperno et al. 2001:127 (Ponce de Leon Inlet, FL). Collette 2002:288–289 (Gulf of Maine), fig. 155. Briggs and Waldman 2002:58 (New York, common in summer). Castillo-Rivera et al. 2002:176 (Pueblo Viejo Lagoon, Veracruz, Mexico). Liao 2002: kinematics of swimming. Starnes 2002:312 (Potomac River). Collette 2003a:1109 (figure, description, range). Collette 2003b:7 (synonymy). Menezes and Figueiredo 2003:67 (Brazil). Poulakis et al. 2003:174 (Charlotte Harbor, FL). Sanvicente-Añorve et al. 2003:373 (larvae from Campechén-La Ría estuary, Gulf of Mexico). Smith et al. 2003:14 (Belize). Arceo-Carranza et al. 2004 (fishes predominate prey in Veracruz). Banford et al. 2004 (molecular phylogeny). Boschung and Mayden 2004:374–375 (description, Alabama rivers), pl. 55D. Frota et al. 2004:23 (lengthweight, Brazil). Lasso et al. 2004:144 (Orinoco delta). Lovejoy et al. 2004:369, 377 (molecular phylogeny). O’Neill et al. 2004:86 (range, Alabama rivers). Poulakis et al. 2004:126 (Charlotte Harbor, FL). Reckel and Melzer 2004:15 (curtain-like falciform process in eye). Spach et al. 2004:51 (Brazil). Chávez-López et al. 2005:148 (Alvarado Lagoon estuary, Veracruz, Mexico). Collette 2005:922–923 (early life history, fig. a–e). Idelberger and Greenwood 2005:228 (Myakka River, southwestern Florida). Miller et al. 2005: 203 (range, habitat, Mexico, fig. 6.234). Pinheiro et al. 2005: 34 (Anil River estuary, Maranhão State, Brazil). Schwartz 2006:64 (North Carolina records). Solomon et al. 2006:6 (lower St. Marys River, northeastern Florida). Tuckey and Dehaven 2006:107, 109 (seagrass and tidal creek habitats, Suwanee River estuary, FL). Fahay 2007:776, 782–783 (early stages, figs. a–c). Félix et al. 2007:286 (beaches, Paraná, Brazil). Love and May 2007:267 (Maryland coastal bays). Luque and Tavares 2007:13, 15–16 (parasitic copepods, Brazil). Nóbrega and Lessa 2007:68 (artisanal fishery, northeastern Brazil). Cohen and Kohn 2008:24 (monogene parasite from Venezuela). Zeug et al. 2007:461 (only present in natural marsh-edge habitats, Guadalupe estuary, TX). Gaelzer and Zalmon 2008a:26, 2008b:168, 173 (sandy beaches, southeastern Brazil). González Bencomo et al. 2008:401–402 (larvae, La Palmita Bay, Lake Maracaibo, Venezuela). Nera and Castro 2008:51, 53 (Maranhão, Brazil). Rogers and Allen 2008:16 (Lake Okeechobee, FL). Tolan 2008:282, 290 (larvae, Nueces Bay, TX). Wingate and Secor 2008:1149 (high summer-fall abundance associated with low winter temperatures and high winter flows in Chesapeake Bay). Able et al. 2009:33 (marsh creek habitats, Delaware Bay estuary). García-Hernández et al. 2009:92 (juveniles, northern Yucatán Peninsula). Gómez Gaspar and Hernández Avila 2009:15 (Isla Cubagua, Venezuela). McEachran 2009:1285 (Gulf of Mexico). Miller et al. 2009:226 (range, habitat, fig. 6.234, map 6.200). Rohde et al. 2009:273–274 (photograph, map, South Carolina rivers below the fall line). Able and Fahay 2010:239–241 (ecology, early life history, Atlantic coast). Butler et al. 2010:59–61 (important component of diet of giant Atlantic Bluefin Tuna in North Carolina waters). Contente et al. 2011:44 (Guaraguaçu River, tributary of Paranaguá Bay estuary, southern Brazil). Reis-Filho et al. 2010:305 (Paraguacu River, Todos os Santos Bay, Bahia, Brazil). Gondolo et al. 2011:187 (surf zone, Itamambuca Beach, Ubatuba, São Paulo, Brazil). Kells and Carpenter 2011:152–153 (description,

Family Belonidae type (in parentheses): dorsal fin rays 16 (16); anal fin rays 19 (18); pectoral fin rays 12–12 (12–12). Predorsal scales 250 (295–300). MNHN 4506 was designated as the lectotype because of its excellent condition. Head length 149 mm; snout length 101 mm; lower jaw extends 6.0 mm beyond tip of upper jaw. MNHN 4505 has the snout distorted. Belone almeida Quoy and Gaimard, 1824. Lectotype, designated herein, MNHN 4508 (270 mm BL); Rio de Janeiro, Brazil; Quoy and Gaimard. Paralectotype also from Rio de Janeiro, MNHN 4511 (229 mm BL). Data for lectotype followed by paralectotype (in parentheses): dorsal fin rays 14 (14); anal fin rays 17 (17); pectoral fin rays 11–11 (11–11). Predorsal scales 250 (?). Head length 156 mm (127 mm); snout length 107 mm (85.5 mm); lower jaw extension 5.9 mm (3.9 mm). MNHN 4508 is designated as lectotype because it is in better condition than the other syntype and the predorsal scales can be counted. Belone galeata. Valenciennes in Cuvier and Valenciennes, 1846:429–430. Holotype MNHN A.792 (847 mm SL); Cayenne, French Guiana; Frère (Collette et al. 1997:7). Belone scrutator Girard, 1858. Lectotype, designated by Collette (in Collette et al. 1992:3), USNM 834 (198 mm BL); Texas, St. Josephs Island; G. Wurdemann, 1853. Paralectotype from same collection, USNM 309268 (212 mm BL). Data for lectotype followed by paralectotype (in parentheses): dorsal fin rays 14 (15); anal fin rays 18 (18); pectoral fin rays 12–12 (12–12). Predorsal scales about 245 (?). Vertebrae 45 precaudal + 26 caudal = 71 total, in both. Both females with only the right ovary present. The smaller specimen was designated as lectotype because it is in somewhat better condition and the approximate number of predorsal scales can be counted. Head length 109 mm; snout 73.9 mm; lower jaw extension 4.1 mm. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Atlantic Needlefish (English) and agujón verde (Spanish). Atlantic Needlefish is not a descriptive name, because there are four Atlantic species of Strongylura (plus several other genera), but recent workers have followed this usage. Earlier workers commonly used the names Silver Gar, Billfish, and Green Gar (Bean 1883; Smith 1907; Gudger 1912; Bigelow and Schroeder 1953; and others). Shiino (1976:92) listed 14 different names that have been used. The general name for Strongylura marina and most other needlefishes in Venezuela is marao (Mago Leccia 1965, 1970; Román 1979). In Brazil the general name is agulhão (Paiva and Holanda 1962; Eskinazi 1972) or agulha (Figueiredo and Menezes

color painting). Neves, Teixeira, and Araújo 2011:121 (middle reaches, Mambucaba River, Rio de Janeiro state, Brazil). Starnes et al. 2011:287 (Potomac River, C&O Canal). Abarca-Arenas et al. 2012:848 (weight–length and food in the Alvarado Lagoon system, Veracruz, Mexico). Mol 2012:652, Mol et al. 2012:278 (Suriname River estuary). Passos et al. 2012:230 (Paranaguá Estuarine complex, Brazil). Schwartz 2012:44 (scarce in North Carolina). Vilar et al. 2011:45 (Baia da Babitonga, southern Brazil). Neves, Teixeira, Franco et al. 2011:666 (middle and lower channels of the estuarine mixing zone of a tropical estuary of Mambucaba River, Rio de Janeiro state, Brazil). Arévalo-Frías and Mendoza-Carranza 2012:259 (larvae and juveniles from the Biosfera Pantanos Reserve in Tabasco, Mexico). Gallardo-Torres et al. 2012:277 (present for two to four months in the coastal lagoon Boca de la Carbonera, Yucatán, Mexico). Vega-Cendejas and Santillana 2012:298 (abundance in lagoon and coastal habitats in Yucatán, Mexico). Barbanti et al. 2013:279 (fig. 1p), 282 (Canal de Bertioga, São Paulo, Brazil). Stevens et al. 2013:1650 (Peace River, tributary of Charlotte Harbor, FL). De Angelo et al. 2014:1042 (seagrass beds in eastern Gulf of Mexico). Gallardo Torres et al. 2014:87 (Yucatán, Mexico, photographs). Snyder and Burgess 2016:102 (listed, Florida). Starck et al. 2017:91 (Florida Keys). Tylosurus galeatus. Jordan et al. 1930:195 (range, synonymy). Puyo 1949:161 (description, good to eat, French Guiana). Tylosurus sp. Gordon 1949:25–26 (summer, Coney Island, NY, AMNH 18221, 18229, 18262). Strongilura [sic] marina. Eskinazi 1967:268 (Pernambuco, Brazil).

The following references seem to be based on both Strongylura marina and S. timucu. Tylosurus subtruncatus. Jordan 1891:315 (Bahia, Brazil). Tylosurus marinus. Henshall 1891:375 (Lemon Bay, FL). Fowler 1915a:247 (Tarpon Springs, FL). Miranda Ribeiro 1918:763 (Brazil). Menzel 1956:51 (Apalachee Bay region, FL). Strongylura timucu. Fowler 1941d:141 (Brazil). Reid 1954:31–32 (ecology, Cedar Key, FL). Springer and Woodburn 1960:24 (ecology, females with right gonad only, males with both, Tampa Bay, FL). Strongylura marina. Gunter et al. 1948:314 (mortality, Ft. Myers Beach, FL). Berry and Rivas 1962 (description, range and synonymy, in part).

Misidentifications. As Tylosurus timucu: Fowler (1917b) Panama (ANSP 49356–7). As Strongylura timucu: Schultz (1949), Lake Maracaibo, Venezuela (USNM 121782, 121784); Gunter and Knapp (1951), Gunter (1958), Mustang Island, TX. Types. There are no extant types for either Esox mari­ nus Walbaum, 1792 or E. longirostris Mitchill, 1818, but both are based on needlefish from New York, which were almost certainly Strongylura marina. Belone truncata Lesueur, 1821. Lectotype MNHN 4506 (283 mm BL), designated by Collette (in Collette et al. 1997:8); Philadelphia, PA; Lesueur. Paralectotype, also from Philadelphia, MNHN 4505 (157 mm BL). Data for lectotype followed by data for paralecto25

Fishes of the Western North Atlantic Table 2.7. Fin ray counts for Strongylura marina. Dashes indicate a value of zero. Fin rays Dorsal

Anal

Pectoral

Locality

13

14

15

16

17

18

16

17

18

19

20

21

10

11

12

U.S. East Coast Gulf of Mexico Central America South America Species total

— — — 3 3

19 63 14 60 3 11 17 40 53 174

49 20 10 4 83

17 1 — — 18

1 — — — 1

1 — — 4 5

20 17 8 32 77

64 59 16 27 166

57 18 1 3 79

8 — — — 8

1 — — — 1

— 1 — — 1

20 36 4 49 109

37 15 9 12 73

Bay, NC, De Vane 1978; southeastern United States; Saloman and Naughton 1983); and Little Tunny, Eu­ thynnus alletteratus (Manooch et al. 1985). Strongylura marina constituted 7% by frequency of occurrence, and 3% by weight, of the diet of giant Atlantic Bluefin Tuna off North Carolina in one of three years studied (Butler et al. 2010). The record (as Tylosaurus [sic] marinus) from Atlantic Bluefin Tuna stomachs at Portland, ME (Crane 1936) is highly unlikely; we believe that this record was based on misidentified Atlantic Saury, Scomberesox saurus. Freshwater and estuarine populations are also preyed on by birds such as the Bald Eagle (Haliaeetus leucocephalus) near lakes in north-central Florida (McEwan and Hirth 1980) and the Black Skimmer (Rynchops niger) near the James River in Virginia (Gordon et al. 2000). Parasites. A wide variety of parasites, 22 species total, have been reported from Strongylura marina: six species of copepods, one branchiuran, two isopods, two acanthocephalans, three monogenes, five digenetic trematodes, a leech, a cestode, and a myxosporidian. copepods: Cressey and Collette (1970:401) reported a total of six species of parasitic copepods from Strongylura marina: Ergasilidae, Ergasilus spatulus (Cressey), 23 females from the gill filaments of 5 freshwater needlefish from Lake Pontchartrain, LA, and Tortuguero, Costa Rica; Bomolochidae, Bomolo­ chus bellones Burmeister (genus following Ho and Lin 2009), 116 females and 3 males from the gill chambers and oral valves from 53 specimens from throughout most of the range of the host, Massachusetts to Venezuela; Anthosomatidae, Lernanthropus belones Krøyer, 114 males and 58 females attached to the gill filaments of 37 specimens from the central part of the range, North Carolina to Venezuela, and L. tylosuri Richardi, 3 males and 2 females from 3 specimens from Belize; Lernaeoceridae, Lernaeolophus sultanus (Milne-Edwards), female from the roof of the mouth of a specimen from Belize; and Philichthyidae, Col­ obomatus goodingi Cressey and Collette, 6 females from cephalic lateral line canals of 5 specimens from

1978; Lucena and Lucena 1981; Martins-Juras et al. 1987). Diagnosis. Strongylura marina is similar to S. hubbsi and the eastern Pacific S. exilis in having many predorsal scales (205–305) and in lacking the right ovary. It also clusters with those two species based on molecular data (Lovejoy et al. 2004:369). Male S. marina lack the right testis, whereas male S. hubbsi have both left and right testes. The other two western Atlantic species and the eastern Atlantic S. senegalensis, have both gonads developed in both sexes although the right gonad is generally longer. Strongylura marina differs from all the Atlantic species except S. timucu in having more vertebrae (69–77 rather than 53–68). It differs from S. exilis in having more total vertebrae (69–77 rather than 67–74) and fewer predorsal scales (205–305, x̅ 256, rather than 274–330, x̅ 306). Description. Dorsal fin rays 13–17; anal fin rays 16–21; pectoral fin rays 10–12, usually 11 (Table 2.7). Vertebrae 41–50 precaudal + 23–29 caudal vertebrae, 69–77 total (Table 2.8). Predorsal scales numerous and tiny, 205–305 (Table 2.9). Color. Bluish green above, silvery below. A conspicuous dark blue stripe along the sides; black pigment behind the eyes, usually not extending below middle of orbit, pre-orbital bone slightly pigmented. Size. The maximum observed sizes for Strongylura marina are 64 cm SL and 42 cm BL (commonly to 50 cm SL). The all-tackle gamefish record is 1.84 kg for a fish caught on the Cape May Reef, NJ, in 2004 (IGFA 2018). Biology Predators. Preyed on by a variety of surface-feeding fishes, such as the Long-nosed Gar, Lepisosteus osseus (Mississippi Gulf Coast, Goodyear 1967); Yellowfin Tuna, Thunnus albacares, and Atlantic Bluefin Tuna, T. thynnus (Dragovich 1969); Dolphinfish, Coryphaena hippurus (off Cape Hatteras, NC, Rose and Hassler 1974); King Mackerel, Scomberomorus cavalla (Onslow 26

Table 2.8. Vertebral counts of Strongylura marina. Dashes indicate a value of zero. Vertebrae Precaudal Locality

41

42

43

44

45

46

Caudal 47

48

49

50

23

25

27

28

29

69

70

71

72

73

74

75

76

77

1 2 — — 3

14 39 43 7 30 15 — 7 17 6 25 16 Vertebrae 27 101 91 Caudal

23 — 9 — 32

1 — — — 1

3 5 — 11 19

4 23 — 17 44

15 18 — 11 44

34 6 3 6 49

22 1 22 3 48 Total

11 — 8 — 19

17 — — — 17

10 — — — 10

6 — — — 6

Locality 41 42 43 44 45 46 47 48 49 50 23 24 25 26 27 28 29 Table 2.9. Number of predorsal scales of Strongylura marina. Dashes indicate a value of zero. U.S. East Coast — — 1 4 28 38 22 19 7 2 — 1 14 39 43 23 1 Gulf of Mexico — 1 6 27 16 4 1 — — — — 2 7 30 15 — — Predorsal scales Central America — — — — 5 20 8 — — — — — — 7 17 9 — South America 3 1 13211–220 19 7 221–230 5 1 —231–240 — — 241–250 2 — 251–260 6 25 16 261–270 — — Locality 201–210 Species total 3 2 20 50 56 67 32 19 7 2 2 3 27 101 91 32 1 U.S. East Coast 3 5 6 4 5 10 10 Gulf of Mexico — 2 6 11 14 9 10 Central America — 1 — 1 8 3 — South 2.9. America — 8 indicate a 6value of zero.5 1 Table Number of predorsal scales 2of Strongylura6marina. Dashes Species total 3 10 18 24 33 27 21 Predorsal scales

69

70

71

72

73

74

75

76

77

3 4 5 23 — — 11 17 271–280 19 44 4 8 — 1 13

13

9

3

Locality

27

U.S. East Coast Gulf of Mexico Central America South America Species total

15 34 22 11 17 18 6 1 — — — 3 22 8 — 11 6 3 291–300 — — 281–290 44 49 48 19 17 9 5 2 4 1 — 1 —

10 6 — — — — — — 301–310 10 6 3 — — —

201–210

211–220

221–230

231–240

241–250

251–260

261–270

271–280

281–290

291–300

301–310

3 — — — 3

5 2 1 2 10

6 6 — 6 18

4 11 1 8 24

5 14 8 6 33

10 9 3 5 27

10 10 — 1 21

4 8 — 1 13

9 2 1 1 13

5 4 — — 9

3 — — — 3

Family Belonidae

24

U.S. East Coast — — 1 4 28 38 22 19 7 2 — Gulf of Mexico — 1 6 27 16 4 1 — — — — Table 2.8. Vertebral counts Central America — —of Strongylura — — marina. 5 20Dashes 8 indicate — —a value — of zero. — South America 3 1 13 19 7 5 1 — — — 2 Species total 3 2 20 50 56 67 32 19 7 2 2 Precaudal

26

Total

Fishes of the Western North Atlantic trematodes are known from Strongylura marina. Schikhobalotrema acutum (Linton, 1910), family Haplosplachnidae, was described from Strongylura from the Tortugas (probably Strongylura timucu) and also reported from Apalachee Bay, FL (Nahhas and Short 1965). Linton (1940) reported Podocotyle olssoni Odhner, Allocreadiidae, from Strongylura marina from Woods Hole, MA. Three species are in the family Bucephalidae. Bucephaloides strongylurae Hopkins, 1954 was described from Port Aransas, TX, and also reported from Grand Isle, LA (Sparks 1958). Two species of Rhipidoctoyle are known from Strongylu­ ra marina: Rhipidocotyle transversale Chandler, 1935 and R. lintoni Hopkins, 1954. Both species have the same intermediate and final hosts: metacercariae in silversides, Menidia menidia, and adults in Stron­ gylura marina (Stunkard 1976). Cercariae have been found in the bivalve Lyonsia hyalina but they have not yet been identified to species (Stunkard 1976). There are records of R. transversale from Galveston and Port Aransas, TX; Grand Isle, LA; Apalachee Bay, Bayboro Harbor, and Tampa Bay, FL; York River, VA; and Woods Hole, MA. Records of R. lintoni are less extensive: Grand Isle, LA; York River, VA; and Woods Hole, MA. cestodes: Larval cestodes, identified as Scolex polymorphus Rudolphi have been reported from the intestine of Strongylura marina from Woods Hole, MA (Linton 1901a, 1901b) and Beaufort, NC (Linton 1905). leeches: The marine leech Calliobdella vivida (Verrill), family Piscicolidae, was reported from several new hosts, including Strongylura marina, by Sawyer et al. (1975). myxosporidians: Chloromyxum granulosum Davis was reported from the urinary bladder of Strongylura mari­ na (Kudo 1919). Food. Strongylura marina feeds on a wide variety of small fishes, crustaceans, and insects. Fishes are caught crosswise between the upper and lower jaws and then turned and swallowed head first (Bean 1902). Fishes eaten include Fundulus (Bean 1890a; Tracy 1910; Hildebrand and Schroeder 1928; Woodwell et al. 1967), anchovies (Bean 1890a; Smith 1907; Darnell 1959, 1961), and especially silversides (Linton 1901a, 1901b, 1905; Smith 1907; Tracy 1910; Nichols 1918; Nichols and Breder 1927; Hildebrand and Schroeder 1928; Joseph and Yerger 1956). Silversides, particularly Menidia menidia, are clearly an important item, as shown by the fact that two species of digenetic trematodes use Menidia as an intermediate host and S. marina as the final host (Hopkins 1954; Sparks 1960; Stunkard 1976). Crustacean prey includes shrimps and amphipods (Linton 1905; Tracy 1910;

Florida. Earlier copepod records were summarized by Cressey and Collette: Lepeophtheirus edwardsi Wilson (Wilson 1905, Woods Hole, MA; Wilson 1908, Beaufort, NC); Tuxophorus caligodes Wilson (Wilson 1908, chalimi larvae, Beaufort, NC); Bomolochus concinnus Wilson 1911 = Bomolochus bellones (but considered a valid species by Ho and Lin 2009) (Beaufort, NC); Lernanthropus chlamydotes Wilson = Lernanthropus be­ lones (Wilson 1922, Beaufort, NC). Two records from S. timucu from Lemon Bay, Florida, by Bere (1936) may refer to either S. timucu or S. marina; Bomolochus nitidus = Bomolochus bellones (but considered a valid species by Ho and Lin 2009) and Lernanthropus chlamydotes = Lernanthropus belones. Pillai (1967) and Vervoort (1969) also mentioned these records. branchiurans: Cressey (1978) reported one species, usually parasitic on clupeids: Argulus alosae Gould. isopods: Two species have been reported: Lironeca ovalis from Great South Bay, NY (Briggs 1970) and Mothocya xenobranchiae Bruce from Florida (Bruce 1986). acanthocephalans: There are records of three species from Strongylura marina: Neoechinorhynchidae, Neoechinorhynchus agile (Rudolphi) from the intestine of fish from Woods Hole, MA (Linton 1901b; Yamaguti 1963); Pomphorhynchus lucyi Williams and Rogers, 1984 from freshwater and brackish water fishes from the southeastern U.S. Gulf coast (Williams and Rogers 1984); and Polymorphidae, Hexaglandula mutabilis (Rudophi) from Angra dos Reis, Rio de Janeiro, Brazil (Santos et al. 2008). monogenes: Five species of monogenes have been described from the gill filaments of Strongylura ma­ rina, three from the family Dactylogyridae, one from the Microcotylidae, and one from the Allopyragraphoridae. The dactylogyrids include Ancyrocephalus tylosuri (MacCallum 1917, redescribed by Price 1937) from specimens in the New York Aquarium, A. par­ vus Linton 1940 from specimens from Woods Hole, MA, and A. cornutus Williams and Rogers 1972 from specimens from Tuscaloosa, AL, and Lake Seminole and Pensacola, FL. There seem to be no subsequent records of A. tylosuri (Sproston 1946) but A. parvus has been reported from Alligator Harbor, FL (Hargis 1955), the Chesapeake Bay area (Kingston et al. 1969), and several localities in Alabama and Florida (Williams and Rogers 1972). The microcotylid Nu­ daciraxine gracilis (Linton, 1940) was described from S. marina from Woods Hole, MA. The allopyragraphorid Allopyragraphorus marinae Fuentes Zambrano, 1998 was described from Laguna de La Restinga in northeast Venezuela (Fuentes Zambrano 1998; Cohen and Kohn 2008). digenetic trematodes: Five species of digenetic 28

Family Belonidae tional figures. Development of yolk-sac larvae was illustrated by Hardy (1978: figs. 40–41) at 9.2, 10.1, 12.0, 13.3, and 14.3 mm TL. Newly hatched larvae, at 14.4 mm TL (Foster 1974), are very darkly pigmented and have well-developed pectoral fins that lack fin rays. Pectoral fin rays form by 25.3 mm TL and pelvic fins begin to develop at 30.0 mm TL. Upper and lower jaws are about equal at 25.3 mm TL, but the lower jaw grows much more rapidly, to 30.0 mm TL when the fish is in the halfbeak stage. Álvarez Cadena and Flores-Coto (1981: fig.6) illustrated a 14 mm halfbeak stage juvenile from Campeche, Mexico. Growth of the jaws was first documented by Breder (1926) and the transition from the halfbeak stage at 19 mm SL to the adult condition at 170 and 540 mm SL was illustrated by Breder (1932a: pl. 1, fig. 1; Collette 2005: fig. E). Ecology. This is an estuarine species that moves upstream into freshwater rivers and lakes (Gunter 1942, 1956; Burgess 1980) and probably reproduces in fresh water throughout its range. Early records such as the description of Esox longirostris Mitchill, 1818 from the Hudson River near Albany, NY, and the report of freshly hatched larvae in the Delaware and Raritan Canal (Abbott 1888) show this. It is fairly common in the lower Hudson River as far north as Poughkeepsie, NY (Smith and Lake 1990). It moves into the Bass River, MA (Curley et al. 1975), and is common in the Chesapeake region, being found in all parts of the bay from Havre de Grace, MD, to Cape Henry, VA (Hildebrand and Schroeder 1928). It is found upstream 40 miles above brackish water in five Virginia rivers; running ripe females and young have been taken in the Pamunkey and Chickahominy rivers (Massmann 1954). There are many freshwater records from Florida: Lake Eustis (Carr 1936), Lake Apopka (Fowler 1941a), Silver Springs (Hubbs and Allen 1943), Homosassa Springs (Herald and Strickland 1949), St. Johns River (McClane 1955), Lake Okeechobee (Ager 1971; Rogers and Allen 2008), South Florida canals into the Everglades (Kushlan and Lodge 1974; Loftus and Kushlan 1987), Apalachicola River (Yerger 1977), and Ochlockonee River (Swift et al. 1977). It also enters freshwater rivers in Alabama, Mississippi, and Louisiana, the Escatawpa River (Beckham 1977), 388 river miles from the Gulf of Mexico at Tuscaloosa, AL, on the Black Warrior River (Boschung and Hemphill 1960), throughout the Mobile River basin (Mettee et al. 1996), and in Lake Meurepas, a tributary of Lake Pontchartrain (Hastings et al. 1987). It has traversed the Tennessee– Tombigbee Canal and entered the Tennessee River at the New Johnsonville Steam Plant, a few kilometers below the mouth of the Duck River (Shute and

Hildebrand and Schroeder 1928; Carr and Adams 1973; Subrahmanyam and Drake 1975). At 35–50 mm SL, S. marina near Crystal River, FL, feed on fishes (30%), shrimps (30%), and amphipods (40%), and then shift to almost 100% fishes after 50 mm SL (Carr and Adams 1973). Insects (Diptera and Coleoptera) are also eaten in freshwater and estuarine areas such as Lake Pontchartrain, LA (Darnell 1959, 1961), the Tampa Bay area, FL (Springer and Woodburn 1960), and the Alvarado Lagoon system in Veracruz, Mexico (Abarca-Arenas et al. 2012). Reproduction. Spawning takes place in fresh and brackish waters in shallows with submerged algal masses between May 20 and June 10 in the Potomac River (Foster 1974). Ripe females have been reported in early May in the Chickahominy River, VA (Massman 1954), March to August in Lake Seminole in the Apalachicola drainage, FL (Yerger 1977), and July in Laguna Madre, TX (Simmons 1957). J. K. Galbraith (pers. comm., 9 April 2018) caught ripe females in a marsh at the headwaters of Buzzards Bay, MA, in mid-June 2017; the females seem to leave the marsh after spawning. This is the northernmost record of spawning. Larvae have been reported from Great South Bay, NY, in June and July (Monteleone 1992), “very young specimens” in late June in the Delaware and Raritan Canal (Abbott 1888:72), a larva from the Magothy River, MD, on 8 June (Dovel 1967), a 23 mm juvenile from the Pamunkey River, VA (Massman 1954), one- to two-inch total length young during the summer in Lake Okeechobee, FL (Ager 1971), and 15–25 mm TL young from Lake Seminole, Apalachicola drainage, FL (Yerger 1977). Only the right gonad develops in Strongylura mari­ na as in the eastern Pacific S. exilis. The other marine species of Strongylura in the Atlantic have both left and right gonads developed. Female S. hubbsi also have only the right gonad developed, but males have both gonads. Ryder (1882) was the first to record that S. marina has only one gonad, but the systematic significance of this was not clear until it was found to be a good character to distinguish S. marina from S. timucu (Collette 1968). A female Strongylura marina (294 mm BL) had approximately 1,000 ovarian eggs averaging 2.75 mm in diameter. Mature eggs were reported to be one-seventh of an inch in diameter (3.63 mm) by Ryder (1882:285). Development. Early development was described and illustrated by Ryder (1882). The embryo begins to form in 24 hours, eyes appear at 44 hours, pectoral buds at 70–80 hours, chromatophores at 116 hours, and the finfolds develop at 165 hours. Hardy (1978) republished Ryder’s illustrations with some addi29

Fishes of the Western North Atlantic the Gulf of Mexico, and along the coasts of Central and South America south at least to Río de Janiero, Brazil. The northernmost record is from Wolf’s Neck, Freeport, ME (Kendall 1914), but Strongylura marina is rare north of Cape Cod (Bigelow and Schroeder 1953; Fig. 2.8). We believe that the record from the Portland, ME, Atlantic Bluefin Tuna stomach (Crane 1936) was based on a misidentified saury, Scomb­ eresox saurus. Strongylura marina runs up rivers well above the limit of brackish water and reproduces in freshwater rivers and lakes (see Reproduction and Ecology sections above). Most species of Strongylura are allopatric. Strongylura marina is absent from the Bahamas and West Indies where Strongylura timu­ cu is found, but both species occur together along the coast from Florida to Brazil. The eastern Atlantic species of Strongylura is S. sengalensis and the eastern Pacific species is S. exilis. There seems to be a gap in the range along the southern coast of the Florida peninsula south to Miami. On the basis of collections from this area, Strongylura timucu is far more abundant in coastal waters. Study Material. A total of 790 specimens (22.5–418 mm BL), including the types of Belone truncata, B. almeida, and B. scrutator, from 179 collections originally at AMNH, ANSP, CAS, FMNH, FSU, MCZ, MNHN, MNRJ, MZUSP, NMC, TU, UA, UF, UMMZ, UPR, USNM, YPM, ZMH, and ZMK. The specimens came from four general areas: The east coast of the United States, 253 specimens (22.5–358 mm BL) from 101 collections; the U.S. coast along the Gulf of Mexico, 392 specimens (26.2–400 mm BL) from 119 collections; Central America, 51 specimens (49.4–418 mm BL) from 14 collections; and South America, 94 specimens (43.9–390 mm BL) from 45 collections.

Figure 2.8. Distribution of Atlantic Needlefish, Strongylura marina, and Timucu, Strongylura timucu, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

Etnier 1998; Fuller et al. 1999). It also ascends rivers in Mexico (Alvarez 1950), Venezuela (Mago Leccia 1965; Carvajal Rojas 1972), and Brazil (Starks 1913; Eskinazi 1972). Strongylura marina moves north along the Atlantic coast in the spring, appearing in February at Beaufort, NC (Yarrow 1877), and moving into Albemarle Sound in spring runs (Smith 1907). It arrives in the area from New York to Cape Cod from May to June and commonly remains until October (Bean 1890a; Smith 1898; Tracy 1910; Sumner et al. 1913; Latham 1919; Nichols and Breder 1927). Relationship to Humans. The primary interest in Strongylura marina is as a baitfish for important recreational species such as marlins (Manooch and Raver 1984). It will take a bait and has sometimes been considered a nuisance to anglers because of its bait-stealing habits (Smith and Bean 1899). Puyo (1949:51) noted that it is considered good to eat in French Guiana, “de chair excellente, il est estime des creoles.” The flesh is perfectly good to eat, but it has never found much favor in markets, perhaps partly because of its green bones (Bean 1883, 1890a; Fowler 1911; Hildebrand and Schroeder 1928; Thomson et al. 1971; Carvajal Rojas 1972). Range. Western Atlantic from Massachusetts south to around the Florida peninsula, along the coast in

Strongylura notata (Poey, 1860) Redfin Needlefish Figures 2.9, 2.10 Tables 2.1, 2.10, 2.11, 2.12 Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Redfin Needlefish (English); it is also Redfin Needlefish in the Bahamas (Böhlke and Chaplin 1968). Briggs (1958) used the name Southern Needlefish. In Cuba, its name in Spanish is agujón de aletas rojas (Rodríguez et al. 1984; Claro and Parenti 2001), but Page et al. (2013) list agujón negro. Diagnosis. Strongylura notata differs strongly from the other three species of western Atlantic Strongy­ lura (S. hubbsi, S. marina, and S. timucu) in having the maxilla completely concealed under the pre-orbital 30

Family Belonidae fin rays 10–12. Predorsal scales few and large, 76–117. Vertebrae 34–40 precaudal + 19–23 caudal = 53–61 total. Color. Bluish green above, silvery below. Distinct vertical bar at posterior margin of preopercle. Dorsal, anal, and caudal fins red or orange. Size. The maximum observed size for Strongylura notata is 411 mm SL, 245 mm BL (female, UF 44447, Key Biscayne, FL).

bone instead of exposed posteriorly, in having fewer anal rays (12–15 rather than 16–20), fewer vertebrae (53–61 rather than 64–77), and fewer predorsal scales (76–110 rather than 120–304). Analysis of molecular data (Lovejoy et al. 2004:369) shows that S. notata clusters closer to more primitive needlefishes such as Belone and sauries (Scomberesocidae) rather than to other species of Strongylura. As its common name Redfin Needlefish implies, S. notata is also distinguished by the brick-red color on the tips of the dorsal fin and upper caudal lobe. Two subspecies are recognized: Strongylura nota­ ta notata (Poey, 1860) from Cuba, Jamaica, Mexico, Belize, and Honduras, and S. n. forsythia Breder, 1932 from the Bahamas, Florida Keys, and southern Florida. Virtually all the available information on its biology is based on S. n. forsythia, but there is a little published information on parasitic copepods and predators of S. n. notata. Description. Body elongate, rounded in cross section. Upper and lower jaws greatly elongate and studded with sharp teeth; maxilla concealed under pre-orbital bone when jaws are closed. Gill rakers absent. Anterior parts of dorsal and anal fins not forming prominent lobes; pectoral fin not falcate. Caudal peduncle without lateral keels, deeper than wide; caudal fin emarginate, not deeply forked. Both right and left gonads present, right longer than left. Dorsal fin rays 12–15; anal fin rays 12–15; pectoral

Strongylura notata notata (Poey, 1860) Belone notata Poey, 1860:293–294 (original description, Cuba). Günther 1866:248 (description; after Poey 1860). Jordan and Gilbert 1881:459 (absence of gill rakers). Howell Rivero 1938:179 (type MCZ 32933). Mees 1962:57–58 (description; placed in genus Belone). Paepke and Seegers 1986:147 (Poey type at ZMB). Tylosurus notatus. Jordan and Gilbert 1883a:373 (description; range in part, West Indies). Bean and Dresel 1884:168 (description, Jamaica; USNM 32074). Jordan and Fordice 1887:345 (synonymy, in part, West Indies). Jordan 1887b:568 (listed; West Indies). Bean 1890b:206 (Cozumel, Yucatán, Mexico). Cockerell 1892:14 (listed; Jamaica). Jordan and Evermann 1896a:319 (in part). Jordan and Evermann 1896b:710–711 (description, range, in part). Evermann and Goldsborough 1902:151 (description, Progreso, Yucatán, Mexico). Strongylura notata. Jordan et al. 1930:195 (range, synonymy). Hubbs 1936:207 (description, Progreso, Yucatán, Mexico; UMMZ 102157, 102170). Buen 1940:45 (Yucatán, Mexico, after Hubbs 1936). Alvarez 1950:59 (description; Mexico). Fowler 1950:72 (description; Cayo Largo, Cuba). Springer

Figure 2.9. Redfin Needlefish, Strongylura notata notata. A. UMMZ 158864, 133 mm BL. Collected in Battery Point Beach, Bayport, Florida, U.S.A., 25 April 1948. B. TU 18200, 16.2 mm BL. Collected in Charlotte County, Florida, U.S.A., 5 May 1958. Illustrations by M. H. Carrington.

31

Fishes of the Western North Atlantic Table 2.10. Fin ray counts for Strongylura notata. Dashes indicate a value of zero. Fin rays Dorsal

Anal

Pectoral

Species and locality

12

13

14

15

12

13

14

15

10

11

12

Strongylura notata forsythia Bahamas Florida, U.S.A.

— —

37 51

51 105

5 7

— —

11 9

59 94

24 64

— 3

25 63

2 11

Subspecies total

—

88

156

12

—

20

153

88

3

88

13

Strongylura notata notata Cuba Jamaica Mexico Guatemala, Honduras, Belize

— 1 — 6

17 10 26 21

12 3 23 8

— — 1 —

— 1 1 2

6 9 9 19

17 2 30 15

6 — 10 —

10 — — 5

10 13 26 6

— — 1 —

7

74

46

1

4

43

64

16

15

55

1

Subspecies total

and Bullis 1956:57 (Golfo de Batabano, Cuba). Duarte-Bello 1959:43 (Cuba). Hildebrand et al. 1964:114 (Alecran Reef, Yucatán, Mexico). Miller 1966:796 (range, in part; Yucatán, Mexico, after Hubbs 1936). Birdsong and Emery 1968:192 (Turneffe Island, Belize, UMML 11715). Collette 1968:190 (in part; compared to S. marina and S. timucu). Randall 1968:39 (description, range, in part), fig. 39 (photograph). Cressey and Collette 1970:402 (parasitic copepods), 415 (fig. 179, range of species). Reséndez Medina 1971:24 (Alacran Reef, Yucatán, Mexico; after Hildebrand et al. 1964). Buen 1972:159 (predorsal scales). Duarte-Bello and Buesa 1973:79 (synonymy; range, in part). Guitart 1975:247 (Cuba, synonymy, description, fig. 182). Miller 1976a:148 (range, in part, after Miller 1966). Castro-Aguirre 1978:62 (Veracruz and Yucatán, Mexico). Hensel and Blahák 1978:87 (Rincon de Guarabo, 20 km east of Havana, Cuba). Reséndez Medina 1981a:268–269 (description; Laguna de Términos, Campeche, Mexico), 287 (pl. iv, fig. 29). Flores-Coto et al. 1983:152 (eggs and larvae; Laguna de Tamiahua). Rodríguez et al. 1984:15 (agujón de aletas rojas, Cuba). Kobelkowsky Díaz 1985:155 (Laguna de Tampamachoco, Veracruz, Mexico). Fuentes Mata et al. 1989:258 (listed, Laguna de Sontecomapan, Veracruz, México). Boughton et al. 1991 (ontogeny of upper jaw). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Ley et al. 1994:886–887 (feeds on fishes, insects, and crustaceans in northeastern Florida Bay). Obregón-Barboza et al. 1994:85 (Veracruz, Mexico). Schmitter-Soto and Gamboa-Pérez 1996:204 (Quintana Roo, Mexico). Greenfield and Thomerson 1997:93–95, 266 (description, Belize), fig. 97. McEachran and Fechhelm 1998:937, figure (Gulf of Mexico). Schmitter-Soto 1998:76–77, fig. 34 (upside down) (Quintana Roo, Mexico). Loftus 2000:34 (Everglades National Park). Lovejoy 2000:1352 (fig. 2, molecular phylogeny). Pérez-Hernández and Torres-Orozco 2000:434 (Mexican coastal lagoons). González-Gándara and Arias-González 2001:249 (Alacranes Reef, Gulf of Mexico). Claro and Parenti 2001:43 (agujón de aletas rojas, Cuba). Lovejoy and Collette 2001 (phylogeny). CastilloRivera et al. 2002:176 (Pueblo Viejo Lagoon, Veracruz, Mexico). Arceo-Carranza et al. 2004 (fishes predominate prey in Veracruz). Banford et al. 2004 (molecular phylogeny). González-Gándara 2004:167 (Veracruz). Reckel and Melzer 2004:15 (curtain-like falciform process in eye). ChávezLópez et al. 2005:148 (Alvarado Lagoon estuary, Veracruz,

Mexico). Miller et al. 2005:203–204 (range, habitat, Mexico, fig. 6.235). Tuckey and Dehaven 2006:109 (tidal creek habitat, Suwanee River estuary, FL). Taylor, Reyier, Davis, and McIvor 2007:884 and Taylor, Reyier, McIvor, and Davis 2007:728 (mangroves, Twin Cays, Belize). Campbell et al. 2008:83 (juveniles, mangrove habitat, Quintana Roo, Mexico). García-Hernández et al. 2009:92 (juveniles, northern Yucatán Peninsula). McEachran 2009:1285 (Gulf of Mexico). Miller et al. 2009:226–227 (range, habitat, fig. 6.235, map 6.201). Kells and Carpenter 2011:152–153 (description, color painting). Peralta-Meixueiro and Vega-Cendejas 2011:677 (hyperhaline coastal system: Ría Lagartos, Mexico). Gallardo-Torres et al. 2012:277 (permanent residents of the coastal lagoon Boca de la Carbonera, Yucatán, Mexico). Sosa-Lopéz et al. 2012:317 (abundance in the Laguna de Términos in southern Gulf of Mexico). Vega-Cendejas and Santillana 2012:298 (abundance in lagoon and coastal habitats in Yucatán, Mexico). Stevens et al. 2013:1650 (Peace River, tributary of Charlotte Harbor, FL). De Angelo et al. 2014:1042 (seagrass beds in eastern Gulf of Mexico). Gallardo-Torres et al. 2014:88 (Yucatán, Mexico, photographs). Humann and DeLoach 2014:64–65 (description, range; color photograph). Strongilura [sic] notata. Claro 1981:98 (eaten by Lutjanus syn­ agris; Golfo de Batabano, Cuba). Strongylura notata notata. Collette 2003a:1110 (figure, description, range). Collette 2003b:7 (synonymy). Smith et al. 2003:15 (Belize). Lovejoy et al. 2004:369, 377 (molecular phylogeny). Arévalo-Frías and Mendoza-Carranza 2012:259 (larvae and juveniles from the Biosfera Patanos Reserve in Tabasco, Mexico).

Misidentifications. The specimens from Progreso, Yucatán, Mexico (MCZ 32903), that Barbour and Cole (1906) reported as Tylosurus marinus are Strongy­ lura notata notata. Types. Belone notata Poey, 1860. Lectotype, designated herein, MCZ 32933 (157 mm BL); Havana, Cuba; F. Poey. Dorsal fin rays 13; anal fin rays 13; pectoral fin rays 10–10. Predorsal scales 82. Vertebrae 35 precaudal + 20 caudal = 55 total. Paralectotype ZNB 5498 (158 mm BL). 32

33

— — — — — — — — — — — — 3 — 3 1 — 7 — 8 4 — 14 — 18 11 1 3 10 25 1 4 1 6 12 — — — 1 1 — — — 1 1 — — — — — — — — — — 4 — 3 5 12 10 1 22 8 41 3 4 3 4 14 — — — — — — — — — — 1 — 2 — 3 4 — 6 1 11 — — 1 1 2

2 — 2 11 15

1 — Subspecies total

Strongylura notata notata Cuba Jamaica Mexico Guatemala, Honduras, Belize Subspecies total

10 5 17 5 37

2 25 69 62 23 3 — — — 3 62 17 102 — 1 8 48

1 — — — Strongylura notata forsythia Bahamas Florida, U.S.A.

24 100

5 20 28 41 29 33 10 13 3 — — — — — — — 1 2 31 31 35 67 8 9 — — — 1 — 8 12 36 49 51

22 21 40 39 38 37 36 35 34 Species and locality

Precaudal

Table 2.11. Vertebral counts of Strongylura notata. Dashes indicate a value of zero.

19

20

Caudal

Vertebrae

Predators. The only specific predator reference is to Lutjanus synagris from the Golfo de Batabano, Cuba (Claro 1981), but it is undoubtedly eaten by other fishes. Parasites. Three species of parasites have been reported from western Atlantic Strongylura notata notata, all of which are copepods. See account of S. n. forsythia for other groups of parasites found on S. no­ tata. copepods: Three species were reported by Cressey and Collette (1970): Bomolochidae, Bomolochus bel­ lones Burmeister (genus following Ho and Lin 2009); Anthosomatidae, Lernanthropus belones Krøyer; and Philichthyidae, Colobomatus goodingi Cressey and Collette. Bomolochus bellones was collected under the opercles and under the oral valves in Jamaica, Yucatán, and Tampico, Mexico; L. belones was attached to the gill filaments in specimens from Cuba, Jamaica, and Yucatán; and C. goodingi was in the cephalic lateral line canals of specimens at Providence Island. Food. Feeds on fishes, insects, and crustaceans in northeastern Florida Bay, lunging downward on benthic prey from above, skimming prey from the surface, and leaping out of the water to capture insects from mangroves (Ley et al. 1994). The contents in 89 digestive tracts of specimens from the Alvarado lagoonal system, Veracruz, Mexico, showed a diet of 29 prey types, with the dominant prey consisting of fishes, penaeid shrimps, polychaetes, and hymenopteran insects (Arceo-Carranza et al. 2004). Range. Cuba; Isle of Pines; Jamaica; coastal Mexico from the states of Tamaulipas (35 miles north of Tampico, UMMZ 157319), Veracruz (Tamiahua, UMMZ 167536, 184419), Quintana Roo (Schmitter-Soto and Gamboa-Pérez 1996; Campbell et al. 2008), and Yucatán; Belize; Guatemala; Bonac-

23

Biology

13 11

57 53

54

55

56

Total

58

59

60

61

Diagnosis. Strongylura notata notata differs from S. n. forsythia primarily in having fewer predorsal scales, usually less than 90 (x̅  84.2, rather than x̅  95.3), and fewer vertebrae, usually less than 57 (x̅  56.4, rather than x̅  58.6). It averages slightly fewer dorsal fin rays (12–15, x̅  13.3, rather than 13–15, x̅  13.7), and anal fin rays (12–15, x̅  13.8, rather than 13–15, x̅  14.2). Description. Dorsal fin rays 12–15; anal fin rays 12–15; pectoral fin rays 10–12, usually 11 (Table 2.10). Vertebrae 34–38 precaudal + 19–22 caudal = 53–59 total (Table 2.11). Predorsal scales 79–96 (Table 2.12). Size. The maximum observed size for Stron­ gylura notata notata is 200 mm BL (UMMZ 102170, Yucatán, Mexico).

— 2

Family Belonidae

Fishes of the Western North Atlantic Table 2.12. Number of predorsal scales of Strongylura notata. Dashes indicate a value of zero. Predorsal scales Species and locality Strongylura notata forsythia Bahamas Florida, U.S.A. Subspecies total Strongylura notata notata Cuba Jamaica Mexico Guatemala, Honduras, Belize

Subspecies total

76–77

78–79

80–81

82–83

84–85

86–87

88–89

90–91

92–93

94–95

— 1

— —

— 1

— 5

2 6

5 10

9 10

2 9

6 11

7 8

1

—

1

5

8

15

19

11

17

15

1 — 1

2 2 2

4 — 3

6 — 6

6 2 6

6 3 7

1 3 7

1 2 4

— — —

1 — —

2 4

4 10

7 14

9 21

2 16

3 19

1 12

—

— —

—

7

1

Predorsal scales Species and locality Strongylura notata forsythia Bahamas Florida, U.S.A. Subspecies total Strongylura notata notata Cuba Jamaica Mexico Guatemala, Honduras, Belize

Subspecies total

96–97

98–99

100–101

102–103

104–105

7 10

3 7

2 7

— 6

1 2

17

10

9

6

— — 1

— — —

— — —

— — —

106–107

108–109

110

2 2

— 4

— 1

3

4

4

1

— — —

— — —

— — —

— — —

—

—

—

—

—

—

—

—

1

—

—

—

—

—

—

—

ca and Utila islands, Honduras; and Providence Island off Nicaragua (ANSP uncatalogued; Fig. 2.10). Records from Puerto Rico (Silvester 1918; Fowler 1928a) and Martinique (Louis et al. 1992) have not been verified and are presumably attributable to Strongylura timucu. The report from Trinidad by Fowler (1931) was based on specimens of S. timucu (ANSP 72850, 88540). Perhaps this misidentification led Román (1979) to include S. notata in his key to Venezuelan fishes. Study Material. A total of 100 specimens (8.6–200 mm BL), including the lectotype of Belone notata Poey (MCZ 32933), originally in AMNH, ANSP, CAS, FMNH, UCLA, LACM, MCZ, UF, UMMZ, and USNM. Specimens come from five areas: 24 specimens from Cuba (52.0–183 mm BL); 13 specimens from Jamaica (36.6–167 mm BL); 38 specimens from Mexico (35–200 mm BL); 21 specimens from Belize and Honduras (8.6–150 mm BL); and 3 specimens from Old Providence Island off Nicaragua (61.7–63.4 mm BL).

Strongylura notata forsythia Breder, 1932 Tylosurus notatus. Bean 1883:450 (Charlotte Harbor, FL). Jordan 1884:111 (Key West, FL, USNM 34998). Jordan and Fordice 1887:345–346 (description, Key West). Jordan and Bollman 1889:550 (Green Turtle Cay, Bahamas, CAS IU 8818–9). Henshall 1891:375 (Card’s Sound and Black Sound, FL). Henshall 1895:212 (Key West and Tampa, FL). Smith 1896:175 (Key Biscayne, FL). Jordan and Evermann 1896a:319 (in part). Jordan and Evermann 1896b:710–711 (description, range, in part). Evermann and Bean 1898:242 (description, Titusville, FL). Evermann and Kendall 1899:61 (Florida). Bean 1905:300 (Bahamas). Fowler 1906:90 (description, Florida Keys, fig. 6). Rosen 1911:51 (Bahamas). Fowler 1915a:247 (Palm Springs, FL). Nichols 1921:22 (Turk Islands, listed). Schroeder 1924:5 (Key West, FL). Parr 1930:19 (description, Crooked Island, Bahamas, YPM ICH 002618). Strongylura notata. Fowler 1919a:3 (nine southern Florida localities). Fowler 1919b:152 (Useppa Island, FL). Breder 1927:8 (Bahamas). Breder 1929:279–280 (mature at 300 mm SL, Tortugas). Breder 1932a:3–5 (growth, FL), pl. 1, fig. 2 (jaw development), pl. 3, fig. 2 (tails). Bere 1936:598 (parasitic copepod Lernanthropus chlamydotus, Lemon Bay, FL).

34

Family Belonidae Storey and Gudger 1936:643 (mortality due to cold, January 1925, Sanibel Island, FL). Storey 1937:16 (cold mortality, range north to Pensacola, FL). Shuler 1938:56 (cestode Ptychobothrium belones, Tortugas). Fowler 1940b:11 (Boca Grande, FL). Meehean 1940:517–518 (Argulus bicolor, Lemon Bay, FL). Fowler 1941c:83 (Coconut Grove and Sanibel Island, FL). Longley and Hildebrand 1941:27 (ecology, Tortugas). Fowler 1945:281 (FL localities). Breder 1946:4–5 (behavior, Palmetto Key, FL). Manter 1947:376 (no trematodes in 22 specimens, Tortugas). Chandler 1954:353 (cestode Ptychobothrium belones, Tortugas, after Shuler 1938). Reid 1954:21 (ecology, food, Cedar Keys, FL). Kilby 1955:194–195 (ecology, Cedar Keys, FL). Joseph and Yerger 1956:126–127 (description, ecology, most abundant needlefish, Alligator Harbor, FL, several FSU collections). Briggs 1958:264 (range, in part). Breder 1959a:141–147 (spawning behavior, egg development, Pine Island Sound and Lemon Bay, FL). Breder 1959b:422, 428 (no departure from randomness when given choice between lighted and dark chambers). Springer 1960a:27 (ecology, Caloosahatchie River area, Florida). Springer and Woodburn 1960:24 (ecology, Tampa Bay, FL), 53 (seasonality, Matecumbe Key, FL). Moss 1961:347 (acellular bone, Bimini). Tabb 1961:20 (in stomachs of 7 of 84 Cynoscion nebulosus, Florida). Berry and Rivas 1962 (synonymy, description, range, in part). Tabb and Manning 1962:611 (ecology, food, Florida Bay). Tabb et al. 1962:51 (salinity, Florida Bay). Gunter and Hall 1963:597 (salinity, St. Lucie Inlet, FL). Rinckey and Saloman 1964:14 (Tampa Bay, FL). Gooding and Collette 1965 (parasitic copepod Colobomatus). Nahhas and Short 1965:41 (digenetic trematode Rhipidocotyle transversale, Apalachee Bay, FL). Breder and Rosen 1966:302 (reproduction, after Breder 1932a, 1959a). Collette 1966:17 (fig. 5a, caudal skeleton of Bimini specimen). Miller 1966:796 (range, in part). Böhlke and Chaplin 1968:117 (description, figure, Bahamas). Moe et al. 1966:16 (10 Florida collections). Starck 1968:17 (Alligator Reef, FL). Finucane 1969:290 (mortality from antimycin, Tampa Bay, FL). Rees 1969:539–540 (cestode Ptychobothri­ um belones, Tortugas, after Shuler 1938). Swarts 1969:329 (blood studies, Bahamas). Cressey and Collette 1970:402 (parasitic copepods), 415 (fig. 179, range of species). de Sylva 1970:51 (Biscayne Bay, FL). Hudson et al. 1970:10 (central Florida Bay). Roessler 1970:864 (seasonal abundance), 883 (temperature and salinity, Buttonwood Canal, FL). Wang and Raney 1971:26 (ecology, Charlotte Harbor, FL). Moe 1972:10 (movements, Florida). Odum and Heald 1972:675 (North River, FL). Powell et al. 1972:119 (12 Florida collections). Kushlan and Lodge 1974:119 (St. Lucie River, FL, after Gunter and Hall 1963). McNulty et al. 1974:108, 120– 121, 192–193 (range in Gulf of Mexico along Florida). Subrahmanyam and Drake 1975:451–456 (Wakulla, FL). Brook 1977:226 (Card Sound, FL, fish larger than 200 mm SL are piscivores, preying particularly on silversides). Stevenson 1977:182 (fresh water records, southern Florida). Gilmore 1977:132 (Indian River area, Florida). Gilmore 1978:147 (spawn February–July, Indian River Lagoon, FL). Gilmore et al. 1978:94 (killed by cold, central Florida, previous records). Loftus and Kushlan 1987:202 (fresh water records, southern Florida), 203 (fig. 24, distribution records). Naughton and Saloman 1978:46, 50 (St. Andrew Bay, FL). Skinner 1978:590, 593, 594 (parasites, Biscayne Bay, FL). Snelson and Bradley 1978:6 (cold mortality, Mosquito Lagoon, FL). Hastings 1979:7 (jetties, St. Andrew, FL). Love 1980:401 (reference to Swarts 1969). Subrahmanyam and Coultas

1980:800 (seasonality, Apalachee Bay, FL). Gilmore et al. 1981:10 and Gilmore et al. 1982:34–36 (Indian River Lagoon, FL). Snelson 1983:192 (ecology, Indian River Lagoon, FL). Bruce 1986:1116 (parasitic isopod Mothocya xenobran­ chia, Florida). Provancha et al. 1986 (cold mortality, Merritt Island, FL). Gilmore 1987:122, 129 (seagrass communities). Sogard et al. 1987:3 (Florida Bay). Schmidt 1989:167 (eaten by young barracuda, Everglades National Park, FL). Sogard et al. 1989a:58, 1989b:195 (Florida Bay). Thayer and Chester 1989:210 (Florida Bay). Brown-Peterson and Eames 1990:230 (associated with spoil islands, Indian River Lagoon, FL). Cressey 1991:4 (parasitic copepod Caligus bery­ chis). Szedlmayer 1991:78 (juveniles, Anclote River estuary, Florida). Smith 1997:393 (description, range). Poulakis et al. 2003:174 (Charlotte Harbor, FL). Banford et al. 2004 (molecular phylogeny). Porter and Motta 2004:993–994 (feeding kinematics). Poulakis et al. 2004:126 (Charlotte Harbor, FL). Collette 2005:924–925 (early life history, figs. A–C). Idelberger and Greenwood 2005:228 (Myakka and Peace rivers, southwestern Florida). Powell et al. 2007:74–75, 209 (densities and monthly length-frequencies, Florida Bay). Vega-Cendejas et al. 2012:141 (length-weight, Yucatán peninsula). Snyder and Burgess 2016:102 (listed, Florida, color photograph). Strongylura notatus. Breder 1929:279–280 (habits, mature at 300 mm SL, Tortugas). Breder 1948:88 (description, Florida north to Pensacola). Strongylura notata forsythia Breder, 1932b:3–4 (original description, Lake Forsyth, Andros Island, Bahamas), fig. 2 (type). Breder 1934:70, 73 (biology, Lake Forsyth, Bahamas), fig. 32. Collette 2003a:1110 (figure, description, range). Collette 2003b:7 (synonymy). Lovejoy et al. 2004:369, 377 (molecular phylogeny). Starck et al. 2017:91 (Florida Keys). Tylosurus notata. Menzel 1956:51 (Apalachee Bay, FL). Stronglura [sic] notatus. Voss 1953:229–230 (eaten by Sailfish, Florida).

Types. Strongylura notata forsythia Breder, 1932. Holotype, AMNH 10108 (144 mm BL); Bacon-Andros Expedition; February 1932. Diagnosis. Strongylura notata forsythia differs from S. n. notata primarily in having more predorsal scales, usually more than 90 (x̅  95.3 rather than x̅  84.2) and more vertebrae, usually more than 57 (x̅  58.6 rather than x̅  56.4). It averages slightly more dorsal fin rays (13–15, x̅  13.7, rather than 12–15, x̅  13.3), and anal fin rays (13–15, x̅  14.2, rather than 12–15, x̅  13.8). Description. Dorsal fin rays 13–15; anal fin rays 13–15; pectoral fin rays 10–12, usually 11 (Table 2.10). Vertebrae 35–40 precaudal + 20–23 caudal = 56–61 total (Table 2.11). Predorsal scales 84–117 (Table 2.12). Size. The maximum observed size for Strongylura notata forsythia is 245 mm BL (411 mm SL female, UF 44447, Key Biscayne, FL). Biology Predators. Preyed on by a variety of surface-feeding fishes, such as Spanish Mackerel, Kingfish, saltwater 35

Fishes of the Western North Atlantic isopod, Cymothoidae, Mothocya xenobranchia from Strongylura notata from Florida. branchiurans: One species of branchiuran is known from Strongylura notata: Argulus bicolor Bere, 1936, from Lemon Bay, FL (Bere 1936; Meehean 1940; Skinner 1978). cestodes: Ptychobothridae, Ptychobothrium belones (Dujardin), from the Tortugas (Shuler 1938; Chandler 1954; Rees 1969). trematodes: Bucephalidae, Rhipidocotyle trans­ versale Chandler, from the intestine of Strongylura notata, Apalachee Bay, FL (Nahhas and Short 1965). A digenetic trematode, Steganoderma elongatum, was described by Manter (1947) from three species of needlefishes from the Tortugas, Strongylura timucu, Tylosurus acus, and T. crocodilus, but the trematode was absent in the 23 specimens of Strongylura notata examined. monogenes: Dactylogyridae, Ancyrocephalus parvus Linton, was reported as a new host record from the gills of Strongylura notata from Biscayne Bay, FL (Skinner 1978). Food. Feeds on small fishes (Breder 1934; Reid 1954; Subrahmanyam and Drake 1975), including larval anchovies and Ladyfish (Tabb and Manning 1962) and shrimps (Tabb and Manning 1962; Subrahmanyan and Drake 1975), and there is a record of a dragonfly naiad from the stomach of a Strongylura notata forsythia from the type locality, Lake Forsyth, Andros Island, Bahamas (Breder 1934). Feeding kinematics were described and illustrated by Porter and Motta (2004). Reproduction. Spawning takes place from February to July in the Indian River Lagoon (Gilmore 1978). Both left and right gonads are developed, ratio of left to right is 0.8–1.4. A 248 mm BL female contained 428 eggs in the left ovary (1.8–2.2 mm in diameter) and 470 eggs in the right ovary (1.6–1.9 mm in diameter), a total of 898 eggs (x̅  1.85 mm in diameter). Diameter of ripe eggs ranged from 3.67 to 4.18 mm (x̅  3.95 mm in diameter) and the eggs were covered with uniformly distributed filaments (Collette et al. 1984:336). Breder (1959a) described spawning behavior and egg development. Development. The halfbeak stage, in which the length of the lower jaw exceeds that of the upper jaw by a significant distance, occurs from about 14 mm SL (Breder 1932a) to at least 16.2 mm BL (Collette 2005:925, fig. C). Breder (1932a: pl. 1, fig. 2) illustrated the relative lengths of upper and lower jaws at 14, 123, and 405 mm SL. Relationship to Humans. Needlefishes are not commonly eaten by people in the United States, but are certainly good to eat, as is well known in Australia,

Figure 2.10. Distribution of Redfin Needlefish, Strongylura notata notata and S. notata forsythia, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

trout (Henshall 1891), Cynoscion nebulosus (present in 7 of 84 stomachs; Tabb 1961), Sailfish (Voss 1953), and young barracuda (Schmidt 1989). Parasites. Ten species of parasites have been reported from western Atlantic Strongylura notata for­ sythia: four copepods, one isopod, one branchiuran, one cestode, two trematodes and a monogene. copepods: Cressey and Collette (1970) found four species of copepods: Bomolochidae, Bomolochus bel­ lones (genus following Ho and Lin 2009); Caligidae, Caligus sp. B; Anthosomatidae, Lernanthropus belones; and Philichthyidae, Colobomatus goodingi. Bomolochus bellones was common in the gill chambers and under the oral values at Andros Island, Bahamas, and Sarasota, New Smyrna, Pensacola, Bayport, Cape Haze, and Key Biscayne, FL; both sexes of L. belones were common on the gill filaments at Bimini and Andros Island, Bahamas, and at Sanibel Island, Cape Haze, Sarasota, Pensacola, and Key Biscayne, FL; Colobo­ matus goodingi lives in the cephalic lateral line canals and was collected at Bimini, Bahamas, and Tampa Bay, Cape Haze, Sanibel Island, and Key Biscayne, FL; Caligus sp. B was found on only two specimens, from Sarasota, FL. Later, this Caligus was identified as Caligus berychis Wilson, 1936 by Cressey (1991). Bere (1936) also reported L. bellones (as L. chlamydotus Wilson) from Lemon Bay, FL. isopods: Bruce (1986) reported one species of 36

Family Belonidae India, and elsewhere. There is one early report that the steward of the Grampus cooked Strongylura notata from Charlotte Harbor and that they were “greatly relished by the crew,” reminding one “somewhat of smelts” when fried (Henshall 1891:375). Range. Bahama Islands, Florida Keys, and southern Florida north to New Smyrna (USNM 62663) on the U.S. Atlantic coast and north and west to Pensacola, FL (USNM 188849), and Mobile Bay, AL (UMMZ 157824) on the U.S. Gulf coast (Fig. 2.10). Reports from Texas (Baughman 1950; Murdy 1983) are probably based on misidentified Strongylura marina. A report from Bermuda (Beebe and Tee-Van 1933) is highly unlikely and there has been no subsequent verification (Smith-Vaniz et al. 1999). Study Material. A total of 334 specimens (16.2–245 mm BL), including the holotype of Strongylura notata forsythia (AMNH 10108), originally at AMNH, ANSP, CAS, CU, FSU, MCZ, TU, UF, UMMZ, USNM, and YPM. From the Bahamas, 95 specimens (52.3–173 mm BL); from Florida, 239 specimens (16.2–245 mm BL).

dan and Evermann 1896a:320 (listed, Cuba and Jamaica). Jordan and Evermann 1896b:711–712 (description, Cuba and Jamaica). McIndoo 1907:487 (Batabano, Cuba). Tylosurus diplotaenia. Jordan and Fordice 1887:346 (after Cope 1871). Jordan 1887b:568 (listed, West Indies). Jordan and Evermann 1896a:320 (listed, after Cope 1871). Jordan and Evermann 1896b:712 (description, after Cope 1871). Tylosurus almeida. Jordan and Fordice 1887:352–353 (synonymy, in part, description, range). Jordan 1887b:568 (listed, West Indies). Eigenmann and Eigenmann 1891:65 (listed, Suriname). Jordan and Evermann 1896a:320 (listed, Rio de Janeiro, Brazil, to the West Indies). Jordan and Evermann 1896b:715 (description, range). Starks 1913:40 (description, in part, Lake Papary, Natal, Brazil, based on Strongylura timucu and S. marina, CAS-SU 64919). Tylosurus timucu. Jordan and Evermann 1896a:320 (listed, Florida Keys to Brazil). Jordan and Evermann 1896b:711 (description, synonymy, range). Evermann and Kendall 1889:61 (Florida). Evermann and Marsh 1902:99 (Puerto Rico). Miranda Ribeiro 1915:17–18 (description, Pernambuco, Brazil). Fowler 1915b:50 (Santo Domingo). Meek and Hildebrand 1923:229–230 (description, synonymy, Panama, USNM 79634, 79648, 79644, 79645). Schroeder 1924:5 (Key West, FL). Paiva and Holanda 1962:3 (description, Paracuru, Ceará, Brazil). Belone (Tylosurus) timucu. Metzelaar 1919:30–31 (description, Curaçao), fig. 10. Strongylura timucu. Fowler 1919a:3 (Colon, Panama and Santa Domingo). Fowler 1919b:136–137, 146 (Jamaica, type of Be­ lone diplotaenia). Fowler 1926:250 (description, Boca Grande, FL). Jordan et al. 1930:195 (range, synonymy, in part). Breder 1932a:7 (description, Panama, Tortugas). Breder 1932b:4 (Lake Forsyth, Andros Island, Bahamas). Breder 1934:70, 73 (food, maturity, Andros Island, Bahamas). Bere 1936:582, 598 (parasitic copepods, Lemon Bay, FL). Fowler 1940c:755 (Rio de Janeiro, Brazil, Wilkes Expedition, USNM 83428), fig. 25. Fowler 1941d:141 (Brazilian records). Longley and Hildebrand 1941:28–29 (description, allopatric to Strongylura marina in the Tortugas). Gunter 1942:317 (occurrence in fresh water in Central America). Manter 1947:375 (digenetic trematode described from Tortugas specimen). Fowler 1948:50 (color, Puerto Rico). Breder 1948:88 (description, Florida Keys). Alvarez 1950:59 (in key to Mexican fishes). Fowler 1951a:26 (description, St. Croix), fig. 2. Fowler 1952b:87 (Hispaniola, previous records). Fowler 1952c:10–11 (description, Rio de Janeiro, Brazil, USNM 83428). Manter 1954:339–340 (trematodes based on previous records). Reid 1954:21–22 (ecology, Cedar Key, FL, based on S. timucu and S. marina). Kilby 1955:194–195 (ecology, Cedar Key and Bayport, FL, based on S. timucu and S. marina). Gunter 1956:350 (listed, North American euryhaline species). Briggs 1958:264 (range). Yamaguti 1958:56 (digene Steganoderma elongatum). Duarte-​ Bello 1959:43 (Cuba). Sogandares-Bernal and Hutton 1959:259–260 and Hutton and Sogandares-Bernal 1960: 288 (digenetic trematode, Bayboro Harbor, FL). Springer 1960a:27 (ecology, Caloosahatchie River, FL). Springer and Woodburn 1960:24 (ecology, Tampa Bay, FL). Randall and Randall 1960:444, 471–472 (immature resemble floating manatee grass, Virgin Islands). Springer 1960b:19 (ecology, St. Lucie River and Indian River, Florida). Tabb and Manning 1962:611 (Florida Bay). Springer and McEarlean 1962:53 (seasonality, Matecumbe Key, FL). Nomura and Menezes 1964:354 (previous records, Brazil).

Strongylura timucu (Walbaum, 1792) Timucu Figures 2.8, 2.11 Tables 2.1, 2.13, 2.14, 2.15 Esox timucu Walbaum, 1792:88 (original description based on Marcgrav). Collette 1968 (designation of neotype). Belone cigonella Valenciennes in Cuvier and Valenciennes, 1846:436–437 (original description, Puerto Rico). Jordan 1887a:529 (types could not be found at MNHN). Cockerell 1892:14 (listed, Jamaica). Poey 1881:337 (Puerto Rico, listed after Valenciennes in Cuvier and Valenciennes, 1846). Belona timucu. Castelnau 1855:52 (description, common in markets of Rio de Janeiro and Bahia, Brazil). Belone subtruncata Poey, 1860:295–296 (original description, Havana, Cuba). Günther 1866:245 (after Poey 1860). Belone depressa Poey, 1860:296 (original description, Havana, Cuba, USNM 4745). Günther 1866:235–236 (in part, Dominica, Jamaica). Cockerell 1892:14 (listed, Jamaica). Belone diplotaenia Cope, 1871:481 (original description, St. Martins, West Indies, ANSP 7572). Tylosurus sagitta Jordan and Gilbert, 1884:25–26 (original description, Key West, FL, USNM 34965). Jordan 1884:111– 112 (Key West, CAS-SU 1891). Tylosurus subtruncatus. Jordan and Fordice 1887:346–347 (synonymy, range, description). Jordan 1887b:568 (listed, West Indies). Jordan 1891:315 (Bahia, Brazil). Belone timucu. Kner 1867:322 (description, Rio de Janeiro, Brazil). Jordan 1887a:529–530 (description, Cuba, Cayenne, French Guiana). Tylosurus euryops Bean and Dresel, 1884:168 (original description, Jamaica, USNM 32073). Jordan 1886b:30 (Havana, Cuba, synonym of Belone depressa Poey?). Jordan and Fordice 1887:347–348 (description, range). Jordan 1887b:568 (listed, West Indies). Jordan 1890:647 (Port Castries, St. Lucia, West Indies). Cockerell 1892:14 (listed, Jamaica). Jor-

37

Fishes of the Western North Atlantic Nahhas and Cable 1964:223 (digenetic trematodes, Curaçao). Carvalho 1964:7 (common names). Cable and Michaels 1967 (cestode, Jamaica). Collette 1967 (validity of S. timucu). Randall 1967:683–684 (food 96% fishes, 4% shrimps, Virgin Islands). Collette 1968 (validity of S. timu­ cu, synonymy, description, designation of neotype) fig. 1 (distribution). Böhlke and Chaplin 1968:118 (description, Bahamas, figure). Birdsong and Emery 1968:192 (Turneffe Island, Belize). Randall 1968:39 (description), fig. 38 (color photograph). Finucane 1969:290 (antimycin as a toxicant, Tampa Bay, FL). Overstreet 1969:131, 171 (trematode Schik­ hobalotrema acutum, Biscayne Bay, FL). Cressey and Collette 1970:402–403 (parasitic copepods). de Sylva 1970:51 (Biscayne Bay, FL). Collette and Parin 1970:23, 25–27 (comparison with Strongylura marina and S. senegalensis). Mago Leccia 1970:89 (listed, Venezuela). Roessler 1970:864, 883 (ecology, Everglades National Park). Dahl 1971:180 (Isla del Rosario, Colombia, curajota). Gilbert and Kelso 1971:27 (Tortuguero, Costa Rica). Iversen et al. 1971:84 (Myxosporidia, Buttonwood Canal and Biscayne Bay, FL). Wang and Raney 1971:26 (abundance, Charlotte Harbor, FL). Buen 1972:159 (135 predorsal scales, Mexico). Kähsbauer 1972:717 (description, Puerto Cortés, Honduras). Roux 1973:72 (description, north of Rio de Janeiro, Brazil). Collette 1974c:612 (table 1, comparison with other three western Atlantic species, figure). Brownell and Guzmán 1974:128 (Isla de Aves, Lesser Antilles). Kushlan and Lodge 1974:119 (fresh water records, southern Florida). Guitart 1975:248 (Cuba, synonymy, description, fig. 183). Miller 1976a:155 (distribution after Collette [1968], enters rivers). Miller 1976b:18 (penetrates fresh water, San Francisco, near Veracruz, Mexico, FMNH 4568). Erdman 1976: 23 (ripe in June, northeastern Caribbean Sea). Shiino 1976:92 (common name, timucu). Gilmore 1977:132 (Indian River Lagoon, FL, previous records). Stevenson 1977: 183 (description, range). Castro-Aguirre 1978:61–62 (Atlantic Mexican freshwater records). Collette 1978 (description, comparisons, range, figure). Figueiredo and Menezes 1978:64 (description, Brazil), fig. 110. Gilmore et al. 1978:82 (occasional hypothermal mortality, Indian River and Tampa Bay, FL). Lindén et al. 1978:254 (Cartagena Bay, Colombia). Sazima and Uieda 1979 (feeding, camouflage, Rio Escuro, Ubatuba, São Paulo, Brazil). Román 1979:83 (in key, Venezuela, figure). Gilmore et al. 1981:10 (Indian River Lagoon, FL). Gilmore et al. 1982:33, 34, 36 (inpoundments, Indian River Lagoon, FL). Lucena and Lucena 1981:41–42 (Recife and Lagoa do Mirim, Tubarão, Brazil). Skinner 1982:275 (trematode Ancyrocephalus parvus, Biscayne Bay, FL). Snelson 1983:192 (rare in Indian River Lagoon, FL). Böhlke 1984:27 (type of Belone diplotaenia). Collins and Finucane 1984:45 (ichthyoplankton abundance off Everglades City, FL). Rodríguez et al. 1984:15 (timúcu, Cuba). Acero P. et al. 1984:41 (reefs, Caribbean Colombia). Acero P. and Garzón F. 1987:90 (Colombia). Corrêa et al. 1986:18 (Brazil). Gilmore 1987:129 (seagrass communities). Loftus and Kushlan 1987:203–204 (commonly enters fresh water in southern Florida, fig. 24, locality records). Martins-Juras et al. 1987:109 (Ilha de São Luis, Maranhão, Brazil). Paiva Filho and Toscano 1987:157 (São Paulo, Brazil). Thayer et al. 1987:30 (abundant in red mangrove prop root habitat, southern Florida). Garzón F. 1989:155 (Colombia). Lopes 1989:211 (distribution; Brazil). Sogard et al. 1989a:58, 1989b:195 (Florida Bay). Boughton et al. 1991 (ontogeny of upper jaw). Cervigón 1991:230–231 (description, Venezuela, fig. 171 [upside down]). Cervigón et al. 1993:269 (north-

ern coast of South America). Bunkley-Williams and Williams 1994:35, 89, 131 (parasites in Puerto Rico). Araújo 1995:119, 122 (baixo, Paraíba do Sul River, Rio de Janeiro state, Brazil). Giannini and Paivo Filho 1995:144 (São Paulo, Brazil). Schmitter-Soto and Gamboa-Pérez 1996:204 (Quintana Roo, Mexico). Greenfield and Thomerson 1997:94–96, 266 (description, Belize), fig. 99. Smith 1997:393 (description, range). McEachran and Fechhelm 1998:938, figure (Gulf of Mexico). Schmitter-Soto 1998:77– 78 (Quintana Roo, Mexico). M. E. de Araújo et al. 2000:140 (estuaries, Ceará, Brazil). Loftus 2000:34 (Everglades National Park). Lovejoy 2000:1352 (fig. 2, molecular phylogeny). Pérez-Hernández and Torres-Orozco 2000:434 (Mexican coastal lagoons). Vendel et al. 2000:180 (tidal flat, Paraná, Brazil). Claro and Parenti 2001:43 (timucú, Cuba). Lovejoy and Collette 2001 (phylogeny). Paperno et al. 2001:127 (Ponce de Leon Inlet, FL). Castillo-Rivera et al. 2002:176 (Pueblo Viejo Lagoon, Veracruz, Mexico). Collette 2003a:1111 (figure, description, range). Collette 2003b:8 (synonymy). Menezes and Figueiredo 2003:67 (Brazil). Pessanha et al. 2003:1049 (Itacuruçá beach, southeastern Brazil). Poulakis et al. 2003:174 (Charlotte Harbor, FL). Smith et al. 2003:15 (Belize). Banford et al. 2004 (molecular phylogeny). Frota et al. 2004:23 (length-weight, Brazil). Lovejoy et al. 2004:369, 377 (molecular phylogeny). Poulakis et al. 2004:126 (Charlotte Harbor, FL). Spach et al. 2004:51 (Brazil). Springer and Johnson 2004:129–130 (dorsal gill arch musculature). Chávez-López et al. 2005:148 (Alvarado Lagoon estuary, Veracruz, Mexico). Collette 2005:926–927 (early life history, fig. A). Idelberger and Greenwood 2005:228 (Myakka and Peace rivers, southwestern Florida). Miller et al. 2005:204 (range, habitat, Mexico, fig. 6.236). Giarrizzo et al. 2006:326 (weightlength relationships, Curuçá mangrove estuary, northern Brazil). Kohn et al. 2006:70 (monogene from Puerto Rico). Tuckey and Dehaven 2006:109 (tidal creek habitat, Suwanee River estuary, Florida). Barletta and Blaber 2007:651–652 (Caeté estuary, northern Brazil). Giarrizzo and Krumme 2007:746–747 (Curuçá mangrove estuary, northern Brazil). Greenwood et al. 2007:812, 814 (mangroves, Charlotte Harbor, FL). Krebs et al. 2007:846 (mangroves, Tampa Bay, FL). Luque and Tavares 2007:13, 16 (parasitic copepods, Brazil). Powell et al. 2007:74–75, 209 (densities and monthly length-frequencies, Florida Bay). Rehage and Loftus 2007:631 (mangrove creeks, southwestern Everglades). Félix et al. 2007:286 (beaches, Paraná, Brazil). Taylor, Reyier, McIvor, and Davis 2007:728 (mangroves, Twin Cays, Belize). Gaelzer and Zalmon 2008a:26, 2008b:168, 173 (sandy beaches, southeastern Brazil). García-Hernández et al. 2009:92 (juveniles, northern Yucatán Peninsula). Gómez Gaspar and Hernández Avila 2009:15 (Isla Cubagua, Venezuela). McEachran 2009:1285 (Gulf of Mexico). Miller et al. 2009:227 (range, habitat, fig. 6.236, map 6.202). Barros et al. 2011:371 (estuary, Pará, Brazil). Benz et al. 2011:51 (copepod record from Bere 1936). Kells and Carpenter 2011:152–153 (description, color painting). Neves, Teixeira, and Araújo 2011:121 (middle reaches, Mambucaba River, Rio de Janeiro state, Brazil). Peralta-Meixueiro and Vega-Cendejas 2011:677 (hyperhaline coastal system: Ría Lagartos, Mexico). Reis-Filho et al. 2010:305 (Paraguacu River, Todos os Santos Bay, Bahia, Brazil). Vilar et al. 2011:45 (Baia da Babitonga, southern Brazil). Passos et al. 2012:230 (Paranaguá Estuarine complex, Brazil). Vega-Cendejas et al. 2012:141 (length-weight, Yucatán peninsula). Neves, Teixeira, Franco et al. 2011:

38

Family Belonidae 666–670 (lagoon, middle and lower channels of the estuarine mixing zone of a tropical estuary of the Mambucaba River, Rio de Janeiro state, Brazil). Gallardo-Torres et al. 2012:277 (present for five to nine months in the coastal lagoon Boca de la Carbonera Yucatán, Mexico). Vega-Cendejas and Santillana 2012:298 (abundance in lagoon and coastal habitats in Yucatán, Mexico). Barbanti et al. 2013:279 (fig. 1p), 282 (Canal de Bertioga, São Paulo, Brazil). Santana et al. 2013:185 (Itamaracá Island, Pernambuco, Brazil). Stevens et al. 2013:1650 (Peace River, tributary of Charlotte Harbor, FL). De Angelo et al. 2014:1043 (seagrass beds in eastern Gulf of Mexico). Gallardo Torres et al. 2014:89 (Yucatán, Mexico, photographs). Smith-Vaniz and Jelks 2014:32 (St. Croix, U.S. Virgin Islands records). Snyder and Burgess 2016:102 (listed, Florida). Strongylura euryops. Jordan et al. 1930:196 (range, synonymy). Duarte-Bello 1959:43 (Cuba and Jamaica). Strongylura diplotaenia. Jordan et al. 1930:195 (range, synonymy). Howell Rivero 1936:1 (description, Port Antonio, Jamaica, MCZ 33856). Strongylura almeida. Fowler 1941d:141 (previous records, Brazil). Strongylura timocu [sic]. Santos 1952:92 (description, range, Pernanbuco and Rio de Janeiro, Brazil). Strongylura timueu [sic]. Dragovich 1969:15 (eaten by Euthynnus alletteratus). Strongylura ticumu [sic]. Austin 1971:32 (mangroves, western Puerto Rico). Strongylura timicu [sic]. Swift et al. 1977:35 (enters springfed Wakulla River, FL).

Strongylura scrutator (not of Girard 1858a). Hubbs 1936:207– 209 (Yucatán Peninsula, UMMZ 102191, 102208). Gunter 1942:317, 1956:350 (after Hubbs 1936). Belone houttuyni. Mees 1962:32–38 (description, range, synonymy, in part). Mees 1966:318 (Tylosurus euryops an additional synonym). Belone marina (not of Walbaum 1792). Erdman 1967:45 (description, often found in rivers, Puerto Rico). Erdman 1972:45 (repeat of 1967 edition), 92 (B. marina = Strongylura timucu).

Types. Esox timucu Walbaum, 1792. Neotype CASSU 22073, female (219 mm BL); Brazil, Natal, Lake Papary; E. C. Starks; 1911; designated by Collette (1968). Dorsal fin rays 17; anal fin rays 19; pectoral fin rays 10–10. Predorsal scales 148. Two ovaries, left 69 mm, right 89 mm. Belone cigonella Valenciennes in Cuvier and Valenciennes, 1846. Puerto Rico. No type specimens found at MNHN (Collette et al. 1997) and none were present in 1887 (Jordan 1887a). Belone subtruncata Poey, 1860. No types extant; Cuba, Havana. Dorsal and anal fin ray counts from original description 17–17. Belone depressa Poey, 1860. Holotype USNM 4745 (190 mm BL); Cuba, Havana; Poey no. 332. Dorsal fin rays 17; anal fin rays 19; pectoral fin rays 11–11. Predorsal scales 140. Vertebrae 47 precaudal + 26 caudal = 73 total (Collette et al. 1992:3). Belone diplotaenia Cope, 1871. Holotype ANSP 7572 (175 mm BL); West Indies, St. Martins; R. E. van Rijgersma. Dorsal fin rays 16; anal fin rays 17; pectoral fin rays 11–11. Tylosurus sagitta Jordan and Gilbert, 1884. Lectotype USNM 34965 (194 mm BL); Florida, Key West; designated by Collette (1968:191). Dorsal fin rays 16; anal fin rays 19; pectoral fin rays 12–12. Vertebrae 48 precaudal + 25 caudal = 73 total. Paralectotypes CASSU 1891, CAS 27614. Tylosurus euryops Bean and Dresel, 1884. Holotype USNM 32073 (181 mm BL); Jamaica (Collette et al. 1992:5). Dorsal fin rays 16; anal fin rays 18; pectoral fin rays 11–11. Predorsal scales ca. 130. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Timucu (English) and agujón timucú (Spanish, Mexico). It is also known as timucú in Cuba (Duarte-Bello 1959; Rodríguez et al. 1984; Claro and Parenti 2001). Needlefishes are called carajota in Colombia (Dahl 1971). They are called marao in Venezuela (Román 1979), but this name is also used for several other needlefishes. The general name for needlefishes, agulha, has been used for both species of Strongylura in Brazil (Figueiredo and Menezes 1978; Martins-Juras et al. 1987), but Corrêa et al. (1986) used timucu for S. timucu in southern Brazil.

The following references are based at least in part, sometimes in entirety, on specimens of Strongylura timucu confounded with S. marina. Tylosurus marinus (not of Walbaum 1792). Jordan and Swain 1884:231 (Cedar Keys, FL). Henshall 1891:375 (Florida). Meek 1904:160–161 (San Francisco, Vera Cruz City, Mexico, FMNH 4568). Linton 1910:19 (food), 68–69 (helminth fauna, Tortugas). Gudger 1929:157 (favorite food of jacks, Tortugas). Menzel 1956:51 (Apalachee Bay region, FL). Sogandares-Bernal 1959:93, 95 (digenetic trematode, after Manter 1947). Strongylura marina (not of Walbaum 1792). Borodin 1934:109 (Florida Keys and Miami, FL). Gunter et al. 1948:304 (mortality, Ft. Myers Beach, FL). Chandler 1954:358 (nematodes: Tortugas, based on Linton papers). Joseph and Yerger 1956:126 (ecology, Alligator Harbor, FL). Caldwell et al. 1959:17 (Tortugero, Costa Rica, UF 7175, 7180, 7186). Duarte-Bello 1959:43 (range, in part). Berry and Rivas 1962 (description, range, synonymy, in part). Caldwell 1963:5 (Puerto Limon, Costa Rica). Caldwell and Caldwell 1964:12 (Tobago). Cervigón 1965:25 (listed, Venezuela). Caldwell 1966a:31 (Jamaica). Cervigón 1966:210–211 (description, Venezuela). Miller 1966:796 (range, in part, West Indies). Moe et al. 1966:16 (Florida collections). Reséndez Medina 1970:102–103 (description, 150 predorsal scales, Laguna de Tamiahua, Veracruz, Mexico). Moe 1972:10 (movements, southern Florida). Powell et al. 1972:48–49 (29 Florida collections). Carr and Adams 1973:512, 516–517 (diet of juveniles, Crystal River, FL). Duarte-Bello and Buesa 1973:78– 79 (Cuba).

39

Fishes of the Western North Atlantic

Figure 2.11. Timucu, Strongylura timucu. USNM 184270, 138 mm BL. Collected by Victor Springer, in Pass-A-Grill, Florida, U.S.A., 10 October 1958. Illustration by M. H. Carrington.

