UNITED STATES DEPARTMENT OF THE INTERIOR, Douglas McKay, Secretary FISH AND WILDLIFE SERVICE, John L. Farley, Director //S FISHES OF THE GULF OF MAINE BY HENRY B. BIGELOW AND WILLIAM C. SCHROEDER First Revision FISHERY BULLETIN 74 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE VOLUME 53 [Contribution No. 592, Woods Hole Oceanographic Institution] UNITED STATES GOVERNMENT PRINTING OFFICE . Washington : 1953 For sale by the Superintendent of Documents, U. S. Government Printing Office Washington, 25, D. C. Price $4.25 (Buckram) CONTENTS Page Introduction 1 Area covered 1 Scope of the work 1 Sources of information 2 Use of the keys 4 Key to Gulf of Maine fishes 5 The Cyclostomes. Class Agnatha 9 Hagfishes and lampreys. Families Myxinidae and Petromyzomdae 9 Hag Myxine glutinosa Linnaeus 10 Sea lamprey Pelromyzon marinus Linnaeus 12 Cartilaginous fishes. Class Chondrichthyes 15 Sharks, torpedoes, skates and rays. Subclass Elasmobranchii 15 Sharks. Order Selachii 15 Sand sharks. Family Carchariidae 18 Sand shark Carcharias laurus Rafinesque 18 Mackerel sharks. Family Isuridae 20 Mackerel shark Lamna nasus (Bonnaterre) 20 Sharp-nosed mackerel shark, Mako Isurus oxyrinchus Rafinesque 23 Maneater, White shark Carcharodon carcharias (Linnaeus) 25 Basking sharks. Family Cetorhinidac 28 Basking shark Cetorhinus maximus (Gunnerus) 28 Thresher sharks. Family Alopiidae 32 Thresher Alopias vulpinus (Bonnaterre) 32 Cat sharks. Family Scyliorhinidae 34 Chain dogfish Scyliorhinus retifcr (Garman) 34 Smooth dogfishes. Family Triakidae 34 Smooth dogfish Mustelus canis (Mitchill) 34 Requiem sharks. Family Carcharhinidae 3f> Tiger shark Galeocerdo cuvier (LeSueur) 37 Blue shark Prionace glauca (Linnaeus) 38 Sharp-nosed shark Scoliodon terrae-novae (Richardson) 40 Dusky shark (Carcharhinus obscurus (LeSueur) 41 Brown shark Carcharhinus milberli (Milller and Henle) 43 Hammerhead sharks. Family Sphyrnidae 44 Bonnet shark, Shovelhead Sphyrna tiburo (Linnaeus) 44 Hammerhead Sphyrna zygaena (Linnaeus) 45 Spiny dogfishes. Family Squalidae 47 Spiny dogfish Squalus acanthias Linnaeus 47 Black dogfish Centroscyllium fabricii (Reinhardt) 51 Portuguese shark Centroscymnus coelolepis Bocage and Brito Capello.. 52 Gurry sharks. Family Dalatiidae 53 Greenland shark Somniosus microcephalia (Bloch and Schneider) __ 53 Dalatias licha (Bonnaterre) 55 Bramble sharks. Family Echinorhinidae 56 Bramble shark Echinorhinus brucus (Bonnaterre) 56 Torpedoes, skates and rays. Order Batoidei 57 Torpedo or electric rays. Family Torpedinidae 58 Torpedo Torpedo nobiliana Bonaparte 58 Skates. Family Rajidae 60 Barn-door skate Raja laevis Mitchill 61 Big skate Raja ocellata Mitchill 63 Brier skate Raja eglanteria Bosc 65 Leopard skate Raja garmani Whitley 66 Little skate Raja erinacea Mitlhill 67 Smooth-tailed skate Raja senla Garman 70 Thornv skate Raja radiata Donovan _ _ _ _ 72 m jy CONTENTS Cartilaginous fishes. Class Chondrichthyes— Continued Sharks, torpedoes, skates and rays. Subclass Elasmobranchii — Continued Torpedoes, skates and, rays. Order Batoidei— Continued Page Whip-tailed sting rays. Family Dasy atidae 74 Sting ray Dasyatis centroura (Mitchill) 74 Cow-nosed rays. Family Rhinopteridae 76 Cow-nosed ray Rhinoptera bonasus (Mitchill) 70 Devil rays. Family Mobulidae 77 Devil ray Mania birostris (Donndorff) 77 Chimaeroids. Subclass Holocephali 79 Chimaeras. Order Chimaerae 79 Family Chimaeridae 79 Chimaera Hydrolagus affinis (Brito Capello) 79 Bony fishes. Class Osteichthyes 80 Sturgeons. Family Acipenseridae 80 Sea sturgeon Acipenser sturio Linnaeus 81 Short-nosed sturgeon Acipenser brevirostrum LeSueur 84 Herring and Tarpon tribes. Families Clupeidae, Dussumieriidae, and Elopidae_ 85 Ten-pounder Elops saurus Linnaeus SG Tarpon Tarpon atlanticus (Cuvier and Valenciennes) 87 Round herring Etrumeus sadina (Mitchill) 87 Herring Clupea harengus Linnaeus 88 Hickory shad Pomolobus mediocris (Mitchill) 100 Alewife Pomolobus pseudoharengus (Wilson) 101 Blueback Pomolobus aestivalis (Mitchill) 100 Shad Alosa sapidissima (Wilson) 108 Thread herring Opisthonema oglinum (LeSueur) 112 Menhaden Brevoortia tyrannus (Latrobe) 113 Anchovies. Family Engraulidae U8 Anchovy Anchoa mitchilli (Cuvier and Valenciennes) 118 Striped anchovy Anchoa hepsetus (Linnaeus) 11" Salmons. Family Salmonidae H'-1 Brook trout Salvelinus fontinalis (Mitchill) 120 Salmon Salmo salar Linnaeus 121 Humpback salmon Oncorhynchus gorbuscha (Walbaum) 131 Silver salmon Oncorhynchus kisulch (Walbaum) 133 Smelts. Family Osmeridae 133 Capelin Mallotus villosus (Miiller) 134 Smelt Osmerus mordax (Mitchill) 135 Argentines. Family Argentinidae 139 Argentine Argentina silus Ascanius 13.) Luminescent fishes Lanternfishes. Family Myctophidae I41 Headlight-fish Diaphus effulgens (Goode and Bean) 142 Lanternfish Myclophum affine (Lutken) Pearlsides. Family Maurolicidae Pearlsides Maurolicus pennanti (Walbaum) 144 Viper Fishes. Family Chauliodontidae I45 Viperfish Chauliodus sloani Bloch and Schneider 145 Stomiatids. Families Gonostomidae and Stomiatidae 140 Cyclothone Cyclothone signata Garman 140 Slomias Stomias ferox Rheinhardt 147 Stomioides nicholsi Parr Trigonolampa miriceps Regan and Trewavas 148 Hatchet fishes. Family Sternoptychidae 149 Silver hatchet fish Argyropelecus aculeatus Cuvier and Valenciennes 149 Eels. Families Anguillidae, Congridae, Simenchelyidae, Synaphobranchidae, Nemichthyidae, and Ophichthyidae 150 Eel Anguilla rostrala (LeSueur) **>1 American conger Conger ocean ica (Mitchill) 154 CONTENTS Bony fishes. Class Osteichthyes — Continued Page Eels — Continued Slime eel Simenchelys parasiticus Gill 157 Long-nosed eel Synaphobranchus pinnalus (Gronow) 158 Snake eel Omochelys cruentifer (Goode and Bean) 159 Snipe eel Nemichthys scolopaceus Richardson 159 Lancetfishes. Family Alepisauridae 160 Lancetfish Alepisaurus ferox Lowe 161 Mummiehogs or killifishes. Family Poeciliidae 162 Common mummichog Fundulus heteroclitus (Linnaeus) 162 Striped mummichog Fundulus majalis (Walbaum) 164 Sheepshead minnow Cyprinodon variegatus Lacepede 165 Billfishes. Family Belonidae 167 Silver gar Tylosurus marinus (Walbaum) 167 Garfish Ablennes hians (Cuvier and Valenciennes) 168 Half beaks. Family Hemiramphidae 169 Halfbeak Hyporhamphus unifasciatus (Ranzani) 169 Needlefishes. Family Scomberesocidae 170 Needlefish Scomberesox saurus (Walbaum) 170 Flying fishes. Family Exocoetidae 171 Flying fish. Cypselurus heterurus (Rafinesque) 172 Silver hake and Cod families. Families Merlucciidae and Gadidae 173 Silver hake Merluccius bilinearis (Mitchill) 173 Co A Gadus collar ias Linnaeus 182 Tomcod Microgadus tomcod (Walbaum) 196 Haddock Melanogrartimus aeglefinus (Linnaeus) 199 American pollock Pollachius virens (Linnaeus) 213 White hake Urophycis tenuis (Mitchill) 221 Squirrel hake Urophycis chuss (Walbaum) 223 Spotted hake Urophycis rcgius (Walbaum) .. 230 Long-finned hake Urophycis chesteri (Goode and Bean) - 232 Blue hake Anlimora roslrala Gunthcr .. 233 Hakeling Physiculus fulvus Bean 233 Four-bearded rockling Enchclyopus cimbrius (Linnaeus)-.... 234 Cusk Brosme brosme (Muller) 238 Grenadiers. Family Macrouridae 243 Common grenadier Macrourus bairdii Goode and Bean 243 Rough-headed grenadier Macrourus berglax Lacepede 245 Long-nosed grenadier Coelorhynchus carminatus (Goode) .. 246 Opahs. Family Lampridae . 247 Opah Lampris regius (Bonnaterre) 247 Flounders and Soles. Families Hippoglossidae, Paralichthyidae, Pleuronectidae, Bothidae, and Achiridae 248 Halibut Hippoglossus hippoglossus (Linnaeus) 249 Greenland halibut Reinhardlius hippoglossoidcs (Walbaum) 258 American dab Hippoglossoides platessoides (Fabricius) 259 Summer flounder Paralichthys denlatus (Linnaeus) 267 Four-spotted flounder Paralichthys oblongus (Mitchill) 270 Yellow-tail Limanda ferruginea (Storer) 271 Winter flounder Pseudopleuronecles americanus (Walbaum)__ 276 Smooth flounder Liopsetta putnami (Gill) - 283 Witch flounder Glyptocephalus cynoglossus (Linnaeus) 285 Sand flounder Lophopsetla maculata (Mitchill) 290 Gulf Stream flounder Cilharichlhys arclifrons Goode 294 Hogchoker Achirus fascialus Lacepede 296 John Dories. Family Zeidae 297 American John Dory Zenopsis ocellata (Storer) 297 Grammicolepid fishes. Family Grammicolepidae 299 Grammicolepid Xenolepidichthys americanus Nichols and Firth 299 Snipe fishes. Family Macrorhamphosidae 301 Snipe fish Macrorhamphosus scolopax (Linnaeus) 301 VI CONTENTS Bony fishes. Class Osteichthyes — Continued Page Silversides. Family Atherinidae 302 Silverside Menidia menidia (Linnaeus) 302 Waxen silverside Menidia beryllina (Cope) 304 Mullets. Family Mugilidae 305 Mullet Mugil cephalus Linnaeus 305 Barracudas. Family Sphyraenidae 306 Northern barracuda Sphyraena borealis DeKay 306 Sticklebacks. Family Gasterosteidae 307 Nine-spined stickleback Pungitius pungitius (Linnaeus) 307 Three-spined stickleback Gasterosteus aculeatus Linnaeus 308 Two-spined stickleback Gasterosteus wheallandi Putnam 310 Four-spined stickleback Apelies quadracus (Mitchill) 311 Pipefishes. Family Syngnathidae 312 Pipefish Syngnathus fuscus Storer 312 Pelagic pipefish Syngnathus pelagicus Linnaeus 314 Seahorses. Family Hippocampidae 315 Sea horse Hippocampus hudsonius DeKay . 315 Trumpetfishes. Family Fistulariidae 316 Trumpetfish Fistularia tabacaria Linnaeus 316 Mackerels. Family Scombridae 317 Mackerel Scomber scornbrus Linnaeus 317 Chub mackerel Pneumatophorus colias (Gmelin) 333 Striped bonito Euthynnus pelamis (Linnaeus) 335 False albacore Euthynnus alleleratus (Rafinesque) 336 Common bonito Sarda sarda (Bloch) 337 Tuna Thunnus thynnus (Linnaeus) 338 Spanish mackerel Scomberomorus maculatus (Mitchill) 347 King mackerel Scomberomorus regalis (Bloch) 348 Cavalla Scomberomorus cavalla (Cuvier) 349 Escolars. Family Gempylidae 349 Escolar Ruvettus pretiosus Cocco 349 Cutlassfishes. Family Trichiuridae 350 Cutlassfish Trichiurus lepturus Linnaeus 350 Swordfishes. Family Xiphiidae 351 Swordfish Xiphias gladius Linnaeus 351 Spearfishes or Marlins and Sailfishes. Family Istiophoridae 357 Blue marlin Makaira ampla (Poey) 358 White marlin Makaira albida (Poey) 360 Dolphins. Family Coryphaenidae 360 Dolphin Coryphaena hippurus Linnaeus 360 Seabreams or pomfrets. Family Bramidae 361 Johnson's Sea bream Taractes princeps (Johnson) 361 Butterfishes. Family Stromateidae 363 Butterfish Poronotus triacanthus (Peck) 363 Harvestfish Peprilus alepidotus (Linnaeus) 368 Rudderfishes. Family Centrolophidae 369 Barrelfish Palinurichthys perciformis (Mitchill) 369 Black ruff Centroluphus niger (Gmelin) 370 Pompanos and Jacks. Family Carangidae 371 Pilotfish Naucrales ductor (Linnaeus) 372 Rudderfish Seriola zonata (Mitchill) 373 Mackerel scad Decapterus macarellus (Cuvier and Valenciennes) 374 Crevalle Caranx hippos (Linnaeus) 375 Hardtail Caranx crysos (Mitchill) 376 Saurel Trachurus trachurus (Linnaeus) 377 Goggle-eyed scad Trachurops crumenopthalmus (Bloch) 377 Moonfish Vomer sclapinnis (Mitchill) 378 Lookdown Selene vomer (Linnaeus) 279 Leatherjacket Oligoplites saurus (Bloch and Schneider) 380 Threadfin Alectis crin itus (Mitchill) 38 1 CONTENTS VII Bony fishes. Class Osteichthyes — -Continued Page Bluefishes. Family Pomatomidae 382 Bluefish Pomatomus sallatrix (Linnaeus) 383 Sea basses. Family Serranidae 389 Striped bass Roccus saxatilis (Walbaum) 389 White perch Morone americana (Gmelin) 405 Sea bass Cenlroprisles striatus (Linnaeus) 407 Wreckfish Polyprion americanus (Bloch and Schneider) 409 Catalufas or Big Eyes. Family Priacanthidae 410 Short big-eye Pseudopriacanthus altus (Gill) 410 Porgies. Family Sparidae 411 Scup Slenotomus versicolor (Mitchill) 411 Sheepshead Archosargus probatocephalus (Walbaum) 416 Croakers, Drums, and Weakfishes. Family Sciaenidae 417 Weakfish Cynoscion regalis (Bloch and Schneider) 417 Spot Leiostomus xanthurus Lac<5pede 423 Kingfish Menticirrhus saxatilis (Bloch and Schneider) 423 Black drum Pogonias cromis (Linnaeus) 425 Tilefishes. Family Branchiostegidae 426 Tilefish Lopholatilus chamaeleonticeps Goode and Bean 426 Rockfishes. Family Scorpaenidae 430 Rosefish Sebasles marinus (Linnaeus) 430 Black-bellied rosefish Helicolenus daclylopterus (De La Roche) 437 Boarfishes. Family Caproidae 438 Boarfish Antigonia capros Lowe 438 Sculpins and Sea Ravens. Families Cottidae and Hemitripteridae 439 Hook-eared sculpin Artediellus uncinatus (Reinhardt) 440 Mailed sculpin Triglops ommatistius Gilbert 441 Grubby Myoxocephalus aeneus (Mitchill) 443 Shorthorn sculpin Myoxocephalus scorpius (Linnaeus) 445 Longhorn sculpin Myoxocephalus oclodecemspinosus (Mitchill) 449 Staghorn sculpin Gymnocanthus Iricuspis (Reinhardt) 452 Arctic sculpin Cottunculus microps Collett 453 Sea raven Hemitripterus americanus (Gmelin) . 454 Alligat orfishes. Family Agonidae 457 Alligatorfish Aspidophoroides monopterygius (Bloch) 457 Lumpfishes. Family Cyclopteridae 459 Lumpfish Cyclopterus lumpus Linnaeus 459 Spiny lumpfish Eumicrotremus spinosus (Miiller) 463 Sea snails. Family Liparidae 464 Sea snail Neoliparis atlanticus Jordan and Evermann 464 Striped sea snail Liparis liparis (Linnaeus) 466 Sea robins or Gurnards and Armored sea robins. Families Triglidae and Peristediidae.. 467 Common sea robin Prionolus carolinus (Linnaeus) 467 Striped sea robin Prionolus evolans (Linnaeus) 470 Armored sea robin Peristedion miniatum Goode 471 Flying gurnards. Family Dactylopleridae 472 Flying gurnard Daclylopterus volitans (Linnaeus) 472 Cunner Tribe or Wrasses. Family Labridae 473 Cunner Tautogolabrus adspersus (Walbaum) 473 Tautog Tautoga onitis (Linnaeus) 478 Remoras. Family Echeneidae 484 Shark sucker Echeneis naucrates Linnaeus 485 Swordfish sucker Remora brachyptera (Lowe) 486 Remora Remora remora (Linnaeus) 487 Sand launces. Family Ammody t idae 487 Sand launce Ammodytes americanus DeKay 488 VIII CONTENTS Bony fishes. Class Osteichthyes — Continued Pag» Blenny-like fishes. Families Lumpenidae, Pholidae, and Stichaeidae 491 Rock eel Pholis gunnellus (Linnaeus) 492 Snake blenny Lumpenus lumpretaeformis (Walbaum) 494 Shanny Leptoclinus maculatus (Fries) 497 Arctic shanny Stichaeus punctalus (Fabricius) 497 Radiated shanny Vivaria subbifurcata (Storer) 498 Wrymouths. Family Cryptacanthodidae 500 Wry mouth Cryptacanthodes maculatus Storer 500 Wolffishes. Family Anarhichadidae 502 WolfEsh Anarhichas lupus Linnaeus ._ 503 Spotted Wolffish Anarhichas minor Olafsen 507 Ocean pouts and Wolf eels. Family Zoarcidae 508 Ocean pout Macrozoarces americanus (Bloch and Schneider) 510 Wolf eel Lycenchelys verrillii (Goode and Bean) 515 Arctic eelpout Lycodes reticulatus Reinhardt 516 Cusk eels. Family Ophidiidae 517 Cusk eel Lepophidium cervinum (Goode and Bean) 517 Toadfishes. Family Batrachoididae 5 IS Toadfish Opsanus lau (Linnaeus) 518 Triggerfishes. Family Balistidae 520 Triggerfish Batistes carolinensis Gmelin 520 Filefishes. Family Monacanthidae 521 Filefish Monacanthus hispidus (Linnaeus) 522 Filefish Monacanthus c Hiatus (Mitchill) 523 Orange filefish Alulera schoepfii (Walbaum) 524 Unicornfish Alutera scripta (Gmelin) 525 Puffers and Porcupine-fishes. Families Tetraodontidae and Diodontidae 525 Puffer Sphaeroides maculatus (Bloch and Schneider) 526 Burrfish Chilomyctcrus schoepfii (Walbaum) 527 Ocean Sunfishes or Headfishes. Family Molidae 528 Sunfish Mola mola (Linnaeus) 529 Sharp-tailed sunfish Maslurus lanceolatus (Lifinard) 531 Anglers. Family Lophiidae 532 American Goosefish Lophius americanus Cuvier and Valenciennes 532 Sargassum fishes. Family Antennariidae 541 Sargassum fish Histrio pictus (Cuvier and Valenciennes) 541 Deep sea anglers. Family Ceratiidae 542 Deep sea angler Ceratias holbolli Kr0yer 543 Bibliography 545 Index 561 FISHES OF THE GULF OF MAINE By HENRY B. BIGELOW and WILLIAM C. SCHROEDER Museum of Comparative Zoology, Harvard University, and Woods Hole Oceanographic Institution During the summer of 1912 the Bureau of Fisheries, with the cooperation of the Museum of Comparative Zoology of Harvard University, commenced an oceanographic and biological survey of the Gulf of Maine, with special refer- ence to its fishes, to its floating plants and animals (plankton), to the physical and chemical state of its waters, and to the circulation of the latter. Cruises were made on the Fisheries schooner Grampus during the summers and autumns of 1912, 1913, 1914, 1915 and 1916, and during the winters and