Diagnosis. Strongylura timucu differs from S. marina in two major characters: both left and right gonads are developed instead of only the right one, and there are fewer predorsal scales, 120–185 (x̅  156, N 254) rather than 205–304 (x̅  256, N 145). Description. Body elongate, rounded in cross section. Upper and lower jaws greatly elongate and studded with sharp teeth; maxilla exposed. Gill rakers absent. Anterior parts of dorsal and anal fins not forming prominent lobes; pectoral fin not falcate. Caudal peduncle without lateral keels, deeper than wide; caudal fin emarginate, not deeply forked. Both right and left gonads present, right longer than left. Dorsal fin rays 14–17, usually 16; anal fin rays 16–20, usually 18; pectoral fin rays 11, rarely 10 or 12 (Table 2.13). Vertebrae 44–49 precaudal + 24–27 caudal = 69–75 total (Table 2.14). Predorsal scales relatively few and large, 120–185 (Table 2.15). Color. Strongylura timucu is a heavily pigmented needlefish (Fig. 2.11). Melanophores usually extend to the ventral margin of the orbit, whereas they do not extend past the middle of the orbit in S. marina. Melanophores are prominent along the lateral line in S. timucu, absent in S. marina. Bluish green above, silvery below. A broad lateral dusky band on sides. Black pigment behind eyes usually extending to ventral margin of orbit, pre-orbital bone densely pigmented. Size. The maximum observed size for Strongylura timucu is 285 mm BL (MCZ 8793, female, Haiti), 418 mm SL (USNM 114314, female, Guatemala). Lengthweight information (using total length) was provided for 15 specimens from the Curucá Estuary, northern Brazil, by Giarizzo et al. (2006).

identified only to genus, probably refer to S. timucu: Noddy and Sooty Terns, Bird Key Rookery, Tortugas (Longley 1929); Blue Marlin, Makaira nigricans, off Puerto Rico (Erdman 1962); Skipjack Tuna, Katsu­ wonus pelamis, and Blackfin Tuna, Thunnus atlanticus, off Cuba (Guevara Carrió 1984a, 1984b). Parasites. Fourteen species of parasites have been reported from Strongylura timucu: five species of copepods, four digenetic trematodes, two monogenes, a cestode, a nematode, and a myxosporidian. copepods: Five species were reported by Cressey and Collette (1970:402): Ergasilidae, Ergasilus spatulus Cressey, from Tortugero, Costa Rica; Bomolochidae, Bomolochus bellones Burmeister (genus following Ho and Lin 2009), from Florida, the West Indies, and Central America; Anthosomatidae, Lernanthropus belones Krøyer, from Florida, the Bahamas, the West Indies, Central America, and Brazil, and L. tylosuri Richardi, six males and seven females, from one specimen from Rio de Janeiro, Brazil; and Philichthyidae, Coloboma­ tus goodingi Cressey and Collette, from Florida, the West Indies, and Curaçao. The records of Bomolochus nitidus and Lernanthropus chlamydotes from Lemon Bay, FL (Bere 1936) are referrable to B. bellones and L. belones, respectively (Cressey and Collette 1970:407). Bunkley-Williams and Williams (1994:89) reported Er­ gasilus lizae Krøyer from Puerto Rican specimens and Table 2.13. Fin ray counts for Strongylura timucu. Dashes indicate a value of zero. Fin rays Dorsal

Biology Predators. Probably eaten by a variety of surface-feeding fishes and birds, but there are few published records. Reported to be a favorite food of jacks in the Tortugas (Gudger 1929). Found in stomachs of the Little Tunny, Euthynnus alletteratus, by Dragovich (1969). Several reports of Strongylura,

Locality

14

15

16

17

10

11

12

Bahamas Florida, U.S.A. West Indies Central America Brazil

— 1 4 1 —

5 21 68 30 2

10 54 48 60 15

2 10 4 18 1

— 1 8 1 4

19 58 71 44 14

— 5 2 — —

6 126 187

35

14 206

7

Species total

40

Pectoral

Family Belonidae Table 2.14. Vertebral counts of Strongylura timucu. Dashes indicate a value of zero. Vertebrae Precaudal

Total

Caudal

Locality

44

45

46

47

48

49

24

25

26

27

69

70

71

72

73

74

75

Bahamas Florida, U.S.A. West Indies Central America Brazil

— — 2 2 —

1 1 17 7 1

2 5 28 31 2

4 19 2 24 2

1 5 1 6 —

— — — 2 —

— — 7 5 —

1 4 28 26 2

6 19 14 38 2

1 7 — 5 2

— — 4 2 —

— — 17 7 —

1 2 18 11 1

2 14 1 16 2

4 14 1 16 1

1 8 1 6 1

— 1 — — —

Species total

4

27

68

51

13

2

12

61

79

15

6

24

33

35

36

17

1

Table 2.15. Number of predorsal scales of Strongylura timucu. Dashes indicate a value of zero. Predorsal scales Locality Bahamas Florida, U.S.A. West Indies Central America Brazil Species total

121–130

131–140

141–150

151–160

161–170

171–180

181–190

— — 1 1 —

— — 15 8 1

— 9 37 24 6

10 39 40 39 8

5 49 13 12 7

3 13 2 4 2

— 2 — — —

2

24

76

136

86

24

2

postulated that Cressey’s record of E. spatulus might be either E. arthrosis Roberts or E. lizae. Bere’s 1936 record of Caligus rufimaculatus Wilson 1905 from Lemon Bay, FL is supported by Benz et al. (2011), noting that this copepod has been reported from 22 species of fishes as well as from a bottlenose dolphin. digenetic trematodes: Four species have been reported: Bucephalidae, Rhipidocotyle transversale Chandler, from Tampa Bay, FL (Sogandares-Bernal and Hutton 1959; Hutton and Sogandares-Bernal 1960); Haplosplanchnidae, Schikhobalatrema acutum (Linton 1910), originally described from Tylosurus marinus, from the Tortugas (= Strongylura timucu), and also reported from Biscayne Bay, FL (Overstreet 1969), Puerto Rico (Siddiqi and Cable 1960), and Curaçao (Nahhas and Cable 1964); Zoogonidae, Steganoderma elongatum Manter 1947, from Strongy­ lura timucu, Tylosurus crocodilus, and T. acus, from the Tortugas; and Zoogonidae, Steganoderma atherinae (Price), from Curaçao (Nahhas and Cable 1964). monogenes: Ancyrocephalidae, Ancyrocephalus par­ vus Linton, from Biscayne Bay, FL (Skinner 1982), and Murraytrematoides sp., from Puerto Rico (Bunkley-Williams and Williams 1994). cestodes: Ptychobothridae, Ptychobothrium belones (Dujardin), from Jamaica (Cable and Michaels 1967). nematodes: Philometridae, Philomitra sp., from the Tortugas (Chandler 1954).

myxosporidians: Myxidium sphaericum Théleham, in the bile and bladder of needlefish, probably Strongy­ lura timucu, from Buttonwood Canal, FL (Iversen et al. 1971). Food. Like other needlefishes, Strongylura timucu feeds mainly on small fishes, but will eat crustaceans and insects as well. In the Virgin Islands, small clupeoids such as Anchoa parva and Jenkinsia made up 96% of the diet by volume, shrimps the other 4% (Randall 1967). Small fishes (perhaps Gambusia or Cy­ prinodon) were found in S. timucu from Lake Forsyth, Andros Island, Bahamas (Breder 1934), in stomachs of specimens from Cedar Key, FL (Reid 1954), and in Tampa Bay along with copepods and insects (Springer and Woodburn 1960). Reproduction. Both left and right gonads are developed, whereas only the right one is developed in Strongylura marina (Collette 1968). Right gonad of S. timucu longer than left; ratio of left to right in males 1.0–1.6 (x̅ 1.21 for 52 specimens); in females 1.0–1.7 (x̅ 1.25 for 70 specimens). Ripe females were reported from the Caribbean in June (Erdman 1976). A 250 mm BL female had 405 eggs (1.6–2.1 mm in diameter) in the left ovary and 970 eggs (1.4–1.9 mm in diameter) in the right ovary, for a total of 1,375 eggs (x̅ 1.76 mm). The halfbeak stage extends beyond 35.5 mm BL. Ecology. This is a largely marine species that moves 41

Fishes of the Western North Atlantic into brackish and even fresh waters in southern Florida (Wakulla River [Swift et al. 1977]; southeast Dade County and near the northern Ten Thousand Islands area [Kushlan and Lodge 1974]) and Mexico, near Veracruz City (Miller 1976b). Range. This is the only species of Strongylura in the West Indies, where it is very abundant (Fig. 2.8). It extends north along the Atlantic coast of Florida to Ponce de Leon Inlet (Paperno et al. 2001) and St. Lucie Inlet (UF 4298), and along the Gulf Coast to Pensacola (USNM 125700). It also occurs along the coasts of Central and South America from Quintana Roo, Mexico (Schmitter-Soto and Gamboa-Pérez 1996), and Yucatán to Río de Janeiro, Brazil (SU, AMNH, MCZ). A record from Long Island, NY (Hickey and Lester 1983), is certainly a misidentification, presumably of S. marina, and a record from St. Paul’s Rocks, Brazil (Feiroza et al. 2003), is also probably a misidentification, possibly of Platybelone argalus ar­ galus. Study Material. A total of 689 specimens (20–310 mm BL), including the types of Esox timucu, Belone depressa, B. diplotaenia, Tylosurus sagitta, and T. eu­ ryops, originally at AMNH, ANSP, CAS, CU, FMNH, FSU, GCRL, LACM, MCZ, MZUSP, TU, UF, UMML, UMMZ, UPR, and USNM. Specimens came from nine areas: 336 specimens from Florida, U.S.A. (20–310 mm BL); 35 specimens from the Bahamas (28.0–248 mm BL); 13 specimens from Cuba (52.0– 200 mm BL); 27 specimens from Jamaica (39.3–359 mm BL); 20 specimens from Hispaniola (170–285 mm BL); 52 specimens from Puerto Rico, U.S.A. (43.6–259 mm BL); 15 specimens from the Lesser Antilles (51.8–180 mm BL); 131 specimens from Central America (43.0–275 mm BL); and 60 specimens from South America (49.5–300 mm BL).

veloped in the Australian T. gavialoides. Tylosurus differs from Ablennes and Strongylura in having more rays in the dorsal fin than in the anal fin, rather than the reverse. Analysis of molecular data suggests species of Tylosurus cluster with Ablennes (Lovejoy et al. 2004:369). Species. Tylosurus contains two polytypic species that are found nearly worldwide in tropical and subtropical marine waters: T. acus and T. crocodilus. In addition, there are four Indo-Pacific species: T. choram, T. punctulatus, T. gavialoides, and T. pacificus. Tylosurus acus acus (Lacepède, 1803) Atlantic Agujon Figures 2.12, 2.13 Tables 2.1, 2.16, 2.17, 2.18 Sphyraena acus Lacepède, 1803:325, 327 (original description based on a drawing by Plumier and therefore from the West Indies, probably Martinique), pl. 1, fig. 3. ICZN 2007 (name reinstated as a valid name for Tylosurus acus). Belona carribea Lesueur, 1821:127–128 (original description; Basseterre near Guadaloupe). Belone caribaea. Valenciennes in Cuvier and Valenciennes 1846:430–432 (description; West Indies; MNHN 4509, A.7619; see Collette et al. 1997:26). Jordan 1887a:529 (description, Martinique). Belone latimana Poey, 1860:292–293 (original description; Havana, Cuba). Jordan and Gilbert 1881 (gill rakers absent). Howell Rivero 1938:179 (holotype MCZ 622). Belone altipinne Poey, 1860:293 (original description; Havana, Cuba). Belone caribbea. Günther 1866:241 (description; Dominica, Jamaica). Cope 1871:481 (listed). Cockerell 1892:14 (listed; Jamaica). Metzelaar 1919:31–32 (description; Curaçao), fig. 11. Belone Jonesii Goode, 1877:295–296 (original description; Bermuda). Belone Jonesii Günther, 1879:151 (original description; Bermuda). Belone latimanus. Goode and Bean 1879:46 (Buzzards Bay, MA). Bean 1880:103 (Woods Hole, MA). Belone jonesi. Günther 1880:10–11 (synonymy; redescription). Tylosurus latimanus. Jordan and Gilbert 1883a:373 (description; Buzzards Bay, MA, and Cuba). Tylosurus caribbaeus. Bean 1884:342 (Woods Hole, MA; USNM 28748, 31438). Jordan 1886a:26 (Beaufort, NC). Jordan and Fordice 1887:357 (description; Martinique). Linton 1890:722–723 (cestode Dibothrium restiforme; Buzzard’s Bay, MA). Linton 1891:531–532 (Echinorhynchus pristus var. tenuicornis in intestine; Woods Hole, MA). Jordan and Evermann 1896a:321 (West Indies). Jordan and Evermann 1896b:717 (description; West Indies; synonomy). Linton 1897:802 (cestode Rhynchobthrium speciosum). Linton 1898:534–535 (trematode Distomum nitens; Woods Hole, MA). Ariola 1900:422–423 (cestode Bothriocephalus resti­ formis; Buzzards Bay, MA). Bean 1905:300 (Nassau, Bahamas; USNM 53090). Linton 1905:358 (stomach contents, copepod on gills; Beaufort, NC). Smith 1907:159 (Cape Lookout, NC). Jordan et al. 1930:197 (range, synonymy). Butsch 1939:19 (common; Barbados). Linton 1941:432 (cestode Bothriocephalus restiformis; Woods Hole, MA).

Genus Tylosurus Cocco, 1833 Tylosurus Cocco, 1833:20 (original description, type species Tylosurus cantrainei Cocco by monotypy = Tylosurus acus im­ perialis Rafinesque, 1810).

Diagnosis. Tylosurus shares several specializations with Ablennes, a deeply forked caudal fin with the lower lobe longer than the upper lobe, many vertebrae and dorsal and anal rays, and a melanistic posterior lobe in the dorsal fin. This lobe is lost in adult Tylosurus, but retained in adult Ablennes. Loss of this lobe is achieved in different ways in Tylosurus species; it is sloughed off in T. crocodilus, but resorbed in T. acus (Breder and Rasquin 1952, 1954). Five of the six species of Tylosurus have a narrow black raised keel on the caudal peduncle, but this is not well de42

Family Belonidae Tylosurus caribaeus. Jordan 1887b:568 (listed; West Indies). Rosen 1911:51 (Nassau; after Bean 1905). Tylosurus acus. Jordan and Fordice 1887:355–357 (synonymy, range, description). Jordan 1887b:568 (listed; West Indies). Jordan and Evermann 1896a:320 (West Indies). Jordan and Evermann 1896b:716–717 (description, range, synonymy). Smith 1898:93 (Woods Hole, MA). Linton 1901b:442–443 (parasites; Buzzards Bay, MA). Barbour 1905:113–114 (abundant, parasites; Bermuda). Bean 1905:300 (Spanish Wells and Clarence Harbor, Bahamas; USNM 53091, 53092), color pl. LIII. Bean 1906:35 (Bermuda). Smith 1907:158–159 (Beaufort, NC), fig. 62. Kendall 1908:57 (Woods Hole, Buzzards Bay, Nantucket, MA). Linton 1908:86, 96, 100, 107, 115 (parasites; Bermuda). Wilson 1908:626 (parasitic copepods). Rosen 1911:51 (after Bean 1905). Gudger 1912:168 (only one ovary well developed; Beaufort, NC). Sumner et al. 1913:744–745 (parasites, previous records; Woods Hole, MA area). Cockerell 1916:53 (scales; Woods Hole, MA). Starks 1916:9, 23 (fig. 11, sesamoid articular). Fowler 1917b:121 (Nantucket, MA). Van Cleave 1918:20 (acanthocephalans; Woods Hole, MA). Fowler 1919a:13 (previous ANSP records of T. raphidoma reidentified as T. acus; New Jersey). Nichols 1921:22 (listed; Turks Islands, Bahamas). Schroeder 1924:5 (sometimes used for food; Key West, FL). Nichols and Breder 1927:58 (description; southern New England). Hildebrand and Schroeder 1928:149–150 (description; Lynnhaven Roads and Buckroe Beach at mouth of Chesapeake Bay), fig. 80. Jordan et al. 1930:197 (range, synonymy). Breder 1932a:14 (mature at 650 mm SL; Tortugas), 33–35 (tables 1–3, size, growth of jaws, eye angle), pl. 2, fig. 3 (development of upper and lower jaws), pl. 5 (development of dorsal and caudal fins). Beebe and Tee-Van 1933:60–61 (description, common; Bermuda). Manter 1940:534 (western Atlantic host of trematode Haplosplanchnus acutus). Mirsky and Ris 1951:457 (DNA content of erythrocytes). Breder and Rasquin 1954 (post-larval transformation). Saunders 1959:379 (Haemogrega­ rina absent; Bermuda). Sogandares-Bernal 1959:93, 95 (trematode Haplosplanchus acutus; after Linton 1908). Berry and Rivas 1962 (synonymy, description, range in western Atlantic). Cervigón 1965:26 (marao ojon; Venezuela). Collette and Berry 1965:391 (validity of name). Roithmayr 1965:20 (listed among industrial bottom fishes; northern Gulf of Mexico). Cervigón 1966:214–216 (description, size, food value; Venezuela), fig. 84 (photograph of 800 mm TL specimens). Collette and Berry 1966:326–327 (validity of name). Moe et al. 1966:16 (Okaloosa Co., FL). Collette 1967:197–198 (validity of name). Randall 1967:684 (stomach contents; Virgin Islands). Böhlke and Chaplin 1968:120 (Bahamas, description, figure). Starck 1968:17 (agujon; listed; Alligator Reef, FL). Dragovich 1969:16 (eaten by Euthynnus allettratus). de Sylva 1970:51 (Biscayne Bay area, FL). Love 1970:403 (reference to Mirsky and Ris 1951). Mago Leccia 1970:89 (listed; Venezuela). Rees 1970:208 (trematode Haplosplanchus acutus; Bermuda). Dooley 1972:9 (Sargassum community). Powell et al. 1972:119 (Ft. Walton Beach, FL). Duarte-Bello and Buesa 1973:79 (synonymy, range). Palacio 1974:35 (Caribbean Colombia). Belyanina 1975:140 (ichthyoplankton; Gulf of Mexico). Fahay 1975:4, 19 (larvae; South Atlantic Bight). Guitart 1975:245–246 (Cuba, synonymy, description, fig. 180). Instituto Nacional de Pesca 1976:6 (range, common names). Shiino 1976:92 (common names). Matthews et al. 1977:14 (eaten by Thunnus albacares). Hardy 1978:96– 101 (description, range, development, figures). Figueiredo and Menezes 1978:65 (description; Brazil), fig. 11. Hastings 1979:7 (jetty; St. Andrew Bay, FL). Román 1979:82 (description, common name, figure; Venezuela). Fedoryako 1980:581 (Sargasso Sea). Gilmore et al. 1981:10 (Indian River Lagoon

area, Florida). Reséndez Medina 1981a:269 (description; Laguna de Términos, Campeche, Mexico), pl. IV, fig. 30. Sacchi et al. 1981:2 (abundant November to December, French Antilles). Tortonese 1982:197 (circumtropical polytypic species). Fahay 1983:212 (counts), 213 (figure of T. a. melanotus from Mito in Uchida et al. 1958). Murdy 1983:88 (outline figure), 90 (in key; Texas). Bravo-Hollis 1984:65–66 (monogene Nuda­ ciraxine gracilis; Veracruz). Collette et al. 1984:336 (egg size). Rodríguez et al. 1984:15 (agujón; Cuba). Acero P. and Garzón F. 1987:90 (Colombia). Rodríguez and Valdés 1987:82 (Cuba). Schwartz 1989:340 (North Carolina). Boughton et al. 1991 (ontogeny of upper jaw). Cervigón 1991:227–228 (description, Venezuela). Able 1992:4 (New Jersey, rare). Boschung 1992:83, 84 (Dauphin Island, AL). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Murdy et al. 1997:117, 286 (Chesapeake Bay, occasional visitor). Smith 1997:393 (description, range). Ho 1998:251 (parasitic copepod, Brazil). McEachran and Fechhelm 1998:939, figure (Gulf of Mexico). Mejía Aristizábal et al. 1998:31 (Roncador Island, Caribbean Colombia). Carvalho Filho 1999:80–81 (description, Brazil). Lovejoy 2000:1352 (fig. 2, molecular phylogeny). Aguilar et al. 2001:131 (Bahía de Cienfuegos, Cuba). Briggs and Waldman 2002:58 (rare in lower New York Bay). Carrasquillo-Casado et al. 2002:65 (beak penetration led to death of bottlenose dolphin in Puerto Rico). McBride and Styer 2002:23 (common year-round in south Florida lampara net fishery). Menezes and Figueiredo 2003:67 (Brazil). Tavares et al. 2004 (metazoan parasites, Rio de Janeiro, Brazil). Collette and Parin 2005 (request for reinstatement of T. acus as valid name for the species). ICZN 2007 (T. acus reinstated as valid name). Casazza and Ross 2008:351 (associated with Sargassum, Gulf Stream off North Carolina). Collette 2010:32 (reproduction and development). Kells and Carpenter 2011:152–153 (description, color painting). Snyder and Burgess 2016:102 (listed, Florida). Belone (Tylosurus) acus. Metzelaar 1919:31 (description; Dutch West Indies). Strongylura acus. Fowler 1919a:3 (Massachusetts, New Jersey, and Pennsylvania). Fowler 1920:155 (Atlantic and Cape May counties, New Jersey). Breder 1929:279–280; (Tortugas). Longley and Hildebrand 1941:29 (Tortugas; after Breder 1932a). Manter 1947:324–325 (trematode host; Tortugas). Pearse 1947:9 (parasitic copepod Caligus schistonyx; Beaufort, NC). Breder 1948:89 (description, range). Pearse 1949:34 (trematode Lecithaster gib­ bosus; Beaufort, NC). Fowler 1952a:112 (Atlantic and Cape May counties, New Jersey). Springer and Bullis 1956:57 (Arcas Cay, Campeche Bank, Mexico). Briggs 1958:264 (agujon, range). Yamaguti 1958:56 (digene Steganoderma nitens). Duarte-Bello 1959:42 (range; Cuba). Schwartz 1962 (eats small fishes and shrimps; MD). Breder and Rosen 1966:302–303 (spawning; after Breder and Rasquin 1952, 1954). Reséndez Medina 1971:24 (Campeche; after Springer and Bullis 1956). Golovanj 1973:561 (pelagic ichthyofauna; Cuba). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Strongylura caribbea. Fowler 1944:81–82 (synonymy, description; Roncador Bank, Caribbean Sea), figs. 96–97. Manter 1947:375 (trematode Steganoderma elongatum; Tortugas). Fowler 1953:52 (Roncador Bank). Duarte-Bello 1959:42 (Cuba; West Indies). de Sylva and Higman 1979:153 (occasionally ciguateric). Strongylura caribaea. Yamaguti 1958:56 (digene Steganoderma nitens). Belone imperialis. Mees 1962:38–41 (synonymy, description, range). Mees 1966:149–150 (validity of name). Belone imperialis imperialis. Mees 1964:319–320 (Atlantic subspecies). Tylosurus acus acus. Parin 1967:53 (three subspecies recognized). Collette and Parin 1970:41–46 (synonymy, types; T. a. acus

43

Fishes of the Western North Atlantic anal fin rays 23; pectoral fin rays 13–13. Vertebrae 63 precaudal + 31 caudal = 94 total. Paralectotypes USNM 202591 (2, 358–376 mm BL). Dorsal fin rays 23, 24; anal fin rays 21, 22. Vertebrae total 90, 91. Belone jonesii Günther, 1879. Holotype BMNH 1879.1.8.2 (551 mm BL, 800 mm SL); Bermuda; Jones. Dorsal fin rays 26; anal fin rays 22; pectoral fin rays 14–14. Common Names. Early references used the name Houndfish for both Tylosurus acus and T. crocodilus in the United States (Smith 1907; Nichols and Breder 1927). Since Breder (1948), the name agujon has been generally used and Atlantic Agujon is recommended by the Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013). It is also used, with an accent (agujón) in Cuba (Duarte-Bello 1959; Rodríguez et al. 1984; Claro and Parenti 2001). Page et al. (2013) lists agujón del Atlántico for the Spanish name in Mexico. The name marao is used for most needlefishes in Venezuela, marao ojon for T. acus (Cervigón 1965, 1966; Román 1979). Agulha is used for needlefishes in Brazil, or sometimes timbale (Figueiredo and Menezes 1978). Diagnosis. This is the species of Tylosurus with the highest number of vertebrae, and dorsal and anal fin rays. Western Atlantic specimens of T. acus have 90– 95 total vertebrae, 22–26 dorsal fin rays, and 20–24 anal fin rays, compared with Atlantic specimens of T. crocodilus, which have 79–84 total vertebrae, 21–24 dorsal fin rays (mode 22), and 17–22 anal fin rays (mode 21). Tylosurus acus has a longer, more slender beak, head, and body than T. crocodilus. Large specimens of T. acus are long, graceful fish, whereas large specimens of T. crocodilus are much stubbier, heavier-bodied fish. Tylosurus crocodilus has larger body proportions at a given body length because it is a shorter fish (i.e., has fewer vertebrae). Tylosurus acus has the left gonad greatly reduced or absent in both sexes. The ratio of left gonad length to right ranges from 2.3 to more than 15.5 compared to 1.1 to 1.7 in T. crocodilus. The western Atlantic subspecies, Tylosurus acus acus, is most similar meristically to T. a. imperialis of the Mediterranean and T. a. melanotus of the Indo-West Pacific. These three subspecies have more vertebrae (89–96) than T. a. rafale from the Gulf of Guinea (82–88) or T. pacificus from the eastern Pacific (74–81) (Collette and Banford 2001). The Mediterranean T. a. imperialis has more vertebrae (93–96, x̅  94.2) than T. a. acus (90–95, x̅  91.9) and more predorsal scales (370–430, x̅  397, rather than 325–389, x̅  347). The three Atlantic subspecies also differ morphometrically (Collette and Parin 1970: table 15) with T. a. acus differing significantly (at the 99.9% level) in 7 out

recognized as western Atlantic subspecies; comparisons with other subspecies). Cressey and Collette 1970:403 (parasitic copepods), 413 (fig. 177, widespread distribution). Collette 1978 (description, range, figure). Collette et al. 1984:344 (fig. 179G, 130 mm specimen). Cervigón et al. 1993:270 (northern coast of South America). Collette et al. 1997:23 (no types of Sphyraena acus Lacepède at NMNH). Smith-Vaniz et al. 1999:166–167 (Bermuda). Claro and Parenti 2001:43 (Cuba). Collette and Banford 2001 (comparison with other subspecies of T. acus and with T. pacificus). Collette 2003a:1112 (figure, description, range in west-central Atlantic). Collette 2003b:9 (synonymy). Banford et al. 2004 (molecular phylogeny). Lovejoy et al. 2004:369, 377 (molecular phylogeny). Collette 2005:928–929 (early life history, figs. A–F). Kohn et al. 2006:70 (monogene from Mexico). Schwartz 2006:64 (North Carolina records). Fahay 2007:776, 784–785 (early stages, figs. A–H). Luque and Tavares 2007:9, 11, 12, 14, 16 (parasitic copepods, Brazil). Nóbrega and Lessa 2007:68 (artisanal fishery, northeastern Brazil). Cohen and Kohn 2008:8, 13, 25 (monogene parasites from Venezuela). Méndez et al. 2008:371 (Gulf of Cariaco, Venezuela). McEachran 2009:1285 (Gulf of Mexico). Smith-Vaniz and Jelks 2014:32 (St. Croix, U.S. Virgin Islands records). Starck et al. 2017:91 (Florida Keys). Tylosaurus [sic] acus. Haedrich et al. 1973:594 (falciform process). Burnett-Herkes 1986:590 (description; Bermuda), 591 (fig. 200). Tylosurus imperialis. Schmitter-Soto et al. 2000:154 (Caribbean Mexico). González-Gándara and Arias-González 2001:249 (Alacranes Reef, Gulf of Mexico). Tylosaurus [sic] acus acus. Medina and Yunda 2008:61 (beaches, Santa Marta, Caribbean Colombia; lechero).

Misidentifications. Several earlier Fowler records of Tylosurus raphidoma from the Atlantic coast of the United States (e.g., Fowler 1914) were reidentified as T. acus (Fowler 1919a). Types of Atlantic Nominal Species. Sphyraena acus Lacepède, 1803. No type specimens extant. The original description is based on a poor drawing by Plumier (Collette et al. 1997). The high number of dorsal fin rays (25) and anal fin rays (24) indicates the description was of the species Tylosurus acus. Belona caribbea Lesueur, 1821. No type specimens were located. The original description compares the long-snouted B. caribbea with the short-snouted B. crocodilus. Belone latimana Poey, 1860. Holotype MCZ 622 (470 mm BL, 679 mm SL). Dorsal fin rays 25; anal fin rays 23; pectoral fin rays 14–14. Vertebrae total 93. Belone altipinna Poey, 1860. No types located and none in the Howell Rivero (1938) list of Poey types in MCZ. Original description gave dorsal fin rays 24, anal fin rays 23. This is a common count for Tylosurus acus in the western Atlantic, but falls out of the anal ray count for T. crocodilus in the Atlantic. Belone jonesii Goode, 1877. Lectotype USNM 21392 (391 mm BL, 568 mm SL); Bermuda; Jones; designated by Collette and Parin (1970:44). Dorsal fin rays 25; 44

Family Belonidae

Figure 2.12. Atlantic Agujon, Tylosurus acus acus. A. USNM 221687, 382 mm BL. Collected by B. B. Collette in Puerto Rico, U.S.A., 3 June 1965. B. USNM 198402, 130 mm BL. Collected in the North Atlantic from M/V Delaware, 1 May 1960. Illustrations by M. H. Carrington.

of 10 characters from T. a. imperialis and 2 out of 10 from T. a. rafale. Description. Body elongate, rounded in cross section. Upper and lower jaws greatly elongate and studded with sharp teeth; teeth straight at all sizes. Gill rakers absent. Anterior part of dorsal fin with a low lobe, contained 10.5–13.3 times in BL; anal fin lobe low, contained 9.7–11.7 times in BL; pectoral and pelvic fins relatively short, 8.0–12.4 and 10.0–14.1 times in BL, respectively; a small black lateral keel is present on the caudal peduncle, caudal peduncle deeper than wide; caudal fin deeply forked, lower lobe much longer than upper. Left gonad absent or greatly reduced in length in both sexes. Dorsal fin rays numerous, 22–26, usually 24; anal fin rays numerous, 20–24, usually 21 or 22; pectoral fin rays 13 or 14 (Table 2.16). Total vertebrae 90–95 (Table 2.17). Predorsal scales tiny and very numerous, 320–390 (x̅ 347.3). Color. Dark bluish above, silvery white below. A dark blue stripe along sides. Juveniles have an elevated black lobe in the posterior part of the dorsal fin that is lost with growth. Size. The maximum observed size for Tylosurus acus acus is 128.5 cm SL and 95 cm BL; commonly to 90 cm SL.

albacares (Matthews et al. 1977), and Bottlenose Dolphin, Tursiops truncatus (Carrasquillo-Casado et al. 2002), but other fishes, including tunas and billfishes, undoubtedly eat Tylosurus acus acus. Parasites. About 21 species of parasites have been reported from Tylosurus acus acus: six species of copepods, two acanthocephalans, two monogenes, three trematodes, three cestodes, a nematode, and a barnacle. copepods: Cressey and Collette (1970) reported eight species of copepods from Tylosurus acus worldwide, but only five of these were from T. a. acus: Bomolochidae, Bomolochus bellones Burmeister (genus following Ho and Lin 2009), 7 females and 3 males from the gill chambers and oral valves of 4 specimens from Florida and Honduras; Caligidae, Caligodes lacinatus (Krøyer), 40 females and 1 male from under the oral valves of 17 specimens from Virginia south to Belize and the Virgin Islands; Anthosomatidae, Lernanthropus tylosuri Richardi, 3 females and 2 males from gill filaments of 3 specimens from Belize, Puerto Rico, and the Virgin Islands; Lernaeoceridae, Lernaeolophus sul­ tanus (Milne-Edwards), 2 females from the roof of the mouth of a specimen from Haiti; and Philichthyidae, Colobomatus goodingi Cressey and Collette, 10 females from the cephalic lateral line canals of 8 specimens from North Carolina, the Bahamas, and Belize. Previous copepod records include Tuxophorus caligodes Wilson, 1908, chalimus larvae, Beaufort; Caligodes megacephalus Wilson, 1905 from the underside of the mouth of a silver gar “Lepistosteus osseus” = Caligodes laciniatus, probably from T. a. acus; the undescribed

Biology Predators. The only published records of predators in the western Atlantic are Little Tunny, Euthynnus al­ letteratus (Dragovich 1969), Yellowfin Tuna, Thunnus 45

Fishes of the Western North Atlantic Table 2.16. Fin ray counts for Tylosurus acus and T. pacificus, based on Collette and Banford (2001). Dashes indicate a value of zero. Fin rays Dorsal Species and locality Tylosurus acus acus Western Atlantic Tylosurus acus imperialis Mediterranean Sea

Anal

20

21

22

23

24

25

26

27

18

19

20

21

22

23

24

—

—

1

23

65

14

3

—

—

—

4

52

49

9

1

—

—

—

2

14

8

2

—

—

—

—

—

16

8

—

Tylosurus acus rafale Gulf of Guinea Tylosurus acus melanotus Indo-West Pacific Eastern Pacific Species total

1

7

31

27

3

—

—

—

—

1

14

42

12

—

—

— — 1

— — 7

— — 32

— — 52

7 — 89

46 — 68

39 — 44

3 — 3

— — —

— — 1

— — 18

— 1 95

16 2 95

49 2 68

20 1 22

Tylosurus pacificus Eastern Pacific

—

9

13

2

—

—

—

—

1

9

13

2

—

—

—

Table 2.17. Total vertebral counts of Tylosurus acus and T. pacificus. Dashes indicate a value of zero. Vertebrae Species and locality

74 75 76 77 78 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96

Tylosurus acus acus Western Atlantic

—

—

—

—

—

—

—

—

—

—

—

—

—

—

—

Tylosurus acus imperialis Mediterranean Sea

—

—

—

—

—

—

—

—

—

—

—

—

—

—

—

—

— —

Tylosurus acus rafale Gulf of Guinea

—

—

—

—

—

—

—

1

1

3 13 24

9

3

—

—

— —

Tylosurus acus melanotus Indo-West Pacific Eastern Pacific

— —

— —

— —

— —

— —

— —

— —

— —

— —

— —

— —

— —

— 1

1 1

Species total

—

—

—

—

—

—

—

1

1

3 13 24

9

3

2

3

3

3

6

2

3

—

—

—

—

Tylosurus pacificus Eastern Pacific

species of Lernanthropus and Bomolochus mentioned by Wilson (1908) = Lernanthropus tylosuri and B. bel­ lones; and Caligus schistonyx, 3 chalimi reported from Beaufort, NC, Cressey and Collette (1970) considered these unidentifiable. Tavares et al. (2004) also found Caligodes lacinatus and Lernanthropus tylosuri on specimens from Rio de Janeiro and added Caligus malabaricus Pillai, Caligus sp. D, and Metacaligus sp. to the list of copepods from T. a. acus. isopods: No cymothoid isopods of the genus Motho­ cya were found on Tylosurus acus (Bruce 1986), unlike the situation in many other needlefishes. acanthocephalans: Two species of Rhadinorhynchus

— —

—

— —

—

—

9 27 29 14

7

2

—

6 11

9

1

—

—

—

—

8 —

8 —

5 —

1 —

1 11 32 36 28 26 16

2

—

—

3 2

5 2

— —

—

—

—

—

were described from Woods Hole, MA, from Tylo­ surus acus: R. tenuicornis (Linton 1891) and R. ornatus Van Cleave, 1917. monogenes: One monogene, Axinidae, Nudaciraxine gracilis (Linton), was reported from the gill filaments of a Tylosurus acus acus from Veracruz, Mexico (Bravo-Hollis 1984; Kohn et al. 2006). Two monogenes were reported from the gills of specimens from Rio de Janeiro: Chlamydaxine sp. and Nudaciraxine sp. (Tavares et al. 2004; Cohen and Kohn 2008). digenetic trematodes: Five species have been recorded. Two were first reported as species of Distomum by Linton: D. nitens Linton 1898 from a Woods Hole, 46

Family Belonidae MA, Tylosurus acus acus and Distomum sp. Linton 1908 from Bermuda, later described as Deradena acutus from Strongylura marina at Tortugas. The first species was known as Steganoderma nitens (Overstreet 1969), Zoogonidae, and now as Lecithostaphylus nitens (Dyer et al. 1985). The second became Schikhobalotrema acuta (Siddiqi and Cable 1960), Haplosplanchnidae, then Haplosplanchus acutus (Manter 1940, 1947; Sogandares-Bernal 1959; Rees 1970) and was also reported from Rio de Janeiro, as Schikhobalotrema acuta by Tavares et al. (2004). The third species, Lecithaster gibbosus (Rudolphi), Hemiuridae, was reported from T. a. acus from Beaufort, NC (Pearse 1949). Tavares et al. (2004) reported two more species from Rio de Janeiro specimens: Parahemiurus merus and Rhipidocotyle sp. cestodes: Four species have been recorded: Ptycho­ bothrium restiforme (Linton 1890; Rees 1969) from Buzzards Bay, MA; Rhynchobothrium speciosum Linton 1897 from Woods Hole, MA; Otobothrium penetrans Linton 1908 from Bermuda; and Scolex pleuronectis (Müller, 1758) from Rio de Janeiro (Tavares et al. 2004). nematodes: The flesh of a Bermuda Tylosurus acus acus was filled with worms, particularly near the vertebral column; these were identified as Ichthyonema globiceps Rudolphi (Linton 1908). Larval Hysterothy­ lacium sp. and Pseudoterranova sp. were found in the mesentaries of specimens from Rio de Janeiro (Tavares et al. 2004). barnacles: Although not parasites, there are several reports of gooseneck barnacles, Conchoderma virga­ tum, attached to the interorbital region of fish from Woods Hole and Buzzards Bay, MA (Smith 1898; Linton 1901b; Sumner et al. 1913). Food. Feeds on small fishes, crustaceans, and insects (Linton 1905, 1908; Schwartz 1962; Randall 1967). In Bermuda, anchovies (Anchoa choerostoma) and silversides (Allanetta harringtonensis) are particularly important (Goode 1877; Barbour 1905). Randall (1967) examined 13 specimens from the Virgin Islands, six were empty, the others contained mostly fishes (96.3% by volume), carangids, and Tylosurus; hermit crabs (1.4% by volume), insects (1.4% by volume), and stomatopods (0.9% by volume). Reproduction. Catching young and juveniles by dip net with night lights indicates that Tylosurus acus probably spawns offshore (Berry and Rivas 1962). The left gonad is either greatly reduced or absent in both sexes, as first noted by Gudger (1912). The ratio of left gonad length to right ranges from 2.3 to more than 15.5 compared to 1.1 to 1.7 in T. crocodi­ lus (Collette and Parin 1970). Ovarian egg counts in specimens of T. a. melanotus (485 mm BL) and T. a. rafale (500 mm BL) were 116 (2.3–3.0 mm diameter)

and 196 (1.0–1.2 mm diameter) in the left ovary, and 1,676 (2.5–2.9 mm diameter) and 12,017 (0.9–1.3 mm diameter) in the right ovary, for totals of 1,792 (x̅  2.72) and 12,213 (x̅  1.05), respectively. Fertilized eggs are probably larger and are 3.2–4.0 mm in T. a. melanotus (Mito in Uchida et al. 1958). Eggs of T. a. melanotus have tufts of long filaments (Mito in Uchida et al. 1958). Development. Two features are of special interest in the development of Tylosurus acus: the melanistic posterior lobe of the dorsal fin and the halfbeak stage. In both cases, development proceeds differently in T. acus and the sympatric T. crocodilus. Loss of the elevated posterior lobe of the dorsal fin is accomplished by resorption in T. acus, in comparison to T. crocodilus in which the loss is accomplished by sloughing off the area (Breder and Rasquin 1954). The posterior lobe reaches its maximum observed size of 15–20 mm in specimens about 200 mm SL and then regresses (Breder and Rasquin 1954: fig. 5). There is a distinct halfbeak stage that continues to about 200 mm SL in the development of Tylosurus acus (Breder and Rasquin 1954: fig. 7), but is absent in T. crocodilus (Breder 1932a: pl. 2) and other species of Tylosurus. The upper jaw has not yet grown out to the tip of the lower jaw in a 130 mm BL specimen of T. a. acus, whereas both jaws are about equal in length in much smaller specimens (72–96 mm BL) of T. gavia­ loides, T. choram, and T. crocodilus (Collette et al. 1984: fig. 179 G–I). Ecology. Although both Tylosurus acus and T. croc­ odilus are widespread in tropical and subtropical waters of the world, there are differences in distribution that correlate with the ecological preferences of the species. Juvenile Tylosurus acus acus occur farther offshore than juvenile Tylosurus crocodilus crocodilus in the western Atlantic and are much less common inshore (Berry and Rivas 1962: fig. 3). Tylosurus a. acus is a member of the pelagic Sargassam complex (Dooley 1972; Fedoryako 1980). The three more oceanic species (Ablennes hians, Platybelone argalus, and T. a. acus) occur at Bermuda; the more inshore species of Strongylura do not. Throughout its range, T. acus extends farther north and south into colder waters than does T. crocodilus (compare Cressey and Collette 1970: figs. 177, 178). Thus, in the western Atlantic, there are many records of T. a. acus north as far as the southern coast of Massachusetts, whereas T. c. crocodilus generally does not occur north of North Carolina. Relationship to Humans. Schroeder (1924) wrote that both Tylosurus acus and T. crocodilus were sometimes used as food in Key West and other Florida 47

Fishes of the Western North Atlantic Tylosurus crocodilus crocodilus (Peron and Lesueur in Lesueur, 1821) Houndfish Figures 2.14, 2.15 Tables 2.1, 2.18, 2.19, 2.20 Belona crocodila Peron and Lesueur in Lesueur, 1821:129–130 (original description; Mauritius). ICZN 1970:213 (name to be placed on the Official Lists and Indexes of Names and Works in Zoology). Belone Raphidoma Ranzani, 1842:359–362 (original description; Brazil), pl. 37, figs. 1–5, pl. 44. Günther 1866:249 (after Ranzani 1842). Belone gerania Valenciennes in Cuvier and Valenciennes, 1846:437– 438 (original description; Martinique). Günther 1866:241 (after Valenciennes in Cuvier and Valenciennes, 1846). Cockerell 1892:14 (listed; Jamaica). Belone crassa Poey, 1860:291–292 (original description; Cuba). Poey 1881:337 (after Poey 1860). Howell Rivero 1938:179 (MCZ 623 type). Belone melanochira Poey, 1860:294–295 (original description; Cuba). Jordan and Gilbert 1881 (no gill rakers). Howell Rivero 1938:179 (MCZ 624 type). Paepke and Seegers 1986:146 (ZMB 5167 type). Tylosurus gladius Bean in Goode and Bean, 1882:430 (original description; Pensacola, FL). Bean and Dresel 1884:168 (Jamaica; USNM 32077). Cockerell 1892:14 (listed; Jamaica). Tylosurus crassus. Jordan 1884:112 (description; Key West, FL; USNM 35039). Smith 1896:175 (Biscayne Bay, FL). Tylosurus raphidoma. Jordan 1886b:35 (Havana, Cuba). Jordan and Fordice 1887:353–354 (synonymy, description; Florida, West Indies). Jordan 1887b:568 (listed; West Indies). Jordan 1890:647 (St. Lucia). Henshall 1895:212 (Florida Keys). Jordan and Evermann 1896a:320 (range). Jordan and Evermann 1896b:715–716 (description, synonymy). Jordan and Rutter 1897:97 (Jamaica). Evermann and Kendall 1899:61 (Florida). Evermann and Marsh 1902:99–100 (Puerto Rico, fig. 17). Jordan and Thompson 1905:234 (Tortugas). Linton 1905:357–358 (stomach contents, parasites; Beaufort, NC). Smith 1907:158 (Beaufort, NC). Nichols 1912:182 (common in market; Cuba). Miranda Ribeiro 1915:1–10 (description; Brazil). Meek and Hildebrand 1923:226–227 (synonymy, description; Panama). Schroeder 1924:5 (sometimes used as food; Florida Keys). Gudger 1929:157–158 (description of juveniles; Tortugas). Jordan et al. 1930:196 (range, synonymy). Parr 1930:19 (Bahamas). Breder 1932a:14–19 (description, behavior, growth, spawning season, parasites; Tortugas), pl. 2, fig. 2 (development of jaws), pl. 4 (development of dorsal and caudal fins), pl. 10 (eye; parasites). Shuler 1938:56 (cestode Ptychobothrium belones; Tortugas). Butsch 1939:19 (common; Barbados). Hildebrand 1939:26 (west side of Gatun Locks, Panama Canal). Manter 1940:534 (trematode Haplosplanchus acutus). Röhl 1942:375 (dangerous to fishermen; bait; Venezuela). Breder 1946:4–5 (juveniles resemble bits of plants; Tortugas). Norman and Fraser 1949:102 (Florida to Brazil). Breder and Rasquin 1952 (black posterior dorsal lobe sloughs off with growth). Breder and Rasquin 1954 (comparison of transformation with T. acus). Menzel 1956:51 (Apalachee Bay region, Florida). Sogandares-Bernal 1959:93, 95 (trematode Haplosplanchnus acutus; Bimini). Miranda Ribeiro 1961:5 (Brazil). Berry and Rivas 1962 (description, synonymy, ecology). Hildebrand et al. 1964:114 (Alacran Reef, Yucatán). Cervigón 1965:26 (listed; Venezuela). Cervigón 1966:213–214 (description; food value; Venezuela). Moe et al. 1966:17 (11 Florida collections). Caballe-

Figure 2.13. Atlantic distribution of Atlantic Agujon, Tylosurus acus, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

Keys. Both T. acus, and especially T. crocodilus, are considered edible and of good quality in Venezuela (Cervigón 1966). The species has been listed as occasionally ciguateric (de Sylva and Higman 1979). Range. Both Tylosurus acus and T. crocodilus are worldwide polytypic species in tropical and subtropical seas within the 23.9 °C isothere (Cressey and Collette 1970: figs. 177, 178). In the western Atlantic, T. a. acus is known from Woods Hole, Buzzards Bay, and Nantucket, MA, south along the Atlantic coast of the United States, at Bermuda, and throughout the Gulf of Mexico and Caribbean Sea south at least to Rio de Janeiro and São Sebastião, Brazil (Fig. 2.13). Study Material. A total of 151 specimens (21.9–950 mm BL), including the types of Belone latimana Poey, B. jonesii Goode, and B. jonesii Günther from 103 western Atlantic collections originally at AMNH, ANSP, BMNH, FMNH, FSU, LACM, MCZ, MNRJ, MZUSP, ROM, SIO, TU, UMML, UMMZ, USNM, and ZMK. Comparative material from the eastern Atlantic (see Collette and Parin 1970) includes 27 specimens (156–495 mm BL) from 8 collections from the Mediterranean, a specimen of T. a. imperialis (505 mm BL) from the Cape Verdes Islands, and 58 specimens of T. a. rafale (207–550) from 30 collections from the Gulf of Guinea. 48

Family Belonidae ro y C. and Bravo-Hollis 1969 (three species of Monogenea found on gills of specimens from Veracruz, Mexico). Swingle 1971:27 (Dauphin Island, AL). Buen 1972:159 (Mexico). Duarte-Bello and Buesa 1973:79–80 (synonymy, range, Cuba). Guitart 1975:244–245 (Cuba, synonymy, description, fig. 179). Instituto Nacional de Pesca 1976:6, 74 (distribution, common names), 164 (fig. 104, photograph). Román 1979:82 (description, common name; Venezuela). Sacchi et al. 1981:2 (French Antilles). Rodríguez et al. 1984 (agujón de costa; Cuba). Pérez Martínez et al. 1995:266 (Cuba). ValdésMuñoz and Garrido 1987:10 (Cuba). Aguilar et al. 2001:131 (Bahía de Cienfuegos, Cuba). Strongylura raphidoma. Fowler 1919a:5 (Jamaica, Puerto Rico). Fowler 1919b:146 (Jamaica). Fowler 1926:250 (Boca Grande, FL). Breder 1927:18 (Bahamas). Beebe and Tee-Van 1928:63– 65 (description; growth, habits; Port-au-Prince Bay, Haiti). Breder 1929:279–280 (behavior, maturity; Tortugas), fig. 5 (young and adult), fig. 6. Fowler 1941d:141 (Brazilian records). Longley and Hildebrand 1941:29 (description, behavior; Tortugas). Fowler 1945:281 (synonymy; Key West, FL). Manter 1947:311–312, 324–325, 375 (trematodes; Tortugas). Breder 1948:48 (description, range, figure). Schultz 1949:80 (listed after Röhl 1942). Fowler 1952b:87 (Hispaniola). Santos 1952:92 (Brazil). Woods 1952:40 (Alacran Reef, Yucatán). Chandler 1954:353 (cestode record after Schuler 1938). Manter 1954:340 (trematode). Joseph and Yerger 1956:126 (juveniles; Alligator Harbor, FL). Springer and Bullis 1956:57 (Gulf of Mexico). Briggs 1958:264 (distribution). Robins 1958:77 (both sides of Atlantic). DuarteBello 1959:43 (Cuba; both sides of Atlantic). Hoese 1959:325 (Port Aransas, TX). Randall 1960 (records of people injured by leaping needlefish; photographs of 52-inch female; Virgin Islands). Randall and Randall 1960:471 (young resemble broken bits of grass, previous records). Springer and Woodburn 1960:24 (Tampa Bay area, Florida). Springer and McErlean 1962:53 (seasonality; Matecumbe Key, FL). Nahhas and Cable 1964:223 (trematodes). Breder and Rosen 1966:303 (reproduction; after Breder and Rasquin 1952, 1954). Dahl 1971:180 (description, food value; Colombia). Moe 1972:10 (migrations; western coast of Florida). Belone (Tylosurus) raphidoma. Metzelaar 1919:32–33 (description; West Indies). Belone maris-rubri maris-rubri. Mees 1962:44–50 (synonymy, description, range). Belone marisrubri marisrubri. Mees 1964:322, 325 (listed). Tylosurus crocodilus. Collette and Berry 1965:390–391 (validity of name). Gooding and Collette 1965 (parasitic copepod). Caldwell 1966a:31 (Jamaica; LACM 5878). Collette and Berry 1966:325–326 and Collette 1967:197 (validity of name). Randall 1967:684 (stomach contents). Böhlke and Chaplin 1968:119 (description, range, figure). Randall 1968:40 (description), fig. 40 (photograph). Starck 1968:17 (Alligator Reef, FL). Dragovich 1969:17 (eaten by Euthynnus alletter­ atus). Overstreet 1969:152, 172 (trematode Steganoderma nitens; Biscayne Bay, FL). Collette and Parin 1970:52–55 (synonymy, description), fig. 13 (480 mm BL specimen from Jamaica). de Sylva 1970:51 (Biscayne Bay area, Florida). Mago Leccia 1970:89 (marao; Venezuela; listed). Powell et al. 1972:49 (14 Florida collections). Duarte-Bello and Buesa 1973:79 (questionable occurrence in Cuba). Palacio 1974:35 (Caribbean Colombia). Belyanina 1975:140 (ichthyoplankton; Gulf of Mexico). Smith et al. 1975:5 (Florida Middle Grounds). Erdman 1976:23 (ripe females, October–November; northeastern Caribbean). Shiino 1976:92 (common names). Erdman 1977:159 (juveniles off La Parguera, Puer-

to Rico). Gilmore 1977:132 (Indian River Lagoon area, Florida). Hoese and Moore 1977:153 (description), pl. 132 (color photograph). Gilmore et al. 1978:94 (previous records of hypothermal mortality, Florida). Hardy 1978:102–108 (description, distribution, ecology, spawning, development), fig. 51 (larvae), fig. 52 (juveniles), fig. 53 (development of beak), fig. 54 (development of dorsal and caudal fin). Lima and Oliveira 1978:15 (local names; Ceará, Brazil). Lindén et al. 1978:254 (Cartagena Bay, Colombia). Mota Alves and Lima 1978:3, 6 (common name, spawning season; Brazil). Naughton and Saloman 1978:51 (St. Andrew Bay, FL). Terrero and Bonnelly de Calventi 1978:7 (Dominican Republic). Hastings 1979:7 (St. Andrew Bay jetties, Florida). Castro-Aguirre and Márquez-Espinoza 1981:13 (Isla Lobos, Veracruz, Mexico). Gilmore et al. 1981:10 (Indian River Lagoon area, Florida). Williams, Brusher, and Trent 1984:3 (listed, recreational fisheries; southeastern United States). Fahay 1983:212 (counts). Murdy 1983:88 (figure), 90 (in key, Texas). Acero P. et al. 1984:41 (Caribbean Colombia). Brusher et al. 1984:50; Williams, Brusher, Palko, and Trent 1984:4; Brusher and Palko 1985:57–58, 1986b:6; Williams et al. 1985:6 (caught trolling on recreational charter boats; southeastern United States). Acero P. and Garzón F. 1987:90 (Colombia). Maldonado and Yáñez-Arancibia 1987:191 (sea grass beds, Laguna de Términos, Campeche, Mexico. Fuentes Mata et al. 1989:258 (2 juveniles from Laguna de Sontecomapan, Veracruz, Mexico). Garzón F. 1989:155 (Colombia). Humann 1989:52–53 (description, range; color photograph). Schwartz 1989:340 (North Carolina). Sogard et al. 1989a:58, 1989b:195 (Florida Bay). Smith and Lake 1990:9 (one specimen from the lower Hudson River). Boughton et al. 1991 (ontogeny of upper jaw). Cervigón 1991:225–227 (description, Venezuela, fig. 169). Boschung 1992:84–85 (Dauphin Island, AL). Bouchon-Navaro et al. 1992:313 (Martinique). Louis et al. 1992:296 (Martinique). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Humann 1994:58–9 (description, range; color photograph). Debelius 1997:58–59 (description, color underwater photograph). Murdy et al. 1997:119 (description, Chesapeake Bay, fig. 98). Smith 1997:393 (description, range; color pl. 111). Ho 1998:251 (parasitic copepod, Brazil). Loftus 2000:34 (Everglades National Park). Lovejoy 2000:1352 (fig. 2, molecular phylogeny). González-Gándara and Arias-González 2001:249 (Alacranes Reef, Gulf of Mexico). Paperno et al. 2001:127 (Ponce de Leon Inlet, FL). Briggs and Waldman 2002:58 (New York after Breder 1938). Humann and Deloach 2002:60–1 (description, range; color photograph). McBride and Styer 2002:23 (common year-round in south Florida lampara net fishery). Menezes and Figueiredo 2003:67 (Brazil). Alvarez-León and Celis-Rincón 2004:77 (Archipiélago Islas del Rosario, Colombia). González-Gándara 2004:167 (Veracruz, Mexico). Lovejoy et al. 2004:369, 377 (molecular phylogeny). Poulakis et al. 2004:126 (Charlotte Harbor, FL). Springer and Johnson 2004:129–130, pl. 99 (dorsal gill arch musculature). Casazza and Ross 2008:351 (associated with Sargassum, Gulf Stream off North Carolina). Collette 2010:32–33 (reproduction and development). De Angelo et al. 2014:1043 (seagrass beds in eastern Gulf of Mexico). Gallardo-Torres et al. 2014:89 (Yucatán, Mexico, photographs). Humann and DeLoach 2014:62–63 (description, range; color photograph). Snyder and Burgess 2016:102 (listed, Florida). Belone raphidoma. Erdman 1967:45, 1972:45 (Aguas Prietas Lagoon; Puerto Rico). Tylosurus crocodilus crocodilus. Parin 1967:58–65 (synonymy, description, range). Cressey and Collette 1970:404–405

49

Fishes of the Western North Atlantic 83 total (56 + 28 = 84, 55 + 29 = 84, 54 + 29 = 83). MCZ 624 is designated as the lectotype because it is the largest syntype and is in good condition. Tylosurus gladius Bean in Goode and Bean, 1882. Holotype USNM 30151 (460 mm BL, 672 mm SL); Pensacola, FL (Collette et al. 1992:5). Dorsal fin rays 23; anal fin rays 21; pectoral fin rays 14–14. Vertebrae 54 precaudal + 38 caudal = 82 total. Belone gerania Valenciennes in Cuvier and Valenciennes, 1846. Syntype MNHN A.8896 (665 mm BL); Martinique; Plée (Collette et al. 1997:7). Common Names. In the United States, the name Houndfish has been used for Tylosurus crocodilus almost uniformly since H. M. Smith (1907). The Common and Scientific Names of Fishes from the Unit­ ed States, Canada, and Mexico (Page et al. 2013) lists Houndfish for T. crocodilus and Atlantic Agujon for T. acus, which has also been called Houndfish. Page et al. (2013) lists agujón lisero as the Spanish name in Mexico. Like other needlefishes, T. crocodilus is generally known as marao in Venezuela (Cervigón 1965, 1966; Mago Leccia 1970; Román 1979), but it has sometimes been called mono (Röhl 1942; Mago Leccia 1970). All three needlefishes reported for Colombia by Dahl (1971), T. crocodilus, Strongylura timucu, and Ablennes, are called carajota. In Cuba, it is called agujón de costa (Duarte-Bello 1959; Rodríguez et al. 1984) or agujón crocodilo (Claro and Parenti 2001) to distinguish it from other needlefishes. It has local names of zambaiorolico and zamboque in Ceará, Brazil (Lima and Oliveira 1978; Mota Alves and Lima 1978). In the Yucatán T. crocodilus is known as argujón cocodrilo. Diagnosis. A species of Tylosurus which, in the Atlantic, has fewer vertebrae (79–84), dorsal fin rays (20–24), and anal fin rays (17–22) than T. acus (vertebrae 82–96, dorsal rays 20–26, anal rays 19–24). Tylosurus crocodilus is a short-snouted, heavy-bodied species of Tylosurus compared with T. acus, the Atlantic congener. At a given body length, T. crocodilus has relatively longer pectoral and pelvic fins and higher dorsal and anal lobes than T. acus. Juveniles and subadults of T. crocodilus differ from T. acus and T. choram in having the teeth in both jaws directed prominently anterior. Tylosurus crocodilus has the left gonad better developed than does T. acus. For both sexes, the ratio of left gonad length to right varies from 1.1 to 1.7 compared to 2.3 to more than 15.5 in T. acus. Description. Body elongate, rounded in cross section. Upper and lower jaws greatly elongate and studded with sharp teeth, teeth in both jaws point anteriorly in juveniles. Gill rakers absent. Anterior part of dorsal and anal fins with relatively high

(parasitic copepods), 414 (fig. 178, worldwide distribution). Collette 1978 (description, range, figure). Collette et al. 1984:344 (fig. 179-J, 96.3 mm BL juvenile). Alvarez-Guillen et al. 1988:320 (Quintano Roo, Mexico). Cervigón et al. 1993:270 (northern coast of South America). McEachran and Fechhelm 1998:940, figure (Gulf of Mexico). Claro and Parenti 2001:43 (agujón crocodilo, Cuba). Harper et al. 2001:11 (landings, recreational fishery, Biscayne National Park, FL). Collette 2003a:1113 (figure, description, range in west-central Atlantic). Collette 2003b:10 (synonymy). Banford et al. 2004 (molecular phylogeny). Collette 2005:930– 931 (early life history, figs. a–h). Kohn et al. 2006:70 (three monogenes from Mexico). Schwartz 2006:64 (North Carolina records). Fahay 2007:776, 786–787 (early stages, figs. a–g). Gómez Gaspar and Hernández Avila 2009:15 (Isla Cubagua, Venezuela). McEachran 2009:1285 (Gulf of Mexico). Campos et al. 2010:66 (Rio Grande do Norte, Brazil). Pepperell 2010:232 (color figure). Kells and Carpenter 2011:152–153 (description, color painting). Smith-Vaniz and Jelks 2014:32 (St. Croix, U.S. Virgin Islands records). Starck et al. 2017:91 (Florida Keys). Tylosurus raphydoma [sic]. Reséndez Medina 1971:14 (La Blanquilla reef off Veracruz, Mexico). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Tylosurus croccodilus [sic]. Mejía Aristizábal et al. 1998:30 (Colombian Caribbean oceanic atolls, listed). Tylosurus crocodylus [sic]. Reckel and Melzer 2004:15 (curtain-like falciform process in eye).

Misidentifications. The specimen reported by Bean (1890b) from Cozumel, Yucatán, Mexico (USNM 37124), as Tylosurus caribbaeus is T. crocodilus crocodilus. Types. Belona crocodila Peron and Lesueur in Lesueur, 1821; Mauritius. No types were located. The counts of dorsal fin rays (22) and anal fin rays (21) indicate that the name crocodila refers to the species under discussion. Lesueur’s comparison with Tylosurus acus (as Belona carribaeus) confirms this. Types of Atlantic Nominal Species. Belone raphidoma Ranzani. Holotype MZUB (The Zoology Museum, University of Bologna) (about 320 mm BL); Brazil. After examination through the bottle, the figure in the original description, the number of dorsal rays (22), and the anteriorly curved teeth in the jaws, we conclude that Belone raphidoma is a synonym of Tylo­ surus crocodilus and not of T. acus. Belone crassa Poey, 1860. Holotype MCZ 623 (467 mm BL, 669 mm SL); Cuba; Poey. Dorsal fin rays 23; anal fin rays 22. Vertebrae total 83. Belone melanochira Poey, 1860. Lectotype, designated herein, MCZ 624 (322 mm BL); Cuba; Poey. Paralectotype MCZ 72048 (1, 300 mm BL), USNM 4744 (1, 243 mm BL), and ZMB 5467 (1, 246 mm BL); Cuba. Data for lectotype followed by paralectotypes (in parentheses): dorsal fin rays 22 (22, 24, 23); anal fin rays 21 (21, 21, 21); pectoral fin rays 14–15 (?–15, 14–14, 14–14). Vertebrae 54 precaudal + 29 caudal = 50

Family Belonidae 1941), Smooth Hammerhead, Sphyrna zygaena (Breder 1932a), Little Tunny, Euthynnus alletteratus (Dragovich 1969), and terns (Breder 1932a). Parasites. Fifteen species of parasites have been reported from western Atlantic Tylosurus crocodilus crocodilus: five copepods, an isopod, a branchiuran, three digenetic trematodes, three monogenes, a cestode, and a nematode. copepods: Eight species of copepods were reported from Tylosurus crocodilus by Cressey and Collette (1970), but only five of these were found on western Atlantic individuals: Bomolochidae, Bomolochus bel­ lones Burmeister (genus following Ho and Lin 2009), 12 females and 5 males from the gill chambers and under the oral valves of 8 specimens from Florida, Cuba, and Venezuela; Caligidae, Caligus sp., 4 chalimus larvae from the fins of 3 specimens from Florida and Venezuela, and Caligodes laciniatus (Krøyer), 19 females from under the oral valves of 7 specimens from Florida, the West Indies, and Venezuela; Anthosomatidae, Lernanthropus tylosuri Richardi, 40 females and 39 males attached to the gill filaments of 20 specimens from Florida, the Bahamas, and Venezuela; and Philichthyidae, Colobomatus goodingi Cressey and Collette, 23 females from cephalic lateral line canals of 18 specimens from Florida, Bahamas, Trinidad, and Venezuela. isopods: Although Bruce (1986) reported several

lobes, contained 5.4–10.6 and 5.5–8.0 times in BL, respectively; pectoral and pelvic fins relatively long, contained 6.6–8.3 and 7.3–10.6 times in BL, respectively. A small black lateral keel on caudal peduncle, caudal peduncle deeper than wide; caudal fin deeply forked, lower lobe much longer than upper. Both right and left gonads present, right longer than left. Dorsal fin rays 20–24, usually 22 or 23; anal fin rays 17–22, usually 20 or 21; pectoral fin rays 13–15, usually 14 or 15 (Table 2.18). Total vertebrae 80–84 (Table 2.19). Predorsal scales tiny and very numerous, 240–290 (Table 2.20). Color. Dark bluish green above, silvery white below. A dark blue stripe along sides. Juveniles have an elevated black lobe in the posterior part of the dorsal fin that is lost with growth. Size. The maximum observed size for Tylosurus crocodilus crocodilus is 101.3 cm SL and 71.5 cm BL in the west-central Atlantic; commonly to 90 cm SL. The all-tackle gamefish record is 4.88 kg for a fish caught April 2013 in Goulding Cay, Bahamas (IGFA 2018). Biology Predators. In the western Atlantic, Tylosurus croc­ odilus crocodilus is eaten by barracuda (Jordan and Thompson 1905), sharks (Longley and Hildebrand

Figure 2.14. Houndfish, Tylosurus crocodilus crocodilus. A. USNM 32077, 480 mm BL. Collected in Jamaica, 10 May 1882. B. USNM 198407, 96.3 mm BL. Collected at 37°08′N, 68°14′W from M/V Delaware, 12 June 1958. Illustrations by M. H. Carrington.

51

Fishes of the Western North Atlantic species of cymothoid isopods from Tylosurus crocodi­ lus, only Mothocya xenobranchia Bruce was found on western Atlantic individuals. branchiurans: One specimen of Argulidae was found on a specimen of Tylosurus crocodilus crocodilus from Sarasota, FL (Cressey and Collette 1970:404). digenetic trematodes: Three species of trematodes in two families have been reported from Tylosurus crocodilus crocodilus in the western Atlantic: Haplosplanchnidae, Schikhobalotrema acutum (Linton) was originally described from T. acus acus and subsequently reported from T. c. crocodilus from the Tortugas (Manter 1947), Bimini (Sogandares-Bernal 1959), and Jamaica (Nahhas and Cable 1964); Zoogonidae, two species of Steganoderma, S. elongatum Manter 1947, from the Tortugas, and S. nitens (Linton 1898), originally described from T. a. acus, from Woods Hole, MA, and then reported from T. c. croco­ dilus in Biscayne Bay, FL (Overstreet 1969). monogenes: Three species of Monogenea have been reported from Tylosurus crocodilus crocodilus in Mexico (Caballero y C. and Bravo-Hollis 1969; Kohn et al. 2006): Axinidae, two species of Axinoides, A. jimenezi Caballero y C. and Bravo-Hollis and A. ra­ phidoma Hargis; and Dactylogyridae, one species, Bychowskymonogenea sogandaresi Caballero y C. and Bravo-Hollis. cestodes: One cestode, Ptychobothridae, Ptychobo­ thrium belones (Dujardin), is known from a variety of needlefishes from both sides of the Atlantic (Rees 1969). One specimen was collected from a Tylosurus crocodilus crocodilus from the Tortugas (Shuler 1938). nematodes: Specimens from the Tortugas were parasitized by a filarid ventral to the kidney and in the ovaries (Breder 1932a:17, pl. 10, figs. 2, 3). This was identified as belonging to the genus Philometra and perhaps close to those identified from Tylosurus acus acus at Bermuda as Philometra (Ichthyonema) globiceps Rudolphi by Linton (1908). Food. Like other needlefishes, Tylosurus crocodilus crocodilus feeds mostly on small fishes plus some crustaceans (Jordan and Thompson 1905; Linton 1905; Randall 1967). In the West Indies, fishes made up 91% of its diet by volume and included Acanthurus, Anchoa, Ctengraulis, Harengula, and Mugil (Randall 1967). They catch fish in their jaws, manipulate them, and swallow them head first (Jordan and Thompson 1905). Reproduction. Ovaries were well developed in females from Haiti in March and April, leading to a prediction that spawning would occur a couple of months later (Beebe and Tee-Van 1928). The spawning season has been reported as May–June in Ceará, Brazil (Mota Alves and Lima 1978). Erdman (1976),

however, reported ripe females in Puerto Rico in October and November. Both gonads are developed, but the right is longer than the left (ratio of left to right 1.1–1.5). An 860 mm female had 7,535 eggs in the left ovary (3.6–4.4 mm in diameter) and 23,721 eggs in the right ovary (3.9–4.8 mm in diameter), for a total of 31,256 eggs (x̅ 4.15 mm in diameter). A 52-inch female from the Virgin Islands weighed 10 pounds, 9 ounces, and had a 2-pound ovary containing more than 25,000 eggs that were 4.1–4.6 mm in diameter (Randall 1960). Development. Early development in Tylosurus has not been studied in the Atlantic, but there is an account of the eggs and developmental stages of T. c. crocodilus based on a large mass of large eggs (4.0–4.1 mm in diameter) washed ashore in October in India (Masurekar 1968). Hatching takes place about 8–10 days after fertilization. The newly hatched larvae are 10.7–11.0 mm, type of length unspecified, but likely total length. The yolk sac is absorbed about 24 hours after hatching, at a length of 12.9 mm. Formation of rays in the dorsal, anal, and caudal fins, but not the pelvics, is complete at this time. The jaws begin to elongate, teeth develop, and the pelvics are still fin buds at 44 hours. Rays appear in the pelvic fins by 7.5 days, at a length of 21 mm. Masurekar’s figures of specimens at 10.9, 15.2, 19.3, and 21.0 mm TL are reproduced in Hardy (1978: fig. 51). The same two features of later development discussed in the account of Tylosurus acus are of interest here. Both anterior and posterior lobes of the dorsal fin reach a maximum height of about 20 mm between 200 and 300 mm SL (Breder and Rasquin 1954: fig. 6). Then the melanistic posterior lobe of the dorsal fin sloughs off (Breder and Rasquin 1952), whereas the lobe is resorbed in T. acus (Breder and Rasquin 1954). The anterior lobe continues to increase in height with the length of the fish. Both upper and lower jaws grow at the same rate in Tylosurus crocodilus (Breder 1932a: pl. 2, fig. 2), so there is no halfbeak stage as there is in T. acus and many other needlefishes (Collette et al. 1984: fig. 179). There is another character that changes with growth in Tylosurus crocodilus as first pointed out by Mees (1962). The teeth in both the upper and lower jaws point distinctly anteriorly in juvenile T. crocodi­ lus until about 500–600 mm TL, when they begin to straighten out. In specimens over 60 cm TL the teeth are always erect and straight as in other needlefishes, both juveniles and adults. Ecology. Although both Tylosurus crocodilus and T. acus occur worldwide in tropical and subtropical marine waters, T. crocodilus does not occur as far off52

Family Belonidae Table 2.18. Fin ray counts for Tylosurus crocodilus. Dashes indicate a value of zero. Fin rays Dorsal Species and locality

Anal

18

19

20

21

22

23

24

25

17

18

19

20

21

22

23

— — —

— — —

4 — —

13 — 9

67 55 4 3 65 125

1 — 60

— — 1

1 — —

4 — —

7 — 5

47 74 5 2 50 181

7 — 69

— — 1

1

6

23

1

—

—

6

26

—

—

—

—

—

Tylosurus crocodilus crocodilus Western Atlantic Gulf of Guinea Indo-West Pacific Tylosurus crocodilus fodiator Eastern Pacific

1

—

Table 2.19. Vertebral counts of Tylosurus crocodilus. Dashes indicate a value of zero. Vertebrae Species and locality

67

68

69

70

71

72

— — —

— — —

— — —

— — —

— — —

1

10

13

6

1

Tylosurus crocodilus crocodilus Western Atlantic Gulf of Guinea Indo-West Pacific Tylosurus crocodilus fodiator Eastern Pacific

//

78

79

80

81

82

83

84

85

86

— — —

— — —

— 3 —

4 2 2

7 — 3

25 1 15

22 — 26

8 — 36

— — 23

— — 5

1

—

—

—

—

—

—

—

—

—

Table 2.20. Number of predorsal scales of Tylosurus crocodilus. Dashes indicate a value of zero. Predorsal scales Species and locality

241–250

251–260 261–270

271–280

281–290

291–300

301–310

311–320

Tylosurus crocodilus crocodilus Western Atlantic Gulf of Guinea Indo-West Pacific

4 — —

8 — —

8 1 2

1 3 8

1 — 20

— — 25

— — 27

— — 31

Tylosurus crocodilus fodiator Eastern Pacific

—

—

—

5

10

12

10

13

Predorsal scales Species and locality Tylosurus crocodilus crocodilus Western Atlantic Gulf of Guinea Indo-West Pacific Tylosurus crocodilus fodiator Eastern Pacific

321–330

331–340

341–350

351–360

361–370

371–380

381–390

391–400

— — 11

— — 6

— — 0

— — 1

— — —

— — —

— — —

— — —

4

12

5

6

1

1

—

—

shore as T. acus does and seems to be more closely restricted to tropical waters (compare Cressey and Collette 1970: figs. 178, 177). Thus, in the western Atlantic, T. crocodilus apparently does not occur north of North Carolina, whereas T. acus regularly reaches southern Massachusetts. Likewise, T. crocodilus is rare at Bermuda. (See Range, below, for discussion

of early records from both areas.) There are regular northward migrations of T. crocodilus along the west coast of Florida (Moe 1972). Juveniles resemble broken bits of eelgrass in both their color pattern and their habit of drifting, motionless, at the surface (Gudger 1929; Breder 1932a; Breder and Rasquin 1952). 53

Fishes of the Western North Atlantic Gulf of Mexico and Caribbean south at least as far as Salvador, Brazil (Fig. 2.15). An early record from Bermuda (Barbour 1905) included an anal fin ray count of 22 that is more likely to have been Tylosurus acus acus than T. crocodilus. This record was cited by Bean (1906), who did not find the species, and it seems to be the only record that led Beebe and Tee-Van (1933:60) to report it as “not uncommon.” B.B. Collette was unaware of any other records until on a field trip with his class at the Bermuda Biological Station, where John Galbraith landed a fine specimen on hook and line (SmithVaniz and Collette 2013). There are also several early records from the Atlantic coast north of North Carolina. Some of these, such as that of a large specimen from Sea Isle City, NJ (Fowler 1914), were later redetermined as Tylosurus acus (Fowler 1919a). There is a record of a dark-colored needlefish different from Strongylura marina from Sandy Hook Bay, NJ (Breder 1926), that was later (Breder 1932a; Nichols 1949) identified as T. raphidoma, a synonym of T. crocodilus. However, Breder (1926) reported dorsal and anal fin ray counts of 17–17 for this specimen, so it must have been an unusually dark S. marina, certainly not a Tylosurus. We have not seen T. crocodilus from north of North Carolina. Study Material. A total of 168 specimens (45.1–715 mm BL), including the types of Belone raphidoma Ranzani, B. gerania Valenciennes, B. crassa Poey, B. melanochira Poey, and Tylosurus gladius Bean, from 96 collections originally at AMNH, ANSP, BZUB, CAS, CU, FSU, LACM, MCZ, MNHN, MNRJ, MZUSP, ROM, TU, UF, UMML, USNM, ZMB, ZMH, and ZMK. Comparative material from the eastern Atlantic (see Collette and Parin 1970) includes six specimens (102–284 mm BL) from six collections from Liberia, Cameroon, Fernando Po (Equatorial Guinea), and Ascension Island.

Figure 2.15. Atlantic distribution of Houndfish, Tylosurus crocodilus, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

Relationship to Humans. There are a few early records of Tylosurus crocodilus used for food in the Florida Keys (Henshall 1895; Schroeder 1924). It has been reported from Cuban markets (Nichols 1912). In Colombia the meat is considered excellent, although the green bones worry some people (Dahl 1971). It is considered edible and of good quality in Venezuela by Cervigón (1966), but dry and useful only as bait according to Röhl (1942). There is a fishery for it with trammel nets (mandingos) and seines off Venezuela (Collette 1978). Recreational fishermen have shown some interest in trolling for T. crocodilus in the southeastern United States (Brusher et al. 1984; Williams, Brusher, Palko, and Trent 1984; Williams, Brusher, and Trent 1984; Brusher and Palko 1985, 1986; Williams et al. 1985; Bohnsack et al. 1994). Needlefish will readily take a surface lure such as a spoon and give a fair fight on rod and reel (Erdman 1967, 1972). Large needlefishes, particularly Tylosurus crocodilus can be dangerous to humans because they occasionally leap out of the water, particularly when attracted to lights at night. There are several records of injuries and a few fatalities (see Randall 1960). Range. Both Tylosurus crocodilus and T. acus are worldwide polytypic species of tropical and subtropical seas within the 23.9 °C isothere (Cressey and Collette 1970: figs. 178, 177). In the western Atlantic, T. crocodilus occurs from North Carolina through the

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Family Scomberesocidae SAURIES

KATHERINE E. BEMIS and BRUCE B. COLLETTE

Introduction Scomberesocidae is the sister group of the Belonidae, which together form the superfamily Scomberesocoidea. All members of Scomberesocoidea share three derived characters: presence of a premaxillary canal, upper jaw at least slightly elongate, and interdigitation of bony processes from each side of the lower jaw (Collette et al. 1984; also E. M. G. Kokkelmans, “The functional morphology and phylogenetic implications of lower jaw structure of the Scomberesocoidea,” unpublished manuscript). Analyses of the pharyngeal jaw apparatus support the concept that sauries are most closely related to needlefishes (Aschliman et al. 2005). Scomberesocidae is defined by one derived character: the dorsal and anal fins are followed by a series of four to seven finlets. Other diagnostic characters include: third pair of upper pharyngeal bones separate, fourth upper pharyngeal bone usually present, and scales on the body are relatively small. There are no spines in the fins. The dorsal, with 14–18 rays, including the finlets, and anal fins, with 16–21 rays, including the finlets, are posterior in position; the pelvic fins, with 6 soft rays, are abdominal in position; pectoral fins are short, with 8–15 rays. The caudal fin is deeply forked and symmetrical, similar to a mackerel tail. The lateral line, when present, runs along the ventral margin of the body. Gill raker counts by Parin (1968) are clearly for lower first arch gill rakers and we presume the same is true for Hubbs and Wisner (1980), although this was not stated. Maximum observed size of the two

large species, Cololabis saira and Scomberesox saurus, is 350–400 mm SL; maximum observed size of the two dwarf species is 68 mm SL for C. adocoetus and 126 mm SL for S. simulans (Hubbs and Wisner 1980). Hubbs and Wisner (1980) placed the four species of sauries in four monotypic genera: Scomberesox and its dwarf derivative Nanichthys, and Cololabis and its dwarf derivative Elassichthys. Rather than recognize four monotypic genera, we recognize two evolutionary lines in the family by considering Nanichthys a synonym of Scomberesox and Elassichthys a synonym of Cololabis (Collette et al. 1984). Both species of Scomberesox develop an elongate beak in comparison to Cololabis; the snout increases in length in S. simulans throughout its life span and in S. saurus until a length of about 200 mm SL. The two dwarf species Cololabis adocoetus and Scomberesox simulans differ convergently from the two larger species, C. saira and S. saurus, in being much smaller, losing one ovary and the swim bladder, and in having fewer vertebrae, branchiostegal rays, pectoral fin rays, and gill rakers. Scomberesox inhabits the Atlantic, Pacific, and Indian oceans; Cololabis is restricted to the Pacific Ocean. Sauries spend most of their life in warm homogeneous surface layers of the open sea, far from shallow continental shelf waters. They live close to the surface, so close that in English waters, where Atlantic Saury are plentiful in summer, few are caught in nets as deep as 2 m. Atlantic Saury are one of the most abundant epipelagic planktivores inhabiting the open part of the Atlantic Ocean, feeding mainly on copepods, euphausiids, and amphipods

Fishes of the Western North Atlantic (Dudnik et al. 1981). The great abundance of sauries and their wide distribution make them an important link in the epipelagic food chain of the ocean, by transferring energy from lower to higher trophic levels. Sauries are valuable food fishes in some parts of the world; the Pacific Saury (Cololabis saira) is commercially one of the most important small pelagic fishery resources in the northwestern Pacific Ocean (Tseng et al. 2011). Modern catch of Pacific Saury by all countries is 400,000 to 600,000 tons, and increasing because of the expansion of fishing areas (Baitaliuk et al. 2013). Atlantic Saury are important in the Mediterranean (Potoschi 1996). At present, there is no fishery for saury in the northwest Atlantic, but an experimental fishery was conducted by Russian vessels from 1969 to 1974. Sauries were attracted by bright lights and caught in nets suspended from booms along the sides of the vessels (Zilanov and Bogdanov 1969). There are reports of a Miocene fossil, †Scomberesox acutillus Jordan and Gilbert (1919:36–37, pl. 14, fig. 3), from the family Scomberesocidae. However, in

their revision of the sauries Hubbs and Wisner (1980) removed this species from the family because of the absence of finlets, significantly fewer vertebrae, and discrepancy in the lengths of the caudal peduncle and anal fin base. †Praescomberesox pacifi­ cus David (1946:58–59) also seems to be referred to Scomberesocidae in error (Hubbs and Wisner 1980). There is no otolith-based fossil record for the family (Nolf 2013). Materials and Methods We did not examine many specimens of Scomberesocidae. This chapter is primarily based on a review of the literature, especially from Hubbs and Wisner (1980). Unless otherwise indicated, all body lengths are reported as standard length (SL) or total length (TL) in millimeters. Standard two-letter postal codes are used for U.S. locations. Standard symbolic codes for institutions housing museum specimens follow Sabaj (2016).

Key to the North Atlantic Sauries (modified from Hubbs and Wisner 1980) 1a. Both jaws produced into long, slender beaks in specimens more than 100 mm SL; gill rakers on first arch numerous (34–45); pectoral rays 12–15; ovaries paired. . . . . . . . . . . . . . . . . . . . . . Scomberesox saurus saurus 1b. Jaws of adults produced as slender beaks, the lower about twice the length of the upper; gill rakers on first arch fewer (19–26); pectoral rays 10–11; ovary single. . . . . . . . . . . . . . . . . . . . . . . . . . . . Scomberesox simulans

Species. There are two species of Scomberesox, S. saurus, which has two subspecies (S. saurus saurus and S. saurus scombroides), and S. simulans. Scomberesox saurus has paired gonads whereas S. simulans only has one ovary, although the testes are paired but form a coherent mass with the left overtopping the right. Juveniles can be distinguished by the beak, which in S. saurus develops at 130–150 mm SL, whereas in S. simulans it develops at around 60 mm SL. Range. Scomberesox saurus is antitropical and S. simulans occurs in warm temperate waters of the Atlantic and Indian oceans and, at least in Atlantic waters, is somewhat antitropical as well. In the northwest Atlantic, sauries occur from Cape Hatteras to Newfoundland. Their preferred habitat is to the west of the Gulf Stream in the open ocean. There are scattered records from Bermuda, northern Florida, the Bahamas, and Cuba.

Genus Scomberesox Lacepède, 1803 Scomberesox Lacepède, 1803:344. Type species Scomberesox camperii Lacepède, 1803. Type by monotypy. Sayris Rafinesque, 1810a:60. Type species Sayris recurvirostra Rafinesque, 1810. Type by subsequent designation of Jordan and Evermann 1896:725. Grammiconotus Costa, 1862:55. Type species Grammiconotus bicolor Costa, 1862. Type by monotypy. Nanichthys Hubbs and Wisner, 1980:530. Type species Nan­ ichthys simulans Hubbs and Wisner, 1980. Type by original designation (also monotypic).

Diagnosis. Both species of Scomberesox develop an elongate beak; the snout increases in length in S. simulans throughout its life span and in S. saurus until a length of about 200 mm SL. Scomberesox also have uniserial and biserial teeth in both the upper and lower jaw (Cololabis have only uniserial teeth) (Hubbs and Wisner 1980). 80

Family Scomberesocidae Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Atlantic Saury (English) and balaou (French). It has also historically been called “skipjack” or “skipper” (Goode 1884). Diagnosis. Scomberesox saurus saurus differs from S. s. scombroides primarily in having fewer gill rakers (34–45, usually 38–40, rather than 39–51, usually 42–46, respectively). Description. Upper and lower jaws prolonged, lower jaw slightly longer in adults, much longer in juveniles (Hardy 1978). Lower jaw with small buttresses interdigitating between the left and right halves of the jaw (Kokkelmans, unpublished manuscript). Teeth small and pointed. Five to seven small separate finlets, dorsally between dorsal and caudal fin, ventrally between anal and caudal fin. Body about nine times as long as deep, laterally flattened, tapering toward head and tail, with slender caudal peduncle. All fins small. Dorsal fin originates slightly behind origin of anal; these fins similar in outline, originating posteriorly. Pelvic fins about midway along body. Caudal fin deeply forked and symmetrical, similar to a mackerel tail. Trunk covered with small scales. Patch of scales on each gill cover. Lateral line scales of adults (270 mm) lack circuli; circuli present weakly in smaller individuals, most defined in the smallest specimens (Hubbs and Wisner 1980). Dorsal fin rays 15–18, including 5–6 finlets; anal fin rays 17–21, including 5–7 finlets; pectoral fin rays 12–15; procurrent caudal rays 5–7 (Table 3.1). Vertebrae 39–43 precaudal + 24–28 caudal = 64–70 total (Table 3.2). Predorsal scales 73–81. Gill rakers on first arch 34–45, usually 39–41 (Table 3.3). The number of gill rakers and their sizes increase as the fish grows and stabilizes at about 150 mm SL (Chigirinskiy and Volkov 1971). Color. Olive green above with a silver band on each side at the level of the eye and about as broad as the latter. There is a dark green spot above the base of

Scomberesox saurus saurus (Walbaum, 1792) Atlantic Saury Figures 3.1, 3.2 Tables 3.1, 3.2, 3.3 Esox brasiliensis Pen na nt, 1787:145 (Croque Ha rbor, Newfoundland, Canada, 51°03ˇN, 55°48ˇW). Esox saurus Walbaum, 1792:93 (Cornwall and British seas). Sayris recurvirostris Rafinesque, 1810a:61 (original description); 1810b:33 (Messina, Italy). Scomberesox camperii Lacepède, 1803:344, 345, pl. 6 (fig. 3) (no locality). Scomberesox scutellatum Lesueu r, 1821:132 (ba n k s of Newfoundland, Canada [stomach content]). Discussed by Hubbs and Wisner (1980). Scomberesox equirostrum Lesueur, 1821:132 (no locality). Scomberesox storeri DeKay, 1842:229, pl. 35 (fig. 111) (no locality). Scombresox Camperi [sic]. Cuvier and Valenciennes 1846:464–472 (description, European seas). Scombresox Rondeletii Cuvier and Valenciennes, 1846:472–476 (description, Mediterranean Sea). Scombresox scutellatus [sic]. Cuvier and Valenciennes, 1846:477– 479 (description, Saint Helena). Scomberesox saurus. Bean 1902:407 (Long Island, NY). Fowler 1917:121 (Newport, RI). D’Ancona 1931:150–153, pl. 9, figs. 1319 (Naples, description, development). Sauskan and Semenov 1969 (North Atlantic distribution). Zilanov and Bogdanov 1969 (reproductive biology). Chigirinskiy and Volkov 1971 (gill rakers, pharyngeal). Hardy 1978:79–81 (development). Wisner 1981 (description, figure, map). Andres and John 1984 (central North Atlantic). Scott and Scott 1988:310–312 (biology, distribution in Canadian waters). McCord and Campana 2003:60 (occasional in stomachs of Blue Shark [Prionace glau­ ca], Nova Scotia, Canada). Aschliman et al. 2005 (pharyngeal jaw apparatus). Fahay 2007:778–779 (development). Nolf 2013:75, pl. 156 (otolith, European seas). Scomberesox saurus saurus. Hubbs and Wisner 1980 (recognition of northern and southern subspecies). Collette 2002:285–287 (description, biology, Gulf of Maine). Hardy and Collette 2006:905–907 (development). Agüera and Brophy 2012:60–66 (age and growth of northeastern Atlantic Ocean population). Collette 2016:2130 (description, figure, map).

Types. There are no known types of any of the synonyms of the western Atlantic subspecies Scomberesox saurus saurus.

Table 3.1. Number of fin rays of Scomberesocidae, based on Hubbs and Wisner (1980). Total dorsal and anal rays include finlets. Subspecies are combined for Scomberesox saurus because there is no variation. Dashes indicate a value of zero. Fin rays Dorsal

Anal

Pectoral

Species

14 15 16 17 18

17 18 19 20 21

10 11 12 13 14 15

Scomberesox saurus Scomberesox simulans

6 45 28 14 49 16

1

1 18 84 30 11 1 9 48 20

8 108 37 99 54

1

Procurrent caudal rays 5–7 4 (rarely 3 or 5)

81

ts of Scomberesocidae, based on Hubbs and Wisner (1980). Subspecies are combined for Scomberesox saurus because there is no variation. f zero.

Fishes of the Western North Atlantic

Figure 3.1. Atlantic Saury, Scomberesox saurus saurus. A. USNM 19853, 274 mm TL. Illustration by H. L. Todd for Goode (1884: pl. 181); reproduced from the collections of the Division of Fishes, National Museum of Natural History, Smithsonian Institution. B. 59 mm SL. From Parin (1968:279, fig. 2). Illustrator unknown.

each pectoral fin; the dorsal fin is greenish; the lower parts are silvery with a golden gloss. Young fry, which live in the surface waters of the open Atlantic, have dark blue backs and silvery sides. Size and Age. Maximum confirmed size in western Atlantic is 294 mm SL (Parin 1968). Bigelow and Schroder (1953) reported they reach 18 inches (45.7 cm), but there are no museum specimens associated with this size record. The record of 762 mm SL first reported by Nichols and Breder (1927) is a misidentification. Sauries are difficult to age because annual growth rings are indistinct in the scales and otoliths, but size-frequency data indicates three distinct peaks (5–15 cm, 17–25 cm, and 27–33 cm) that probably correspond to year classes (Sauskan and Semenov 1969; Zilanov and Bogdanov 1969). In the northeast Atlantic, the population available for fishery exploitation is predominately less than one year in age (Agüera and Brophy 2012).

Peninsula S. s. saurus was the most abundant teleost prey (87%) of Shortfin Mako, Isurus oxyrinchus (Biton Porsmoguer et al. 2014). Birds, such as the Northern Gannet (Garthe et al. 2011), Cory’s Shearwater (Neves et al. 2012), and Roseate Tern (Martins et al. 2004), are also known to prey on S. s. saurus. Food. Scomberesox saurus feeds mainly on copepods, euphausids, and amphipods. Larvae of polychaetes, decapods, isopods, ostracods, cirripeds, siphonophores, and fish eggs, fish larvae, and protozoans are consumed in lesser quantities (Dudnik et al. 1981). Reproduction. Oviparous, both ovaries developed; eggs slightly oval, moderately large. Greatest egg diameter 2.32–2.52 mm; no oil globules; yolk clear, nonvesicular; shell with numerous, uniformly spaced, short, rigid bristles, apparently remnants of chorionic filaments (Boehlert 1984; Hardy and Collette 2006). In the Atlantic, spawning grounds expand to lower latitudes as water temperature increases during the annual cycle. Scomberesox saurus saurus is a serial spawner, meaning that a single female reproduces in different localities as suitable spawning area expands (Agüera and Brophy 2011). They reach maturity at 29–32 cm (Zilanov and Bogdanov 1969: table 226) at an age of two years (Sauskan and Semenov 1969). Spawning takes

Biology Predators. Scomberesox saurus saurus was found in stomachs of two Blue Sharks, Prionace glauca, off Nova Scotia, Canada (McCord and Campana 2003). In the eastern Atlantic from the Azores to the Iberian 82

Family Scomberesocidae Range. Marine, epipelagic, usually offshore. Antitropical in temperate parts of the Atlantic, Pacific, and Indian oceans (Fig. 3.2). Two subspecies are recognized (Hubbs and Wisner 1980): the nominal subspecies, Scomberesox saurus saurus, is broadly distributed in the North Atlantic Ocean, mostly north of 30°N (Sauskan and Semenov 1969) and throughout the Mediterranean Sea, whereas S. s. scombroides (Richardson) is found in the Southern Hemisphere. In the northwest Atlantic, sauries are found from Cape Hatteras to Newfoundland. The area to the west of the Gulf Stream core is their main habitat in the open sea of the northwest Atlantic, although they have been taken east of the Gulf Stream. The area of distribution extends from coastal waters eastward to 40°W and from 32°N northward to 50°N. There are scattered records at the southern limit of the range, from Bermuda, northern Florida, the Bahamas, and Cuba (Hubbs and Wisner 1980). In the central North Atlantic, offshore from the Azores and Cape Verde Islands, S. s. saurus made up 26.4% of the total fish and fish larvae caught by neuston nets (Andres and John 1984).

Figure 3.2. Atlantic distribution of Atlantic Saury, Scomberesox saurus, based on specimens examined by Hubbs and Wisner (1980). Map by W. E. Bemis.

Scomberesox simulans (Hubbs and Wisner, 1980) Dwarf Atlantic Saury Figures 3.3, 3.4 Tables 3.1, 3.2, 3.3

place mostly offshore, between the 16.5° isotherm in the north and the 23.5° isotherm in the south. Incubation lasts 14–18 days or more, apparently dependent on water temperature. Eggs and larvae are planktonic (Hardy and Collette 2006). Development. Development was described and illustrated by Sanzo (1940) and Nesterov and Shiganova (1976). Length at hatching is 6.0–9.5 mm; pigmentation at hatching is deep blue, except fins and yolk. Length of lower jaw is initially variable, extending beyond upper jaw in some hatchlings lacking yolk, and shorter than upper jaw in other individuals up to 9.5 mm. Both jaws begin to elongate at about 38 mm. Between 100–150 mm jaws resemble those of halfbeaks; adult beak developed by about 160 mm. Ten or fewer teeth appear on the jaws when fish are about 80–90 mm SL, increasing in number with size to 28–60 teeth on the upper jaw and 19–52 on the lower jaw (Chigirinskiy and Volkov 1971). Finlets first become evident behind dorsal and anal at 15.0–18.0 mm (Hardy 1978; Hardy and Collette 2006). Relationship to Humans. There is a small traditional fishery for Scomberesox saurus saurus in southern Italy and a seasonal fishery in the Mediterranean and Atlantic off the coast of Spain (Agüera and Brophy 2011).