springs of 191 .'3 and 1915. The work was interrupted by the war, but was resumed with a cruise of the Fisheries steamer Albatross in the late winter and spring of 1920, and was continued by the Fisheries steamer Halcyon during the winter and spring of 1920-21, and the summers of 1921 and 1922. The first part of the general report, dealing with the fishes, was published in 1925, as Bulletin 40 (Pt. 1) of the United States Bureau of Fisher- ies;1 subsequent parts describing the plankton of the offshore waters of the Gulf and the physical characteristics of its waters were published in 1920-27, as Part 2. The preparation of the section on the fishes was assigned originally to W. W. Welsh, who bad gathered a large body of original observations on the growth, reproduction, diet, and other phases of the lives of many of the more important species. The report was far advanced when it was inter- rupted by his untimely death, and H. B. Bigelow undertook to carry it to publication along the lines originally laid down. The new edition, entailing a general revision and the addition of much new material, has been prepared jointly by H. B. Bigelow and by W. C. Schroeder. ' The Bureau of Fisheries was transferred on July 1, 1939, from the Depart- ment of Commerce to the Department of the Interior, and on July 30, 1940, It was consolidated with the Bureau of Biological Survey to form the Fish and Wildlife Service. 210941 — 53 2 AREA COVERED The term "Gulf of Maine" covers the oceanic bight from Nantucket Shoals and Cape Cod on the west, to Cape Sable on the east. Thus it includes the shore lines of northern Massachu- setts, New Hampshire, Maine, and parts of New Brunswick and of Nova Scotia. The eastern and western boundaries adopted in this paper are 65° and 70° West longitude, respectively. South- ern strays, or northern, which have no real status in the Gulf of Maine except by accident, are mentioned only briefly, or are relegated to foot- notes. The Gulf of Maine has a natural seaward rim formed by Nantucket Shoals, by Georges Bunk, and by Browns Bank. We have chosen the 150-fathom contour as the arbitrary offshore boundary, because this will include all of the species that are likely to be caught by commer- cial fishermen but will exclude almost the entire category of the so-called "deep-sea" fishes, which are numerous in the basin of the open Atlantic but are not constituents of the fauna of the Gulf of Maine, properly speaking. The general oceanography of this area has been the subject of another report, but it may not be nniiss to point out that the temperature of the Gulf and its fauna are boreal, and that its south- ern and western boundaries are the northern limit to common occurrence of many southern species of fishes and of invertebrates. SCOPE OF THE WORK Our aim has been a handbook for the easy identification of the fishes that occur in the Gulf of Maine, with summaries of what is known of the distribution, relative abundance, and more significant facts in the life history of each. The descriptions are as little technical as is com- patible with scientific accuracy, and are limited 1 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE chiefly to such of the external features of each kind of fish as may serve for identification in the field. References to more detailed descriptions and synonymies are given to Bigelow and Schroeder (Fishes of the Western North Atlantic, Parts 1 and 2, 1948, 1953) for the cartilaginous fishes; also to Garman's beautiful plates for such of these as he pictured in his classic monograph, published in 1913, in vol. 36, of the Memoirs of the Museum of Comparative Zoology. References for the various species of bony fishes are to Jordan and Ever- mann's Fishes of North and Middle America (Bulletin 47, U. S. National Museum, 1896-1900, Parts 1-4), which still remains the only compre- hensive work on the bony fishes of North America. Many of the illustrations have been borrowed from earlier publications, but some of them are original. Keys are provided for all species as a further aid to identification. In most cases the sizes of larval fish and eggs are given in millimeters (1 inch equals 25.4 mm.); the sizes of the larger fishes are in inches and feet; weights are in pounds. The scientific nomenclature of the cyclostomes, of the elasmobranchs, and of the chimaeroids, follows Bigelow and Schroeder (Fishes of the Western North Atlantic, No. 1, Parts 1 and 2, 1948; 1953) that of the bony fishes follows Jordan, Evermann, and Clark's Check List of the Fishes and Fishlike Vertebrates of North and Middle America (Report, U. S. Commissioner of Fisheries for 1928 (1930), Part 2), unless otherwise noted. The families of bony fishes are arranged for the most part in the sequence employed by Jordan, Evermann, and Clark, except that the several families of luminescent fishes are grouped together, in the hope of making it easier for the nontechnical observer to identify such of them as may come to hand. SOURCES OF INFORMATION The literature dealing with the fishes of the Gulf of Maine begins with the earliest descriptions of New England. Captain John Smith, for in- stance, commented on the abundance of sturgeon, cod, hake, haddock, cole (the American pollock), cusks, sharks, mackerel, herring, dinners, eels, salmon, and striped bass, in his Generall Historie of Virginia, New England and the Summer Isles, published in 1616, while Wood in his New Eng- land's Prospect, 1634, gave much interesting infor- mation, some of which we quote hereafter. The sea fishes of northern New England and of the Maritime Provinces had begun to attract scientific attention by the early part of the nine- teenth century, and many local faunal lists have been published since then. The following are the most important of these, in chronological arrange- ment: 1850. Report on the sea and river fisheries of New Brunswick, within the Gulf of St. Lawrence and Bay of Chaleur, M. H. Perley, 137 pp., 1850. Fredericton, New Brunswick. 