Scomberesox sp. Parin 1968: fig. 1 (undescribed species under study by Hubbs and Wisner). Dudnik 1975 (overview, comparison with S. saurus range and characters; one ovary developed, second rudimentary; spawning biology). Nanichthys simulans Hubbs and Wisner, 1980:531, fig. 5 (original description; south-central Atlantic, 24°02.5ˇS, 15°32ˇW). Wisner 1981 (description, figure, map). Parin and Astakhov 1982:276, figs. 2, 3. Andres and John 1984:147 (central North Atlantic). Scomberesox simulans. Collette et al. 1984:336 placed in Scomberesox. Collette et al. 1992 (USNM types). Collette 2004:3 (synonymy). Collette 2016:2131 (description, figure, map).

Types. Nanichthys simulans Hubbs and Wisner, 1980. Holotype SIO 63-546 (89.5 mm SL), an adult male dipnetted at surface under a light in the south-central Atlantic Ocean, 24°02.5ˇS, 15°32ˇW; 9 June 1963. See Hubbs and Wisner (1980) for full list of paratypes. Common Names. No official common name has been suggested, however Dwarf Atlantic Saury is used by both Dudnik (1975) and Hubbs and Wisner (1980). Diagnosis. A dwarf saury. Swim bladder absent. Single ovary; testes paired, but form coherent mass 83

of zero. Species

14 15 16 17 18

Scomberesox saurus Scomberesox simulans

6 45 28 14 49Dorsal 16

1

1 18 Fin 84 rays 30 11 1 9 Anal 48 20

Species

14 15 16 17 18

17 18 19 20 21

17 18 19 20 21

caudal rays

10 11 12 13 14 15 8 108 37 99 54 Pectoral

1

5–7 4 (rarely 3 or 5) Procurrent caudal rays

10 11 12 13 14 15

Scomberesox saurus 6 45 28 1 1 18 84 30 11 8 108 37 1 5–7 Scomberesox simulans 14 49 16 1 9 48 20 99 54 4 (rarely 3 or 5) Table 3.2. Vertebral counts of Scomberesocidae, based on Hubbs and Wisner (1980). Subspecies are combined for Scomberesox saurus because there is no variation. Dashes indicate a value of zero. Vertebrae Precaudal Table 3.2. Vertebral counts of Scomberesocidae, based on Hubbs and Wisner (1980).Caudal Subspecies are combined for Scomberesox saurusTotal because there is no variation. Dashes indicate a value of zero. Species 35 36 37 38 39 40 41 42 43 22 23 24 25 26 27 28 58 59 60 61 62 63 64 65 66 67 68 69 70 Scomberesox saurus Scomberesox simulans Species

2 25 112 33 3 40 71 11 Precaudal

4

35 36 37 38 39 40 41 42 43

2 11

25 71

22 23

24

97 Vertebrae 50 3 39 3 Caudal 25 26

1

2 11 30 46 21

27 28

9 73 149 83 20

3

1

Total

58 59 60 61 62 63 64 65 66 67 68 69 70

First arch gill rakers

84

Species and localityof gill rakers on 19 22 23 24 25based 26 on // Hubbs 34 35 37 (1980). 38 39 40 41 are42shown 43 for 44Scomberesox 45 46 saurus 47 48because 49 50 51 Table 3.3. Number first 20 arch21 of Scomberesocidae, and 36 Wisner Subspecies of the variation in gill rakers by collection locality. Dashes indicate a value of zero. Scomberesox saurus saurus Northern Hemisphere Scomberesox saurus scombroides Species and locality Southern Hemisphere Scomberesox Species totalsaurus saurus Northern Hemisphere Scomberesox simulans Scomberesox saurus scombroides Southern Hemisphere

19 20

1

3

21

8

22

24

23

19

24

12

25

8

26

4

Species total Scomberesox simulans

1

5

// 34 1 1

1 1

3

8

24

19

12

8

4

11 9 17gill 18 20 First arch rakers

18

35

36

37 38

39 40 6 12

5 5

11 11

9 17 9 17

5

11

9 17

6

5

3

1

41 28

42 43 36 47

44 41

45 43

46 47 48 49 35 19 11 11

24 32 18 20

46 18

42 52 6 5

44 3

44 1

35 19

11 11

4

3

6 12

28

36 47

41

43

35 19

11 11

4

3

24 32

46

42 52

44

44

35 19

11 11

4

3

50 51 4 3

Fishes of the Western North Atlantic

9 73 149 83 20 3 1 Scomberesox saurus 2 25 112 33 4 25 97 50 3 1 2 11 30 46 21 Scomberesox simulans 3 40 71 11 2 11 71 39 3 Table 3.3. Number of gill rakers on first arch of Scomberesocidae, based on Hubbs and Wisner (1980). Subspecies are shown for Scomberesox saurus because of the variation in gill rakers by collection locality. Dashes indicate a value of zero.

Family Scomberesocidae

Figure 3.3. Atlantic Dwarf Saury, Scomberesox simulans. A. Atlantide P342349, 90 mm SL. From Parin (1968:277, fig. 1). B. Atlantide P342350 or P342351, 56 mm SL. Both specimens collected off the Canary Islands, 26°57′N, 17°10′W. From Parin (1968:279, fig. 2). Illustrator unknown.

with the left overtopping the right in ventral view; developed gonads are dorsolateral to the gut and unattached. Scomberesox simulans can be distinguished from juvenile S. saurus because in specimens larger than 60 mm long a beak is present; in S. saurus the beak does not develop until 130–150 mm. Lower numbers of meristic characters are also diagnostic: 10–11 pectoral rays (12–15 in S. saurus), gill rakers 19–26 (39–51 in S. saurus), and vertebrae 58–62 (64– 70 in S. saurus). Description. Both jaws of adults prolonged as slender, fragile beaks, the upper jaw is about half the length of the lower jaw. Teeth in the upper jaw are uniserial behind and biserial forward; in the lower jaw teeth biserial near gape and uniserial forward. Lateral line only developed anteriorly, extending slightly past pelvic base. Long caudal peduncle, especially in comparison to Scomberesox saurus; 77–91 lateral midline scales; lateral line scales of S. simu­ lans differ from the two larger species of sauries in shape and circuli, which are much more numerous and strongly developed, extending over most of the scale except the central part (Hubbs and Wisner 1980:548, fig. 11). Dorsal fin rays 14–17, including about six finlets; anal fin rays 17–21, including about seven finlets; pectoral fin rays 10–11; procurrent caudal rays 4, rarely 3 or 5 (Table 3.1). Vertebrae 35–38 precaudal + 22–26 caudal = 58–62, usually 59–62 total (Table 3.2). Gill rakers on first arch 19–26, usually 22–24 total (Table 3.3).

Size. Size and maturity data unclear. Parin (1968) described a 90 mm SL mature female (Scomberesox sp., NP342349); Dudnik (1975) reported another to be 112 mm; Hubbs and Wisner (1980) in the original description list the two maximum observed lengths as 121 mm (MMF [Museu Municipal do Funchal] 2866) and 126 mm SL (BMNH 1953.3.7). These lengths seem rare; both large fish came from the same locality, Funchal Harbor, Madeira, Portugal. Hubbs and Wisner (1980) examined hundreds of specimens and found no other specimens exceeding 101 mm SL (USNM 204257). Color. Similar in coloration to other sauries; silver ventrally and laterally, becoming greenish with brown specks dorsally (Hubbs and Wisner 1980). Biology Reproduction. Eggs of Scomberesox simulans lack filaments (Hubbs and Wisner 1980). Dudnik (1975) reported that female S. simulans had three size classes of eggs: 0.1–0.5, 0.7–1.1, and 2.0–2.7 mm in diameter. There were 100–150 eggs in each female, but the number of large eggs (2.0–2.7 mm in diameter) was far fewer (8–30 large eggs; Dudnik 1975). This suggests S. simulans is an asynchronous batch spawner. The relatively large diameter of the oocytes, as characteristic of the sauries, is present in S. simulans despite the reduced size of the abdominal cavity. This permits only one ovary to develop, which lowers absolute fecundity of S. simulans (Dudnik 85

Fishes of the Western North Atlantic Literature Cited Agüera, A. and D. Brophy. 2011. Use of sagittal otolith shape analysis to discriminate Northeast Atlantic and Western Mediterranean stocks of Atlantic Saury, Scomberesox saurus saurus (Walbaum). Fisheries Research 110(3):465–471. ——— . 2012. Growth and age of Atlantic Saury, Scomberesox sau­ rus saurus (Walbaum), in the northeastern Atlantic Ocean. Fisheries Research 131–133:60–66. Andres, H. G. and H. C. John. 1984. Results of some neuston net catches in the warmer central North Atlantic— fish larvae and selected invertebrates. Meeresforschung 30(3):144–154. Aschliman, N. C., I. R. Tibbetts, and B. B. Collette. 2005. Relationships of sauries and needlefishes (Teleostei: Scomberesocoidea) to the internally fertilizing halfbeaks (Zenarchopteridae) based on the pharyngeal jaw apparatus. Proceedings of the Biological Society of Washington 118(2):416–427. Baitaliuk, A. A., A. M. Orlov, and Y. K. Ermakov. 2013. Characteristic features of ecology of the Pacific Saury Cololabis saira (Scomberesocidae, Beloniformes) in open waters and in the Northeast Pacific Ocean. Journal of Ichthyology 33(11):899–913. Bean, T. H. 1902. The catalogue of the fishes of Long Island. Sixth Annual Report of the Forest, Fish and Game Commission of the State of New York. pp. 373–478. Bigelow, H. B. and W. C. Schroeder. 1953. Fishes of the Gulf of Maine. United States Fish and Wildlife Service Fishery Bulletin 53(74):1–577. Biton Porsmoguer, S., D. Bănaru, P. Béarez, I. Dekeyser, M. Merchán Fornelino, and C. F. Boudouresque. 2014. Unexpected headless and tailless fish in the stomach content of Shortfin Mako, Isurus oxyrinchus. PLoS ONE 9(2):e88488. Boehlert, G. W. 1984. Scanning electron microscopy. In National Marine Fisheries Service, National Oceanic and Atmospheric Administration; H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Kendall, Jr., and S. L. Richardson, eds. Ontogeny and Systematics of Fishes: based on an international symposium dedicated to the memory of Elbert Halvor Ahlstrom. [Lawrence, KS]: American Society of Ichthyologists and Herpetologists. Special Publication 1. pp. 43–48. Chigirinskiy, A. I and A. F. Volkov. 1971. Some morphological features and age-related changes in the feeding habits of the Atlantic Saury [Scomberesox saurus (Walb]). Voprosy Ikhtiologii 11(5):868–876. [In Russian, English translation in Journal of Ichthyology 11(5):751–759.] Collette, B. B. 2002. Flyingfishes and allies, order Beloniformes. In B. B. Collette and G. Klein-MacPhee, eds. Bigelow and Schroeder’s Fishes of the Gulf of Maine. 3rd ed. Washington, DC: Smithsonian Institution Press. pp. 284–292. ——— . 2004. Family Scomberesocidae Müller 1846—sauries. California Academy of Sciences Annotated Checklists of Fishes 21. 5 pp. ——— . 2016. Scomberesocidae, sauries. In K. E. Carpenter and N. de Angelis, eds. The Living Marine Resources of the Eastern Central Atlantic. Volume 3, Bony fishes, part 1 (Elopiformes to Scorpaeniformes). Rome: Food and Agriculture Organization of the United Nations. FAO Species Identification Guide for Fishery Purposes. pp. 2128–2131. Collette, B. B., G. E. McGowen, N. V. Parin, and S. Mito. 1984. Beloniformes: development and relationships. In National Marine Fisheries Service, National Oceanic and Atmospheric Administration; H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Kendall, Jr., and S. L. Richardson, eds. Ontogeny and

Figure 3.4. Atlantic distribution of Atlantic Dwarf Saury, Scomberesox simulans, based on specimens examined by Hubbs and Wisner (1980). Map by W. E. Bemis.

1975). Dudnik describes the second ovary as being rudimentary and barely discernible; however, in the species description by Hubbs and Wisner (1980) they note that none of the specimens examined had a second ovary. Development. Until Scomberesox simulans reaches 30 mm SL, the jaws are scarcely prolonged. The premaxillaries then are pointed forward and begin to elongate slowly. The dentaries become very slender and elongate far beyond the premaxillaries. At 60 mm SL the lower beak is more than twice the length of the upper, looking most like a halfbeak of all the sauries (Hubbs and Wisner 1980). Range. Prefers warm temperate waters; common in central parts of the north and south Atlantic, but also known from the Indian Ocean (Hubbs and Wisner 1980; Collette 2004). Southern border of the range is ca. 10°N and northern border at ca. 35°N. The occurrence farthest west is near St. Thomas, West Indies (Hubbs and Wisner 1980). In the central North Atlantic, offshore from the Azores and Cape Verde Islands, Scomberesox simulans made up 28.8% of the total fish and fish larvae caught by neuston nets (Andres and John 1984). Like S. saurus, the distribution of S. simulans in the Atlantic is antitropical (Fig. 3.4). 86

Family Scomberesocidae Systematics of Fishes: based on an international symposium dedicated to the memory of Elbert Halvor Ahlstrom. [Lawrence, KS]: American Society of Ichthyologists and Herpetologists. Special Publication 1. pp. 334–354. Collette, B. B., N. V. Parin, and M. S. Nizinski. 1992. Catalog of type specimens of Recent fishes in the National Museum of Natural History, Smithsonian Institution. 3. Beloniformes (Teleostei). Washington, DC: Smithsonian Institution Press. Smithsonian Contributions to Zoology 525. 16 pp. Costa, A. 1862. Di un novello genere di pesci Esocetidei. Annuario del Museo Zoologico della Università di Napoli 1:54–57. Cuvier, G. and A. Valenciennes. 1846. Chapitre 8, Des Scombresésoces. In Histoire naturelle des poissons, Volume 18. Paris: P. Bertrand. pp. 459–482. D’Ancona, U. 1931. Ordine: Synentognathi. In S. Lo Bianco. Uova, larve e stadi giovanili di Teleostei. Roma: G. Bardi. Fauna e Flora del Golfo di Napoli, Monografia 38. pp. 156–176. [In Italian, translated to English for Smithsonian Institution and the National Science Foundation, Washington, DC, by the Israel Program for Scientific Translations, Jerusalem, 1969.] David, L. R. 1946. Some typical Upper Eocene fish scales from California. In E. L. Furlongo. Fossil Vertebrates from Western North America and Mexico. Contributions to Paleontology. [Washington, DC: Carnegie Institute of Washington]. Publication 551. pp. 47–79. DeKay, J. E. 1842. Zoology of New-York; or the New-York Fauna; comprising detailed descriptions of all the animals hitherto observed within the state of New-York, with brief notices of those occasionally found near its borders... Part 4, Fishes. Albany: W. & A. White, J. Visscher. Natural History of New York. 415 pp., pls. 1–79. Dudnik, Y. I. 1975. Contribution to the biology of Dwarf Atlantic Saury Scomberesox sp. Parin (Pisces, Scomberesocidae). Okeanologii 15:738–743. [In Russian, English translation in Oceanology 15(4):503–506.] Dudnik, Y. I., V. K. Zilanov, V. D. Kudrin, V. A. Nesvetov, and A. S. Nesterov. 1981. Distribution and biology of Atlantic Saury, Scomberesox saurus (Walbaum), in the northwest Atlantic. NAFO Scientific Council Studies 1:23–29. Fahay, M. P. 2007. Early Stages of Fishes in the Western North Atlantic Ocean (Davis Strait, southern Greenland and Flemish Cap to Cape Hatteras). Volume 1, Acipenseriformes through Syngnathiformes. Dartmouth, Nova Scotia: Northwest Atlantic Fisheries Organization. 931 pp. Fowler, H. W. 1917. Notes on New England fishes. Proceedings of the Boston Society of Natural History 35(4):109–138. Garthe, S., W. A. Montevecchi, and G. K. Davoren. 2011. Annual changes in prey fields trigger different foraging tactics in a large marine predator. Limnology and Oceanography 56(3):802–812. Goode, G. B. 1884. The Fisheries and Fishery Industries of the United States. Section I, Natural History of Useful Aquatic Animals with an atlas of two hundred and seventy-seven plates. Washington, DC: Government Printing Office. 2 vols. Hardy, J. D., Jr. 1978. Scomberesocidae—sauries. In Development of Fishes of the Mid-Atlantic Bight: an atlas of egg, larval and juvenile stages. Volume 2, Anguillidae through Syngnathidae. [Fort Collins, CO]: U.S. Fish and Wildlife Service. Biological Services Program, FWS/OBS-78/12-02. pp. 71–81. Hardy, J. D., Jr. and B. B. Collette. 2006. Chapter 76, Scomberesocidae: sauries. In W. J. Richards, ed. Early Stages of Atlantic Fishes. Boca Raton, FL: Taylor & Francis. pp. 905–908.

Hubbs, C. L. and R. L. Wisner. 1980. Revision of the sauries (Pisces, Scomberesocidae) with descriptions of two new genera and one new species. Fishery Bulletin 77:521–566. Jordan, D. S. and B. W. Evermann. 1896. Fishes of North and Middle America: a descriptive catalogue of the species of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, Part 1. Bulletin of the United States National Museum 47:1–1240. Jordan, D. S. and J. Z. Gilbert. 1919. Fossil Fishes of the Miocene (Monterey) Formations. In D. S. Jordan. Fossil Fishes of Southern California. [Palo Alto, CA]: Stanford University. Leland Stanford Junior University Publications, University Series 38. pp. 13–60. Lacepède, B. G. E. 1803. Histoire naturelle des poissons, Volume 5. Paris: Plassan. 803 pp. Lesueur, C. A. 1821. Observations on several genera and species of fish, belonging to the natural family of the Esoces. Journal of the Academy of Natural Sciences of Philadelphia 2(1):124–138. Martins, I., J. C. Pereira, J. A. Ramos, and S. E. Jørgensen. 2004. Modelling the effects of different quality prey fish species and of food supply reduction on growth performance of Roseate Tern chicks. Ecological Modelling 177(1–2):95–106. McCord, M. E. and S. E. Campana. 2003. A qualitative assessment of the diet of the Blue Shark (Prionace glauca) off Nova Scotia, Canada. Journal of the Northwest Atlantic Fishery Science 32:57–63. Nesterov, A. A. and T. A. Shiganova. 1976. The eggs and larvae of the Atlantic Saury, Scomberesox saurus of the North Atlantic. Voprosy Ikhtiologii 16(2):315–322. [In Russian, English translation in Journal of Ichthyology 16(2):277–283.] Neves, V., D. Nolf, and M. Clarke. 2012. Spatio-temporal variation in the diet of Cory’s Shearwater Calonectris diomedea in the Azores archipelago, northeast Atlantic. Deep-Sea Research I 70:1–13. Nichols, J. T. and C. M. Breder, Jr. 1927. The Marine Fishes of New York and Southern New England. New York: New York Zoological Society. Zoologica 9. 192 pp. Nolf, D. 2013. The Diversity of Fish Otoliths, Past and Present. Brussels: Royal Belgian Institute of Natural Sciences. 222 pp. Page, L. M., H. Espinosa-Pérez, L. T. Findley, C. R. Gilbert, R. N. Lea, N. E. Mandrak, R. L. Mayden, and J. S. Nelson, eds. 2013. Common and Scientific Names of Fishes from the United States, Canada, and Mexico. 7th ed. Bethesda, MD: American Fisheries Society. Special Publication 34. 384 pp. Parin, N. V. 1968. Scomberesocidae (Pisces, Synentognathi) of the eastern Atlantic Ocean. Atlantide-Report: scientific results of the Danish Expedition to the Coasts of Tropical West Africa, 1945–1946, 10:275–290. Parin, N. V. and D. A. Astakhov. 1982. Studies on the acoustico-​ lateralis system of beloniform fishes with their systematics. Copeia 1982(2):276–291. Pennant, T. 1787. Supplement to the Arctic Zoology. London: H. Hughs. 163 pp. Potoschi, A. 1996. Observations about some biological aspects of Scomberesox saurus (Walbaum, 1792) in the area of the Straits of Messina (Italy). Ophelia 22:139–146. Rafinesque, C. S. 1810a. Caratteri di alcuni nuovi generi e nuove specie di animali e piante della Sicilia, con varie osservazioni sopra i medesimi. Palermo: Sanfilippo. Part 1, pp. [i–iv], 3–69; Part 2, pp. [ia–iva], 71–105. ——— . 1810b. Indice d’ittiologia siciliana; ossia, catalogo metodico dei nomi latini, italiani, e siciliani dei pesci, che si rinvengono in Sicilia disposti secondo un metodo naturale

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Fishes of the Western North Atlantic e seguito da un appendice che contiene la descrizione de alcuni nuovi pesci siciliani. Messina: Giovanni del Nobolo. 70 pp. Sabaj, M. H. 2016. Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an online reference. Version 6.5 (16 August 2016). Washington, DC: American Society of Ichthyologists and Herpetologists. http://www.asih.org/ Sanzo, L. 1940. Sviluppo embrionale e larva appena schiusa di Scomberesox saurus (Flem.). Venezia: C. Ferrari. Regio Comitato Talassografico Italiano Memoria 276. 8 pp., 1 pl. Sauskan, V. I. and G. N. Semenov. 1969. The North Atlantic Saury. Annales Biologiques 25:250–252. Scott, W. B. and M. G. Scott. 1988. Atlantic fishes of Canada. Toronto: University of Toronto Press. Canadian Bulletin of Fisheries and Aquatic Sciences 219. 731 pp. Tseng, C. T., C. L. Sun, S. Z. Yeh, S. C. Chen, W. C. Su, and D. C. Liu. 2011. Influence of climate-driven sea surface tempera-

ture increase on potential habitats of the Pacific Saury (Colo­ labis saira). ICES Journal of Marine Science 68(6):1105–1113. Walbaum, J. J. 1792. Petri Artedi sueci Genera piscium: in quibus systema totum ichthyologiae proponitur cum classibus, ordinibus, generum characteribus, specierum differentiis, observationibus plurimis. Redactis speciebus 242 ad Genera 52. Pars III, Ichthyologiae. Grypeswaldiae: Ant. Ferdin. Röse. 723 pp. Wisner, R. L. 1981. Scomberesocidae, sauries. In W. Fischer, G. Bianchi, and W. B. Scott, eds. FAO Species Identification Sheets for Fishery Purposes: Eastern Central Atlantic (Fishing Areas 34, 47), Volume 3. Ottawa: Department of Fisheries and Oceans Canada with Food and Agriculture Organization of the United Nations. 6 pp. Zilanov, N. V. and S. I. Bogdanov. 1969. Results of research on Scomberesox saurus in the north-eastern Atlantic in 1968. Annales Biologiques 25:252–255.

88

Family Hemiramphidae HALFBEAKS

BRUCE B. COLLETTE and KATHERINE E. BEMIS

Introduction The Hemiramphidae, the halfbeaks, is one of six families of the order Beloniformes. It is the sister group of the Exocoetidae, the flyingfishes, together forming the superfamily Exocoetoidea (Collette et al. 1984). (The Indo-West Pacific family Zenarchopteridae, containing five genera and 54 sexually dimorphic Indo-West Pacific estuarine or freshwater species, was previously considered to be a subfamily of the Hemiramphidae, but is now recognized as a distinct family.) Most halfbeaks have an elongate lower jaw that distinguishes them from the flyingfishes, which have lost the elongate lower jaw, and from the needlefishes (Belonidae) and sauries (Scomberesocidae), which have both jaws elongate. The family is defined by one derived character, a third pair of upper pharyngeal bones anklylosed into a plate. Other diagnostic characters include pectoral fins short or moderately long; premaxillae pointed anteriorly, forming a triangular upper jaw (except in Oxyporhamphus); lower jaw elongate in juveniles of all genera, adults of most genera; parapophyses forked; and swim bladder not extending into haemal canal. The tip of the lower jaw is bright red or orange in most species, due to carotenoid pigments, especially zeaxanthin, astaxanthin, and beta-doradexanthin. Most halfbeaks have tricuspid teeth in their jaws; some have unicuspid teeth. Halfbeaks, like other beloniforms, lack a true stomach, but manage to digest their mostly herbivorous food items with their simple, short gut using enzymes (Day et al. 2011). The Hemiramphidae contains nine genera and subgenera and at least 64 species and subspecies (Col-

lette 2004; Banford 2010). Four genera, the first three monotypic (Arrhamphus, Chriodorus, Melapedalion, and Oxyporhamphus), have very short or no beaks. Euleptorhamphus and Oxyporhamphus contain two offshore species each. Hemiramphus (with 10 species) is a widespread marine genus. Rhynchorhamphus (with 4 species) has fimbriate nasal papillae and is confined to Indo-West Pacific marine waters. Hyporhamphus, the most speciose genus, includes two subgenera: Hyporhamphus with 25 species is widespread and Reporhamphus with 11 species occurs only in the Indo-West Pacific. Some Hyporhamphus species are marine, some estuarine, and some freshwater. All genera are characterized by particular lateral line characters (Parin and Astakhov 1982). We include Oxyporhamphus in the family Hemiramphidae, not the Exocoetidae (Collette 1966; Parin et al. 1980; Collette et al. 1984; Lovejoy et al. 2004; Lewallen et al. 2011). In external morphology, Oxyporhamphus, with short jaws and elongate pectoral fins, resembles the true flyingfishes. Because of this, Oxyporhamphus was at one time placed in the family Exocoetidae (Nichols and Breder 1928; Hubbs 1933; and others) and has been so placed once again by Dasilao et al. (1997). This similarity is a condition not so much of close relationship as of parallel development of the ability to glide in flight over the water. In most of the anatomical characters (presence of forked parapophyses, direct connection of the basioccipital and cleithrum, incomplete fusion of the frontals and parietals, failure of interradial foramina in the pectoral fin skeleton to close, and nonprojection of the swim bladder into the haemal canal [Parin 1961]), and also from the evidence of ontogeny, there is

Fishes of the Western North Atlantic derivatives of marine genera and the rivers in which they are found drain into the western Atlantic. The first attempt at a revision of the western Atlantic halfbeaks was by Meek and Goss (1884) on the genus Hemiramphus (including Hyporhamphus). Confusion over the validity of Hyporhamphus was ended by Miller (1945a), who clearly distinguished the two genera on the basis of characters of the type species of Hemiramphus (brasiliensis) and Hyporhamphus (unifasciatus). Meek and Goss (1884) pointed out that the common inshore halfbeaks of the Atlantic and Gulf coasts of the United States were not the same as those in the West Indies, Central America, and South America. They used the name roberti Valenciennes for the northern species and restricted unifasciatus Ranzani to the southern species. However, this use of names was in error, as shown by Miller (1945b), who clearly distinguished Hyporhamphus roberti (type locality French Guiana) from H. unifasciatus and placed H. hildebrandi Jordan and Evermann (type locality Panama) in the synonymy of roberti. Berry and Collette discovered in 1960 that Meek and Goss were correct in distinguishing between northern and southern species of Hyporhamphus unifasciatus, but that the common halfbeak on the Atlantic and Gulf coasts of the United States was undescribed. This situation was further complicated by the use of the name unifasciatus for halfbeaks in the eastern Pacific, eastern Atlantic, and various localities in the Indo-West Pacific. The eastern Atlantic species is H. picarti (Valenciennes) as shown by Collette (1965) and the Indo-West Pacific species is H. limbatus (Valenciennes) as shown by Parin et al. (1980). The species on the Atlantic and Gulf coasts of the United States was described as H. meeki by Banford and Collette (1993); the eastern Pacific species as H. naos by Banford and Collette (2001); and the Bermuda population as H. collettei by Banford (2010). Relationships to Humans. Halfbeaks are valued food fishes in many parts of the world, such as Australia and New Zealand (Collette 1974). They represent an important and increasingly exploited resource in Australia (Tibbetts et al. 2008). They are also used for food in the West Indies and South America, but are not currently considered an important food fish resource in the United States. Their value in the western Atlantic is largely as forage and bait for a wide variety of important food and game species such as tunas, Spanish mackerels, billfishes, and dolphinfishes (as detailed in the species accounts). Halfbeaks are forage for a wide variety of other fishes and also for seabirds and porpoises. The most direct use of halfbeaks in the western North Atlantic is as bait for game species. Ballyhoo, Hemiramphus brasiliensis

complete similarity to other genera of the Hemiramphidae, particularly Hemiramphus. The Indo-Pacific Oxyporhamphus convexus (but not the widespread O. micropterus) and all 10 species of Hemiramphus share vesicular swimbladders (Tibbetts et al. 2007), a character found elsewhere only in billfishes (Istiophoridae). Analysis of molecular data (Lovejoy et al. 2004; Lewallen et al. 2011) also shows Oxyporhamphus clustering with Hemiramphus. Even the elongate pectoral fins and short (in adult fish) lower jaw of the flying halfbeaks do not distinguish Oxyporhamphus from the other Hemiramphidae, because analogous features are found in several other halfbeaks (pectoral fins elongate in Euleptorhamphus, lower jaw barely elongate in Arrhamphus and Melapedalion and not in the least in Chriodorus). As in other Hemiramphidae, young Oxyporhamphus have two folds of skin ventrally along the midline on each side of the elongate lower jaw. A vestige of the latter is also present in adult fish (after loss of the elongate lower jaw); it has the appearance of a light or pigmented bifurcated structure and when partially broken resembles barbels. From other representatives of the Hemiramphidae, Oxyporhamphus differs only in two anatomical characters that it shares with the Exocoetidae: upper pharyngeal bones of the third gill arch separate, not fused into a single plate, and absence of a triangular projection from the anterior margin of the upper jaw. We consider these differences insufficient to recognize a separate family, Oxyporhamphidae distinguished by Bruun (1935) and accepted by Parin (1961). The only fossil record from the Americas of which we are aware is that of Hulbert (2001) for four vertebrae questionably referred to Hyporhamphus from the early Pleistocene, ca. 1.3 Ma, from the Leisey Shell Pit 3A fauna, Hillsborough Co., Florida, U.S.A. (R. C. Hulbert, pers. comm.) The halfbeak fauna of the western Atlantic includes 10 marine species classified in five genera (Collette 2003b): Chriodorus (1 species), Euleptorhamphus (1 species), Oxyporhamphus (1 species), Hemiramphus (3 species), and Hyporhamphus (4 species), plus two freshwater species of Hyporhamphus in rivers draining into the western Atlantic. The distribution of western North Atlantic marine halfbeaks (except for Oxyporhamphus) is marginal on the Caribbean Plate; they are not widespread throughout the Caribbean seas like flyingfishes. All the Atlantic species of Hyporhamphus belong to the subgenus Hyporhamphus. The two freshwater species, Hyporhamphus mexicanus Alvarez and Hyporhamphus brederi (Fernández-Yépez), from rivers in Central America and South America, respectively, are included in this account of western Atlantic species because we consider these species to be freshwater 90

Family Hemiramphidae out much of the larval stage. During postembryonic development, halfbeaks, like other beloniform fishes, undergo several complex changes in beak length, melanistic dorsal fin lobe, body bars, and pelvic fin pigmentation. Adults of four genera lack the elongate lower jaw that characterizes most members of the family. Juveniles of these four genera, including Chriodorus and Oxyporhamphus in the western Atlantic, have a distinct beak. Juveniles of Hemiramphus and Oxyporhamphus develop a darkened posterior lobe on the dorsal fin similar to that present in two genera of needlefishes, Ablennes and Tylosurus. The 10 species of Hemiramphus have a series of broad vertical dark bars on the body during some stages of their development. Body bars are retained for different periods during development: all body bars are lost before 120 mm SL in H. bermudensis and H. brasiliensis, but are retained past 175 mm in H. balao, and all are retained throughout life in the Indo-Pacific H. far. The species of Hemiramphus also have melanistic pelvic and caudal fins as juveniles. Patterns of pelvic fin pigmentation divide the genus into two species groups, one with pigment concentrated proximally on the fin (H. balao group), and the other with pigment absent basally and concentrated distally (H. brasiliensis group, including H. bermudensis).

(and also H. balao), are considered the best bait for billfishes (see, for example, Sosin 1967). Biology. The biology and life history of most species of halfbeaks has not been well studied. In the western Atlantic, the best-studied species are Hemiramphus brasiliensis and H. balao (Santos 1970; Berkeley et al. 1975; Berkeley and Houde 1978; McBride et al. 1996; McBride 2001), because of a fear that numbers of Hemiramphus for the sport fishery in southern Florida might be diminished by overfishing. There is also pertinent literature on related species, particularly from India, Australia, and Japan. Talwar (1962a) studied the food and feeding relationships of four species of Indian halfbeaks: Hyporhamphus georgii (= Rhynchorhamphus malabaricus), Hyporhamphus quoyi, Hemiramphus marginatus (= Hemiramphus lutkei), and Hemiramphus far. He also reported on the biology of “Rhynchorhamphus malabaricus” and “Hemiramphus lutkei” (Talwar 1962b, 1967). Thomson (1959) investigated feeding habits of fishes in Lake Macquarie, New South Wales, Australia, including two halfbeaks, Hemiramphus ardelio (= Hyporhamphus regularis ardelio) and Hemiramphus australis (= Hyporhamphus australis). The biology of another Australian halfbeak, Reporhamphus melanochir (= Hyporhamphus melanochir) was studied by Ling (1958). Tibbetts and Carseldine (2005) showed that three Australian halfbeaks (Arrhamphus sclerolepis krefftii, Hyporhamphus regularis ardelio, and Hyporhamphus quoyi) undergo a marked ontogenetic shift from an animal to plant diet between 50 and 110 mm SL. This shift is correlated with development of the toothed area on the pharyngeal tooth pads (Tibbetts et al. 2008). Halfbeaks have no true stomach and lack the enzyme cellulase, but the pharyngeal mill efficiently triturates seagrass tissue (Klumpp and Nichols 1983; Day et al. 2011). Age and growth of Hemiramphus sajori (= Hyporhamphus sajori) from Japan was investigated by Hattori and Seki (1959). Species names in cited literature have been corrected or revised. Development. Halfbeak eggs are typically 1.5– 2.5 mm in diameter and have attaching filaments, although these are greatly reduced in length in the pelagic eggs of Oxyporhamphus. (Egg sizes given in the species accounts are a minimum size, as fully ripe, hydrated eggs would be larger.) Halfbeaks hatch at 4.8–11 mm, smaller than needlefishes but larger than flyingfishes and sauries (Collette et al. 1984). Larvae are well developed at hatching with partially to fully pigmented eyes, an open mouth, fully flexed notochord, developing rays in the dorsal, anal, and caudal fins, and a small to moderate yolk sac (Watson 1996). A pre-anal finfold typically is present through-

Materials and Methods Counts and measurements follow other papers by B. B. Collette (e.g., Collette 1974) and are summarized here. All elements in the dorsal, anal, and pectoral fins were counted. Gill rakers were counted on both the first and second arches. Counts are presented in the text as “number on upper arch + number on lower arch = total.” Vertebrae were counted from radiographs and counts are given in the text as “precaudal + caudal = total.” Predorsal scales were counted from the origin of the dorsal fin anteriorly to the posterior end of the head. Ranges of counts are given in the text. Morphometric data collected to distinguish species include standard length (SL); total length (TL); head length, tip of upper jaw to posterior end of opercular membrane; lower jaw length, tip of upper jaw to tip of lower jaw; P1 length, base of uppermost pectoral ray to tip of longest ray; P1–P2, distance from base of uppermost pectoral ray to base of anteriormost pelvic ray; P2–C, distance from base of anteriormost pelvic ray to caudal base (C); length of dorsal fin base; length of anal fin base. Ratios of several of these measurements have proved particularly useful: P1–P2/P2–C; head length in standard length; lower jaw length in head length; pectoral and pelvic fin length in standard length; length of 91

Fishes of the Western North Atlantic anal fin base in dorsal fin base. Minimum egg size is reported, but fully ripe, hydrated eggs would be larger. These measurements were recorded by selecting a large female from each species, calculating the volume of each ovary, and counting and measuring a

subsample of the eggs to estimate totals for each ovary. Standard two-letter postal codes are used for U.S. locations. Standard symbolic codes for institutions housing museum specimens at the time they were examined follow Sabaj (2016).

Key to the Hemiramphidae of the Western Atlantic 1a. Lower jaw not noticeably elongate (except in small juveniles). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Lower jaw distinctly elongate; total gill rakers on first arch 25–46; pectoral fin rays usually 7–12. . . . . . . . . . . . 3 2a. Total gill rakers on first arch 19–24; pectoral fins short; pectoral rays usually 13–14. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chriodorus atherinoides (Florida, U.S.A.; Mexico; Cuba) 2b. Total gill rakers on first arch 30–35; pectoral fins long (30%–35% SL); pectoral rays 11–13. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxyporhamphus micropterus similis 3a. Dorsal fin rays 21–24; anal fin rays 20–24; pectoral fins very long; pectoral fin rays 7–9. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euleptorhamphus velox 3b. Dorsal fin rays 12–17; anal fin rays 10–18; pectoral fins short to moderate; pectoral fin rays 9–12. . . . . . . . . . . . 4 4a. Caudal fin deeply forked; scales absent on upper jaw, preorbital ridge absent; anal fin rays usually 10–13 (range 11–15). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiramphus … 5 4b. Caudal fin emarginate or only slightly forked, scales present on upper jaw, preorbital ridge well developed; anal fin rays usually 14–17 (range 14–18) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus … 7 5a. Pectoral fin longer (4.4–5.4 times in SL) reaching beyond anterior margin of nasal pit when folded forward; anal fin rays 10–14, usually 11 or 12; upper caudal lobe blue in life; juveniles with the pelvic fin pigmentation concentrated proximally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiramphus balao (Atlantic) 5b. Pectoral fin shorter (5.2–6.7 times in SL), its length not extending to the anterior margin of the nasal fossa; anal rays 12–15, usually 13; upper lobe of caudal of adults reddish orange in life; juveniles with the tip of the lower lobe of the caudal pigmented and the pelvic fin pigmentation concentrated distally. . . . . . . . . . . . . . . . . . 6 6a. Total gill rakers on first arch 37–44. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiramphus bermudensis (Bermuda) 6b. Total gill rakers on first arch 28–36 (usually 30–32) . . . . . . . . . . . . . . . . . . . . . Hemiramphus brasiliensis (Atlantic) 7a. Scales covering dorsal and anal fins of adults; lower jaw short. . . . . . . . . . . . . . . . . . . . . . . . 8 (with three choices) 7b. Scales absent or only a few present on anterior part of dorsal and anal fins of adults . . . . . . . . . . . . . . . . . . . . . . . 9 8a. Dorsal fin rays 13–16, usually 15; pectoral rays 9–12, almost always 11; total gill rakers on first arch 27–35, usually 29–32; total gill rakers on second arch 19–28, usually 23–25; lower jaw short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus unifasciatus (western Atlantic) 8b. Dorsal fin rays 12–16, usually 14 or 15; pectoral rays 11 or 12; total gill rakers on first arch 31–43, usually more than 32; total gill rakers on second arch usually more than 24; lower jaw long . . . . . . . . . . . Hyporhamphus meeki (U.S. Atlantic and Gulf coasts) 8c. Dorsal fin rays 13–16, usually 14 or 15; pectoral rays 10, rarely 9; total gill rakers on first arch 26–32, usually 29–30; total gill rakers on second arch 20–25, usually 21 or 22 . . . . . .Hyporhamphus collettei (only in Bermuda) 9a. Total gill rakers on first arch 22–32. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9b. Total gill rakers on first arch 29–48 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus roberti … 11 10a. Total gill rakers on first arch 28–32; lower jaw equal to about one-half standard length, pectoral fin rays 9–11, usually 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus brederi (Orinoco and Amazon rivers) 10b. Total gill rakers on first arch 22–27; pectoral fin rays 11–12, usually 11. . . . . . . . . . . . . . Hyporhamphus mexicanus (freshwater Atlantic drainage, Central America) 11a. Total gill rakers on first arch usually more than 38; total of dorsal plus anal rays usually more than 30; lower jaw longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus roberti hildebrandi (Belize to Panama) 11b. Total gill rakers on first arch usually less than 39; total of dorsal plus anal rays usually less than 31; lower jaw shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus roberti roberti (Venezuela to Brazil)

92

Family Hemiramphidae Genus Chriodorus Goode and Bean, 1882

otolith with fossil species, Chriodorus miocaenicus). DuarteBello and Suárez-Caabro 1957:2 (Río Chambas o Los Perros, Camagüey, Cuba; used as food). Norman 1957:172 (listed). Briggs 1958:264 (range). Tabb and Manning 1962:611 (least common Florida Bay halfbeak, never in brackish areas, abundant in clear water around Florida Keys). Collette 1966:5–7 (meristic data). Miller 1966:796 (range). Böhlke and Chaplin 1968:123 (feeds on Batophora in Lake Forsyth). Randall 1968:42 (mentioned). Starck 1968:17 (Alligator Reef, FL). Overstreet 1969:169 (Biscayne Bay, FL). de Sylva 1970:55 (Biscayne Bay, FL). Hudson et al. 1970:10 (listed, Florida Bay). Roessler 1970:864 (Buttonwood Canal, Everglades Park, FL; year-round). De Buen 1972:162 (listed). Duarte-Bello and Buesa 1973:80 (range). Guitart 1975:244–245 (Cuba, synonymy, description, fig. 185). Shiino 1976:89 (common name Hardhead Halfbeak). Instituto Nacional de Pesca 1976:75 (common names pajarito and manjúa; range), 165 (fig. 107). Erdman 1976:23, 1977:159 (ovaries subripe, January– February). Castro-Aguirre 1978:57 (Campeche, after Hubbs 1936). Collette 1978 (description, range, figure). Rodríguez et al. 1984:14 (listed, common name pejerrey; Cuba). Collette et al. 1984:352 (meristic characters). Castro-Aguirre et al. 1986:164 (Veracruz). Robins and Ray 1986:101–102 (description, range), pl. 17. Gilmore 1987:122, 129 (seagrass communities). Maldonado and Yáñez-Arancibia 1987:189 (Campeche). Thayer et al. 1987:30 (few found in seagrasses near red mangrove prop root habitat, southern Florida). Schmidt 1989:167 (eaten by young barracuda, Everglades National Park, FL). Sogard et al. 1989a:58, 1989b:195 (Florida Bay). Thayer and Chester 1989:210 (Florida Bay). Collette et al. 1992:8 (type). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Smith 1997:389 (description, range). McEachran and Fechhelm 1998:957, figure (Gulf of Mexico). SchmitterSoto 1998:79–80 (description, Quintana Roo, Mexico), fig. 36. Claro and Parenti 2001:43 (pejerrey, Cuba, listed). Collette 2003b:1143 (range, figure). McBride and Styer 2003:24–25 (uncommon in southern Florida lampara net fishery). Collette 2004:2–3 (synonymy). Lovejoy et al. 2004:369, 376 (molecular phylogeny). Aschliman et al. 2005:427 (relationships). Collette 2005:936–937 (early life history, figure). Powell et al. 2007:58–60, 209 (densities and monthly length-frequencies, Florida Bay). Tibbetts et al. 2007:809 (single-chambered swim bladder). McEachran 2009:1286 (Gulf of Mexico). Miller et al. 2009:227 (UMMZ specimens from Laguna de Balcar, Quinta Roo, Mexico). Kells and Carpenter 2011:150–151 (description, color painting). Peralta-Meixueiro and Vega-Cendejas 2011:677 (hyperhaline coastal system: Ría Lagartos, Mexico). Gallardo-Torres et al. 2012:279 (permanent residents of the

Chriodorus Goode and Bean, 1882:432 (type species Chriodorus atherinoides by original designation).

Diagnosis. Chriodorus lacks the elongate lower jaw characteristic of most halfbeaks; young specimens have a very short beak that disappears with growth. Chriodorus has the fewest gill rakers of the western Atlantic halfbeaks (20–24), overlapping only with Hyporhamphus mexicanus. Chriodorus also has the most pectoral rays (12–14). Species. A single species, Chriodorus atherinoides. Range. Marine, endemic to the western Atlantic: southern Florida, U.S.A., south through Florida Keys, to the Bahamas, Cuba, Campeche and Yucatán, Mexico, and Belize (Collette 2004). Chriodorus atherinoides Goode and Bean, 1882 Hardhead Halfbeak Figures 4.1, 4.2 Tables 4.1, 4.2, 4.3 Chriodorus atherinoides Goode and Bean, 1882:432 (original description; Key West, FL). Jordan and Gilbert 1883a:903 (description). Jordan 1884a:114 (Key West; hardhead; “an excellent panfish”). Jordan 1884b:77 (abundant at Key West). Bean 1891:300 (first Bahamas record, Spanish Wells). Lönnberg 1894:119 (Key West, FL). Jordan and Evermann 1896b:719 (description, synonymy, range, size to 10 inches, feeds on green algae). Jordan and Evermann 1896a:321 (listed). Evermann and Kendall 1899:150 (Key West). Fowler 1906:91 (Marquesas Keys, FL, description; ANSP 30606). Regan 1911:7 (Chriodorus possible synonym of fossil Cobitopsis). Rosen 1911:51 (Bahamas, after Bean 1891). Fowler 1919a:6 (Marquesas Keys and Hailer’s Rock, FL, ANSP 30721-7). Frost 1926:472 (otolith, pl. 20, fig. 20). Jordan et al. 1930:199 (listed). Breder 1932b:5 (common but difficult to seine; Lake Forsyth, Andros Island, Bahamas). Weed 1933:48 (description). Breder 1934:70 (Lake Forsyth, leap over seine, manipulate peculiar brackish-water algae Batophora to remove diatoms and other organisms). Hubbs 1936:209 (Río Champoton, Campeche, Mexico, description). Fowler 1945:282 (Key West, FL). Breder 1948:90 (Florida Keys). Weinfurter 1952:464 (comparison of

Figure 4.1. Hardhead Halfbeak, Chriodorus atherinoides. USNM 125631, 108 mm SL. Key West, Florida, U.S.A., 22 October 1896. Illustration by M. H. Carrington.

93

Fishes of the Western North Atlantic

94

Family Hemiramphidae

Park, U.S.A. (Schmidt 1989), but Chriodorus atherinoides is undoubtedly eaten by other fishes. Food. Chriodorus atherinoides in Lake Forsythe, Andros Island, Bahamas, manipulates the alga Batophora in its mouth and apparently retains adherent organisms such as diatoms (Breder 1934). Reproduction. A large female (156 mm SL, USNM 295008) collected from Camaguey, Cuba, had 140 eggs in the left ovary (0.75–1.7 mm in diameter, x̅  1.24 mm for 20 eggs) and 140 eggs in the right ovary (1.05–1.80 mm in diameter, x̅  1.35 for 20 eggs). Range. Southern Florida in the United States, from Bonito Beach, Fort Myers (USNM 179876), on the west coast and Miami on the east coast, south through the Florida Keys; Bahamas; Cuba (Camaguey); Campeche and Yucatán, Mexico; Chetumal Bay, Belize (ANSP 96866) (Fig. 4.2). The record from off Charleston, SC (Fahay 1975), was based on a specimen of Hyporhamphus meeki (G. E. McGowen, pers.

coastal lagoon Boca de la Carbonera, Yucatán, Mexico). SosaLopéz et al. 2012:317 (abundance in the Terminos Lagoon in southern Gulf of Mexico). Vega-Cendejas and Santillana 2012:298 (abundance in lagoon and coastal habitats in Yucatán, Mexico). Gallardo Torres et al. 2014:84 (Yucatán, Mexico, photographs). Snyder and Burgess 2016:100 (listed, Florida). Starck et al. 2017:91 (Florida Keys).

Types. Chriodorus atherinoides Goode and Bean, 1882. Holotype USNM 26593 (114 mm SL); Key West, FL; S. Stearns, 1880. Dorsal fin rays 17; anal fin rays 17; pectoral fin rays 13–14; gill rakers on first arch 6 + 17 = 23 (Collette et al. 1992:8). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Hardhead Halfbeak (English), a name used in the United States since Jordan (1884b), and pajarito cabezidura (Spanish, Mexico). It is known as pejerrey in Cuba (Duarte-Bello 1959; Rodríguez et al. 1984; Claro and Parenti 2001). Diagnosis. Chriodorus atherinoides lacks the elongate lower jaw characteristic of most halfbeaks; young specimens have a very short beak that disappears with growth. As its specific name atherinoides implies, Chriodorus looks like a silverside. Chriodorus atherinoides has the fewest gill rakers of the western Atlantic halfbeaks (20–24, x̅ 21.4), overlapping only with Hyporhamphus mexicanus. Chriodorus atherinoides also has the most pectoral rays (12–14, x̅ 13.1). Description. Dorsal fin rays 15–18; anal fin rays 15–17; pectoral fin rays 12–14, usually 13 (Table 4.3). Vertebrae 31–32 (occasionally 33) precaudal + 18–19 caudal = 49–51 total. Gill rakers on first arch 4–6 + 15–18 = 20–24; gill rakers on second arch 2–4 + 13–14 = 16–18. Color. Silver-white with dark silver streak that extends the length of the body. Silver patch on and just behind operculum; dorsal fin light yellow-green (Gallardo Torres et al. 2014:84) Size. The maximum observed size for Chriodorus atherinoides is 164 mm SL. Biology

Figure 4.2. Distribution of Hardhead Halfbeak, Chriodorus atherinoides, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

Predators. There is one record of predation, by a young barracuda, in Florida Bay, Everglades National 95

Fishes of the Western North Atlantic pid in the middle and posterior parts. Teeth present on vomer and on the tongue (in an elongate strip, sometimes distributed along the center line). Gill rakers on first arch 24–35; gill rakers on on second arch 16–24. Olfactory papilla not fimbriate. Dorsal fin rays 20–25; anal fin rays 20–25; pectoral fin rays 7–9 (lower ray sometimes very short, and when its length is less than a fourth the length of the preceding ray, it is recorded as one-half), upper unbranched ray very thick. Pectoral fins rather long (3.6–4.0 times in SL). Pelvic fins short (23–26 times in SL), located in the posterior part of the body appreciably closer to the base of the caudal fin than to the origin of the pectoral fins (P1–P2/P2–C = 1.1–1.3). Origin of anal fin under the fourth to sixth ray of the dorsal fin. Caudal fin with greatly enlarged lower lobe. No scales on fins. Species. Euleptorhamphus contains two very similar species, E. viridis (van Hasselt) in the Indo-Pacific and E. velox Poey in the Atlantic. Parin (1964), having limited material from the Atlantic, considered the genus monotypic. Collette (1965) recognized E. velox from the Atlantic as distinct, because of fewer pectoral rays (7–7½ rather than 8–8½ in E. viridis), fewer rays in the dorsal and anal fins, and slightly more gill rakers. Parin et al. (1980) confirmed the differences in pectoral ray number. Of 304 Indo-Pacific specimens, 294 had 8 pectoral rays, 89 of these had an additional short ray (P1 8½), 2 specimens had 9 rays, and only 8 had 7½ rays. Of 89 Atlantic specimens, 14 had 7 rays and 75 had 7½ rays. Range. Both species are marine: Euleptorhamphus velox in the western Atlantic, from New England south through the Gulf of Mexico and Caribbean Sea to Recife, Brazil, and in the eastern Atlantic from the Cape Verde Islands, Sierra Leone, and Nigeria; E. viridis is widely distributed in tropical and subtropical waters of the Indo-Pacific (Parin et al. 1980).

comm.). Records from the Tuxpan–Tampamachoco estuarine system of Veracruz, Mexico (Castro-Aguirre et al. 1986; Pérez-Hernández and Torres-Orozco B. 2000) need to be verified. There are several verified records from Mexico (H. Espinosa, pers. comm.): one off the coast of Chetumal Bay, in Othon P. Blanco municipality (CNPE-IBUNAM 3210), and two records from Campeche, one off the coast of Champotón (CNPE-IBUNAM 4593) and one in Tenabo (CNPE-IBUNAM 4610). Study Material. A total of 175 specimens (25.5–164 mm SL) originally at ANSP, CAS-SU, FMNH, FSU, TU, UF, UMML, UMMZ, and USNM. The specimens came from five general regions: 85 specimens from southern Florida and the Florida Keys (25.5–150 mm SL); 80 specimens from Campeche and Yucatán, Mexico (56.6–88.5 mm SL); 4 specimens from Belize (94.0–147 mm SL); and 2 specimens from Cuba (143– 164 mm SL). Genus Euleptorhamphus Gill, 1859 Oceanic Halfbeaks Euleptorhamphus Gill, 1859b:155 (original description; type species Euleptorhamphus brevoortii Gill = ? E. viridis van Hasselt by original designation). In his original description, Gill (1859b:157) wrote that “the habitat of this species is unknown.” Counts of 22 dorsal and 22 anal fin rays are not adequate evidence to decide between an Atlantic and an Indo-Pacific origin for his specimen. The pectoral ray count of eight makes it more likely that the type specimen of E. brevoortii came from the Indo-Pacific rather than the Atlantic. We conclude that E. brevoortii is best considered a junior synonym of the Indo-Pacific E. viridis.

Diagnosis. Body very elongate, strongly compressed laterally, almost ribbon-shaped. Greatest depth in adult fish 11.3–13.7 times in SL (in young of about 80 mm to 26 times in SL), width of body in its depth 1.7–2.0 times in SL. Vertebrae 70–75, of which 26–29 are caudal. Scales rather large, predorsal scales 49–72. Opercle, suborbital, and interorbital regions of head covered with scales. Scales present on upper jaw. Pectoral branch of lateral line single. Lacrimal canal T-shaped. Superficial head canals absent (Parin and Astakhov 1982). Swim bladder simple, single-chambered (Tibbetts et al. 2007). Analysis of molecular data (Lovejoy et al. 2004:369) shows Euleptorhamphus clustering with Hemiramphus and Oxyporhamphus. Head length 6.5–7.4 times in SL. Upper jaw with triangular projection from anterior margin. Interorbital region concave. Lower jaw very long (2.2–2.7 times in SL). Preorbital shelf absent. Preorbital canal without posterior branch. Jaw teeth arranged in three rows, unicuspid in anterior part of both jaws, tricus-

Euleptorhamphus velox Poey, 1868 Atlantic Ribbon Halfbeak Figures 4.3, 4.4 Tables 4.1, 4.2 Euleptorhamphus velox Poey, 1868:383–384 (original description; Cuba). Jordan and Evermann 1896b:724 (description, synonymy, range). Jordan and Evermann 1896a:321 (listed). Evermann and Kendall 1899:62 (after Lönnberg 1894). Bean 1902:407 (north to Massachusetts). Bean 1903:326–327 (synonymy, description; Newport, RI, and Cape Cod, MA). Kendall 1908:121–122 (off Nantucket, MA, and Newport, RI). Tracy 1910:89 (off Nantucket). Sumner et al. 1913:745 (Newport and Nantucket). Fowler 1919a:10 (Atlantic City, NJ, ANSP 22443). Nichols and Breder 1927:59 (Woods Hole, Nantucket, MA, and Newport, RI). Beebe and Tee-Van 1928:71 (sighted en route to Gonave Island, Haiti). Breder 1929:280, 1932a:20

96

Family Hemiramphidae (left by terns, Bird Key, Tortugas). Longley 1929:289 (Tortugas, FL). Jordan et al. 1930:198 (listed). Beebe and Hollister 1935:212 (seen skipping over the waters, Union Island, Grenadine Islands). Howell Rivero 1938:180 (MCZ 8779 holotype of E. velox). Longley and Hildebrand 1941:30 (Tortugas). Breder 1948:90 (West Indies to Massachusetts). Woods and Kanazawa 1951:630–631 (description, Bermuda; FMNH 48437). Fowler 1952b:87 (Hispaniola). Erdman 1956:324 (in stomach of Coryphaena hippurus, Puerto Rico). Springer and Bullis 1956:58 (Gulf of Mexico). Briggs 1958:264 (range). Hoese 1959:325 (after Springer and Bullis 1956). Breder 1960:73 (Long Island, NY; AMNH 20402). Gordon 1960:31–32, 87 (Rhode Island, after Tracy 1910). Collette 1965:218–221 (eastern Atlantic, differences from E. viridis), fig. 2. Collette 1966:3, 5–7 (meristic data). Moe et al. 1966:17 (Gulf of Mexico). Randall 1968:41 (gliding). Starck 1968:17 (Alligator Reef, FL). De Buen 1972:162 (Antilles). Powell et al. 1972:47 (Gulf of Mexico). Bright and Cashman 1974:342, 348 (description; Flower Gardens, Gulf of Mexico), 378 (fig. 12, photograph). Rose and Hassler 1974:96 (four specimens in stomachs of two Coryphaena hippurus). Fahay 1975:4, 16 (off Savannah, GA). Guitart 1975:251–252 (Cuba, synonymy, description, fig. 186). Smith et al. 1975:5 (Florida middle ground). Shiino 1976:90 (common name Flying Halfbeak). Hoese and Moore 1977:150 (Gulf of Mexico), pl. 118. Pequegnat et al. 1977:11 (Gulf of Mexico). Hardy 1978:128 (development). Collette 1978 (description, range, figure). Vitvitsky 1978:662 (collagen stability). Grier et al. 1980:332 (restricted lobular testis). Gilmore et al. 1981:10 (Indian River area, Florida). Murdy 1983:90 (in key, Texas). Manooch et al. 1984:1517 (food of Coryphaena hippurus). Rodríguez et al. 1984:14 (escribano de Golfo; Cuba). Robins and Ray 1986:102 (description, range), pl. 17. Rodríguez and Valdés 1987:95 (Cuba). Schwartz 1989:340 (North Carolina). Cervigón 1991:219–220 (Venezuela, fig. 165). Able 1992:4 (New Jersey, rare). Boschung 1992:83 (Alabama). Nolf and Stringer 1992:70, pl. 12, fig. 21 (otolith from off Trinidad). Cervigón et al. 1993:337 (northern coast of South America). Hensley and Hensley 1995:814 (eaten by Sooty Terns and Brown Noddies, Dry Tortugas, FL). Smith 1997:389 (description, range). McEachran and Fechhelm 1998:958, figure (Gulf of Mexico). Mejía Aristizábal et al. 1998:30 (Colombian Caribbean oceanic atolls, listed). Carvalho Filho 1999:70 (Brazil). Smith-Vaniz et al. 1999:175 (Bermuda records), fig. 100a. Claro and Parenti 2001:43 (Cuba, listed). Collette 2003b:1143 (range, figure). Menezes 2003:68 (Brazil). Collette 2004:3 (synonymy). Parenti and Grier 2004:336 (restricted lobular testis). Collette 2005:938–939 (early life history, figure). Schwartz 2006:66 (North Carolina records). Fahay 2007: 776, 788–789 (early stages, figs. A–D). Casazza and Ross 2008:351 (associated with Sargassum, Gulf Stream off North Carolina). McEachran 2009:1286 (Gulf of Mexico). Pepperell 2010: 234 (color figure).

Collette 2010:34 (reproduction and development). Kells and Carpenter 2011:150–151 (description, color painting). Snyder and Burgess 2016:100 (listed, Florida). Starck et al. 2017:91 (Florida Keys). Euleptorhamphus longirostris (not of Valenciennes in Cuvier and Valenciennes 1847). Putnam 1870b:236–239 (Nantucket, MA; E. brevoortii and E. velox synonyms). Gill 1873:31 (Florida to Cape Cod). Goode 1879:16 (Florida). Jordan 1887b:569 (West Indies). Hemirhamphus macrorhynchus (not of Valenciennes in Cuvier and Valenciennes 1847). Cope 1870:121 (Atlantic City, NJ). Hemiramphus longirostris (not of Valenciennes in Cuvier and Valenciennes 1847). Lönnberg 1894:119 (Coronado Beach, FL). Miranda Ribeiro 1961b:10–11 (description, Salvador, Brazil), fig. 5. Euleptorhamphus viridis (not of van Hasselt 1824). Fowler 1940a:12 (Ventnor, NJ; ANSP 86213). Parin 1964:186 (in part, E. velox considered a synonym of Indo-Pacific E. viridis). Duarte-Bello and Buesa 1973:80 (Cuba). Erdman 1976:23, 1977:159 (ripe females, March–April, northeastern Caribbean). Gilmore 1977:132 (Indian River area, Florida). Fedoryako 1980:581 (Sargasso Sea). Manooch et al. 1985:1210 (in stomach of a Euthynnus alletteratus). Euleptorhamphus brevoorti. Fowler 1920:155 (Atlantic City, NJ; ANSP 22443). Weed 1933:54–55 (generic description; type species of Euleptorhamphus). Fowler 1944:95 (Courtown Key, Caribbean Sea, Colombia, ANSP 72786), fig. 105. Fowler 1952a:112 (Atlantic and Cape May counties, New Jersey). Fowler 1953:52 (Courtown Key, after Fowler 1944).

Types. Euleptorhamphus velox Poey, 1868. Holotype MCZ 8779 (248 mm SL); Cuba; Poey no. 422. Dorsal fin rays 22; anal fin rays 22; pectoral fin rays 7½–7½. Gill rakers on first arch 6 + 22 = 28. Lower jaw length 96.5 mm; pectoral fin length 61.1 mm. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Flying Halfbeak (English); however, we choose to call it Atlantic Ribbon Halfbeak to parallel the name Pacific Ribbon Halfbeak for Euleptorhamphus viridis. Page et al. (2013) lists agujeta voladora (Spanish, Mexico); it is known as as escribano volador in Cuba (Claro and Parenti 2001). Diagnosis. Euleptorhamphus velox is very closely related to E. viridis and differs mainly in having either seven pectoral rays or seven plus a tiny splint, whereas E. viridis has eight or eight plus a tiny splint. Euleptorhamphus velox averages about one dorsal and

Figure 4.3. Atlantic Ribbon Halfbeak, Euleptorhamphus velox. USNM 188747, 135 mm SL. Collected from R/V Silver Bay, off North Carolina, U.S.A., 24 July 1960. Illustration by M. H. Carrington.

97

Fishes of the Western North Atlantic and Caribbean Sea to Recife, Brazil (CAS-SU 51794), and 25°30′05″S, 45°26′08″W (MZUSP 67710) (Fig. 4.4). Study Material. A total of 79 specimens (27.5–310 mm SL) from the western Atlantic, including the holotype (MCZ 8779), originally at ANSP, CAS-SU, CM, FMNH, MCZ, MZUSP, ROM, TU, UF, UMML, USNM, and ZMK. The specimens come from five general regions: 24 specimens (27.5–277 mm SL) from the east coast of the United States; 3 specimens (128–148 mm SL) from Bermuda; 18 specimens (51.5–280 mm SL) from the Gulf of Mexico; 23 specimens (62.5–266 mm SL) from the Caribbean Sea and Atlantic east of the Leeward Islands; and 12 specimens (62.6–281 mm SL) from South America. Comparative material from the eastern Atlantic includes six specimens (53.4–255 mm SL) (see Collette 1965). Genus Hemiramphus Cuvier, 1816 Ballyhoos Hemiramphus Cuvier, 1816:186 (type species Esox brasiliensis Linnaeus, 1758 by subsequent designation of Gill 1863). Farhians Whitley, 1930:250 (type species by original designation Hemiramphus commersoni Cuvier, 1829 = Hemiramphus far (Forsskål). Ardeapiscis Whitley, 1931:314 (type species by original designation Hemiramphus welsbyi Ogilby, 1908 = Hemiramphus robustus Günther).

Figure 4.4. Distribution of Atlantic Ribbon Halfbeak, Euleptorhamphus velox, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

one anal ray less than E. viridis and has slightly more gill rakers. Description. Dorsal fin rays 21–24; anal fin rays 20–24; pectoral fin rays 7 large rays plus a very small one. Vertebrae 45–46 precaudal + 26–27 caudal = 71–73 total. Gill rakers on first arch 6–9 + 20–27 = 26–35; gill rakers on second arch 2–6 + 15–20 = 17–25. Size. The maximum observed size for Euleptorhamphus velox is 310 mm SL, 415 mm TL (ROM 23338, Bahamas).

Diagnosis. The characteristic features of Hemiramphus, distinguishing it from all other marine halfbeaks, are the absence of scales on the upper jaw, cellular swim bladder (the same swim bladder structure is present in one species of Oxyporhamphus), and significant posterior displacement of the anal fin relative to a vertical with the origin of the dorsal fin. Juveniles of Hemiramphus (and Oxyporhamphus) have colored bands on the body. Analysis of molecular data (Lovejoy et al. 2004:369) clusters Hemiramphus with Oxyporhamphus and Euleptorhamphus. This genus differs also from Hyporhamphus in the absence of the preorbital shelf and presence of two pectoral branches of the lateral line (from Hyporhamphus sensu stricto also in the presence of a posterior branch to the preorbital canal); from Rhynchorhamphus in the shape of the upper jaw, the structure of the preorbital canal, the nonfimbriate nasal papilla, and the lower number of gill rakers; from Euleptorhamphus in the relatively short body, the absence of teeth in the mouth, fewer vertebrae and rays in the unpaired fins, and more rays in the pectoral fins; and from Oxyporhamphus in the presence of a triangular upper jaw, shorter pectoral fins, and a series of other characters. Body rather deep, laterally compressed, its crosssectional diameter elongate-oval. Greatest body

Biology Predators. Eaten by oceanic fishes and birds. There are several reports of Euleptorhamphus velox remains near nests of Sooty Terns and Brown Noddies at the Bird Key Rookery in the Tortugas (Breder 1929, 1932a; Longley 1929; Hensley and Hensley 1995). Dolphinfish, Coryphaena hippurus, have been reported to eat E. velox, in low frequencies, in several areas in the western Atlantic: Puerto Rico (Erdman 1956), off Hatteras, NC (Rose and Hassler 1974), and the southeastern United States (Manooch et al. 1984). One specimen was also found in the stomach of a Little Tunny, Euthynnus alletteratus, from the southeastern United States (Manooch et al. 1985). Range. Atlantic Ocean, in the western Atlantic from South Dartmouth, MA (MCZ 46567), and Newport, RI (USNM 15648), south through the Gulf of Mexico 98

Family Hemiramphidae depth 5.9–7.0 times in SL, width of body in its depth 1.3–2.05 times in SL. Vertebrae 50–59, of which 30–41 are precaudal and 16–19 are caudal. Scales medium-sized, predorsal scales 32–40. Pectoral branch of the lateral line double, with the anterior part shorter and proceeding along the edge of the opercle (at the anterior margin of the cleithrum); the posterior branches under the posterior edge of the pectoral fin and extends to its base. Swim bladder vesicular as in Oxyporhamphus convexus (Tibbetts et al. 2007). Head length 3.7–4.7 times in SL. Upper jaw blunt, length much less than width. Interorbital region flat. Lower jaw long (3.0–4.5 times in SL). Preorbital shelf absent. Preorbital canal with posterior branch. Jaw teeth distributed in two to four rows, all tricuspid. No teeth in jaws. Pharyngeal teeth mostly sharp, tricuspid, but in the posterior rows on the lower pharyngeal and third upper pharyngeal bones with thickened or recurved edges. Gill rakers on first arch 25–48, on second arch 20–37. Nasal papilla not fimbriate. Opercle, suborbital, and interorbital parts of head covered with scales. No scales on upper jaw. Dorsal fin rays 11–15; anal fin rays 9–14; pectoral fin rays 10–13. Pectoral fins of medium length (4.5–6.8 times in SL). Pelvic fins short (8.8–10.7 times in SL), located significantly closer to the base of the caudal fin than to the origin of the pectoral fins (P1–P2/P2–C = 1.2–1.6). Origin of anal fin located under fourth to eighth ray of dorsal. Caudal fin deeply forked, with enlarged lower lobe. Dorsal fin with or without well-developed anterior lobe. Anterior rays of dorsal and anal fins covered with scales. Juveniles have transverse bands of dark pigment on the body and to some degree, pigmented pelvic, dorsal, anal, and caudal fins. Species. Ten species belong to the genus Hemiramphus (Collette 2004): H. archipelagicus, H. balao, H. bermudensis, H. brasiliensis, H. depauperatus, H. far, H. lutkei, H. marginatus, H. robustus, and H. saltator. Range. Hemiramphus is a widespread marine genus (Collette 2004). Three species—H. balao, H. brasiliensis, and H. bermudensis—are endemic to the Atlantic Ocean (Collette 1962, 1965, 2003b) and one species, H. saltator, is found along the Pacific shores of the Americas. The remaining six species occur in the Indo-West Pacific region (see Parin et al. 1980).

and Evermann 1896b:723 (synonymy, description, range). Jordan and Evermann 1896a:321 (listed). Evermann and Kendall 1899:62 (Florida). Jordan 1919:397–398 (description, Cuba, CAS-SU 23586), figure. Fowler 1944:94–95 (synonymy, description; Courtown Keys, Colombia; ANSP 72787). Fowler 1953:52 (Courtown Keys, Colombia, after Fowler 1944). Anderson and Gehringer 1959:44, 48 (30°N, 77°W). Collette 1962:434–436 (comparison with H. bermudensis and H. brasiliensis), fig. 1. Cervigón 1965:26 (Venezuela). Collette 1965:221–224 (synonymy, description, range), fig. 3. Caldwell 1966:32 (Jamaica). Randall 1967:684–686 (food: fishes, pteropods, polychaetes). Böhlke and Chaplin 1968:125 (Bahamas), figure. Randall 1968:42 (Caribbean), fig. 43 (photograph). Starck 1968:17 (Alligator Reef, FL). Dragovich 1969:12 (eaten by Katsuwonus pelamis). de Sylva 1970:55 (Biscayne Bay, FL). Mago Leccia 1970:89 (Venezuela). Dooley 1972:9–10 (member of Sargassum complex). Berkeley et al. 1975 (fishery and biology, southeastern Florida). Erdman 1976:23, 1977:159 (ovaries subripe, January–April). Shiino 1976:90 (common name Balao). Gilmore 1977:132 (Indian River vicinity, Florida). Hoese and Moore 1977:150 (description, Gulf of Mexico), pl. 119. Berkeley and Houde 1978:624–644 (biology, Florida). Collette 1978 (description, range), figure. Figueiredo and Menezes 1978:57 (southeastern Brazil), fig. 97. Vitvitsky 1978:662 (collagen thermostability). Román 1979:83 (Venezuela, marao fosforo), figure. Burgess et al. 1980:77 (juvenile, 34°44.5′N, 75°22.5′W). Gilmore et al. 1981:10 (Indian River area, Florida). Lucena and Lucena 1981:40 (agulha, Porto Belo, SC, Brazil; southern range extension). Sacchi et al. 1981:2 (balaou bleu, French Antilles). Acero P. et al. 1984:41 (Caribbean Colombia). Collins and Finucane 1984:47 (ichthyoplankton, Florida Everglades). Collette et al. 1984:346 (fig. 180, juvenile pigmentation), 357 (fig. 183, pelvic fin pigmentation). Robins and Ray 1986:102 (description, range), pl. 17. Gilmore 1987:129 (seagrass communities). Dennis and Bright 1988:4 (above East Flower Garden bank, northwestern Gulf of Mexico). Ditty et al. 1988:813 (larvae, northern Gulf of Mexico). Garzón F. 1989:155 (Colombia). Lopes 1989:211 (Brazil). Schwartz 1989:340 (North Carolina). Cervigón et al. 1993:338 (northern coast of South America). Humann 1994:58–59 (description, range; color photograph). Begossi and Figueiredo 1995:714 (local name in southern Brazil panaguaiú; consumed). Hensley and Hensley 1995:814 (eaten by Sooty Terns and Brown Noddies, Dry Tortugas, FL). McBride et al. 1996 (Florida bait fishery). Debelius 1997:58–59 (description, color underwater photograph). Smith 1997:386 (description, range). McEachran and Fechhelm 1998:959, figure (Gulf of Mexico). Mejía Aristizábal et al. 1998:30 (Colombian Caribbean oceanic atolls, listed). Carvalho Filho 1999:70 (Brazil). Smith-Vaniz et al. 1999:175–176 (Bermuda records), fig. 100b. Schmitter-Soto et al. 2000:154 (Caribbean Mexico). Claro and Parenti 2001:43 (Cuba, listed). McBride 2001 (landings in south Florida lampara net fishery). Humann and DeLoach 2002:62–63 (description, range; color photograph). Collette 2003b:1138 (description, range, figure). McBride and Styer 2003:24–26 (second most common species in south Florida lampara net fishery). McBride and Thurman 2003 (reproductive biology). McBride et al. 2003 (spawning cycles in south Florida). Menezes 2003:68 (Brazil). Collette 2004:3–4 (synonymy). Lovejoy et al. 2004:369, 376 (molecular phylogeny). Collette 2005:940–941 (early life history, figs. A–G). Schwartz 2006:66 (North Carolina records). Walsh et al. 2006:276 (juveniles collected off Georgia coast). Fahay 2007:776, 790–791 (early stages, figs. A–C). Tibbetts et al. 2007:809, 812 (vesicular swim bladder). Casazza and Ross 2008:351 (associated with

Hemiramphus balao Lesueur, 1821 Balao Figures 4.5, 4.6 Tables 4.1, 4.2, 4.4, 4.5 Hemiramphus balao Lesueur, 1821:136–137 (original description; Lesser Antilles, Guadeloupe, Martinique). Meek and Goss 1884:221 (in key). Jordan 1887b:568 (West Indies). Jordan

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Figure 4.5. Balao, Hemiramphus balao. A. ZMK P.341511, 220 mm SL. Collected near the Isle of Rolas, Sao Tomé, from the R/V Galathea, Sta. 49. B. USNM 187566, 117 mm SL. Dipnetted at night from M/V Delaware off North Carolina, U.S.A., 19 July 1958. C. USNM 200592, 53.7 mm SL. Dipnetted at night by R. H. Gibbs, Jr. off Cape Hatteras, North Carolina, U.S.A., from M/V Delaware, 10 October 1957. Illustrations by M. H. Carrington. Sargassum, Gulf Stream off North Carolina). Headley et al. 2009:672 (table 1, eaten by Blackfin Tuna, Thunnus atlanticus, in Tobago). McEachran 2009:1286 (Gulf of Mexico). Collette 2010:35 (reproduction and development). Pepperell 2010:234 (bait for sports fishers). Kells and Carpenter 2011:150–151 (description, color painting). Smith-Vaniz and Jelks 2014:32 (St. Croix, U.S. Virgin Islands records). Snyder and Burgess 2016:100 (listed, Florida). Starck et al. 2017:91 (Florida Keys). Hemiramphus vittatus Valenciennes, 1843:70–71 (original description; Lancerotte and Arecife, Canary Islands). Hemiramphus Pleii Valenciennes in Cuvier and Valenciennes, 1847:21 (original description; Cuba, Martinique). Collette et al. 1997:18 (types). Hemirhamphus macrochirus Poey, 1860:299 (original description; Cuba). Poey 1868:383 (description; Cuba). Collette et al. 1992:9 (type). Hemiramphus guineënsis Bleeker, 1863:119–120 (original description; Boutry, Gold Coast), pl. 25, fig. 2. Hemirhamphus balao. Henshall 1895:212 (Key West, FL). Evermann and Kendall 1899:62 (Florida records). Jordan et al. 1930:199 (listed). Cervigón 1966:220 (description, Venezuela). Duarte-Bello and Buesa 1973:81 (synonymy, Cuba). Fischthal and Nasir 1974:75 (trematode Monorchiaponurus hemirhamphi

described from H. balao small intestine; Los Testigos Island, Venezuela). Belyanina 1975:140 (ichthyoplankton; Gulf of Mexico). Guitart 1975:253–254 (Cuba, synonymy, description, fig. 189). Fink 1981:171 (jaw and pharyngeal teeth with small area of unmineralized collagen at base). Rodríguez et al. 1984:14 (escribano balao; Cuba). Rodríguez and Valdés 1987:95 (escribano balao, Cuba). Cervigón 1991:221 (Venezuela). Lara-Domínguez et al. 1993:364 (Gulf of Mexico).

Misidentifications. As Hemiramphus brasiliensis: Miranda Ribeiro (1961a:8), MNRJ 9246 Angra dos Reis, Brazil; Fowler (1944:71–72, 82, 1953:52), ANSP 74153, 74172, Serranilla Bank and Roncador Reef; Fowler (1928:468), ANSP 79291, St. Lucia. Types. Hemiramphus vittatus Valenciennes, 1843. Holotype MNHN B.2183 (284 mm SL), Canary Islands; Webb and Berthelot; half a skin mounted on a board. Dorsal fin rays 14; anal fin rays 10. Lower jaw length at least 70 mm SL; pectoral fin length about 47 mm SL. 100

Family Hemiramphidae Hemiramphus pleii Valenciennes in Cuvier and Valenciennes, 1847. Lectotype (designated by Collette 1965:223) MNHN B.1050 (180 mm SL); Martinique; Choris. Dorsal fin rays 13; anal fin rays 11. Gill rakers on first arch 9 + 25 = 34; gill rakers on second arch 4 + 21 = 25. Lower jaw length 50.1 mm; pectoral fin length 35.7 mm. Paralectotypes MNHN B.1051 (180 mm SL) and MNHN B.1048 (173 mm SL). Hemirhamphus macrochirus Poey, 1860. Lectotype (designated by Collette 1965: 223) MCZ 693 (186 mm SL); Cuba; Poey no. 199. Dorsal fin rays 13; anal fin rays 12. Gill rakers on first arch 9 + 25 = 34. Pectoral fin length 37.5 mm. Paralectotype USNM 4748 (242 mm SL). Hemiramphus guineensis Bleeker, 1863. Lectotype RMNH 2823 (232 mm SL); Boutry, Gold Coast; designated herein. Dorsal fin rays 13; anal fin rays 11. Gill rakers on first arch 10 + 27 = 37; gill rakers on second arch 6 + 23 = 29. Lower jaw length 67.1 mm; pectoral fin length 45.7 mm. Paralectotype RMNH 2823 (256 mm SL). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico

(Page et al. 2013) lists Balao (English) and agujeta balao (Spanish, Mexico); it is known as escribano balao in Cuba (Claro and Parenti 2001). Diagnosis. Hemiramphus balao differs from H. brasiliensis and H. bermudensis primarily in its long pectoral fin, which, when extended forward, reaches beyond the anterior margin of the nasal fossa. In this respect, H. balao is very similar to H. saltator from the eastern Pacific, but differs from that species in having fewer total gill rakers (H. saltator 38–46, x̅ 40.8). Adult H. balao differ from H. brasiliensis and H. bermudensis in having a bluish tip to the upper caudal lobe in life instead of a reddish orange tip. Description. Upper jaw short, triangular; snout scaleless; preorbital ridge (bony ridge under nostril) absent. Dorsal fin rays 11–15, usually 13 or 14; anal fin rays 10–13, usually 11 or 12; pectoral fins long, reaching beyond anterior margin of nasal pit when folded forward, and with 10–12 rays; caudal fin deeply forked, lower lobe much longer than upper. Vertebrae 37–39 precaudal + 16–17 caudal = 54–56 total. Gill rakers on first arch 7–10 + 22–29 = 31–38 (Table 4.4); gill rakers on second arch 4–7 +19–24 = 25–30 (Table 4.5).

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Fishes of the Western North Atlantic Color. Dark bluish above, silvery white below. Beak dark with fleshy red tip; upper and lower lobes of caudal fin bluish violet. Size. The maximum observed size for Hemiramphus balao is 40 cm TL, about 28 cm SL, commonly to 35 cm TL.

Food. Unlike Hemiramphus brasiliensis, H. balao is largely a carnivore. Neither Randall (1967) nor Berkeley and Houde (1978) found seagrasses in stomach contents. In southeastern Florida, Berkeley and Houde (1978) found copepods, decapods, siphonophores, and polychaetes to be the most common food items. Randall (1967) reported that the diet in the Virgin Islands consisted of 39.1% fishes (such as Jenkinsia), 31.4% pteropods, 8.9% polychaetes, 4.7% crab larvae, 2.9% shrimp larvae, and 1.2% copepods. Reproduction. Cyclic patterns of gonadosomatic indices indicate that Hemiramphus balao spawns during spring and summer months in south Florida (McBride et al. 2003). Hydration of oocytes began in the morning and spawning occurred at dusk the same day. All mature females spawned daily in June (McBride and Thurman 2003). Batch fecundity is higher than in H. brasiliensis, an average of 3,734 rather than 1,164 hydrated oocytes in a 100 g female. Spawning is in deeper, more offshore waters than for H. brasiliensis. A large female (USNM 294082, 157 mm SL) collected from the Caribbean Sea had 100 eggs in the left ovary (0.95–1.25 mm in diameter, x̅ 1.14 mm for 20 eggs) and 100 eggs in the right ovary (1.00–1.25 mm in diameter, x̅ 1.15 mm for 20 eggs). Range. Atlantic Ocean, in the western Atlantic from off New York, U.S.A., south throughout the Gulf of Mexico and the Caribbean Sea to Santos, Brazil (Collette 2003b), and in the eastern Atlantic from the Canary Islands south to Luanda, Angola (Collette 1965) (Fig. 4.6). Study Material. A total of 297 specimens from the western Atlantic (13.7–258 mm SL), including the types of Hemiramphus pleii (MNHN B.1050, B.1051, B.1048) and H. macrochirus (MCZ 693); originally at ANSP, BMNH, FMNH, GCRL, MACN, MCZ, MNHN, MNRJ, MZUSP, SMF, SU, TU, UMML, USNM, and ZMK. The specimens come from seven general regions: 133 specimens from the east coast of the United States (13.7–171 mm SL); 8 specimens from Bermuda (34.2–95.5 mm SL); 53 specimens from the Bahamas (22.5–219 mm SL); 13 specimens from the Gulf of Mexico (14.6–174 mm SL); 51 specimens from the Antilles (41.9–207 mm SL); 22 specimens from the Caribbean coast of Central America (26.5–222 mm SL); and 17 specimens from South America (119–258 mm SL). Comparative material from the eastern Atlantic: 22 specimens (175–284 mm SL), including the types of Hemiramphus vittatus (MNHN B.2183) and H. guineensis (RMNH 2823). This total includes 14 specimens (206–284 mm SL) previously reported (Collette 1965), plus the following: UMML 21717 (4, 177–221

Biology Predators. Reported from stomachs of Skipjack Tuna, Katsuwonus pelamis (Dragovich 1969), and Blackfin Tuna, Thunnus atlanticus, in Tobago (Headley et al. 2009). There are also records of Hemiramphus sp. (some of which are probably H. balao) from several other fishes: Sailfish, Istiophorus platypterus, by Jolley (1977); Little Tunny, Euthynnus alletteratus, by Dragovich (1969); and Blue Marlin, Makaira nigricans, by Erdman (1962). Both species of Hemiramphus are eaten by Sooty Terns and Brown Noddies in the Dry Tortugas (Hensley and Hensley 1995). Both species are important bait in the sport fishery for billfishes, Dolphinfish, and King Mackerel off the coast of Florida (Berkeley and Houde 1978; Pepperell 2010). Parasites. One species of digenetic trematode, Monorchiaponurus hemirhamphi was described from the small intestine of Hemiramphus balao from Los Testigos Island, Venezuela, by Fischthal and Nasir (1974).

Figure 4.6. Distribution of Balao, Hemiramphus balao, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

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Family Hemiramphidae another halfbeak endemic to Bermuda, has been described and thus a more appropriate name for Hemiramphus bermudensis is Bermuda Ballyhoo. Diagnosis. Hemiramphus bermudensis differs from H. brasiliensis primarily in having more gill rakers. Small juveniles of H. bermudensis have fewer gill rakers (35 and 34, respectively, in specimens 41.5 and 32.1 mm SL) than adults, but more than juveniles of H. brasiliensis (29–31 gill rakers in specimens 32–50 mm SL). From H. balao, the third Atlantic species of the genus, H. bermudensis differs primarily in its shorter pectoral fin, more anal rays (usually 13 rather than usually 11 or 12), and more gill rakers (37–45 rather than 31–37). In number of gill rakers, H. bermudensis is close to H. saltator Gilbert and Starks, of the eastern Pacific (gill rakers 35–44), but differs from it by having a much shorter pectoral fin (5.6–6.7 times in SL rather than 4.4–5.1 in specimens over 150 mm SL). Of the four species of Hemiramphus mentioned, H. bermudensis has the least body depth, differing most strongly from H. brasiliensis in this regard. Description. Upper jaw short, triangular; snout scaleless; preorbital ridge (bony ridge under nostril) absent. Pectoral fins short, not reaching to nasal pit when folded forward and caudal fin deeply forked, lower lobe much longer than upper. Dorsal fin rays 13–15, usually 14; anal fin rays 12–14, usually 13; pectoral fin rays 10–12, usually 11. Vertebrae 36–37 precaudal + 17–19 caudal = 53–56, usually 54 total. Gill rakers on first arch 9–13 + 26–33 = 37–44; gill rakers on second arch 5–10 + 21–28 = 28–35. Pectoral insertion to pelvic insertion 2.1–2.4 times in SL; pelvic insertion to caudal base 2.8–3.0 times in SL; lower jaw length 3.1–3.6 times in SL; pectoral length 5.2–5.7 times in SL in specimens 100–149 mm SL, 5.6–5.9 times in SL in specimens 150–221 mm SL, and 6.0–6.7 times in SL in specimens 226–300 mm SL; body depth at pectoral origin 7.3–8.5 times in SL; depth at pectoral tip 6.0–8.7 times in SL; depth at pelvic origin 8.2–9.1 times in SL in specimens 100– 122 mm SL and 6.5–8.2 times in SL in specimens 137–300 mm SL. Color. Dark bluish above, silvery white below. Beak black with bright red fleshy tip; caudal fin lobe reddish orange in adults. Size. The maximum observed size for Hemiramphus bermudensis is 310 mm SL, about 440 mm TL, commonly to 350 mm TL.

mm SL), Pillsbury 274, Annobón Island, 19 May 1965; UMML 16658 (1, 175 mm SL), Pillsbury 19, 05°12′N to 04°58′N, 00°12′E, 26–27 May 1964; MNHN 84-1089, 84-1090 (2, 273–275 mm SL), Talisman Expedition, Porto Praia, Cape Verde Islands; ZMK uncat. (1, 246 mm SL) Atlantide Expedition, Sta. 44, 10°22′N, 16°22′W, 17 December 1945. Hemiramphus bermudensis Collette, 1962 Bermuda Ballyhoo Figures 4.7, 4.8 Tables 4.1, 4.2, 4.4, 4.5 Hemiramphus braziliensis (not of Linnaeus 1758). Jones 1859:103 (listed as common). Hemirhamphus Pleii (not of Valenciennes in Cuvier and Valenciennes 1847). Goode 1876:11 (taken in large numbers in Bermuda bays), 64 (“common gar-fish” seined in large quantities for the market, two sizes sizes, about 20 inches and about 8 inches [the latter were probably Hyporhamphus collettei]). Goode 1877:292 (listed). Bean 1880:103 (duplicates distributed to other museums). Hemiramphus pleii (not of Valenciennes in Cuvier and Valenciennes 1847). Jones 1876:135 (very common). Hemiramphus brasiliensis (not of Linnaeus 1758). Barbour 1905:114 (Bermuda, MCZ 8774). Bean 1906:36 (Bermuda, FMNH 50513). Beebe and Tee-Van 1933:62 (common in Bermuda). Saunders 1959:376 (none of 45 specimens infected with Haemogregarina bigemina). Hemirhamphus sp. Richardson 1905:265 (Irona nana on gills of specimens from Harrington Sound, Bermuda). Hemirhamphus brasiliensis (not of Linnaeus 1785). Rees 1970:217 (no helminths on six Bermuda specimens). Hemiramphus bermudensis Collette, 1962:432–437 (original description; Bermuda), fig. 1. Collette 1965:226 (allopatric to H. brasiliensis at Bermuda). Collette 1978 (description), figure. Burnett-Herkes 1986:589 (description after Collette 1962), fig. 200. Bruce and Bowman 1989:19 (isopod Glossobius hemiramphi found on one Bermuda specimen. Collette et al. 1992:8 (types). Smith 1997:386 (description, range). Smith-Vaniz et al. 1999:176–177 (Bermuda records), fig. 100c. Collette 2003b:1139 (description, range, figure). Collette 2004:4 (synonymy). Collette 2005:942–943 (early life history, fig. A–B). Tibbetts et al. 2007:809 (vesicular swim bladder).