1853-1867. A history of the fishes of Massachusetts, David Humphreys Storer. Memoirs, American Academy of Arts and Sciences, New Series, vol. 5, pp. 49-92, 122-168, and 257-296; vol. 6, pp. 309-372; vol. 8, pp. 389-439; vol. 9, pp. 217-256, 39 pis. (Also in book form with supple- ment, 1867), Cambridge and Boston. 1879. A list of the fishes of Essex County, including those of Massachusetts Bay, George Brown Goode, and Tarleton H. Bean. Bulletin, Essex Institute, vol. 11, No. 1, pp. 1-38. Salem. 1884. Natural history of useful aquatic animals, George Brown Goode and associates, Section I, The Fisheries and Fishery Industries of the United States. Published jointly by the U. S. Fish Commission and the U. S. Bureau of the Census, 895 pp. Washington. 1908. Fauna of New England. 8. List of the Pisces, William C. Kendall. Occasional Papers, Boston Society of Natural History, vol. 7, No. 8, April 1908, pp. 1-52. Boston. 1914. An annotated catalogue of the fishes of Maine, William C. Kendall. Proceedings, Portland Society of Natural History, vol. 3, 1914, Part 1, pp. 1-198. Portland. 1922. The fishes of the Bay of Fundy, A. G. Huntsman. Contributions to Canadian Biology (1921), 1922, No. 3, pp. 1-24 (51-72). Ottawa. These lists contain all the early published local- ity records of the rarer species, either first hand, or by reference to original sources, while the last two, with a paper by Gill,2 and the first edition of the present book give complete bibliographies for the Canadian coasts of the Gulf and for the coasts of Maine and of Massachusetts. A similar list of the captures of deep water fishes along the outer part of the continental shelf is to be found in Goode and Bean's "Oceanic Ichthyology."3 The most pertinent extralimital lists are Smith's4 and Sumner, Osburn and Cole's 6 lists of Woods J Rept. U. S. Comm. Fish., (1904) 1905, pp. 163-188. 3 Smithsonian Contribution to Knowledge, vol. 30, 1895. < Bull. U. S. Fish Comm., Vol. 17, 1898, pp. 85-111. 1 Bull. V. S. Bur. Fish., vol. 31, Pt. 2, 1913, pp. 549-794. FISHES OF THE GULF OF MAINE Hole fishes; Halket's 6 Checklist of the fishes of Canada and of Newfoundland, and Vladykov and McKenzie's The Marine Fishes of Nova Scotia.7 The literature dealing with the habits of the fishes of the Gulf of Maine is very extensive, for most of the important commercial species, and many of the others also, are common to both sides of the North Atlantic. Among general European manuals, Day's Fishes of Great Britain and Ire- land,8 Smitt's "Scandinavian Fishes," 9andEhren- baum's summary of the many scattered accounts of the eggs and larvae of northern fishes 10 have been especialty helpful. A large amount of information as to local dis- tribution and abundance of various fishes has been gleaned from unpublished material in the files of the U. S. Fish and Wildlife Service, as well as from the fishery statistics published by the Fisheries Branch, U. S. Fish and Wildlife Service (formerly the U. S. Bureau of Fisheries), by the Dominion of Canada, and by the Commonwealth of Massa- chusetts. The superintendents of the Woods Hole, Gloucester, and Boothbay hatcheries have supplied much valuable information, as have other members of the U. S. Fish and Wildlife Service. Among these, Leslie Scatlergood has given many interesting pieces of information for Maine waters, while Howard Schuck has contributed authentic- ity to the account of the haddock. Dr. A. G. Huntsman has contributed his unpublished notes on the fishes of the Bay of Fundy and Gulf of St. Lawrence. Dr. A. H. Leim, Mr. R. A. McKen- zie, and Dr. Vadim D. Vladykov have supplied us with pertinent information on certain species from the Nova Scotian-St. Lawrence River regions. The late Prof. J. P. McMurrich permitted the use of his unpublished plankton records, and a number of Newfoundland records were furnished by Drs. George W. Jeffers and E. Templeman. The late W. F. Clapp has contributed many interesting notes gleaned during his experience as a fisherman before entering the scientific field. Harry Piers of the Provincial Museum of Halifax, • Checklist of the Fishes of the Dominion of Canada and Newfoundland 1913, 138 pp. ' Proc. Nova Scotia Inst, of Science, vol. IP, Pt. 1, 1935, pp. 17-113. 