Types. Hemiramphus bermudensis Collette, 1962. Holotype FMNH 48115 (272 mm SL); Bermuda, St. Georges Island, Whalebone Bay; Bermuda Oceanographic Expedition, 22 June 1948. Dorsal fin rays 15; anal fin rays 12; pectoral fin rays 11. Gill rakers on first arch 11 + 30 = 41; gill rakers on second arch 7 + 24 = 31. Lower jaw length 79.0 mm; pectoral fin length 42.5 mm. Paratypes 39 specimens (109–310 mm SL) at USNM, FMNH, MCZ, ZMK, and UMMZ (see Collette 1962:436). Common Names. Smith-Vaniz et al. (1999) referred to this species as the Bermuda Halfbeak when it was the only known Bermuda endemic of the Hemiramphidae. However, since then, Hyporhamphus collettei,

Biology Parasites. An isopod identified as Irona nana Schioedte and Meinert was reported from the gills 103

Fishes of the Western North Atlantic

Figure 4.7. Bermuda Ballyhoo, Hemiramphus bermudensis. A. Holotype FMNH 48115, 272 mm SL. Whalebone Bay, Bermuda, 1948. B. Paratype, USNM 187565, 120 mm SL. Collected by R. H. Gibbs, Jr., St. George Harbor, Bermuda, 4 February 1958. Illustrations by M. H. Carrington.

of specimens from Harrington Sound, Bermuda (Richardson 1905:265). A pair of the isopod Glossobius hemiramphi was found in the mouth of a 139 mm SL specimen from Ferry Reach, Bermuda (Bruce and Bowman 1989). The female was on top of the tongue and extended back to the gill rakers on the second gill arch on the right side (B. B. Collette, pers. observ.). The small male was to the side of the female anteriorly in the mouth. Saunders (1959) found that none of 45 specimens were infected with Haemogregarina bigemina and Rees (1970) did not find any helminths on 6 specimens. Food. Adults observed feeding on floating seagrass in Harrington Sound, Bermuda (B. B. Collette, pers. observ.). Range. Endemic to the waters around Bermuda, where it is allopatric to the wide-ranging Hemiramphus brasiliensis (Fig. 4.8). Study Material. A total of 67 specimens from Bermuda (19.2–310 mm SL), including the holotype (FMNH 48115), and 39 paratypes (109–310 mm SL) (see Collette 1962). Also five other adults (230–289 mm SL), in BMNH and FMNH, and seven juveniles (19.2–41.5 mm SL) in FMNH, UMMZ, and YPM, and 15 recently collected specimens (USNM 292592, 135–181 mm SL).

Figur e 4.8. Dist r ibut ion of Ber muda Ba l lyhoo, Hemiramphus ber mudensis, and Redtail Ballyhoo, Hemiramphus brasiliensis, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

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Family Hemiramphidae Hemiramphus brasiliensis (Linnaeus, 1758) Redtail Ballyhoo Figures 4.8, 4.9 Tables 4.1, 4.2, 4.4, 4.5

Indies 81% by volume of seagrasses Cymodocea manatorum and Thalassia testudinium, 19% Jenkinsia), 754 (eaten by Scomberomorus regalis). Starck and Davis 1967:318 (abundant day and night, Alligator Reef, Florida Keys). Böhlke and Chaplin 1968:124 (description; Bahamas), figure. Randall 1968:141–142 (description), fig. 42. Starck 1968:17 (Alligator Reef, FL). de Sylva 1970:55 (Biscayne Bay area, Florida). Mago Leccia 1970:89 (listed; Venezuela). Iversen et al. 1971:84 (spore of myxosporidium Myxidium sp. in gall bladder; southern Florida). De Buen 1972:162 (Antilles). Moe 1972:9–10 (southern Florida). Powell et al. 1972:47 (seven collections, FSBC). Roux 1973:71 (description, Brazil). Brownell and Guzmán 1974:128 (abundant, Isla de Aves, Venezuela). Hardy and Johnson 1974:242–243 (description of larvae and juveniles), fig. 1. Rose and Hassler 1974:96 (in stomachs of Coryphaena hippurus, North Carolina). Berkeley et al. 1975 (fishery, biology, southeastern Florida). Fahay 1975:4–5, 17, 19 (larvae and juveniles, South Atlantic Bight). Sawyer et al. 1975:660 (marine leech Malmiana philotherma, Puerto Rico). Smith et al. 1975:5 (Florida middle ground). Davies and Bortone 1976:250 (food of billfishes, Gulf of Mexico). Erdman 1976:23, 1977:159 (northeastern Caribbean). Shiino 1976:90 (common names, Ballyhoo). Gilmore 1977:132 (Indian River area, Florida). Hoese and Moore 1977:150 (description, Gulf of Mexico), pl. 120. Valdés-Muñoz and Silva Lee 1977:5 (food 100% plants, Cuba). Wallace and Sawyer 1977 (marine leech Malmiana philotherma, Puerto Rico). Castro-Aguirre 1978:58 (Veracruz, Campeche, and Yucatán, Mexico). Collette 1978 (description, range), figure. Figueiredo and Menezes 1978:56–57 (southeastern Brazil), fig. 96. Hardy 1978:129–132 (description, development), figs. 67–69 (development). Terrero and Bonnelly de Calventi 1978:7 (Dominican Republic). de Sylva and Higman 1979:153 (occasionally ciguateric). Hastings 1979:7 (St. Andrew, FL). Román 1979:83 (Venezuela), figure. Grier et al. 1980:332 (restricted lobular testis). Modde and Ross 1980:914 (Gulf of Mexico). Gilmore et al. 1981:10 (Indian River area, Florida). Lucena and Lucena 1981:40 (agulha-preta, Porto Belo, SC, Brazil; southern range extension). Sacchi et al. 1981:2 (balaou queue jaune; present year-round, French Antilles). Cooper 1982:151–152 (used as bait for Scomberomorus regalis, Jamaica). Flores-Coto and Alvarez Cadena 1982:74 (ichthyoplankton, Laguna de Terminos, Campeche, Mexico). Schwartz et al. 1982:27 (13.7 to 31.4 °C, 7.0‰ to 32.0‰ salinity; Cape Fear Estuary, NC). Fahay 1983:210–211 (larvae, after Hardy and Johnson 1974). Fernholm and Wheeler 1983:228– 229 (types of Esox brasiliensis). Saloman and Naughton 1983 (10% occurrence in stomachs of Scomberomorus cavalla off southeastern United States). Acero P. et al. 1984:41 (Caribbean Colombia). Collins and Finucane 1984:47 (ichthyoplankton, Florida Everglades area). Manooch et al. 1984:1517 (in stomachs of Coryphaena hippurus). Naughton and Saloman 1984:15 (in stomachs of Pomatomus saltatrix). Salomon and Naughton 1984:11, 20 (in stomachs of Caranx hippos, Gulf of Mexico). Vasconcelos Filho et al. 1984:241–243, table 4 (diet, Brazil). Manooch et al. 1985:1212 (in stomachs of two Euthynnus alletteratus). Williams and Williams 1985 (isopod Glossobius hemiramphi described from mouths of H. brasiliensis, Puerto Rico). Robins and Ray 1986:102 (description, range), pl. 17. Acero P. and Garzón F. 1987:90 (Colombia). Gilmore 1987:129 (seagrass communities). Paiva Filho and Toscano 1987:157 (São Paulo). Ditty et al. 1988:813 (larvae, northern Gulf of Mexico). Bruce and Bowman 1989:19 (isopod Glossobius hemiramphi). Schwartz 1989:340 (North Carolina). Sogard et al. 1989a:58, 1989b:195 (Florida Bay). McAllister 1990:127 (off Nova Scotia). Able 1992:4 (New Jersey, rare). Boschung

Esox brasiliensis Linnaeus, 1758:314 (original description after Browne; Jamaica). Walbaum 1792:92–93 (description). Hemiramphus brasiliensis. Cuvier 1817:286 (description of genus). Schinz 1822:313 (after Cuvier 1817). Castelnau 1855:52 (description, Rio de Janeiro to Bahia, Brazil, good food fish). Gray 1859:11 (Brazil). Lönnberg 1894:119 (Key West, FL). Jordan and Evermann 1896b:722–723 (synonymy, description, range). Jordan and Evermann 1896a:321 (listed). Jordan and Rutter 1897:97 (Jamaica). Evermann and Kendall 1899:61 (Florida). Evermann and Goldsborough 1902:152 (description, Yucatán). Jordan and Thompson 1905:234 (common, Tortugas, tongue parasite). Fowler 1906:91 (Florida Keys). Smith 1907:16 (upper caudal lobe orange-yellow; Beaufort Harbor, NC). Rosen 1911:51 (Bahamas). Nichols 1912:182 (Havana market, Cuba). Sumner et al. 1913:745 (Buzzards Bay, MA). Fowler 1915:531 (Trinidad). Fowler 1917b:23 (Virgin Islands). Fowler 1919a:7 (in part; western Atlantic). Fowler 1919b:144 (St. Croix), 145 (St. Christopher), 146 (Jamaica), 152–153 (Florida). Fowler 1920:154 (Sea Isle City, NJ; ANSP 327202). Nichols 1921:22 (Turks Island, Bahamas). Nichols and Breder 1927:59 (Woods Hole, MA). Borodin 1928:13 (Florida). Fowler 1928:457 (Haiti). Hildebrand and Schroeder 1928:153–154 (description; previous Chesapeake Bay localities). Nichols and Breder 1928:424–425 (Atlantic). Fowler 1930a:270 (Grenada, ANSP 52409). Breder 1932a:19–20 (description). Breder 1934:59 (Andros Island, Bahamas). Fowler 1936:434–435 (synonymy; description based on specimens from New Jersey and West Indies). Howell Rivero 1936:1 (description, Jamaica). Fowler 1937:309 (Haiti). Howell Rivero 1938:180 (MCZ 8775, types of H. filamentosus). Butsch 1939:19 (Barbados). Fowler 1941b:142 (listed; Brazil). Longley and Hildebrand 1941:30 (Tortugas). Fowler 1945:281–282 (synonymy, Pt. Everglades, FL). Miller 1945a:186–187 (diagnostic characters of Hemiramphus). Manter 1947:311 (trematode Steganoderma hemiramphi described from intestine of Tortugas specimens). Rivas 1949:8 (common names, range). Schultz 1949:81 (Venezuela, after Röhl 1942). Burkholder et al. 1959:92 (feed on turtle grass, Thalassia testudinium). Fowler 1952a:87 (Hispaniola), 112 (Cape May County, NJ). Woods 1952:40 (attracted to surface light at night; Campeche Banks), 52 (Caribbean). Manter 1954:339 (parasitic trematode, Steganoderma hemiramphi). Springer and Bullis 1956:58 (nine Gulf of Mexico stations). Robins 1958:14 (range, common names). Anderson and Gehringer 1959:44, 48 (Tongue of the Ocean). Schwartz 1961:5 (Patuxent River, MD). Springer 1961:481 (Tampa Bay area, Florida). Collette 1962:434–437 (comparison with H. bermudensis), fig. 1. Erdman 1962:227 (used as bait for Blue Marlin; may be food of Blue Marlin). Schwartz 1962 (Maryland). Tabb and Manning 1962:611–612 (Florida Bay area). Nahhas and Cable 1964:262 (trematodes Haplosplanchnoides hemiramphi and Lepocreadium hemiramphi described from H. brasiliense [sic] from Jamaica and Curaçao, respectively). Bane 1965:56 (common tuna bait species, Mona Passage). Cervigón 1965:26 (marao fosforo, Venezuela). Collette 1965:225–227 (synonymy, description, eastern Atlantic), fig. 6. Randall 1965:258–259 (feed on Cymodocea manatorum and Jenkinsia, St. Johns, Virgin Islands). Caldwell 1966:32 (Jamaica). Moe et al. 1966:17 (Pinellas and Munroe counties, Florida). Saunders 1966:430 (blood cell counts; Puerto Rico). Randall 1967:684–686 (diet in West

105

Fishes of the Western North Atlantic 1992:83 (Alabama). Nolf and Stringer 1992:70, pl. 12, fig. 22 (otolith from off Cuba). Cervigón et al. 1993:338 (northern coast of South America). Bohnsack et al. 1994:985–986 (landings, Monroe Count y, Florida). Humann 1994:58–59 (description, range; color photograph). Hensley and Hensley 1995:814 (eaten by Sooty Terns and Brown Noddies, Dry Tortugas, FL). McBride et al. 1996 (Florida bait fishery). Debelius 1997:58–59 (description, color underwater photograph). Murdy et al. 1997:120 (Chesapeake Bay, fig. 99). Smith 1997:386 (description, range; color pl. 108). McEachran and Fechhelm 1998:960, figure (Gulf of Mexico). Mejía Aristizábal et al. 1998:30 (Colombian Caribbean oceanic atolls). Carvalho Filho 1999:70 (Brazil). Allué et al. 2000:83 (Bahamas). Loftus 2000:34 (Everglades National Park, FL). Pérez-Hernández and Torres-Orozco 2000:434 (Mexican coastal lagoons). Schmitter-Soto et al. 2000:154 (Caribbean Mexico). Claro and Parenti 2001:43 (Cuba, listed). Gasparini and Floeter 2001:1643 (Trindade Island, South Atlantic). González-Gándara and Arias-Gonzáles 2001:249 (Alacran Reef, Yucatán, Mexico). McBride 2001 (landings in south Florida lampara net fishery). Rocha and Rosa 2001:989 (northeastern Brazil). Collette 2003b:1140 (description, range, figure). McBride and Styer 2003:24–26 (most common species in south Florida lampara net fishery). McBride and Thurman 2003 (reproductive biology). McBride et al. 2003 (spawning cycles in south Florida). Menezes 2003:68 (Brazil). Smith et al. 2003:15 (Belize). Alvarez-León and Celis-Rincón 2004:73, 77 (Archipiélago Islas del Rosario, Colombia). Collette 2004:4 (synonymy). Lovejoy et al. 2004:369, 376 (molecular phylogeny). Parenti and Grier 2004:336 (restricted lobular testis). Reckel and Melzer 2004 (falciform process in eye). Spach et al. 2004:51 (Brazil). Collette 2005:944–945 (early life history, figs. A–F). Fariña et al. 2005:585, 589 (one of most abundant species on a reef in Los Monjes Archipelago, Venezuela). Grier et al. 2005:208, fig. 19c (germinal eipithelium). Pimenta et al. 2005:1595 (in stomach contents of Sailf ish of f R io de Janeiro state, Brazil). Bouchon-Navaro et al. 2006:323 (weight-length relationships, seagrass beds of Guadeloupe and Martinique). Schwartz 2006:66 (North Carolina records). Smith-Vaniz et al. 2006:29 (Buck Island Reef National Monument, St. Croix). Walsh et al. 2006:260, 276 (juveniles collected off the U.S. Georgia coast). Fahay 2007:776, 792–793 (early stages, figs. A–D). Powell et al. 2007:61–63, 209 (densities and monthly length-frequencies, Florida Bay). Tibbetts et al. 2007:809 (vesicular swim bladder). Nóbrega and Lessa 2007:70 (artisanal fishery, northeastern Brazil). Casazza and Ross 2008:351 (associated with Sargassum, Gulf Stream off North Carolina). García-Hernández et al. 2009:92 (juveniles, northern Yucatán Peninsula). McEachran 2009:1286 (Gulf of Mexico). Campos et al. 2010:66 (Rio Grande do Norte, Brazil). Collette 2010:35 (reproduction and development). Pepperell 2010:234 (bait for sports fishers). Barros et al. 2011:371 (estuary, Pará, Brazil). Kells and Carpenter 2011:150– 151 (description, color painting). Vilar et al. 2011:45 (Baia da Babitonga, southern Brazil). Arévalo-Frías and Mendoza-Carranza 2012:258, table 3 (larvae and juveniles from the Pantanos de Centla Biosphere Reserve in Tabasco, Mexico). Passos et al. 2012:230 (Paranaguá Estuarine complex, Brazil). Rudershausen et al. 2012:158 (photograph of use in different hook arrangement for trolling). Costa and Camara 2012:475– 477 (more abundant in dry season, Guaraíras lagoon, Rio Grande do Norte, Brazil). Del Moral-Flores et al. 2013:832 (Veracruz Coral Reef System). De Angelo et al. 2014:1043 (seagrass beds in eastern Gulf of Mexico). Gallardo Torres et al. 2014:86 (Yucatán, Mexico, photographs). Smith-Vaniz and Jelks 2014:32 (St. Croix, U.S. Virgin Islands). Pinheiro et al.

2015:10 (underwater photograph, Trindade Island). Brewton et al. 2016:844 (in stomachs of Dolphinfish, Coryphaena hippurus). Snyder and Burgess 2016:100–101 (listed, Florida, color photograph). Starck et al. 2017:91 (Florida Keys). Hemiramphus marginatus Lesueur, 1821:135–136 (original description; Lesser Antilles, Guadeloupe, Martinique). Hemirhamphus brasiliensis. Lay and Bennett 1839:45 (specimen from gullet of Booby, Pelecanus sula, Rio de Janeiro, Brazil). Gill 1863:273 (type species). Evermann and Kendall 1899:62 (Florida records). Steindachner 1902:145 (description, Martinique). Evermann and Marsh 1902:102 (Puerto Rico, fig. 19). Jenkins 1903:434 (in part, West Indian specimens, not Hawaiian specimens that are Hemiramphus depauperatus). Bean 1905:299 (good food fish; Bahamas). Miranda Ribeiro 1915:20 (description, Brazil). Nichols and Mowbray 1917:79 (Puerto Rico). Miranda Ribeiro 1918:763 (Ilha Grande, Brazil). Meek and Hildebrand 1923:234–235 (description; Panama). Breder 1927:19 (Bahamas and Cuba). Beebe and Tee-Van 1928:66–68 (description, food; Haiti). Breder 1929:280 (Tortugas, FL). Jordan et al. 1930:199 (listed). Parr 1930:20 (Bahamas). Bigelow and Schroeder 1940:139 (Chatham, MA). Galloway 1941:119 (hypothermal mortality; Key West, FL). Röhl 1942:374 (balajú, description, Venezuela), fig. 186. Breder 1948:91 (description, range). Carvalho 1950:99, 101, 107–108 (Trindade Island, Brazil; food fish). Röhl 1959:456–457 (description, Venezuela), fig. 22. Miranda Ribeiro 1961a:8 (Fernando Noronha [MNRJ 1335] and Trindade Island [MNRJ 1338], Brazil). Nomura 1965:103–104 (length-weight; northeastern Brazil). Cervigón 1966:218–219 description, Venezuela), fig. 85. Menezes 1968:10 (agulha, Ceará, Brazil). Menezes 1969:17, 20 (44% occurrence in diet of 798 specimens of Scomberomorus cavalla, Ceará, Brazil). Santos 1970:285–304 (length-weight, sex ratio, age, sexual maturity, herbivore, fishery; Pernambuco, Brazil). Dahl 1971:181 (edible, but little appreciated; northern Colombia), figure. Duarte-Bello and Buesa 1973:81 (synonymy, range; Cuba). Fischthal and Nasir 1974:72–73, 77, 79 (three new species of digenetic trematodes described from small intestine, Los Roques Island, Venezuela; Bucephalopsis hemirhamphi, Paraproctotrema spinoacetabulum, and Conohelmins venezuelensis). Belyanina 1975:140 (ichthyoplankton, Caribbean Sea and Gulf of Mexico). Guitart 1975:253 (Cuba, synonymy, description, fig. 188). Subrahmanyam and Drake 1975:465 (northern Florida salt marshes). Instituto Nacional de Pesca 1976:75 (common names, range), 164 (fig. 106). Pequegnat et al. 1977:11 (Gulf of Mexico). Lima and Oliveira 1978:20 (agulha-preta; Ceará and Pernambuco states, Brazil). Mota Alves and Lima 1978:1 (spawning season February–April, Ceará, Brazil). Murdy 1983:90 (Texas). Rodríguez et al. 1984:14 (escribano de aletas rojas, Cuba). Perez and Rylander 1985:643 (hemoglobin heterogeneity, Venezuela). Castro-Aguirre et al. 1986:164 (Veracruz, Mexico). Maldonado and Yáñez-Arancibia 1987:191 (Campeche). Rodríguez and Valdés 1987:95 (escribano de aletas rojas, Cuba). Valdés-Muñoz and Garrido 1987:6, 10 (Cuba). Fuentes Mata et al. 1989:258 (Veracruz). Cervigón 1991:220–221 (Venezuela, fig. 166). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Méndez 2004:24 (Mochima National Park, Venezuela). Chávez-López et al. 2005:148 (Alvarado Lagoon estuary, Veracruz, Mexico). Méndez et al. 2006:686 (fringing reef Parque Nacional Mochima, Venezuela). Longart et al. 2011 (reproduction, Isla de Cubagua, Venezuela). Hemiramphus Brownii Valenciennes in Cuvier and Valenciennes, 1847:13–20 (original description, Martinique, Guadeloupe, Rio de Janeiro, Gorée). Kner 1867:324 (Rio de Janeiro, Brazil). Posada Arango 1909:297 (Colombia). Collette et al. 1997:15 (types).

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Family Hemiramphidae Color. Dark bluish green above, silvery white below. Beak black with fleshy red tip; entire upper lobe of caudal fin yellowish orange, lower lobe dusky. Size. The maximum observed size for Hemiramphus brasiliensis is 350 mm SL (405 mm TL, commonly to 350 mm TL). Bouchon-Navaro et al. (2006) provided length-weight information for 34 specimens from the Lesser Antilles.

Hemirhamphus filamentosus Poey, 1860:297–298 (original description; Cuba). Poey 1868:382 (description, Cuba). Poey 1881:337 (Puerto Rico). Collette et al. 1992:9 (type). Hemirhamphus browni. Jordan 1887a:529 (examination of type). Hemiramphus pleei Valenciennes in Cuvier and Valenciennes, 1847:21–24 (original description; Antilles). Meek and Goss 1884:222 (in key). Jordan 1891:315 (description; probably identical with Hemiramphus brasiliensis; Bahia, Brazil). Hemirhamphus pleei. Jordan 1885:546 (correct identification of Key West, FL, specimens previously identified as H. balao by Jordan [1885]). Jordan 1886b:31 (Cuba). Bean 1891:92 (abundant; Cape Charles City, VA). Cockerell 1892:14 (Jamaica). Haemiramphus braziliensis [sic]. Hill 1881:125 (Jamaica).

Biology Predators. A wide variety of pelagic fishes feed on species of Hemiramphus and there is a fishery for H. brasiliensis used as bait for billfishes, Dolphinfish, and King Mackerel (Berkeley and Houde 1978; Pepperell 2010). Dolphinfish, Coryphaena hippurus, three specimens in three stomachs off Hatteras, NC (Rose and Hasler 1974:96), in 23 of 2,219 individuals, 2.3% by volume, southeastern and Gulf coasts of the United States (Manooch et al. 1984:1517), and occasional in the Gulf of Mexico (Brewton et al. 2016); Bluefish, Pomatomus saltatrix, 0.8% frequency in southern Florida (Naughton and Saloman 1984:14); Caranx hippos, 0.1% frequency in northwest Florida, but unidentified Hemiramphus sp. with 10.7% frequency in southern Florida (Saloman and Naughton 1984:11, 20); Little Tunny, Euthynnus alletteratus, 0.2% frequency (Manooch et al. 1985:1212); King Mackerel, Scomberomorus cavalla, 4.4% occurrence in the state of Ceará, Brazil (Menezes 1969), and 10% occurrence off southeastern United States (Saloman and Naughton 1983); Cero, Scomberomorus regalis, in the West Indies (Randall 1967); and various species of billfishes, particularly the Sailfish, Istiophorus platypterus, usually identified only as Hemiramphus sp. (Voss 1953; Jolley 1977), White Marlin, Kajikia albida (Davies and Bortone 1976), and Blue Marlin, Makaira nigricans (Erdman 1962). Tabb and Manning (1962) observed small Carcharhinus limbatus feeding on H. brasiliensis in Florida Bay in the Everglades. A specimen was found in the gullet of a Booby, Pelecanus sula, in Rio de Janeiro (Lay and Bennett 1839). They are eaten by Sooty Terns and Brown Noddies in the Dry Tortugas (Hensley and Hensley 1995). Parasites. Ten species of parasites have been reported from the western Atlantic Hemiramphus brasiliensis: one isopod, one leech, seven trematodes, and one myxosporidian. isopods: The most common parasite of Hemiramphus brasiliensis is the isopod Glossobius hemiramphi Williams and Williams, which is found on top of the tongue in nearly 50% of the specimens in the Tortugas (Jordan and Thompson 1905), and at a high percentage in southeast Florida (Berkeley

Misidentifications. As Hemiramphus balao: Jamaica (Bean and Dresel 1884, in part, USNM 30077, 30100) and the Florida Keys (Jordan 1884a:114, USNM 35044; Jordan 1884b:79). Types. Hemiramphus Brownii Valenciennes in Cuvier and Valenciennes, 1847. Lectotype (designated by Collette 1965:226) MNHN 4585 (195 mm SL); Martinique; Plée, 1826. Dorsal fin rays 13; anal fin rays 12. Gill rakers on first arch 10 + 23 = 33; gill rakers on second arch 11 + 20 = 24. Lower jaw length 55.6 mm; pectoral fin length 33.7 mm. Paralectotypes MNHN A.7602 (249 mm SL) and B.1052 (3, 194–201 mm SL). Hemirhamphus filamentosus Poey, 1860. Lectotype (designated by Collette 1965:226) MCZ 8775 (237 mm SL); Cuba; Poey no. 50. Dorsal fin rays 14; anal fin rays 13. Gill rakers on first arch 7 + 22 = 29; gill rakers on second arch 5 + 20 = 25. Lower jaw and pectoral fins broken. Paralectotypes MCZ 43030 (2, 195–225 mm SL) and USNM 4746 (1, 182 mm SL). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Ballyhoo (English), agujeta brasileña (Spanish, Mexico), and demi-bec brésilien (French). The Spanish common name in Cuba is escribano de aletas rojas (Claro and Parenti 2001). We prefer Redtail Ballyhoo as the common name in English because it is more descriptive. Diagnosis. Hemiramphus brasiliensis differs from H. balao and H. saltator in its shorter pectoral fin, which does not reach the nasal fossa when folded forward. Differs from H. bermudensis and H. saltator in having fewer gill rakers (37–44 and 38–44, respectively). Adults differ from H. balao in having a reddish instead of a bluish tip to the upper caudal lobe. Description. Dorsal fin rays 12–15, usually 13 or 14; anal fin rays 11–15, usually 13; pectoral fin rays 10–12, usually 11. Vertebrae 34–37 precaudal + 16–18 caudal = 52–55 total, usually 36 precaudal + 17–18 caudal = 53–54 total. Gill rakers on first arch 7–10 + 20–26 = 28–36 (Table 4.4); gill rakers on second arch 3–7 + 18–23 = 20–29 (Table 4.5). Upper jaw short, scaleless; preorbital ridge (bony ridge under nostril) absent. 107

Fishes of the Western North Atlantic

Figure 4.9. Redtail Ballyhoo, Hemiramphus brasiliensis. A. ANSP 103311, 272 mm SL. Nassau Harbor, Hog Island, Bahamas, 20 March 1958. B. USNM 188749, 119 mm SL. Collected on the M/V Theodore N. Gill off Grand Bahama Island, 29 August 1954. C. USNM 188748, 50 mm SL. Collected on the M/V Silver Bay off North Carolina, U.S.A., 18 July 1960. Illustrations by M. H. Carrington.

and Houde 1978), although it was not formally described until 1985 by Williams and Williams on the basis of 35 isopods from 18 halfbeaks from Puerto Rico. It is known from H. brasiliensis from both the western and eastern Atlantic (Bruce and Bowman 1989). leeches: A marine leech of the family Piscicolidae, Malmia philotherma, was described from several species of fishes, including Hemiramphus brasiliensis from Puerto Rico (Sawyer et al. 1975; Wallace and Sawyer 1977). trematodes: Several trematodes have been reported from Hemiramphus brasiliensis: Zoogonidae,

Steganoderma hemiramphi Manter, 1947, from Tortugas (Manter 1947), Curaçao, and Jamaica (Nahhas and Cable 1964); Haplosplanchnidae, Haplosplanchnoides hemiramphi Nahhas and Cable, 1964, from Jamaica; Lepocreadiidae, Lepocreadium hemiramphi Nahhas and Cable, 1964, from Curaçao; Haplosplanchnidae, Schikhobalotrema adacuta (Manter), from Curaçao and Jamaica (Nahhas and Cable 1964); Bucephalidae, Bucephalopsis hemirhamphi Fischthal and Nasir 1974, from Venezuela; Monorchiidae, Paraproctotrema spinoacetabulum Fischthal and Nasir, 1974, from Venezuela; and Waretrematidae, Conohelmins venezuelensis Fischthal and Nasir, 1974, from Venezuela. 108

Family Hemiramphidae myxosporidians: Spores of a myxosporidian, Myxidium sp., were reported from the gall bladder of Hemiramphus brasiliensis from south Florida (Iverson et al. 1971). Food. The food of this species consists largely of seagrasses, manatee grass, Cymodocea manatorum, and turtle grass, Thalassia testudinium: Beebe and TeeVan 1928 (Haiti); Burkholder et al. 1959; Randall 1965, 1967 (Virgin Islands and Florida); Valdés-Muñoz and Silva Lee 1977 (Cuba). Randall (1965, 1967) reported stomach contents of Hemiramphus brasiliensis to be Cymodocea and Jenkinsia, sometimes noting that the contents were ground up. However, one Brazilian study (Vasconcelos Filho et al. 1984) showed the diet to consist mostly of diatoms and other plant material. Beebe and Tee-Van (1928) also noted green algae, insects, spiders, and small fishes. The lower jaw is used to guide the blades of floating seagrasses into the mouth, where the pharyngeal mills draw the blades in at a uniform rate (Randall 1965). Occasionally, two individuals will start at opposite ends of the same blade of grass and “fight” over it (Randall 1965; B. B. Collette, pers. observ.). Reproduction. Cyclic patterns of gonadosomatic indices indicate that Hemiramphus brasiliensis spawns during spring and summer months in south Florida (McBride et al. 2003). Hydration of oocytes began in the morning and spawning occurred at dusk the same day. All mature females spawned daily in April (McBride and Thurman 2003). Batch fecundity is lower than in H. balao, an average of 1,164 hydrated oocytes, rather than 3,734, in a 100 g female. Spawning occurs all along the coral reef tract of the Atlantic Ocean, inshore of the reef tract, in shallower, more inshore waters than in H. balao. A large female (USNM 344910, 246 mm SL) collected from Cayo Hachuela, Cuba, had 100 eggs in the left ovary (1.00–1.45 mm in diameter, x̅  1.17 mm for 20 eggs) and 120 eggs in the right ovary (0.90–1.50 mm in diameter, x̅  1.15 mm for 20 eggs). Relationship to Humans. Hemiramphus brasiliensis is important as a baitfish for billfishes, particularly in the Florida Keys. Commercial landings in Monroe County, FL, in 1992 were 0.27 × 106 kg (Bohnsack et al. 1994). Caught, sold, and consumed at Búzios Island off the coast of São Paulo state, Brazil (Begossi and de Figueiredo 1995). Range. Atlantic Ocean, in the western Atlantic from Buzzards Bay (Sumner et al. 1913) and Woods Hole, MA (Nichols and Breder 1927), south through the Gulf of Mexico and Caribbean Sea to Rio de Janeiro and Porto Belo, Brazil (Lucena and Lucena 1981), and Trindade Island (Pinheiro et al. 2015) (Fig. 4.8). Absent from Bermuda. Present in the eastern

Alantic from the Canary Islands and Dakar, Senegal, south to Luanda, Angola (Collette 1965). Study Material. A total of 727 specimens (14.1–290 mm SL) from the western Atlantic, including the types of Hemiramphus browni (MNHN 4585, A.7602, B.1052) and H. filamentosus (MCZ 8775, 43030), originally at AMNH, ANSP, BMNH, CAS, FMNH, FSBC, ISH, MCZ, MNHN, MNRJ, MZUSP, TU, UMML, USNM, VIMS, YPM, and ZMK. The specimens came from six general regions: 215 specimens (14.1–274 mm SL) from the east coast of the United States; 152 specimens (36.5–272 mm SL) from the Bahamas; 44 specimens (55.0–263 mm SL) from the Gulf of Mexico; 181 specimens (31.1–261 mm SL) from the Antilles; 29 specimens (32.2–248 mm SL) from Central America; and 104 specimens (132–290 mm SL) from South America. Comparative material from the Gulf of Guinea includes 221 specimens (17–223 mm SL) reported previously (Collette 1965), plus 7 specimens (115–216 mm SL) from ISH and RMNH. Genus Hyporhamphus Gill, 1859 Subgenus Hyporhamphus Gill, 1859 Inshore Halfbeaks Hyporhamphus Gill, 1859a:131 (type species Hyporhamphus tricuspidatus Gill = Hyporhamphus unifasciatus (Ranzani) by monotypy). Ichthyacus Fernández-Yépez, 1948 (type species I. breederi by original designation and monotypy).

Diagnosis. Hyporhamphus sensu stricto differs from the subgenus Reporhamphus Whitley in the structure of the preorbital canal (no posterior branch) of the lateral line system, the shape of the caudal fin (truncate or only slightly forked instead of deeply forked), and the shape of the body (slightly compressed laterally). Description. Body fairly small, moderately compressed; oval in cross section. Greatest body depth 6.3–14 times in SL (increasing with growth), width of body 1.1–1.8 times in depth. Vertebrae 28–37 precaudal + 16–20 caudal = 45–55 total. Scales moderately large; predorsal scales 30–63. Pectoral branch of lateral line single. Swim bladder simple (one-celled). Head length 3.9–5.7 times in SL. Upper jaw blunt or pointed anteriorly; its length less or greater than its width, length divided by width 0.6–1.3. Interorbital region flat. Lower jaw elongate (2.9–7.6 times in SL, decreasing with growth), lower jaw length in head length 0.65–1.8. Preorbital ridge present. Preorbital canal simple, without posterior branch. Jaw teeth arranged in three to eight rows, tricuspid or unicuspid. Pharyngeal teeth on lower pharyngeal bone and third upper pharyngeal bones tricuspid 109

Fishes of the Western North Atlantic in the anterior part of the plate, tricuspid or blunt in the center part, and blunt in the posterior part; on upper pharyngeal bone on third gill arch sharp with a single cusp or tricuspid with one cusp significantly larger. Opercle, suborbital, and interorbital regions of the head covered with scales. Scales present on the upper jaw and scales present or absent on anterior parts of dorsal and anal fins. Nasal papilla not fimbriate. Dorsal fin rays 12–17; anal fin rays 13–19; pectoral fin rays 10–13. Vertebrae 28–37 precaudal + 16–20 caudal = 45–55 total. Gill rakers on first arch 19–53; gill rakers on second arch 15–47. Pectoral fins short (5.8–9.2 times in SL). Pelvic fins short (9.8–19.5 times in SL), outer rays slightly longer than inner rays, located closer to the origin of the pectoral fins than to the origin of the caudal fin, or about midway between them (P1–P2/P2–C = 0.5–1.15). Origin of anal fin located under rays 1–3 of the dorsal fin or in advance of its origin (origin of dorsal fin may be above rays 1–3 of anal fin). Caudal fin truncate or weakly forked with slightly longer lower lobe. Species. All Atlantic species belong to the subgenus Hyporhamphus, which has 25 species, four of which occur in the western North Atlantic, plus two freshwater species (Collette 2004). Range. Circumtropical distribution, with many species migrating to higher latitudes during seasonal warming (Banford 2010). Representatives of the subgenus Hyporhamphus occur in tropical and temperate waters along the continental coasts of all oceans, but are generally absent from oceanic islands (except for Bermuda and the Galápagos). The species live in close proximity to shore, mainly in bays, gulfs, and estuaries; some species occur in fresh waters. All eastern Pacific, western Atlantic, and eastern Atlantic species of Hyporhamphus belong to the subgenus Hyporhamphus. Species in the subgenus Reporhamphus are restricted to the Indo-West Pacific (Collette 2004).

2003a:589 (Orinoco and Amazon rivers). Collette 2004:6–7 (Orinoco and Amazon rivers). Lasso et al. 2004:144 (Venturai and Delta, Orinoco Basin).

Types. Hyporhamphus brederi (Fernández-Yépez, 1948). Holotype (45.5 mm SL); Venezuela, Orinoco River, Boca del Cano Orape “entre Pto Páez y La Urbana” (Fernández-Yépez 1948:2); A. FernándezYépez and F. J. Fernández-Yépez no. 48.051; 8 May 1948. According to the original description, the type was in the personal collection of Augustin FernándezYépez. The present whereabouts of this specimen are unknown. From the original description: dorsal fin rays 12; anal fin rays 15; lower jaw length 28.1 mm; head length 11.1 mm. Common Names. No of ficial common name has been suggested, so we recommend Breder’s Halfbeak. Diagnosis. A species of Hyporhamphus without scales covering the dorsal and anal fins; a very long beak, longer than half the standard length; and a moderate number of gill rakers on the first arch (28– 32). Differs from all other species of Hyporhamphus under consideration in its greatly elongate beak. Description. Dorsal fin rays 13–15; anal fin rays 13–16; pectoral fin rays 9–11 (Table 4.6). Vertebrae 31–32 precaudal + 15–17 caudal = 47–48 total. Gill rakers on first arch 6–8 + 21–23 = 28–32; gill rakers on second arch 3–5 + 19–22 = 23–26 (Table 4.7). Distance from pelvic fin origin to caudal base extended forward, falls between the posterior tip of the opercle and the pectoral fin base. Preorbital canal narrow, median pore directed anteriorly, sharp angle dorsal to median pore, ventral pore very slightly enlarged. Size. The maximum observed size of Hyporhamphus brederi is 104 mm SL (MZUSP 5733). Biology Reproduction. Males appear ripe by 97 mm SL, females by 85 mm SL. Range. This species is restricted to fresh water and to the Orinoco River (type locality), and the Amazon River at Santarem (Collette 2003a). Geographic Variation. Specimens collected in the Brazilian Amazon have fewer dorsal and more anal rays than those collected in Venezuela’s Orinoco River (Table 4.6), but this is based on limited data. Study Material. A total of 154 specimens (25.0– 81.7 mm SL) from Brazil from five collections: MNRJ 1897 (24, 25.0–81.7 mm SL); Amazon River at Santarem, Estada Pará. CAS 81339 (1, 76.7 mm SL); Lago Grande; C. Ternetz, November 1924. MZUSP

Hyporhamphus brederi (Fernández-Yépez, 1948) Breder’s Halfbeak Figures 2.6, 4.10 Tables 4.1, 4.2, 4.6, 4.7 Ichthyacus breederi Fernández-Yépez, 1948 (original description; Orinoco River between the mouths of the Meta and Apure rivers; named breederi after C. M. Breder, Jr., of the American Museum of Natural History, emended to brederi), figure of type, Fernández-Yépez no. 48.051. Hyporhamphus brederi. Collette 1966:1 (South America). Mago Leccia 1970:20, 89 (listed, fresh water, Venezuela). Collette

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Family Hemiramphidae

Figure 4.10. Breder’s Halfbeak, Hyporhamphus brederi. MNRJ 1897, 81.7 mm SL. Santarem, Brazil, 11 September 1927. Illustration by M. H. Carrington.

backwater of Cano Araguao, 112 nautical miles, 08°38′N, 61°43′W; J. N. Baskin et al., JNB 50-78, 21 February 1978. USNM 233543 (1, 37.4 mm SL); Delta Amacuro territory, Orinoco River, inlet, 82 nautical miles; J. N. Baskin et al., JNB 6478, 22 February 1978. USNM 233548 (1, 62.3 mm SL); Bolivar state, laguna on southern side of Isla Isabela, between Palua and Ciudad Bolivar, 201 nautical miles, 08°13′N, 65°56′52″W; J. Lundberg et al., JNB 1779, 7 November 1979. USNM 233544 (1, 39.0 mm SL); Delta Amacuro territory, small cano and swamp off Orinoco River on northern side of Isla Carosimo, 85.7 nautical miles, 08°25′54″N, 61°20′54″W; D. J. Stewart et al., DJS 1579, 21 November 1979. USNM 233555 (2, 56.6–93.2 mm SL); Bolivar state, laguna on southern side of Isla Isabela, between Palua and Ciudad Bolivar, 201 nautical miles, 08°18′43″N, 65°56′52″W; J. Lundberg et al., JNB 1779, 7 November 1979.

5733 (127, 50–104 mm SL); Rio Tapajos at Santarem, 02°25′S, 54°44′W; February–March 1967. MZUSP 31689 (1, 84.5 mm SL) and MZUSP 58987 (1, 91 mm SL); Rio Tapajos at Santarem; M. Goulding, 25 November 1983. A total of 21 specimens (18.3–99.6 mm SL) from Venezuela’s Orinoco drainage from eight collections: USNM 227643 (8, 19.7–32.0 mm SL); Bolivar state, Orinoco River at dock at Palua, 182 nautical miles from sea buoy, 08°22′N, 62°42′W; D. Taphorn et al., JNB 978, 13 February 1978. USNM 227644 (1, 99.6 mm SL); Monagas State, Barrancas River, 141 nautical miles, 08°43′N, 62°08′W; J. N. Baskin et al., JNB 2878, 17 February 1978. USNM 227645 (6, 19.0–97.0 mm SL); Delta Amacuro territory, Cano el Consejo, 134 nautical miles, 08°41′N, 62°03′W; J. N. Baskin et al., JNB 4678, 19 February 1978. USNM 233542 (1, 18.3 mm SL); Delta Amacuro territory,

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Fishes of the Western North Atlantic in comparison to H. unifasciatus, which has 19–28, usually 23–25, or H. meeki, which has 20–30, usually 26–27 (Table 4.10). Sequence data for 800 base pairs of mtDNA Cyt b gene clearly separate the Bermudan species from other species of Hyporhamphus by a minimum genetic distance of 0.034 (Banford 2010). Description. Dorsal fin rays 13–16, usually 14 or 15; anal fin rays 15–18, usually 16 or 17; pectoral fin rays 9–10, usually 10 (Table 4.8). Vertebrae 32–33 precaudal + 18–19 caudal = 50–52 total. Gill rakers on first arch 26–32, usually 29 or 30 (Table 4.9); gill rakers on second arch 20–25, usually 21 or 22 (Table 4.10). Adults with dorsal and anal fin bases covered with scales. Color. Green-tan on dorsal portion of body, lateral silver stripe from opercle opening onto caudal peduncle and silver-white on ventral body. Fleshy tip of lower jaw bright reddish orange (Banford 2010). Size. The maximum observed size for Hyporhamphus collettei is 168 mm SL (MCZ 34890, Bermuda; Banford 2010).

Hyporhamphus collettei Banford, 2010 Collette’s Bermuda Halfbeak Figures 4.11, 4.12 Tables 4.1, 4.2, 4.8, 4.9, 4.10 Hyporhamphus unifasciatus (not of Ranzani 1842). Barbour 1905:114 (description; Flatts Inlet, Bermuda). Bean 1906:36 (Bermuda; FMNH 5096–5103). Linton 1908:86 (green algae in stomach, Bermuda). Beebe and Tee-Van 1933:63–64 (description, Bermuda). Saunders 1959:374, 376 (Bermuda specimen parasitized by Haemogregarina bigemina). Rees 1970:217 (no helminths in 12 Bermuda specimens). Bruce 1986:1133 (parasitic isopod Mothocya bermudensis on Hyporhamphus unifasciatus from Bermuda). Parrish 1989:80 (feed on floating seagrasses, Bermuda). Smith-Vaniz et al. 1999:177–178 (Bermuda records). Hyporhamphus roberti (not of Valenciennes in Cuvier and Valenciennes 1847). Bean 1906:36 (Bermuda). Hyporhamphus hildebrandi (not of Jordan and Evermann 1927). Beebe and Tee-Van 1933:63 (rare; Bermuda). Hemiramphus unifasciatus (not of Ranzani 1842). Fowler 1930b:146 (Bermuda; ANSP 5234045, 123715). Hyporhamphus collettei Banford, 2010 (original description).

Misidentifications. As Hemiraphas [sic] braziliensis from Bermuda (Borodin 1934). Types. Holotype USNM 391198 (135 mm SL); Bermuda. See Banford (2010) for full list of paratypes. Common Names. The common name Collette’s Halfbeak was suggested by Banford (2010). Because another halfbeak was described with the same species name, Dermogenys collettei Meisner, 2001, we emend the common name to Collette’s Bermuda Halfbeak to avoid confusion. Diagnosis. A small, slender species with a ratio of body depth at pectoral fin origin to standard length usually less than 0.12; ratio of body depth at pelvic fin to standard length usually less than 0.12. Hyporhamphus unifasciatus has a deeper body at both pectoral and pelvic fin origins. Pectoral fin rays almost always 10 (rarely 9), whereas H. unifasciatus usually has 11 pectoral fin rays and H. meeki usually has 11 or 12 (Table 4.8). Hyporhamphus collettei has 20–25 total gill rakers on the second arch, usually 21–22,

Biology Parasites. Saunders (1959) found one specimen f rom Bermuda infected with Haemog regarina bigemina Laveran and Mesnil. Rees (1970) did not find any helminths on 12 specimens. In his revision of the cymothoid isopods of the genus Mothocya, Bruce (1986) described a new species, M. bermudensis, from Hyporhamphus collettei from Bermuda and H. unifasciatus from Haiti. Food. In Bermuda, Hyporhamphus collettei actively feed on floating pieces of seagrasses (Parrish 1989; B. B. Collette, pers. observ.). Habitat. Marine inshore, sandy areas with seagrasses. Range. Endemic to Bermuda (Fig. 4.12; Banford 2010). Study material. A total of 94 specimens (37.0– 168 mm SL) from Bermuda, as listed in Banford (2010:355), including the holotype (USNM 391198).

Figure 4.11. Collette’s Bermuda Halfbeak, Hyporhamphus collettei. Holotype, USNM 391198, 135 mm SL. Collected from Ferry Reach at Biological Station Bermuda, 3 August 1990. Illustration by H. M. Banford.

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Family Hemiramphidae 1932c:180, Breder and Nigrelli 1934:194 (Sandy Hook Bay, NJ). Gordon 1949:24 (August, Coney Island, NY). Caballero y Caballero and Bravo Hollis 1965:299 (trematode Lobatostoma ringens from intestine; Veracruz, Mexico). Hendrix and Overstreet 1977:814 (reference to Caballero y Caballero and Bravo Hollis 1965). Gilmore et al. 1981:10 (Indian River lagoon, Florida). Smith 1997:386 (in part, description, range). Hemiramphus sp. Putnam 1870a:171 (Danversport, North River, MA). Schwartz 1964:180–181, 191 (preyed on by Bluefish and Summer Flounder; Ocean City, MD). Hemirhamphus unifasciatus (not of Ranzani 1842). Cope 1870:119 (Newport, RI, ANSP 750912). Gill 1873:31 (Cape Cod, MA, to Florida). Yarrow 1877:214 (abundant late summer, eaten by Bluefish, Fort Macon, NC). Goode 1879:116 (St. Johns River, FL). Jordan and Gilbert 1879:383 (abundant; Beaufort Harbor, NC). Jordan 1880a:20 (Indian River, FL). Jordan 1880b:222 (St. Johns River, FL). Jordan and Gilbert 1882a:262 (Galveston, TX, USNM 31027). Bean 1883:450 (range, in part). Jordan and Gilbert 1883b:588 (Charleston, SC, specimen agrees with Beaufort, NC, specimens). Jordan and Swain 1884:231 (Cedar Keys, FL). Evermann and Kendall 1894:108 (Galveston, TX). Hemirhamphus roberti (not of Valenciennes in Cuvier and Valenciennes 1847). Bean 1890:40 (description, rare in New York waters; Fire Island, Great South Bay). Bean 1891:92 (Cape Charles City, VA and Point Lookout, MD). Hemorhamphus [sic] unifasciatus (not of Ranzani 1842). Goode 1879:116 (St. John’s River, FL). Hemiramphus roberti (not of Valenciennes in Cuvier and Valenciennes 1847). Meek and Goss 1884:221–223 (specimens from Atlantic coast of the United States north of Florida Keys differ from West Indian Hyporhamphus unifasciatus in slenderness of body and greater length of lower jaw). Jordan 1886a:26 (Beaufort, NC). Hyporhamphus unifasciatus (not of Ranzani 1842). Barbour and Cole 1906:157 (Progreso, Yucatán, Mexico, MCZ 32881). Cockerell 1913:52 (in part, scales of Woods Hole, MA, specimens similar to Panamanian specimens and those of Hyporhamphus quoyi). Fowler 1916b:41 (Sandy Hook, NJ). Fowler 1917a:120–121 (description; Newport, RI, ANSP 750912). Fowler 1920:154–155 (Atlantic, Cape May, and Monmouth counties, New Jersey). Hildebrand and Schroeder 1928:152–153 (description, food; Chesapeake Bay). Burkenroad 1931:20–21 (sound production, Louisiana). Fowler 1938:60 (between Longport and Ocean City, NJ). Greeley 1939:84 (Long Island, NY). Pearse et al. 1942:188 (Beaufort, NC). Gunter 1945:47 (Aransas Bay and Mustang Island, TX). Baughman 1950:135 (previous Texas records; USNM 120056). Fowler 1950:68 (Hampton Roads, VA, ANSP 979801). Fowler 1951b:74 (40 miles southeast of Atlantic City, NJ). Fowler 1952a:112 (same as Fowler 1920). Scattergood 1952:205 (Small Point, ME). Bigelow and Schroeder 1953:169 (description; rare stray in Gulf of Maine), fig. 81. Mansueti and Scheltema 1953:4 (Choptank River opposite Cambridge, MD, Cape Charles, VA; salinities 13.2‰–23.9‰). Reid 1954:22 (March–December, gonads developing in March; Cedar Keys, FL). Joseph and Yerger 1956:127 (young first appear May–July; Alligator Harbor, FL). Kilby 1955:194 (open waters near shores of bays; May–November, salinities 7.5‰–27.2‰; Cedar Keys and Bayport, FL). Menzel 1956:51 (St. George’s Sound, Apalachee Bay, FL). Simmons 1957:183, 191 (salinities 25‰–45‰; upper Laguna Madre, TX). Gunter 1958:190 (southern Texas). Anderson and Gehringer 1959:48 (southern Atlantic coast of the United States.). Hoese 1959:325 (common in surface waters at night, Texas). Leim and Day

Figure 4.12. Distribution of Collette’s Bermuda Halfbeak, Hyporhamphus collettei, Meek’s Halfbeak, Hyporhamphus meeki, and Atlantic Silverstripe Halfbeak, Hyporhamphus unifasciatus, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

Hyporhamphus meeki Banford and Collette, 1993 Meek’s Halfbeak Figures 4.12, 4.13 Tables 4.1, 4.2, 4.8, 4.9, 4.10 Hyporhamphus roberti (not of Valenciennes in Cuvier and Valenciennes 1847). Jordan and Evermann 1896a:321 (records from the east coast of the United States, in part). Smith 1898:93 (Vineyard Sound, Menemsha, and West Falmouth, MA; July– September). Evermann and Kendall 1899:62 (Tarpon Springs, FL). Bean 1900:100, 1902:406 (Great South Bay, NY). Linton 1905:358 (herbivore, parasites; Beaufort, NC). Smith 1907:161 (description; visits North Carolina in spring and remains in summer; feeds on green algae and small crustaceans). Kendall 1908:121 (previous records from Massachusetts and Rhode Island). Tracy 1910:88–89 (occasional in Rhode Island, summer and early fall). Sumner et al. 1913:745 (summarizes previous New England records). Nichols 1913:96 (June–December; near New York City). Wiegmann and Nichols 1915:44 (Coney Island, NY). Kendall 1914:41 (Machias and Casco Bay, ME). Latham 1917:19, 1918:54, 1919:56 (Orient, Long Island Sound, NY). Nichols 1918:53, 107 (common in New York City area, August–October). Fowler 1919a:7 (Newport, RI, ANSP 750912; Corson’s Inlet, NJ, ANSP 36652; Volusia, FL, ANSP 18894; St. Augustine, FL, ANSP 20343; Point Pinellas, FL, ANSP 7515). Latham 1920:91, 1923:61 (Long Island, NY). Nichols and Breder 1927:59 (summer and early fall resident, New York and southern New England). Breder

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Fishes of the Western North Atlantic 1959:507 (Chamcook, New Brunswick). Arnold et al. 1960:11 (June–October, Galveston Island, TX). Gordon 1960:31 (coastal waters in summer and full; Rhode Island). McAllister 1960:19 (listed after Leim and Day, 1959). Springer and Woodburn 1960:25 (salinities 25.2‰–25.8‰, temperatures 28.9–31.0 °C; widespread in Tampa Bay, FL). Schwartz 1961:5 (Patuxent River, MD). Hellier 1962:16 (late summer and early fall, Laguna Madre, TX). Legendre et al. 1964:52 (listed, Canadian Atlantic). Alperin and Schaefer 1965:3 (previous New York records; salinities 23.2‰–25.4‰, September, Long Island, NY). Parker 1965:208 (Galveston Bay, TX). Roithmayr 1965:20 (northern Gulf of Mexico). Fox and Mock 1968:46, 51 (spring and summer; Berateria Bay, LA). Dawson 1969:58 (barnacle Conchoderma virgatum attached to dorsal fin, Round Island, Mississippi Sound). Kingston et al. 1969:545 (monogenetic trematode Axinoides hyporhamphi). Laird and Bullock 1969:1080 (one Woods Hole, MA, specimen lacked haematozoa). Lunz and Schwartz 1970:22 (off Borders Island, 8 miles up Ashepoo River, SC). Reséndez Medina 1970:102 (description; Laguna de Tamiahua, Veracruz, Mexico), fig. 16. Grimes 1971:42 (Crystal River, FL). Perret et al. 1971:45 (Louisiana estuaries, salinities 15.0‰–18.4‰). Swingle 1971:29 (salinities 10.0‰–31.6‰, July–August; Alabama). Mountain 1972:53 (Crystal River, FL). Carr and Adams 1973:512, 517 (juveniles planktivores, adults omnivores; Crystal River, FL). Derickson and Price 1973:555–558 (summer, Delaware). Nixon and Oviatt 1973:486 (September–November; Narraganset Bay, RI). Tagatz and Wilkens 1973:8 (northwestern Florida estuary). Turner and Johnson 1973:7 (Newport River, NC). Burgess and MacPherson 1974:163 (Bogue Sound, NC). McNulty et al. 1974:104–105, 126–127, 182–183 (Tampa Bay, Cedar Keys, St. Marks, FL). Williams and Gaines 1974:139 (no Acanthocephala found; Mobile Bay). Wittenberg and Haedrich 1974:143 (choroid rete and pseudobranch present). Fahay 1975:4, 5, 17, 19 (larvae and juveniles, southern Atlantic Bight). Grimes 1975:174 (Crystal River, FL). Subrahmanyam and Drake 1975:455–456, 465 (occurrence, food; northern Florida salt marshes). Adams 1976:274 (occurrence in eelgrass communities; Phillips Island and Bogue Sound, NC). Gilmore 1977:132 (records from Indian River area, Florida). Hoese and Moore 1977:150 (description, most common inshore halfbeak in northwestern Gulf of Mexico), pl. 119. Cooley 1978:102 (adults common spring and summer, juveniles rare in winter; Santa Rosa Sound, FL). De Vane 1978:584 (eaten by Scomberomorus cavalla, Onslow Bay, NC). Hardy 1978:133–138 (description, development, figures). Naughton and Saloman 1978:46–50 (July–October; St. Andrew Bay, FL). Gilmore et al. 1979:82 (hypothermal mortality; Indian River Lagoon, FL). Hastings 1979:7 (St. Andrew jetties, Florida). Modde and Ross 1980:914 (6.3% frequency of occurrence, surf zone habitat, northern Gulf of Mexico). Subrahmanyam and Coultas 1980:797, 800 (St. Marks and Wakulla tidal marshes, Florida). Williams 1980:384–385 (new monogenetic trematode, Ancyrocephalus trullae, described from specimens from Dauphin Island, AL). Gilmore et al. 1981:10 (common; Indian River Lagoon, FL). Reséndez Medina 1981:267–268 (description; Laguna de Terminos, Campeche, Mexico), pl. IV, fig. 27. Guillory 1982:5 (common; Grand Isle area, Louisiana). Schwartz et al. 1982:27 (temperature 13.7–31.4 °C, salinity 7.0‰–32.0‰; Cape Fear Estuary, NC). Fahay 1983:210–211 (larvae, after Hardy and Johnson 1974). Flores-Coto et al. 1983:252 (larvae caught spring, summer, and fall; Tamiahua Lagoon, Mexico). Manooch and Hogarth 1983:231 (five specimens found in two stomachs of Wahoo, Acanthocybium

solandri). Murdy 1983:90 (Texas). Snelson 1983:192 (ecology; Indian River Lagoon system, Florida). Bruce 1986:1112 (parasitic isopod Mothocya nana on specimens from Chesapeake Bay, Georgia, and Florida). Robins and Ray 1986:102 (description, range in part), pl. 17. Ditty 1987:945 (ichthyoplankton off Louisiana, September). Gomez-Soto and Contreras-Balderas 1988 (in part, Tamaulipas, Mexico). Ditty et al. 1988:813 (larvae, northern Gulf of Mexico). Olney and Boehlert 1988:36 (eggs, Chesapeake Bay). Schwartz 1989:340 (North Carolina). Scott and Scott 1988:310 (Chamcook, New Brunswick, Canada). McAllister 1990:127 (Canada). Wenner and Sedberry 1989:48 (southeastern United States). Norcross and Hata 1990:448 (Virginia Barrier Islands). Able 1992:4 (New Jersey, occasional). Boschung 1992:83 (Mobile Bay, AL). Brown-Peterson et al. 1993:181 (Indian River Lagoon, FL). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Casazza and Ross 2008:351 (one individual associated with Sargassum, Gulf Stream off North Carolina). Wingate and Secor 2008:1149 (occasional in Chesapeake Bay). Schwartz 2012:44 (rare in North Carolina). Hyporhamphus hildebrandi (not of Jordan and Evermann 1927). Pearse et al. 1942:144 (Beaufort, NC). Breder 1948:91 (in part, description; part of range). Nicol and Arnott 1973:71 (neither eyeshine nor tapetum lucidum present; Port Aransas, TX). Hyporhampus [sic] sp. Price 1962:4–6 (new monogenetic trematode, Axine hyporhamphi described from Woods Hole, MA, specimen). Schwartz 1962: (Maryland). Collette 1966:3 (comparative material). Richards and Castagna 1970:243 (common in Chesapeake Bay tributaries). Wiley 1970:156 (Potomac River below Great Falls). Lewis and Wilkens 1971:186 (larvae; lower Newport River estuary, NC). Hardy and Johnson 1974:243–245 (larvae, Chesapeake Bay), fig. 2 (larvae and prejuvenile). Hemirhamphus sp. Thomson et al. 1971:79–80 (juvenile, Guilford, CT, Long Island Sound, NY). Hyporhamphus meeki (nomen nudum). Bruce 1986:1112 (infested with parasitic isopod, Mothocya nana, Florida). Hyporhamphus meeki Banford and Collette, 1993:371–382, fig. 2A (original description; Morehead City, NC). Contreras-Balderas et al. 1997 (Mexican Gulf of Mexico). Murdy et al. 1997:120–121 (Chesapeake Bay, fig. 100). Paperno et al. 2001:127 (Ponce de Leon Inlet, FL). Collette 2002:290–291 (Gulf of Maine), fig. 156. Gibson and Myers 2002 (Nova Scotia, Canada). Humann and DeLoach 2002:62–63 (description, range; color photograph). Collette 2003b:1141 (description, range, figure). Collette 2004:8–9 (synonymy). Poulakis et al. 2004:126 (Charlotte Harbor, FL). Collette 2005:946–947 (early life history, figs. A–F). Schwartz 2006:66 (North Carolina records). Tuckey and Dehaven 2006:107 (seagrass habitat, Suwanee River estuary, FL). Fahay 2007:776, 794–795 (early stages, figs. A–E). Love and May 2007:255 (rare, Maryland coastal bays). McEachran 2009:1286 (Gulf of Mexico). Kells and Carpenter 2011:150–151 (description, color painting). Schwartz 2012:35 (North Carolina). Vega-Cendejas and Santillana 2012:298 (abundance in lagoon and coastal habitats in Yucatán, Mexico). De Angelo et al. 2014:1042 (seagrass beds in eastern Gulf of Mexico). Snyder and Burgess 2016:100 (listed, Florida). Starck et al. 2017:91 (Florida Keys).

Misidentifications. As Chriodorus atherinoides off Charleston, SC, by Fahay (1975); specimen reidentified by G. E. McGowen (pers. comm.). Types. Hyporhamphus meeki Banford and Collette, 1993. Holotype USNM 294369 (1, 160 mm SL); Morehead City, NC; 5 September 1964. Dorsal fin rays 17; 114

Family Hemiramphidae

Figure 4.13. Meek’s Halfbeak, Hyporhamphus meeki. Holotype, USNM 294369, 160 mm SL. Collected by B. B. Collette, Morehead City, North Carolina, U.S.A., 5 September 1964. Illustration by M. H. Carrington.

than 0.70 in H. meeki, less than 0.70 in H. unifasciatus. Hyporhamphus meeki differs from H. collettei, in pectoral fin rays (H. meeki usually has 11–12, H. collettei almost always has 10; Table 4.8). Description. An elongate fish with a greatly prolonged beak-like lower jaw. Upper jaw short, scaly; preorbital ridge (bony ridge under nostril) present; ratio of preorbital length to orbit diameter usually greater than 0.70. Dorsal fin rays 12–17, usually 14 or 15; anal fin rays 14–18, usually 15–17; pectoral fins short, not reaching to nasal pit when folded forward and with 10–13 rays, usually 11 or 12 (Table 4.8). Caudal fin emarginate to slightly forked. Bases of dorsal and anal fins covered with scales. Vertebrae 31–35 precaudal + 16–19 caudal = 49–53 total. Predorsal

anal fin rays 17; pectoral fin rays 13–14. Gill rakers on first arch 6 + 17 = 23. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists False Silverstripe Halfbeak (English) and agujeta flaca (Spanish, Mexico). We do not follow the limitations of Page et al. (2013) on patronymics and prefer the logical “common” name derived from the scientific name, Hyporhamphus meeki, Meek’s Halfbeak (Collette 2003b). Diagnosis. Differs from its southern relative, Hyporhamphus unifasciatus, in having more first arch gill rakers (31–40, usually 33–36, rather than 27–35, usually 30–32; Table 4.9), a longer beak, and the ratio of preorbital length to orbit diameter is usually greater

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scales 34–39, usually 35–37. Gill rakers on first arch 8–12 + 20–29 = 31–40, usually 33–37 (Table 4.9); gill rakers on second arch 2–6 + 20–26 = 20–30, usually 25–28 (Table 4.13) Color. Greenish above, silvery white below; three distinct narrow black lines along from head to dorsal fin; fleshy tip of beak red; caudal fin pale, dark-edged. Size. The maximum observed size of Hyporhamphus meeki is 208 mm SL (MCZ 695, Key Biscayne, FL).

Kingston et al., 1969); and Ancyrocephalus trullae Williams, Dactylogyridae, Dauphin Island, AL (Williams, 1980). Bruce (1986) reported the cymothoid isopod Mothocya nana (Schioedte and Meinert) from specimens from the Chesapeake Bay, Georgia, and Florida. A specimen with a barnacle, Conchoderma virgatum, attached to the dorsal fin was seined near Round Island, Mississippi Sound (Dawson 1969). Food. Hyporhamphus meeki is largely an herbivore as an adult, feeding on algae and seagrasses plus a few small invertebrates. Linton (1905) found algae, sea lettuce, and a few young crustaceans in specimens from North Carolina. Tracy (1910) reported that the food of specimens in Rhode Island was almost exclusively algae. Hildebrand and Schroeder (1928) found vegetable matter, small crustaceans, and small mollusks in eight small specimens from Chesapeake Bay. Plants, detritus, polychaetes, amphipods, and crabs were found in specimens from two north Florida salt marshes (Subrahmanyam and Drake 1975). Carr and Adams (1973) found epiphytic algae and detritus (51%) and seagrass (49%) in 77 specimens (130–199 mm SL) from Crystal River, Florida. A 77 mm juvenile from this area had eaten mostly zooplankton (crab megalopa, veligers, copepods, and two insects). Reproduction. Eggs are attached to floating eelgrass blades (Zostera) or other plant material over vegetated habitats during summer months in Chesapeake Bay (Olney and Boehlert 1988).

Biology Predators. Hyporhamphus meeki has been recorded as eaten by Bluefish, Pomatomus saltatrix, at Fort Macon, NC (Yarrow 1877), and Ocean City, MD (Schwartz 1964); King Mackerel, Scomberomorus cavalla, present in 5 of 113 specimens, Onslow Bay, NC (De Vane 1978); Wahoo, Acanthocybium solandri (Manooch and Hogarth 1983); and Jaegers in the vicinity of Woods Hole, MA (Smith 1898). Parasites. Linton (1905) reported immature nematodes (Ascaris), cestodes (Scolex polymorphus), and larvae similar to Dibothrium in specimens from Beaufort, NC. There are several species of trematodes known from this halfbeak: Lobatostoma ringens (Linton), Aspidogastridae, from the intestine of specimens from Veracruz, Mexico (Caballero y Caballero and Bravo Hollis 1965; Hendrix and Overstreet 1977); Axinoides hyporhamphi Price (Price 1962; 116

Family Hemiramphidae

Egg diameter 2.0 mm, chorion with several very long attachment filaments (Hardy 1978). Length at hatching unknown, but smallest known specimen is 3.0 mm. Illustrations of 7.0 and 15.8 mm larvae, and of a 23.5 mm prejuvenile, were presented by Hardy and Johnson (1974). A large female (USNM 294394, 140 mm SL) collected from Maryland had 60 eggs in the left ovary (0.65–1.05 mm in diameter, x̅ 0.84 mm for 20 eggs) and 60 eggs in the right ovary (0.75–1.00 mm in diameter, x̅ 0.87 mm for 20 eggs). Range. Hyporhamphus meeki occurs on the Atlantic coast of the United States from Miami, FL, north to the Annapolis Basin, Nova Scotia (Gibson and Myers 2002), and Chamcook, New Brunswick, Canada (Leim and Day 1959), and on the Gulf coast from Everglades Park, FL, west to Galveston, TX. Hyporhamphus meeki and H. unifasciatus are sympatric on the east coast of Florida from St. Lucie Inlet south to Miami, on the west coast from Tampa Bay to Everglades Park, and on the Mexican Gulf of Mexico coast (Fig. 4.12). Hyporhamphus meeki is the only one of the two species known to occur north of St. Lucie Inlet and Tampa Bay. Records of H. unifasciatus (such as Castro-Aguirre et al. 1986) and H. roberti (such as Chávez-López et al. 2005) from the Mexican Gulf of Mexico could be based on either H. meeki or H. unifasciatus. Study Material. A total of 670 specimens (24.0–208 mm SL) originally at AMNH, ANSP, USNM, CAS,

CU, FMNH, FSU, FSBC, MCZ, TU, UMML, UMMZ, UF, VIMS, YPM, and the Texas Game and Fish Commission. The specimens came from five general areas: 207 specimens (24.0–187 mm SL) from the Atlantic coast of the United States from Massachusetts to Georgia; 44 specimens (27.0–208 mm SL) from the east coast of Florida; 200 specimens (47.0–170 mm SL) from the west coast of peninsular Florida; 204 specimens (34.0–207 mm SL) from the Florida panhandle to Texas; 15 specimens (64.2–152 mm SL) from Yucatán, Mexico. See Banford and Collette (1993) for full data on 338 specimens. Hyporhamphus mexicanus Alvarez, 1959 El Petén Halfbeak Figures 2.6, 4.14 Tables 4.1, 4.2 Hyporhamphus mexicanus Alvarez, 1959:71–73 (original description; Coatzacoalcos River, Oaxaca, Mexico). Miller 1966:777, 796, Usumacinta Province). Espinosa Pérez et al. 1993:32 (Coatzacoalcos River). Lara-Domínguez et al. 1993:364 (Gulf of Mexico). Contreras-Balderas et al. 1997:330 (marine beach east of mouth of Coatzacoalcos River). Collette 2003a:589, 2004:9 (fresh water, Guatemala and Mexico). Miller et al. 2005:205 (range, habitat, Mexico, fig. 6.238). Salgado-Maldonado 2006:217, 346 (acanthocephalan parasite, Tonalá River basin). Lozano-Vilano et al. 2007:46, 48 (Lacantún River, Chiapas, Mexico). González-Díaz et al. 2008:50 (La Venta River, Chiapas, Mexico). Miller et al. 2009:228 (range, habitat, fig. 6.238, map 6.204). Salgado-Maldonado 2008:42 (nematode parasite). Arévalo-Frías and Mendoza-Carranza 2012:258,

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table 3 (larvae and juveniles from the Pantanos de Centla Biosphere Reserve, Tabasco, Mexico).

Parasites. An intestinal acanthocephalan, Neoechinorhynchus golvani Salgado-Maldonado, family Neoechinorhynchidae, was reported from Hyporhamphus mexicanus from Lago El Rosario in the Tonalá River basin (Salgado-Maldonado 2006, 2008). Range. Hyporhamphus mexicanus is restricted to freshwater streams of Mexico and Guatemala draining into Campeche Bay, Gulf of Mexico (Fig. 2.6). The holotype is from the Coatzacoalcos River, which enters Campeche Bay at its southernmost point at Coatzacoalcos. One specimen was reported from “probably marine waters” from a beachside pool 1 km east of the mouth of the Coatzacoalcos River (Contreras-Balderas et al. 1997:330). Two Tulane University specimens are from the Nuevo Gonzalez River. All other known specimens are from the El Petén section of Guatemala, which is drained by the Usumacinta River that empties into Campeche Bay some 190 km east of Coatzacoalcos. Study Material. A total of 39 specimens (20.0–161 mm SL), including the holotype. Also from Mexico: TU 87566 (2, 161 mm SL); Tabasco, Nuevo Gonzalez River on the north side of Rt. 180, 26 miles southwest of Alvero Obregon; R.D. Suttkus 3014, 1 June 1961. AMNH 25204 (1, 88.7 mm SL); near mouth of Laguna del Ronpiro, Mexico; Rosen and party, 2 March 1966. AMNH 25570 (1, 81.5 mm SL); San Ramon River; R. M. Bailey and party, 27 March 1966. The rest of the study material is from the Usumacinta River drainage, El Petén, Guatemala: UMMZ 143517 (7, 83–147 mm SL); San Pedro de Martír River within 2 miles of El Paso de los Caballos; C. L. Hubbs and party, 10–14 March 1935. UMMZ 143519 (11, 20–126 mm SL); lowest league of Arroyo de Petexbatúm above Sayaxche; C. L. Hubbs and party, 22 April 1935. UMMZ 143520 (2, 96–151 mm SL); about 13 miles by river above Sayaxche; C. L. Hubbs and party, 4 April 1935. UMMZ 143516 (11, 28–133 mm SL); Pasión River at Tres Islas near southern border

Types. Hyporhamphus mexicanus Alvarez, 1959:71– 73. Holotype PNCP 403 (65.2 mm SL); Mexico, Oaxaca, Palomares, Rancho San Carlos, 24 km east of Palomares, 1 km upstream from the confluence of the Coatzacoalcos and Saribia rivers; J. Alvarez et al., 25 May 1959. Dorsal fin rays 14; anal fin rays 13; pectoral fin rays 11–11. Vertebrae 34 precaudal + 16 caudal = 50 total. Gill rakers on first arch 7 + 19 = 26; gill rakers on second arch 3 + 19 = 22. Head length 14.8 mm; lower jaw length 21.0 mm; P1–P2 is 25.3 mm, P2–C is 25.0 mm; P2–C extension falls just posterior to P1 base. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Mexican Halfbeak (English) and parajito mexicano (Spanish). However, because almost all specimens have come from Guatemala, we use the name El Petén Halfbeak. Diagnosis. A species of Hyporhamphus with very few scales on the dorsal and anal fins compared with H. meeki, H. collettei, and H. unifasciatus. Hyporhamphus mexicanus has very few gill rakers on the first arch (22–27) in comparison to H. roberti. It has few anal rays (13–15), a moderately long beak, and the pelvic fins are displaced far posteriorly so that distance from the pelvic origin to caudal base extended forward does not quite reach to the pectoral fin base. Description. Dorsal fin rays 14–15, usually 14; anal fin rays 13–15, usually 14; pectoral fin rays 11 (1 specimen of 24 with 12). Vertebrae 32–34 precaudal + 15–17 caudal = 48–51 total. Gill rakers on first arch 6–7 + 17–19 = 23–26; gill rakers on second arch 2–4 + 14–19 = 17–22. Several scales are present on the first few dorsal and anal fin membranes in large specimens. Median pore of preorbital canal on anterior part of canal; ventral pore enlarged. Size. The maximum observed size for Hyporhamphus mexicanus is 161 mm SL.

Figure 4.14. El Petén Halfbeak, Hyporhamphus mexicanus. UMMZ 143519, 126 mm SL. Collected by C. L. Hubbs in El Petén, Guatemala, 22 April 1935. Illustration by M. H. Carrington.

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Family Hemiramphidae of El Petén; C. L. Hubbs and party, 20 April 1935. UMMZ 143518 (1, 21 mm SL); Arroyo San Martín, tributary of Pasión River; C. L. Hubbs and party, 22 April 1935. UMMZ 143515 (2, 114–141 mm SL); Santa Amelia River, between Santa Amelia and Tres Islas; C. L. Hubbs and party, 4 April 1935. AMNH 18831 (1, 141 mm SL); Pasión River, Santa Amelia; B. Brown, March 1949.

1978:58 (southeastern Brazil), fig. 99. Román 1979:84 (listed; Venezuela). Cervigón 1991:223 (Venezuela). Cervigón et al. 1993:338 (northern coast of South America). Rodenas and López-Rojas 1993:75 (Laguna de Tacarigua, Venezuela). González Bencomo 1997:39–40 (juveniles, Strait of Lake Maracaibo, Venezuela). Carvalho Filho 1999:70 (Brazil). Menezes 2003:69 (Brazil). Miller et al. 2005:205–206 (range, habitat, Mexico, fig. 6.239). Giarrizzo et al. 2006:326 (weightlength relationships, Curucá mangrove estuary, northern Brazil). Giarrizzo and Krumme 2007:746–747 (Curuçá mangrove estuary, northern Brazil). González-Bencomo et al. 2008:401–403 (larvae and juveniles, La Palmita Bay, Lake Maracaibo, Venezuela). Neves, Teixeira, and Araújo 2011:121 (middle reaches, Mambucaba River, Rio de Janeiro state, Brazil). Neves, Teixeira, Franco et al. 2011:666 (middle and lower channels of the estuarine mixing zone of a tropical estuary of Mambucaba River, Rio de Janeiro state, Brazil). Santana et al. 2013:184 (Itamaracá Island, Pernambuco, Brazil). Hyporhamphus roberti roberti. Collette 2003b:1143 (range, figure). Collette 2004:10–11 (synonymy). Collette 2005:948–949 (early life history, figure). Lasso et al. 2004:144 (Orinoco delta). Mol 2012:655 (color photographs of 39 mm TL juvenile), 656 (early stages collected in vegetation at mid-river sand banks in the Suriname River. Mol et al. 2012:278 (Suriname rivers). Passos et al. 2012:230 (Paranaguá Estuarine complex, Brazil).

Hyporhamphus roberti (Valenciennes in Cuvier and Valenciennes, 1847) Pajarito Figures 4.15, 4.16 Tables 4.1, 4.2, 4.11, 4.12, 4.13, 4.14 Common Name. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Slender Halfbeak (English); however, because this species does not occur in the United States, we prefer Pajarito (Miller et al. 2005). The common name in Spanish is agujeta larga (Page et al. 2013). Diagnosis. Hyporhamphus roberti is a species of Hyporhamphus that lacks scales on the dorsal and anal fins (H. meeki, H. collettei, and H. unifasciatus all have scales on the fins) and have a moderate to high number of gill rakers on the first arch; few pectoral fin rays (usually 9 or 10); and a long beak. Two subspecies are recognized, the southern H. roberti roberti and the Central American H. r. hildebrandi. Range. Hyporhamphus roberti is marine and estuarine along the coasts of Central and South America (Fig. 4.16).

Misidentifications. As Hyporhamphus unifasciatus, Miranda Ribeiro 1961a:8 (MNRJ 5378, S. Mateus). Types. Hemiramphus roberti Valenciennes in Cuvier and Valenciennes, 1847. Lectotype MNHN 5634 (1, 113 mm SL); Cayenne, designated in Collette et al. (1997:19). Other syntypes from Cayenne are MNHN 3204 (1, 116 mm SL) and MNHN B.1073 (2, 86–91 mm SL). Both specimens in MNHN B.1073 have a low number of gill rakers (31, 32) and a slightly shorter beak than H. roberti roberti usually does. The smaller specimen has 12 pectoral rays and could be a specimen of H. unifasciatus; its poor condition does not permit accurate determination. MNHN 3204 has a slightly longer beak than normal and more dorsal and anal rays (15–16) and gill rakers (35) than H. r. roberti usually does; it could easily be mistaken for a specimen of H. r. hildebrandi. Hyporhamphus kronei Mirando Ribeiro, 1915. Holotype MNRJ 1970 (134 mm SL); Brazil, Iguape, São Paulo; Dr. R. Krone, 2 February 1966. Dorsal, anal, and pectoral fin rays uncountable. Gill rakers on first arch 10 + 25 = 35; lower jaw length 35.6 mm; P1–P2 53.5 mm; P2–C extension falls anterior to P1 base. Specimen badly dried and shrunken in 1966. Hyporhamphus salvatoris Fowler and Bean, 1923. Holotype USNM 83286 (137 mm SL); Brazil; Wilkes Expedition. Dorsal fin rays 15; anal fin rays about 15; pectoral fin rays about 12. Vertebrae 33 precaudal + 17 caudal = 50. Gill rakers on first arch 12 + 25 = 37; gill rakers on second arch 4 + 24= 28. P1–P2 is 50.5 mm; P2–C is 54.5 mm; P2–C extension falls on rear of opercle.

Hyporhamphus roberti roberti (Valenciennes in Cuvier and Valenciennes, 1847) Hemiramphus Roberti Valenciennes in Cuvier and Valenciennes, 1847:24–25 (original description; Cayenne, French Guiana). Gilbert 1900:164 (Pernambuco, Brazil). Fowler 1941b:142 (reference to Gilbert 1900). Collette et al. 1997:19 (types). Hyporhamphus kronei Miranda Ribeiro, 1915:19 (unnumbered) (original description, Iguape, São Paulo, Brazil). Fowler and Bean 1923:9–10 (comparison with H. salvatoris). Fowler 1940c:757 (listed, Rio de Janeiro, Brazil). Fowler 1941b:142 (listed). Miranda Ribeiro 1953:398–399 (holotype listed as no. 1970). Miranda Ribeiro 1961a:9 (listed). Hyporhamphus salvatoris Fowler and Bean, 1923:9–10 (original description; Rio de Janeiro, Brazil; USNM 83286). Fowler 1940c:757, 1941b:142, figs. 28, 1950:11 (refers to original description). Collette et al. 1992:10 (holotype). Hyporhamphus roberti. Jordan et al. 1930:198 (listed). Puyo 1936:172–174 (description, ecology, food value; French Guiana), fig. 34H. Fowler 1941b:142 (Brazil). Puyo 1949:160–161 (based on Puyo 1936), fig. 82. Schultz 1949:80–81 (50 USNM specimens, Lago de Maracaibo, Venezuela). Roux 1963 (type MNHN 5634). Mago Leccia 1970:89 (listed; Venezuela). Shiino 1976:91 (common name pajarito). Figueiredo and Menezes

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Fishes of the Western North Atlantic Diagnosis. Hyporhamphus roberti roberti differs from H. r. hildebrandi primarily in having fewer gill rakers on the first arch (usually 35–37 rather than 39–42; Table 4.13). About 92% of the specimens examined can be distinguished with this character, using a line of separation between 38 and 39. Hyporhamphus r. roberti also has fewer gill rakers than H. r. hildebrandi on the second arch (usually 28–30 rather than 31–33). About 88% of the H. roberti examined can be separated to subspecies by drawing a line between 30 and 31 second arch gill rakers (Table 4.14). Hyporhamphus r. roberti has fewer vertebrae than H. r. hildebrandi (usually 48–49 rather than 51–52; Table 4.12). About 98% of the specimens examined can be separated by drawing a line between 49 and 50 vertebrae. Hyporhamphus r. roberti usually has 14 or 15 dorsal fin rays and 15 anal fin rays; H. r. hildebrandi usually has 15 dorsal fin rays and 16 anal fin rays. About 74% of the specimens examined can be separated by plotting the number of dorsal rays against the number of anal rays, specimen by specimen. Another way of expressing this difference is to add the dorsal and anal rays of each specimen together. By this method, about 75% of the specimens examined can be separated by a line between 30 and 31 total rays. Most H. r. roberti have 30 or fewer (x̅ 29.7), whereas most H. r. hildebrandi have 31 or more (x̅ 30.9). Also, H. r. roberti has a shorter beak than H. r. hildebrandi. Description. Dorsal fin rays 13–16, usually 14 or 15; anal fin rays 14–17, usually 15; dorsal plus anal rays 28–32, usually 30 or less; pectoral fin rays 9–11, usually 10 (Table 4.11). Vertebrae 31–33, usually 32, precaudal + 16–18, usually 17, caudal = 47–50 total (Table 4.12). Gill rakers on the first arch 8–12 + 21–29 = 29–41 (Table 4.13); gill rakers on the second arch 3–6 + 19–28 = 23–33 (Table 4.14). Size. The maximum observed size for Hyporhamphus roberti roberti is 140 mm SL (USNM 121817, Lake Maracaibo). Length-weight information was provided for 94 specimens from the Curucá Estuary, northern Brazil, by Giarizzo et al. (2006).

From Venezuela, Lago de Maracaibo, L. P. Schultz: USNM 121816 (1, 105 mm SL); Los Monitos, Río Limón, 11 March 1942. USNM 121817 (12, 85.5–140 mm SL); Maracaibo Yacht Club, 5 March 1942. USNM 121818 (1, 56.5 mm SL); Maracaibo Yacht Club, 16 May 1942. USNM 121819 (1, 96 mm SL); 7 km south of Maracaibo, 6 March 1942. USNM 121820 (33, 51.5– 103 mm SL); 1 km off Pueblo Viejo, 7–8 April 1942. USNM 121725 (2, 107–121 mm SL); Maracaibo Yacht Club, 27 February 1942. From Trinidad, FMNH 3769 (9, 72.2–100 mm SL); O. Riddle. From the Guianas, AMNH 13433 (1, 82.2 mm SL); Essequibo River, Guyana; Pinkus, 1933. MNHN 5634 (1, 113 mm SL), lectotype of Hemiramphus roberti; Cayenne, French Guiana. MNHN 3204 (1, 116 mm SL), paralectotype of H. roberti; Cayenne, French Guiana; Poiteau. MNHN B.1073 (2, 85.5–90.5 mm SL), paralectotypes of H. roberti; Cayenne, French Guiana; Robert. ZMA 105.042 (1, 125 mm SL); Suriname, Maroni River, 25 km south of Albina; H. Nijssen, 17 June 1966. From Brazil, 101 specimens (38–137 mm SL): CAS-SU 51786 (1, 88.3 mm SL); Fortaleza. MCZ 678 (1, 104 mm SL); Pernambuco (= Recife). MCZ 4685 (1, 80.5 mm SL); Thayer Expedition, Pernambuco (= Recife); Agassiz and Bourget. FMNH 3849 (1, 108 mm SL); Pernambuco (= Recife); Greely, 1899. FMNH 92319 (1, 111 mm SL); Bay of Bahia; J. D. Haseman 1873, 13 April 1908. CAS-SU 51784 (2, 69.5–78.6 mm SL); Carapetal. MACN 1630 (1, 125 mm SL); Jatoba River, San Francisco. MNRJ 5378 (2, 128–133 mm SL); Lagoa Juparana, Linhares, Estado Espirito Santo. ANSP 121205 (2, 68.7–69.1 mm SL); Ilha da Conociencia, Atafona, R.J.; J. Saskin et al., July–August 1963. USNM 83286 (1, 137 mm SL), holotype of Hyporhamphus salvatoris Fowler and Bean; Rio de Janeiro; Wilkes Expedition. FMNH 76973 (1, 112 mm SL) and FMNH 76988 (1, 133 mm SL); Santos; 3 July 1908. FMNH 76984 (1, 93.2 mm SL); mouth of Rio Caite; 1 January 1910. MNRJ 8702 (1, 81.0 mm SL); Surví, Praia de San Francisco, Rio de Janeiro. MNRJ 1970 (1, 134 mm SL), holotype of Hyporhamphus kronei A. Miranda Ribeiro; Iguape, Estado São Paulo; R. Krone. MZUSP 13382 (1, 104 mm SL); São Paulo, Cananeia; Clarimundo de Jesus, 5 November 1952. MZUSP 67775 (10, 107–128 mm SL); São Paulo, Cananeia; 11 May 1982. MZUSP 102659 (1, 92 mm SL); Bahia, Ilheus, Lago Encantada, na Vila Formosa, 14°37′12″S, 39°08′30″W; S.M.O. Lima et al., 13 February 2009. MZUSP 3669 (20, 54.5–96 mm SL); Rio de Janeiro, Angra dos Reis; L. Travassos, 16 July 1945. MZUSP 67763 (12, 38–88 mm SL); Rio de Janeiro, Angra dos Reis; L. Travassos, 29 June 1945. MZUSP 67764 (40,

Biology Range. This subspecies is known from Lake Maracaibo, Venezuela, Trinidad, the Guianas, and Brazil south to Iguape, Estado São Paulo, south of Rio de Janeiro (Fig. 4.16). All records of Hyporhamphus roberti from the West Indies or continental United States refer to either H. unifasciatus or H. meeki. Study Material. A total of 97 specimens (38–140 mm SL) from four areas: 47 specimens 56.5–140 mm SL) from Venezuela, and 101 specimens from Trinidad, Guianas, and Brazil (38–137 mm SL). 120

Family Hemiramphidae

121

Fishes of the Western North Atlantic

61–98 mm SL); Rio de Janeiro, Angra dos Reis; L. Travassos, 1945.

2003b:1143 (range, figure). Collette 2005:948–949 (early life history, figure). Smith et al. 2003:15 (Belize). Collette 2004:10– 11 (synonymy). Matamoros et al. 2007 (inland reservoir, Honduras).

Hyporhamphus roberti hildebrandi Jordan and Evermann, 1927

Types. Hyporhamphus hildebrandi Jordan and Evermann, 1927. Lectotype FMNH 8260, designated by Weed (1933:61); Toro Point, Fox Bay, Colon, Panama. Paralectotypes, 10 lots at FMNH and USNM. Diagnosis. Hyporhamphus roberti hildebrandi differs from H. r. roberti in having more gill rakers on the first arch (usually 39–42 rather than 35–37; Table 4.13). About 92% of the specimens examined can be distinguished with this character, using a line of separation between 38 and 39. Hyporhamphus r. hildebrandi also has more gill rakers than H. r. roberti on the second arch (usually 31–33 rather than 28–30). About 88% of the H. roberti examined can be separated to subspecies by a line drawn between 30 and 31 second arch gill rakers (Table 4.14). Hyporhamphus r. hildebrandi has more vertebrae than H. r. roberti (usually 51–52 rather than 48–49; Table 4.12). About 98% of the specimens examined can be separated by a line between 49 and 50 vertebrae. Hyporhamphus r. hildebrandi usually has 15 dorsal fin rays and 16 anal fin rays.; H. r. roberti usually has 14 or 15 dorsal fin

Hyporhamphus roberti. Meek and Hildebrand 1923:239 (description; Toro Point and Fox Bay, Colon, Canal Zone), pl. 16, fig. 2. Miller 1945a:192 (comparison with species of Hyporhamphus from the Americas). Miller 1945b:235 (H. hildebrandi synonymized under H. roberti). Caldwell et al. 1959:17–18 (18 UF specimens, first record from Costa Rica; report of specimens collected by Miller in Guatemala). McEachran and Fechhelm 1998:961, figure (Gulf of Mexico). Schmitter-Soto 1998:80–82 (description, Quintana Roo, Mexico), fig. 37 (photograph upside down). Miller et al. 2005:205–206 (range, habitat, fig. 6.239, map 6.205). Miller et al. 2009:228–229 (range, habitat, fig. 6.239, map 6.205). Hyporhamphus hildebrandi Jordan and Evermann, 1927:503–504 (original description based on Meek and Hildebrand 1923; type specimens collected by Meek and Hildebrand at Toro Point, Fox Bay, Colon, Panama). Weed 1933:60–62 (FMNH 8260 designated lectotype of H. hildebrandi, fig. 2 (dorsal view of head). Grey 1947:159 (types). Shiino 1976:91 (common name pajarito). Ibarra and Stewart 1987:49 (types). Collette et al. 1992:9 (types). Hyporhamphus roberti hildebrandi. Miller 1966:796 (Río Sarstún to Panama; B. B. Collette, pers. comm.). Gilbert and Kelso 1971:27 (Tortuguero, Costa Rica). Greenfield and Thomerson 1997:91–92, 262 (description, Belize), fig. 96. Collette

Figure 4.15. Pajarito, Hyporhamphus roberti hildebrandi. UF 7174, 169 mm SL. Tortuguero Lagoon, Costa Rica, 11 July 1958. Illustration by M. H. Carrington.

122

Family Hemiramphidae rays and 15 anal fin rays. About 74% of the specimens examined can be separated by plotting the number of dorsal rays against the number of anal rays, specimen by specimen. Another way of expressing this difference is to add the dorsal and anal rays of each specimen together. By this method, about 75% of the specimens examined can be separated by a line between 30 and 31 total rays. Most H. r. hildebrandi have 31 or more (x̅  30.9), whereas most H. r. roberti have fewer than 30 (x̅  29.7). Also, H. r. hildebrandi has a longer beak than H. r. roberti. Description. Dorsal fin rays 14–16, usually 15; anal fin rays 14–17, usually 16; pectoral fin rays 8–11, usually 10 (Table 4.11). Vertebrae 32–35 precaudal + 16–18 caudal = 49–53 total (Table 4.12). Gill rakers on first arch 9–14 + 26–34 = 38–48 (Table 4.13); gill rakers on second arch 3–7 + 24–33 = 29–38 (Table 4.14). Size. The maximum observed size for Hypo­ rhamphus roberti hildebrandi is 186 mm SL (UF 7174, Tortuguero, Costa Rica). Biology

Figure 4.16. Distribution of Pajarito, Hyporhamphus roberti roberti and H. roberti hildebrandi, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

Reproduction. A large female (USNM 114315, 131 mm SL) collected from Honduras had 20 eggs in the left ovary (0.40–0.75 mm in diameter, x̅  0.56 for 20 eggs) and 20 eggs in the right ovary (0.35–0.75 mm in diameter, x̅  0.54 mm for 20 eggs). Range. Hyporhamphus roberti hildebrandi is known from Quintana Roo, Mexico (Schmitter-Soto 1998), and Belize (FMNH 50031) southeast along the coast of Central America to the Gulf of Uraba, Colombia (Fig. 4.16). The report of H. roberti from Alvarado Lagoon Estuary, Veracruz, Mexico, is probably based on specimens of either H. meeki or H. unifasciatus. The record of H. hildebrandi from Old Providence Island off the coast of Nicaragua (Fowler 1944, 1953; Garzón F. and Acero P. 1983) is based on two small specimens of H. unifasciatus (ANSP 72753). Three specimens were collected along the shore of the Francisco Morazán El Cajón Reservoir in Honduras (Matamoros et al. 2007). There is a gap between the Gulf of Uraba and Lake Maracaibo, Venezuela, where the nominal subspecies occurs. Additional collecting is needed to determine which form occupies the intervening area. All references to H. hildebrandi from Bermuda refer to H. collettei and those from the continental United States refer either to H. unifasciatus or H. meeki. Study Material. A total of 125 specimens (35.5–186 mm SL) from 31 collections, including 20 paralectotypes. The material (from six countries) is listed from Belize south to Colombia. From Belize: FMNH 50031 (2, 53.0–60.3 mm SL); Belize; B. H. Bailey. FMNH 77713 (3, 81.4–84.9

mm SL); Belize City, St. Johns College Bench; D. W. Greenfield, T. A. Greenfield, G73-25, 3 July 1973. From Guatemala: USNM 114261 (12, 35.5–90.5 mm SL); Lake Yzabal; R. R. Miller, 24–27 April 1947. USNM 134705 (5, 80.5–103 mm SL); Lake Yzabal; R. R. Miller. USNM 134706 (1, 112 mm SL); El Estor; Holloway. USNM 114315 (10, 96.5–137 mm SL); Río Sarstún; 29–30 April 1947. AMNH 35043 (1, 79.8 mm SL); Izabal state; Río Dulce at mouth of Río Cienaga; R. M. Bailey et al., 9 April 1974. From Honduras: LACM 31009-6 (1, 165 mm SL); Colon, bayou of unnamed river 30 miles east of Trujillo; C. Swift, J. Rudloe, A. Rogorin, 16 August 1969. FMNH 84984 (1, 133 mm SL); tributary of Río Patuca; R. R. Miller et al., RRM 75-24, 9 May 1975. USM 31216 (3, 120–124 mm SL); Francisco Morazán El Cajón reservoir; 1 January 2003. From Costa Rica and Tortuguero: UF 7174 (14, 85.0–186 mm SL) and USNM 204220; L. H. Ogren, 11 July 1958. UF 9268 (5, 133–160 mm SL); 8 September 1959. UF 5375 (1, 116 mm SL); 24 July 1957. UF 7185 (1, 68.5 mm SL); 3 September 1958. BMNH 1939.7.10.15 (1, 112 mm SL); Port Parker; Sopwith. UF 16438 (5, 80.8–84.1 mm SL); D. D. Kelso et al., 5 May 1964. From Panama, paralectotypes from Fox Bay, Colon, 1911–1912: FMNH 8261 (1, 92.0 mm SL). FMNH 17408 (1, 72.0 mm SL). FMNH 17407 (1, 79.2 mm SL). FMNH 8262 (1, 85.5 mm SL). FMNH 8263 123

Fishes of the Western North Atlantic (1, 75.0 mm SL). FMNH 17406 (1, 59.0 mm SL). USNM 79664 (2, 99.0–100 mm SL). USNM 79663 (2, 58.5– 86.0 mm SL). USNM 82111 (1, 104 mm SL). USNM 79662 (2, 87.5–91.0 mm SL). Also from Panama: YPM ICH 003288 (2, 69.5–88.5 mm SL); Limon Bay; 12 February 1934. USNM 128519 (5, 51.5–94.5 mm SL); Fort Sherman. MCZ 43075 (5, 61.5–79.9 mm SL); Canal Zone; I. Rubinoff. From Colombia, Gulf of Uraba: UMML 24592 (7, 56.1–87.5 mm SL); 08°17′N, 76°49.5′W; Pillsbury 354, 11 July 1966. USNM 203825 (25, 61.7–93.5 mm SL); 08°17.5′N, 76°51.9′W; 11 July 1966.

1924:189 (Montevideo, Uruguay). Breder 1927:19 (Bahamas). Devincenzi and Barattini 1928: pl. 18, figs. 2, 3 (Uruguay). Beebe and Tee-Van 1928:69–71 (description, development; Haiti). Breder 1929:280, 1932a:20 (Tortugas). Jordan et al. 1930:198 (listed). Pozzi and Bordale 1935:157 (35°30′S, Argentina). Beebe 1936:204 (in stomach of Thunnus albacares, St. Lucia). Fowler 1941b:142 (Brazil). Herre 1942:291 (description, Antig u a). M i l le r 194 5a:187 (d i ag nos t ic c h a r ac t e r s of Hyporhamphus). Breder 1948:91 (description; range in part, northern records refer to H. meeki). Schultz 1949:81 (Venezuela; USNM 123283). De Buen 1950:92 (synonymy, references from Uruguay). Fowler 1952b:87 (description, Hispaniola; ANSP 73315, 77221). Santos 1952:93 (range). Fowler 1953:52 (Caribbean Colombia). Martin 1956:93 (description, Los Roques Island, Venezuela). Schultz 1949:81 (Estanques Bay, Venezuela; USNM 12385). Siddiqi and Cable 1960:274 (digenetic trematode Schikhobalotrema acuta found in intestine; Puerto Rico). Miranda Ribeiro 1961a:8 (Brazil, in part; NMRJ 3134). Tabb et al. 1962:51 (Coot Bay, Florida Bay). Nomura and Menezes 1963:355 (range, omnivore; Brazil). Roux 1963:399 (type of Hemiramphus richardi). Caldwell and Caldwell 1964:12–13 (Panama, Venezuela). Bane 1965:56 (common tuna bait species, Mona Passage). Cervigón 1965:26 (marao blanco, Venezuela). Caldwell 1966:32 (Jamaica). Cervigón 1966:221–222 (description, Venezuela). Böhlke and Chaplin 1968:126 (description, Bahamas). Mago Leccia 1970:89 (Venezuela). Austin 1971:32 (abundant, mangroves, western Puerto Rico). Dahl 1971:182 (Colombia). De Buen 1972:162 (South America). Hinegardner and Rosen 1972:639 (cellular DNA). Oliveira 1972:37, 40 (not found at salinities under 18‰, northeastern Brazil). Ali et al. 1973:102–103, 107, 111 (retinal adaptations; Brazil). Duarte-Bello and Buesa 1973:80 (synonymy, range; Cuba). Eskinazi 1972:287, 292 (temperature 25–31 °C, salinity 17‰–34‰; canal 50 km north of Recife, Brazil). Roux 1973:72 (description; Brazil). Guitart 1975:252 (Cuba, synonymy, description, fig. 187). Instituto Nacional de Pesca 1976:75 (range, in part). Erdman 1976:23, 1977:159 (northeastern Caribbean). Oliveira 1976:10, 12, 14, 16 (salinity; Jaguaribe River estuary, Ceará, Brazil). Shiino 1976:91 (common names). Castro-Aguirre 1978:58–59 (Atlantic Mexican records). Collette 1978 (synonymy, description, range) figure. Figueiredo and Menezes 1978:57–58 (description; southeastern Brazil), fig. 98. Lima and Oliveira 1978:1, 20 (common name agulhabranca, Ceará and Pernambuco, Brazil). de Sylva and Higman 1979:153 (occasionally ciguateric). Román 1979:84 (marao blanco, in key; Venezuela), figure. Amezcua-Linares and Yáñez-Arancibia 1980:85, 87, 89 (ecology; Laguna de Terminos, Campeche, Mexico). Álvarez Cadena and Flores-Coto 1981:20 (larvae; Laguna de Terminos, Campeche, Mexico), fig. 5 (9.0 mm larva). Lucena and Lucena 1981:41 (Porto Belo, Florianopolis, and Garepaba, Santa Caterina, Brazil, and Torres and Rio Grande, Rio Grande do Sul, Brazil). Flores-Coto and Álvarez- Cadena 1982:74 (ichthyoplankton, Laguna de Terminos, Campeche, Mexico). Reséndez Medina 1982:394 (Laguna de Zontecompanan, Veracruz, Mexico). Rodríguez et al. 1984:14 (Cuba). Vasconcelos Filho et al. 1984:239–241, table 3, diet, Brazil). Perez and Rylander 1985:643 (hemoglobin heterogeneity; Venezuela). Bruce 1986:1099 (parasitic isopod Mothocya argenosa on Hyporhamphus unifasciatus from Cuba and the Virgin Islands), 1112 (M. nana, Panama), 1113 (M. bermudensis, Haiti), 1127 (M. omidaptria, Brazil and Curaçao). Robins and Ray 1986:102 (description, range in part), pl. 17. Acero P. and Garzón F. 1987:90 (Colombia). Gilmore 1987:129 (seagrass communities). Paiva Filho and Toscano 1987:157 (São Paulo, Brazil). Rodríguez and Valdés 1987:95 (Cuba).

Hyporhamphus unifasciatus (Ranzani, 1842) Atlantic Silverstripe Halfbeak Figures 4.12, 4.17 Tables 4.1, 4.2, 4.8, 4.9, 4.10 Hemirhamphus unifasciatus Ranzani, 1842:326 (original description; Brazil). Jordan 1884a:112–113 (description, abundant; Key West, FL; USNM 34999). Ihering 1897:46–47 (description; Rio Grande do Sul, Brazil). Hemiramphus Richardi Valenciennes in Cuvier and Valenciennes, 1847:26–28 (original description; St. Croix; Cayenne, French Guiana; Bahia, Rio de Janeiro, Brazil). Collette et al. 1997:19 (types). Hyporhamphus tricuspidatus Gill, 1859a:131 (original description; Barbados). Collette et al. 1992:10 (holotype). Hemirhamphus fasciatus Poey, 1860:299–300 (original description; Cuba). Günther 1866:262 (H. poeyi, replacement name for H. fasciatus). Howell Rivero 1938:180 (MCZ 32934 is the co-type of H. fasciatus). Hemirhamphus poeyi Günther, 1866:262 (replacement name for H. fasciatus Poey, preoccupied by H. fasciatus Bleeker). Poey 1868:383 (description, Cuba). Jordan and Gilbert 1882b:373 (Panama). Hemiramphus unifasciatus. Jordan 1886b:35 (Cuba). Jordan 1887b:568 (West Indies). Jordan 1890:6 (St. Lucia). Jordan 1891:315 (Bahia, Brazil). Perugia 1890–1891:654 (Montevideo, Uruguay). Cockerell 1892:14 (Jamaica). Berg 1895:25–26 (Montevideo, Uruguay). Posada Arango 1909:297 (Colombia). Devincenzi 1920:101, 106 (after Perugia 1890–1891 and Berg 1895). Fowler 1928:463 (Puerto Rico, Guanica; ANSP 2364553). Fowler 1931:396 (Trinidad; ANSP 76397, 93350). Hyporhamphus unifasciatus. Jordan and Evermann 1896a:321 (Key West, FL, to Rio de Janeiro, Brazil). Jordan and Rutter 1897:97 (Jamaica). Evermann and Kendall 1899:62 (in part; Florida records). Fowler 1899:118 (Port Antonio, Jamaica; ANSP 19989). Gilbert 1900:164 (Maceio, Brazil). Evermann and Marsh 1902:101 (Puerto Rico, fig. 18). Jordan and Thompson 1905:234 (Tortugas). Starks 1913:40 (genus misspelled Hyporhynchus; Natal, Brazil). Cockerell 1915:157 (scales from Balboa, Panama specimens differ from scales of Woods Hole, MA, H. roberti [= H. meeki]). Fowler 1915:151 (Trinidad, Port­ of‑Spain; ANSP 45043). Miranda R ibeiro 1915:18–19 (description; south to Rio de Janeiro). Fowler 1916a:401 (Colon, Panama; ANSP 452178). Miranda Ribeiro 1918:763 (Rio de Janeiro). Fowler 1919a:6 (West Indies; Panama). Fowler 1919b:137 (St. Martins, West Indies; ANSP 75215). Nichols 1921:22 (Turks Island, Bahamas). Meek and Hildebrand 1923:237–239 (description; Panama), pl. 16, fig. 1. Devincenzi

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Family Hemiramphidae Thayer et al. 1987:30 (few found in seagrasses near red mangrove prop root habitat, southern Florida). Alvarez-Guillen et al. 1988:320 (Quintano Roo, Mexico). Gomez-Soto and Contreras-Balderas 1988 (in part, Tamaulipas, Mexico). Sogard et al. 1989a:58, 1989b:195 (Florida Bay). Thayer and Chester 1989:210 (Florida Bay). Cervigón 1991:221–222 (Venezuela, fig. 167). Nolf and Stringer 1992:70, pl. 12, fig. 17–18 (otoliths from off Acapulco, Mexico, and Margarita Island, Venezuela). Cervigón et al. 1993:337 (northern coast of South America). Begossi and Figueiredo 1995:714 (local name in southern Brazil is peixe-agulha). Giannini and Paivo Filho 1995:144 (São Paulo, Brazil). Contreras-Balderas et al. 1997 (Mexican Gulf of Mexico). Greenfield and Thomerson 1997:92, 262 (description, Belize), fig. 95. Smith 1997:386 (description, range). McEachran and Fechhelm 1998:962 (Gulf of Mexico). Carvalho Filho 1999:70 (Brazil). Arãújo, Teixeira et al. 2000:140 (estuaries, Ceará, Brazil). Arãújo, Cunha et al. 2000:135 (sandstone reefs, Ceará, Brazil). Loftus 2000:34 (Everglades National Park, FL). Pérez-Hernández and Torres-Orozco 2000:434 (Mexican coastal lagoons). Schmitter-Soto et al. 2000:154 (Caribbean Mexico). Vendel et al. 2000:180 (tidal flat, Paraná, Brazil). Claro and Parenti 2001:43 (Cuba, listed). Castillo-Rivera et al. 2002:176 (Pueblo Viejo Lagoon, Veracruz, Mexico). Collette 2003b:1142 (description, range, figure). Menezes 2003:69 (Brazil). Sanvicente-Añorve et al. 2003:373 (larvae from Campechén-La Ría estuary, Gulf of Mexico). Smith et al. 2003:15 (Belize). Collette 2004:12 (synonymy). Poulakis et al. 2004:126 (Charlotte Harbor, FL). Spach et al. 2004:49, 51, 55 (ecology, Brazil). Collette 2005:950–951 (early life history, figure). Bouchon-Navaro et al. 2006:323 (weightlength relationships, seagrass beds of Guadeloupe and Martinique). Powell et al. 2007:64–68, 209 (densities and monthly length-frequencies, Florida Bay). Nóbrega and Lessa 2007:68 (artisanal fishery, northeastern Brazil). Félix et al. 2007:287 (beach, Paraná, Brazil). Campbell et al. 2008:83–84 (juveniles, mangrove habitat, Quintana Roo, Mexico). Medina and Yunda 2008:61 (beaches, Santa Marta, Caribbean Colombia). García-Hernández et al. 2009:92 (juveniles, northern Yucatán Peninsula). Gómez Gaspar and Hernández Avila 2009:15 (Isla Cubagua, Venezuela). McEachran 2009:1286 (Gulf of Mexico). Cousseau et al. 2010:29 (Argentina, between 34°S and 55°S). Reis-Filho et al. 2010:305 (Paraguacu River, Todos os Santos Bay, Bahia, Brazil). Contente et al. 2011:43 (Guaraguaçu River, tributary of Paranaguá Bay Estuary, southern Brazil). Gondolo et al. 2011:187 (surf zone, Itamambuca Beach, Ubatuba, São Paulo, Brazil). Kells and Carpenter 2011:150–151 (description, color painting). Neves, Teixeira,

and Araújo 2011:121 (middle reaches, Mambucaba River, Rio de Janeiro state, Brazil). Peralta-Meixueiro and Vega-Cendejas 2011:677 (hyperhaline coastal system: Ría Lagartos, Mexico). Vilar et al. 2011:45 (Baia da Babitonga, southern Brazil). Neves, Teixeira, Franco et al. 2011:666–670 (lagoon, middle and lower channels of the estuarine mixing zone of a tropical estuary of the Mambucaba River, Rio de Janeiro state, Brazil). Gallardo-Torres et al. 2012:280 (present for five to nine months in the coastal lagoon Boca de la Carbonera Yucatán, Mexico). Vega-Cendejas and Santillana 2012:298 (abundance in lagoon and coastal habitats in Yucatán, Mexico). Del MoralFlores et al. 2013:832 (Veracruz Coral Reef System). De Angelo et al. 2014:1044 (seagrass beds in eastern Gulf of Mexico). Gallardo Torres et al. 2014:85 (Yucatán, Mexico, photographs). Smith-Vaniz and Jelks 2014:32 (St. Croix, U.S. Virgin Islands records). Snyder and Burgess 2016:100 (listed, Florida). Hemirhamphus roberti (not of Valenciennes in Cuvier and Valenciennes 1847). Henshall 1891:375 (Cape Sable, western Florida). Hyporhamphus roberti (not of Valenciennes in Cuvier and Valenciennes 1847). Fowler 1919a:7 (Key West, FL, ANSP 165389). Fowler 1951a:26 (St. Croix). Hiporhamphis [sic] unifasciatus. Kobelkowsky Díaz 1985:155 (Laguna de Tampamachoco, Veracruz, Mexico). Hyporamphus [sic] unifasciatus. Louis et al. 1992:296, 298 (Martinique). Hyporhamphus aff. unifasciatus. Nolf and Stringer 1992:70, pl. 12, fig. 19 (otolith from Rio Gurabo, Cercado Formation, Late Miocene of the Dominican Republic). Nolf and Aguilera 1998:238 (otoliths from the Cantaure Formation, Early Miocene of Venezuela).

Misidentifications. As Hyporhamphus hildebrandi from Old Providence Island, Caribbean Sea (Fowler 1944, 1953) and based on Fowler (Garzón F. and Acero P. 1983), ANSP 72753 (2, 35.4–41.0 mm SL). Types. Hemiramphus unifasciatus Ranzani, 1842. Holotype MZUB (145 mm SL); Brazil. Dorsal fin rays about 14; anal fin rays about 15. Gill rakers on first arch at least 8 + 21 = 29; head length 31.1 mm; lower jaw length 31.3 mm; P1–P2 is 52.8 mm; P2–C is 62.3 mm; P2–C extension falls on posterior third of preopercle. Hemiramphus Richardi Valenciennes in Cuvier and

Figure 4.17. Atlantic Silverstripe Halfbeak, Hyporhamphus unifasciatus. USNM 79666, 164 mm SL. Collected by S. E. Meek and S. F. Hildebrand in Fox Bay, Colon, Panama, 27 January 1912. Illustration by M. H. Carrington.

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Fishes of the Western North Atlantic 2–6 + 17–23 = 19–28 (Table 4.10). Upper jaw short, scaly; preorbital ridge (bony ridge under nostril) present; ratio of preorbital length to orbit diameter usually less than 0.70. Pectoral fins short, not reaching to nasal pit when folded forward; caudal fin emarginate to slightly forked. Color. Greenish above, silvery white below. Three distinct narrow black lines along from head to dorsal fin; fleshy tip of beak red; caudal fin pale, dark-edged. Size. The maximum observed size for Hyporhamphus unifasciatus is 240 mm SL (MCZ 4684, Rio de Janeiro, Brazil); about 270 mm TL, commonly to 200 mm SL. Length-weight information was provided for 201 specimens from the Lesser Antilles by Bouchon-Navaro et al. (2006).

Valenciennes, 1847. Lectotype designated in Collette et al. (1997:19), MNHN B.1072 (1, 144 mm SL); St. Croix; Richard. Paralectotypes MNHN 4341 (1, 174 mm SL), Brazil, B.1070 (2, 150–155 mm SL), Cayenne, and B.1071 (1, 154 mm SL), Rio de Janeiro. MNHN B.1072 was designated as lectotype because it is the first specimen mentioned in the original description; it is the only syntype collected by Richard, after whom the species was named; it adequately fits both the original description and our concept of Hyporhamphus unifasciatus; and it is in good condition. The lectotype has the following characters: dorsal fin rays 14; anal fin rays 15; pectoral fin rays 11–11. Gill rakers on first arch 8 + 21 = 29; gill rakers on second arch 4 + 21 = 25. Lower jaw length 30.2 mm; head length 32.9; P1–P2 is 52.1 mm; P2–C is 60.0 mm; P2–C extension falls on anterior margin of preopercle. Hyporhamphus tricuspidatus Gill, 1859. Holotype USNM 3407 (1, about 180 mm SL). Dorsal fin rays 15; anal fin rays 15; pectoral fin rays 11–11. Gill rakers on first arch 10 + 23 = 33. Lower jaw length 35.3 mm. Hemiramphus fasciatus Poey, 1860. Lectotype MCZ 32934 (1, 174 mm SL); Cuba; Poey no. 194, designated herein because it has the original Poey number tied on tail. Paralectotype MCZ 47954 (1, 178 mm SL). Characters of both (paralectotype in parentheses): dorsal fin rays 15 (16); anal fin rays 16 (17); pectoral fin rays 11–11 (12–12). Gill rakers on first arch 9 + 22 = 31 (8 + 23 = 31). Lower jaw length 23.0 mm (28.9 mm). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Atlantic Silverstripe Halfbeak (English) and parajito blanco del Atlántico (Spanish). Other common names include pajarito plateado in Mexico (Gallardo Torres et al. 2014), escribano in Cuba (Claro and Parenti 2001), marao blanco in Venezuela (Román 1979), and agulhabranca in Ceará and Pernambuco, Brazil (Lima and Oliveira 1978). Diagnosis. Hyporhamphus unifasciatus has 19–28, usually 23–25, gill rakers on the second arch in comparison to the Bermuda species H. collettei, which has 20–25, usually 21–22, and H. meeki, which has 20–30, usually, 26–27 (Table 4.10). Hyporhamphus unifasciatus also differs from H. collettei in the number of pectoral fin rays (usually 11 rather than 10; Table 4.8). Bases of dorsal and anal fins covered with scales (compared to the bases of the fins of H. roberti, which are scaleless). Description. Dorsal fin rays 13–16, usually 15; anal fin rays 14–18; pectoral fin rays usually 11, rarely 9 or 12 (Table 4.8). Vertebrae 32–34 precaudal + 17–19 caudal = 50–51 total. Gill rakers on first arch 7–11 + 18–25 = 27–35 (Table 4.9); gill rakers on second arch

Biology Predators. Hyporhamphus unifasciatus was reported from the stomach of a Yellowfin Tuna, Thunnus albacares, from St. Lucia (Beebe 1936:204) and is undoubtedly eaten by many more species of fishes. Parasites. Three of six specimens of Hyporhamphus unifasciatus from Buttonwood Canal, south Florida were infected with free spores and trophozoites of Ceratomyxa aggregata Davis (Iversen et al. 1971). The digenetic trematode Schikobalotrema acuta Linton, Haplosplanchnidae, was reported from the intestine of a specimen from Puerto Rico (Siddiqi and Cable 1960). In his revision of the cymothoid isopods of the genus Mothocya, Bruce (1986) reported one described and three new species from western Atlantic H. unifasciatus: M. argenosa Bruce from Cuba and the Virgin Islands, M. nana (Schioedte and Meinert) from Panama, M. bermudensis Bruce from Haiti, and M. omidaptria Bruce from Curaçao and Rio de Janeiro. Mothocya bermudensis was also described from H. collettei from Bermuda (as H. unifasciatus). Food. Seagrasses, algae, crustaceans, and small molluscs were found in stomachs of Brazilian specimens (Nomura and Menezes 1963; Rodrigues and Watanabe 1988), but another Brazilian investigation (Vasconcelos Filho et al. 1984) showed the diet to consist mostly of diatoms. Reproduction. A large female Hyporhamphus unifasciatus (USNM 342900,161 mm SL) from Cayo Hachuela, Cuba, had 60 eggs in the left ovary (1.00– 1.45 mm in diameter, x̅ 1.17 mm for 20 eggs) and 120 eggs in the right ovary (0.90–1.50 mm in diameter, x̅  1.15 mm for 20 eggs). Range. Confined to the western Atlantic Ocean from Uruguay north along the coast of South America and Central America, throughout the Caribbean Sea, 126

Family Hemiramphidae elongate (2.6–3.2 times in SL), barely reaching origin of pelvics. Pelvic fins short (7.3–10.6 times in SL), located approximately midway between the origin of the pectoral fins and the origin of the caudal fin or closer to the latter (P1–P2/P2–C = 0.9–1.3). Origin of the anal fin located under dorsal fin rays 2–4. Caudal fin with greatly elongate lower lobe. No scales present on fins. Swim bladder simple (single-chambered) or alveolar. Head length 4.0–4.6 times in SL. In contrast to other halfbeaks, in Oxyporhamphus the measurement is not of the lower jaw anterior to the end of the upper, but the snout length (from the anterior margin of the orbit to the tip of the lower jaw). Interorbital region convex. Lower jaw in adult fish very short (snout length 11–17 times in SL), but in the young halfbeak stage there is a strong projection, up to 4.0 times in SL at a length of 35–45 mm, apparently breaking off on attaining lengths of 50–100 mm. Preorbital shelf absent. Preorbital canal without posterior branch. There are more pores in the roof of the supraorbital canal than in other genera found in the Atlantic, three or four in canalis nasalis and six to eight in canalis frontalis. Jaw teeth with supplementary cusps, frequently tricuspid. Olfactory papilla not fimbriate. Opercle, suborbital, and interorbital parts of head covered with scales. Development. Eggs pelagic, large (1.9–2.1 mm) without an oil droplet; membrane covered with short (0.06–0.08 mm) filaments, equally distributed over the surface (Bruun 1935; Boehlert 1984: fig. 20a). Larvae of lengths less than 10 mm without elongate lower jaw. Pelvic fins develop on reaching a length of 7–8 mm (Khrapkova-Kovalevskaya 1963). Species. Two species are recognized (Parin et al. 1980; Collette 2004): Oxyporhamphus convexus and O. micropterus. Both have two subspecies; the Atlantic subspecies is O. micropterus similis. Range. Both species are marine; Oxyporhamphus convexus is restricted to the Indo-Pacific, whereas O. micropterus is widespread in tropical and subtropical seas (Collette 2004).

and reaching southern Florida (Fig. 4.12). Sympatric with Hyporhamphus meeki on the east coast of Florida from Miami north to St. Lucie Inlet, on the west coast from the Everglades north to Tampa Bay, and on the Mexican Gulf of Mexico (Contreras-Balderas et al. 1997). Reports of H. roberti and H. unifasciatus from the Mexican Gulf of Mexico, such as Castro-Aguirre (1978) and Castro-Aguirre et al. (1986), could be based on either H. meeki or H. unifasciatus. Literature records of Hyporhamphus unifasciatus from outside of this range refer to other superficially similar species of Hyporhamphus. The species previously considered as H. unifasciatus in the eastern Pacific from Baja California to Panama and the Galápagos Islands was described as H. naos by Banford and Collette (2001). Study Material. A total of 628 specimens from the western Atlantic (19.5–240 mm SL), including the types of Hemiramphus unifasciatus (MZB), Hemiramphus richardi (MNHN), Hyporhamphus tricuspidatus (USNM), and Hemiramphus fasciatus (MCZ), originally at AMNH, ANSP, BMNH, CAS, CHML, FMNH, FSU, FSBC, LACM, MCZ, MNHN, MNRJ, MZUSP, SU, UF, UMML, UMMZ, UPR, USNM, YPM, and ZMK. The specimens came from seven general regions: 105 specimens from southern Florida (19.5–205 mm SL); 193 specimens from the Antilles (16.0–195 mm SL); 55 specimens from Central America (31.2–183 mm SL); 103 specimens from the Caribbean coast of South America (104–210 mm SL); 161 specimens from Brazil (68–240 mm SL); 10 specimens from Fernando de Noronha (92.9–144 mm SL); and 1 specimen from Uruguay (191 mm SL). Genus Oxyporhamphus Gill, 1863 Bigwing Halfbeaks Oxyporhamphus Gill, 1863:273 (type species Hemiramphus cuspidatus by monotypy). Evolantia Heller and Snodgrass, 1903:189 (type species Exocoetus micropterus Valenciennes by monotypy).

Diagnosis. Oxyporhamphus differs from all other halfbeak genera in the shape of the upper jaw, which is short without a triangular projection from anterior margin. The lower jaw of Oxyporhamphus is distinguishable from most other genera (except Arrhamphus, Chriodorus, and Melapedalion) in having the lower jaw not being noticeably elongate (except in small juveniles). Description. Body elongate, laterally compressed; in cross section elongate oval. Greatest depth 5.5–6.0 times in SL, width of body in its depth 1.4–1.6 times. Pectoral branch of lateral line single. Pectoral fins

Oxyporhamphus micropterus similis Bruun, 1935 Atlantic Bigwing Halfbeak Figures 4.18, 4.19 Tables 4.1, 4.2 Exocoetus micropterus. Lütken 1876:102, 396 (three Atlantic specimens, 11°N, 26°W and 31°N, 76°W). Collette et al. 1997:20 (types). Evolantia microptera. Jordan et al. 1930:199 (listed). Oxyporhamphus micropterus similis Bruun, 1935:17–70 (original description), fig. 2 (teeth), fig. 3 (ovarian egg), pl. 2, fig. 1 (holotype). Beebe 1936:204 (in stomach of Thunnus albacares; St. Lucia). Breder 1938:4–12 (description, change in proportions

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Fishes of the Western North Atlantic 1977:150–151 (description, range), pl. 121. John 1983:24–25 (central Atlantic distribution between 20°N and 20°S), fig. 2 (distribution). Shiino 1976:91 (common name, Smallwing Flyingfish). Murdy 1983:90 (in key to Texas fishes). Rodríguez et al. 1984:14 (escribano volader; Cuba). Robins and Ray 1986:103 (description, range), pl. 18. Rodríguez and Valdés 1987:95 (Cuba). Boschung 1992:83 (Horn Island, AL). Davenport 1992, 1994 (wing-loading, comparison with Exocoetidae). Hensley and Hensley 1995:814 (eaten by Sooty Terns and Brown Noddies, Dry Tortugas, FL). Dasilao et al. 1997 (transferred to Exocoetidae). McEachran and Fechhelm 1998:963, figure (Gulf of Mexico). Mejía Aristizábal et al. 1998:30 (Colombian Caribbean oceanic atolls, listed). Schmitter-Soto et al. 2000:154 (Caribbean Mexico). Claro and Parenti 2001:43 (Cuba, listed). Menezes 2003:69 (Brazil). Lovejoy et al. 2004 (molecular phylogeny). Vaske et al. 2006 (in stomachs of Elagatis bipinnulata from São Pedro and São

with length), fig. 1 (adult and juvenile, 138 and 40 mm SL), fig. 3 (reduction of beak length with growth), fig. 4 (distribution in western Atlantic). Fowler 1944:96 (Courtown Key, Caribbean Sea, ANSP 93836). Parin 1961:148 (comparison with O. m. micropterus and O. meristocystis). Dragovich 1969:14 (eaten by Katsuwonus pelamis and Thunnus albacares). Duarte-Bello and Buesa 1973:84 (Cuba). Belyanina 1975:140 (ichthyoplankton; Caribbean Sea). Parin et al. 1980:146–156 (comparison with Indo-Pacific Oxyporhamphus), tables 22–25 (meristic data), fig. 47 (worldwide range). Collette 2005:952–953 (early life history, figs. A–G). Kells and Carpenter 2011:148–149 (description, color painting). Lewallen et al. 2011:165 (relationships to Hemiramphus). Schwartz 2012:35 (North Carolina). Oxyporhamphus micropterus. Fahay 1975:18 (juveniles; South Atlantic Bight). Guitart 1975:250 (Cuba, synonymy, description, fig. 184). Erdman 1976:23, 1977:159 (ripe females, March, August, November, Caribbean Sea). Hoese and Moore

Figure 4.18. Atlantic Bigwing Halfbeak, Oxyporhamphus micropterus similis. A. USNM 159032, 146 mm SL. B. USNM 159032, 70 mm SL. C. USNM 159032, 42 mm SL. All three specimens dipnetted from M/V Oregon in the Gulf of Mexico, south of Terrebonne Bay and New Orleans, Louisiana, U.S.A., 26 September 1954. Illustrations by M. H. Carrington.

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Family Hemiramphidae phus micropterus similis differs from all other western Atlantic halfbeaks, except Chriodorus atherinoides, in the lower jaw not being noticeably elongate (except in small juveniles). Oxyporhamphus m. similis differs from C. atherinoides in having more gill rakers on the first arch (30–35 in O. m. similis rather than 19–24 in C. atherinoides) and fewer pectoral fin rays (11–13 in O. m. similis rather than 12–14 in C. atherinoides). Description. Dorsal fin rays 13–15; anal fin rays 13–16; pectoral fin rays 11–13. Vertebrae 30–32 (usually 31) precaudal + 18–20 (usually 19) caudal = 49–51 (usually 50) total. Gill rakers on first arch 8–9 + 22–25 = 30–35. Pectoral fins long 30%–35% SL. Caudal fin well developed, lower lobe slightly larger than upper (Bruun 1935). Color. Oxyporhamphus micropterus similis is dark above, silver-white below, with a darker streak subdorsally that extends the length of the body. Juvenile stages are more pigmented than adults (Bruun 1935). Size. The maximum observed size for Oxyporhamphus micropterus is 185 mm SL. Figure 4.19. Distribution of Atlantic Bigwing Halfbeak, Oxyporhamphus micropterus similis, based on specimens examined by B. B. Collette. Map by W. E. Bemis.

Biology Predators. Oxyporhamphus micropterus similis has been recorded from the stomachs of a wide variety of fishes: Yellowfin Tuna, Thunnus albacares (Beebe 1936; Perrin et al. 1973); Skipjack Tuna, Katsuwonus pelamis (Dragovich 1969); Blackfin Tuna, Thunnus atlanticus (Dragovich 1969; Headley et al. 2009); Rainbow Runner, Elagatis bipinnulata (Vaske et al. 2006); and sea birds such as Brown Noddies and Sooty Terns in the Dry Tortugas (Hensley and Hensley 1995). Parasites. The cymothoid isopod Lironeca has been reported from Oxyporhamphus micropterus from the eastern Pacific (Aguilar-Palomino et al. 1997). Food. The diet of Oxyporhamphus micropterus similis is composed mainly of copepods and some amphipods (Gorelova and Gurdstev 1987), as it is also for the eastern Pacific population of the nominal subspecies O. m. micropterus (Van Noord et al. 2013). Larvae and 4–16 mm young from across the Pacific Ocean fed largely on the nauplii, copepodites, and adult stages of small copepods, accounting for more than 90% by weight and 80% numerically of the diet (Gorelova 1980). In addition to copepods, siphonophores were also significant in the diet of young O. m. micropterus in the Pacific, but not in the Indian Ocean (Lipskaya 1980). The young feed only during daylight hours (Gorelova 1980; Lipskaya 1980). Reproduction. Females of Oxyporhamphus micropterus similis are ripe by 120 mm. They are ready to shed eggs in November between Barbados and St.

Paulo, Arquipelago, Brazil). Cotten and Comyns 2005:960– 961 (figures of 10 juveniles, 3.4–41.2 mm). Fahay 2007:776, 796–797 (early stages, figs. A–I). Tibbetts et al. 2007:809 (simple, single-chambered swim bladder). Casazza and Ross 2008:351 (associated with Sargassum, Gulf Stream off North Carolina). Headley et al. 2009:672 (table 1, eaten by Blackfin Tuna, Thunnus atlanticus in Tobago). McEachran 2009:1286 (Gulf of Mexico). Collette 2010:34 (reproduction and development). Nolf 2013:74, pl. 154 (illustrations of several otoliths from Acapulco, Mexico). Snyder and Burgess 2016:100 (listed, Florida). Oxyporhamphus similis. Starck et al. 2017:91 (Florida Keys).

Types. Oxyporhamphus micropterus similis Bruun, 1935. Lectotype designated by Bruun (1937:186); MNHN A.9925 (120 mm SL). Paralectotypes MNHN 3408, 3409, B.840 (Collette et al. 1997:20). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Smallwing Flyingfish (English) and volador alita (Spanish). However, because we recognize this species as a halfbeak, not a flyingfish, we choose to call it Atlantic Bigwing Halfbeak. The common name in Cuba is volador de alas negras (Claro and Parenti 2001). Diagnosis. Anterior margin of upper jaw straight, not forming a prominent triangular anterior projection. Swimbladder a simple, single chamber, not vesicular, composed of many small bubbles as in its sister species Oxyporhamphus convexus. Oxyporham129

Fishes of the Western North Atlantic Lucia, in March at 10°00′W, 03°45′S, and in August at 16°52′W, 03°05′N. Variation in timing, despite differences in localities, suggests that spawning periods are not restricted. Ripe females occur far from coasts and the eggs, therefore, are pelagic (Bruun 1935). A large female (USNM 216329, 149 mm SL) collected from the Caribbean Sea had 500 eggs in the left ovary (0.95–1.25 mm in diameter, x̅ 1.09 for 100 eggs) and 320 eggs in the right ovary (0.90–1.30 mm in diameter, x̅ 1.09 mm for 64 eggs). Range. Oxyporhamphus micropterus similis is widespread in tropical and subtropical waters of the Atlantic, in the western Atlantic north to 40°N, in the Gulf of Mexico and Caribbean Sea, and the Atlantic south past the equator (Fig. 4.19); to 27°07′05″S, 46°39′29″W (MZUSP 78209). In the eastern Atlantic it occurs from 20°N south to 20°S (John 1983: fig. 2). The nominal subspecies, O. m. micropterus, is widespread throughout the Indo-Pacific from 35°N off the coasts of Japan to 25°S in the Central Pacific (Parin et al. 1980: fig. 47). Geographic Variation. The Atlantic subspecies Oxyporhamphus micropterus similis was described by Bruun (1935) because the transition to the definitive state of the lower jaw takes place in O. micropterus micropterus at a larger size (90–100 mm SL) than in O. micropterus similis (about 60 mm SL). Breder (1938) also noted that the two subspecies differ in the color of the pelvic fins in adult fish (usually light in O. m. similis and black in O. m. micropterus) and in the relative length of the base of the dorsal fin (greater in the Atlantic subspecies). There are small differences between the subspecies in the number of gill rakers on the first arch (x̅  30.7 in O. m. micropterus and 32.0 in O. m. similis) and predorsal scales (x̅  31.1 and x̅  29.9, respectively), and also in the position of the pelvic fins (in O. m. micropterus almost always closer to the base of the lower lobe of the caudal fin than to margin of the opercle and in O. m. similis most often closer to the latter) according to Parin et al. (1980). Study Material. A total of 148 specimens (16.3–146 mm SL) from 52 western Atlantic collections at MZUSP and USNM.

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Family Exocoetidae FLYINGFISHES NIKOLAY V. PARIN ILIA B. SHAKHOVSKOY KATHERINE E. BEMIS and BRUCE B. COLLETTE Introduction The Exocoetidae, the flyingfishes, is one of six families of the order Beloniformes. It is the sister group of the Hemiramphidae, the halfbeaks, forming the superfamily Exocoetoidea (Collette et al. 1984). Flyingfishes lack the elongate lower jaw (except for juveniles of Fodiator) present in most halfbeaks and can be distinguished from the needlefishes (Belonidae) and sauries (Scomberesocidae), which have both jaws elongate. The Exocoetidae contains seven genera and about 65–70 species (Shakhovskoy and Collette, in press). Flyingfishes are small- to medium-sized (to 480 mm total length) with elongate, broadly cylindrical (round or elliptical in cross section) bodies, flattened ventrally in some species. The head is short, snout usually blunt (produced only in Fodiator), and shorter than the eye in all western Atlantic species. Flyingfishes have a small mouth and very small or no jaw teeth, but have well-developed pharyngeal teeth. The upper pharyngeal bones of the third gill arches are close together, but not fused into a single plate. There are no spines in the fins. The dorsal and anal fins are set equally far back on the body, their bases are short and opposed. The caudal fin is deeply forked, its lower lobe longer than the upper. The pectoral fins insert high on the body, are strikingly long, and always extend beyond the origin of the dorsal fin in adults. The pelvic fins are abdominal in position, and greatly enlarged in many, but not all, species. The lateral line is low on the body, and provides diagnostic characters for the genera as described by Parin and Astakhov (1982). The large cycloid scales are easily shed. The swim bladder is large, extending posteriorly beyond the body cavity. Adult flyingfishes are usually dark above and pale below. The dark colors are typically iridescent blue or green in life; the dorsal fin in some species has black

pigment, and the pectoral fins in some species have a pale crossband and outer margin. Otolith and tagging studies suggest that flyingfishes are short lived, with individuals usually living no more than 1.5 years. All species are oviparous. The eggs of some species are buoyant and float on the surface, whereas the eggs of other species have specialized filaments to attach to floating debris and seaweed. Juveniles (to about 100–150 mm SL) differ in appearance from adults, having shorter pectoral fins, a higher dorsal fin, and a conspicuous single or paired chin barbel in many species. Color patterns of juvenile flyingfishes are highly variable, and they often have spots and bars not found in adults. Exocoetids are well known for leaping out of the water and gliding through the air to escape predators below the surface (see the amazing photographs of flyingfishes in Howell [2014]). The two-winged flyingfishes use enlarged pectoral fins, whereas the four-wing flyingfishes have enlarged pelvic as well as pectoral fins that allow them to glide relatively longer distances. Fins are not flapped during flight, but rather erected quickly as the fish leaves the water and held perpendicular to the body during the glide; the fish is propelled into the air by a strong thrust of the caudal fin (Davenport 1994, 2003). Flyingfishes inhabit the surface of open oceans as well as neritic and inshore areas and are abundant in most tropical seas. Many species of flyingfishes school in large groups, but do not undertake extensive migrations. Flyingfishes are important components of pelagic food webs and serve as prey species for dolphinfishes, tunas, billfishes, sharks, cetaceans, and pelagic seabirds (Perrin et al. 1973; Richard and Barbeau 1994; Andrew et al. 1995; Vaske et al. 2005; Alonso et al. 2014; Herzing and Elliser 2014). Little specific information is known about the diet of western Atlantic species of flyingfishes, which predominantly

Fishes of the Western North Atlantic consists of zooplankton. Larger species of flyingfishes also prey on smaller fishes. Plastics are becoming more common in oceanic ecosystems and 60% of all flyingfishes examined in the eastern Pacific were found to have had consumed plastics, up to 75% of individuals of some species (Van Noord et al. 2013). Khachaturov (1983) studied the digestive systems of 10 species of flyingfishes from seven genera. He found that the premaxillary and dentary bones have few weak teeth. There are a moderate number of gill rakers on the anterior side of the first gill arch (19–37), differing by genera. The shape of the gill rakers also differs among genera. Absence of a true stomach and presence of a short intestine (47%–55% of body length) are correlated with a pharyngeal apparatus that processes food before swallowing. The pharyngeal apparatus contains well-developed upper and lower pharyngeal bones with teeth. The lower pharyngeal bones are fused into a triangular plate that bears teeth of different shapes. The number of teeth increases as the fish grows. All material found in the intestines had undergone mechanical processing by pharyngeal teeth. The mucosa of the gut is thick, 60%–70% of the entire thickness of the walls of the gut. The liver is 2%–3% of a fish’s weight; it is bilobate with the right lobe larger than the left. The gall bladder lies under the right lobe of the liver. Kurochkin (1980) provided a summary of the species of parasites reported from flyingfishes: myxosporidians (1 species), trematodes (36 species), monogenes (13 species), cestodes (5 species), acanthocephalans (3 species), and crustaceans (24 species). The didymozoid trematode Gonapodasmius cypseluri is known from 10 species of flyingfishes and Gonapodasmius spilonotopteri from 4 species (Nikolaeva and Mordvinova 1988). Bannikov et al. (1985) described a new genus and species, †Rhamphexocoetus volans, from the lower Eocene of Italy that they provisionally placed in Exocoetidae. There is no otolith-based fossil record for the Exocoetidae (Nolf 2013). Although flyingfishes are edible and some species are highly prized, few commercial fisheries have been developed. In the western Atlantic, there are target fisheries in Brazil and the Caribbean, especially Barbados and Tobago, based on drift gill nets in localities of unusually heavy spawning concentrations. The catch from this fishery is mostly fresh frozen. In other regions, especially around the Antilles, flyingfishes are consumed fresh or used as bait. Statistics from the Food and Agriculture Organization of the United Nations report landings ranging from 3,954 to 7,297 tons from 2011 to 2013 from the Atlantic Ocean and adjacent seas.

Parin (1961) and Collette et al. (1984) proposed a subfamilial classification based on morphological characters: Fodiatorinae, including only Fodiator (2 species); Parexocoetinae, including only Parexocoetus (3 species); Exocoetinae, with only Exocoetus (5 species); and Cypselurinae with four genera, Prognichthys (6 species), Cypselurus (ca. 11 species), Cheilopogon (ca. 25–30 species) and Hirundichthys (ca. 10 species). Molecular analyses strongly support the monophyly of Exocoetidae, all four subfamilies, and all existing genera except the most diverse genus, Cheilopogon, which is divided among two or three separate clades (Lewallen et al. 2011: fig. 2; Gordeeva and Shakhovskoy 2017). The flyingfish fauna of the western Atlantic includes 16 species in six genera: Exocoetus (2 species); Cheilopogon (6 species); Cypselurus (1 species); Hirundichthys (4 species); Parexocoetus (1 species); and Prognichthys (2 species). Two of these species (Cypselurus comatus and Prognichthys occidentalis) are endemic to the western Atlantic and 8 species (Parexocoetus hillianus, E. obtusirostris, H. affinis, H. volador, Prognichthys glaphyrae, Cheilopogon exsiliens, Cheilopogon heterurus, and Cheilopogon melanurus) are endemic to the Atlantic. Six species—E. volitans, H. speculiger, H. rufipinnis, Cheilopogon cyanopterus, Cheilopogon pinnatibarbatus, and Cheilopogon furcatus—are also in other oceans, but the last two species are represented in the Atlantic by endemic subspecies. Five species are endemic to the eastern Atlantic: Fodiator acutus, Cheilopogon nigricans, Cheilopogon milleri, H. rondeletii, and Prognichthys gibbifrons. Fodiator acutus was originally described by Valenciennes in Cuvier and Valenciennes (1847) as E. acutus and the type is listed as from Suriname. There are a few other records of F. acutus from the western Atlantic, including the Caribbean Sea (Morrow 1957) and the Gulf of Mexico (Castro-Aguirre and Marquez-Espinosa 1981). However, subsequent reliable records of the Atlantic species have only come from the Gulf of Guinea. Therefore, records of F. acutus in the western Atlantic are likely erroneous or the result of occasional larval drift; Parin and Belyanina (2002a) provide a thorough summary of the matter. We include F. acutus in the key, but not a full species account. Materials and Methods Unless otherwise indicated, all body lengths are reported as standard length (SL) in millimeters. Methods and measurements follow Parin (1960, 1961); dental indices follow methods of Bruun (1935) and Parin (1961). A scale smaller than half the size of a normal scale is designated as a half (½) scale. The 150

Family Exocoetidae urostyle is included in vertebral counts. The bulk of study material was examined by N. V. Parin, T. N. Belyanina, and I. B. Shakhovskoy. Archive data (meristic and morphometric sheets) of R. H. Gibbs, Jr. and J. C. Staiger have also been used. Many specimens of Parexocoetus hillianus were examined by A. Khachaturov, T. N. Steyker, and G. N. Pokhilskaya. Most of the flyingfishes examined for this report are at the Smithsonian Institution or the Museum of Comparative Zoology, Harvard University, where

Parin visited in 1978, 1990, and 1992 to study and borrow materials. Standard two-letter postal codes are used for U.S. locations. Standard symbolic codes for institutions housing museum specimens follow Sabaj (2016). We thank all those who have studied specimens used in our work. Irina Eidus (Zoological Museum, University of Hamburg) kindly provided us with photographs and X-rays of ZMH specimens of Cheilopogon pinnatibarbatus.

Key to the Species of Exocoetidae in the Western Atlantic 1a. Pectoral fins moderately long, not reaching beyond posterior part of anal fin base; pectoral branch of lateral line present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Pectoral fins very long, reaching beyond anal fin base to or almost to caudal fin base; pectoral branch of lateral line absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2a. Snout long and pointed, much longer than eye diameter; upper jaw not protrusible; juveniles without barbels, with lower jaw greatly elongate (recorded only twice from the area, off Suriname and in the Caribbean Sea, but both records seem to be erroneous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fodiator acutus 2b. Snout short and blunt, shorter than eye diameter; upper jaw protrusible; juveniles with paired chin barbels (easily lost) and lower jaw not elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parexocoetus hillianus 3a. Pelvic fins short, not reaching anal fin origin (barely reaching in juveniles), inserted nearer to pectoral fin insertion than anal fin origin; juveniles without barbels. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3b. Pelvic fins long, reaching well beyond anal fin origin, inserted nearer anal fin origin than pectoral fin insertion; juveniles with or without barbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4a. Gill rakers on first gill arch 22–29 (usually 25–27); 7 or 8 (usually 7½) scales in transverse row between dorsal fin origin and lateral line; juveniles humpbacked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Exocoetus obtusirostris 4b. Gill rakers on the first gill arch 29–37 (usually 32–34); 6 or 6½ scales in transverse row between dorsal fin origin and lateral line; juveniles elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Exocoetus volitans 5a. Origin of anal fin three rays or more behind origin of dorsal fin; dorsal fin two to five rays more than anal fin; juveniles with or without barbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5b. Origin of anal fin slightly before, under, or rarely, not more than two rays behind origin of dorsal fin; dorsal fin usually with less or equal number of rays than anal fin; juveniles without barbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 6a. First two pectoral fin rays unbranched; juveniles without barbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6b. Only the first pectoral fin ray unbranched; juveniles with barbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7a. Pectoral fins chiefly pale with central part gray; eye diameter 1.2–1.5 times in postorbital part of head; anal fin rays 8–11 (usually 10); juveniles with pectoral fins pale with two large black blotches, one at base and the other in outer portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prognichthys glaphyrae 7b. Pectoral fins brownish or gray with light tip, light posterior margin and light area near fin base up to seventh to ninth rays; eye diameter 1.4–1.8 times in postorbital part of head; anal fin rays 8–10 (usually 9); juveniles with pectoral fins almost uniformly pigmented, usually with lighter oblique transverse stripe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prognichthys occidentalis 8a. Pelvic fins inserted about midway between posterior head margin and origin of lower caudal fin lobe; lower jaw slightly shorter than upper jaw and included beneath the latter when mouth closed; pectoral fins lightly and almost uniformly pigmented; juveniles with a single long chin barbel . . . . . . . . . . . . . . . . . . . . . . . . Cypselurus comatus 8b. Pelvic fins inserted much nearer to posterior head margin than origin of lower caudal fin lobe (except in Cheilopogon pinnatibarbatus and C. cyanopterus); both jaws of equal length or lower jaw a little longer than upper jaw; pectoral fins more densely pigmented, usually (except in C. cyanopterus) with paler transverse strip or basal triangle; juveniles with paired chin barbels or one semicircular fringed appendage. . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

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Fishes of the Western North Atlantic 9a. Vertebrae 49–52; least caudal peduncle depth 17–18.5 times in SL; predorsal scales 38–47; head length 4.4–5.0 times in SL; juveniles up to 120–140 mm SL with semicircular fringed chin barbel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilopogon pinnatibarbatus 9b. Vertebrae 42–47 (46–50 in Cheilopogon heterurus); least caudal peduncle depth 12.5–18 times in SL; predorsal scales 23–38 (32–43 in C. cyanopterus); head length 3.8–4.6 times in SL (in C. heterurus up to 4.9); juveniles with paired chin barbels. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10a. Dorsal fin relatively high (longest ray usually less than 10 times in SL), bearing a prominent black spot; palatine teeth present; jaw teeth noticeable (conspicuous to the touch) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10b. Dorsal fin relatively low (longest ray usually more than 10 times in SL), without black spot; palatine teeth absent; jaw teeth minute (not conspicuous to the touch). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11a. Predorsal scales 23–30; pectoral fins black with a definite pale crossband; lower caudal fin lobe obviously darker than upper lobe; juveniles with moderately long flat barbels, their length more than 2.2 times in SL. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilopogon exsiliens 11b. Predorsal scales 32–43; pectoral fins uniformly bluish black except for outer margin and lowermost rays; both caudal fin lobes equally pigmented; juveniles with very long barbels with fleshy axis, their length less than 1.7 times in SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cheilopogon cyanopterus 12a. Pectoral fin usually blackish, with definite pale crossband and broad outer margin; juveniles with moderately long barbels, their length usually less than 7 times in SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilopogon furcatus 12b. Pectoral fins gray, with indefinite pale triangular crossband and narrow outer margin; juveniles with short barbels, their length more than 7 times in SL. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13a. Head length 3.9–4.6 times, usually less than 4.3 times in SL; predorsal scales 25–33, usually 27–30; vertebrae 45–47, usually 46 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilopogon melanurus 13b. Head length 4.3–4.9 times, usually more than 4.5 times in SL; predorsal scales 28–38, usually 30–35; vertebrae 46–50, usually 48. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilopogon heterurus 14a. First 2 pectoral fin rays unbranched; pectoral fin uniformly black without pale crossband; head length usually 4.5–5.0 times in SL; pectoral fin length usually 1.25–1.35 times in SL. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14b. Only the first pectoral fin ray unbranched; pectoral fins with pale triangular crossband; head length usually 3.9–4.6 times in SL; pectoral fin length 1.4–1.6 times in SL. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 15a. Pelvic fins with prominent black spot; scales in oblique transverse row 6½–8½, usually 7–7½; juveniles with banded body and, usually, mottled pectoral and pelvic fins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hirundichthys volador 15b. Pelvic fins without black spot; scales in oblique transverse row 5½–7½, usually 6½; juveniles with almost uniformly colored body and dark pectoral and pelvic fins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hirundichthys rufipinnis 16a. Palatine teeth usually present; pectoral fins with broad pale outer margin . . . . . . . . . . . . . . . Hirundichthys speculiger 16b. Palatine teeth usually absent; pectoral fins with narrow pale outer margin . . . . . . . . . . . . . . . . . . Hirundichthys affinis

Genus Cheilopogon Lowe, 1841

in number of unbranched rays of the pectoral fin. Description. Dorsal fin rays 9–17; anal fin rays 7–12; dorsal fin rays minus anal rays 1–6 (rarely 0 or –1); pectoral fin rays I (II in Cheilopogon agoo) 12–17. Gill rakers 17–28; vertebrae 26–35 + 12–17 = 41–52. Body elongate, depth 4.8–7.6 times in SL. Pelvic fins long, their base closer to caudal fin base than to snout tip. Pectoral fins long, reaching end of dorsal fin base or beyond. Anal fin origin under or behind third dorsal fin ray. Lower jaw longer than upper jaw or equal to it. Jaw teeth usually unicuspid. Mouth not protrusible. Pectoral branch of lateral line absent. Eggs with filaments. Juveniles with two chin barbels, sometimes fused into a wide fringed

Cheilopogon Lowe, 1841:38 (type species Cypselurus pulchellus Lowe, 1841 [type by subsequent monotypy]).

Diagnosis. Species of Cheilopogon differ from all other flyingfishes (except for Parexocoetus) in the structure of the juvenile chin barbels. Cheilopogon differ from species of Exocoetus and Parexocoetus in length and position of the pelvic fins as well as in relative position of the anal and dorsal fin origins, from Cypselurus in structure of jaw teeth and length of lower jaw, from Hirundichthys in relative position of the anal and dorsal fin origins, and from Prognichthys 152

Family Exocoetidae appendage (Parin 1961; Shakhovskoy and Collette, in press). This genus needs further study, as Cheilopogon has no known synapomorphies and may be paraphyletic based on analysis of molecular data (Lovejoy et al. 2004; Lewallen et al. 2011; Gordeeva and Shakhovskoy 2017). Species. About 25–30 species worldwide. Six of eight Atlantic species found in the western Atlantic. Range. Found in the Atlantic, Indian, and Pacific in open oceans and neritic waters (Parin 1961; Shakhovskoy and Collette, in press).

EXOC. Cyp.2 (western Atlantic). Hunte et al. 1995:28–35 (eggs and larval distribution; eastern Caribbean). Oxenford et al. 1995a:16–21 (distribution of adults; eastern Caribbean). Monteiro et al. 1998:399 (distribution; northeastern Brazil). ?Cypselurus hewitti Smith, 1935:181–182 (original description; South Africa; holotype RUSI 28). Smith 1961:126 (South Africa). Cheilopogon cyanopterus. Parin 1961:168 (listed). Kovalevskaya 1977:292–297 (early life history; Atlantic Ocean, Indo-West Pacific). Fedoryako 1980:581 (Sargasso Sea). Heemstra and Parin 1986:392 (short description; South Africa). Gorelova and Grudtsev 1987:481 (feeding; Atlantic). Nikolsky and Luschina 1990:135–136 (eastern equatorial Atlantic). Parin and Gibbs 1990:583 (eastern Atlantic). Parin and Belyanina 1996:755 (distribution; Atlantic Ocean). McEachran and Fechhelm 1998:944, figure (Gulf of Mexico). Smith-Vaniz et al. 1999:170–171 (Bermuda). Parin 2003:1121, figure, map (description, western Atlantic). Cotten and Comyns 2006:968–969, figs. A–E (early life history). Fahay 2007:800– 801, figs. A–E (early life history stages). Casazza and Ross 2008:352, 355 (associated with Sargassum, Gulf Stream off North Carolina). Lewallen et al. 2011 (molecular systematics). Parin and Shakhovskoy 2016:2138 (description, distribution; east-central Atlantic). ?Cheilopogon furcatus. (not of Mitchill 1815). Collette et al. 2003: fig. 33 (juvenile; Navassa Island, West Indies).

Cheilopogon cyanopterus (Valenciennes in Cuvier and Valenciennes, 1847) Margined Flyingfish Figures 5.1, 5.2 Tables 5.1, 5.2, 5.3, 5.4, 5.5 Exocoetus cyanopterus Valenciennes in Cuvier and Valenciennes, 1847:97–98 (original description; Brazil; three syntypes MNHN B 769, 1996-230, A 7605). Günther 1866:294 (coast of Brazil). Jordan and Meek 1885:63 (coast of Brazil). Jordan and Evermann 1896a:739 (in part; description; North America). Bruun 1937:182 (type description). Collette et al. 1997:11 (types). Exocoetus dussumieri Valenciennes in Cuvier and Valenciennes, 1847:132–133 (original description; South Africa; holotype MNHN 1930). Bruun 1937:186 (description of type). Collette et al. 1997:11 (type). Exocoetus vermiculatus Poey, 1860:300–301 (original description; Cuba; no types known). Exocoetus albidactylus Gill, 1863:167–168 (original description; Caribbean Sea; no types known). Exocoetus bahiensis. (not (?) of Ranzani 1842). Günther 1866:293 (Atlantic Ocean). Jordan and Meek 1885:64 (tropical seas, north to Cuba). Osorio 1898:199 (listed; Annobon Island). Exocoetus parrae Poey, 1868:385 (original description; between Hispaniola and Cuba; no types known). Cypsilurus bahiensis. (not (?) of Ranzani 1842). Jordan and Evermann 1898:2836 (listed). Smith 1949:126 (South Africa). Cypsilurus cyanopterus. Jordan and Evermann 1898:2836 (species considered valid). Bruun 1935:40–44, pl. 3, figs. 1, 4 (description; Atlantic Ocean). Cypselurus bahiensis. (not (?) of Ranzani 1842). Evermann and Marsh 1900:104 (short description; Puerto Rico). Breder 1927:22–23 (Caribbean Sea). Jordan et al. 1930:202 (listed). Nichols and Breder 1930:1–8 (Caribbean Sea). Parr 1930:26 (Bahamas). Beebe and Tee-Van 1933:69 (in part; Bermuda). Breder and Nichols 1934:40–44. Smith 1961:126 (South Africa). Cypselurus monroei Nichols and Breder, 1928:432–433 (original description; Florida; holotype AMNH 8678). Breder and Nichols 1930:1–3, fig. 1 (description; Florida). Nichols and Breder 1930:5–7 (in key). Breder and Nichols 1934:40–44 (discussion). Cypselurus cyanopterus. Jordan et al. 1930:202 (listed). Fowler 1936:1244 (Canary Islands). Breder 1938:39–46, figs. 18–20 (description; western Atlantic). Cadenat and Marchal 1963:1254–1255 (description; eastern Atlantic [Ascension Island]). Staiger 1965:685–691, figs. 3–6 (description; western Atlantic). Gibbs and Staiger 1970:449–452 (eastern Atlantic). Fahay 1975:16 (juvenile off Florida). Gibbs 1978:

Types. No type specimens known to be extant for these three species: Exocoetus vermiculatus Poey, 1860, E. albidactylus Gill, 1863, and E. parrae Poey, 1868. Exocoetus cyanopterus Valenciennes in Cuvier and Valenciennes 1847. Lectotype (designated by Bruun 1937:182) MNHN B.769 (306 mm SL); coast of Brazil, Bahia. Dorsal fin rays 13; anal fin rays 10; pectoral fin rays I 13. Predorsal scales 37; transverse scales between dorsal fin origin and lateral line 7. Vertebrae 30 + 15 = 45. Jaw teeth conspicuous. Pectoral fin black to tenth membrane; dorsal fin with large black blotch in third to tenth membranes. Paralectotypes MNHN 1996-230 (out of B.769, 283 mm SL) and A.7605 (296 mm SL). Exocoetus dussumieri Valenciennes in Cuvier and Valenciennes, 1847. Holotype MNHN 1930 (130 mm SL); Cape of Good Hope. Dorsal fin rays 13; anal fin rays 9; pectoral fin rays I 12. Predorsal scales about 34; transverse scales between dorsal fin origin and lateral line 7. Gill rakers 7 + 18 = 25. Vertebrae 30 + 14 = 44. Barbel length 35% SL. Cypselurus monroei Nichols and Breder, 1928. Holotype AMNH 8678 (45.5 mm SL); Miami, Florida, Coconut Grove. Dorsal fin rays 13; anal fin rays 9; pectoral fin rays I 12; dorsal fin very high. Barbel length 47.9% SL. Cypselurus hewitti Smith, 1935. Holotype RUSI 28 (174 mm SL); Port Alfred, South Africa. Dorsal fin rays 14; anal fin rays 10; pectoral fin rays I 12; Predorsal scales 35; scales between dorsal fin base and lateral line 8; dorsal fin height 16.5% SL. Gill rakers 7 + 19 = 26. Barbel length 12.7% SL. 153

Fishes of the Western North Atlantic

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Family Exocoetidae

Figure 5.1. Margined Flyingfish, Cheilopogon cyanopterus. A. USNM 203890, 267.2 mm SL. Collected off the island of Annobón, 12 December 1965. From Gibbs and Staiger (1970). B. USNM 198378, 178.5 mm SL. Collected by R. H. Gibbs, Jr. and B. B. Collette, fish flew aboard M/V Delaware at ca. 39°N, 68°W, 23 September 1957. C. USNM 198455, 82.6 mm SL. Collected by R. H. Gibbs, Jr. at 39°28′N, 69°30′W from M/V Delaware, 30 October 1956. From Gibbs and Staiger (1970). Illustrations by H. E. Hamman.

155

Fishes of the Western North Atlantic of juveniles (25–120 mm SL) is less elongate (depth 5.8–7.1 times in SL) and the head is shorter (4.1–5.0 times in SL). Eye diameter 2.8–3.3 times in head length and 1.0–1.2 times in interorbital space. Dorsal fin greatly elevated, much higher than in adults, decreasing in relative size with growth, its longest rays (fifth to seventh) 3.9–5.8 times in SL. Pectoral fins 1.4–1.7 times in SL. Pelvic fins 2.4– 3.0 times in SL. Paired chin barbels coalescent at base with a firm pointed median stem and two soft skin folds along the anterior and posterior sides (the longest barbeled specimen is 190 mm SL, the shortest unbarbeled about 165 mm SL); barbels very long, usually exceeding standard length at 20–80 mm SL. No pigmented transverse bars on body. Stem of barbel not pigmented, lateral folds black. Dorsal fin black. Anal fin dark gray, gradually losing pigment. Pectoral fins uniformly gray (in small juveniles), darkening at about 40–50 mm SL when numerous oval light spots appear along the anterior edge, entirely black at more than 100 mm SL. Pelvic fins partly pigmented, remnants of pigmentation on distal part disappear at 150–160 (rarely 178) mm SL. Size. Cheilopogon cyanopterus is one of the largest species of the Atlantic flyingfishes, with a maximum observed size of 335 mm SL.

Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Margined Flyingfish (English) and volador azul (Spanish). Locally it is known as “guineaman” in the Barbados, “den” in Dominica, and codène in Grenada (Oxenford et al. 2007). Diagnostic Characters. Cheilopogon cyanopterus differs from C. exsiliens in predorsal scale count (32–43 rather than 23–30, respectively; Table 5.1) as well as pectoral and caudal fin coloration. Juvenile C. cyanopterus (20–80 mm SL) have long paired chin barbels 1.7 times in SL with fleshy axis, compared to flat barbels more than 2.5 times in SL that characterize C. exsiliens. Cheilopogon cyanopterus differs from all other species of Cheilopogon in pectoral fin coloration and structure of juvenile chin barbels. Description. Dorsal fin rays 11–14, usually 12–13; anal fin rays 8–11; dorsal rays minus anal rays 1–4; pectoral fin rays I 12–15 (Table 5.2). Vertebrae 27–31 precaudal + 14–16 caudal = 42–46, usually 43–45 total (Table 5.3). Predorsal scales 32–43, usually 34–40 (Table 5.4); scales in transverse row from dorsal fin origin to lateral line 7–9½, usually 7½–8½ (Table 5.4). Total gill rakers on the first arch 5–8 upper + 14–19 lower = 20–28, usually 23–25 (Table 5.5). Measurements based on Atlantic specimens (247–323 mm SL). Body elongate (greatest depth 5.3–6.0 times in SL); slightly compressed (width 1.1–1.4 times in depth). Head 4.1–4.4 times in SL and 1.0–1.3 times in dorso-caudal distance. Eye diameter 3.1–3.6 times in head length, 1.1–1.4 times in interorbital space. Jaws subequal. Jaw teeth comparatively large, sharp, often slightly curved; dental index 0.15–0.28. Palatine teeth usually present. Dorsal fin rather high, its longest (second or third) ray 7.6–13.8 times in SL. Anal fin originates under fifth to seventh dorsal fin ray. Pectoral fins 1.4–1.5 times in SL, reaching from middle to end of caudal peduncle, its first ray 1.5–1.8 times shorter than total fin length, first ray unbranched. Pelvic fins inserted at equal distance to posterior margin of opercle and origin of lower caudal fin lobe (opercle to pelvic distance 0.9–1.1 times in pelvic to caudal fin distance), moderately long (3.2–3.8 times in SL); reaching middle to end of anal fin base. Color. Body dark above, pale below. Dorsal fin gray with a prominent large blue-black blotch between third and seventh to eleventh rays, lower margin of pigmented area semicircular, usually not extending to the base of fin. Anal fin transparent. Caudal fin dark gray to almost black. Pectoral fins bluish black except narrow outer margin and lowermost portion. Pelvic fins unpigmented. Development. In comparison to adults, the body

Biology Predators. Eaten by a wide variety of fishes, including Rainbow Runner, Elagatis bipinnulata (Vaske et al. 2006). Cheilopogon cyanopterus is a major part of the diets of sea birds (Sula leucogaster, Anous stolidus, and A. minutus) at Saint Paul Archipelago (Vaske et al. 2005). Food. Feeds on zooplankton and small fishes (Vaske et al. 2006). Gorelova and Grudtsev (1987) found that copepods were the dominant prey of Cheilopogon cyanopterus in the Atlantic from 20°N to 30°S and 40°W to 6°E. Febyanty and Syahailatua (2010) reported that in the Macassar Strait, C. cyanopterus fed on copepods (68.75%), algae (12%), chaetognaths (6.2%), and Malacostraca (3.1%). Reproduction. Mature at about 270 mm SL. The largest known ovarian eggs (1.2 mm in diameter) were from a female that was not fully ripe (Gibbs and Staiger 1970). The ovaries of a large female of Cheilopogon cyanopterus from the Indian Ocean near Madagascar had 24,300 eggs (0.65–0.75 mm in diameter) with long filaments, about 4.5 mm in length, covering the entire surface (Parin and Gorbunova 1964). These authors suggested a single spawning event for C. cyanopterus. Spawning behavior was reported by Bane (1965) in the eastern 156

Family Exocoetidae

157

Fishes of the Western North Atlantic Atlantic (the Gulf of Guinea) just before sunset in August. The sea surface temperature was 24.4 °C. Relationship to Humans. Cheilopogon cyanopterus is caught in smaller numbers in the Caribbean than the targeted Hirundichthys affinis, but it is of commercial importance in St. Lucia, St. Vincent, the Grenadines, Martinique, and Dominica. It is usually caught by hook and line, rather than with the gill nets that are used for H. affinis (Oxenford et al. 2007). Range. The range of Cheilopogon cyanopterus includes both the tropical Atlantic and Indo-West Pacific in coastal waters and the open sea. In the western Atlantic it is distributed along the shores from 40°N (juveniles expatriated from the Gulf Stream) to Rio de Janeiro, Brazil, including the Gulf of Mexico and the Caribbean Sea (Fig. 5.2). In the eastern Atlantic, C. cyanopterus is known from the Canary Islands to the southern part of the Gulf of Guinea (3°S). Eastern and western Atlantic populations are not in complete isolation, because C. cyanopterus is found in the equatorial zone (Parin and Belyanina 1996). Also known from off South Africa (Heemstra and Parin 1986), in the tropical Indian Ocean, and western Pacific, with the most eastern records off southern Japan, Taiwan, the Mariana and Caroline Islands, and Queensland, Australia (Parin 1996). Geographic Variation. Gibbs and Staiger (1970) found some minor differences between western and eastern Atlantic populations of Cheilopogon cyanopterus (considered at that time as disjunct) in the number of vertebrae and number of predorsal scales. Vertebral counts of 44 and 45 are almost equally common in the western Atlantic, but have a strong mode at 44 in the eastern Atlantic. The range of predorsal scales ranges from 30–41 (x̅  36.5) in the west and from 33–43 (x̅  38.1) in the east; however, in the case of the western Atlantic, counts for small juveniles with underdeveloped scales were included following Staiger (1965). Atlantic and Indo-West Pacific populations of C. cyanopterus have not been compared. Study Material. Description according to Bruun (1935), Breder (1938), Staiger (1965), Gibbs and Staiger (1970), and Kovalevskaya (1977). Additional material includes meristics of 153 Atlantic specimens examined by N. V. Parin, R. H. Gibbs, Jr., J. C. Staiger, T. N. Belyanina, and I. B. Shakhovskoy. From the northwestern Atlantic, 90 specimens (15.7–321.5 mm SL) from four collections. AMNH 8678 (1, 45.5 mm SL); Florida, Dade County Miami, Coconut Grove. Holotype of Cypselurus monroei; 22 January 1921. MCZ 58473 (1, 211 mm SL); Strait

Figure 5.2. Atlantic distribution of Margined Flyingfish, Cheilopogon cyanopterus, based on Parin and Belyanina (1996). Map by W. E. Bemis.

of Florida? MCZ 41847 (1, 274 mm SL); 18°07′N, 65°05′W; 1 May 1962. MCZ 34712 (1, 146.5 mm SL); 38°20′N, 71°04′W, 5 August 1937. ZMH 12730 (1, 144 mm SL); St. Thomas. ZMH (ISH 11/52) (1, 291 mm SL); 19°32′N, 84°04′W; 20 March 1952. USNM (?) (1, 42.5 mm SL); R/V Crawford Cruise 7, 12°00′N, 76°00′W; 26 October 1956. USNM 197843 (3, 199.6– 223.7 mm SL); Oregon Sta. 1602, 20°10′N, 92°25′W; 24 November 1956. USNM 197844 (1, 286.5 mm SL); Oregon Sta. 1841, 20°12′N, 91°59′W, 12 July 1957. USNM 197845 (1, 187.9 mm SL); Oregon Sta. 1605, 20°50′N, 93°00′W; 25 November 1956. USNM 197846 (1, 200.9 mm SL); Oregon Sta. 1599–1600, ca. 21°N, 91°W; 21 November 1956. USNM 197840 (1, 179.6 mm SL); Delaware, 39°07′N, 65°58′W; 21 September 1957. USNM 198378 (1, SL-?); Delaware; 23 September 1957. USNM 198378 (1, 180 mm SL); 39°N, 68°W. USNM 5996 (1, 181 mm SL); Jamaica. USNM 164810 (1, SL-?); near Jamaica; 23 June 1933. USNM 198455 (1, 82.6 mm SL); Delaware, 39°28′N, 69°30′W; 30 October 1956. USNM 196609 (2, SL-?); Oregon Sta. 1385; 7 September 1955. USNM 94721 (1, 321.5 mm SL); Barbados. USNM 294702 and 294703 (10, 270–288 mm SL); M/V Fregata 68-2 Sta. 103, Antilles, Bonaire; 18 February 1968. USNM 294696 (1, 285.0 mm SL); Oregon 103 Sta. 5541, 28°14′N, 80°03′W; 19 August 1965. USNM 294699 158

Family Exocoetidae uncat. (1, 24.6 mm SL); Oregon Sta. 1066, 21°42′N, 93°35′W; 22 May 1954. USNM 197841 (1, 171.5 mm SL); Delaware, 36°42′N, 70°00′W; 3 October 1957. From the southwestern Atlantic, 14 specimens (192.8–318 mm SL) from seven collections. AtlantNIRO uncat. (1, 313 mm SL); SRTM 8016, 17°27′S, 34°32′W; 6 January 1978. INBYuM uncat. (1, 306 mm SL); R/V Vodyanitsky Cruise 2 Sta. 166, 10°00′S, 30°15′W; 11 July 1977. INBYuM uncat. (1, 318 mm SL); R/V Vodyanitsky Cruise 2 Sta. 176, 07°00′S, 30°14′W; 13–14 July 1977. INBYuM uncat. (1, 304 mm SL); R/V Vodyanitsky Cruise 2 Sta. 85, 01°00′S, 25°44′W; 11 June 1977. INBYuM uncat. (1, 257 mm SL); R/V Vodyanitsky Cruise 2 Sta. 86, 01°00′S, 25°44′W; 11 June 1977. USNM 299777 (1, 301 mm SL); 07°18′S, 34°27′W; 18 January 1967. USNM 294700 (1, 308.2 mm SL); Oregon Sta. 4274, 01°49′S, 40°32′W; 16 March 1963. MNHN B.769 (2, 283.4–305.7 mm SL) Brazil, types. MNHN A.7605 (1, 295.9 mm SL); Rio de Janeiro, type. ISH 28-1950 (1, 218 mm SL); Walfangreise “Olimpic Challenger” 1950–1951, 09°34′S, 34°31′W; 18 November 1950. BMNH 1906.7.20.3 (1, 192.8 mm SL); Capt. George Caie, off coast of Brazil. CAS 167731 (1, 292 mm SL); Brazil, Bahia, Salvador; 4 May 1945. From the eastern Atlantic, 49 specimens (147– 290 mm SL) from six collections. AtlantNIRO uncat. (1, 147 mm SL); SRTM 8016, 10°19′N, 23°12′W; 28 December 1977. AtlantNIRO uncat. (1, 257 mm SL); SRTM 8060, 01°30′N, 06°59′E; 13 November 1976. AtlantNIRO uncat. (1, 263 mm SL); SRTM 8060, 03°52′N, 05°30′E; 10 November 1976. AtlantNIRO uncat. (1, 287 mm SL); SRTM 8060, 00°34′S, 04°07′E; 14 September 1978. AtlantNIRO uncat. (1, 255 mm SL); Iskra, 13°56′N, 21°30′W; 14 July 1977. ZIN 3567 (1, SL-?); St. Jago. INBYuM uncat. (1, 278 mm SL); Kerchenskyi pioner Trawl 30, 00°20′S, 19°46′W; 26 September 1977. INBYuM uncat. (1, 289 mm SL); Kerchenskyi pioner Trawl 25, 00°42′N, 20°09′W; 24 September 1977. INBYuM uncat. (1, 169 mm SL); R/V Vodyanitsky Cruise 1 Sta. 42, 05°30′N, 22°51′W; 19 January 1977. MCZ 55502 (1, 290 mm SL); 14°N, 21°W. MCZ uncat. (1, 264 mm SL); Chain 35, 01°56′N, 25°32′W; 24 February 1963. UMML 21715 (28, SL-?) Annobón. USNM 204375 (1, SL-?); GTS tr. 9; 19 November 1963. USNM 204373 (1, SL-?); GTS tr. 13 Sta. 5; 11 November 1963. USNM 204374 (1, SL-?); La Rafale, trans 9; 18 November 1963. USNM uncat. (1, SL-?); TABL 68-220 (western Africa). USNM uncat. (1, SL-?); TABL 68-217 (western Africa). USNM uncat. (1, SL-?); TRSNB 9227 (eastern Atlantic). USNM uncat. (3, SL-?); 01°24′S, 09°37′E. USNM uncat. (1, SL-?); eastern Atlantic, IG: 16501 Reg. 8265.

(1, 267 mm SL); Oregon Sta. 3595, 09°02′N, 81°26′W; 30 May 1962. USNM 294697 (1, 308.5 mm SL); Oregon 84 Sta. 4283, 01°52′N, 47°16′W; 18 March 1963. USNM 154014 (1, 247.7 mm SL); Albatross TT 11268, Yucatán Channel; 22 November 1919. USNM uncat. (1, 73.4 mm SL); Oregon Sta. 1063, 19°13′N, 95°34′W; 17 May 1954. USNM 294698 (1, 287 mm SL); R/V Gill, Cruise 6, Sta. 5, 30°00′N, 77°00′W; 16 April 1954. USNM uncat. (1, 180.4 mm SL); R/V Shoyo Maru, 13°04′N, 76°04′W; 4 February 1967. USNM 251937-F58 (1, 298.1 mm SL); R/V Crawford Cruise 16, ca. 16°N, 59°W; 28 November 1957. USNM 204591 (1, 315.8 mm SL); R/V Crawford Cruise 15, 11°00′N, 52°25′W; 7 August 1957. USNM uncat. (2, 58.7–60.8 mm SL); Silver Bay 2201, 34°34′N, 75°40′W; 15 August 1961. USNM uncat. (4, 22.1– 36.7 mm SL); Silver Bay 2201, 34°34′N, 75°40′W. USNM uncat. (1, 108.7 mm SL); Oregon Sta. 1593, UF 7245, 28°50′N, 87°58′W; 25 July 1956. USNM uncat. (4, 15.7–241 mm SL); Oregon Sta. 1052, UF 3673, 20°01′N, 92°27′W; 14 May 1954. USNM uncat. (1, 96.2 mm SL); Gill 3, Reg. 26, 30°18′N, 80°12′W; 29 July 1963. USNM uncat. (1, 90.3 mm SL); Combat Sta. 443, 25°11′N, 79°56′W; 15 May 1959. USNM uncat. (1, 41.5 mm SL); 40 miles east of Miami, 19–20 June 1956. USNM uncat. (1, 44.7 mm SL); 10 miles east of Miami, 6 May 1958. USNM uncat. (1, 113.7 mm SL); 7 miles east-southeast of Alligator Light; 24 August 1961. USNM uncat. (1, 103.1 mm SL); Oregon Sta. 1076, 24°54′N, 96°05′W; 27 May 1954. USNM uncat. (1, 24.3 mm SL); Silver Bay 2172, 35°00′N, 75°19′W; 18 July 1960. USNM uncat (1, 21.5 mm SL); Gill 3 Reg. 62, 32°39′N, 76°46′W; 10 August 1953. USNM uncat. (2, 20.9–22.6 mm SL); Silver Bay 2139, 29°55′N, 80°38′W; 12 June 1960. USNM uncat. (1, 37.3 mm SL); Gill 8, Reg. 29-30, 31°N, 79°25′W; 14 September 1954. USNM uncat (1, 111 mm SL); 09°54′N, 79°26′W; 7 July 1966. USNM uncat. (1, 25.1 mm SL); Oregon Sta. 1372, 28°55′N, 87°57′W; 21 August 1955. USNM uncat. (1, 80.1 mm SL); Silver Bay 483, 29°56′N, 80°10′W; 12 May 1959. USNM uncat. (2, 48–72.6 mm SL); Cruise IV Reg. 61, 32°53′N, 77°04′W; 8 November 1955. USNM uncat. (3, 19.5–68.3 mm SL); G-70, 25°23′N, 79°41′W. USNM 299758 (1, 175 mm SL); 12°52′N, 70°04′W; 26 November 1970. USNM 294682 (1, 265 mm SL); 09°18′N, 80°26′W. USNM 228633 (1, 290 mm SL); 09°24′N, 82°19′W. USNM 197814 (1, 68.6 mm SL); Delaware, 34°45′N, 73°41′W; 10 October 1957. USNM uncat. (2, 25.1–25.2 mm SL); Delaware, 38°00′N, 63°25′W; 1 May 1960. USNM uncat. (1, 37.9 mm SL); Oregon Sta. 1044, 23°10′N, 92°00′W; 11 May 1954. USNM 197813 (1, 80.6 mm SL); Delaware, 37°45′N, 71°49′W; 7 October 1957. USNM 159

Fishes of the Western North Atlantic Types. Exocoetus exsiliens Linnaeus, 1771. Holotype ZSL 165, dry specimen in poor condition (ca. 121 mm SL); not examined. Exocoetus spilopus Valenciennes in Cuvier and Valenciennes, 1847. Lectotype MNHN B.828 (135 mm SL) (designated by Bruun 1937); La Rochelle, France. Dorsal fin rays 15; anal fin rays 10; pectoral fin rays I 14. Predorsal scales 30; transverse scales between origin of dorsal fin and lateral line 6. Gill rakers 7 + 17 = 24. Vertebrae 30 + 15 = 45. Dorsal fin with black spot; dorsal fin height 21.3% SL; pelvic fins dark in posterior part. Paralectotypes MNHN 178 (191 mm SL) and B.829 (230 mm SL). Exocoetus bicolor Valenciennes in Cuvier and Valenciennes, 1847. Holotype MNHN B.833 (244 mm SL); Atlantic Ocean. Dorsal fin rays 15; anal fin rays 10; pectoral fin rays I 13. Predorsal scales 30; scales between origin of dorsal fin and lateral line 6. Gill rakers 7 + 18 = 25. Vertebrae 30 + 15 = 45. Dorsal fin with large black blotch; its height 15.1% SL; pectoral fins with wide unpigmented band; pelvic fins with small black spot near posterior margin. Cypselurus smithi Breder and Nichols, 1930. Holotype AMNH 9673 (201 mm SL); north of Bahamas. Dorsal fin rays 15, anal fin rays 10, pectoral fin rays I 13. Predorsal scales 26; scales between origin of dorsal fin and lateral line 7. Gill rakers 7 + 16 = 23. Dorsal fin with black spot; its height 14.2% SL; pelvic fins not pigmented. Paratype AMNH 20939 (158 mm SL); Bahamas. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Bandwing Flyingfish (English) and volador campechano (Spanish). Diagnostic Characters. Cheilopogon exsiliens differs from all other western Atlantic species of Cheilopogon in coloration of caudal fin with dark lower lobe, probably blackish in life, and white upper lobe, as well as in structure of chin barbels of juveniles (see Development section below). Description. Dorsal fin rays 13–16 (very rarely 17); anal fin rays 8–11 (usually 9–10); dorsal rays minus anal rays 3–6 (usually 5); pectoral fin rays I 12–15 (Table 5.6). Vertebrae 29–31 precaudal + 14–16 caudal = 42–46 total vertebrae (Table 5.7). Predorsal scales 23–30 (Table 5.8); scales in transverse row from dorsal fin origin to lateral line 6–8½ (Table 5.8). Gill rakers on the first arch 4–9 + 14–20 = 20–28 (Table 5.9). Measurements based on specimens 180–237 mm SL. Body elongate (greatest depth 5.6–6.6 times in SL); slightly compressed (width 1.1–1.2 in depth), nearly rectangular in cross section, almost flat ventrally. Head 3.8–4.2 times in SL and 1.1–1.3 times in

Cheilopogon exsiliens (Linnaeus, 1771) Bandwing Flyingfish Figures 5.3, 5.4 Tables 5.1, 5.6, 5.7, 5.8, 5.9 Exocoethus [sic] exsiliens Linnaeus, 1771:529 (original description; Carolina; holotype ZSL 165). Exocoetus exiliens [sic]. Statius Müller 1776:209 (Carolina). Gmelin 1789:1400 (Carolina). Valenciennes in Cuvier and Valenciennes 1847:114–115 (New Jersey). Jordan and Meek 1885:54–55 (western Atlantic). Günther 1909:147–149 (description of holotype). Exocoetus bicolor Valenciennes in Cuvier and Valenciennes, 1847:111–112 (original description; Atlantic Ocean; holotype MNHN B.833). Bruun 1937:184 (description of type). Collette et al. 1997:10 (type). Exocoetus spilopus Valenciennes in Cuvier and Valenciennes, 1847:118 (in part: original description, Atlantic Ocean). Bruun 1937:184–185 (description of type). Collette et al. 1997:13–14 (types). Exocoetus nigricans (not of Bennett 1840). Jordan and Meek 1885:63–64 (Atlantic Ocean). Jordan and Evermann 1896a:737–738 (short description; Atlantic Ocean). Exocoetus exsiliens. Lilljeborg 1891:466–469 (on Linnaean type specimen). Bruun 1934:134–135 (on Linnaean type specimen). Exonautes exiliens. Jordan and Evermann 1896b:322 (type of new genus). Jordan and Evermann 1898 (listed). Cypsilurus nigricans (not of Bennett 1840). Jordan and Evermann 1898:2836 (listed). Cypselurus nigricans (not of Bennett 1840). Jordan et al. 1930:201 (listed). Duarte-Bello 1959:46 (listed; Cuba). Cypselurus smithi Breder and Nichols, 1930:4–6, fig. 3 (original description; north of Bahamas; holotype AMNH 9673). Nichols and Breder 1930:5–7 (in key). ?Cypselurus pinnatibarbatus (not of Bennett 1831). Fowler 1936:428–429 (description; New Jersey). Cypsilurus exsiliens. Bruun 1935:44–47, pl. 3, figs. 2, 3 (description; Atlantic Ocean). Cypselurus exsiliens. Breder 1938:46–48, figs. 23–24 (description; western Atlantic). Duarte-Bello 1959:45 (listed; Cuba). Staiger 1965:691–697, figs. 7–9 (description; western Atlantic). Gibbs and Staiger 1970:445–449, fig. 4 (comparison with Cheilopogon nigricans; western Atlantic). Fahay 1975:16 (juvenile off Florida). Cypselurus cyanopterus (not of Valenciennes in Cuvier and Valenciennes 1847). Breder 1938:43, fig. 20 (in part: only specimen YPM ICH 001590). Cheilopogon exsiliens. Parin 1961:168 (listed). Tsokur 1975:5, fig. 2 (eggs; 23°54′N, 91°37′W). Fedoryako 1980:581 (Sargasso Sea). Nikolsky and Luschina 1990:135–136 (eastern equatorial Atlantic). Shiganova and Kovalevskaya 1991:97–99 (larvae and juveniles; central Atlantic). Parin and Belyanina 1996:754–758, figs. 1a, 2 (ontogeny, western Atlantic, distribution). McEachran and Fechhelm 1998:945, figure (Gulf of Mexico). Smith-Vaniz et al. 1999:171–172 (Bermuda). Parin 2003:1122, figure, map (description, western Atlantic distribution). Cotten and Comyns 2006:970–971, figs. a, f, g (in part; early life history). Fahay 2007:802–803, figs. a–f (early life history stages). Casazza and Ross 2008:352, 355 (associated with Sargassum, Gulf Stream off North Carolina). Lewallen et al. 2011 (molecular systematics). Parin and Shakhovskoy, 2016:2139 (description, distribution; east-central Atlantic).

160

Family Exocoetidae

Figure 5.3. Bandwing Flyingfish, Cheilopogon exsiliens. A. USNM 198388, 201.4 mm SL. Collected at ca. 31°01′N, 48°30′W from M/V Delaware, 21 May 1963. B. USNM 197825, 44.5 mm SL. Collected at 19°13′N, 95°34′W from M/V Oregon, Sta. 1063, July 1954. C. Pectoral and pelvic fins of USNM 197824, 71.1 mm. Collected at 19°05′N, 94°10′W from M/V Oregon, 17 July 1954. D. USNM 294889, 41.3 mm SL. Collected at 36°07′N, 73°25′W from M/V Delaware, 22 May 1960. Illustrations by H. E. Hamman.