1 The fishes of Great Britain and Ireland, by F. Day, Text vol. 1, CX1I + 336 pp., vol. 2, 388 pp., and atlas, 179 plates, 1880-1884. London and Edin- burgh. ' A history of Scandinavian fishes. Second edition, vol. 1, 1892; vol. 2, 1895; 1,240 pp., 53 pis. Stockholm. ,0 Eier und Larven von Fischen. Nordisches Plankton, vol. I, 413 pp., 148 figs.; appeared in two parts as Lief. 4, 1905, and Lief. 10, 1919. has supplied interesting information on the occur- rence of the blue shark. John Worthington has furnished us with pound-records for the Truro- Provincetown region covering a recent span of about fifteen years and has given us specimens of three species heretofore unreported in the Gulf of Maine. Benjamin H. Morrow has supplied inter- esting data from the vicinity of Sandwich, Mass. We have received much information about the striped bass in Nova Scotia from Major Howard Scott, through the kind offices of Henry Lyman. And we owe it to consultation with Dr. A. Vedel Timing of the Marine Biological Laboratory, Charlottenlund, Denmark, and the specimens contributed by Dr. C. E. Lucas of the Scottish Fisheries Laboratory, Aberdeen, that we have dared to reach a conclusion as to the relationship between the rosefish of our gulf and of north European waters. Francis Sargent, also of the Division of Marine Fisheries of Massachusetts, and Henry Lyman, editor of the Salt Water Sportsman, have been unfailing in their response to our many inquiries. Myvanwy Dick of the Harvard Museum of Comparative Zoology has been of assistance in the handling of certain of our study material and in the preparation of a number of illustrations. The illustrations of the hagfish and lamprey and most of those of the sharks, slates, rays, and chimaera arc reprinted here through the courtesy of the Sears Foundation for Marine Research, publisher of the Fishes of the Western Atlantic Memoir 1, Parts 1 and 2, in which the illustrations originally appeared. Claude Ronne of the Woods Hole Oceanograpliic Institu- tion prepared many photographs from both original and published drawings, which were used to illustrate this book. We owe a debt of gratitude, also, to the late Dr. Samuel Garman, who was ever ready with assistance until the time of his death, and to W. C. Adams, former director of the division of fisheries and game of the State of Massachusetts. We wish to express our hearty thanks to the many commercial fishermen and to the many salt water anglers of our acquaintance who have met our inquiries in the most cordial way and who have supplied us with a vast amount of first-hand information on the habits, distribution, and abundance of the commercial and game fishes, which could be had from no other source. The 4 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE preparation of this book would have been out of the question without their help. Finally, we have ourselves gathered a large body of data as to distribution, habits, spawning seasons, and like matters, through many years, at many localities, both inshore and on the offshore banks. USE OF THE KEYS The various fins and other structures mentioned in the keys are named in the accompanying out- lines of a haddock and of a typical shark (fig. 1). A simple way to explain the use of the keys is to use the haddock as an example, running it down with the illustration at hand for reference. Turning to Key A (p. 5), we find that our fish fits the second alternative under section 1, since it has bony jaws and pectoral fins, and is not shaped like an eel. This refers us to section 3. There being only one gill opening on each side, we go from section 3 to section 5. As our fish does not have a tubular snout section 5 refers us to section 6, and this in turn to section 7, since neither the upper jaw nor the lower is greatly 1st Dorsal Fin prolonged. Since the body is not square-cut close behind the dorsal and anal fins, but has a definite tail part, we proceed from section 7 to section 8, and from section 8 to section 11, for our fish has no sucking plate or disc, either on top of the head, or on the chest. Section 11 refers us in turn to section 12 because the tail fin is nearly symmetri- cal in outline. The anal fin being clearly and definitely separated from the caudal fin, we go from section 12 to section 13; and from section 13 to section 14, for our fish does not have any evi- dent light-producing ("luminescent") spots either on its sides or on its head. Our fish does not have a fleshy fin or flap either in front of the ordi- nary dorsal fins or behind them, but all of its dorsal fins are supported by rays that are visible if held against the light. Consequently, we proceed from section 14 to section 18, and this refers us to section 22, there being no flaps or tags of skin on the sides of the head." Our fish obviously does not lie flat on one side, i. e., it is not one of the flat fishes, which brings us to section 23, and n There is a barbel on its chin, but this is very different in appearance from the skin flaps around the jaws that are characteristic of the few species that fall under the first, alternative of section 18. 2nd Dorsal Fin 1 Baibel Ventral fin tst.Ana.lfin Figure 1.— Diagrams of a haddock (below) and of a typical shark (above) with terms used in the keys and descriptions. FISHES OF THE GULF OF MAINE 5 this in turn carries us to Key E (p. 7) because which refers it to section 2. And here the black it has three separate, well developed dorsal fins. lateral line and the dark blotch on each shoulder Since there are 3 dorsal fins and 2 anal fins, name it a haddock, section 1 of Key E sends us to the key to the cod Any other Gulf of Maine species is to be named and silver hake families (p. 173). Turning to the in the same way, starting with Key A, section 1, first section of the latter we find that our fish fits and following tlirough the appropriate alternatives the first alternative (3 dorsal fins and 2 anals), as thev refer it from section to section. KEY TO GULF OF MAINE FISHES Key A 1. Mouth soft, with no firm jaws; no pectoral fins; form