161

Fishes of the Western North Atlantic part paler or clear. Dorsal fin light, with gray spot at the middle in smaller juveniles; almost entirely pigmented in specimens up to 100 mm SL. Anal fin slightly pigmented. Both caudal lobes clear to about 100 mm SL (lower lobe with a few gray spots); in larger juveniles lower lobe densely pigmented. Pectoral fins in specimens less than 50 mm SL lightly pigmented, with two darker crossbands; in larger specimens the area between crossbands begins to clear and by 100 mm SL the light band crossing the fin and two dark bands expanding until the anterior and posterior areas are evenly and darkly pigmented. Pelvic fins in smaller juveniles almost transparent, with pigment mostly concentrated along the rays; at about 60–120 mm SL a dark spot begins at the inner posterior corner and gradually extends to entire posterior two-thirds of the fin. Size. The maximum observed size for Cheilopogon exsiliens is 238 mm SL in the western North Atlantic, 273 mm SL elsewhere.

dorso-caudal distance. Eye diameter 3.1–3.5 times in head length, 1.0–1.2 times in interorbital space. Jaws subequal. Jaw teeth comparatively large, conical and slightly curved; dental index 0.13–0.20. Palatine teeth usually present. Dorsal fin moderately high, its longest (second or third) ray 5.1–8.0 times in SL. Anal fin originates under sixth to ninth dorsal fin ray. Pectoral fins 1.3–1.5 times in SL, reaching from middle of caudal peduncle to end of squamation, first ray unbranched, 1.7–1.9 times shorter than total fin length. Pelvic fins inserted nearer to posterior margin of opercle than to origin of caudal fin lobe (operculo-pelvic distance 1.1–1.4 times in pelvic to caudal distance), rather long (2.8–3.8 times in SL); reaching middle of anal fin base to one-fourth of caudal peduncle. Color. Body dark above, pale below. Top of snout sometimes blackish. Dorsal fin gray anteriorly and posteriorly, with a large black blotch covering the central part between third to fifth and twelfth to penultimate ray; lower margin of pigmented area often semicircular, reaching base of the fin in most specimens. Anal fin unpigmented. Caudal fin with upper lobe almost unpigmented, lower lobe black. Pectoral fins bluish black with much lighter, often not pigmented, oblique band, crossing the fin. Pelvic fins light gray on the rays, membranes transparent, often with a black spot or spots of different sizes posteriorly. Development. Parin and Belyanina (1996) described and figured juveniles (13.5, 28.0, and 50.5 mm SL). Staiger (1965) described 23 specimens (15–100 mm SL) and Shiganova and Kovalevskaya (1991) described three specimens (12.5–24.5 mm SL). Body slightly deeper than in adults (5.1–6.3 times in SL in specimens 28–126 mm SL). Head of similar proportions (3.7–4.4 times in SL). Eye larger (2.5–3.1 times in head). Dorsal fin greatly elevated, its longest ray (second or third) 2.9–4.7 times in SL. Pectoral fins 1.3–1.8 times in SL. Pelvic fins 2.2–2.8 times in SL. Paired chin barbels coalescent at bases, flat, with anterior edges straight, posterior edges serrated, bearing a flap-like appendage especially well developed in specimens smaller than 60 mm SL; length of barbels 2.2–9.8 times in SL, increasing proportionally to about 40% at 40 mm SL, then decreasing to about 12% SL at 70–100 mm SL and totally disappearing at 105–115 mm SL. In smaller juveniles (up to 50 mm SL); body with six transverse dark bars on a light background; in larger barbeled juveniles three front bars faded, but bars at pelvic fin insertion, above anal fin base, and on caudal peduncle remain more or less noticeable. Barbels densely pigmented along the edges, with central

Biology Food. Cheilopogon exsiliens feeds on zooplankton (Parin 2003). Reproduction. Matures at 185–200 mm SL. Largest ovarian eggs (about 1.6 mm in diameter) found in a spent female (201 mm SL). Larvae and small juveniles are common offshore. One ripe female (218 mm SL) was captured in January at 01°00′S, 31°30′W (Bruun 1935). A spent female (190 mm SL) was captured in the Gulf of Mexico at water temperature 23.5 °C (Tsokur 1975). Unfertilized hydrated eggs of this female were 1.7–1.85 mm in diameter, white, and evenly covered with about 53 short (0.7–0.8 mm) filaments. The structure of Cheilopogon exsiliens eggs indicates that they are probably pelagic. Range. Widely distributed in warm waters of the Atlantic Ocean. Cheilopogon exsiliens is common in the western Atlantic from about 40°N to 25°S, including the Gulf Stream and the Gulf of Mexico, but rather rare in the Caribbean Sea and in the Southern Hemisphere (Fig. 5.4). This species is absent in the Gulf of Guinea and adjacent areas, where a closely related species, C. nigricans, is found as well along the equator. Most eastern Atlantic records have been reported from around the Cape Verde Islands. The most southern record is 25°50′S, 26°45′W (Parin and Belyanina 1996), and the most northern is La Rochelle, France (lectotype of Exocoetus spilopus). Geographic Variation. Northern and southern Atlantic populations of Cheilopogon exsiliens may be spatially isolated. One of the few known South 162

Family Exocoetidae

163

Fishes of the Western North Atlantic IORAS uncat. (1, 21.3 mm SL); R/V Vityaz Cruise 2 Sta. 152, 19°40′N, 76°42′W; 12 June 1982. IORAS uncat. (1, 178 mm SL); R/V Vityaz Cruise 2 Sta. 151, 19°16′N, 76°58′W; 8 June 1982. IORAS uncat (1, 113 mm SL); R/V Vityaz Cruise 63 Sample 7, ca. 30°N, 70°30′W; 19 June 1978. IORAS uncat. (1, 111 mm SL); R/V Vityaz Cruise 63 Sta. 7881, 29°58′N, 71°02′W; 10 August 1978. IORAS uncat. (1, 125 mm SL); 38°53′N, 66°15′W. IORAS uncat (2, 172–200 mm SL); R/V Ioffe Sta. 1611, 08°58′N, 38°12′W; 26 October 2004. AMNH 9673 (1, 200.5 mm SL); north of Bahamas; 30 September 1929; holotype of Cypselurus smithi. AMNH 20939 (1, 158 mm SL); Bahamas, Diamond Shoal Light; 19 September 1929; paratype of C. smithi. UMML uncat. (1, 105.3 mm SL); Delaware, 39°28′N, 69°30′W; 30 October 1956. UMML 15281 (1, SL-?); near Bermuda. UMML 14036 (2, SL-?); Florida. MSNG 36937 (1, 183 mm SL); Porto Rico; 8 December 1886. ANSP 45466 (1, 131 mm SL); Sea Isle City, NJ; 14 September1915. RMNH 24583 (1, 197.3 mm SL); 19°10′N, 43°05′W; 26 February 1961. ZMH 12780 (1, 171 mm SL); zwischen Trinidad und San Domingo. ZMH 12744 (1, 232 mm SL); Laguayra, 1859. ZMH 8832 (1, 208 mm SL); St. Thomas. MCZ 156571 (1, 19.5 mm SL); 30°50′N, 80°21′W; 23 July 1980. USNM 294730 (4, 195.7–225.3 mm SL); Discoverer, 13°24′N, 39°30′W; 7–8 February 1969. USNM 294733 (1, 225 mm SL); Discoverer, 10°51′N, 42°29′W; 14–15 February 1969. USNM uncat. (1, 218.8 mm SL); Discoverer, 12°45′N, 40°38′W; 10–11 February 1969. USNM 294701 (6, 179.4–237.9 mm SL); Discoverer, 13°49′N, 39°03′W; 5–6 February 1969. USNM uncat. (1, SL-?); ATEX drift; 8–10 February 1969. USNM 294692 (1, 213.3 mm SL); Discoverer, 12°57′N, 40°09′W; 9–10 February 1969. USNM 200853 (3, 195.1–235 mm SL); Anton Bruun Cruise 9, Sta. 479, 24°49′–25°02′N, 62°10′–30°08′W; 16–23 January 1965. USNM 198388 (1, 200.3 mm SL); Delaware, 31°00′N, 48°30′W; 21 May 1963. USNM 30975 (1, SL-?); Caribbean Sea, Panama. USNM (?) uncat. (1, 60.5 mm SL); Oregon 4301, 07°34′N, 54°13′W. USNM (?) uncat. (1, 52.7 mm SL); Oregon Sta. 1616; 6–7 December 1956. USNM (?) uncat. (2, 88.6–110.5 mm SL); ca. 6 miles southeast of Alligator (Florida); 30 June 1960. USNM (?) uncat. (1, 121.4 mm SL); Oregon Sta. 1587, 28°17′N, 88°27′W; 22 July 1956. USNM (?) uncat. (1, 42.7 mm SL); Gill 7, 26°30′N, 76°40′W; 12–13 June 1954. USNM (?) uncat. (1, 18.8 mm SL); Silver Bay 3327, 32°40′N, 78°16′W; 24 October 1962. USNM (?) uncat. (1, 18.2 mm SL); Cruise 2 Sta. 72, 33°49′N, 75°59′W; 5–10 May 1953. USNM (?) uncat. (1, 15.3 mm SL); Cruise 2 Sta. 63, 33°14′N, 76°25′W; 8 May 1953. USNM (?) uncat. (1, 45.2 mm SL); Oregon Sta. 1069, 19°13′N, 95°34′W; 17

Figure 5.4. Distribution of Bandwing Flying f ish, Cheilopogon exsiliens, based on Parin and Belyanina (1996). Map by W. E. Bemis.

Hemisphere specimens (USNM 299769, from 18°42′S, 04°14′W) is much larger (273 mm SL) and has more dorsal fin rays (17) than any northern C. exsiliens examined. These populations need to be compared. Study Material. Description according to Bruun (1935), Breder (1938), Staiger (1965), Gibbs and Staiger (1970), Shiganova and Kovalevskaya (1991), and Parin and Belyanina (1996). Additional material includes meristics of 90 specimens examined by N. V. Parin, R. H. Gibbs, Jr., J. C. Staiger, T. N. Belyanina, and I. B. Shakhovskoy. From the northern Atlantic, 83 specimens (15.3– 237.9 mm SL) from 10 collections. INBYuM uncat. (2, 223–228 mm SL); Kerchenskyi pioner, Trawl 4, 16°42′N, 32°57′W; 9 September 1977. IORAS uncat. (1, 28 mm SL); R/V Mendeleev Cruise 1 Sta. 60, 07°54′N, 53°16′W; 3 April 1969. IORAS uncat. (1, 38 mm SL); R/V Kurchatov Cruise 14 Sta. 1259, 19°16′N, 80°29′W; 21–22 March 1973. IORAS uncat. (2, 48–53 mm SL); R/V Vityaz Cruise 63 Sta. 7848, 30°35′N, 67°20′W; 17 July 1978. IORAS uncat. (2, 52–53 mm SL); R/V Vityaz Cruise 63 Sta. T-ka L, off Bermuda; 27 June 1978. IORAS uncat. (1, 50.5 mm SL); R/V Vityaz Cruise 63 Sta. 7893, 31°22′N, 66°40′W; 12–13 August 1978. IORAS uncat. (1, 120 mm SL); R/V Vityaz Cruise 63 Sta. B, 30°44′N, 68°39′W; 28–29 July 1978. IORAS uncat. (1, 123.8 mm SL); R/V Vernadsky Cruise 27 Sta. 2229, 04°43′N, 43°47′W. 164

Family Exocoetidae 1847:135–136 (Gulf of Mexico). Günther 1866:286 (Atlantic and Indian oceans). Cypselurus furcatus. Jordan and Gilbert 1883:380–381 (in part; description; Atlantic Ocean). Breder 1938:69–73, figs. 35–36 (in part: specimen YPM ICH 001636 is Cheilopogon melanurus; description of juveniles, western Atlantic). Duarte-Bello 1959:45 (listed; Cuba). Staiger 1965:697–705, figs. 13–15 (juveniles; Atlantic Ocean). Gibbs and Staiger 1970:461 (comparison with Cheilopogon heterurus and Cheilopogon melanurus; Atlantic). Hoese and Moore 1977:151 (Gulf of Mexico). Gibbs 1978: EXOC Cyp 4 (western Atlantic). Cypsilurus furcatus. Bruun 1935:58–60, pl. 5, figs. 2, 4 (description, Atlantic distribution). Cheilopogon furcatus. Parin 1961:174 (listed). Fedoryako 1980:581 (Sargasso Sea). Heemstra and Parin 1986:393, figure (short description; South Africa). Gorelova and Grudtsev 1987:481 (feeding; Atlantic). Kovalevskaya and Shiganova 1988:88– 95, figs. 1, 2b, 2d–f (early life history [in part(?)]): only larva and juveniles [4.3 mm SL and 12.5–34.5 mm SL]); central Atlantic). Nikolsky and Luschina 1990:135–136 (eastern equatorial Atlantic). Parin and Gibbs 1990:548 (Atlantic Ocean). Shiganova and Kovalevskaya 1991:99 (early life history; central Atlantic). McEachran and Fechhelm 1998:946, figure (Gulf of Mexico). Parin and Belyanina 2000:162, 164 (distribution and comparison with Cheilopogon heterurus and C. melanurus; Atlantic). Collette et al. 2003:102 (Navassa Island, West Indies). Parin 2003:1123, figure, map (description, western Atlantic). Cotten and Comyns 2006:972–973, figs. a–h (early life history). Fahay 2007:804–805, figs. a–e (early life history stages). Casazza and Ross 2008:352, 355 (associated with Sargassum, Gulf Stream off North Carolina). Gallardo-Torres et al. 2014:82 (Yucatán, Mexico, photographs). Parin and Shakhovskoy 2016:2140 (description, east-central Atlantic). Cheilopogon furcatus furcatus. Parin and Belyanina 1998:569, figs. 1–6 (separation of nominal subspecies in the Atlantic from new subspecies in the Indian and Pacific oceans).

May 1954. USNM 197822 (1, 34.5 mm SL); Oregon Sta. 1074, 24°00′N, 96°50′W; 25 May 1954. USNM 197826 (1, 100.5 mm SL); Oregon Sta. 1035, 26°40′N, 92°00′W; 8 May 1954. USNM 197824 (1, 71.0 mm SL); Oregon, 19°05′N, 94°10′W; 17 May 1954. USNM (?) uncat. (1, 148.5 mm SL); ca. 23°30′N, 82°40′W; 12 September 1967. USNM (?) uncat. (1, 183.5 mm SL); Silver Bay 467–468, 29°50′–29°46′N, 80°11′–12′W; 16–17 June 1958. USNM 294689 (1, 197.3 mm SL); Oregon Sta. 5478, 27°02′N, 77°10′W; 16 June 1965. USNM 294695 (1, 203.8 mm SL); Oregon Sta. 3531, 25°00′N, 86°09′W; 10 May 1962. USNM 294714 (1, 207 mm SL); Gill Cruise 5, Reg. 15, 29°00′N, 79°48′W; 2 February 1954. USNM 197847 (1, 185.2 mm SL); Oregon Sta. 1068, 22°10′N, 96°20′W; 23 May 1954. USNM (?) uncat. (2, 205.5–230.8 mm SL); Discoverer, 12°27′N, 41°21′W; 12–13 February 1969. USNM (?) uncat. (1, 202.3 mm SL); Discoverer, 13°07′N, 39°59′W; 9 February 1969. USNM 197815 (1, 94.5 mm SL); Delaware, 39°07′N, 65°58′W; 21 September 1957. USNM 251937-F51 (1, 42.5 mm SL); 36°07′N, 73°25′W; 22 April 1960. USNM 197825 (1, 40 mm SL); 19°13′N, 95°34′W. USNM (?) uncat. (1, 147 mm SL); Gerda 908. USNM 251621 (1, 123 mm SL); 32°N, 64°W. USNM uncat. (1, 33.5 mm SL); Delaware, 38°00′N, 63°25′W; 1 May 1960. USNM uncat. (1, 22.1 mm SL); Atlantis, 32°00′N, 63°20′W. USNM uncat. (1, 18 mm SL); Delaware, 38°58′N, 66°26′W; 9 June 1957. USNM 257606 (1, 19 mm SL); 32°10′N, 64°10′W. From the southern Atlantic, seven specimens (13.5–273 mm SL) from four collections. ZMH (ISH 239/51) (1, 214 mm SL); 18°49′S, 38°32′W; 17 December 1951. ZMH (ISH 380/53) (1, 253 mm SL); 31°25′S, 29°04′W; 8 December 1952. IORAS uncat. (1, 13.5 mm SL); R/V Kurchatov Cruise 11 Sta. 826, 25°18′S, 26°33′W; 15 November 1971. IORAS uncat. (1, 37 mm SL); R/V Kurchatov Cruise 11 Sta. 985, 16°22′S, 34°58′W; 10 January 1972. IORAS uncat. (1, 32 mm SL); SRTM 8018, 17°48′S, 28°02′W; 1 May 1977. MCZ 155998 (1, 184.4 mm SL); Atlantis II Cruise 37, 13°55′S, 30°39′W; 2 March 1967. USNM 299769 (1, 273 mm SL); 18°43′S, 04°14′W; 4 April 1971.

Types. Type specimens of both Exocoetus furcatus Mitchill, 1815 and E. nuttallii Lesueur, 1821 are not known to be extant (Eschmeyer et al. 2018). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Spotfin Flyingfish (English) and volador ala manchada (Spanish). Diagnostic Characters. Cheilopogon furcatus differs from other Atlantic Cheilopogon species (except for C. heterurus and C. melanurus) in having minute jaw teeth, a lower dorsal fin, and differences in the structure of chin barbels in juveniles. It differs from C. heterurus and C. melanurus in pectoral fin coloration, length of first pectoral fin ray, and length of chin barbels in juveniles. Description. Dorsal fin rays 12–14 (very rarely 15); anal fin rays 8–11, usually 9–11; dorsal rays minus anal rays 2–5; pectoral fin rays I 13–16 (Table 5.10). Vertebrae 27–30 precaudal + 15–17 caudal = 43–46 total, usually 44 or 45 (Table. 5.11). Predorsal scales 26–33, usually 27–31; scales in transverse row 7–9, usually 7½–8½ (Table 5.12). Gill rakers on first arch

Cheilopogon furcatus furcatus (Mitchill, 1815) Spotfin Flyingfish Figures 5.5, 5.6 Tables 5.1, 5.10, 5.11, 5.12, 5.13 Exocoetus furcatus Mitchill, 1815:449, pl. 5, fig. 2 (original description; northwestern Atlantic, “New York”). ?Exocoetus nuttallii Lesueur, 1821:10–11, pl. 4, fig. 1 (original description, Gulf of Mexico). Exocoetus furcatus. Valenciennes in Cuvier and Valenciennes

165

Fishes of the Western North Atlantic

166

Family Exocoetidae

Figure 5.5. Spotfin Flyingfish, Cheilopogon furcatus furcatus. A. USNM 198374, 161.6 mm SL. Dipnetted at night at 24°54′N, 96°05′W from M/V Oregon, Sta. 1076, 27 May 1954. B. USNM 199243, 81.3 mm SL. Collected at 39°07′N, 65°58′W from M/V Delaware, 14 August 1957. C. USNM 309903, 47.0 mm SL. Collected at 26°40′N, 92°00′W from M/V Oregon, Sta. 1035. D. Pectoral and pelvic fins of USNM 197816, 29.9 mm SL. Collected at 32°00′N, 63°20′W from R/V Atlantis, 14 May 1957. E. Pectoral and pelvic fins of USNM 196627, 18.8 mm SL. Collected from Gulf of Mexico off the Mississippi River Delta at 28°46′N, 88°40′W from M/V Oregon, 30 August 1954. Illustrations by H. E. Hamman.

167

Fishes of the Western North Atlantic pale with black spots; pectoral fins with dark lower part and one or two oblique dark bands; pelvic fins also with dark spots and bands; caudal fin with one spot at the end of upper lobe and two spots on lower lobe. Juveniles with paired chin barbels consisting of firm pointed stem and with skin folds along the outer margins; barbel length 2.5–11.1 times in SL at 50–150 mm SL. Barbels disappear at 112–175 mm SL. Range. Cheilopogon furcatus is a true oceanic species not connected with inshore waters during any period of life. It is widespread in all tropical oceans except for the easternmost Pacific Ocean (Parin and Belyanina 1998: fig. 7). The ranges of the three subspecies of C. furcatus represent spatially isolated environments separated by the relatively colder waters of the Benguela Current off southwest Africa and the shallow-water inner seas of Indonesia and the Philippines. Most records of Cheilopogon furcatus furcatus in the Atlantic Ocean are from the warm-water northwestern area (Fig. 5.6). It is most common in the North Central Gyre, including the Gulf Stream (individual juvenile strays are carried as far north as 44°30′N), the Gulf of Mexico, and the Caribbean Sea; adults have been recorded between 15°N and 5°S. The most eastern record on the equator is at 01°W. South of the equator, there are records from the southwestern Atlantic south to 17°48′S. Eastern

4–8 upper + 14–17 lower = 18–25, usually 20–23, total (Table 5.13). Measurements based on specimens 198–270 mm SL. Body elongate (greatest depth 4.8–6.3 times in SL); nearly rectangular in cross section, almost flat ventrally. Head length 4.0–4.5 times in SL, slightly less than dorso-caudal distance (1.1–1.3 times in the latter). Eye 2.8–3.1 times in head length, 0.9–1.2 times in interorbital distance. Lower jaw projects slightly beyond upper jaw when mouth is closed. Jaw teeth diminutive, mostly conical, tooth index 0.06–0.10. Palatine teeth absent. Dorsal fin low, its longest ray (second or third) 8.3–13.8 times in SL. Anal fin originating under fifth to seventh dorsal fin ray. Pectoral fins 1.3–1.5 times in SL, extending from middle to end of caudal peduncle, its first ray unbranched. Pelvic fins 2.7–3.3 times in SL, inserted nearer to posterior margin of opercle than origin of caudal fin base. Color. Body dark iridescent blue above, pale silvery below in life. Dorsal and caudal fins gray; anal fin transparent; pectoral fins black to gray (deep blue in life) with unpigmented oblique crossband reaching or nearly reaching upper margin of the fin and very broad unpigmented margin along rear edge of the fin. Size. The maximum observed size for Cheilopogon furcatus furcatus is 293 mm SL in the western Atlantic (Bruun 1935). Biology Food. In the Atlantic, from 20°N to 30°S and 40°W to 06°E, Cheilopogon furcatus feeds mainly on small fishes such as Vinciguerria, as well as on amphipods and siphonophores (Gorelova and Grudtsev 1987). Reproduction. Breeding occurs throughout the year in tropical waters of the Indo-Pacific; embryonic development takes about five to seven days, eggs are demersal and laid on drifting algae or other floating objects (Gorbunova and Parin 1963; Kovalevskaya 1980). The smallest recorded ripe male was 193 mm SL (Bruun 1935). The eggs of Cheilopogon furcatus from the central Atlantic are 1.7–1.9 mm in diameter, with two bunches of filaments arranged at opposite poles: one with 15–17 very short filaments and the other with 13–15 filaments of different lengths and widths (the longest more than 20 mm). Eggs and early larvae were captured in this area at water temperatures of 22.6 to 22.8 °C (Kovalevskaya and Shiganova 1988; Shiganova and Kovalevskaya 1991). Development. Juveniles (30–115 mm SL) have six transverse vertical bars on body. Dorsal and anal fins

Figure 5.6. Distribution of Spotfin Flyingfish, Cheilopogon furcatus furcatus, based on Parin and Belyanina (1998). Map by W. E. Bemis.

168

Family Exocoetidae

and southern boundaries for the species need to be clarified (Parin and Belyanina 1998). The first records of C. f. furcatus from the Mediterranean Sea were reported from two specimens from northeast off Zarzis in the Gulf of Gabes in 2004 (Ben Souissi et al. 2005). However, these specimens are most likely C. heterurus and not C. f. furcatus (Shakhovskoy and Malikova 2018).

Geographic Variation. Parin and Belyanina (1998) recognized three subspecies of Cheilopogon furcatus: C. f. furcatus, C. f. neimani Parin and Belyanina and C. f. fisunovi Parin and Belyanina. The Atlantic subspecies, C. f. furcatus differs in having a relatively larger eye and a pectoral fin pigmentation pattern in juveniles that lacks the transverse triangular pigmented edge in the anterior third of the pectoral fin 169

Fishes of the Western North Atlantic, Part 10 Table 5.13. Number of first arch gill rakers of Cheilopogon furcatus, based on Parin and Belyanina (1998). Dashes indicate a value of zero. First arch gill rakers Species and locality Cheilopogon furcatus furcatus Atlantic Ocean Cheilopogon furcatus neimani Indian Ocean Cheilopogon furcatus fisunovi Pacific Ocean Species total

17

18

19

20

21

22

23

24

25

26

—

2

4

15

17

13

15

1

1

—

1

3

4

16

15

13

6

3

1

2

— 1

1 6

14 22

13 44

18 50

9 35

10 31

2 6

— 2

1 3

(see Parin and Belyanina 1998: fig. 5). Cheilopogon f. neimani, known from almost all warm parts of the Indian Ocean, has significantly longer chin barbels (up to 50% SL with x̅ 38% compared to not more than 34%–37% with x̅ of 25%–26% for Atlantic and Pacific populations). The Pacific Ocean subspecies C. f. fisunovi has more vertebrae (usually 45–46, x̅ 45.3, rather than 44–45, x̅ 44.4). Study Material. A total of 58 specimens (35–270 mm SL) from the Atlantic were examined (see Parin and Belyanina [1998] for full list and localities). Data also used from Bruun (1935), Breder (1938), and Staiger (1965).

Springer 1959:166 (comparison with Cheilopogon melanurus). Staiger 1965:703–714 (in part; western Atlantic). Gibbs and Staiger 1970:456 (distribution; Atlantic). Gibbs 1978: EXOC Cyp 5 (Bermuda). Cheilopogon furcatus (not of Mitchill 1815). Ben Souissi et al. 2005:1144–1149, fig. 1 (description; Tunis, Mediterranean). Cheilopogon heterurus. Parin 1961:174 (listed). Parin 1973: 265 (northeastern Atlantic and Mediterranean Sea). Wheeler 1978:182–183 (northeastern Atlantic). Fedoryako 1980:580–581 (in part?; Sargasso Sea). Smith-Vaniz et al. 1999:171–172, color pl. 3–28 (Bermuda). Parin and Belyanina 2000:215–230, figs. 3, 4a (description; northeastern Atlantic and off Bermuda). Parin 2003:1124, figure, map (description, western Atlantic). Cotten and Comyns 2006:974–975, figs. D–G (in part; early life history). Parin and Shakhovskoy 2016:2141 (description, east-central Atlantic). Shakhovskoy and Malikova 2018:232–234, figs. 1, 3 (eastern Mediterranean Sea).

Cheilopogon heterurus (Rafinesque, 1810) Blotchwing Flyingfish Figures 5.7, 5.8 Tables 5.1, 5.14, 5.15, 5.16, 5.17

Types. Exocoetus heterurus Rafinesque, 1810. No type specimens extant. Exocoetus procne De Filippi and Verany, 1859. Syntypes (3) MZUT 451, not examined. Exocoetus maculipinnis Vincig uerra, 1883. Holotype MSNG 13998 (126 mm SL). Dorsal fin rays 14; anal fin rays 10; pectoral fin rays I 15. Predorsal scales 36; transverse scales between dorsal fin and lateral line 8. Head length 21.3% SL; eye diameter 6.9% SL; body depth 17.7% SL; no barbel. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Blotchwing Flyingfish (English) and volador ala lunada (Spanish). Diagnostic Characters. Cheilopogon heterurus differs from other species of Cheilopogon (except for C. melanurus and C. furcatus) in having minute jaw teeth, a lower dorsal fin, and in the structure of chin barbels of juveniles. It differs from C. furcatus in pectoral fin coloration and length of chin barbels. Cheilopogon heterurus is very similar in appearance to C. melanurus, differing mainly in maximum body length (350 mm SL and 265 mm SL, respectively), number of vertebrae (46–50 rather than 44–47),

Exocoetus heterurus Rafinesque, 1810:58–59 (original description; Palermo, Sicily; no type specimens extant). Jordan and Meek 1885:59–61 (in part; western North Atlantic). D’Ancona 1931:168–173, pl. 10, figs. 14, 15; pl. 11, figs. 2–12 (in part; development). Exocoetus volitans not of Linnaeus 1758. Valenciennes in Cuvier and Valenciennes 1847:83–92 (Mediterranean Sea, Atlantic coast of France, to the English Channel). Fries et al. 1892:357– 361, fig 94 (northern Europe). Exocoetus procne De Filippi and Verany 1859:194–195 (original description; Nice, France: syntypes MZUT 451). Exocoetus maculipinnis Vinciguerra, 1883:577–582 (original description; Tunis, Tunisia: holotype MSNG 13998). Exocoetus rondeletii (not of Valenciennes in Cuvier and Valenciennes 1847). D’Ancona 1931:165–168, pl. 10, figs. 1–7 (in part; eggs and larvae; Mediterranean). Exocoetus exsiliens (not of Linnaeus 1771). Lilljeborg 1891:466 (northern Europe). Cypsilurus heterurus. Bruun 1935:61–64, pl. 4, fig. 2 (Mediterranean Sea, Bay of Biscay, Bermuda). Bruun 1936: pl. 234 (Mediterranean Sea, eastern North Atlantic). Tortonese 1937:234 (distribution; Mediterranean). Bruun 1938:295–299, fig. 1 (description; Oslo Fjord). Nielsen 1963:156 (Denmark). Cypselurus heterurus. Fowler 1936:1242–1243 (northwestern Africa). Breder 1938:56–69 (in part, western Atlantic).

170

Family Exocoetidae number of predorsal scales (usually 30–36 rather than 26–31), and in head length (usually less than 22.5% SL compared to more than 22% SL). Description. Dorsal fin rays 12–15, usually 13–14; anal fin rays 8–10; pectoral fin rays I 13–16, usually I 14–16 (Table 5.14). Vertebrae 30–33 + 15–17 = 46–50, usually 47–49 (Table 5.15). Predorsal scales 28–38; scales in transverse row from dorsal fin origin to lateral line 7–9½, usually 8–9 (Table 5.16). Gill rakers 5–7 + 15–17 = 19–26, usually 20–24 (Table 5.17). Measurements are based on 18 western North Atlantic specimens (290–348 mm SL) from Parin and Belyanina (2000). Body elongate (greatest depth 5.3–6.5 times in SL); slightly compressed (width 1.1–1.4 times in depth). Head short (4.3–4.9 times in SL and 1.2–1.3 times in dorso-caudal distance). Eye diameter 3.1–3.8 times in head length, 1.0–1.2 times in interorbital space. Lower jaw pointed, projecting slightly beyond the upper jaw when mouth is closed. Jaw teeth diminutive, conical and blunt, rarely with one or two supplementary cusps; dental index 0.06–0.08. Palatine teeth absent. Dorsal fin low, its longest (second or third) ray 7.9–10.8 times in SL. Anal fin originates under fifth to seventh dorsal fin ray. Pectoral fin 1.3–1.5 times in SL, reaching from middle to end of caudal peduncle, its first ray unbranched, 1.8–2.1 times shorter than total fin length. Pelvic fins inserted nearer to posterior margin of opercle than to origin of lower caudal fin lobe (opercle to pelvic distance 1.1–1.4 times shorter than pelvic to lower caudal-lobe origin distance), very long (2.7–3.3 times in SL), reaching from middle to end of anal fin base. Color. In adults, body dark above, pale below. Dorsal and anal fins gray, lacking pigment on membranes. Caudal fin uniformly dark. Pectoral fins from very light gray to dark gray, with indefinite pale triangular crossband reaching third to fifth ray, and a narrow unpigmented margin along the posterior edge of the fin. Pelvic fins pale, sometimes (in smaller adults) with a black spot distally. Development. Body of juveniles (25–145 mm SL) a little more elongate than in adults, maximum depth 5.4–7.2 times in SL, slightly increasing with growth. Head short (4.4–5.1 times in SL); its length not changing with growth. Eye diameter 2.1–3.5 times in head length and 0.9–1.3 times in interorbital space. Dorsal fin higher than in adults, its longest ray 5.0–7.2 times in SL. Pectoral fins 1.4–2.0 times in SL. Pelvic fins 1.8–2.3 times in SL, gradually displaced posteriorly with growth. Paired chin barbels consisting of firm pointed stem with soft skin fold along posterior edge; in small juveniles, fold is semicircular and much broader than stem width; in

larger specimens from the eastern North Atlantic it gradually tapers to a point (the largest barbeled specimen is 84 mm SL, whereas all specimens more than 90 mm SL lack barbels); length of barbels 8–50 (usually 10–25) times in SL. Pigmentation of larvae and juveniles changes significantly during development and is reviewed by Parin and Belyanina (2000: figs. 3, 4a). At 4.5 mm SL, when the finfold and yolk sac are still present, the body and head are evenly covered by large stellate melanophores, and the pectoral and pelvic fins and finfold are not pigmented. From 6.0–7.5 mm SL the body is evenly covered by melanophores present on the membranes of the upper rays of pectoral fins; the pelvic fins are evenly dark in color; melanophores are present near the bases of the anterior rays of the dorsal fin; anal and caudal fins without pigment. At 9–10 mm SL, lightly colored barbels appear. In small juveniles (13–15 mm SL) the barbel tips are black and the lobes are not developed. From 9–15 mm SL, the body is evenly covered by large stellate melanophores; the pelvic fins are almost black; the pectoral fins also intensely pigmented except for the membranes of the lower rays. Most specimens in this size range have a light transverse spot in the anterior part of the pectoral fin. Pigmentation in the anterior part of the dorsal fin is followed by the appearance of pigment near the bases of rays in its posterior part; anal fin without pigment or with melanophores near the bases of the anterior rays; caudal fin without pigment, although some melanophores are present along the lower rays in larger specimens. From 19–35 mm SL, transverse bands of pigment are visible on the body. The most notable bands are under the bases of pectoral fins, in the middle of the pectoral-pelvic distance, and above the bases of pelvic fins. Bands posterior to these are usually poorly visible. The position of bands on the body are consistent with pigment spots on pectoral fins if pectoral fins are pressed to the body. These spots form characteristically concentric semi-rings near the bases of rays and at the level of the middle of the first ray (usually more weakly developed or interrupted in its middle part), and the third, less visible semiring is situated at the level of the end of the first ray. A large dark spot covers the posterior lower part of the pectoral fin, upper posterior part of fin lightly colored with small spots near the ends of the second and sixth to seventh rays and on the membranes of the third to fourth rays. Pelvic fins dark, with lightly colored regions present at the ends of rays. Anterior part of the dorsal fin pigmented as well as the margins of the membrane on the posterior part. Anal 171

Fishes of the Western North Atlantic

172

Family Exocoetidae

Figure 5.7. Blotchwing Flyingfish, Cheilopogon heterurus. A. No. 469, 298 mm SL. From Bruun (1935), illustration by P. H. Winther. B. IORAS uncatalogued, 101 mm SL. Collected at 18°55′N, 18°26′W. From Parin and Belyanina (2000), illustration by T. N. Belyanina. C. SMF 240, 75 mm SL. Collected from Mediterranean. From Parin and Belyanina (2000), illustration by T. N. Belyanina. D. USNM 257579, 28.5 mm SL. Collected at 31°48′N, 63°43′W. From Parin and Belyanina (2000), illustration by T. N. Belyanina. E. USNM 299702, 13.4 mm SL. Collected at 33°36′N, 65°00′W. From Parin and Belyanina (2000), illustration by T. N. Belyanina.

173

Fishes of the Western North Atlantic

Table 5.15. Vertebral counts of Cheilopogon heterurus and C. melanurus, based on Parin and Belyanina (2000). Dashes indicate a value of zero. Vertebrae Precaudal Species and locality

29

30

Cheilopogon heterurus Northeastern Atlantic Northwestern Atlantic

— —

1 —

Species total

—

31

Caudal

Total

32

33

14

15

16

17

44

45

46

47

48

49

50

4 9 5 16

1 1

— —

2 —

10 11

3 11

— —

— —

1 —

7 4

6 19

1 9

— 1

1

9

25

2

—

2

21

14

—

—

1

11

25

10

1

Cheilopogon melanurus Northwestern Atlantic Southwestern Atlantic Eastern Atlantic

16 108 2 9 2 6

86 9 5

6 2 —

— — —

4 — 1

72 127 3 18 6 5

13 1 1

— — 1

51 215 2 8 4 9

79 12 2

— — —

— — —

— — —

Species total

20 123 100

8

—

5

81 150

15

1

57 232

93

—

—

—

174

Family Exocoetidae

fin without pigment or with melanophores near the bases of the anterior and the ends of posterior rays. Caudal fin lightly colored with few melanophores along the lower rays of the lower lobe. In larger juveniles (50–80 mm SL), bars on pectoral fins merge in the lower portion of fin to form several spots: one spot near base, two along the anterior margin, and one near the tip of the fin. Pelvic fins gradually brighten during development. The anal fin is clear, and the dorsal fin has pigment at bases of anterior and posterior rays and near the tips of rays 3 to 9. Caudal fin with spots on upper and lower lobes (Parin and Belyanina 2000). Size. The maximum observed size for Cheilopogon heterurus is 350 mm SL in the western North Atlantic, 340 mm SL in the eastern North Atlantic.

(some eggs were still present in ovaries). Larvae and small juveniles were captured off Bermuda from April to July (Parin and Belyanina 2000). Eggs are demersal, laid on aquatic vegetation. Development of fertilized eggs takes about 17 days. Diameter of eggs in the Mediterranean 1.4–1.5 mm, the shell evenly covered with about 90 thin filaments 1.3–1.4 mm in length (D’Ancona 1931; note that there is confusion in D’Ancona’s descriptions of eggs and larvae of C. heterurus and Hirundichthys rondeletii). Relationship to Humans. A man who was wading in the Mediterranean Sea was reportedly struck by a flyingfish that caused a temporal bone fracture (Goldenberg et al. 1998). A school of three fish leapt out of the water, one of which hit the man in the right ear. The species was reported to be Cheilopogon heterurus, but this identification could not be confirmed because the specimen was not collected. Range. A neritic species, rarely found far from the coast. Distributed in the subtropical and temperate North Atlantic with two disjunct populations (Fig. 5.8). In the western Atlantic known from Bermuda (where larvae and small juveniles are abundant in coastal waters); Bruun (1938) reported one specimen from the Gulf of Maine near 40°N. In the eastern North Atlantic, Cheilopogon heterurus is found in the western Mediterranean Sea (including the Adriatic Sea) and the eastern Mediterranean (Shakhovskoy and Malikova 2018), along the shores of northwest Africa (from Mauritania to Gibraltar), and off the Azores (Parin and Belyanina 2000). Individual stragglers known as far north as Denmark and southern Norway (Bruun 1938; Nielsen 1963). This (or closely related) species may also be distributed in the southwestern subtropical Atlantic (see Cheilopogon melanurus account below). Geographic Variation. No differences were found

Biology Parasites. A cymothoid isopod, Glossobius albinae, was described by Kononenko (1986) from the mouth of a flyingfish identified as Cheilopogon heterurus from the east-central part of the Atlantic. This isopod name was later placed in the synonymy of Glossobius auritus Bovallius by Bruce and Bowman (1989). Interestingly, the hosts of most of their isopod material were recorded as Cypselurus comatus. Food. Cheilopogon heterurus feeds on zooplankton (Parin 2003). Reproduction. Matures by about 280–300 mm SL. Spawns in inshore waters (Bruun 1935). A ripe female (304 mm SL) was captured in May off Arzilla, Morocco, where Cheilopogon heterurus spawns in June and July (Bruun 1935). Eggs and larvae were found in the Mediterranean (Gulf of Naples) in June and July (D’Ancona 1931). Specimens caught off of Bermuda (USNM 294784) in May had just spawned 175

Fishes of the Western North Atlantic between populations in the western and eastern North Atlantic. Small shifts in limits and average values of a few meristics (vertebrae 47–50, x̅ 48.2, rather than 46–49, x̅ 47.5; predorsal scales 29–38, x̅ 33.6, rather than 28–34, x̅ 32.0; gill rakers 19–23, x̅ 20.7, rather than 20–26, x̅ 22.0) are considered to be of little significance (Bruun 1935; Parin and Belyanina 2000). Study Material. Description based on 55 specimens (28–348 mm SL; see Parin and Belyanina [2000] for full list and localities) and data of Bruun (1935). Additional material includes meristics of 33 western Atlantic specimens examined by N. V. Parin.

Gibbs 1978: EXOC Cyp.6 (tropical western Atlantic). Monteiro et al. 1998:399–400 (distribution; northeastern Brazil). Cypselurus comatus (not of Mitchill 1815). Staiger 1965:681 (in part: only the specimen 242.5 mm SL from Gulf of Mexico). Cheilopogon comatus (not of Mitchill 1815). McEachran and Fechhelm 1998:943 (in part; Gulf of Mexico). Cheilopogon melanurus. Parin and Gibbs 1990:584 (tropical eastern Atlantic). McEachran and Fechhelm 1998:947, figure (Gulf of Mexico). Smith-Vaniz et al. 1999:172 (Bermuda). Parin and Belyanina 2000:220–227, figs. 4b, 6 (description, tropical Atlantic distribution). Collette 2002:291–292, figure, map (description, Gulf of Maine). Collette et al. 2003:102, fig. 32 (Navassa Island, West Indies). Parin 2003:1125, figure, map (description, western Atlantic). Casazza et al. 2005:363–375, pl. 1, 2 (reproduction and mating behavior; North Carolina). Cotten and Comyns 2006:976–977, figs. A–F (early life history). Fahay 2007:806–807, figs. A–E (early life history stages). Casazza and Ross 2008:352–359 (associated with Sargassum, Gulf Stream off North Carolina). Parin and Shakhovskoy 2016:2142 (description, east-central Atlantic). Shakhovskoy and Malikova 2018:230-232, figs. 2, 3 (north-central Atlantic).

Cheilopogon melanurus (Valenciennes in Cuvier and Valenciennes, 1847) ATLANTIC FLYINGFISH

Figures 5.8, 5.9 Tables 5.1, 5.14, 5.15, 5.16, 5.17

Types. Exocoetus melanurus Valenciennes in Cuvier and Valenciennes, 1847. Holotype MNHN A. 7603 (215 mm SL); New York. Dorsal fin rays 13; anal fin rays 10; pectoral fin rays I 15. Predorsal scales 30; transverse scales between dorsal fin origin and lateral line 7. Gill rakers 5 + 16 = 21. Vertebrae 46. Head length 23.2% SL; eye diameter 7.7% SL. Pectoral fins pigmented, darker posteriorly, with lighter oblique strip to sixth ray. Exocoetus lutkeni Jordan and Evermann, 1896. Holotype ANSP 7502 (180 mm SL); Cape San Antonio, Cuba (later changed for California). Dorsal fin rays 14; anal fin rays 9; pectoral fin rays I 14. Predorsal scales 29; scales between dorsal fin origin and lateral line 7½. Gill rakers 6 + 18 = 24. Vertebrae 30 + 15 = 45. Head length 24.4% SL; eye diameter 8.3% SL; body depth 17.3% SL. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Atlantic Flyingfish (English) and volador blanquito (Spanish). Diagnostic Characters. Cheilopogon melanurus differs from other species of Cheilopogon (except for C. heterurus and C. furcatus) in having minute jaw teeth, lower dorsal fin, and the structure of the chin barbels of juveniles. It differs from C. furcatus in pectoral fin coloration and length of chin barbels. Cheilopogon melanurus is very similar in appearance to C. heterurus (see differences in account for Cheilopogon heterurus above). Description. Dorsal fin rays 11–14, usually 12–14; anal fin rays 7–11, usually 8–10; pectoral rays I 13–17, usually I 14–16 (Table 5.14). Vertebrae 29–32 + 14–17 = 44–47 (Table 5.15). Predorsal scales 25–33 (up to

Exocoetus melanurus Valenciennes in Cuvier and Valenciennes, 1847:101–102 (original description; North Atlantic: “New York”; holotype MNHN A.7603). Bruun 1937:182 (redescription of type). Collette et al. 1997:12 (type). Exocoetus heterurus (not of Rafinesque 1810). Jordan and Meek 1885:59–61 (in part; North America). Jordan and Evermann 1896a:735–736 (in part; short description; western Atlantic). Bean 1903:331–332 (New York). Exocoetus furcatus (not of Mitchill 1815). Jordan and Meek 1885:61–63 (description; North America). Exocoetus robustus (not of Günther 1866). Jordan and Meek 1885:61 (description; Cuba). Exocoetus lutkeni Jordan and Evermann, 1896a:736 (original description; Cuba; holotype ANSP 7502). Cypsilurus lutkeni. Jordan and Evermann 1898:2836 (listed). Bruun 1935:56–58, pl. 4, fig. 1 (description, tropical Atlantic). Cypsilurus heterurus (not of Rafinesque 1810). Jordan and Evermann 1898:2836 (listed). Cypselurus lutkeni. Smith 1907:167 (description; North Carolina). Breder and Nichols 1930:6–7, fig. 4 (description; West Indies). Jordan et al. 1930:201 (listed). Nichols and Breder 1930:5–6 (in key). Breder and Nichols 1934:40–44. Duarte-Bello 1959:45 (listed; Cuba). Cypselurus heterurus (not of Rafinesque 1810). Nichols and Breder 1927:61 (New York). Jordan et al. 1930:20 (listed). Nichols and Breder 1930:5 (in key). Parr 1930:21–25, fig. 4 (description, northwestern Atlantic; in part: specimen 320 mm SL is Cheilopogon heterurus). Breder and Nichols 1934:40– 44. Breder 1938:56–69, figs. 29–31, 34 (description, in part: Caribbean, Gulf of Mexico, Gulf Stream). ?Bigelow and Schroeder 1953:172 (Gulf of Maine). Duarte-Bello 1959:45 (listed; Cuba). Staiger 1965:705–714, figs. 16–18 (in part; juveniles; western Atlantic). Mago Leccia 1970:88 (listed; Venezuela). Monteiro et al. 1998:399 (distribution; northeastern Brazil). Cypselurus luetkeni. Springer 1959:166 (comparison with Cheilopogon heterurus). Cervigón 1966:224–225 (Venezuela). Cheilopogon luetkeni. Parin 1961:174 (listed). Cypselurus melanurus. Gibbs and Staiger 1970:452–457, figs. 7a–c, 8 (in part; description, tropical Atlantic Ocean distribution).

176

Family Exocoetidae 36 in eastern Atlantic), usually 27–30; scales in transverse row from dorsal fin origin to lateral line 6–8½, usually 7–8 (Table 5.16). Gill rakers on the first arch 5–7 + 13–18 = 16–24 (Table 5.17). Measurements based on western Atlantic specimens (170–258 mm SL). Body elongate (greatest depth 4.9–6.2 times in SL); slightly compressed (width 1.2–1.4 times in depth), nearly rectangular in cross section, almost flat ventrally. Head length 3.9–4.6 times in SL and 1.1–1.2 times in dorso-caudal distance. Eye diameter 2.8–3.6 times in head length, 0.8–1.3 times in interorbital space. Lower jaw pointed, slightly projecting beyond upper when mouth is closed. Jaw teeth diminutive, conical, and blunt, rarely with one or two supplementary cusps; dental index 0.06–0.10. Palatine teeth absent. Dorsal fin low, its longest (second or third) ray 8.0–12.2 times in SL. Anal fin originates under fifth to seventh dorsal fin ray. Pectoral fins 1.4–1.6 times in SL, reaching from middle to end of caudal peduncle, its first ray unbranched, 1.8–2.0 times shorter than total fin length. Pelvic fins inserted nearer to posterior margin of opercle than to origin of lower caudal fin lobe (opercle to pelvic fin distance 1.1–1.4 times shorter than pelvic fin to lower caudal-lobe origin distance), very long (2.5–3.6 times in SL), reaching from middle to end of anal fin base. Color. Body dark above, pale below. Dorsal and anal fins with no pigment on membranes. Caudal fin uniformly dark. Pectoral fin gray, with indefinite pale triangular crossband reaching third to fifth ray, and narrow unpigmented margin along the rear edge of the fin. Pelvic fins unpigmented. Development. Body of juveniles (25–164 mm SL) somewhat more elongate than in adults, maximum depth 5.1–6.4 times in SL, slightly increasing with growth. Head 3.9–4.7 times in SL, its length not changing with growth. Eye diameter 2.0–3.1 times in head length and 0.8–1.3 in interorbital space. Dorsal fin higher than in adults, its longest ray 4.6–9.3 times in SL. Pectoral fins 1.5–1.8 times in SL. Pelvic fins 1.9–2.7 times in SL, gradually displaced posteriorly with growth. Paired chin barbels consisting of firm pointed stem with soft skin fold along posterior edge; in smaller specimens the fold is broad and semicircular, in larger juveniles it gradually tapers to a point (barbels lost at 80–130 mm SL); length of barbels 7–26 times, usually 8–20 times in SL. Pigmentation of larvae and juveniles changes significantly during development; for more details, see Parin and Belyanina (2000: figs. 4b, 6). At 8.3–12.5 mm SL, the body is evenly covered with stellate melanophores, the pectoral and pelvic fins

Figure 5.8. Distributions of Blotchwing Flyingfish, Cheilopogon heterurus, and Atlantic Flyingfish, Cheilopogon melanurus, based on Parin and Belyanina (2000) and Shakhovskoy and Malikova (2018). Map by W. E. Bemis.

are evenly dark in color, pigment is present in the anterior part of the dorsal fin, and the anal and caudal fins are lightly colored. At 17–22 mm SL, large melanophores form relatively distinct bands under the bases of pectoral fins, in between the pectoral-pelvic distance, above the bases of pelvic fins, in front of the anal fin, above its posterior part, and near the base of rays of the caudal fin. A light transverse spot appears on the pectoral fins in the lower proximal part and a second larger spot is present in the central part of the fin. The pectoral fins become mottled at 23–25 mm SL and the position of the dark spots aligned with the bands on the body. Pelvic fins dark with pale transverse spot proximally and light areas of membranes distally in juveniles above 40 mm SL. Paired fins become more lightly colored by 70–85 mm SL. Pigment usually present on distal regions of middle and posterior part of anal fin. Two dark spots present on both the lower and upper lobe of the caudal fin. Transverse pigment bands occur on the belly in juveniles up to 100–120 mm SL. These bands become less visible with increased growth, the anterior bands persisting the longest. In fish 147–155 mm SL, dorsal and anal fins without pigment, caudal fin dark, pelvic fins lightly colored, and pectoral fins gray with narrow light posterior border and oblique light stripe crossing the fin. 177

Fishes of the Western North Atlantic

Figure 5.9. Atlantic Flyingfish, Cheilopogon melanurus. A. USNM 198387, 244.5 mm SL. Dipnetted at night by R. H. Gibbs, Jr. at 20°50′N, 93°00′W from M/V Oregon, Sta. 1605, 25 November 1956. B. USNM 198376, 218 mm SL. Dipnetted at night by R. H. Gibbs, Jr. at 20°10′N, 92°25′W from M/V Oregon, 24 November 1956. C. USNM 198389, 197.5 mm SL. Dipnetted at night by R. H. Gibbs, Jr. at 35°50′N, 72°35′W from M/V Delaware, 19 July 1958. D. USNM 159038, 121.3 mm SL. Collected

178

Family Exocoetidae

off the Yucatán Peninsula from M/V Oregon, Sta. 1297. E. USNM 198386, 84.4 mm SL. Collected in the Gulf of Mexico from M/V Oregon, Sta. 1074. F. Pectoral and pelvic fins of USNM 294841, 123.8 mm SL. Collected at 34°45′N, 73°41′W from M/V Delaware, 10 October 1957. G. Pectoral and pelvic fins of USNM 197807, 38.3 mm SL. Collected by R. H. Gibbs, Jr. at 36°15′N, 72°28′W from M/V Delaware, 23 January 1959. Illustrations by H. E. Hamman.

179

Fishes of the Western North Atlantic Size. The maximum observed size for Cheilopogon melanurus is 265 mm SL (Parin and Belyanina 2000). According to Breder (1938) and Staiger (1965), maximum length is about 330 mm SL (USNM 10409, Brazil); however, we have studied this specimen and reidentified it as Cheilopogon cf. heterurus.

Mexico and the Caribbean Sea (Fig. 5.8). Found offshore only in the area between the Gulf Stream and Bermuda. Also known from the coast of Brazil from the equator to about 30°S. A separate population inhabits coastal Africa from Dakar, Senegal, to Luanda, Angola. One specimen of C. melanurus has recently been found in the north-central Atlantic (26°34′N, 33°55′W) and is probably an expatriate from the northwestern Atlantic population (Shakhovskoy and Malikova 2018). Geographic Variation. Slight differences were found in head length, eye diameter, and snout length between samples of Cheilopogon melanurus from the western North Atlantic (including the Gulf of Mexico and Caribbean Sea) and the western South Atlantic (off Brazil). Parin and Belyanina (2000) examined 15 specimens from the eastern Atlantic and found few differences; however, this may have been due to the small available sample size. Gibbs and Staiger (1970) showed differences in pectoral fin pigmentation and barbel structure between juvenile specimens (24–100 mm SL) of Cheilopogon melanurus from the eastern tropical Atlantic and similarly sized western Atlantic specimens. Although they did not find any distinctions between adults of these two populations, they suspected the presence of an undescribed species near the shores of West Africa. Parin and Belyanina (2000) reexamined three of these smaller (24–56 mm SL) eastern Atlantic specimens (USNM 203903, 203905, and 218912) and reidentified them as C. milleri. However, the western and eastern populations are somewhat different in predorsal scale number (25–33, x̅ 28.4, rather than 27–36, x̅ 31.7; see Table 5.16), and may represent two valid subspecies. Study Material. Description based on 92 western Atlantic specimens (23–258 mm SL; see Parin and Belyanina [2000] for full list and localities), as well as data of Bruun (1935) and Gibbs and Staiger (1970). Additional material includes meristics of 227 western Atlantic specimens examined by N. V. Parin.

Biology Food. Cheilopogon melanurus feeds on zooplankton (Parin 2003). Reproduction. Mature at about 200 mm SL (Bruun 1935). In the western Atlantic, females caught in June and August have ripe ovarian eggs up to 1.8– 1.9 mm in diameter (Gibbs and Staiger 1970). Eggs are demersal with filaments covering entire surfaces (Breder 1938). One ripe female (195 mm SL) and two ripe males (198 and 204 mm SL) were captured in late March at 22°05′N, 87°58′W. Sea surface temperature was 23 °C. Development of eggs took about 10–12 days. Unfertilized, hydrated eggs were 1.8–1.9 mm in diameter and had about 56 fine, very long (up to 21 mm) filaments (Tsokur 1975). The eggs of Cheilopogon melanurus are negatively buoyant and have long adhesive filaments (Casazza et al. 2005). Egg diameters ranged from 0.30 to 2.20 mm and eggs 1.40 mm and larger had long filaments present. The number of eggs increased with increasing size of females, from 3,837 eggs (178 mm SL female) to 28,202 eggs (257 mm SL). It is likely that C. melanurus is a batch spawner. Overall female gonadosomatic indices were highest in June and July, but on the basis of captured mature females, the spawning season off North Carolina lasts from at least June to August. The ratio of females to males did not differ from 1:1 (Casazza et al. 2005). Previous descriptions of spawning events suggested that flyingfishes spawn mainly in large schools at the surface, depositing eggs on floating substrates (Bane 1965; Parin and Lakshminaraina 1993; Stevens et al. 2003; Casazza et al. 2005). A single-pair spawning event off Cape Fear, North Carolina, showed unusual coloration and swimming behavior of Cheilopogon melanurus (Casazza et al. 2005). The female was stationary with pectoral fins fully extended, with three bright white bars. Small clumps of floating Sargassum were present in the spawning area, but the fish did not seem to be using it. Range. Cheilopogon melanurus is a neritic species restricted in the western North Atlantic along the coast from about 42°N (probably zone of expatriation) south to Trinidad, including the Gulf of

Cheilopogon pinnatibarbatus (Bennett, 1831) Bennett’s Flyingfish Figures 5.10, 5.11 Tables 5.1, 5.18, 5.19, 5.20, 5.21 Exocoetus pinnatibarbatus Bennett, 1831:146 (original description; Atlantic coast of northern Africa; no types known). Günther 1866:284 (description of juvenile; Atlantic coast of northern Africa). Exocoetus pulchellus Lowe, 1841:38 (original description; off Madeira; no types known).

180

Family Exocoetidae

Figure 5.10. Bennett’s Flyingfish, Cheilopogon pinnatibarbatus. A. IFAN 1966-89, 380.0 mm SL. B. IRSNB 9207, 162.5 mm SL. C. MMF, 25.0 mm SL. D. Same specimen, MMF, 25.0 mm SL, barbel in ventral view. Illustrations by H. E. Hamman, from Gibbs and Staiger (1970).

181

Fishes of the Western North Atlantic Bennett’s Flyingfish and Giant Flyingfish are more commonly used. Diagnostic Characters. Cheilopogon pinnatibarbatus differs from all other flyingfishes in the region because of its slender body (body depth 13.1%– 17.5% SL and depth of caudal peduncle 5.4%–5.9% SL); small head (head length 20.0%–22.7% SL); more vertebrae (49–52), and more predorsal scales (38– 47). Juveniles have a unique chin barbel: short and wide with fringed margin (Fig. 5.10). Adults of this species are often confused with C. heterurus because of slight overlap of characters and similar pigmentation. However, pelvic fin origin is more posterior in C. pinnatibarbatus. Description. Dorsal fin rays (10?) 11–14, usually 12–13; anal fin rays 9–12, usually 10–11; pectoral fin rays I 12–15 (Table 5.18). Vertebrae 33–35 + 16–18 = 49–52 (Table 5.19). Predorsal scales 38–47; scales in transverse row from dorsal fin origin to lateral line 6–9 (Table 5.20). Gill rakers on the first arch 5–8 + 14–18 = 19–26 (Table 5.21). Measurements based on specimens 200–400 mm SL. Body elongate (greatest depth 5.7–7.6 times in SL); nearly rectangular in cross section (width 1.1–1.4 times in depth), almost flat ventrally. Head 4.4–5.0 times in SL. Eye diameter 3.5–4.2 times in head length, 1.2–1.5 times in interorbital space. Jaws subequal. Jaw teeth conspicuous, most conical, some with additional cusps; dental index 0.11–0.17 (up to 0.22 in Indo-Pacific). Palatine teeth usually absent (very small and rare if present). Dorsal fin low to medium in height, its longest (usually second or third) ray 7.2–12.8 times in SL. Anal fin originates under third to seventh dorsal fin ray. Pectoral fins 1.3–1.5 times in SL, reaching end of dorsal to origin of caudal; first ray unbranched. Pelvic fins inserted closer to posterior margin of opercle than to origin of caudal fin lobe or midway (operculo-pelvic distance 1.0–1.1 times in pelvic to caudal distance), rather long (2.6–3.7 times in SL); reaching at least middle of anal fin base. Color. Body dark above, pale below. Dorsal and caudal fins gray (in juvenile specimens dorsal fin with black spot), anal fin transparent. Pectoral fins uniformly dark gray with pale outer margin, sometimes with a small pale triangle (juvenile specimens have a lighter crossband throughout the fin); pelvic fins gray basally, transparent elsewhere. Development. Juveniles have a single, wide chin barbel with a fringed margin (Fig. 5.10C, D); it disappears at about 140 mm SL. Dorsal, pectoral, and pelvic fins mottled with gray spots and bars. Gibbs and Staiger (1970) described three juveniles (25.0, 42.3, and 162.5 mm SL) from the eastern

Exocoetus lineatus Valenciennes in Cuvier and Valenciennes, 1847:92–94 (original description; lectotype MNHN A.7604). Günther 1866:287 (description; Madeira). Bruun 1937:181 (description of type). Collette et al. 1997:12 (types). Exocoetus altipennis Valenciennes in Cuvier and Valenciennes, 1847:109, pl. 560 (original description; Mers de l’Inde; lectotype MNHN A-9921). Bruun 1937:183–184 (description of type [as altipinnis]). Collette et al. 1997:10 (types). Exocoetus orbignianus Valenciennes in Cuvier and Valenciennes, 1847:131 (original description; Montevideo, Uruguay; no types known). Exocoetus longipinnis Castelnau, 1861:64 (original description; Agulhas Bank, South Africa; no types known). Exocoetus furcatus (not of Mitchill 1815). Roule and Angel 1930:85, pl. 4, fig. 110 (description of a juvenile; Azores). Cypsilurus lineatus. Bruun 1935:47–52, pl. 4, fig. 1 (description; Atlantic). Lozano Rey 1947:623–625, fig. 176 (after Breder, 1938) (description; Atlantic coast of Spain and Portugal). Poll 1953:187–189, fig. 78 (description; northeastern Atlantic). Cypselurus minos Nichols and Breder, 1930:1, fig. 2 (original description; Mindelo, St. Vincent, Cape Verde Islands; holotype Cleveland Museum [CMNH] 72). Cypselurus lineatus. Fowler 1936:429–431 (description; Madeira). Breder 1938:49–50, fig. 24 (description, eastern Atlantic). Cypsilurus heterurus (not of Rafinesque 1810). Poll 1953:190–191, fig. 80 (description; off northwestern Africa). Cheilopogon pinnatibarbatus. Parin 1961:171 (listed). Parin 1973:264 (Eastern Atlantic). Gorelova and Grudtsev 1987:481 (feeding; Atlantic). Andrew et al. 1995:14 (short description; Tristan da Cunha Island). Shakhovskoy 2007:259–266, figs. 1–6 (Atlantic; cranial osteology). Parin and Shakhovskoy 2016:2145 (description, eastern central Atlantic). Cypselurus pinnatibarbatus. Staiger, 1965:714–717 (description and distribution [compiled]; Atlantic). Wheeler 1969:234– 235 (brief description; northern Europe and British Isles). Gibbs and Staiger 1970:457–461, figs. 9–10 (description, Atlantic distribution).

Types. Exocoetus lineatus Valenciennes in Cuvier and Valenciennes, 1847. Lectotype MNHN A.7604 (366 mm SL), designated and described by Bruun (1937); Gorée, Senegal. Dorsal fin rays 13; anal fin rays 11; pectoral fin rays I 13. Predorsal scales about 40; transverse scale rows 7½. Gill rakers 6 + 16 = 22. Palatine teeth absent; first anal fin ray under sixth dorsal fin ray. Exocoetus altipennis Valenciennes in Cuvier and Valenciennes, 1847. Lectotype MNHN A-9921 (284 mm SL), designated and described by Bruun (1937); Mers de l’Inde. Dorsal fin rays 13; anal fin rays 12; pectoral fin rays I 14–15. Predorsal scales about 42; transverse scale rows 7½. Gill rakers 6 + 14 = 20. Palatine teeth present; first anal fin ray under fifth dorsal fin ray; dorsal fin with remains of a large black spot. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Smallhead Flyingfish (English) and volador cabecita (Spanish). However, 182

Family Exocoetidae Table 5.18. Fin ray counts for Cheilopogon pinnatibarbatus, based on specimens examined by N. V. Parin and I. B. Shakhovskoy, and literature data from Bruun (1935) and Breder (1938). Dashes indicate a value of zero. Fin rays Dorsal

Anal

Pectoral

Locality

10

11

12

13

14

9

10

11

12

14

15

16

Northwestern Atlantic (?) Southwestern Atlantic Northeastern Atlantic Southeastern Atlantic Species total

1 — — — 1

1 1 1 — 3

— 3 23 2 28

— 6 38 2 46

— — 4 — 4

— — 2 — 2

— 7 21 — 28

2 3 36 2 43

— — 6 2 8

— — 19 2 21

— 4 26 2 32

— — 2 — 2

Table 5.19. Vertebral counts of Cheilopogon pinnatibarbatus, based on specimens examined by N. V. Parin and I. B. Shakhovskoy, as well as literature data from Bruun (1935) and Breder (1938). Dashes indicate a value of zero. Vertebrae Precaudal

Caudal

Total

Locality

33

34

35

16

17

18

50

51

52

Northwestern Atlantic (?) Southwestern Atlantic Northeastern Atlantic Southeastern Atlantic Species total

1 1 3 — 5

1 — 2 — 3

— — — 1 1

— — 1 — 1

2 1 3 1 7

— — 1 — 1

1 1 4 2 8

1 — 3 1 5

— — 1 1 2

Table 5.20. Number of predorsal and transverse scales of Cheilopogon pinnatibarbatus, based on specimens examined by N. V. Parin and I. B. Shakhovskoy, as well as literature data from Bruun (1935) and Breder (1938). Dashes indicate a value of zero. Scales Predorsal

Transverse

Locality

38

39

40

41

42

43

44

45

46

47

6½

7

7½

8

8½

9

Southwestern Atlantic Northeastern Atlantic Southeastern Atlantic Species total

1 3 — 4

3 2 — 5

4 10 — 14

2 8 — 10

— 14 1 15

— 12 — 12

— 8 1 9

— 3 1 4

— 3 — 3

— 1 — 1

— 1 — 1

1 1 1 3

2 12 3 17

— 5 — 5

— 6 — 6

— 2 — 2

Table 5.21. Number of gill rakers on the first arch for Cheilopogon pinnatibarbatus, based on specimens examined by N. V. Parin and I. B. Shakhovskoy, as well as literature data from Bruun (1935) and Breder (1938). Dashes indicate a value of zero. First arch gill rakers Upper Locality Southwestern Atlantic Northeastern Atlantic Southeastern Atlantic Species total

Lower

Total

5

6

7

8

14

15

16

17

18

19

20

21

22

23

24

25

— 2 — 2

— 16 1 17

1 18 1 20

— 6 1 7

— 6 1 7

1 12 1 14

— 15 — 15

— 7 1 8

— 2 — 2

— 1 — 1

— 5 1 6

— 5 — 5

1 12 1 14

— 8 — 8

— 8 — 8

— 3 1 4

183

Fishes of the Western North Atlantic California, small crustaceans and larval fishes are dominant prey items (Fitch and Lavenberg 1971). Reproduction. Cheilopogon pinnatibarbatus matures at about 320–350 mm SL (Bruun 1935; Breder 1938). Near the Cape Verde Islands, a mature male (343 mm SL) was captured in late December (Breder 1938). Two females (380–394 mm SL) from off Senegal captured in September and December had eggs measuring up to 1.8 mm in diameter, with filaments over the entire egg surface (Gibbs and Staiger 1970). California specimens had eggs 1.57– 1.70 mm in diameter with approximate 60 very long filaments uniformly distributed over the entire egg surface (Hubbs and Kampa 1946). Japanese specimens contained eggs 1.95–2.10 mm in diameter (Imai 1959). Fecundity in Japanese specimens was estimated to be 16,000 to 26,000 eggs (Imai 1959). Spawning of Cheilopogon pinnatibarbatus off California takes place at the surface of the ocean from late June into September, and the eggs usually adhere to floating debris (Jordan and Meek 1885; Fitch and Lavenberg 1971). Spawning occurs in Japanese waters (Hachijo Island) from February to May, with surface temperature at the beginning of spawning at about 19 °C (Abe 1954). Spawning occurs near Kyushu in December and January (Abe 1960). Paired spawning was reported off California (Miller 1952). Hatching occurred 14–16 days after fertilization (Miller 1952; Imai 1959). Larvae were captured south to Boso in March–April, when temperatures were 17.5–22.5 °C (Chen 1987). The largest individuals in waters off California seem to be five years old, according to the growth zones noted on otoliths (Fitch and Lavenberg 1971); however, such a long life span seems to be an overestimation. Range. Cheilopogon pinnatibarbatus occurs in subtropical parts of all oceans, mainly in neritic waters (distribution of this species in the Atlantic Ocean is shown on Fig. 5.11). In the eastern Atlantic it mainly occurs at 16.5–23 °C (Bruun 1935) or, in the Pacific, at 15–25 °C (Parin 1959). In the eastern Atlantic, C. pinnatibarbatus has been reported from England (but this report needs confirmation) and Spain south to Edina, Liberia, as well as around St. Helena, Ascension Island, and the Cape of Good Hope (Bruun 1935; Wheeler 1969; Gibbs and Staiger 1970; Parin 1973). In the southwestern Atlantic it is known from Rio de Janeiro (Gibbs and Staiger 1970) and Cape Frio, Brazil (Bruun 1935), south to Montevideo, Uruguay (as Exocoetus orbignianus Valenciennes in Cuvier and Valenciennes, 1847). The southernmost occurrence is from 39°54′S, 53°32′W (IORAS, no number, SRTM 8018, 22.03.1977, 225 and 320 mm

Atlantic. The dorsal fin height is about 4.0 times in SL in the 25.0 mm SL specimen, about 6.5 times in the 162.5 mm SL specimen. In the 25.0 mm SL specimen the dorsal fin is heavily speckled throughout. At somewhat larger sizes it presumably becomes almost entirely gray or black. Still later (probably at more than 200 mm SL) black pigment becomes restricted to a spot in the distal part of the middle rays. The pectoral fins of the 25.0 and 42.3 mm SL specimens have two prominent spots, one in the middle of the proximal part of the fin and the other near its postero-mesial part. In larger specimens, the pectoral spots presumably expand across the width of the fin and approach one another at the midpoint of the fin, leaving a narrow, pale stripe between them. In the 162.5 mm SL specimen, the distal part of pectoral fin is considerably darker than the proximal part. The only known juvenile specimen of Cheilopogon pinnatibarbatus from the western Atlantic was from off Uruguay and described as Exocoetus orbignianus by Valenciennes in Cuvier and Valenciennes (1847) based on a drawing by Alcide d’Orbigny. This specimen was about 127 mm long and had a short and wide barbel, but the divisions of its margin into filaments were not drawn. Body elongate, head short. Pectoral fins rounded and short, not reaching first anal fin ray. Pelvic fins inserted in the middle of belly, long, reaching caudal fin. Dorsal fin pointed, triangular, its height more than twice the body depth. Belly with three dark spots, two of which in front of pelvic fins. Pelvic fins pale blue above, white below, with blue rays. Rays of dorsal, anal and caudal fins also blue. Larvae and juveniles of Cheilopogon pinnatibarbatus have been described from three Indo-Pacific regions: Japan (Abe 1954; Imai 1959; Chen 1987); California (Hubbs and Kampa 1946); and Australia (Belyanina 1993). Size. The maximum observed size of Cheilopogon pinnatibarbatus is about 400 mm SL (Parin and Shakhovskoy 2016). Biology Predators. At Tristan Island Cheilopogon pinnatibarbatus is one of the major prey items for the Great Hammerhead, Sphyrna mokarran, and the Snoek, Thyrsites atun (Andrew et al. 1995). Food. In the eastern Atlantic, Cheilopogon pinnatibarbatus feeds mainly on copepods (Poll 1953). Atlantic specimens (303–330 mm SL) mainly consumed siphonophores and rarely amphipods (Gorelova and Grudtsev 1987). In waters off 184

Family Exocoetidae near the Gulf of Guinea: ZMH 8838 (236 mm SL), Westküste Afrikas (Gold küste), Leg. Merten, October 1950; and ZMH 12764 (ca. 270 mm SL), Gabon, H. Petersen, 1885). Geographic Variation. Several different subspecies have been recognized (Parin 1973): Cheilopogon pinnatibarbatus pinnatibarbatus (eastern North Atlantic); C. p. altipennis (southern Indian Ocean); C. p. melanocercus (southern Australia and New Zealand); C. p. japonicus (western North Pacific); C. p. californicus (eastern North Pacific); and C. p. fernandezianus (eastern South Pacific). The taxonomic status of the western South Atlantic population is uncertain. Study Material. The description is based mainly on Bruun (1935), Breder (1938), Gibbs and Staiger (1970), and Parin and Shakhovskoy (2016). Additional material includes meristics of 63 specimens examined by N. V. Parin and I. B. Shakhovskoy. From the northeastern Atlantic, 49 specimens (171–388 mm SL) from five collections. MCZ 55487 (1, 347 mm SL); 24°05′N, 18°34′W; 15 October 1970. MCZ 42996 (1, 353 mm SL); 21°09′N, 16°04′W; 1 September 1959. ZMH 8841 (1, 343 mm SL); Leg. Kpt. Dahlstrom, 08°52′N, 18°49′W; 26 April 1963. ZMH 8838 (1, 236 mm SL); Westküste Afrikas (Gold küste); Leg. Merten, October 1950. ZMH 8840 (1, 365 mm SL); 24°30′N, 17°20′W; 1 March 1911. ZMH 12745 (1, 345 mm SL); Edina, West Africa; 1851. ZMH 12737 (1, 362 mm SL); St. Vincent, Cape Verde Islands; 29 December 1900. ZMH (ISH 360/53) (1, 279 mm SL); Olympic Victor, 18°40′N, 17°40′W; 21 April 1953. ZMH (ISH 20/54) (1, 318 mm SL); Nördl. Kanar. Inseln. ZMH (ISH 1534/64) (1, 388 mm SL); La Rafale, GTS II; 1964. MNHN A.7604 (1, 362.2 mm SL); Gorée; holotype of Exocoetus lineatus. AtlantNIRO uncat. (3, 262–360 mm SL); SRTM 8016, 19°56′N, 18°05′W; 25 December 1977. AtlantNIRO uncat. (1, 337 mm SL); SRTM 8016, 21°34′N, 17°56′W; 7 April 1978. AtlantNIRO uncat. (1, 322 mm SL); SRTM 8016, 18°01′N, 19°02′W; 6 April 1978. AtlantNIRO uncat. (1, 369 mm SL); SRTM 1246, 21°02′N, 17°43′W; 29 September 1979. AtlantNIRO uncat. (1, 370 mm SL); SRTM 8060, 23°30′N, 17°09′W; 26 October 1976. AtlantNIRO uncat. (1, 340 mm SL); SRTM 8060, 20°19′N 17°27′W; 28 July 1978. AtlantNIRO uncat. (5, 230–375 mm SL); SRTM 8018, 20°47′N, 17°54′W; 18 January 1977. INBYuM uncat. (9, 171–336 mm SL); R/V Vodyanitsky Cruise 2, 19°35′N, 18°06′W; 5 June 1977. INBYuM uncat. (8, 241–326 mm SL); R/V Vodyanitsky Cruise 3 Sta. 296, 20°36′N, 19°44′W; 1 November 1977. INBYuM uncat. (1, 213 mm SL); R/V Vodyanitsky Cruise 3 Sta. 299, 23°20′N, 21°00′W; 3 November 1977. INBYuM uncat. (1, 192 mm SL); R/V Vodyanitsky Cruise 3;

Figure 5.11. Distribution of Bennett ’s Flyingf ish, Cheilopogon pinnatibarbatus, based on specimens identified by N. V. Parin and I. B. Shakhovskoy, as well as data from the scientific literature. Map by W. E. Bemis.

SL). Andrew et al. (1995) described C. pinnatibarbatus from Tristan da Cunha. The only western North Atlantic record, from off Bermuda (Goode 1876; Beebe and Tee-Van 1933), was regarded as probably erroneous (see Breder 1938; Gibbs and Staiger 1970). However, Parin (unpublished data) found two specimens of C. pinnatibarbatus from the northwest Atlantic (ISH 11/52 [ZMH 100173], ca. 280 and ca. 290 mm SL; 19°32′N, 84°04′W; collected by G. Krefft, 20 March 1952) in the collection of the Zoological Museum (Hamburg). Also found in the same sample were C. cyanopterus (ca. 295 mm SL); C. nigricans or C. exsiliens (SL 230 mm), Hirundichthys speculiger (220 and 230 mm SL) and H. (rondeletii) sp. (215 mm SL). These two C. pinnatibarbatus specimens differ from other Atlantic conspecifics in fewer dorsal fin rays (10 or 11; see Table 5.18) and are similar in this respect to specimens of C. pinnatibarbatus californicus (see Hubbs and Kampa 1946). This suggests that these fishes might have been captured in the eastern Pacific and mislabeled as from the Caribbean Sea. However, species composition of this sample counts in favor of Atlantic instead of eastern Pacific. Thus, occurrence of C. pinnatibarbatus in the northwestern Atlantic requires confirmation. We also regard as probable mislabeling two reported captures of Cheilopogon pinnatibarbatus 185

Fishes of the Western North Atlantic 30 October 1977. INBYuM uncat. (1, 325 mm SL); R/V Vodyanitsky Cruise 1 Sta. 29; 7 January 1977. INBYuM uncat. (1, 357 mm SL); R/V Vodyanitsky Cruise 1. INBYuM uncat. (3, 238–356 mm SL); R/V Vodyanitsky Cruise 1 Sta. 27, 21°46′N, 18°10′W; 5 January 1977. IORAS uncat. (1, 207 mm SL); SRTM 8003, 20°50′N, 18°29′W; 21 August 1974. From the northwestern Atlantic, two specimens from one collection. ZMH 100173 (ISH 11/52) (2, ca. 280–290 mm SL); 19°32′N, 84°04′W, 20 March 1952. From the southeastern Atlantic, one specimen. MNHN A9920 (1, 269.4 mm SL); Cap de Bonne Esperance, Verreaux; paralectotype of Exocoetus altipennis. From the southwestern Atlantic, 12 specimens (189–338) from three collections. ZMH 12739 (1, 338 mm SL); Cap Frio, Brazil; 12 June 1901. IORAS uncat. (3, 189–261 mm SL); SRTM 8018, 39°54′S, 53°32′W; 21 March 1977. IORAS, uncat. (1, 311 mm SL); SRTM 8016, 37°48′S, 47°54′W; 25 January 1978. AtlantNIRO uncat. (2, 269–289 mm SL); SRTM 8016, 40°04′–39°10′S, 49°57′–49°19′W; 4 March 1978. AtlantNIRO uncat. (2, 225–320) SRTM 8018, 39°58′S, 53°12′W; 22 March 1977. AtlantNIRO uncat. (1, SL-?); SRTM 8018, 38°02′S, 50°29′W; 3 March 1977. AtlantNIRO (2, 225–320 mm SL); SRTM 8018, 39°58′S, 53°12′W; 22 March 1977.

transverse scales 7–10½. Pectoral branch of lateral line absent. Swim bladder extends into haemal canal, reaching middle of caudal peduncle or farther posterior. Head 3.8–4.8 times in SL. Snout blunt, shorter than eye. Lower jaw usually a little shorter than the upper and included beneath the upper jaw with mouth closed; upper jaw not protrusible. Jaw teeth medium-sized (dental index 0.09–0.22), conical, with one to two additional cusps or (most often) tricuspid (unicuspid in some species). Gill rakers on first arch 17–31. Dorsal fin rays 10–14, anal fin rays 7–11, dorsal rays minus anal rays 1–5, pectoral fin rays I 12–16; pectoral fins long (1.4–1.6 times in SL), usually extending to end of dorsal fin base or farther posteriorly. Pelvic fins long (2.9–3.8 times in SL), with third ray the longest, inserted before to behind the middle of distance between posterior margin of opercle and caudal fin origin. Dorsal fin uniformly gray, sometimes with small dark spots, low or of medium height, anterior rays the longest. Anal fin originates four to seven rays behind origin of dorsal fin. Eggs demersal (probably pelagic in a few species), uniformly covered with short or long filaments. Juveniles either with a single chin barbel or without barbels. Species. At least 11 species, all but one distributed in the Pacific and Indian oceans. The only Atlantic species, Cypselurus comatus, belongs to the subgenus Cypselurus, together with C. angusticeps, C. hiraii, C. naresii, and C. opisthopus.

Genus Cypselurus Swainson, 1838 Cypselurus Swainson 1838:299 (type species Exocoetus appendiculatus Wood = Cypselurus comatus (Mitchill) by original designation).

Cypselurus comatus (Mitchill, 1815) Clearwing Flyingfish Figures 5.12, 5.13 Tables 5.1, 5.22, 5.23, 5.24, 5.25

Diagnosis. Cypselurus, like other genera of the Cypselurinae, has long pectoral fins reaching the end of the dorsal fin base and elongate pelvic fins with the third ray the longest and inserted nearer to the anal fin origin than to the pectoral fin insertion. Cypselurus differs from Prognichthys in having only the first pectoral fin ray unbranched, from Cheilopogon in having the lower jaw a little shorter than the upper and included beneath the upper jaw, and from Hirundichthys in having the anal fin origin three rays or more behind origin of dorsal fin. Pectoral branch of the lateral line absent. Jaw teeth usually tricuspid, mouth not protrusible. Juveniles with one or no chin barbel (Parin 1961; Shakhovskoy and Collette, in press). Description. The body of Cypselurus is nearly rectangular in cross section in adults, almost flat ventrally. Greatest body depth 4.3–6.0 times in SL, width of body 1.2–1.4 times in its depth. Vertebrae 25–32 + 12–16 = 39–48. Predorsal scales 23–38;

Exocoetus comatus Mitchill, 1815:448, pl. 5, fig. 1 (original description; New York). DeKay 1842:231 (short description; New York). Storer 1846:440 (New York). Valenciennes in Cuvier and Valenciennes 1847:133–135 (South Carolina). Günther 1866:286 (western South Atlantic). Lütken 1876:399, pl. 10, fig. 1. Exocoetus appendiculatus Wood, 1825:283, pl. 17, fig. 2 (original description, figure, locality unknown). ?Exocoetus bahiensis Ranzani, 1842:362–365 (original description; Bahia, Brazil; syntypes MZUB 950, MZUB 981). Considered as a probable synonym of Cheilopogon cyanopterus by I. B. Shakhovskoy. Exocoetus heterurus (not of Rafinesque 1810). Jordan and Meek 1885:59–61 (in part). Jordan and Evermann 1896a:735–736 (in part; western Atlantic). Cypselurus vitropinna Breder, 1927:20–21, fig. 11 (original description, illustration; West Indies; holotype YPM ICH 000459). Parr 1930:25, 27 (Bahamas). Breder and Nichols 1930:7–9 (description of jaw teeth). Nichols and Breder 1930:5, 7; Breder and Nichols 1934:40, 42, 44.

186

Family Exocoetidae

Figure 5.12. Clearwing Flyingfish, Cypselurus comatus. A. No. 418, 193 mm SL. From Bruun (1935), illustration by P. H. Winther. B. USNM 251937-F53, 71.1 mm SL. Collected at 25°20′N, 80°00′W, 30 August 1962. Illustration by H. E. Hamman. Cypselurus antarei Beebe and Hollister, 1933:83–85, fig. 15 (original description; 21°50′N, 63°32′W; holotype USNM 170921). Collette et al. 1992:6 (holotype). Cypsilurus comatus. Bruun 1933:382 (not synonymous with Cheilopogon heterurus). Bruun 1935:52–56, text fig. 10, pl. 2, fig. 3 (description, illustrations; western Atlantic). Cypselurus comatus. Jordan and Gilbert 1883:381 (description; Atlantic). Breder and Nichols 1934:40, 42 (discussion). Breder 1938:51–55, fig. 25 (description, distribution). DuarteBello 1959:45 (listed; Cuba). Parin 1961:165 (listed). Staiger 1965:679–685, figs. 1–2 (description, western Atlantic;

in part: specimen from the Gulf of Mexico is Cheilopogon melanurus). Gibbs and Staiger 1970:461 (distribution in Atlantic). Gibbs 1978: EXOC Cyp.1 (description of juveniles, figure, range). Kovalevskaya 1982:108–110, fig. 2 (larvae and juveniles). Bruce and Bowman 1989:16 (Cymothoidae parasites). Monteiro et al. 1998:399 (distribution; northeastern Brazil). Parin 2003:1126, figure, map (description, western Atlantic). Quattrini et al. 2004:161–162, pl. 1, fig. e (juvenile; North Carolina, 34°17′N, 75°47′W). Cotten and Comyns 2006:978–979, figs. A–E (early life history). Fahay 2007:808– 809, figs. A–D (early life history stages). Casazza and Ross

187

Fishes of the Western North Atlantic

(Page et al. 2013) lists Clearwing Flyingfish (English). Diagnostic Characters. Cypselurus comatus is most similar to the Indo-West Pacific Cypselurus naresii, from which it differs in having fewer vertebrae (40–43 rather than 42–45), fewer predorsal scales (23–30, usually 25–28, rather than 27–33), more dorsal fin rays (11–13, usually 12–13, rather than 10–12) and in lighter and less extended pigmented area of pectoral fins (usually to seventh ray rather than to ninth ray). Besides generic distinctions, Cypselurus comatus differs from all western Atlantic species of Cheilopogon (except for Cheilopogon cyanopterus and Cheilopogon pinnatibarbatus) in the more posterior pelvic insertion (operculo-pelvic distance 0.9–1.1 times in pelvic-caudal distance) and in the pigment pattern of pectoral fins, which are entirely darkly pigmented in Cheilopogon cyanopterus and with a pale transverse strip or basal triangle in others. Furthermore, a prominent black spot on the dorsal fin is present in Cheilopogon cyanopterus and C. exsiliens, and palatine teeth are absent in C. furcatus, C. heterurus, and C. melanurus. Juvenile Cypselurus comatus also differ from all other Atlantic flyingfishes in having a single long chin barbel. Description. Dorsal fin rays 11–13, usually 12–13; anal fin rays 7–9; pectoral fin rays I 11–15 (Table 5.22). Vertebrae 25–28 + 13–17 = 40–43 (Table 5.23). Predorsal scales 23–30, usually 25–28; scales in

2008:352, 355 (associated with Sargassum, Gulf Stream off North Carolina). Cypselurus alienus Herre, 1935:392–393 (original description; south coast of Cuba; holotype FMNH 17224). Herre 1936:11, 13, fig. 1 (listed).

Types. No type specimens are extant for these three species: Exocoetus comatus Mitchill, 1815, E. appendiculatus Wood, 1825, and E. bahiensis Ranzani, 1842. Cypselurus vitropinna Breder, 1927. Holotype YPM ICH 000459 (177 mm SL); West Indies. Dorsal fin rays 12; anal fin rays 9. Pectoral fin rays I 12. Predorsal scales 26; scales between dorsal fin origin and lateral line 8½. Gill rakers 5 + 17 = 22. Head length equals dorso-caudal distance; pectoral fin lightly pigmented to seventh ray. Paratypes YPM ICH 000167 (5, 176–210 mm SL). Cypselurus antarei Beebe and Hollister, 1933. Holotype USNM 170921 (67 mm SL); 21°50′N, 63°32′W. Dorsal fin rays 13; anal fin rays 9; pectoral fin rays I 14/I 13. Predorsal scales 24; transverse scales between dorsal fin origin and lateral line 7½. Gill rakers 7 + 16 = 23. Vertebrae 28 + 15 = 43. Head 1.1 times in dorso-caudal distance, operculo-pelvic distance equal to pelvic-caudal distance; chin barbel equal to standard length (Collette et al. 1992:6). Cypselurus alienus Herre, 1935. FMNH 17224 (not seen). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico 188

Family Exocoetidae

transverse row from dorsal fin origin to lateral line 7–8½, usually 7½ (Table 5.24). Gill rakers on the first arch 4–7 + 15–20 = 19–25, usually 22–23 (Table 5.25). Measurements based on specimens 158–210 mm SL. Body elongate (greatest depth 5.0–6.0 times in SL); moderately compressed (width 1.3–1.4 times in depth). Head moderate (3.8–4.2 times in SL); approximately equal to dorso-caudal distance (0.9– 1.1 times in the latter). Eye diameter 2.8–3.5 times in head length, equal to interorbital space (0.9–1.1 times in the latter). Jaw teeth rather strong, often with one or two additional cusps, sometimes tricuspid, dental index 0.13–0.16. Palatine teeth usually present. Dorsal fin low, its longest ray (second or third) 8.9–11.1 times in SL. Anal fin originates under fourth to sixth dorsal fin ray. Pectoral fin 1.5–1.6 times in SL, reaching from base of penultimate dorsal fin ray to middle of caudal peduncle, first ray unbranched. Pelvic fins inserted near middle of distance between posterior margin of opercle and origin of lower caudal lobe (operculo-pelvic distance 0.9–1.1 times in pelvic-caudal distance), long (3.0–3.7 times in SL); reaching from base of penultimate anal fin ray to behind end of anal fin base. Color. Body dark iridescent blue above, pale silvery below in life. Dorsal and caudal fins gray; anal fin transparent; pectoral fins uniformly light gray to

gray to seventh (rarely eighth) ray, their lower part and narrow hind margin transparent; pelvic fins gray near the base, transparent elsewhere, sometimes a black spot of different sizes is present on one or both fins. Development. Many proportions of juvenile Cypselurus comatus (25–98 mm SL) differ significantly from those of adult specimens: head and eye are larger, pelvic fins inserted more anteriorly (operculopelvic distance is much shorter than pelvic-caudal distance at 20–30 mm SL); dorsal fin higher, pectoral fins shorter, and pelvic fins longer (Breder 1938; Staiger 1965; Kovalevskaya 1982). Juveniles have a single median barbel with a strong fleshy axis and two lateral folds extending for its entire length and tapering to a point distally. Relative length of barbel varies from 1.6–4.0 times in SL at 20–50 mm SL to 0.9–2.3 times at 50–80 mm SL and, again, to 1.5–3.8 times at 80–160 mm SL. The largest known barbeled specimen is 161 mm SL (Bruun 1935: fig. 10) and the smallest specimen without barbels is 151 mm SL (USNM 197842). In smaller juveniles (below 45 mm SL), the body is uniformly blackish and pelvic fins heavily pigmented, darkest posteriorly; at 70–100 mm SL a dark spot is retained only at the central part of fins. Medial axis of barbel light, lateral folds blackish during entire development. 189

Fishes of the Western North Atlantic our opinion belongs to Cheilopogon melanurus (see Fig. 5.9A). The northernmost published record is off North Carolina (Quattrini et al. 2004). It is also recorded off Brazil between 01°S and 20°S (Ranzani 1842; Bruun 1935; Kovalevskaya 1982). According to our data, the southernmost occurrence is 22°51′S, 35°08′W (IORAS, no number, SRTM 8016, 62 mm SL, 7 January 1978). Sauskan (1973) reported C. comatus as the most abundant flyingfish in shelf waters of the eastern tropical Atlantic, but gave no characters and undoubtedly misidentified the flyingfish as C. comatus. Study Material. Description based on Bruun (1935), Breder (1938), Staiger (1965), and Parin (2003). Additional material includes meristics of 64 specimens examined by N. V. Parin, R. H. Gibbs, Jr., J. C. Staiger, and I. B. Shakhovskoy. From the northwestern Atlantic, 55 specimens (27.4–204.4 mm SL) from seven collections. UMML uncat. (3, 182–204.4 mm SL); Sea Hunter, Cab Cay, West End; 21 May–3 June 1962. UMML uncat. (4, 169.5–179.7 mm SL); D. deS. 351a, off Bimini; 6–7 July 1961. UMML uncat. (1, 96.2 mm SL); Jara Sta. 90, 22°35′N, 78°08′W; 29 November 1951. UMML G-50 (2, 31.1–46.5 mm SL); 25°38′N, 79°25′W. UMML G-70 (2, 34.7–55.5 mm SL); 25°23′N, 79°41′W. UMML uncat. (1, 60.1 mm SL); Cruise 7 Reg 28, 30°21′N, 79°26′W; 27 June 1954. UMML uncat. (1, 180.8 mm SL); Gill 7 Reg 7, 27°40′N, 79°18′W; 23 June 1954. UMML uncat. (1, 43.8 mm SL); Gill 2 Reg 8, 28°19′N, 79°26′W; 24 April 1953. UMML uncat. (1, 29.4 mm SL); Gill 5 Std Sta., 26°25′N, 76°40′W; 24 January 1954. UMML uncat. (1, 38.1 mm SL); Gill 2 Sta. 1, 27°00′N, 79°18′W; 22 April 1953. UMML uncat. (1, 60.1 mm SL); Silver Bay 3501, 20°43′N, 73°33′W; 22 December 1961. UMML uncat. (1, 27.4 mm SL); Cruise 8. Reg. 7, 27°40′N, 79°18′W; 11 September 1954. UMML uncat. (1, 40.4 mm SL); Gill 7 Std Sta., 26°25′N, 76°40′W; 13–14 June 1954. UMML 13816 (3, adults); Bahamas. UMML 16364 (6, adults); Bahamas. USNM 170321 (3, 181.5–197 mm SL); Sta. 168-56, 17°27′N, 63°13′W; 13 April 1956. USNM 251937-F52 (1, 82.4 mm SL); Atlantis 229, 20°10′N, 60°05′W; 10 December 1956. USNM 197842 (1, 151 mm SL); M/V Irmay, between Grand Cayman Island and Swam Island; 15 January 1950. USNM 200802 (1, juvenile); Florida Straits. USNM (?) uncat (1, 172 mm SL); TABL 2237, Geronimo Cruise 7 Sta. 23, 13°35′N, 59°54′W; 6 February 1966. USNM 251937F53 (1, 71.1 mm SL); G-64, 25°20′N, 80°00′W; 30 August 1962. USNM 268483 (1, 81 mm SL); 20°10′N, 60°05′W; 10 December 1956. USNM 294693 (7, 149–181 mm SL); 20°40′N, 69°56′W. USNM 170921 (1, 67 mm SL); north of Sombrero Island; type of

Figure 5.13. Distribution of Clearwing Flyingfish, Cypselurus comatus, based on specimens identified by N. V. Parin and I. B. Shakhovskoy, as well as data from the scientific literature. Map by W. E. Bemis.