eel-like 2 Mouth has firm jaws; pectoral fins are present even if the form is eel-like .3 2. Two separate fins on the back; no barbels on the snout Lamprey, p. 12 Only one fin on the back; with barbels on the snout Hag, p. 10 3. Five gill openings on each side 4 Only one gill opening on each side 5 4. General form cylindrical in all Gulf of Maine species; the forward edges of the pectoral fins are not attached to the sides of the head forward, past the gill openings; the gill openings are not confined to the lower surface; the upper margin of each orbit is free from the eyeball, as a free eyelid Sharks, key, p. 16 General form very flat, disclike; the forward edges of the pectoral fins are attached to the sides of the head forward past all of the gill openings; the gill openings are confined to the lower surface; the upper margin of each orbit i.- not free from the eyeball (no free eyelid) Skates and Rays, key. p. 57 5. The bones of the head are fused in a tubular snout, with the mouth at its lip refer to Key B, p. 6 No tubular snout 6 6. One or both jaws are prolonged as a bony sword or bill refer to Key C, p. 6 Neither jaw is greatly prolonged 7 7. Body abruptly square-cut, close behind the very high dorsal and anal fins refer to Sunfishes, key, p. 529 Body with distinct tail part ..8 8. There is a sucking plate or disc, either on the top of the head or on the chest 9 There is no sucking disc or plate 1] 9. The sucking plate is on the top of the head refer to Remora family, key, p. 485 The sucking disc is on the chest ..10 10. General form is like a tadpole; the anal fin originates about as far back as the tips of the pectorals. refer to Sea snail family, key, p. 464 General form is not like a tadpole, but is high arched, with longitudinal ridges; the anal fin originates far behind the tips of the pectorals refer to Lumpfish family, key, p. 459 1 1. Tail like a shark, i. e., with the upper lobe much longer than the lower Sturgeons, key, p. 81 Tail with the upper and lower lobes of equal lengths, or nearly so 12 12. No clear separation between the anal and the caudal fins, which together form one continuous fin (the anal portion may be either long or short) refer t o Key D, p. 6 Anal and caudal fins are separated by a deep notch, or by a space 13 13. Sides of body and head, or both, with luminescent spots or patches, easily seen if not damaged. refer to Luminescent fishes, key, p. 141 No luminescent organs 14 14. There is a fleshy ("adipose") fin, with neither rays nor spines, either in front of the rayed dorsal fin, or behind it.. 15 There is no fleshy ("adipose") fin, but both the dorsals (if there are two) are supported by rays or by spines that can be felt, if not seen ]8 15. The adipose fin is on the nape of the neck, in front of the dorsal fin Tilefish, p. 426 The adipose fin is behind the dorsal fin 16 16. The dorsal fin extends nearly the entire length of the body Lancetfish, p. 161 The dorsal fin is short, standing about midway of the body 17 17. Tail deeply forked refer to Smelts and Argentine Key, p. 133 Tail nearly square or only slightly forked refer to Salmon key, p. 120 18. The head is fringed with fleshy tags or flaps 19 The head is not fringed with fleshy tags or flaps .22 19. The pectorals are armlike 20 The pectorals are not armlike 21 6 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Key A — Continued 20. Body very broad and flat; mouth enormous Goosefish, p. 532 Body deep and flattened sidewise; mouth small Sargassum fish, p. 541 21. The first (spiny) dorsal fin is longer than the second (sofVrayed dorsal) ; neither dorsal fin is fleshy Sea raven, p. 454 The first (spiny) dorsal fin is much shorter than the second (soft-rayed dorsal) ; both of the dorsals are thick and fleshy Toadfish, p. 518 22. Fishes which lie flat on the one side, with both of their eyes on the other side; the upper side is dark, the lower side normally is pale _. refer to Flatfish tribe key, p. 248 Fishes which do not lie flat on one side 23 23. Two or more separate and well-developed dorsal fins, each with continuous membrane refer to Key E, p. 7 Only one well-developed dorsal fin with continuous membrane (this, however, may be preceded by isolated spines or rays) 24 24. Top of snout with several barbels or beards Rockling (cod family in part) , p. 234 No barbels or beards on the top of the snout 25 25. Jaws with very large canine tusks refer to Wolffishes key, p. 503 No large canine tusks in either jaw 26 26. Dorsal fin soft^rayed throughout its length, except that it may be preceded by a few separate spines. refer to Key F, p. 8 At least the forward one-third of the dorsal fin is with stiff sharp rays or spines refer to Key G, p. 9 Key B Fishes with tubular snouts (from No. 5, p. 5). 1. Head is horselike; rear portion of trunk is slender, prehensile; no caudal fin Sea horse, p. 315 Head is not horselike; rear part of trunk is not prehensile; there is a caudal fin 2 2. Body and head (measured from tip of snout) are only about 4 times as long as deep; the dorsal fin has a long, strong, saw-edged spine Snipefish, p. 30 1 Body and head (measured from tip of snout) are at least 25 times as long as deep; the dorsal fin does not have a large spine 3 3. The snout is not longer than the dorsal fin; the anal fin is very small; no ventral fins; the caudal fin is rounded Pipefishes, key, p. 312 The snout is more than 6 times as long as the dorsal fin; the anal fin is about as large as the dorsal; ventral fins are present though small; the caudal fin is forked Trumpetfish, p. 316 KeyC Fishes with bills or swords (from No. 6, p. 5) 1. Both of the jaws are elongated 4 Only one of the jaws is elongated 2 2. Upper jaw elongated, as a sword.. 