Size. The maximum observed size for Cypselurus comatus is 212 mm SL. Biology Parasites. Glossobius auritus (Cymothoidae) reported from Cypselurus comatus in the Bahamas three miles west of Bimini, Saba Bank in the Lesser Antilles, southwest of Barbados and off Honduras (Bruce and Bowman 1989). Food. Cypselurus comatus feeds on zooplankton (Parin 2003). Reproduction. The smallest ripe female is 178 mm SL (Bruun 1935). The breeding season in the West Indies extends from January to April; ovarian eggs (1.2–1.45 mm) have filaments over the entire surface (Bruun 1935; Breder 1938). Eggs demersal. Range. Cypselurus comatus is a neritic species found usually within 300 miles of land. It is endemic to the western tropical Atlantic in two discrete areas (Fig. 5.13). North of the equator, C. comatus is rather abundant off Florida, the Bahamas, and the Antilles, and less common in the Caribbean. We reexamined the only reported (Staiger 1965) Gulf of Mexico specimen of C. comatus (USNM 198387, SL 242 mm, 20°50′N, 93°00′W). This specimen has 14 dorsal fin rays and 47 (30 + 17) vertebrae, and in 190

Family Exocoetidae in SL), extending to caudal fin base. Pelvic fins short (6.5–8.6 times in SL), inserted nearer snout than caudal fin base and nearer pectoral fin insertion than anal fin origin. Dorsal fin uniformly gray, low, with anterior rays the longest. Anal fin originates approximately at the same vertical line as dorsal fin origin. Osteological characters for Exocoetus published by Parin (1961) and Parin and Shakhovskoy (2000). Eggs pelagic, without filaments. Juveniles with a single chin barbel (Exocoetus monocirrhus) or without barbels. Species. Five oceanic species of Exocoetus are recognized (Parin and Shakhovskoy 2000; Lewallen et al. 2017): circumtropical E. volitans, Indo-Pacific E. monocirrhus, Atlantic E. obtusirostris, E. gibbosus from the southern subtropical Pacific, and E. peruvianus from open ocean off Peru and Ecuador in the eastern Pacific. Molecular analysis indicates that E. volitans is sister to the other four species and that the Atlantic E. obtusirostris is sister to the IndoPacific E. monocirrhus (Lewallen et al. 2011, 2017; Gordeeva and Shakhovskoy 2017). The two allopatric South Pacific species, E. gibbosus and E. peruvianus, lack genetic distinctness in Cyt b and Rag 2 genes (Lewallen et al. 2017). However, the specimens of E. gibbosus used in Lewallen et al. (2017) were subsequently identified by I. B. Shakhovskoy as E. peruvianus based on photographs kindly provided by R. Winterbottom and E. Holm (ROM). Range. The genus Exocoetus is widely distributed in warm marine waters, especially offshore. It dominates the mid-size nekton of the tropical epipelagic zone outside the neritic zone (Parin and Shakhovskoy 2000). In open waters, Exocoetus is the most abundant flyingfish and has been reported to be as high as 80% to 90% of the total catch (Bruun 1935; Nesterov and Grudtsev 1980). Two species (Exocoetus volitans and E. obtusirostris) occur in the western Atlantic. One specimen of E. monocirrhus was reported off Brazil (paratype of E. georgianus MNHN B.841), but it was probably mislabeled (see Parin and Shakhovskoy 2000).

Cypselurus antarei; 30 July 1932. USNM 53085 (1, 180 mm SL); near Cat Island, Bahamas. MCZ 41190 (2, 181–186 mm SL); Bahamas; June 1945. CAS 124419 (1, 165 mm SL); Bahama Islands, Caicos Passage; paratype of Cypselurus alienus; 27 November 1928. ANSP 150180 (1, 172 mm SL); Bahamas, Crooked Island passage; 26 May 1962. ANSP 150179 (1, 197 mm SL); Bahamas, Hog Island, northern shore near Wend, on the beach; 1 August 1967. AMNH 27113 (1, 169 mm SL); West Indies; Harry Payne Bingham, February–April 1925. YPM ICH 000459 (1, 177 mm SL); West Indies; holotype of Cypselurus vitropinna. From the southwestern Atlantic, nine specimens (184.5–207.9) from three collections. ZMH (ISH 238/51) (1, 188 mm SL); Olympic Challenger, 18°49′S, 38°32′W; 17 December 1951. USNM 294694 (1, 175 mm SL); Undaunted Cruise 5, Sta. 215; 5 October 1966. USNM (?) uncat. (2, 198.7–207.9 mm SL); TABL 103136, Undaunted Cruise 5 Sta. 179, 14°29′S, 38°51′W; 16 September 1966. USNM 294694 (1, 184.5 mm SL); 18°05′S, 35°57′W. IORAS uncat. (4, 192–203 mm SL); R/V Ioffe Cruise 16 Sta. 1623–1624, 20°49′–21°00′S, 34°46′–48′W; 4–5 November 2004. Genus Exocoetus Linnaeus, 1758 Exocoetus Linnaeus, 1758:316 (type species E. volitans Linnaeus by monotypy). Halocypselus Weinland, 1858:385 (type species H. mesogaster Weinland = Exocoetus volitans by monotypy).

Diagnosis. Exocoetus differs from all other flyingfishes in having short pelvic fins, with the first or second rays the longest and inserted nearer to the snout than to caudal fin base, and nearer to pectoral fin insertion than to anal fin origin. Teeth on jaws absent or rudimentary. Upper jaw protrusible. Pectoral branch of lateral line absent, except some specimens of E. monocirrhus may have a rudiment present (Parin 1961). Description. Body round, slightly flattened ventrally. Greatest body depth 4.6–6.5 times in SL, width of body 1.2–1.4 times in its depth. Vertebrae (24–27) + (16–20) = 42–46. Predorsal scales 16–23; transverse scales 6–8½. Pectoral branch of lateral line absent (rarely present on one scale in Exocoetus monocirrhus). Swim bladder extends into haemal canal, reaching fifth to eighth caudal vertebra. Head 3.6–4.4 times in SL. Snout blunt, shorter than eye. Jaws subequal; jaw teeth usually absent (rarely few scattered teeth present on one or both jaws). Gill rakers on first arch 22–37. Dorsal fin rays 12–16; anal fin rays 12–15; pectoral fin rays I 12–16. Pectoral fins very long (1.2–1.4 times

Exocoetus obtusirostris Günther, 1866 Oceanic Two-wing Flyingfish Figures 5.14, 5.15 Tables 5.1, 5.26, 5.27, 5.28, 5.29 Exocoetus obtusirostris Günther, 1866:283 (original description; Cape Verde Islands; holotype BMNH 1864.6.6.51). Jordan et al. 1930:200 (listed). Bruun 1935:34–40, pl. 2, fig. 2 (description, Atlantic Ocean). Priol 1937:367–368 (description; eastern Atlantic). Breder 1938:33–37, fig. 16 (juveniles, western Atlantic). Poll 1949:255 (San Salvador). Poll 1953:184 (in part; description; eastern Atlantic). Duarte-Bello 1959:46

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Fishes of the Western North Atlantic

Figure 5.14. Oceanic Two-wing Flyingfish, Exocoetus obtusirostris. A. USNM 198384, 169.4 mm SL. Dipnetted at night by R. H. Gibbs, Jr. at 36°07′N, 73°25′W from M/V Delaware, 22 April 1960. B. USNM 198381, 28.5 mm SL. Dipnetted at night by R. H. Gibbs, Jr. at 35°50′N, 72°35′W from M/V Delaware, 19 July 1958. C. Pectoral fin of USNM 198384, 116.4 mm SL. Dipnetted at night by R. H. Gibbs, Jr. at 36°07′N, 73°25′W from M/V Delaware, 22 April 1960. Illustrations by H. E. Hamman.

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Family Exocoetidae differs from E. volitans in having fewer gill rakers (22–29 rather than 29–37), more transverse scales (usually 7½ rather than 6½) and anal fin usually originating slightly before dorsal fin; juveniles are humpbacked (elongate in E. volitans). As adults, E. obtusirostris is similar to the Indo-Pacific E. monocirrhus (the only distinctive character is lighter pectoral fins coloration), but juveniles of the latter have a single chin barbel (absent in E. obtusirostris). Exocoetus obtusirostris is similar to the two South Pacific species E. gibbosus and E. peruvianus that were recently separated from E. obtusirostris (Parin and Shakhovskoy 2000). Exocoetus obtusirostris differs primarily in a less developed postero-lateral process of the cleithrum, number of pectoral rays (mode of I 15 in E. obtusirostris rather than I 14 in both Pacific species) and in morphometrics (Parin and Shakhovskoy 2000). Description. Dorsal fin rays 12–15; anal fin rays 12–15; pectoral fin rays I 14–16 (Table 5.26). Vertebrae 24–27 + 16–19 = 42–46 (Table 5.27). Predorsal scales 18–23, usually 19–21; scales in transverse row from dorsal fin origin to lateral line 7–8, usually 7½ (Table 5.28). Gill rakers on the first arch 4–7 + 18–23 = 22–29, usually 24–28 (Table 5.29). Measurements based on specimens 140–185 mm SL. Body elongate, roundish in cross section, slightly flattened ventrally. Greatest depth 4.6–6.1 times in SL, width about 1.2–1.4 times in depth. Head moderate (3.6–4.1 times in SL); usually slightly shorter than dorso-caudal distance (1.0–1.2 times in the latter). Eye diameter 3.2–3.6 times in head length, shorter than interorbital space. Upper and lower jaws of nearly equal length, usually without teeth. Dorsal fin low, its longest ray (second or third) 9.0–11.0 times in SL. Anal fin usually originates just before dorsal fin origin (rarely as far posterior as under its second ray). Pectoral fins 1.3–1.4 times in SL, usually reaching behind origin of upper caudal lobe. Pelvic fins inserted much nearer to posterior margin of opercle than to origin of lower caudal lobe (operculo-pelvic distance 2.8–4.0 in pelvic-caudal distance), very short (6.5–8.2 times in SL); reaching middle of pelvic-anal distance or slightly posterior. Color. Body dark iridescent blue above, pale silvery below in life. Dorsal and caudal fins gray, anal and pelvic fins transparent, pectoral fins brownish gray with a rather broad colorless margin. Development. Juveniles (20–100 mm SL), body humpbacked; upper profile of head steeply sloping. No chin barbel. Maximum body depth 3.4–4.3 times in SL at 20–40 mm SL, 3.9–4.8 times in SL at 30–80 mm SL. Pelvic fins relatively longer than in adults (at SL 30–80 mm 4.0–6.7 times in SL), reaching (or

(listed; Cuba). Evans and Sharma 1963:55 (juveniles, central Atlantic). Caldwell and Caldwell 1964:13 (Caribbean Sea, waters of Colombia). Kovalevskaya 1964:217–222 (description of juveniles; in part: Atlantic material only). McKenney 1965:26–31 (description of juveniles; western Atlantic). Parin 1973:263 (northeastern Atlantic and Mediterranean Sea). Belyanina 1975:140 (larvae and juveniles, Caribbean Sea and Gulf of Mexico). Fahay 1975:16 (juveniles off Florida). John 1976:121–123 (larvae in central and western South Atlantic). Hoese and Moore 1977:151 (Gulf of Mexico). Gibbs 1978: EXOC Exoc 1 (distribution, figure, range in western Atlantic). Fedoryako 1980:581 (Sargasso Sea). Grudtsev et al. 1986:920–928 (morphometrics; distribution in Atlantic). Gorelova and Grudtsev 1987:481 (feeding; Atlantic). Nikolsky and Luschina 1990:135–136 (eastern equatorial Atlantic). Shiganova and Kovalevskaya 1991:94 (eggs, larvae and juveniles; northeastern Atlantic). Hunte et al. 1995:28 (eggs and larvae distribution; eastern Caribbean). McEachran and Fechhelm 1998:948, figure (Gulf of Mexico). Smith-Vaniz et al. 1999:172 (Bermuda). Parin and Shakhovskoy 2000:S51–S55 (description, Atlantic distribution). Parin 2003:1127, figure, map (description, western Atlantic). Cotten and Comyns 2006:964–965, figs. D, E (in part; early life history). Fahay 2007:810–811, fig. E (in part; early life history). Casazza and Ross 2008:352 (association with Sargassum, Gulf Stream off North Carolina). Collette 2010:35–36 (reproduction). Lewallen et al. 2011 (molecular systematics). Parin and Shakhovskoy 2016:2146 (description, east-central Atlantic). Gordeeva and Shakhovskoy 2017:289 (fig. 1 maximum likehood tree; DNA-barcoding). Lewallen et al. 2017 (phylogeny). Exocoetus holubii Steindachner, 1882:54 (original description; equatorial Atlantic; holotype NMW 76604). Halocypselus obtusirostris Jordan and Gilbert, 1883:378 (short description; north to Louisiana). Nichols 1922:49–51 (description of juvenile; Florida). Beebe and Tee-Van 1933:65 (Bermuda). Halocypselus evolans (not of Linnaeus 1766). Jordan and Meek 1885:49–50 (in part). Jordan and Evermann 1896a:729–730 (in part; listed). Exocoetus evolans (not of Linnaeus 1766). Günther 1910:363–364 (in part; description; Atlantic). Exocoetus gaussianus Lampe, 1914:224, pl. XI, fig. 6 (original description; 02°09′N, 17°38′W). Cypselurus holubi. Jordan et al. 1930:202 (listed).

Types. Exocoetus obtusirostris Günther 1866. Holotype BMNH 1864.6.6.51 (161 mm SL). Dorsal fin rays 13; anal fin rays 13. Predorsal scales 24; scales in transverse row from dorsal fin origin to lateral line 7. Paratype BMNH 1864.4.10.2 (117 mm SL). Collett (1896) redescribed and made drawings of the holotype of Exocoetus holubii Steindachner, 1882. Parin examined the holotype of Exocoetus gaussianus Lampe, 1914 (see Parin and Shakhovskoy 2000). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Oceanic Two-winged Flyingfish (English) and volador flecha (Spanish). Diagnostic Characters. Exocoetus obtusirostris 193

Fishes of the Western North Atlantic

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Family Exocoetidae

nearly reaching) origin of anal fin at 20–25 mm SL. Body evenly pigmented, without pigmented crossbars. Dorsal and anal fins blackish posteriorly. Pectoral fins of smaller juveniles usually transparent, at above 25–45 mm SL pigmented proximally. Pelvic fins in fishes under 34–47 mm SL mostly blackish, colorless in some specimens. Size. The maximum observed size for Exocoetus obtusirostris is 196 mm SL.

reproductive period is at least 80 days. Estimated total fecundity averages 10,300 eggs (Alekseev et al. 1989). Range. Exocoetus obtusirostris is endemic to the tropical and subtropical Atlantic Ocean (Fig. 5.15). In the western Atlantic, it is distributed between 46°N and 40°S (extreme occurrences in the North Atlantic current at 46°16′N, 35°29′W: MCZ 156524, a juvenile, 31 mm SL; in the Brazil current at 39°54′S, 53°13′W: AtlantNIRO, no number, 191 mm SL); in the eastern Atlantic between 30°N and 30°S. Common in the Gulf of Mexico; occasional strays

Biology Predators. Exocoetus obtusirostris are consumed by a variety of predatory fishes, including dolphinfishes, tunas, snake mackerels, and Rainbow Runner, Elagatis bipinnulata (Vaske et al. 2006), and also by omastrephid squids, seabirds, and dolphins. Food. Exocoetus obtusirostris feeds on small zooplankton, predominantly copepods (Nannocalanus minor, Euchaeta marina, Scolecithryx danae, Candacia pachidactyla, C. curta, C. aethiopica, Undinula vulgaris, Labidocera acutifrons, Farranula spp., and Oncaea spp.) and also chaetognaths and amphipods (Gorelova and Grudtsev 1987). Eastern Atlantic specimens of E. obtusirostris are reported to feed on copepods, gastropod larvae, and fry of Mysidacea (Poll 1953). Reproduction. Females and males are sexually mature at about 140–170 mm SL and all females larger than 170 mm SL are in spawning condition during the whole year (Alekseev et al. 1989). Otoliths of mature fishes, including the largest specimens, consistently have one opaque and one hyaline zone, indicating that longevity is 1 to 1.5 years (Grudtsev et al. 1986), and nearly all fishes die after the first reproductive season. Spawning is intermittent, with up to 20 batches laid in five or more days; egg diameter 2.8–3.4 mm, each batch consisting of 420–890 (x̅  630) eggs. Mean duration of individual

Figure 5.15. Distribution of Oceanic Two-wing Flyingfish, Exocoetus obtusirostris, based on Parin and Shakhovskoy (2000). Map by W. E. Bemis.

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Fishes of the Western North Atlantic 1998:949 (Gulf of Mexico). Monteiro et al. 1998:400 (distribution; northeastern Brazil). Smith-Vaniz et al. 1999:172–173 (Bermuda). Parin and Shakhovskoy 2000:S33–S44 (description, distribution). Parin 2003:1128, figure, map (description, western Atlantic). Cotten and Comyns 2006:966–967, fig. A–H (early life history). Fahay 2007:812–813, figs. A–H (early life history). Collette 2010:35–36 (reproduction). Lewallen et al. 2011 (molecular systematics). Lewallen et al. 2016 (population genetic structure). Parin and Shakhovskoy 2016:2147 (description, east-central Atlantic distribution). Gordeeva and Shakhovskoy 2017:289 (fig. 1 maximum likehood tree; DNA-barcoding). Lewallen et al. 2017 (phylogeny; sister to the other four species in the genus). Halocypselus mesogaster (not of Bloch 1795). Weinland 1858:385. Halocypselus evolans. Jordan and Gilbert 1883:377–378 (description; Atlantic coast of North America). Jordan and Meek 1885:48–50 (in part). Jordan and Evermann 1896a:729–730 (in part; short description; North America). Nichols 1922:49– 51 (juveniles off Florida). Nichols and Breder 1928:427–428, figures (30°N, 60°W). Breder 1932:22 (Bird Key, Tortugas). Beebe and Tee-Van 1933:64–65 (Bermuda). Longley and Hildebrand 1941:31 (Tortugas). Erdman 1956:324 (listed; Vieques Passage, Puerto Rico).

to Portugal and Mediterranean Sea (Wheeler 1969; Parin 1973). Exocoetus obtusirostris is present in a wide range of water temperature, from 17.6 to 29.2 °C, and is more abundant than E. volitans in productive areas adjacent to the zones of coastal upwelling, the Gulf of Guinea and in the Caribbean Sea (Grudtsev et al. 1986). Geographical Variation. There is no significant variation in morphometrics between samples from the eastern and western Atlantic Ocean, but the Caribbean population differs in having a slightly larger head, eye and snout, and shorter pelvic and pectoral fins (Grudtsev et al. 1986). The northern and southern Atlantic populations also do not differ significantly (Parin and Shakhovskoy 2000). Study Material. Description based on 68 northern Atlantic specimens (31–184 mm SL) and 29 southern Atlantic specimens (27–184 mm SL; see Parin and Shakhovskoy [2000] for full list and localities), as well as data of Bruun (1935) and Breder (1938). Additional material includes meristics of 55 (148– 172 mm SL) western Atlantic specimens examined by N. V. Parin.

Types. Exocoetus volitans Linnaeus, 1758. Holotype in Museum of the University of Uppsala (UUZM) (fide Bruun 1934:133–134). Gill rakers 30 or more. Exocoetus evolans Linnaeus, 1766. Holotype BMNH (in dry condition). Scales between dorsal fin origin and lateral line 6 (fide Bruun 1935:32). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Tropical Two-winged Flyingfish (English) and volador tropical (Spanish). Diagnostic Characters. Exocoetus volitans differs from E. obtusirostris in having more gill rakers (29–37 rather than 22–29), fewer transverse scales (usually 6½ rather than 7½), and anal fin usually originating under first or second dorsal fin ray; juveniles elongate (humpbacked in E. obtusirostris). It also differs in many osteological characters (see Parin and Shakhovskoy 2000) and is genetically distinct (Lewallen et al. 2011, 2017; Gordeeva and Shakhovskoy 2017). Description. Dorsal fin rays 13–15; anal fin rays 12–15; pectoral fin rays I 13–15 (Table 5.26). Vertebrae 24–26 + 18–20 = 43–45 (Table 5.27). Predorsal scales 16–21, usually 18–20; scales in transverse row from dorsal fin origin to lateral line 6 or 6½, usually 6½ (Table 5.28). Gill rakers on the first arch 6–9 + 24–29 = 29–37, usually 31–34 (Table 5.29). Measurements based on specimens (140–179 mm SL) from the Atlantic Ocean. Body elongate (greatest depth 5.0–6.5 times in SL); moderately compressed (greatest width 1.2–1.3 times in depth). Head moderate (3.6–4.0 times in SL); slightly shorter than dorso-caudal distance (1.1–1.3 times in the

Exocoetus volitans Linnaeus, 1758 Tropical Two-wing Flyingfish Figures 5.16, 5.17 Tables 5.1, 5.26, 5.27, 5.28, 5.29 Exocoetus volitans Linnaeus 1758:316 (original description, “habitat in alto Pelago, Europaeo et Americano”; holotype ZMUU Linn. Coll. 59 (1)). Jordan and Evermann 1896b:322 (in part; listed). Jordan et al. 1930:200 (in part: listed). Bruun 1934:133–134 (on Linnaean type specimen). Bruun 1935:28–34, pl. 2, fig. 1 (description, distribution; Atlantic). Priol 1937:366–367 (description; eastern Atlantic). Breder 1938:30–53, fig. 15 (juveniles, distribution; western Atlantic). Duarte-Bello 1959:46 (listed; Cuba). Evans and Sharma 1963:53–55 (juveniles, central Atlantic). McKenney 1965:17–25 (description of juveniles; western Atlantic). Mago Leccia 1970:89 (listed, Venezuela). Parin 1973:263 (northeastern Atlantic and Mediterranean Sea). Sauskan 1973:151–152 (eastern Atlantic). Belyanina 1975:140 (larvae and juveniles off Puerto Rico and in Caribbean). Fahay 1975:16–18 (juveniles off Florida). John 1976:121– 123 (larvae in central and western South Atlantic). Gibbs 1978: EXOC Exoc2 (description, figure, range in western Atlantic). Fedoryako 1980:581 (Sargasso Sea). Khachaturov 1983 (digestive system, gill rakers and pharyngeal teeth). Grudtsev et al. 1986:920–928 (morphometrics; distribution in Atlantic). Heemstra and Parin 1986:394 (short description; South Africa). Gorelova and Grudtsev 1987:481 (feeding; Atlantic). Nikolsky and Luschina 1990:135–136 (eastern equatorial Atlantic). Shiganova and Kovalevskaya 1991:94– 95 (eggs, larvae and juveniles; northeastern Atlantic). Hunte et al. 1995:28–35 (eggs and larvae distribution; eastern Caribbean). Oxenford et al. 1995b:44 (distribution of juveniles; eastern Caribbean). ?McEachran and Fechhelm

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Family Exocoetidae

Figure 5.16. Tropical Two-wing Flyingfish, Exocoetus volitans. A. USNM 159035, 161.9 mm SL. Collected south of Walton County, Florida, U.S.A. from M/V Oregon, Sta. 1297, 22 April 1960. B. USNM 198375, 35.9 mm SL. Dipnetted at night by R. H. Gibbs, Jr. at 36°07′N, 73°25′W from M/V Delaware, 22 April 1960. Illustrations by H. E. Hamman.

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Fishes of the Western North Atlantic latter). Eye diameter 3.5–3.9 times in head length, shorter than interorbital space. Jaws subequal, usually without teeth. Dorsal fin low, the longest ray (usually second) 7.9–11.0 times in SL. Anal fin originates under first to third dorsal fin ray. Pectoral fins 1.3–1.4 times in SL, reaching posterior to origin of upper caudal lobe. Pelvic fins inserted much nearer to posterior margin of opercle than to origin of lower caudal lobe (operculo-pelvic distance 2.9–3.6 times in pelvic-caudal distance), very short (6.5–7.7 times in SL); reaching middle of pelvic-anal distance or slightly posteriorly. Several otoliths of Exocoetus volitans from Madeira were illustrated by Nolf (2013: pl. 153). Color. Body dark iridescent blue above, pale silvery below in life. Dorsal and caudal fins gray, anal and pelvic fins transparent, pectoral fins gray with a narrow colorless margin. Development. Body of juveniles (19–98 mm SL) elongate, fusiform; upper profile of head gently sloping. No chin barbel. Maximum body depth 4.8–6.0 times in SL at less than 40 mm SL, and 5.1–5.8 times in SL at larger size. Pelvic fins relatively longer than in adults (5.8–7.5 times in SL). Body pale, at 20–60 mm SL usually with two vertical pigmented bars on tail. Dorsal and anal fins transparent, blackish posteriorly at 20–40 mm SL. Pectoral fins transparent proximally and slightly

pigmented distally in smaller juveniles, entirely gray in specimens larger than 40 mm SL. Pelvic fins transparent. Size. The maximum known size for Exocoetus volitans in the Atlantic Ocean is 189 mm SL. Biology Predators. Exocoetus volitans is consumed by a large variety of predatory fishes, such as dolphinfishes, tunas, snake mackerels, and Rainbow Runner, Elagatis bipinnulata (Vaske et al. 2006), and also by omastrephid squids, seabirds, and dolphins. In the eastern Pacific, porpoise (Stenella sp.) and Yellowfin Tuna, Thunnus albacares, have been recorded as feeding on Exocoetus volitans (Perrin et al. 1973). Exocoetus volitans is important in the diet of sea birds (Sula leucogaster, Anous stolidus, and A. minutus) off Saint Paul Archipelago (Vaske et al. 2005). Flyingfishes consumed by Gempylus serpens in the Indo-Pacific were almost all Exocoetus volitans (I. B. Shakhovskoy, unpublished data). Food. Exocoetus volitans feeds mainly on copepods in the Atlantic from 20°N to 30°S and 40°W to 6°E (Gorelova and Grudtsev 1987). The diet of E. volitans in the eastern Pacific population was specialized and composed almost exclusively of copepods (Van Noord et al. 2013). Reproduction. As in Exocoetus obtusirostris, all specimens of E. volitans smaller than 140 mm SL are immature and all those larger than 170 mm SL are ripe (Alekseev et al. 1989). Otolith annuli show that most fish live less than 1.5 years (Grudtsev et al. 1986, Alekseev et al. 1989). Spawning of E. volitans occurs throughout the year, but in the eastern tropical Atlantic it is concentrated from October to April. Spawning is intermittent; egg diameter 2.4–2.8 mm (2.7–3.0 mm in Indo-Pacific), each batch consisting of 327–418 eggs (x̅  370) (Alekseev et al. 1989). Range. Exocoetus volitans is an oceanic species, and undoubtedly the most abundant flyingfish in offshore tropical waters of all oceans at temperatures of 20.0 to 29.0 °C (Grudtsev et al. 1986). In the Atlantic Ocean, it is common between 35°N and 30°S in the west and between 20°N and 20°S in the east, but rare in the Gulf of Mexico, where only juvenile specimens accidentally drawn by currents are known (Fig. 5.17; see also Lewallen et al. 2016: fig. 1). Occasional adult stragglers may be found farther north and south, a few specimens reported even from the western Mediterranean (Bruun 1935; Collette et al. 1997; Abdul Malak et al. 2011). In the eastern North Atlantic, offshore from the Azores and Cape Verde Islands, E. volitans made up 10.4%

Figure 5.17. Atlantic distribution of Tropical Two-wing Flyingfish, Exocoetus volitans, based on Parin and Shakhovskoy (2000). Map by W. E. Bemis.

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Family Exocoetidae farther than a vertical through the second, rarely third, ray of dorsal fin). Dorsal fin low. Pectoral fins long, reaching the posterior end of dorsal fin base or farther, their first or first and second rays unbranched. Pelvic fins long, reaching past middle of the anal fin base; situated somewhat closer to posterior margin of operculum than to the origin of caudal fin lower lobe or between these levels (very rarely a little closer to the latter). In adults, head 3.7–5.2 times in SL, eye diameter 2.4–4.1 times in head length, the greatest body depth 5.0–7.4 times in SL, body width 1.0–1.4 times in its depth. Jaw teeth are small, unicuspid, and conical. Mouth not protrusible. Lower jaw a little longer than upper jaw or equal to it. Jaw teeth unicuspid. Palatine teeth usually absent (present in Hirundichthys speculiger). Pectoral branch of lateral line absent. Filaments on eggs have a bipolar arrangement. Juveniles elongate with mottled paired fins; chin barbels absent (Shakhovskoy and Collette, in press). Species. About ten species are recognized in two subgenera, Hirundichthys and Danichthys (Parin and Belyanina 2002b; Shakhovskoy and Parin 2013a, 2013b; Shakhovskoy and Collette, in press). Five species in the Atlantic Ocean, four in the western Atlantic. Range. Species of Hirundichthys are pelagic in neritic and oceanic areas of tropical and subtropical Atlantic, Indian, and Pacific oceans (Parin 1961, 2003; Shakhovskoy and Collette, in press).

of the total fish and fish larvae caught by neuston nets (Andres and John 1984). Geographical Variation. Atlantic and Indo-Pacific populations of Exocoetus volitans can differ in length of pelvic fins, being on the average 2%–3% of SL longer in the former (Parin 1960; McKenney 1965), but this was not confirmed (Parin and Shakhovskoy 2000) and it is probably a result of differences in methods. No significant morphometric differences were found between samples of E. volitans from the western Atlantic off northern Brazil and from the eastern Atlantic off Cape Verde and Gulf of Guinea (Grudtsev et al. 1986). A parsimony-based haplotype network did not reveal population subdivision among ocean basins, but an analysis of molecular variance revealed limited, but statistically significant, population structure between Pacific and Atlantic oceans and a spatially unbiased Bayesian approach identified two circumtropical population clusters north and south of the equator (Lewallen et al. 2016). Study Mater ial. Description based on 69 Atlantic specimens (29.5–176 mm SL; see Parin and Shakhovskoy [2000] for full list and localities), as well as data of Bruun (1935) and Breder (1938). Additional material includes, from the western Atlantic, 112 specimens (19–168 mm SL) from 65 collections at USNM and IORAS, including 5 specimens (24–39 mm SL) from the Gulf of Mexico (USNM 299092, 299094). From the eastern Atlantic, 96 specimens (19–171 mm SL). Meristics of 51 western Atlantic specimens examined by N. V. Parin.

Hirundichthys affinis (Günther, 1866) Fourwing Flyingfish Figures 5.18, 5.19 Tables 5.1, 5.30, 5.31, 5.32, 5.33

Genus Hirundichthys Breder, 1928 Hirundichthys Breder, 1928:14, 20 (type species Exocoetus rubescens Rafinesque, 1818 by original designation).

Exocoetus af f inis Günther, 1866:288 (orig inal description; Atlantic, western Africa, Cuba; syntypes BMNH 1855.12.26.455, Cuba, BMNH ?.6.22.59, Atlantic; BMNH ?1843.1.10.21, West Africa). Exocoetus roberti Müller and Troschel, 1848:675 (original description, Barbados; holotype ZMB 2903). Exocoetus scylla Cope, 1871:481 (original description; Gulf of Mexico coast; holotype ANSP 7500). ?Exocoetus volitans (not of Linnaeus). Jordan and Meek 1885:57 (in part: Atlantic, northward to New England). Bean 1903:330 (description; Atlantic, northward to Newfoundland, Grand Banks). Exocoetus vinciguerrae (not of Jordan and Meek 1885). Jordan and Meek 1885:56 (in part: 126.5 mm SL specimen from Gulf of Mexico). Jordan and Evermann 1896a:734 (description; in part: Newfoundland, St. Martin Island, Gulf of Mexico). Exocoetus rufipinnis (not of Valenciennes in Cuvier and Valenciennes 1847). ?Jordan and Meek 1885:58 (description; tropical America). Jordan and Evermann 1896a:735

Diagnosis. Flyingfishes of genus Hirundichthys differ from all other flyingfishes (except for Exocoetus and Parexocoetus) in the position of the anal fin origin under or ahead of the third dorsal fin ray. They differ from Exocoetus mainly in position and length of pelvic fins and from Parexocoetus in length of pectoral and pelvic fins. Description. Dorsal fin rays 9–13, very rarely 8; anal fin rays 10–13, very rarely 9; pectoral fin rays 14–21, very rarely 13. Gill rakers 22–38, usually 25–31. Predorsal scales 25–37; scales in an oblique transverse row between the first ray of dorsal fin and lateral line 5½–8½. Vertebrae 41–48 total. The anal fin origin is on a vertical line with the dorsal fin origin, in front of it, or slightly behind (but not 199

Fishes of the Western North Atlantic

Figure 5.18. Fourwing Flyingfish, Hirundichthys affinis. A. USNM 196611, 177.3 mm SL. Collected at 26°40′N, 88°45′W from M/V Oregon, Sta. 1218, 14 December 1954. B. USNM 198379, 22.5 mm SL. Dipnetted at night by R. H. Gibbs, Jr. and B. B. Collette at 37°44′N, 65°42′W from M/V Delaware, 13 September 1957. C. Pectoral and pelvic fins of USNM 198385, 74.0 mm SL. Dipnetted at night by R. H. Gibbs, Jr. at 36°07′N, 73°25′W from M/V Delaware, 22 April 1960. D. Pectoral and

200

Family Exocoetidae

pelvic fins of USNM 198385, 58.9 mm SL. Dipnetted at night by R. H. Gibbs, Jr. at 36°07′N, 73°25′W from M/V Delaware, 22 April 1960. E. Pectoral and pelvic fins of USNM 198380, 30.3 mm SL. Dipnetted at night by R. H. Gibbs, Jr. and B. B. Collette at 38°17′N, 70°00′W from M/V Delaware, 25 September 1957. Illustrations by H. E. Hamman.

201

Fishes of the Western North Atlantic Schomburgk, 1848. Holotype ZMB 2903 (228 mm SL); Barbados. Dorsal fin rays 11; anal fin rays 12; pectoral fin rays I 16. Predorsal scales 30; scales in transverse row from dorsal fin origin to lateral line 7. Gill rakers 8 + 21= 29. Vertebrae 28 + 18 = 46. Exocoetus vinciguerrae Jordan and Meek, 1885. Paratype ANSP 7496 (126.5 mm SL); Gulf of Mexico. Dorsal fin rays 11; anal fin rays 12; pectoral fin rays I 16. Predorsal scales 32; scales in transverse row from dorsal fin origin to lateral line 7. Gill rakers 7 + 19 = 26. Vertebrae 46. Exocoetus scylla Cope, 1871. Holotype ANSP 7500 (196 mm SL); said to be from near Tabasco, Mexico. Dorsal fin rays 11; anal fin rays 11; pectoral fin rays I 17. Predorsal scales 31; scales in transverse row from dorsal fin origin to lateral line 7. Gill rakers 7 + 18 = 25. Pectoral fins with a small “mirror.” Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Fourwing Flyingfish (English), volador golondrina (Spanish), and exocet à frange blanche (French). Known as volant in Dominica (Oxenford et al. 2007). Diagnostic Characters. Hirundichthys affinis differs from H. speculiger in pectoral fin coloration (small noncontrasting triangular band (so-called mirror) and a narrow posterior edging), lack of palatine teeth (although there are exceptions in both species), fewer pectoral fin rays (15–19, usually 16–18, rather than 17–21, usually 18–19). Juveniles differ in having a less extensive membrane between the first and second rays and the length ratio of the second and third pectoral fin rays (their lengths are approximately equal or the second ray is only slightly shorter than the third), in pigmentation pattern of pelvic fins and lower jaw, in length of pectoral fins and head depth. Hirundichthys affinis differs from H. volador and H. rufipinnis in having only the first pectoral fin ray unbranched, and shorter pectoral fins with lighter coloration. Description. Dorsal fin rays 10–13, usually 10–11; anal fin rays 9–13, usually 11–12; pectoral fin rays I 14–18, usually I 15–17 (Table 5.30). Vertebrae 27–30 precaudal + 16–19 caudal = 44–47 total (Table 5.31). Predorsal scales 28–33, usually 29–32; scales in transverse row from dorsal fin origin to lateral line 5½–8, usually 6½–7½ (Table 5.32). Gill rakers on the first arch 5–9 upper + 17–23 lower = 23–31 total, usually 25–28 (Table 5.33). Measurements based on specimens (160–240 mm SL) from the western Atlantic. Body elongate, nearly rectangular in cross section, almost flat ventrally. Depth 5.4–6.9 times in SL, body width 1.0–1.3 times in depth. Head length 3.8–4.4 times in SL,

(description; in part: tropical America [Tabasco, Panama], Barbados). ?Danichthys rondeletii (not of Valenciennes in Cuvier and Valenciennes 1847). Fowler 1944:96, fig. 120 (in part: one specimen total length 62 mm; Atlantic, Courtown Keys). Hirundichthys affinis. Bruun 1935:67–70, pl. 6, figs. 1, 3 (description, western Atlantic distribution). Breder 1938:78–83, figs. 40–42 (description of juveniles; northwestern Atlantic). Fowler 1944:96–97, fig. 118 (description; Atlantic, Courtown Keys). Poll 1953:194–196, figs. 82–83 (description, early life stages; eastern Atlantic). Evans 1961:483–502 (early life history stages; western Atlantic). Lewis et al. 1962:73–94 (biolog y; Barbados). Caldwell and Caldwell 1964:14 (Caribbean Sea, waters of Colombia). Lewis 1964:381–386 (tagging data; Barbados). Cruz 1965:19 (fishery; northeastern Brazil). McKenney 1965:42–46 (description of juveniles; western Atlantic). Sauskan 1973:151–152 (eastern Atlantic). Fedoryako 1980:581 (Sargasso Sea). Matsuura 1983:198 (short description, photograph; Suriname and Guyana). Gorelova and Grudtsev 1987:481 (feeding; Atlantic Ocean). Nikolsky and Luschina 1990:135–136 (eastern equatorial Atlantic). Campana et al. 1993:211–219 (age determination; western Atlantic). Oxenford 1994:49–61 (migration; eastern Caribbean). Oxenford et al. 1994:585–592 (otolith age validation and growth rate; eastern Caribbean). Belyanina 1994:102–106, figs. 5–9 (early life history stages; comparison with Hirundichthys speculiger); northwestern and northeastern Atlantic). Oxenford et al. 1995a:16–21 (distribution of adults; eastern Caribbean). Oxenford et al. 1995b:45 (distribution of juveniles; eastern Caribbean). Hunte et al. 1995:28– 35 (distribution of eggs and larva; eastern Caribbean). McEachran and Fechhelm 1998:951, figure (Gulf of Mexico). Monteiro et al. 1998:400 (northeastern Brazil). Gomes et al. 1998:1029–1037 (genetic variability of populations (RAPD); west-central Atlantic). Gomes et al. 1999:485–497 (at least three unit stocks in west-central Atlantic based on mitochondrial D-loop variation). Smith-Vaniz et al. 1999:173 (Bermuda). Khokiattiwong et al. 2000:43–60 (seasonal abundance and reproduction; Barbados). Parin 2003:1129 (description; west-central Atlantic). Araújo and Chellappa 2002a:131–134 (histology of gonads; Brazil). Araújo and Chellappa 2002b:691–703 (reproduction; Brazil, Caiçara). Cotton and Comyns 2006:980–981, figs. A–E (early life history stages; North Atlantic). Fahay 2007:814–815, figs. A–H (early life history stages). De Castro et al. 2008:139–142, fig. 2b (in part: two specimens 14 and 15 mm SL; Formoso River estuary, 8°35′S, 35°95′W). Casazza and Ross 2008:352, 355 (associated with Sargassum; Gulf Stream off North Carolina). Lewallen et al. 2011:164 (molecular systematics). Shakhovskoy and Parin 2013b:526–535, figs. 5, 7, 8 (review). Nóbrega et al. 2015:801 (preyed on by Coryphaena hippurus). Oliveira et al. 2015:198–206 (reproductive biology; Brazil). Parin and Shakhovskoy 2016:2149 (description, east-central Atlantic distribution). Gordeeva and Shakhovskoy 2017:289 (fig. 1, maximum likehood tree; DNA-barcoding).

Types. The following type material was studied and identified as Hirundichthys affinis. Exocoetus affinis Günther, 1866. Syntype BMNH 1855.12.26.455 (218 mm SL); Cuba. Dorsal fin rays 11; anal fin rays 11; pectoral fin rays I 17. Predorsal scales 33. Gill rakers 8 + 21 = 29. Exocoetus rober ti Müller and Troschel in 202

Family Exocoetidae Table 5.30. Fin ray counts for Hirundichthys affinis, based on Shakhovskoy and Parin (2013b). Dashes indicate a value of zero. Fin rays Dorsal

Anal

Pectoral

Locality

10

11

12

13

9

10

11

12

13

15

16

17

18

19

Western Atlantic Eastern Atlantic Species total

17 10 27

62 28 90

13 5 18

1 — 1

1 — 1

2 2 4

44 20 64

43 21 64

2 — 2

2 1 3

16 4 20

34 29 63

17 11 28

1 — 1

Table 5.31. Vertebral counts of Hirundichthys affinis, based on Shakhovskoy and Parin (2013b), in which vertebrae were counted without the urostyle. In this work, we have included the urostyle in vertebral counts. Dashes indicate a value of zero. Vertebrae Precaudal

Caudal

Total

Locality

27

28

29

30

16

17

18

19

44

45

46

47

Western Atlantic Eastern Atlantic Species total

— 2 2

10 2 12

10 10 20

— 1 1

1 1 2

12 12 24

6 1 7

1 — 1

— 1 1

4 3 7

15 9 24

1 2 3

0.8–1.1 times in dorso-caudal distance. Eye 3.0–3.8 times in head length, 1.0–1.4 times in interorbital space. Jaws subequal. Jaw teeth conspicuous, conical. Dental index about 0.14. Palatine teeth usually absent (present in one of 66 specimens studied by Shakhovskoy and Parin [2013b]). Dorsal fin low, its longest ray (second or third) 8.8–14.5 times in SL. Anal fin originating in line or one or two rays behind dorsal fin origin (rarely slightly before). Pectoral fin 1.4–1.6 times in SL, reaching past base of dorsal fin, only first ray unbranched. Pelvic fins 3.2–4.3 times in SL, reaching to middle of anal fin base or farther; inserted slightly nearer to posterior margin of opercle than origin of caudal fin base (operculo-pelvic distance 1.0–1.2 times in pelvic-caudal distance). Color. Body dark above, pale below. Dorsal and caudal fins gray; anal fin transparent; pectoral fins dark gray or light brown with small unpigmented triangular crossband (“mirror”) and narrow outer margin; pelvic fins light. Development. Juveniles (40–100 mm SL) differ significantly from adults in proportions and coloration. Interorbital space and eye are larger, and snout shorter; pelvic fins inserted more anteriorly, dorsal and anal fins higher, pectoral fins shorter, and pelvic fins longer (see Shakhovskoy and Parin [2013b] for details). Juveniles without barbel. Body of juveniles up to 50 mm SL, usually with one to four dark bands on ventral side. Underside of lower jaw

usually pigmented in larvae and juveniles. Pectoral and pelvic fins mottled to about 80–100 mm SL. In juveniles 40–110 mm SL, membrane between the first and second pectoral rays relatively narrow, second pectoral ray only slightly shorter than third one or equal to it. Upper part of dorsal fin with black spot to about 100–120 mm SL. Lower lobe of caudal fin darker than upper lobe up to about 60 mm SL. Size. T he ma x i mu m obser ved si ze for Hirundichthys affinis in the western Atlantic is 238 mm SL; up to 262 mm SL in the eastern Atlantic (Shakhovskoy and Parin 2013b). Biology Predators. Consumed by Sailfish, large tunas, dolphinfishes, and Wahoo (Oxenford 1986; Nóbrega et al. 2015). Parasites. The cymothoid isopod Glossobius impressus was reported from Hirundichthys affinis in the Caribbean (Bunkley-Williams and Williams 2000). Food. Hirundichthys affinis feeds mainly on small fishes like Vinciguerria, and also amphipods and copepods (Gorelova and Grudtsev 1987). In the Barbados, fish larvae were the main food followed by large zooplankton (copepods, amphipods, and decapods); small quantities of H. affinis eggs were also consumed (Lewis et al. 1962). Remains of 203

Fishes of the Western North Atlantic Table 5.32. Number of predorsal and transverse scales of Hirundichthys affinis, based on Shakhovskoy and Parin (2013b). Dash indicates a value of zero. Scales Predorsal Locality Western Atlantic Eastern Atlantic Species total

Transverse

28

29

30

31

32

33

6

6½

7

7½

8

4 2 6

7 4 11

21 9 30

28 12 40

11 11 22

3 5 8

4 1 5

16 13 29

31 21 52

13 7 20

1 — 1

Table 5.33. Number of gill rakers on the first arch of Hirundichthys affinis, based on Shakhovskoy and Parin (2013b). Dashes indicate a value of zero. First arch gill rakers Locality

23

24

25

26

27

28

29

30

31

Western Atlantic Eastern Atlantic Species total

1 — 1

3 1 4

10 8 18

21 17 38

18 11 29

8 4 12

5 2 7

1 — 1

1 — 1

pteropods and amphipods were found in the stomachs of eastern Atlantic H. affinis (Poll 1953). Reproduction. Mature ovarian eggs of a female Hirundichthys affinis caught off Barbados had diameters of about 1.5–1.6 mm with 8–14 long (more than 4 mm) and thick filaments (central filament the thickest), projecting from a limited area and a small disk of short threads at the opposite pole (Bruun 1935; Evans 1961). Larger egg diameters (2.0– 2.5 mm) were reported for waters of Brazil (Araujo and Chellappa 2002a). Hatching occurred four to five days after fertilization at 26 °C (Evans 1961). Hirundichthys affinis reaches sexually maturity in seven to eight months at about 190 mm SL (Bruun 1935; Oxenford et al. 1994). It is an annual species, with a maximum age less than two years (Campana et al. 1993). Spawning occurs between November and July, with a small peak from November to January and a major peak from May to July. Individuals probably spawn several times within the season (Oxenford et al. 2007). Total fecundity ranges between 4,100 and 9,200 eggs (Parin 2003). Eggs are negatively buoyant with adhesive filaments for attaching to spawning substrata. Historically, it was thought that eggs were laid on floating objects, but Lao et al. (2007) found a surprisingly low association of flotsam and flyingfish eggs, with flotsam with eggs attached extremely rare. Scarcity of eggs could represent an overall scarcity of spawning substrata, or perhaps flyingfish use the flotsam so heavily that it quickly sinks from the added weight of the eggs. However, to maintain current population sizes, and

considering how little flotsam was observed, either flyingfish are spawning on flotsam elsewhere or flyingfish spawning may not rely on floating material, perhaps using instead submerged substrata or the bottom (Lao et al. 2007). Surface spawning is preferred, but H. affinis may spawn at least 20 m below the surface even when surface substrata are available (Oxenford 2007). It is likely that there is heavy post-spawning mortality (Oxenford et al. 2007). Histology of the gonads of Hirundichthys affinis collected off Rio Grande do Norte, Brazil, indicated that mature males and females were most abundant in May and June (Araújo and Chellappa 2002a). Males and females were found in a ratio of 1:1 and absolute fecundity ranged from 4,400 to 6,400 oocytes (x̅  5,400) (Araújo and Chellappa 2002b). However, fecundity of H. affinis in this region can be much greater, ranging from 7,398 to 10,021 eggs (x̅  9,092) (Oliveira et al. 2015). All life stages are sequentially present through the year off Barbados (Oxenford et al. 2007). Growth rate is variable and juveniles that hatch in warmer months (April to July, toward the end of the spawning season) grow significantly faster than those that hatch in colder months (November to March, toward the beginning of the spawning season). A von Bertalanffy growth curve fitted to juvenile size-at-age data gave daily preliminary growthcurve parameters of to = 2.85 d and k = 0.00854 with an asymptotic length (L∞) of 245 mm fork length (Oxenford et al. 1994). Relationship to Humans. Hirundichthys affinis is a 204

Family Exocoetidae

Figure 5.19. Distribution of Fourwing Flying f ish, Hirundichthys affinis, based on Shakhovskoy and Parin (2013b). Map by W. E. Bemis.

target species of artisanal fisheries off the Antilles, Barbados, Martinique, Dominica, St. Lucia, and Tobago (Ogawa and Alves 1971). This species supports the largest single-species fishery in the eastern Caribbean (Khokiattiwong et al. 2000). It is taken with gill nets, scooping with handheld dip nets, and with baited hooks near land. Gill nets of stretched mesh sizes of 3.18 and 3.81 cm effectively target fish available off Barbados (Mahon et al. 2000). The fishery season extends from January to May in the eastern Caribbean. Hirundichthys affinis is also important as prey and bait for species taken by sport and commercial fisheries (Oxenford et al. 1995a). Hirundichthys affinis is also economically important in Rio Grande do Norte, Brazil (Cruz 1965; Araújo and Chellappa 2002a). Here, the fishery season is from April to July or August when the fish come close to the coast. A triangular net called a jereré is used (see Cruz 1965: fig. 2). The mean fork length in the fishery is 235.5 mm (Cruz 1965). Range. Hirundichthys affinis is mainly neritic and abundant in the western Atlantic from 38°N (USNM 198379, 37°44′N, 65°42′W) to 21°S (IORAS 03181, 21°04′S, 35°57′W), including the Gulf of Mexico and the Caribbean Sea (Fig. 5.19). Juveniles are found in the Gulf Stream north to about 35°N. The species is also known from off Brazil and in the tropical eastern Atlantic. It rarely occurs in open waters, 205

especially during spawning season when adults are most abundant close to islands, especially the leeward side (Oxenford et al. 1993; Shakhovskoy and Parin 2013b). Mapping the distribution and relative abundance in Caribbean waters surrounding Barbados, Dominica, Grenada, Martinique, St. Lucia, St. Vincent, and Tobago revealed that of 15,346 adult flyingfishes, 47% were H. affinis (Oxenford et al. 1995a). The highest abundance was recorded west of the Lesser Antilles island chain (Oxenford et al. 1995a). A tagging experiment of 7,019 individuals (recapture rate 4.5%) suggests that the eastern Caribbean is not segregated into island-specific populations. Dispersal occurred in all directions, although northwest was most common, and dispersion rate was not affected by fish size or sex, but varied with maturity of the fish (maturing and mature fish moving more than running ripe or spent individuals) and time of year released (Oxenford 1994). Around Barbados, H. affinis shows large seasonal variation in abundance, common from December to June and scarce from July to November. Geographic Variation. Samples of Hirundichthys affinis from the eastern and western parts of the Atlantic Ocean are quite similar in meristic and morphometric characters. However, adult fish are somewhat different in coloration of pectoral fins: individuals with light brown fins prevail in the western Atlantic, whereas in the eastern Atlantic individuals with dark brown fins, often without a “mirror,” prevail. In addition, there are small differences in the pigmentation of pectoral and caudal fins in juveniles. These populations probably merit assignment of subspecies status (Shakhovskoy and Parin 2013b). Meristics of a sample of 156 Hirundichthys affinis specimens from off Brazil (Caiçara, Rio Grande do Norte) showed the following counts: dorsal fin rays 9–12, usually 11; anal fin rays 10–13, usually 11–12; pectoral fin rays 16–21, usually 17. Predorsal scales 29–32, usually 30. Gill rakers (6–8) + (18–22) = (24–29), usual count 7 + 20 = 27. Vertebrae 44–47, usually 46 (Cruz and Soares 1966). These fish differ from those mainly from the northwestern Atlantic (Shakhovskoy and Parin 2013b) only in having more pectoral fin rays (16–21 rather than 15–19); however, this difference may be a result of sample contamination by H. speculiger specimens. Variation in the mitochondrial D-loop indicated the existence of at least three unit stocks in the western central Atlantic: in the eastern Caribbean islands, southern Netherlands Antilles, and northeastern Brazil (Gomes et al. 1999).

Fishes of the Western North Atlantic

Vertebrae Precaudal

Caudal

Total

Species

Study Material. Shakhovskoy and Parin (2013b) completed a full morphological study for 56 specimens (41–238 mm SL) and a partial study for 38 specimens (132–228 mm SL) from the western Atlantic. Data from Bruun (1935) and Breder (1938) have also been included.

Danichthys rondeletii (not of Valenciennes in Cuvier and Valenciennes 1847). Bruun, 1935:73–77 (description, distribution; in part: only material from south Atlantic). Hirundichthys rondeletii (not of Valenciennes in Cuvier and Valenciennes 1847). Kovalevskaya 1972:57–64, fig. 13c (early life stages; in part: southern Pacific and Indian oceans). Kovalevskaya 1980:244–246, 269–271 (early life stages; in part: southern Pacific and Indian oceans). Heemstra and Parin 1986:394–395 (in part; short description; South Africa). Parin and Gibbs 1990:589 (in part; eastern tropical Atlantic). Belyanina 1993:142–143 (early life stages; eastern Australia). Andrew et al. 1995:14–15 (short description; Gough Island). Vera and Pequeño 2002:61–67, fig. 1 (in part; description, distribution; Chile). Guardia and Huamani 2013:163, fig. 3b (description of eggs; southern Peru). Hirundichthys rufipinnis. Parin and Belyanina 2002b:S40–S42 (description, distribution, recognition as valid species; southern subtropical waters of all oceans). Parin and Shakhovskoy 2016:2151 (description, east-central Atlantic). Gordeeva and Shakhovskoy 2017:289 (fig. 1 maximum likehood tree; DNA-barcoding).

Hirundichthys rufipinnis (Valenciennes in Cuvier and Valenciennes, 1847) Redfin Flyingfish Figures 5.20, 5.21 Tables 5.1, 5.34, 5.35, 5.36, 5.37 Exocoetus rufipinnis Valenciennes in Cuvier and Valenciennes, 1847:99 (original description; Payta, Peru; holotype MNHN B-0838). Bruun 1937:182 (description of type). Collette et al. 1997:13 (type). Exocoetus lamellifer Kner and Steindachner, 1867:384–385, pl. 2, fig. 11 (original description; 12°S, 33°W; holotype NMW 20603). Exocoetus cribrosa Kner, 1867:325–326 (original description; Sydney, Australia; holotype ?NMW 59899). Exonautes fulvipes Ogilby, 1908:8–10 (original description; Lord Howe Island; holotype AMS I.1955). Exonautes cribrosus. McCulloch 1929:106 (Australia).

Types. Types have been studied by Bruun (1935) and Parin (Collette et al. 1997; Parin and Belyanina 2002b). Exocoetus rufipinnis Valenciennes in Cuvier and 206

Family Exocoetidae

Figure 5.20. Redfin Flyingfish, Hirundichthys rufipinnis. A. 268 mm SL. From Vera and Pequeño (2002). B. Juvenile, about 93 mm SL. From Kner and Steindachner (1867).

Valenciennes, 1847. Holotype MNHN B-838 (256 mm SL); Payta, Peru. Dorsal fin rays 11; anal fin rays 10 (but recorded as 12 by Bruun [1937]); pectoral fin rays II 16. Predorsal scales 30; transverse scales between dorsal fin origin and lateral line 6½. Gill rakers 10 + 20. Vertebrae 27 + 17. Anal fin origin under second dorsal fin ray. Pectoral fins pigmented up to fourteenth ray, with narrow light margin. Pelvic fin with traces of dot pigmentation along rays. Selected

measurements: cephalo-ventral length 40.2% SL; postventral 38.7% SL; head length 21.4% SL; orbit diameter 5.9% SL; interorbital distance 7.9% SL; body depth 14.8% SL; caudal peduncle depth 5.6% SL; dorso-caudal distance 25.5% SL; pectoral fin length 72.1% SL; first pectoral ray length 29.1% SL; second pectoral ray length 43.0% SL; pelvic fin length 28.9% SL; dorsal fin height 9.9% SL. Exocoetus fulvipes Ogilby, 1908. Holotype AMS 207

Fishes of the Western North Atlantic Table 5.37. Number of gill rakers on the first arch of Hirundichthys rufipinnis and H. volador, based on Parin and Belyanina (2002b). Dashes indicate a value of zero.

section, almost flat ventrally. Depth 5.5–7.1 times in SL, body width about 1.1–1.4 times in depth. Head length 4.6–5.1 times in SL. Eye about 2.6–3.6 times in head length. Jaws subequal. Jaw teeth conspicuous, conical. Dental index about 0.09–0.16 (Parin 1961). Palatine teeth absent. Dorsal fin low, its longest ray (second or third) 6.9–12.6 times in SL. Anal fin originating slightly before or one to two rays behind dorsal fin origin. Pectoral fin 1.2–1.4 times in SL, reaching past base of dorsal fin to origin of caudal fin upper lobe, two first rays unbranched. Pelvic fins 2.7–3.5 times in SL, reaching to middle of anal fin base and farther; inserted about midway between posterior margin of opercle and origin of caudal fin base. Color. Body dark above, pale below. Dorsal and caudal fins gray, anal fin transparent. Pectoral fins black with a moderate light outer margin. Pelvic fins usually without black spot. Development. Juveniles (30–100 mm SL) differ significantly from adults in proportions and coloration. Body more prolonged (depth 6.0–8.8 times in SL). Interorbital space and eye are larger, and snout shorter; dorsal and anal fins higher, pelvic fins longer and inserted more anteriorly (see Parin and Belyanina [2002b] for details). Juveniles without barbel. Body of juveniles under 100 mm SL almost

First arch gill rakers Species Hirundichthys rufipinnis Hirundichthys volador

24 25 26 27 28 29 30 31 32 1

4 17 23 20 20 15

2

9 16 14

3

3

—

3

1

—

—

I.1955 (247 mm SL); Lord Howe Island, Tasman Sea. Dorsal fin rays 12; anal fin rays 12; pectoral fin rays II  15. Predorsal scales 28; transverse scales between dorsal fin origin and lateral line 6½. Gill rakers 8 + 18. Anal fin origin and dorsal fin origin at the same level. Selected measurements: cephalo-ventral length 40.4% SL; postventral 36.5% SL; head length 20.6% SL; orbit diameter 6.5% SL; interorbital distance 8.1% SL; body depth 16.0% SL; caudal peduncle depth 6.1% SL; dorso-caudal distance 26.3% SL; pectoral fin length 78.9% SL; first pectoral ray length 27.7% SL; second pectoral ray length 42.5% SL; pelvic fin length 33.0% SL; dorsal fin height 9.7% SL. Common Names. No official common name has been suggested for Hirundichthys rufipinnis. Shakhovskoy and Collette (in press) proposed Redfin Flyingfish. Diagnostic Characters. Hirundichthys rufipinnis differs from H. affinis and H. speculiger in having two unbranched rays at the leading edge of the pectoral fins and in pectoral fin coloration; from the latter species also in absence of palatine teeth. From H. volador it differs in absence of black blotch on pelvic fins of adults, as well as in lower number of transverse scales (usually 6½, rather than usually 7–7½); juveniles of H. rufipinnis are more uniformly pigmented. From H. rondeletii it differs in higher number of branched pectoral fin rays (15–18, usually 16–17, rather than 14–16, usually 15–16). Description. Dorsal fin rays 9–13, usually 11–12; anal fin rays 10–14, usually 11–12; pectoral fin rays II 15–18 (Table 5.34). Vertebrae 27–30 precaudal + 15–17 caudal = 44–47 total (Table 5.35). Predorsal scales 26–32, usually 27–30 (Table 5.36). Scales in transverse row from dorsal fin origin to lateral line 5½–7½, usually 6–6½ (Table 5.36). Total gill rakers on the first arch 24–32, usually 26–30 (Table 5.37). Measurements based on specimens (196–270 mm SL) from the Atlantic, Indian, and Pacific oceans. Body elongate, nearly rectangular in cross

Figure 5.21. Atlantic distribution of Redfin Flyingfish, Hirundichthys rufipinnis, and Atlantic Blackwing Flyingfish, Hirundichthys volador, based on Parin and Belyanina (2002b). Map by W. E. Bemis.

208

Family Exocoetidae uniformly colored (sometimes lightly colored with one to three weakly pronounced spots on belly or pigmented posterior part). Pectoral and pelvic fins usually uniformly dark and not mottled with dark spots and bands. Size. The maximum observed size for Hirundichthys rufipinnis is 270 mm SL (Parin and Belyanina 2002b).

Exocoetus volitans (not of Linnaeus 1758). Jordan and Meek 1885:57 (in part; Atlantic, north to New England). Jordan and Evermann 1896a:734–735 (in part; waters of North America north of the Isthmus of Panama). ?Seale 1940:9 (Gulf of Panama). Cypselurus hyperistius Fowler, 1919:12, fig. 4 (original description; St. Martin Island; holotype ANSP 7485). Cypsilurus (Exonautes) speculiger. Weber and de Beaufort 1922:181 (description; Atlantic, Indian, and Pacific oceans as far south as Australia and New Zealand). Cypselurus rubescens (not(?) of Rafinesque 1818). Fowler 1936: 426–427 (description; open Atlantic). Hirundichthys speculiger. Bruun 1935:70–73, pl. 6, figs. 2, 4 (description, Atlantic distribution). Breder 1938:84–85, fig. 43 (description, Atlantic distribution). Parin 1961:175 (osteology). McKenney 1965:47–52 (description of juveniles; western Atlantic). Boer and Nijssen 1972:151–154, fig. 1 (53°23′N, 04°07′E). Parin 1973:266 (northeastern Atlantic). Fedoryako 1980:581 (Sargasso Sea). Heemstra and Parin 1986:395 (short description; South Africa). Gorelova and Grudtsev 1987:481 (feeding, Atlantic Ocean). Nikolsky and Luschina 1990:135–136 (eastern equatorial Atlantic). Shiganova and Kovalevskaya 1991:99 (early life history stages; northeastern Atlantic). Belyanina 1994:99–102, figs. 1–4 (early life history stages, Atlantic Ocean). Oxenford et al. 1995b:39 (distribution of juveniles, eastern Caribbean). Monteiro et al. 1998:400 (northeast coast of Brazil). Parin 2003:1131 figure, map (description; west-central Atlantic). Cotton and Comyns 2006:984–985, figs. A–G (early life history stages). Fahay 2007:818–819, figs. A–G (early life history stages). Lewallen et al. 2011:164 (molecular systematics; Pacific Ocean). Shakhovskoy and Parin 2013a:117–133, figs. 1a–b, 2–5 (review). Parin and Shakhovskoy 2016:2152 (description, distribution; east-central Atlantic).

Biology Food. Hirundichthys rufipinnis feeds on zooplankton (Parin and Shakhovskoy 2016). Reproduction. Smallest known ripe male 195 mm SL (Bruun 1935). Egg morphology has been described for Hirundichthys rufipinnis by Guardia and Huamani (2013). The eggs are whitish, mean diameter is 1.8 (±0.09) mm. There are about 25 (±4) filaments at one pole of an egg and one filament at the opposite pole. In the south Indian Ocean, spawning of Hirundichthys rufipinnis probably occurs year-round; larvae and juveniles have been captured at sea surface temperatures of 19–23 °C (Kovalevskaya 1972). Range. Hirundichthys rufipinnis is an oceanic species that is circumglobal in subtropical waters of the Southern Hemisphere. It occurs in the western Atlantic from Brazil, Uruguay, and Argentina (La Plata Bay) offshore between 28°S and 40°S (Fig. 5.21). Capture of a juvenile at 12°S, 33°W (holotype of Exocoetus lamellifer) is the result of occasional drift with currents or mislabeling (see Parin and Belyanina 2002b). Geographic Variation. Local populations from the Atlantic, Indian, and Pacific oceans have never been compared. Study Material. Description based mainly on 109 specimens (33–270 mm SL; see Parin and Belyanina [2002b] for full list and localities). Data from Bruun (1935) and Vera and Pequeño (2002) have also been included.

Types. The following type material was studied by N. V. Parin and identified as Hirundichthys speculiger. Cypselurus hyperistius Fowler, 1919. Holotype ANSP 7485 (69 mm SL); St. Martin Island. Dorsal fin rays 11; anal fin rays 11; pectoral fin rays I 17. Vertebrae 46. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Mirrorwing Flyingfish (English) and volador espejo (Spanish). Diagnostic Characters. Hirundichthys speculiger differs from all other species of the genus in having well-developed teeth on the palatines. However, 5.8% (36 out of 626) of specimens studied lacked palatine teeth. Juveniles (40–100 mm SL) have a somewhat larger incidence of being edentulous (8.4%, n = 311) than larger fish (3.2%, n = 315). In the Atlantic Ocean there is a lower rate of fish without palatine teeth (2.4%, n = 122) than in those from the Pacific and Indian oceans (8.3%, n = 179, and 5.5%, n = 325, respectively). Adults of H. speculiger also differ from representatives of other Atlantic Hirundichthys species in having a large contrasting triangular crossband (so called “mirror”) of light

Hirundichthys speculiger (Valenciennes in Cuvier and Valenciennes, 1847) Mirrorwing Flyingfish Figures 5.22, 5.23 Tables 5.1, 5.38, 5.39, 5.40, 5.41 Exocoetus speculiger Valenciennes in Cuvier and Valenciennes, 1847:94 (original description; in part; Indo-Pacific; lectotype MNHN B-0822; Mauritius). Günther 1910:366–367 (Samoa, Solomon Islands). Bruun 1937:181 (redescription of type). Collette et al. 1997:13 (types). Exocoetus polleni Bleeker, 1865:130 (original description; Atlantic, 04°04′S, 32°44′W; holotype RMNH 6974).

209

Fishes of the Western North Atlantic

Figure 5.22. Mirrorwing Flyingfish, Hirundichthys speculiger. A. No. 287, 217 mm SL. Collected from 10°30′S, 7°45′W. From Bruun (1935), illustration by P. H. Winther. B. 46.0 mm SL. Collected from M/V Discoverer at 10°00′N, 44°20′W, 21 February 1969. From Belyanina (1994), illustration by T. N. Belyanina. C. 26.0 mm SL. Collected from Meteor Sta. 880, 11°00′N,

210

Family Exocoetidae

20°46′W. From Belyanina (1994), illustration by T. N. Belyanina. D. IORAS uncatalogued, 13.8 mm SL. Collected from 21°41′N, 22°16′W. From Belyanina (1994), illustration by T. N. Belyanina.

211

Fishes of the Western North Atlantic Table 5.38. Fin ray counts for Hirundichthys speculiger, based on Shakhovskoy and Parin (2013a). Dashes indicate a value of zero. Fin rays Dorsal

Anal

Pectoral

Locality

10

11

12

13

10

11

12

13

16

17

18

19

20

21

Atlantic Ocean Pacific Ocean Indian Ocean

5 10 9

65 62 51

69 28 24

1 2 —

1 1 —

20 22 30

84 64 46

35 14 9

— 13 5

6 54 36

49 24 36

50 4 7

8 — —

3 — —

Species total

24 178 121

3

2

72

194

58

18

96

109

61

8

3

Table 5.39. Vertebral counts of Hirundichthys speculiger, based on Shakhovskoy and Parin (2013a), in which vertebrae were counted without the urostyle. In this work we have included the urostyle in vertebral counts. Dashes indicate a value of zero. Vertebrae Precaudal

Caudal

Total

Locality

27

28

29

30

16

17

18

19

44

45

46

47

Atlantic Ocean Pacific Ocean Indian Ocean

3 — 6

15 21 32

13 12 9

— — 2

— 1 9

13 19 27

16 11 13

2 2 —

— — 8

7 11 39

20 19 15

6 6 3

9

68

34

2

10

59

40

4

8

57

54

15

Species total

times in dorso-caudal distance. Eye 2.9–3.5 times in head length, 0.9–1.4 times in interorbital space. Jaws subequal. Jaw teeth conspicuous, conical. Dental index 0.06–0.14 (Parin 1961). Palatine teeth usually present. Dorsal fin low, its longest ray (second or third) 9.7–14.3 times in SL. Anal fin originating slightly before or in line, and only rarely one to two rays behind dorsal fin origin. Pectoral fins 1.4–1.5 times in SL, reaching past base of dorsal fin, only first ray unbranched. Occasional specimens may have one of their pectoral fins with two unbranched rays (Bruun 1935; Shakhovskoy and Parin 2013a). Pelvic fins 3.2–4.0 times in SL, reaching to middle of anal fin base or farther; inserted slightly nearer to posterior margin of opercle than origin of caudal fin base (operculo-pelvic distance 1.0–1.4 times in pelvic-caudal distance). Color. In adults, body dark above, pale below. Dorsal and caudal fins gray; anal fin transparent; pectoral fins very dark gray with unpigmented contrast triangular crossband (“mirror”) extending to fourth to seventh ray and broad outer margin; pelvic fins light. Development. Shakhovskoy and Parin (2013a) provide a detailed report on the development, especially color changes, from juvenile to adult, including figures of color changes and descriptive details. Juveniles (40–100 mm SL) from the Atlantic

color and wide posterior light edging on the pectoral fin. Juveniles of H. speculiger differ from juveniles of H. affinis in having a broad membrane between the first and second pectoral fin rays, in the ratio of the lengths of the second and third rays of this fin, as well as in the characteristic pigmentation of the lower jaw, pectoral fins, and pelvic fins (Bruun 1935; McKenney 1965; Belyanina 1994; Shakhovskoy and Parin 2013a). Hirundichthys speculiger also differs from H. volador and H. rufipinnis in having shorter pectoral fins with only the first ray unbranched, and from the former also in absence of a black spot on pelvic fins of adults. Description. Dorsal fin rays 10–13, usually 11–12; anal fin rays 10–13, usually 12–13; pectoral fin rays I 16–20 (very rarely 15), usually I 17–18 (Table 5.38). Vertebrae 27–29 precaudal + 16–19 caudal = 44–47 total (Table 5.39). Predorsal scales 28–35, usually 30–32; scales in transverse row from dorsal fin origin to lateral line 5½–7½, usually 6½–7 (Table 5.40). Gill rakers on the first arch 5–9 upper + 15–22 lower = 22–31 total, usually 24–27 (Table 5.41). Measurements based on specimens (160–236 mm SL) from the Atlantic. Body elongate, nearly rectangular in cross section, almost flat ventrally. Depth 5.5–6.8 times in SL, body width 1.1–1.3 times in depth. Head length 3.8–4.4 times in SL, 0.8–1.1 212

Family Exocoetidae

Table 5.41. Number of gill rakers on the first arch of Hirundichthys speculiger, based on Shakhovskoy and Parin (2013a). Dashes indicate a value of zero. First arch gill rakers Locality

22

23

24

25

26

27

28

29

30

31

Atlantic Ocean Pacific Ocean Indian Ocean

2 — 1

5 8 3

10 12 11

16 24 15

27 23 27

26 17 18

6 8 13

1 4 2

— — 1

1 1 —

3

16

33

55

77

61

27

7

1

2

Species total

Ocean differ significantly from adults in proportions and coloration. Greatest body depth does not change significantly with growth (5.8–6.8 times in SL). The eye diameter and interorbital space are larger (eye 2.2–2.7 times in head length), and snout length shorter. Pectoral fins are relatively short (1.4–1.6 times in SL) in small juveniles (40–60 mm SL); then as the fish grows, fins elongate reaching maximum length (1.3–1.4 in SL) at 70–80 mm SL. Subsequently, the length of pectoral fins relative to standard length begins to decrease again as the fish continues to grow. Pelvic fins are longer and their bases are located somewhat closer to the posterior margin of the head. The membrane between the first and second pectoral fin rays is wide, up to SL about 90 mm. In juveniles (up to 120–130 mm SL); the second ray of the pectoral fin is usually much shorter than the third ray, which distinguishes Hirundichthys speculiger from H. affinis (where the second ray is usually equal to the third or a little shorter); however, this character is difficult to use for about 10% of specimens examined by Shakhovskoy and Parin (2013a), in which the second ray is only slightly shorter than the third. Juveniles without barbel. Body of specimens up to 70 mm SL, usually with one to four dark bands on ventral side. Underside of lower jaw usually not pigmented in larvae and juveniles up to about 75 mm SL. Pectoral and pelvic fins mottled to about

70 mm SL. Upper part of dorsal fin with black spot to about 100–120 mm SL. Lower lobe of caudal fin darker than upper lobe up to about 75 mm SL. Size. The maximum observed size for Hirundichthys speculiger is 251 mm SL (Bruun 1935). Biology Parasites. The population of Hirundichthys speculiger from the Atlantic Ocean is heavily infested with parasitic isopods (Cymothoidae) living in the mouth, whereas the infestation rate of populations from the Pacific and Indian oceans is negligible (Shakhovskoy and Parin 2013a). Glossobius impressus (Cymothoidae) was identified from at least one specimen of H. speculiger from the North Atlantic (Bruce and Bowman 1989). Food. Hirundichthys speculiger feeds on pteropods, amphipods, and rarely, small fishes like Vinciguerria (Gorelova and Grudtsev 1987). Larvae and juveniles (7–28 mm SL) of H. speculiger prey mainly on copepods (large Calanoida, Farranula sp., and Oncea sp.) in the equatorial Pacific (Gorelova 1980). The diet of eastern Pacific juveniles (about 119 mm fork length) of H. speculiger is primarily composed of amphipods, larval fishes, and copepods; tunicates and siphonophores are also consumed (Van Noord et al. 2013). Reproduction. Hirundichthys speculiger matures at 188 mm SL (Bruun 1935). Spawning likely occurs year-round (Bruun 1935; Kovalevskaya 1972), but 213

Fishes of the Western North Atlantic 00°45′W in the south. Occasional strays, probably drifting in the Gulf Stream, are found far northward in the Mediterranean (Collette et al. 1997, MNHN B.820) and North Sea (Boer and Nijssen 1972). See Shakhovskoy and Parin (2013a) for discussion of this finding. Hirundichthys speculiger is rare in the Caribbean, except for the eastern part, and extremely rare in the Gulf of Mexico; there is only one known capture (USNM 299261, 33 mm SL, 29°03′N, 88°02′W). Del Moral-Flores et al. (2013) listed (without any data) H. speculiger in the Veracruz reef system; however, this locality seems doubtful. Larvae of H. speculiger are concentrated in the most eastern and southeastern parts of the Caribbean Sea and in adjacent oceanic areas and are not found in the western part of the Caribbean Sea or in Gulf of Mexico (Belyanina 1994). Juveniles of H. speculiger (under 100 mm SL) were reported from the Atlantic in all seasons except summer (Shakhovskoy and Parin 2013b). Geographic Variation. Populations of Hirundichthys speculiger from the Atlantic, Pacific, and Indian oceans have virtually no appreciable differences in morphometric characters. The Atlantic population usually has 18–19 pectoral fin rays, whereas Pacific and Indian ocean fish have 17–18. Atlantic and Pacific populations have a higher mode for total vertebrae than Indian Ocean fish (45 and 44, respectively), but there is some overlap for this and all other meristic characters (see Shakhovskoy and Parin [2013a] for complete tables on meristic variation). Study Material. Shakhovskoy and Parin (2013a) recorded total morphometrics for 67 specimens (43–236 mm SL) and partial data sets for 73 specimens (50–233 mm SL) of Hirundichthys speculiger from the Atlantic. Data from Bruun (1935) have also been included.

Figure 5.23. Atlantic distribution of Mirrorwing Flyingfish, Hirundichthys speculiger, based on Shakhovskoy and Parin (2013a). Map by W. E. Bemis.

is probably more intense in spring and summer months (Shakhovskoy and Parin 2013a). Batch fecundity of a female from the Indian Ocean totaled about 2,690 eggs (Kovalevskaya 1972). Eggs are considered demersal (Parin and Gorbunova 1964; Kovalevskaya 1972, 1980). Ovarian eggs of H. speculiger from the Indian Ocean were 2.05–2.15 mm in diameter, with 7 to 10 very short thin threads at one pole and a single long (35–40 mm) and thick filament at the opposite pole (Parin and Gorbunova 1964; Kovalevskaya 1972, 1980). Munro (1954) provided different data on eggs; however, these data may be from Cheilopogon furcatus and so are not reported here. Range. Hirundichthys speculiger is an oceanic species, widely distributed in the tropical parts of the Atlantic, Pacific, and Indian oceans. However, populations are for the most part geographically isolated (Shakhovskoy and Parin 2013a). In the Atlantic, H. speculiger is found north to 37°N in the western part of the ocean (USNM 38194, 36°35′N, 74°03′W), to 25°N in the eastern part (AtlantNIRO, 24°15′N, 20°08′W, 7 October 1978), south to 30°S in the western part (AtlantNIRO, 30°02′S, 36°18′W, 15–16 February 1978) and to 08°S in the eastern part (SRTM 8024, 07°16′S, 01°23′E) (Fig. 5.23; Shakhovskoy and Parin 2013a). According to Bruun (1935), the range of this species in the eastern Atlantic is broader, 26°15′N, 20°53′W to 10°30′S,

Hirundichthys volador (Jordan, 1884) Atlantic Blackwing Flyingfish Figures 5.21, 5.24 Tables 5.1, 5.34, 5.35, 5.36, 5.37 ?Exocetus fasciatus Lesueur, 1821:9–10, pl. 4, fig. 2 (original description, Gulf Stream and in traverse from St. Croix Island to the United States; no types known). Exocoetus volador Jordan, 1884:34 (original description, Pensacola, FL; holotype USNM 34975). Collette et al. 1992:7 (holotype). Exocoetus exiliens (not of Linnaeus 1771). Goode 1876:64–67 (description of juvenile; Bermuda). Jordan and Meek 1885:54 (description; northwest Atlantic). Jordan and Evermann 1896a:732–733 (description of juvenile; western Atlantic). Exocoetus rondeletii (not of Valenciennes in Cuvier and Valenciennes 1847). Jordan and Meek 1885:55 (northwestern

214

Family Exocoetidae 41°40′N, 62°28′W, Carl Schultz; N. McPhee, 15 October 1878. Dorsal fin rays 10; anal fin rays 11; pectoral fin rays II 15/15. Gill rakers on first arch 8 + 17 = 25. Vertebrae 28 + 17 = 45. Predorsal scales 28. First anal fin ray under first dorsal fin ray; pectoral fins dark brown with a narrow, pale, posterior margin; pelvic fins with a dark blotch (Collette et al. 1992:7; Parin and Belyanina 2002b:S33–34). Cypselurus polyethmus Fowler, 1919. Holotype ANSP 7493 (141.5 mm SL); western Atlantic. Dorsal fin rays 11; anal fin rays 12; pectoral fin rays II 15–16. Predorsal scales 28. Gill rakers on first arch 9 + 21 = 30. Exonautes nonsuchae Beebe and Tee-Van, 1932. Holotype USNM 170912 (20.2 mm SL); Bermuda, St. David’s Island; 15 May 1929. Dorsal fin rays 12; anal fin rays 13; pectoral fin rays 17/17, all rays unbranched (Collette et al. 1992:8; Parin and Belyanina 2002b:S34). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Blackwing Flyingfish (English) and volador ala negra (Spanish). However, this name refers to Hirundichthys rondeletii, not to H. volador, and therefore we prefer Atlantic Blackwing Flyingfish to acknowledge recognition of the Atlantic species (Parin and Belyanina 2002b). Diagnostic Characters. Hirundichthys volador is distinct from all other species in the H. rondeletii group in having a large pigment spot on the pelvic fins in adults and nonuniform coloration of the paired fins in juveniles. Apart from coloration, Hirundichthys volador differs from all other species in the H. rondeletii group in having more scales in the oblique transverse row (6½–8½, usually 7½, rather than 5½–7½, usually 6–6½), as well as a deeper body and smaller eye diameter. Hirundichthys volador differs from H. gilberti and H. rufipinnis in having fewer rays in the pectoral fins (II 14–17, usually II 15–16 rather than II 15–18, usually II 16–17) and from the former species in having a significantly narrower light band along the posterior margin of pectoral fins. Hirundichthys volador differs from H. affinis and H. speculiger in having two unbranched rays in pectoral fins, as well as in pectoral and pelvic fins coloration. From H. speculiger this species differs also in absence of palatine teeth. Description. Dorsal fin rays 10–13, usually 11–12; anal fin rays 10–13, usually 11–12; pectoral fin rays II 14–17, usually II 15–16 (Table 5.34). Vertebrae 28–30 precaudal + 15–18 caudal = 44–47 total (Table 5.35). Predorsal scales 25–31, usually 27–29; scales

Atlantic). Jordan and Evermann 1896a:733 (short description; in part: western Atlantic). Exocoetus vinciguerrae Jordan and Meek, 1885:56 (original description (in part: specimen 126.5 mm SL from Gulf of Mexico is Hirundichthys affinis); off Newfoundland, 41°41′N, 62°28′W; holotype USNM 21870). Jordan and Evermann 1896a:734 (in part; description; northwest Atlantic). Collette et al. 1992:7 (redescription of holotype). Exonautes exsiliens (not of Linnaeus 1771). Jordan and Evermann 1898:2836 (listed). Jordan et al. 1930:200 (northwestern Atlantic). Beebe and Tee-Van 1933:65–66 (Bermuda). Exonautes rondeletii (not of Valenciennes in Cuvier and Valenciennes 1847). Jordan and Evermann 1898:2836 (listed). Jordan et al. 1930:200 (northwestern Atlantic). Beebe and Tee-Van 1933:66 (Bermuda). Exonautes vinciguerrae. Jordan and Evermann 1898:2836 (listed). Jordan et al. 1930:200 (northwest Atlantic). Cypselurus polyethmus Fowler, 1919:10 (original description; “Atlantic Ocean,” holotype ANSP 7493). Exonautes nonsuchae Beebe and Tee-Van, 1932:112 (original description; “St. David’s I., Bermuda”; holotype USNM 170912). Collette et al. 1992:8 (holotype). Danichthys rondeletii (not of Valenciennes in Cuvier and Valenciennes 1847). Bruun 1935:73, pl. 7, fig. 4 (in part; only north subtropical Atlantic). Breder 1938:85, figs. 44–45 (Gulf of Mexico and northwestern Atlantic). Duarte-Bello 1959:46 (in part; Cuba). Cypselurus exsiliens (not of Linnaeus 1847). Fowler 1936:424–425 (description of juvenile; 31°30′N, 36°36′W). Cypselurus rondeletii (not of Valenciennes in Cuvier and Valenciennes 1847). Fowler 1936:425–426 (description; Gulf of Mexico). Hirundichthys rondeletii (not of Valenciennes in Cuvier and Valenciennes 1847). McKenney 1965:32–42 (description of juveniles; western Atlantic). Fahay 1975:18 (juveniles off Florida). Hoese and Moore 1977:151 (Gulf of Mexico). Gibbs 1978: EXOC Hir 2. McEachran and Fechhelm 1998:952, figure (Gulf of Mexico). Arruda 1997:64 (listed; Azores). SmithVaniz et al. 1999:173–174 (Bermuda). Parin 2003:1130, figure, map (description, distribution; western Atlantic). Cotten and Comyns 2006:982–983, figs. D–G (in part; early life history). Gallardo-Torres et al. 2014:83 (Yucatán, Mexico, photographs). Hirundichthys volador. Parin and Belyanina 2002b:S23–S44 (description, distribution, recognition as a valid species; North Atlantic). Parin and Shakhovskoy 2016:2153 (description, east-central Atlantic). Hirundichthys rondeletii volador. Lewallen et al. 2011:164 (molecular systematics).

Types. Exocoetus volador Jordan, 1884. Holotype USNM 34975 (170 mm SL); Gulf of Mexico off Pensacola, FL. Dorsal fin rays 11; anal fin rays 11; pectoral fin rays II 16/16. Gill rakers on first arch 7 + 18 = 25. Predorsal scales 29. Vertebrae 28 + 16 = 44. Pectoral fins black, light posterior facing almost not visible; pelvic fins black in their posterior part (Collette et al. 1992:7; Parin and Belyanina 2002b:S33). Exocoetus vinciguerrae Jordan and Meek, 1885. Holotype USNM 21870 (202 mm SL); western North Atlantic in open sea south of Newfoundland, 215

Fishes of the Western North Atlantic

Figure 5.24. Atlantic Blackwing Flyingfish, Hirundichthys volador. A. USNM 198571, 200.3 mm SL. Collected at 29°40′N, 71°40′W from M/V Delaware, 28 March 1957. From Parin and Belyanina (2002b). B. USNM 198570, 191.6 mm SL. Collected at 36°57′N, 68°05′W from M/V Delaware, 23 September 1957. C. USNM 198383, 41.7 mm SL. Dipnetted at night at 29°40′N, 71°40′W from M/V Delaware. From Parin and Belyanina (2002b). D. Pectoral and pelvic fins of USNM 198370, 97.4 mm SL. Dipnetted at night by R. H. Gibbs, Jr. and B. B. Collette at 38°56′N, 66°20′W from M/V Delaware, 8 June 1957. From Parin and Belyanina (2002b). E. Pectoral and pelvic fins of USNM 198372, 59.2 mm SL. Dipnetted at night by R. H. Gibbs, Jr. at 38°55′N, 70°47′W from M/V Delaware, 29 May 1956. From Parin and Belyanina (2002b). Illustrations by H. E. Hamman.

216

Family Exocoetidae

217

Fishes of the Western North Atlantic the bases of the rays, in the middle part of the fin and near the ends of upper rays. The pelvic fins are also mottled, on light background, pigment spots are found near their bases, in the middle part and near the bases of posterior rays. In larger juveniles, pigmentation of body and fins becomes more intense and more contrasting. Six transverse bands formed by large stellate melanophores are clearly visible on the body (the last band is near the end of caudal peduncle). Dark spots with unclear shape are found over pectoral and pelvic fins and, when fins are pressed to body, conform to the position of transverse pigment bands on body. Along with these mottled specimens, there are also darkly colored specimens of similar size with dark pectoral and pelvic fins and a uniformly pigmented body. In subadult specimens, pigmentation changes to the pelagic coloration, with pectoral fins dark brown or black up to the fourteenth and fifteenth rays with gray “mirror” in the middle part, band along the outer margin practically lacking. Compared to all other species of the Hirundichthys rondeletii group, the bright dark spot occupying most of the pelvic fins is retained even in adult specimens. Dorsal and anal fins without pigment, caudal fin gray. Size. The maximum observed size for Hirundichthys volador is 236 mm SL for a specimen collected in the Gulf of Mexico (Parin and Belyanina 2002b).

in transverse row 6½–8½ (Table 5.36). Gill rakers on first arch 21–29 (Table 5.37). Measurements based on specimens 190–250 mm SL. Body elongate, nearly rectangular in cross section, almost flat ventrally. Depth 5.2–6.5 times in SL, width about 1.2 times in depth. Head length 4.2–5.2 times in SL. Eye 3.1–3.3 times in head length. Jaws subequal. Jaw teeth conspicuous, conical. Dental index about 0.09–0.16 (Parin 1961). Palatine teeth absent. Dorsal fin low, 8.3–12.3 times in SL. Anal fin originating slightly before, or one to two rays behind dorsal fin origin. Pectoral fins 1.2–1.4 times in SL, reaching (or nearly so) origin of caudal fin upper lobe, first two rays unbranched. Pelvic fins 3.0–3.6 times in SL, inserted slightly nearer to posterior margin of opercle than origin of caudal fin base. Color. Body dark above, pale below. Dorsal and caudal fins gray, anal fin transparent; pectoral fins almost uniformly deep black, except the lowermost rays and quite narrow light outer margin. Pelvic fins usually with a black spot. Development. Juveniles of Hirundichthys volador (30–100 mm SL) differ significantly from adults in proportions and coloration. Body more elongate (depth 6.0–7.5 times in SL). Inter-orbital space and eye are larger, and snout shorter; dorsal and anal fins higher, pelvic fins longer and inserted more anteriorly (see Parin and Belyanina [2002b] for details). Juveniles without barbels. Recently hatched larvae (4.8–5.0 mm SL) are darkly colored. Melanophores densely cover the entire body and are especially abundant in the yolk sac region. This intense pigmentation is usually retained even at 10–20 mm SL: body densely covered with large stellate melanophores; pigment present on dorsal fin and near the bases of anal fin rays in some specimens; pectoral and pelvic fins dark, uniformly covered with melanophores. Lightly colored specimens are also found along with such dark juveniles, with small melanophores concentrated mainly in the posterior part of the body and along the body midline, with only dark transverse spots present on the belly. Pectoral fins in such specimens are lightly colored and small melanophores are found mainly in the upper parts of fins, forming dark aggregations near the bases of the rays in the central part and near the outer margin. Pelvic fins nonuniformly pigmented with small melanophores, transparent areas are visible near the ends of rays and on membranes of posterior rays. At 30 mm SL, five distinct transverse dark spots are present on ventral surface of body and near the bases of caudal fin rays. Pectoral fins mottled, most of them lightly colored, but spots are present near

Biology Food. Hirundichthys volador feeds on zooplankton (Parin 2003; Parin and Shakhovskoy 2016). Reproduction. Eggs demersal (Parin 2003). Egg morphology has not been described for Hirundichthys volador. Eggs of other species of the H. rondeletii species group—H. rondeletii (see D’Ancona 1931; note that D’Ancona described eggs of H. rondeletii as Cheilopogon heterurus and vice versa), H. rufipinnis (see Guardia and Huamani 2013), and H. gilberti (see Kovalevskaya 1980)—have a single thick, long filament at one pole and about 15–30 thin filaments at the opposite pole. Probably spawns in coastal areas of the Gulf of Mexico and in the western part of the Sargasso Sea (Parin and Belyanina 2002b). Range. The Hirundichthys rondeletii complex from subtropical waters of all oceans of the northern and southern hemispheres was divided into four separate species (H. rondeletii, H. volador, H. gilberti, and H. rufipinnis) by Parin and Belyanina (2002b). Hirundichthys volador is found mainly in 218

Family Exocoetidae beyond middle part of dorsal fin base. Pelvic fins moderately long (4.5– 5.5 times in SL); with third ray the longest, inserted nearer posterior margin of opercle than caudal fin base and nearer anal fin origin than pectoral fin insertion. Dorsal fin with a large black spot, very high, with middle rays the longest. Anal fin originates before second ray of dorsal fin. Eggs demersal, with filaments. Juveniles with paired chin barbels (Parexocoetus brachypterus and P. hillianus) or without barbels (P. mento). Species. T h ree spec ies a re recog n i zed: Parexocoetus mento and P. brachypterus from the Indo-Pacific, and P. hillianus from the Atlantic. Parexocoetus mento has been considered to have Indo-Pacific and Atlantic subspecies. However, validity of P. mento atlanticus is doubtful. This subspecies is known only from two type specimens (holotype BMNH 1924.8.29.13 and paratype BMNH 1924.8.29.14), reported from west of Cap Blanc and near the Cape Verde Islands, and not differing in any character from Indo-Pacific specimens. Absence of additional records from the well-investigated area off northwestern Africa is hardly understandable if the type specimens were correctly labeled (Parin and Gibbs 1990). The Atlantic species Parexocoetus hillianus (Gosse) has been considered as subspecies of P. brachypterus since Bruun (1935). However, Parin (2003) regarded it as a different species. Range. Pelagic in nearshore and neritic waters, rare in open oceans; found in the Atlantic, Indian, and Pacific oceans.

the northwestern Atlantic, including the Gulf of Mexico (Fig. 5.21). Juveniles are widely dispersed by the Gulf Stream. The northern limit of the range during the warmest parts of the year is Woods Hole, MA, in the United States; there is one specimen (MCZ 39727) from Scarborough, ME, Gulf of Maine. According to Eschmeyer et al. (2018), the species occurs up to 46°N, 61°W (holotype of Exocoetus vinciguerrae). Parin and Belyanina (2002b) also report several records from the eastern Atlantic. To the east of 30°W, records are extremely rare and it is possible that some of these specimens belong to the Mediterranean species H. rondeletii. Study Material. Description mainly based on 73 specimens (34–236 mm SL; see Parin and Belyanina [2002b] for full list and localities). Data from Bruun (1935) and Breder (1938) have also been included. Genus Parexocoetus Bleeker, 1866 Parexocoetus Bleeker, 1866:126 (type species Exocoetus mento Valenciennes by monotypy).

Diagnosis. Parexocoetus differs from other genera of flyingfishes in the western Atlantic in its rather compressed body, moderately long pectoral fins not reaching close to end of dorsal fin base, high dorsal fin with middle rays the longest, well-developed branch of lateral line ascending to pectoral fin base, and presence of vomerine teeth. Parexocoetus shares these characters with Fodiator, but differs in having a blunt and short (rather than pointed and long) snout, protrusible upper jaw, and pectoral fins reaching to or beyond the middle part of dorsal fin base (rather than before middle part of dorsal fin base). Description. Cross-sectional diameter of body elongate-oval, not flattened ventrally. Greatest body depth 4.3–6.5 times in SL, width of body in its depth 1.3–1.7 times. Vertebrae 21–25 + 14–16 = 35–41. Predorsal scales 16–25; transverse scales 4½–5½. Pectoral branch of lateral line present. Swim bladder extends into haemal canal, reaching the tenth to fourteenth caudal vertebrae. Head 3.7–4.7 times in SL. Snout blunt, shorter than eye. Jaws subequal (lower jaw a little extended in Parexocoetus mento), upper jaw protrusible. Jaw teeth small (dental index 0.05–0.15), conical, without supplementary cusps. Teeth present on palatines, vomer, entopterygoids, and tongue. Gill rakers on first arch 23–33. Dorsal fin rays 9–14; anal fin rays 10–14; dorsal rays minus anal rays –2 to –1; pectoral fin rays I 11–14. Pectoral fins moderately long (1.7–2.1 times in SL); extending to or

Parexocoetus hillianus (Gosse, 1851) Atlantic Sailfin Flyingfish Figures 5.25, 5.26 Tables 5.1, 5.42, 5.43, 5.44, 5.45 Exocoetus hillianus Gosse, 1851:11, pl. I, fig. 1 (original description, Jamaica; no types known). Parexocoetus mesogaster (not of Bloch 1795). Jordan and Meek 1885:47 (in part; Gulf Stream to Newport). Jordan and Evermann 1896a:728–729 (description; in part: western Atlantic). Evermann and Marsh 1900:104 (description; Puerto Rico). Fowler 1917:116–118, fig. 1 (description; Newport, RI). Fowler 1919:10 (Newport, RI; West Indies). Breder 1929a:281, fig. 4. Breder 1929b:295. Hildebrand and Cable 1930:445–449, figs. 60–63 (Beaufort, NC, spawning and development). Jordan et al. 1930:200 (listed). Breder 1932:20–22, pl. 8 (biology, larvae, and juveniles; Florida). Bruun 1933:380, fig. 1. (vertebral counts). Breder and Nichols 1934:42 (vertebral counts). Duarte-Bello 1959:47 (listed). Parexocoetus brachypterus hillianus. Bruun 1935:22–26, pl. 1, fig. 3 (description, western Atlantic distribution). Breder 1938:17– 21, fig. 9 (description, Caribbean Sea to 40°N). Fowler 1944:95,

219

Fishes of the Western North Atlantic

Figure 5.25. Atlantic Sailfin Flyingfish, Parexocoetus hillianus. A. USNM 198377, 107.9 mm SL. B. USNM 198377, 73.5 mm SL. C. USNM 198377, 23.3 mm SL. All three specimens dipnetted at night by R. H. Gibbs, Jr. at 20°50′N, 93°00′W from M/V Oregon, Sta. 1605, 25 September 1956. Illustrations by H. E. Hamman.

220

Family Exocoetidae

146–147, fig. 119 (Courtown Keys, St. Andrews Island). Poll 1953:181–184, fig. 76 (description; eastern Atlantic). Lewis 1961:255–266 (food, growth and reproduction; Barbados). Caldwell and Caldwell 1964:14 (Caribbean Sea, waters of Colombia). Cervigón 1966:222 (Venezuela). Kovalevskaya 1967:493–494 (juveniles; Atlantic). Mago Leccia 1970:89 (listed). Smith-Vaniz et al. 1999:174 (Bermuda). Parexocoetus brachypterus littoralis Breder, 1938:22–28, figs. 10–12 (original description, Caribbean Sea to 40°N; comparison with P. b. hillianus; holotype YPM ICH 001567). Duarte-Bello 1959:47 (listed, Cuba). Parexocoetus brachypterus (not of Richardson 1846). Sauskan 1973:151–152 (eastern Atlantic). Fahay 1975:18 (juveniles off Florida). Hoese and Moore 1977:151 (Gulf of Mexico). Gibbs 1978: EXOC Par 1 (western Atlantic). Fedoryako 1980:581 (Sargasso Sea). Hunte et al. 1995:28–35 (eggs and larvae distribution; eastern Caribbean). Oxenford et al. 1995a:16–21 (distribution of adults; eastern Caribbean). Oxenford et al. 1995b:42–44 (distribution of juveniles; eastern Caribbean). McEachran and Fechhelm 1998:953, figure (Gulf of Mexico). Monteiro et al. 1998:400–401 (distribution; northeastern Brazil). Collette et al. 2003:102 (Navassa Island, West Indies). Stevens et al. 2003:71–76 (spawning in Gulf of Mexico). Cotten and Comyns 2006:962–963, figs. A, C, F (in part; early life history). Casazza and Ross 2008:352–359 (associated with Sargassum, Gulf Stream off North Carolina). Parexocoetus hillianus. Günther 1866:284 (description; West Indies). Parin 2003:1132 (description, west-central Atlantic distribution). Fahay 2007:820–821, figs. E–I (early life history stages). Lewallen et al. 2011 (molecular systematics). Parin and Shakhovskoy 2016:2154 (description, east-central Atlantic).

differs from the related Indo-Pacific species P. brachypterus in having a lower dorsal fin, fewer vertebrae (modally 39 rather than 40) and more gill rakers (usually 28–30, x̅ 29.2, rather than usually 25–28, x̅ 26.5). Description. Dorsal fin rays 11–14; anal fin rays 12–15; pectoral I 11–13 (Table 5.42). Vertebrae 22–24 + 15–16 = 37–40, usually 39 (Table 5.43). Predorsal scales 19–25, usually 20–22; scales in transverse row from dorsal fin origin to lateral line 4½–5½ (Table 5.44). Gill rakers on the first arch 7–9 + 19–24 = 26–34, usually 28–30 (Table 5.45) Measurements based on western Atlantic specimens 90–126 mm SL. Body elongate (greatest depth 4.3–6.0 times in SL); elliptical in cross section, somewhat compressed (greatest width 1.3–1.7 times in depth). Head short (3.9–4.9 times in SL); much shorter than dorso-caudal distance. Eye diameter 2.9–3.7 times in head length, usually equal to interorbital length. Snout blunt, shorter than eye. Jaws subequal, with small, conical teeth. Dental index about 0.07. Dorsal fin very high (2.4–3.1 times in SL); reaching beyond the base of caudal fin; the longest dorsal ray (fifth or sixth) 3.8–4.5 times in SL. Anal fin origin before second ray of dorsal fin. Pectoral

Types. Parexocoetus brachypterus littoralis Breder, 1938. Holotype YPM ICH 001567 (126 mm SL). Dorsal fin rays 12; anal fin rays 13. Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Sailfin Flyingfish (English); we have modified the name to Atlantic Sailfin Flyingfish to recognize this species distribution in the Atlantic. It is called volador aletón in Mexico (Page et al. 2013) and is locally known as petite volant in Dominica, which means “small flyingfish” in English (Oxenford et al. 2007). Diagnostic Characters. Parexocoetus hillianus 221

Fishes of the Western North Atlantic

fins 1.7–2.1 times in SL, reaching to dorsal fin ray 5 to 9, first ray unbranched. Pelvic fins inserted much nearer posterior margin of opercle than caudal fin base (operculo-pelvic distance 1.4–1.5 times in pelvic-caudal distance), short (4.5–5.5 times in SL); barely or not reaching anal fin origin. Color. In adults, body dark above, pale below, the dark color usually iridescent bluish green in life. Dorsal fin with a large black blotch distally, all other fins transparent except the very upper portion of pectorals, which is gray. Breder (1932) reported sexual dimorphism in coloration: in males sides of body as well as pelvic and caudal fins are suffused with rose. Development. Juveniles (20–70 mm SL) are similar to adults in general appearance, but differ in some proportions (body more slender, head longer, interorbit narrower, pectoral fins shorter, dorsal fin higher, etc.). Paired chin barbels may be present in specimens 20–107 mm SL, easily lost, highly variable in size, their length up to 2 times in head at 25 mm SL, about equal to head at 80–90 mm SL, and 1.7–4.6 times in head at 95–105 mm SL. In coloration juveniles resemble adults, except pelvic and anal fins bearing much black pigment; barbels black. Size. The maximum observed size for Parexocoetus

hillianus is 126 mm SL (holotype of P. brachypterus littoralis) in the western Atlantic, 134 mm SL in eastern Atlantic (IORAS, no number, Africa, off Conakry, 16 November 1963). Biology Predators. Parexocoetus hillianus is eaten by many predatory fishes and also provides a very important food item for many tropical marine birds (such as terns and noddies). Historically, in the Florida Keys P. hillianus made up about one-third of all fishes captured by birds nesting there (Breder 1932). Parasites. Manter (1947, 1954) noted and described two species of digenetic trematodes of the family Zoogonidae from the intestine of specimens of Tortugas flyingfish identified as Parexocoetus mesogaster (which is probably P. hillianus): Steganoderma parexocoeti and Brachyenteron parexocoeti. It is interesting that five of the eight species in the genus Steganoderma are known from beloniform hosts (four from Belonidae, one from Hemiramphidae, and one from Exocoetidae). Food. Breder (1932) suggested that Parexocoetus hillianus feeds almost entirely on planktonic copepods in the waters of Florida. Lewis (1961) reported that it feeds on plankton, predominantly copepods 222

Family Exocoetidae and other crustaceans, and (rarely) fishes in the Barbados. Reproduction. Parexocoetus hillianus reaches full size in one year, maturing at about 110–130 mm SL (Lewis 1961). Off the Florida Keys and North Carolina, spawning occurs during the summer from at least the middle of May to the middle of July (Hildebrand and Cable 1930; Breder 1932). Spawning probably occurs off Barbados in two pulses: one from September to January, and the other from March to August (Lewis 1961; Khokiattiwong 1988; Oxenford et al. 1995b). Ripe females and males range from 93–126 mm SL and from 88–117 mm SL, respectively, at Tortugas. Eggs are demersal (1.3–1.4 mm in diameter) with many filaments over the surface (Breder 1938). A spawning aggregation was observed in the northeastern Gulf of Mexico over the outer continental shelf in May 2001 just after moonrise two days after the full moon (Stevens et al. 2003). More than a million flyingfish were estimated to be part of this aggregation. Parexocoetus hillianus was observed flying, jumping, and swimming more actively than usual near the surface, extruding ripe sperm and eggs. Male flyingfish were three times more abundant than females (based on dipnetting captures) and there were observations of several males simultaneously pursuing a single female. No flotsam was present in the area, suggesting these fish may not rely on attaching their eggs to floating objects. Range. Parexocoetus hillianus is a neritic species known from both sides of the Atlantic Ocean. In the western Atlantic it is distributed from 40°N in the Gulf Stream south through the Gulf of Mexico and Caribbean Sea to Brazil (Fig. 5.26). The southernmost occurrence is at 20°49′–21°00′S, 34°46′– 48′W (IORAS, no number, 108, 112, 118 mm SL, R/V Akademic Ioffe Cruise 16, 4–5 November 2004). It is very common in the Caribbean Sea and Lesser Antilles area, less common in the Gulf of Mexico, the Gulf Stream, and the western Sargasso Sea. In the Caribbean, adults were found to be most abundant east of Barbados (Oxenford et al. 1995a). A separate eastern Atlantic population (maybe another subspecies) occurs along the west coast of Africa between 12°N and 11°S (Parin and Gibbs 1990). Some specimens probably drifted with the Angola Current far offshore (Fig. 5.26): MCZ 156682 (12 specimens, 19–93 mm SL, R/V Atlantis II Cruise 60, 16°01′S, 02°00′E, 31 May 1971); MCZ 156680 (4 specimens, 61–66 mm SL, R/V Atlantis II Cruise 60, 15°59′S, 02°02′E, 31 May 1971); MCZ 156679 (3 specimens, 48–99 mm SL, R/V Atlantis II Cruise 60,

Figure 5.26. Distribution of Atlantic Sailfin Flyingfish, Parexocoetus hillianus, based on specimens identified by N. V. Parin and I. B. Shakhovskoy. Map by W. E. Bemis.

15°25′S, 03°07′E, 1 June 1971). The related species, P. brachypterus, occurs in the tropical Indo-Pacific. Geographic Variation. The eastern Atlantic population differs from that of the western Atlantic Ocean in having more predorsal scales (usually 21–24 rather than usually 20–22), in having pelvic fins inserted more anteriorly (operculo-pelvic distance 1.5–1.7 times in pelvic-caudal distance) and, as a result, usually not reaching anal fin origin. Chin barbels are absent in all eastern Atlantic specimens more than 60 mm SL. Study Material. Description according to Bruun (1935), Breder (1938), Lewis (1961), and Parin (2003). Additional material includes meristics of 223 western and eastern Atlantic specimens (28–134 mm SL) examined by N. V. Parin, V. A. Khachaturov, G. N. Pokhilskaya, T. N. Steyker, and I. B. Shakhovskoy. From the western Atlantic, 113 specimens (48.5– 126 mm SL) in IORAS, USNM, and ZMH. From the eastern Atlantic, 110 specimens (28.3– 134 mm SL) in IORAS, USNM, ZMH, and BCF. Genus Prognichthys Breder, 1928 Prognichthys Breder, 1928:20 (type species Exocoetus gibbifrons Valenciennes by original designation).

Diagnosis. Prognichthys is a member of the subfamily Cypselurinae characterized by long pectoral 223

Fishes of the Western North Atlantic fins reaching the end of the dorsal fin base and elongate pelvic fins having the third ray the longest and inserted nearer to anal fin origin than to pectoral fin insertion. Prognichthys differs from three other Cypselurinae genera: from Cypselurus and Cheilopogon in having two to four unbranched upper pectoral fin rays and in the absence of chin barbels in juveniles, and from Hirundichthys in having the anal fin origin three rays or more behind the origin of the dorsal fin. Description. Dorsal fin rays 10–13; anal fin rays 8–11; pectoral fin rays II–IV 12–17. Vertebrae 27–30 + 13–16 = 41–45. Predorsal scales 19–29; transverse scales 6½–8½. Gill rakers on first arch 21–31. Cross-sectional diameter of body nearly rectangular in adults, almost flat ventrally. Greatest body depth 5.1–6.3 times in SL, width of body 1.1–1.3 in its depth. Pectoral branch of lateral line absent. Swim bladder extends into haemal canal, reaching middle of caudal peduncle. Head 3.6–4.4 times in SL. Snout blunt, shorter than eye. Lower jaw a little shorter than the upper and included beneath the upper jaw with mouth closed; upper jaw not protrusible. Jaw teeth medium-sized (dental index 0.08–0.17), conical or with one to two additional cusps. Palatine teeth absent. Pectoral fins long (1.4–1.5 times in SL); extended to end of dorsal fin base or farther posteriorly. Pelvic fins long (2.8–3.4 times in SL); with third ray the longest, inserted nearer to hind margin of opercle than to caudal fin origin. Dorsal fin uniformly gray, low, anterior rays the longest. Anal fin originates three to seven rays behind origin of dorsal fin. Eggs probably pelagic, with evenly spaced, rather short filaments. Juveniles robust, without barbels. Species. Of the six species recognized, two, Prognichthys brevipinnis and P. sealei, are Indo-Pacific; one species, P. tringa, is eastern tropical Pacific; and three, P. glaphyrae, P. occidentalis, and P. gibbifrons, are Atlantic Ocean endemics. Indo-Pacific species, which differ from Atlantic ones (and P. tringa) in having three or four rather than two unbranched pectoral fin rays in adults, are also known from the Atlantic by single captures: one specimen of P. brevipinnis (161 mm SL) captured at 33°14′S, 44°30′W (holotype of Exocoetus chloropterus) and one specimen of P. sealei captured at 19°45′S, 17°40′W (see Parin [1999] for details). Food. Amphipods were the predominant food item of fishes of the genus Prognichthys caught in the Atlantic from 20°N to 30°S and 40°W to 6°E (Gorelova and Grudtsev 1987). Prognichthys sealei and P. tringa from the Pacific fed primarily on copepods and amphipods (Van Noord et al. 2013).

Range. Prognichthys contains some neritic and some oceanic species from tropical Atlantic and Indo-Pacific (Parin 1999; Shakhovskoy and Collette, in press). Prognichthys glaphyrae Parin, 1999 Oceanic Bluntnose Flyingfish Figures 5.27, 5.28 Tables 5.1, 5.46, 5.47, 5.48, 5.49 Exocoetus sp. Lütken 1876:407 (description; Atlantic; in part: two specimens, 89–126 mm SL examined by Bruun [1935] and Parin [1999]). Prognichthys gibbifrons (not of Valenciennes in Cuvier and Valenciennes 1847). Bruun 1935:64–67, pl. 7, fig. 3 (in part; central Atlantic). John 1976:125, 128 (in part; larvae and juveniles in central Atlantic). Gibbs 1978: EXOC Prog 1 (in part; figure). Shiganova and Kovalevskaya 1991:95–97, fig. 2 (larvae and juveniles in central Atlantic). Prognichthys glaphyrae Parin, 1999:295–297, fig. 1 (original description, 16°50′S, 26°40′W; holotype ZIN 51826). SmithVaniz et al. 1999:174 (Bermuda). Parin 2003:1133, figure, map (description, western Atlantic). Cotten and Comyns 2006:986–987, fig. A–F (in part(?); early life history). Parin and Shakhovskoy 2016:2156 (description, east-central Atlantic). Gordeeva and Shakhovskoy 2017:289 (fig. 1 maximum likehood tree; DNA-barcoding).

Types. Prog nichthys glaphyrae Parin, 1999. Holotype ZIN 51826 (201 mm SL); 16°50′S, 26°40′W. Dorsal fin rays 13; anal fin rays 10; pectoral fin rays II 16. Predorsal scales 24; transverse scales between dorsal fin origin and lateral line 7½. Gill rakers 5  + 19 = 24. Vertebrae 27 + 16 = 43. Head length 25.8% SL, eye diameter 8.4% SL; postorbital space 11.0% SL. Paratypes 26 specimens (42–185 mm SL) listed by Parin (1999). Common Names. The common name Oceanic Bluntnose Flyingfish was suggested by Parin (2003). Diagnostic Characters. Prognichthys glaphyrae is one of the four species of Prognichthys with two unbranched pectoral fin rays. It differs from both the other Atlantic species, P. gibbifrons and P. occidentalis, and from P. tringa of the eastern tropical Pacific, in having a larger eye (1.2–1.5 rather than 1.4–1.8 times in postorbital part of head), more pectoral fin rays (usually II 16 rather than usually II 14 or 15), and less pigmented pectoral fins (chiefly pale with central part gray rather than chiefly blackish with only distal tip and lowermost portion pale). In juveniles, the pectoral fins are highly contrastingly pigmented, predominantly clear with a large black area posterior-ventrally. Description. Dorsal fin rays 11–13; anal fin rays 8–11, usually 10; pectoral fin rays II 14–17, usually II 16 (Table 5.46). Vertebrae 27–29 + 14–16 = 42–44 (Table 5.47). Predorsal scales 20–25, usually 21–22; 224

Family Exocoetidae Table 5.46. Fin ray counts for Prognichthys glaphyrae and P. occidentalis, based on Parin (1999). Dashes indicate a value of zero. Fin rays Dorsal

Anal

Pectoral

Species

10

11

12

13

8

9

10

11

15

16

17

18

19

Prognichthys glaphyrae Prognichthys occidentalis

— 1

1 39

37 91

42 11

1 15

20 111

57 16

2 —

— 4

9 47

25 38

39 2

7 1

Table 5.47. Vertebral counts of Prognichthys glaphyrae and P. occidentalis, based on Parin (1999). Vertebrae Species Prognichthys glaphyrae Prognichthys occidentalis

Precaudal

Caudal

Total

27 28 29

14 15 16

42 43 44

12

6

2

3 11

6

10 18

2

1 17

2

3 16

1

6 20

4

Table 5.48. Number of predorsal and transverse scales of Prognichthys glaphyrae and P. occidentalis, based on Parin (1999). Scales

Species Prognichthys glaphyrae Prognichthys occidentalis

Predorsal

Transverse

20 21 22 23 24 25

6-6½ 7-7½ 8-8½

5 22 27 14 2

7

1

2

33

8

6 21 23 13

9

3

42

6

Table 5.49. Number of gill rakers on first arch of Prognichthys glaphyrae and P. occidentalis, based on Parin (1999). Table 5.49. Number Dashes indicate a valueofof gill zero.rakers on first arch of Prognichthys glaphyrae and P. occidentalis, based on Parin (1999). Dashes indicate aFirst value ofgill zero. arch rakers Species

20 21First 22 arch 23 gill 24 rakers 25 26 27

Prognichthys glaphyrae Species Prognichthys occidentalis Prognichthys glaphyrae Prognichthys occidentalis

— 21 — 22 3 23 7 24 21 25 26 26 12 27 4 20 5 4 9 22 16 9 5 — — — 3 7 21 26 12 4 5 4 9 22 16 9 5 —

scales in transverse row 6–8 (Table 5.48). Gill rakers on the first arch 4–7 + 15–20 = 22–27, usually 24–25 (Table 5.49). Measurements based on 24 specimens 150–201 mm SL: body elongate (greatest depth 5.1–6.4 times in SL); nearly rectangular in cross section, almost flat ventrally, slightly compressed (width 1.1–1.2 times in depth). Head moderate (3.6–4.0 times in 225

SL); equal to or slightly less than dorso-caudal distance (0.95–1.15 times in the latter). Eye diameter 2.8–3.2 times in head length and 1.2–1.5 times in postorbital part of head, nearly equal to interorbital space (0.9–1.1 in the latter). Lower jaw a little shorter than the upper and included beneath upper jaw when mouth closed. Jaw teeth small, unicuspid; dental index 0.13–0.14. Palatine teeth absent. Dorsal fin low, its longest ray (second or third) 8.3–10.9 times in SL. Anal fin originates under third to fifth dorsal fin ray. Pectoral fins 1.4–1.5 times in SL, two upper rays unbranched; reaching from penultimate dorsal fin ray to middle of caudal peduncle. Pelvic fins inserted nearer posterior margin of opercle than origin of lower caudal lobe (operculo-pelvic distance 1.1–1.4 times in pelvic-caudal distance), extremely long (2.8–3.3 times in SL); reaching from penultimate anal fin ray to middle of caudal peduncle. Color. In adults, body dark above, pale below; the dark color iridescent blue, pale color silvery in life. Dorsal and caudal fins gray; anal fin transparent; pectoral fins gray in central part, but their pointed tips for about one-quarter of fin length, posterior margin and lower portion transparent; pelvic fins mostly gray, especially in the middle (probably both pectoral and pelvic fins are greenish in life). Development. Juvenile Prognichthys glaphyrae (14–128 mm SL) lack barbels. Smaller juveniles (20–60 mm SL) are characteristically robust and bluntsnouted. They have a deep body (up to 4.7 times in SL); large head (to 3.5 times in SL) and eye (to 2.0 times in head), and pectoral and pelvic fins of comparable length (1.5–2.1 and 2.3–2.8 times in SL). At 20–30 mm SL the body is heavily pigmented, especially before the pelvic insertion, the paired fins are mostly black except for the gray transverse band at the middle of pectoral fin and the pale distal portion. In larger juveniles (30–130 mm SL); the pectoral fins are very contrastingly pigmented being mostly pale with a black area at the base and a second, more extensive, black area in the outer part posteriorly.

Fishes of the Western North Atlantic

Figure 5.27. Oceanic Bluntnose Flyingfish, Prognichthys glaphyrae. A. Holotype, ZIN 51826, 201 mm SL. Collected at 16°50′S, 26°40′W. B. IORAS uncatalogued, 62.6 mm SL. Collected at 10°22′S, 27°18′W from R/V Petr Lebedev, 8 November 1964. C. IORAS uncatalogued, 27.2 mm SL. Collected at 12°56′N, 52°56′W from R/V Dmitrii Mendeleev, Sta. 54. D. IORAS

226

Family Exocoetidae

uncatalogued, 11.0 mm SL. Collected at 12°56′N, 52°56′W from R/V Dmitrii Mendeleev, Sta. 54. Illustrations from Parin (1999), drawn by G. N. Pokhilskaya.

227

Fishes of the Western North Atlantic Study Material. A total of 89 specimens (14–201 mm SL) from collections of AtlantNIRO, BAH, INBYuM, IORAS, ISH, USNM, ZIN, ZMMU, and ZMUC (listed in Parin [1999]). Prognichthys occidentalis Parin, 1999 Western Bluntnose Flyingfish Figures 5.28, 5.29 Tables 5.1, 5.46, 5.47, 5.48, 5.49 Exocoetus gibbifrons (not of Valenciennes in Cuvier and Valenciennes 1847). Jordan and Meek 1885:65–66 (description; Newport, RI). Jordan and Evermann 1896a:741 (short description; western Atlantic). Cypsilurus gibbifrons (not of Valenciennes in Cuvier and Valenciennes 1847). Jordan and Evermann 1896b:323 (listed; Newport, RI). Cypselurus gibbifrons (not of Valenciennes in Cuvier and Valenciennes 1847). Fowler 1917:119–120, fig. 3 (description; Newport, RI). Jordan et al. 1930:202 (listed). Prognichthys gibbifrons (not of Valenciennes in Cuvier and Valenciennes 1847). Breder 1928:20–22 (in part; comparison with Prognichthys tringa). Bruun 1935:64–67, pl. 7, fig. 1 (description; in part: four specimens 161–175 mm SL from the Antilles and Brazil). Breder 1938:74–78, figs. 37–38 (in part; description of juveniles from Caribbean, Gulf of Mexico, and near Bahamas). Rogers 1938:45, fig. 1 (Chatham, MA). Fowler 1944:72, 460, fig. 121 (Serranilla Bank, Bahamas). Poll 1949:256 (San Salvador). Duarte-Bello 1959:47 (listed; Cuba). McKenney 1965:52–56 (description of juveniles; western Atlantic) Belyanina 1975:140 (larvae and juveniles in Caribbean and Gulf of Mexico). Fahay 1975:18 (larvae and juveniles in the Gulf Stream from Florida to North Carolina). John 1976:125, 128, Abb. 9 (in part; juveniles from South America, 30°–38°S). Hoese and Moore 1977:151 (Gulf of Mexico). Gibbs 1978: EXOC Prog 1 (in part; west-central Atlantic). McEachran and Fechhelm 1998:954, figure (Gulf of Mexico). Monteiro et al. 1998:401 (distribution; northeastern Brazil). ?Lewallen et al. 2011 (molecular systematics). ?Prognichthys glaphyrae (not of Parin 1999). Lewallen et al. 2011 (molecular systematics). Prognichthys occidentalis Parin, 1999:297–301, fig. 3 (original description, Gulf of Mexico, 22°30′N, 91°24′W; holotype ZIN 51829). Smith-Vaniz et al. 1999:174–175 (Bermuda). Collette et al. 2003:102 (Navassa Island, West Indies). Parin 2003:1134, figure, map (description, western Atlantic). Cotten and Comyns 2006:988–989, figs. A–H (early life history). Fahay 2007:822–823, figs. a–e (early life history stages). Casazza and Ross 2008:352, 355 (associated with Sargassum, Gulf Stream off North Carolina). Lewallen et al. 2011 (molecular systematics). Randall et al. 2015:97–109 (distribution of larvae; northern Gulf of Mexico). Gordeeva and Shakhovskoy 2017:289 (fig. 1 maximum likehood tree; DNA-barcoding).

Figure 5.28. Distribution of Oceanic Bluntnose Flyingfish, Prognichthys glaphyrae, and Western Bluntnose Flyingfish, Prognichthys occidentalis, based on Parin (1999). Map by W. E. Bemis.

Size. The maximum observed size of Prognichthys glaphyrae is 201 mm SL. Biology Food. Prognichthys glaphyrae feeds on zooplankton (Parin 2003). Reproduction. The eggs of Prognichthys glaphyrae are probably pelagic with many filaments over the entire surface. Fish are sexually mature at lengths of about 170 mm SL, and Bruun (1935) reported females with ripe eggs from August. Larvae have been captured from April to May at sea surface temperatures of 22.9–24.3 °C (Shiganova and Kovalevskaya 1991). Relationship to Humans. Of no importance in fisheries. Range. Prognichthys glaphyrae is an Atlantic tropical oceanic species mostly restricted to cores of central oceanic gyres (Fig. 5.28). The northernmost records are from 30°N to 32°N (paratypes USNM 249164, 257623, and ZMUC 24765). The southernmost record is from 20°S, 26°30′W (IORAS, no number). The lack of records from the equatorial zone between 06°N and 6°S suggests possible isolation between northern and southern Atlantic populations; however, this possibility is not based on sufficient material.

Types. Prognichthys occidentalis Parin, 1999. Holotype ZIN 51829 (female, 179 mm SL); R/V Akademik Kurchatov, 14th cruise, Sta. 1290, 22°30′N, 91°24′W; 7 April 1973. Dorsal fin rays 12; anal fin rays 9; pectoral fin rays II 14. Predorsal scales 22; 228

Family Exocoetidae gray; anal fin transparent; pectoral fins brownish with pale distal tip and lowermost portion, extension of which is highly variable (from being limited by two or three lower membranes to reaching seventh ray at the base of fin and ninth ray at its hind margin), pelvic fins usually blackish except the innermost membrane. Development. Juvenile Prognichthys occidentalis (13–138 mm SL) lack barbels. Allometric growth is similar to that of P. glaphyrae (body depth up to 4.6 times in SL; head length up to 3.4 times in SL; eye up to 1.9 times in head; pectoral fin length down to 1.9 times in SL; and pelvic fin length up to 2.3 times in SL in juveniles 20–60 mm SL). Juveniles of these species differ slightly only in postorbital distance to eye diameter ratio starting with length 50–60 mm SL upward (see Parin 1999: fig. 5). In fish 13–25 mm SL, the intensity of pigmentation differs greatly, but in most individuals the body is densely pigmented. Specimens shorter than 60 mm SL usually have black paired fins with small pale patches near the distal tips of upper pectoral rays, but in some juveniles the fins are not so heavyily and uniformly pigmented. In most specimens 60–80 mm SL, much lighter oblique or semicircular cross band is well developed in central part of pectoral fin and traces of this band can be seen even in juvenile specimens up to 100–120 mm SL. Size. The maximum observed size for Prognichthys occidentalis is 190 mm SL from a Gulf of Mexico specimen.

transverse scales 7. Gill rakers 6 + 18 = 24. Vertebrae 28 + 15 = 43. Head length 25.6% SL; eye diameter 7.3% SL; postorbital length 12.5% SL. Paratypes 19 specimens (80–177 mm SL) listed by Parin (1999). Common Names. The Common and Scientific Names of Fishes from the United States, Canada, and Mexico (Page et al. 2013) lists Bluntnose Flyingfish (English) and volador chato (Spanish). Parin (2003) refers to the species as the Western Bluntnose Flyingfish. Diagnostic Characters. Prognichthys occidentalis is most similar to the eastern Atlantic P. gibbifrons from which it differs in having fewer predorsal scales (20–25, usually 22–23, rather than 22–28, usually 24–26) and to the eastern tropical Pacific P. tringa, from which it differs in having more gill rakers (20–26, usually 22–25, rather than 18–22). Prognichthys occidentalis differs from P. glaphyrae in having a smaller eye (1.4–1.8 rather than 1.2–1.5 times in postorbital part of head), fewer pectoral fin rays (usually II 14–15 rather than usually II 16), and more darkly pigmented pectoral fins. Description. Dorsal fin rays 10–13; anal fin rays 8–10, usually 9; pectoral fin rays II 13–17, usually II 14–15 (Table 5.46). Vertebrae 27–29 + 14–16 = 42–44 (Table 5.47). Predorsal scales 20–25, usually 22–23; scales in transverse row 6½–8, usually 7½ (Table 5.48). Gill rakers 4–7 + 15–20 = 20–26, usually 23 or 24 (Table 5.49). Measurements based on 32 specimens (147–179 mm SL). Body elongate (greatest depth 5.2–6.7 times in SL); slightly compressed (width 1.1–1.2 times in depth). Head moderate (3.7–4.1 times in SL); equal to or slightly less than dorso-caudal distance (0.9– 1.1 times in the latter). Eye diameter 3.0–3.6 times in head length and 1.4–1.7 times in postorbital part of head, less than interorbital space (1.1–1.3 times in the latter). Lower jaw a little shorter than the upper and included beneath upper jaw when mouth closed. Jaw teeth small, mostly conical, dental index 0.12– 0.14. Palatine teeth absent. Dorsal fin low, its longest ray (second or third) 8.2–10.9 times in SL. Anal fin originates under fourth to sixth dorsal fin ray. Pectoral fins 1.4–1.6 times in SL, reaching from end of dorsal fin base to middle of caudal peduncle, two upper rays unbranched. Pelvic fins inserted nearer posterior margin of opercle than origin of lower caudal lobe (operculo-pelvic distance 1.1–1.4 times in pelvic-caudal distance), very long (2.8–3.4 times in SL); reaching from middle of anal fin base to slightly beyond its end. Color. In adult Prognichthys occidentalis, the body is dark above, pale below; the dark color iridescent blue, pale color silvery in life. Dorsal and caudal fins

Biology Food. Prognichthys occidentalis feeds on zooplankton (Parin 2003). Reproduction. Larvae of Prognichthys occidentalis were captured in June and July in the northern Gulf of Mexico (Randall et al. 2015). There were significant monthly and interannual changes in the densities of flyingfish larvae. Range. Prognichthys occidentalis is a neritic species that avoids the open sea. It is distributed along the Atlantic coast of the Americas; abundant in the Caribbean Sea, Gulf of Mexico, and near the Bahamas (Parin 1999; Fig. 5.28). Over three years, 9,533 larval P. occidentalis were collected from the outer shelf and slope waters of the Gulf of Mexico; this represented 77% of the flyingfishes collected in the area (Randall et al. 2015). South American records are known from of f Guyana, Brazil (Fortaleza, Rio de Janeiro, and Porto Alegre), and Argentina (south to 40°S). The northernmost and southernmost records result from passive transport 229

Fishes of the Western North Atlantic

Figure 5.29. Western Bluntnose Flyingfish, Prognichthys occidentalis. A. Holotype ZIN 51829, 179 mm SL. Collected at 22°30′N, 91°24′W. From Parin (1999), illustration by T. N. Belyanina and A. N. Kotlyar. B. IORAS uncatalogued, 67.2 mm SL. Collected at 19°40′N, 76°42′W. From Parin (1999), illustration by G. N. Pokhilskaya. C. USNM 198371, 51.5 mm SL.

230

Family Exocoetidae

Dipnetted at night by R. H. Gibbs, Jr. and B. B. Collette at 40°00′N, 68°45′W from M/V Delaware. Illustration by H. E. Hamman. D. USNM 198382, 21.4 mm SL. Dipnetted at night at 38°55′N, 70°25′W from R/V Atlantis, 18 December 1955. Illustration by H. E. Hamman.

231

Fishes of the Western North Atlantic of juveniles by the Gulf Stream (to about 40°N, with occasional juvenile strays reaching Massachusetts) and the Brazil Current. Study Material. Description based on 153 specimens (13–179 mm SL) from collections of AtlantNIRO, IORAS, USNM, ZIN, ZMH, ZMMU, and ZMUC (listed in Parin [1999]).

York Zoological Society 12(9):83–88. [Issue dated March 22, 1931.] Beebe, W. and J. Tee-Van. 1932. New Bermuda fish, including six new species and forty-three species hitherto unrecorded from Bermuda. Zoologica: Scientific Contributions of the New York Zoological Society 13(5):109–120. ——— . 1933. Field Book of the Shore Fishes of Bermuda. New York: G. P. Putnam’s Sons. 337 pp. Belyanina, T. N. 1975. Preliminary results of the study of ichthyoplankton of the Caribbean Sea and the Gulf of Mexico. Trudy Instituta Okeanologii 100:125–146. [In Russian with English abstract.] ——— . 1993. Early stages of development of the East Australian flying fishes (family Exocoetidae). Trudy Instituta Okeanologii 128:108–146. [In Russian with English abstract.] ——— . 1994. Early stages of ontogeny of Hirundichthys speculiger and H. affinis (Exocoetidae) from the Atlantic Ocean. Voprosy Ikhtiologii 34(1):98–107. [In Russian, English translation in Journal of Ichthyology 34(5):60–87.] Bennett, E. T. 1831. Characters of new genera and species of fishes of the Atlantic coast of northern Africa presented by Captain Belcher, R.N. Proceedings of the Zoological Society of London 1830–1831:145–148. Bennett, F. D. 1840. Narrative of a Whaling Voyage Round the Globe, from the Year 1833 to 1836, Volume 2. London: R. Bentley. 395 pp. Ben Souissi, J., D. Golani, H. Mejri, and C. Capapé. 2005. On the occurrence of Cheilopogon furcatus in the Mediterranean Sea. Journal of Fish Biology 67(4):1144–1149. Bigelow, H. B. and W. C. Schroeder. 1953. Fishes of the Gulf of Maine. Fishery Bulletin of the Fish and Wildlife Service 53:1– 577. Bleeker, P. 1865. Description d’une espèce inédite d’Exocet découvent par M. François Pollen. Nederlandsch Tijdschrift voor de Dierkunde 3:130–133. ——— . 1866. Sur les especes d’exocet de l’Inde Archipélagique. Nederlandsch Tijdschrift voor de Dierkunde 3:105–129. ——— . 1866–1872. Exocoetiformes. In P. Bleeker. Atlas ichthyologique des Indes Orientales Néêrlandaises, Volume 6. Amsterdam: Frédéric Muller. pp. 67–78. Bloch, M. E. 1795. Naturgeschichte der ausländischen Fische. Berlin: Im Verlage der Morinoschen Kunsthandlung. Allgemeine Naturgeschichte der Fische, Theil 9. 192 pp., [Atlas 4], pls. 397–429. Boer, P. and H. Nijssen. 1972. The occurrence of the flying fish, Hirundichthys speculiger (Valenciennes, 1847), in the North Sea (Pisces, Atheriniformes, Exocoetidae). Bulletin Zoologisch Museum 2(15):151–154. Breder, C. M., Jr. 1927. Scientific results of the first oceanographic expedition of the “Pawnee” 1925. Article 1, Fishes. Bulletin of the Bingham Oceanographic Collection l(l):1–90. ——— . 1928. Scientific results of the second oceanographic expedition of the “Pawnee.” Bulletin of the Bingham Oceanographic Collection 2(2):1–25. ——— . 1929a. Report on synentognath habits and development. In Carnegie Institution of Washington. Year Book No. 28: July 1, 1928 to June 30, 1929. Washington, DC: Carnegie Institution. pp. 279–282. ——— . 1929b. Field observations on flyingfishes; a suggestion of methods. Zoologica: Scientific Contributions of the New York Zoological Society 9(7):295–312. ——— . 1932. On the habits and development of certain Atlantic Synentognathi. Publication 435, Papers from the Tortugas Laboratory of the Carnegie Institution of Washington 28(1): 1–35.

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Index Principal references are given in parentheses. References to illustrations (f), tables (t), and maps (m) are in italics.

A

Ablennes, 5, 6, (8–12), 42, 91 hians, 4f, 6, 7, (8–12), 8t, 10f, 11m, 12t, 47 hians hians, 10 acanthocephalans Exocoetidae, 150 Hyporhamphus mexicanus, 117, 118 Strongylura hubbsi, 19 Strongylura marina, 24, 28 Tylosurus acus acus, 43, 46 acus Belone, 43 Strongylura, 43 Tylosurus, 4f, 6, 7, 36, 41, (42–48), 46t, 48m, 50, 52–53, 54 acus acus Tylosurus, 8t, (42–48), 45f, 46t, 54 acus imperialis Tylosurus, 44–45, 46t, 48 acus melanotus Tylosurus, 44, 46t, 47 acus rafale Tylosurus, 44–45, 46t, 47, 48 acutillus Scomberesox, 80 acutus Exocoetus, 150 Fodiator, 150, 151 adocoetus Cololabis, 79 affinis Exocoetus, 199, 202 Hirundichthys, 150, 152, 154t, 158, (199–206), 200– 201f, 203t, 204t, 205m, 208, 212, 213, 215 agoo Cheilopogon, 152 agujon, 44 albidactylus Exocoetus, 153 alienus Cypselurus, 188, 191 almeida Belone, 21, 25 Strongylura, 39 Tylosurus, 22, 37 altipennis Exocoetus, 182 altipinna Belone, 44

altipinne Belone, 42 amphipods, 28, 29 anastomella Strongylura, 4f, 18 angusticeps Cypselurus, 186 antarei Cypselurus, 187, 188 appendiculatus Exocoetus, 186, 188 archipelagicus Hemiramphus, 99 Ardeapiscis, 98 ardeola Belone, 13 Strongylura, 13 Tylosurus, 13 ardeola ardeola Belone, 15 ardeolus Tylosurus, 13 argalus Belona, 12 Belone, 13 Platybelone, 4f, 7, (12–18), 15m, 16t, 17t, 47 Strongylura, 13 argalus annobonensis Platybelone, 16t, 17, 17t argalus argalus Platybelone, 8t, (12–18), 13f, 15, 16t, 17t, 42 argalus lovii Platybelone, 16t, 17, 17t argalus platyura Platybelone, 16t, 17t argalus pterura Platybelone, 16t, 17, 17t argalus spp. Platybelone, 16t, 17t argalus trachura Platybelone, 15, 16t, 17, 17t Arrhamphus, 127 sclerolepis krefftii, 91 atherinoides Chriodorus, 92, (93–96), 93f, 94t, 95m, 95t, 114, 129 Atlantic Agujon, (42–48), 45f, 46t, 48m, 50 Atlantic Bigwing Halfbeaks, 92, 94t, (127–130), 128f, 129m Atlantic Blackwing Flyingfish, 150, 152, 154t, 202, 206t, 208m, 208t, 212, (214–219), 216–217f

Fishes of the Western North Atlantic marina, 39 maris-rubri maris-rubri, 49 marisrubri marisrubri, 49 melanochira, 48, 50 notata, 31, 32, 34 raphidoma, 48, 49, 50 scrutator, 21, 25 subtruncata, 37, 39 timucu, 21, 37 trachura, 13 truncata, 21, 25 Belonidae, (5–7), 8t, 149 Ablennes, (8–12) Key, 7 Platybelone, (12–18) Strongylura, (18–42) Tylosurus, (42–54) Beloniformes, (1–4), 2f, 3f, 20m, 86m Belonidae, (5–54), 8t, 149 Exocoetidae, (149–232) Hemiramphidae, (89–130) Key, 4 Scomberesocidae, (79–86) Belonion, 5, 6, 7 Bennett's Flyingfish, 150, 151, 152, 154t, (180–186), 181f, 183t, 185m, 188 Bermuda Ballyhoo, 91, 92, 94t, 99, 101, 101t, (103–104), 104m, 104f, 107 Bermuda Halfbeak, 103 bermudensis Hemiramphus, 91, 92, 94t, 99, 101, 101t, (103–104), 104m, 104f, 107 bicolor Exocoetus, 160 Bigwing Halfbeaks, (127–130) Billfish, 15, 25 See also Sailfish Blackfin Tuna, 15, 40, 102, 129 Black Skimmer, 26 Blackwing Flyingfish, 215 See also Atlantic Blackwing Flyingfish Blotchwing Flyingfish, 150, 152, 154t, 165, 169, (170– 176), 172–173f, 174t, 175t, 177m, 188 Bluefish, 107, 116 Blue Marlin, 11, 40, 102, 105, 107 Blue Shark, 81, 82 Booby, 106, 107 Bottlenose Dolphin, 45 brachypterus Parexocoetus, 219, 221 brachypterus hillianus Parexocoetus, 219, 221 brachypterus littoralis Parexocoetus, 221, 222 branchiurans Strongylura marina, 23, 28 Strongylura notata forsythia, 36 Tylosurus crocodilus crocodilus, 52 brasiliensis Esox, 105 Haemiramphus, 107 Hemiramphus, 90–91, 92, 94t, 99, 101, 101t, 102, 103, 104m, (105–109), 108f braziliensis Hemiraphas, 112

Atlantic Bluefin Tuna, 26 Atlantic Dwarf Saury, 79, 81t, (83–86), 84t, 85f, 86m Atlantic Flyingfish, 150, 152, 154t, 165, 170, 174t, 175t, (176–180), 177m, 178–179f, 187, 188, 190 Atlantic Needlefish, 4t, 6, 7, 8t, 18, 19, (21–30), 22f, 26t, 27t, 30m, 39, 54 Atlantic Ribbon Halfbeaks, 92, 94t, (96–98), 97f, 98m Atlantic Sailfin Flyingfish, 150, 151, 154t, (219–223), 220f, 221t, 222t, 223m Atlantic Saury, 4f, 26, 30, 79, 79–80, 80, (81–83), 81t, 82f, 83m, 84t Atlantic Silverstripe Halfbeak, 90, 92, 94t, 109, 112, 113, 113m, 115, 115t, 116t, 117, 117t, 118, 119, 120, 123, (124–127), 125f

B

bahiensis Cypselurus, 153 Exocoetus, 153, 186, 188 Balao, 91, 92, 94t, (99–103), 100f, 101t, 102m, 107, 109 balao Hemiramphus, 91, 92, 94t, (99–103), 100f, 101t, 102m, 107, 109 Bald Eagle, 26 Ballyhoos, 90–91, (98–109) Bandwing Flyingfish, 150, 152, 154t, 156, (160–165), 161f, 163t, 164m, 185 Belona argalus, 12 caribbaeus, 50 carribea, 42, 44 crocodila, 48, 50 timucu, 37 truncata, 21 Belone, 5, 12 acus, 43 almeida, 21, 25 altipinna, 44 altipinne, 42 ardeola, 13 ardeola ardeola, 15 argalus, 13 belone, 4f, 6 caribaea, 42 caribbea, 42 cigonella, 37, 39 countermani, 6 crassa, 48, 50 depressa, 13, 37, 39 diplotaenia, 37, 38, 39 galeata, 21, 25 gerania, 48, 50 hians, 8, 9, 12 houttuyni, 39 imperialis, 43 imperialis imperialis, 43 jonesi, 42 jonesii, 44 Jonesii Goode, 42, 48 Jonesii Günther, 42, 48 latimana, 42, 44 latimanus, 42 longirostris, 21 maculata, 8, 10

242

Index 180), 177m, 178–179f, 187, 188, 190 milleri, 150 nigricans, 150, 185 pinnatibarbatus, 150, 151, 152, 154t, (180–186), 181f, 183t, 185m, 188 pinnatibarbatus altipennis, 185 pinnatibarbatus californicus, 185 pinnatibarbatus fernandizianus, 185 pinnatibarbatus japonicus, 185 pinnatibarbatus melanocercus, 185 pinnatibarbatus pinnatibarbatus, 185 chloropterus Exocoetus, 224 choram Tylosurus, 42, 47, 50 Chriodorus, 91, (93–96), 127 atherinoides, 92, (93–96), 93f, 94t, 95m, 95t, 114, 129 miocaenicus, 93 cigonella Belone, 37, 39 Clearwing Flyingfish, 150, 151, 154t, 176, (186–191), 187f, 188t, 189t, 190m Cobitopsis, 93 collettei Dermogenys, 112 Hyporhamphus, 90, 94t, 103, (112), 113m, 115, 115t, 116t, 117t, 118, 119, 126 Collette’s Bermuda Halfbeak, 90, 92, 94t, 103, (112), 113m, 115, 115t, 116t, 117t, 118, 119, 126 Cololabis adocoetus, 79 saira, 4f, 79, 80 comatus Cheilopogon, 176 Cypselurus, 150, 151, 154t, 176, (186–191), 187f, 188t, 189t, 190m Cypsilurus, 187 Exocoetus, 186, 188 convexus Oxyporhamphus, 99, 127, 129 copepods Cheilopogon cyanopterus, 156 Cheilopogon pinnatibarbatus, 184 Exocoetus obtusirostris, 195 Exocoetus volitans, 198 Hirundichthys affinis, 202 Hirundichthys speculiger, 213 Oxyporhamphus micropterus similis, 129 Parexocoetus hillianus, 222 Platybelone argalus argalus, 15 Strongylura hubbsi, 19 Strongylura marina, 21, 22, 23, 26, 28 Strongylura notata forsythia, 34, 36 Strongylura notata notata, 33 Strongylura timucu, 37, 38, 40–41 Tylosurus acus acus, 42, 43, 44, 45–46 Tylosurus crocodilus crocodilus, 51 Cory’s Shearwater, 82 countermani Belone, 6 crassa Belone, 48, 50 crassus Tylosurus, 48

brederi Hyporhamphus, 20m, 90, 94t, (110–111), 111f, 111t Breder’s Halfbeak, 20m, 90, 94t, (110–111), 111f, 111t breederi Ichthyacus, 109, 110 brevipinnis Prognichthys, 224 brevoortii Euleptorhamphus, 96, 97 browni Hemiramphus, 107, 109 Brownii Hemiramphus, 106, 107 Brown Noddies Ablennes hians, 11 Euleptorhamphus velox, 98 Hemiramphus balao, 99, 102 Hemiramphus brasiliensis, 106, 107 Oxyporhamphus micropterus similis, 128, 129

C

Camperi Scomberesox, 81 camperii Scomberesox, 81 caribaea Belone, 42 Strongylura, 43 caribaeus Tylosurus, 43 caribbaeus Belona, 50 Tylosurus, 42, 50 caribbea Belone, 42 Strongylura, 43 carribea Belona, 42, 44 Cero, 11, 107 cestodes Ablennes hians, 11 Exocoetidae, 150 Hyporhamphus meeki, 116 Strongylura marina, 28 Strongylura notata forsythia, 35, 36 Strongylura timucu, 41 Tylosurus acus acus, 42, 47 Tylosurus crocodilus crocodilus, 48, 52 Cheilopogon, 150, (152–186), 224 agoo, 152 comatus, 176 cyanopterus, 150, 151, 152, (153–159), 154t, 155f, 157t, 158m, 185, 188 exsiliens, 150, 152, 154t, 156, (160–165), 161f, 163t, 164m, 185 furcatus, (165–170), 169t, 170t, 176, 188, 214 furcatus fisunovi, 169–170, 169t, 170t furcatus furcatus, 150, 152, 153, 154t, (165–170), 166–167f, 168m, 169t, 170t furcatus neimani, 169t, 170, 170t heterurus, 150, 152, 154t, 165, 169, (170–176), 172–173f, 174t, 175t, 177m, 188 luetkeni, 176 melanurus, 150, 152, 154t, 165, 170, 174t, 175t, (176–

243

Fishes of the Western North Atlantic

D

cribrosa Exocoetus, 206 cribrosus Exonautes, 206 croccodilus Tylosurus, 50 crocodila Belona, 48, 50 crocodilus Tylosurus, 4f, 6, 7, 36, 41, 42, 44, 47, (48–54), 53t, 54m crocodilus crocodilus Tylosurus, 8t, 47, 51f, 53t crocodilus fodiator Tylosurus, 53t crocodylus Tylosurus, 50 crustaceans Exocoetidae, 150 cuspidatus Hemiramphus, 127 cyanopterus Cheilopogon, 150, 151, 152, (153–159), 154t, 155f, 157t, 158m, 185, 188 Cypselurus, 153, 160 Exocoetus, 153 Cypselurus, 150, 152, (186–191), 224 alienus, 188, 191 angusticeps, 186 antarei, 187, 188 bahiensis, 153 comatus, 150, 151, 154t, 176, (186–191), 187f, 188t, 189t, 190m cyanopterus, 153, 160 exsiliens, 160, 215 furcatus, 165 gibbifrons, 228 heterurus, 170, 176 hewitti, 153 hiraii, 186 holubi, 193 hyperistius, 209 lineatus, 182 luetkeni, 176 lutkeni, 176 melanurus, 176 minos, 182 monroei, 153, 158 naresii, 186, 188 nigricans, 160, 162 opisthopus, 186 pinnatibarbatus, 160, 182 polyethmus, 215 rondeletii, 215 rubescens, 209 smithi, 160 vitropinna, 186, 188, 191 Cypsilurus comatus, 187 furcatus, 165 gibbifrons, 228 heterurus, 170, 176, 182 lineatus, 182 lutkeni, 176 speculiger, 209

Danichthys, 199 rondeletii, 202, 206, 215 depauperatus Hemiramphus, 99 depressa Belone, 13, 37, 39 Dermogenys collettei, 112 digenetic trematodes Hyporhamphus unifasciatus, 126 Parexocoetus hillianus, 222 Platybelone argalus argalus, 15 Strongylura marina, 28 Strongylura notata forsythia, 35 Strongylura timucu, 41 Tylosurus acus acus, 46–47 Tylosurus crocodilus crocodilus, 52 diplotaenia Belone, 37, 38, 39 Strongylura, 39 Tylosurus, 37 Diptera, 29 Dolphinfish Ablennes hians, 11 Euleptorhamphus velox, 98 Exocoetus obtusirostris, 195 Exocoetus volitans, 198 Hemiramphus balao, 102 Hemiramphus brasiliensis, 106, 107 Hirundichthys affinis, 202 Strongylura timucu, 26 dussumieri Exocoetus, 153 Dwarf Atlantic Saury, 79, 81t, (83–86), 84t, 85f, 86m

E

El Petén Halfbeak, 20, 20m, 90, 92, 93, 94t, 95, (117– 119) equirostrum Scomberesox, 81 Esox belone var. Houttuyni, 21 belone var. marinus, 21 brasiliensis, 81, 105 longirostris, 21, 25, 29 marinus, 21, 25 saurus, 81 timucu, 37, 39 Euleptorhamphus, 90, (96–98) brevoortii, 96, 97 longirostris, 97 velox, 92, 94t, (96–98), 97f, 98m viridis, 96, 97 euryops Strongylura, 39 Tylosurus, 37, 39 Euthynnus alletteratus See Little Tunny evolans Exocoetus, 193, 196 Halocypselus, 193, 196 Evolantia, 127

244

Index microptera, 127 exilis Strongylura, 4f, 18, 26, 29 Exocetus fasciatus, 214 Exocoethus exsiliens, 160 Exocoetidae, 3f, (149–232) Cheilopogon, 150, (152–186), 224 Cypselurus, 150, 152, (186–191), 224 Exocoetus, 150, 152, (191–199) Hirundichthys, 150, 152, 186, (199–219), 224 Key, 4, 151–152 Parexocoetus, 150, 152, 199, (219–223) Prognichthys, 150, 152, 186, (223–232) Exocoetoidea Exocoetidae, (149–232) Hemiramphidae, (89–130) Zenarchopteridae, 1, 89 Exocoetus, 150, 152, (191–199) acutus, 150 affinis, 199, 202 albidactylus, 153 altipennis, 182 appendiculatus, 186, 188 bahiensis, 153, 186, 188 bicolor, 160 chloropterus, 224 comatus, 186, 188 cribrosa, 206 cyanopterus, 153 dussumieri, 153 evolans, 193, 196 exsiliens, 160, 170, 214 fulvipes, 207–208 furcatus, 165, 176, 182 gaussianus, 193 georgianus, 191 gibbifrons, 228 gibbosus, 191, 193 heterurus, 170, 176, 186 hillianus, 219 holubii, 193 lamellifer, 206, 209 lineatus, 182, 185 longipinnis, 182 lutkeni, 176 maculipinnis, 170 melanurus, 176 micropterus, 127 monocirrhus, 191, 193 nigricans, 160 nuttallii, 165 obtusirostris, 150, 151, 154t, (191–196), 192f, 194t, 195m, 195t orbignianus, 182, 184 parrae, 153 peruvianus, 191, 193 pinnatibarbatus, 180 polleni, 209 procne, 170 pulchellus, 180 roberti, 199, 202

robustus, 176 rondeletii, 170, 214–215 rufipinnis, 199, 202, 206–207 scylla, 199, 202 speculiger, 209 spilopus, 160, 162 vermiculatus, 153 vinciguerrae, 199, 202, 215, 219 volador, 214, 215 volitans, 150, 151, 154t, 170, 191, 193, 194t, 195t, (196–199), 197f, 198m, 209 Exocoetus sp., 224 Exonautes cribrosus, 206 exsiliens, 160, 215 fulvipes, 206 nonsuchae, 215 rondeletii, 215 speculiger, 209 vinciguerrae, 215 exsiliens Cheilopogon, 150, 152, 154t, 156, (160–165), 161f, 163t, 164m, 185 Cypselurus, 160, 215 Exocoethus, 160 Exocoetus, 160, 170, 214 Exonautes, 160, 215

F

False Silverstripe Halfbeak, 115 far Hemiramphus, 99 Farhians, 98 fasciatus Exocetus, 214 Hemiramphus, 124, 126, 127 filamentosus Hemiramphus, 107, 109 Flat Needelfish, 4f, 6, 7, (8–12), 8t, 10f, 11m, 12t, 47 fluviatilis Strongylura, 4f, 18 Flying Halfbeaks, 97 See also Atlantic Ribbon Halfbeaks Fodiator, 149, 150, 219 acutus, 150, 151 Fourwing Flyingfish, 150, 152, 154t, 158, (199–206), 200–201f, 203t, 204t, 205m, 208, 212, 213, 215 fulvipes Exocoetus, 207–208 Exonautes, 206 Fundulus, 28 furcatus Cheilopogon, (165–170), 169t, 170t, 176, 188, 214 Cypselurus, 165 Cypsilurus, 165 Exocoetus, 165, 176, 182 furcatus fisunovi Cheilopogon, 169–170, 169t, 170t furcatus furcatus Cheilopogon, 150, 152, 153, 154t, (165–170), 166–167f, 168m, 169t, 170t furcatus neimani Cheilopogon, 169t, 170, 170t

245

Fishes of the Western North Atlantic

G

far, 91, 98 fasciatus, 124, 126, 127 filamentosus, 107, 109 guineënsis, 100, 101, 102 longirostris, 97 lutkei, 91, 99 macrochirus, 100, 101, 102 macrorhynchus, 97 marginatus, 91, 99, 106 pleei, 107 Pleii, 100, 103 pleii, 101, 102, 103 poeyi, 124 Richardi, 124, 125–126 richardi, 127 Roberti, 119 roberti, 113, 119, 125 robustus, 98 sajori, 91 saltator, 99, 101, 107 unifasciatus, 113, 124, 125, 127 vittatus, 100, 102 welsbyi, 98 Hemiraphas braziliensis, 112 Hemirhamphus unifasciatus, 113 Hemirhamphus sp., 103, 113, 114 heterurus Cheilopogon, 150, 152, 154t, 165, 169, (170–176), 172–173f, 174t, 175t, 177m, 188 Cypselurus, 170, 176 Cypsilurus, 170, 176, 182 Exocoetus, 170, 176, 186 hewitti Cypselurus, 153 hians Ablennes, 4f, 6, 7, (8–12), 8t, 10f, 11m, 12t, 47 Belone, 8, 9, 12 Tylosurus, 8–9 hians hians Ablennes, 10 hildebrandi Hyporhamphus, 90, 112, 114, 122, 123, 125 hillianus Exocoetus, 219 Parexocoetus, 150, 151, 154t, (219–223), 220f, 221t, 222t, 223m Hiporhamphis unifasciatus, 125 hiraii Cypselurus, 186 Hirundichthys, 150, 152, 186, (199–219), 224 affinis, 150, 152, 154t, 158, (199–206), 200–201f, 203t, 204t, 205m, 208, 212, 213, 215 gilberti, 215, 218 rondeletii, 150, 175, 185, 206, 208, 215, 218, 219 rondeletii volador, 215 rufipinnis, 150, 152, 154t, 202, (206–209), 206t, 207f, 208m, 208t, 212, 215, 218 speculiger, 150, 152, 154t, 185, 199, 202, 205, 208, (209–214), 210–211f, 212t, 213t, 214m volador, 150, 152, 154t, 202, 206t, 208m, 208t, 212, (214–219), 216–217f

galeata Belone, 21, 25 gaussianus Exocoetus, 193 gavialoides Tylosurus, 4f, 6, 42, 47 georgianus Exocoetus, 191 gerania Belone, 48, 50 Giant Flyingfish, 182 See also Bennett’s Flyingfish gibbifrons Cypselurus, 228 Cypsilurus, 228 Exocoetus, 228 Prognichthys, 150, 224, 228, 229 gibbosus Exocoetus, 191, 193 gilberti Hirundichthys, 215, 218 gladius Tylosurus, 48, 50 glaphyrae Prognichthys, 150, 151, 154t, (224–228), 225t, 226–227f, 228, 228m, 229 Great Barracuda, 11 Great Hammerhead, 184 Green Gar, 25 guineënsis Hemiramphus, 100, 101, 102

H

Haemiramphus brasiliensis, 107 Halocypselus, 191 evolans, 193, 196 mesogaster, 196 obtusirostris, 193 Hardhead Halfbeak, 92, (93–96), 93f, 94t, 95m, 95t, 114, 129 Helminths, 112 Hemiramphidae, 3f, (89–130), 94t Chriodorus, (93–96) Euleptorhamphus, (96–98) Hemiramphus, (98–109), 101t Hyporhamphus, (109–127) Key, 4, 92 Oxyporhamphus, (127–130) Hemiramphus, 92, 96, (98–109), 101t archipelagicus, 99 ardelio, 91 australis, 91 balao, 91, 92, 94t, (99–103), 100f, 101t, 102m, 107, 109 bermudensis, 91, 92, 94t, 99, 101, 101t, (103–104), 104m, 104f, 107 brasiliensis, 90–91, 92, 94t, 99, 101, 101t, 102, 103, 104m, (105–109), 108f browni, 107, 109 Brownii, 106, 107 cuspidatus, 127 depauperatus, 99

246

Index Hyporhamphus collettei, 112 Hyporhamphus meeki, 114, 116 Hyporhamphus unifasciatus, 124, 126 Oxyporhamphus micropterus similis, 129 Platybelone argalus argalus, 15 Strongylura marina, 23, 24, 28 Strongylura notata forsythia, 35, 36 Tylosurus acus acus, 46 Tylosurus crocodilus crocodilus, 51–52

holubi Cypselurus, 193 holubii Exocoetus, 193 Houndfish, 8t, 44, 47, (48–54), 51f, 53t, 53t, 54m houttuyni Belone, 39 hubbsi Strongylura, 4f, 5, 7, 8t, (18–21), 20m, 20f, 26, 29 hyperistius Cypselurus, 209 Hyporamphus unifasciatus, 125 Hyporhamphus, 90, 92, 98, (109–127) aff. unifasciatus, 125 australis, 91 brederi, 20m, 90, 94t, (110–111), 111f, 111t collettei, 90, 92, 94t, 103, (112), 113m, 115, 115t, 116t, 117t, 118, 119, 126 georgii, 91 hildebrandi, 90, 112, 114, 122, 123, 125 kronei, 119, 120 limbatus, 90 meeki, 90, 92, 94t, 95, 112, 113m, (113–117), 115f, 115t, 116t, 117t, 118, 119, 120, 123, 124, 126, 127 meeki (nomen nudum), 114 melanochir, 91 mexicanus, 20, 20m, 90, 92, 93, 94t, 95, (117–119) naos, 90, 127 picarti, 90 quoyi, 91, 113 regularis ardelio, 91 roberti, 90, 92, 112, 113, 117, 118, (119), 120, 121t, 122, 122t, 123m, 125, 126, 127 roberti hildebrandi, 92, 94t, 119, 120, 121t, (122–124), 122f, 122t, 123m roberti roberti, 92, 94t, (119–122), 121t, 122t, 123, 123m sajori, 91 salvatoris, 119, 120 tricuspidatus, 109, 124, 126, 127 unifasciatus, 90, 92, 94t, 109, 112, 113, 113m, 115, 115t, 116t, 117, 117t, 118, 119, 120, 123, (124–127), 125f Hyporhampus sp., 114

J

Jaegers, 116 jonesi Belone, 42 jonesii Belone, 44 Jonesii Goode Belone, 42, 48 Jonesii Günther Belone, 42, 48

K

Kingfish, 35 King Mackerel, 26, 102, 107, 116 krefftii Strongylura, 4f, 18 kronei Hyporhamphus, 119, 120

L

Ladyfish, 36 lamellifer Exocoetus, 206, 209 latimana Belone, 42, 44 latimanus Belone, 42 Tylosurus, 42 Lecithaster gibbosus, 43 leeches Hemiramphus brasiliensis, 105, 108 Strongylura marina, 23, 28 leiura Strongylura, 4f, 18 lineatus Cypselurus, 182 Cypsilurus, 182 Exocoetus, 182, 185 Little Tunny Euleptorhamphus velox, 98 Hemiramphus balao, 102 Hemiramphus brasiliensis, 105, 107 Strongylura timucu, 26, 40 Tylosurus acus acus, 43, 45 Tylosurus crocodilus crocodilus, 49, 51 longipinnis Exocoetus, 182 longirostris Belone, 21 Esox, 21, 25, 29 Euleptorhamphus, 97 Hemiramphus, 97

I

Ichthyacus, 109 breederi, 109, 110 imperialis Belone, 43 Tylosurus, 44 imperialis imperialis Belone, 43 incisa Strongylura, 4f, 18 isopods Ablennes hians, 11 Cheilopogon heterurus, 175 Cypselurus comatus, 190 Hemiramphus bermudensis, 103–104 Hemiramphus brasiliensis, 105–106, 107, 108 Hirundichthys affinis, 202 Hirundichthys speculiger, 213

247

Fishes of the Western North Atlantic Tylosurus, 21 longleyi Strongylura, 6, 13 Long-nosed Gar, 23, 26 luetkeni Cheilopogon, 176 Cypselurus, 176 lutkei Hemiramphus, 99 lutkeni Cypselurus, 176 Cypsilurus, 176 Exocoetus, 176

Mexican Halfbeak, 118 See also El Petén Halfbeak mexicanus Hyporhamphus, 20, 20m, 90, 93, 94t, 95, (117–119) microptera Evolantia, 127 micropterus Exocoetus, 127 Oxyporhamphus, 127, 128, 129 micropterus micropterus Oxyporhamphus, 128 micropterus similis Oxyporhamphus, 92, 94t, (127–130), 128f, 129m milleri Cheilopogon, 150 minos Cypselurus, 182 miocaenicus Chriodorus, 93 Mirrorwing Flyingfish, 150, 152, 154t, 185, 199, 202, 205, 208, (209–214), 210–211f, 212t, 213t, 214m monocirrhus Exocoetus, 191, 193 Monogenea, 52 monogenes Exocoetidae, 150 Strongylura marina, 28 Strongylura notata forsythia, 36 Strongylura timucu, 41 Tylosurus acus acus, 43, 44, 46 Tylosurus crocodilus crocodilus, 52 monroei Cypselurus, 153, 158 myxosporidians Hemiramphus brasiliensis, 105, 109 Strongylura marina, 28 Strongylura timucu, 41

M

macrochirus Hemiramphus, 100, 101, 102 macrorhynchus Hemiramphus, 97 maculata Belone, 8, 10 maculipinnis Exocoetus, 170 marginatus Hemiramphus, 99, 106 Margined Flyingfish, 150, 151, 152, (153–159), 154t, 155f, 157t, 158m, 185, 188 marina Belone, 39 Strongylura, 4f, 6, 7, 8t, 18, 19, (21–30), 22f, 26t, 27t, 30m, 39, 54 marinus Esox, 25 Strongylura, 22 Tylosurus, 18, 21, 22, 39, 41 maris-rubri maris-rubri Belone, 49 marisrubri marisrubri Belone, 49 Maya Needlefish, 4t, 5, 7, 8t, (18–21), 20m, 20f, 26, 29 meeki Hyporhamphus, 90, 94t, 95, 112, 113m, (113–117), 115f, 115t, 116t, 117t, 118, 119, 120, 123, 124, 126, 127 meeki (nomen nudum) Hyporhamphus, 114 Meek’s Halfbeak, 90, 92, 94t, 95, 112, 113m, (113–117), 115f, 115t, 116t, 117t, 118, 119, 120, 123, 124, 126, 127 melanochira Belone, 48, 50 melanurus Cheilopogon, 150, 152, 154t, 165, 170, 174t, 175t, (176–180), 177m, 178–179f, 187, 188, 190 Cypselurus, 176 Exocoetus, 176 Melapedalion, 127 mento Parexocoetus, 219 mento atlanticus Parexocoetus, 219 meristocystis Oxyporhamphus, 128 mesogaster Halocypselus, 196 Parexocoetus, 219

N

Nanichthys, 79 naos Hyporhamphus, 127 naresii Cypselurus, 186, 188 nematodes Hyporhamphus meeki, 116 Strongylura hubbsi, 19 Strongylura timucu, 41 Tylosurus acus acus, 47 Tylosurus crocodilus crocodilus, 52 nigricans Cheilopogon, 150, 185 Cypselurus, 160, 162 Exocoetus, 160 Noddy Terns, 40 nonsuchae Exonautes, 215 North Atlantic Sauries Key, 80 Northern Gannet, 82 notata Belone, 31, 32, 34 Strongylura, 4f, 6, 7, 18, 19, 23, (30–37), 31f, 32t, 33t, 34t, 36m

248

Index pinnatibarbatus californicus Cheilopogon, 185 pinnatibarbatus fernandizianus Cheilopogon, 185 pinnatibarbatus japonicus Cheilopogon, 185 pinnatibarbatus melanocercus Cheilopogon, 185 pinnatibarbatus pinnatibarbatus Cheilopogon, 185 Platybelone, 5, 6, (12–18), 15 argalus, 4f, 7, (12–18), 15m, 16t, 17t, 47 argalus annobonensis, 16t, 17, 17t argalus argalus, 8t, (12–18), 13f, 16t, 17t, 42 argalus lovii, 16t, 17, 17t argalus platyura, 16t, 17t argalus pterura, 16t, 17, 17t argalus spp., 16t, 17t argalus trachura, 15, 16t, 17, 17t pleei Hemiramphus, 107 Pleii Hemiramphus, 100, 103 pleii Hemiramphus, 101, 102, 103 poeyi Hemiramphus, 124 polleni Exocoetus, 209 polyethmus Cypselurus, 215 Praescomberesox pacificus, 80 procne Exocoetus, 170 Prognichthys, 150, 152, 186, (223–232) brevipinnis, 224 gibbifrons, 150, 224, 228, 229 glaphyrae, 150, 151, 154t, (224–228), 225t, 226–227f, 228m, 229 occidentalis, 150, 151, 154t, 224, 225t, 228m, (228– 232), 230–231f sealei, 224 tringa, 224, 229 pulchellus Exocoetus, 180 punctulatus Tylosurus, 4f, 42

Tylosurus, 35 notata forsythia Strongylura, 7, 8t, 31, 32t, 33t, (34–37), 34t, 36m notata notata Strongylura, 7, 8t, (31–34), 31f, 32t, 33t, 34t, 36m notatus Strongylura, 35 Tylosurus, 31, 34 nuttallii Exocoetus, 165

O

obtusirostris Exocoetus, 150, 151, 154t, (191–196), 192f, 194t, 195m, 195t Halocypselus, 193 occidentalis Prognichthys, 150, 151, 154t, 224, 225t, 228m, (228– 232), 230–231f Oceanic Bluntnose Flyingfish, 150, 151, 154t, (224– 228), 225t, 226–227f, 228m, 229 Oceanic Halfbeaks, (96–98) Oceanic Two-Wing Flyingfish, 150, 151, 154t, (191–196), 192f, 194t, 195m, 195t opisthopus Cypselurus, 186 orbignianus Exocoetus, 182, 184 Oxyporhamphus, 89, 91, 96, 98, (127–130) convexus, 99, 127, 129 meristocystis, 128 micropterus, 127, 128, 129 micropterus micropterus, 130 micropterus similis, 92, 94t, (127–130), 128f, 129m similis, 129

P

Pacific Ribbon Halfbeaks, 97 Pacific Saury, 79, 80 pacificus Praescomberesox, 80 Tylosurus, 42, 44, 46t Pajarito, 92, 94t, (119–122), 121t, 122f, 122t, 123, 123m Parexocoetus, 150, 152, 199, (219–223) brachypterus, 219, 221 brachypterus hillianus, 219, 221 brachypterus littoralis, 221, 222 hillianus, 150, 151, 154t, (219–223), 220f, 221t, 222t, 223m mento, 219 mento atlanticus, 219 mesogaster, 219 parrae Exocoetus, 153 peruvianus Exocoetus, 191, 193 pinnatibarbatus Cheilopogon, 150, 151, 152, 154t, (180–186), 181f, 183t, 185m, 188 Cypselurus, 160, 182 Exocoetus, 180 pinnatibarbatus altipennis Cheilopogon, 185

Q

quoyi Hyporhamphus, 113

R

Rainbow Runner, 129, 156, 195, 198 raphidoma Belone, 48, 49, 50 Strongylura, 49 Tylosurus, 6, 43, 44, 48, 49 raphydoma Tylosurus, 50 recurvirostra Rafinesque Sayris, 80, 81

249

Fishes of the Western North Atlantic Redfin Flyingfish, 150, 152, 154t, 202, (206–209), 206t, 207f, 208m, 208t, 212, 215, 218 Redfin Needlefish, 4t, 6, 7, 18, 19, 23, (30–37), 31f, 32t, 33t, 34t, 36m Redtail Ballyhoo, 90–91, 92, 94t, 99, 101, 101t, 102, 103, 104m, (105–109), 108f regalis Scomberomorus, 9, 11 Reporhamphus, 109, 110 melanochir, 91 Rhamphexocoetus volans, 150 Rhynchorhamphus malabaricus, 91 Richardi Hemiramphus, 124, 125–126 richardi Hemiramphus, 127 Roberti Hemiramphus, 119 roberti Exocoetus, 199, 202 Hemiramphus, 113, 119, 125 Hyporhamphus, 112, 113, 117, 118, (119), 120, 121t, 122, 122t, 123m, 125, 126, 127 roberti hildebrandi Hyporhamphus, 92, 94t, 119, 120, 121t, (122–124), 122f, 122t, 123m roberti roberti Hyporhamphus, 92, 94t, (119–122), 121t, 122t, 123, 123m robustus Exocoetus, 176 Hemiramphus, 99 rondeletii Cypselurus, 215 Danichthys, 202, 206, 215 Exocoetus, 170, 214–215 Exonautes, 215 Hirundichthys, 150, 175, 185, 206, 208, 215, 218, 219 Rondeletii Scomberesox, 81 rondeletii volador Hirundichthys, 215 rubescens Cypselurus, 209 rufipinnis Exocoetus, 199, 202, 206–207 Hirundichthys, 150, 152, 154t, 202, (206–209), 206t, 207f, 208m, 208t, 212, 215, 218

saltator Hemiramphus, 99, 101, 107 salvatoris Hyporhamphus, 119, 120 Sargassum, 9, 43, 47, 49 saurus Esox, 81 Scomberesox, 4f, 26, 30, 79, 80, (81–83), 81t, 82f, 83m, 84t saurus saurus Scomberesox, 80, (81–83), 82f, 84t saurus scombroides Scomberesox, 80, 83, 84t Sayris Rafinesque, 80 recurvirostra Rafinesque, 80, 81 scapularis Strongylura, 4f, 18 Scomberesocidae, 3f, (79–86), 81t, 84t Key, 4, 80 Scomberesocoidea, 2f Belonidae, (5–54), 8t, 149 Key, 4, 80 Scomberesocidae, 3f, 4, 8t, (79–86), 81t, 84t Scomberesox, (80–86) acutillus, 80 Camperi, 81 camperii, 81 equirostrum, 81 Rondeletii, 81 saurus, 4f, 26, 30, 79, 80, (81–83), 81t, 82f, 83m, 84t saurus saurus, 80, (81–83), 82f, 84t saurus scombroides, 80, 83, 84t scutellatum, 81 scutellatus, 81 simulans, 79, 81t, (83–86), 84t, 85f, 86m storeri, 81 Scomberesox sp., 83, 85 Scomberomorus regalis, 9, 11 scrutator Belone, 21, 25 Strongylura, 39 scutellatum Scomberesox, 81 scutellatus Scomberesox, 81 scylla Exocoetus, 199, 202 sealei Prognichthys, 224 senegalensis Strongylura, 4f, 18, 23, 26, 30 Shortfin Mako, 82 Silver Gar, 25 similis Oxyporhamphus, 129 simulans Scomberesox, 79, 81t, (83–86), 84t, 85f, 86m Skipjack Tuna, 40, 102, 129 Slender Halfbeak, 119 See also Pajarito Smallhead Flyingfish, 182 See also Bennett’s Flyingfish Smallwing Flyingfish, 128, 129 See also Atlantic Bigwing Halfbeaks

S

sagitta Tylosurus, 37, 39 Sailfin Flyingfish See Atlantic Sailfin Flyingfish Sailfish Ablennes hians, 11 Hemiramphus balao, 102 Hemiramphus brasiliensis, 106, 107 Hirundichthys affinis, 202 Strongylura notata forsythia, 36 saira Cololabis, 4f, 79, 80

250

Index

T

smithi Cypselurus, 160 Smooth Hammerhead, 51 Snoek, 184 Sooty Terns Ablennes hians, 11 Euleptorhamphus velox, 98 Hemiramphus balao, 99, 102 Hemiramphus brasiliensis, 106, 107 Oxyporhamphus micropterus similis, 128, 129 Strongylura timucu, 40 Spanish Mackerel, 35 speculiger Cypsilurus, 209 Exocoetus, 209 Exonautes, 209 Hirundichthys, 150, 152, 154t, 185, 199, 202, 205, 208, (209–214), 210–211f, 212t, 213t, 214m spilopus Exocoetus, 160, 162 Spotfin Flyingfish, 150, 152, 153, 154t, (165–170), 166–167f, 168m, 169t, 170t, 176, 188, 214 storeri Scomberesox, 81 Strongylura, 5–6, 7, (18–42) acus, 43 almeida, 39 anastomella, 4f, 18 ardeola, 13 argalus, 13 caribaea, 43 caribbea, 43 diplotaenia, 39 euryops, 39 exilis, 4f, 18, 26, 29 fluviatilis, 4f, 18 hubbsi, 4f, 5, 7, 8t, (18–21), 20m, 20f, 26, 29 incisa, 4f, 18 krefftii, 4f, 18 leiura, 4f, 18 longleyi, 6, 13 marina, 4f, 6, 7, 8t, 18, 19, (21–30), 22f, 26t, 27t, 30m, 39, 54 marinus, 22 notata, 4f, 6, 7, 18, 19, 23, (30–37), 31f, 32t, 33t, 34t, 36m notata forsythia, 7, 8t, 31, 32t, 33t, (34–37), 34t, 36m notata notata, 7, 8t, (31–34), 31f, 32t, 33t, 34t, 36m notatus, 35 raphidoma, 49 scapularis, 4f, 18 scrutator, 39 senegalensis, 4f, 18, 23, 26, 30 strongylura, 4f, 6, 18 timicu, 39 timocu, 39 timucu, 4f, 7, 8t, 18, 19, 23, 25, 26, 30, 30m, 36, (37–42), 40f, 40t, 41t timueu, 39 urvillii, 18 subtruncata Belone, 37, 39 subtruncatus Tylosurus, 37

timicu Strongylura, 39 timocu Strongylura, 39 Timucu, 4t, 7, 8t, 18, 19, 23, 25, 26, 30, 30m, 36, (37–42), 40f, 40t, 41t timucu Belona, 37 Belone, 21, 37 Esox, 37, 39 Strongylura, 4f, 7, 8t, 18, 19, 23, 25, 26, 30, 30m, 36, (37–42), 40f, 40t, 41t Tylosurus, 18, 37 timueu Strongylura, 39 trachura Belone, 13 trematodes Exocoetidae, 150 Hemiramphus balao, 102 Hemiramphus brasiliensis, 105, 106, 108 Hyporhamphus meeki, 113, 114, 116 Hyporhamphus unifasciatus, 124, 126 Parexocoetus hillianus, 222 Platybelone argalus argalus, 15 Strongylura marina, 21, 22, 28 Strongylura notata forsythia, 35, 36 Strongylura timucu, 37, 38, 41 Tylosurus acus acus, 42, 43, 46–47 Tylosurus crocodilus crocodilus, 48, 52 tricuspidatus Hyporhamphus, 109, 124, 126, 127 tringa Prognichthys, 224, 229 Tropical Two-Wing Flyingfish, 150, 151, 154t, 170, 191, 193, 194t, 195t, (196–199), 197f, 198m, 209 truncata Belone, 21, 25 Tylosurus, 5, 6, 7, (42–54), 91 acus, 4f, 6, 7, 36, 41, (42–48), 46t, 48m, 50, 52–53, 54 acus acus, 8t, (42–48), 45f, 46t, 54 acus imperialis, 44–45, 46t, 48 acus melanotus, 44, 46t, 47 acus rafale, 44–45, 46t, 47, 48 almeida, 22, 37 ardeola, 13 ardeolus, 13 caribaeus, 43 caribbaeus, 42, 50 choram, 42, 47, 50 crassus, 48 croccodilus, 50 crocodilus, 4f, 6, 7, 36, 41, 42, 44, 47, (48–54), 53t, 54m crocodilus crocodilus, 8t, 47, (48–54), 51f, 53t, 53t, 54m crocodilus fodiator, 53t crocodylus, 50 diplotaenia, 37 euryops, 37, 39 galeatus, 25 gavialoides, 4f, 6, 42, 47 gladius, 48, 50 hians, 8–9 imperialis, 44 latimanus, 42

251

Fishes of the Western North Atlantic vitropinna Cypselurus, 186, 188, 191 vittatus Hemiramphus, 100, 102 volador Exocoetus, 214, 215 Hirundichthys, 150, 152, 154t, 202, 206t, 208m, 208t, 212, (214–219), 216–217f volans Rhamphexocoetus, 150 volitans Exocoetus, 150, 151, 154t, 170, 191, 193, 194t, 195t, (196–199), 197f, 198m, 209

longirostris, 21 marianus, 22 marinus, 18, 21, 22, 25, 39, 41 notata, 35 notatus, 31, 34 pacificus, 42, 44, 46t punctulatus, 4f, 42 raphidoma, 6, 43, 44, 48, 49 raphydoma, 50 sagitta, 37, 39 subtruncatus, 25, 37 timucu, 18, 37

U

W

unifasciatus Hemiramphus, 113, 124, 125, 127 Hemirhamphus, 113 Hiporhamphis, 125 Hyporamphus, 125 Hyporhamphus, 90, 94t, 109, 112, 113, 113m, 115, 115t, 116t, 117, 117t, 118, 119, 120, 123, (124–127), 125f urvillii Strongylura, 18

Wahoo, 114, 116, 202 welsbyi Hemiramphus, 99 Western Atlantic Keeltail Needlefish, 8t, (12–18), 13f, 15m, 16t, 17t, 42 Western Bluntnose Flyingfish, 150, 151, 154t, 224, 225t, 228m, (228–232), 230–231f White Marlin, 107

Y

V

Yellowfin Tuna Exocoetus volitans, 198 Hyporhamphus unifasciatus, 126 Oxyporhamphus micropterus similis, 127, 128, 129 Strongylura timucu, 26 Tylosurus acus acus, 45

velox Euleptorhamphus, 92, 94t, (96–98), 97f, 98m vermiculatus Exocoetus, 153 vinciguerrae Exocoetus, 199, 202, 215, 219 Exonautes, 215 viridis Euleptorhamphus, 96, 97

Z

Zenarchopteridae, 1, 89

252