3 Lower jaw elongated Halfbeak, p. 169 3. The sword is flattened dorso-ventrally, and is sharp-edged; the first dorsal fin is shorter than the sword forward of the eyes ; no ventral fins Swordfish, p. 351 The sword is round-edged; the first dorsal fin is nearly twice as long as the sword refer to Spearfishes or Marlins and Sailfish, key,12 p. 358 4. The caudal fin is well developed 5 No caudal fin; the tip of the tail is whip-like Snipe eel, p. 159 5. There are several finlets behind the dorsal and anal fins Needlefish, p. 170 No finlets behind the dorsal and anal fins refer to Billfishes or Silver gars, key, p. 167 Key D Bony fishes with snouts of ordinary form; with only one gill opening on each side, and with the anal fin continuous with the caudal fin around the tip of the tail (from No. 12, p. 5). 1 . Only one dorsal fin 2 Two separate dorsal fins, the first much bigger than the second, but shorter 7 2. Body band-shaped, the tail tapering to a whip-like tip Cutlassfish, p. 350 Body thick, eel-like; the vertical fins continuous around the tip of the tail in a broad band 3 3. The dorsal fin is spiny from end to end 4 The dorsal fin is soft-rayed, at least for almost all its length 5 » The sailflsh would also come under this heading should one ever be taken in the Gulf of Maine. The distinctions between it and the spearfishes are given on page 358. FISHES OF THE GULF OF MAINE 7 Key D — Continued 4. Mouth large and strongly oblique; there are no ventral fins Wrymouth, p. 500 Mouth small and horizontal; with small ventral fins Rock eel, p. 492 5. There are no ventral fins refer to Eel family key, p. 150 With small but distinct ventral fins, situated forward of the pectorals 6 6. The ventrals are situated behind the gill openings and are of ordinary form refer to Eelpout family, key, p. 509 The ventrals are situated on the chin, well in front of the gill openings and are reduced to forked, barbel-like struc- tures Cusk eel, p. 517 7. The ventral fins are situated below the points of origin of the pectorals; the skin is conspicuously scaly. refer to Grenadier family, key, p. 243 The ventral fins are situated far back, behind the tips of the pectorals; the skin is soft, without scales. Chimaera, p. 79 Key E Bony fishes of ordinary form, with 2 or 3 well-developed dorsal fins and with the anal fin and the rearmost dorsal separated from the caudal fin. (from No. 23, p. 6). 1. Three dorsal fins and 2 anal fins .refer to Cod family, key (in part), p. 173 Only 2 dorsal fins and 1 anal fin 2 2. With one or more small finlets between the second dorsal and anal fins and the caudal fin 3 No finlets between the second dorsal and anal fins and the caudal fin 4 3. With more than 3 dorsal finlets and 3 anal finlets refer to Mackerel family, key, p. 317 With only 2 dorsal finlets and 2 anal finlets Escolar, p. 349 With only 1 dorsal finlet and 1 anal finlet Mackerel scad (Pompano family, in part), p. 374 4. Head very broad; top and sides of head bony, with sharp spines or horns 5 Head not noticeably broad; sides of head have no spines or horns 7 5. First (lower) few rays of the pectoral fins are not separate from the remainder of the fin; the mouth is very large refer to Sculpin family, key, p. 440 First (lower) few rays of the pectoral fins are separate from the remainder of the fin; the mouth is not very large - 6 6. Each of the first (lower) 2 or 3 rays of the pectoral fins have the form of a separate feeler; outline of tip of snout is concave; the first few spines of the first dorsal fin are not separate from the remainder of the fin. refer to Sea robin and Armored sea robin, key, p. 467 First (lower) few rays of the pectorals do not have the form of feelers, but are connected, one with the next, by mem- brane, as a separate fin; outline of tip of snout convex; the first few spines of the first dorsal are separate. Flying gurnard, p. 472 7. First spine of first dorsal fin is very much stouter than the other spines, and can be locked erect by the second spine; no ventral fins; skin of the sides is very hard Triggerfish, p. 520 First dorsal spine is not much stouter than the others and cannot be locked erect by the second spine; ventral fins are well-developed; skin of the sides is soft 8 8. The space between the two dorsal fins is nearly as long as the first dorsal fin, or longer; the ventral fins are situated behind the middle of the pectorals -9 There is little or no free space between the two dorsal fins; the ventrals are in front of the middle of the pectorals... 11 9. Jaws long; teeth large and strong; anal with one spine Barracuda, p. 306 Jaws short; teeth weak 10 10. Anal fin is about as long as head (snout to gill openings) and has one weak spine, refer to Silverside family key, p. 302 Anal fin is only about half as long as head and has three stiff spines (only two spines in very young specimens). Mullet, p. 305 11. Caudal peduncle is extremely slender; the caudal fin is deeply forked Pompano family (in part) key, p. 371 Caudal peduncle is at least moderately deep and thick; the caudal fin is only moderately forked, at most 12 12. First dorsal fin is much lower than second dorsal 13 First dorsal fin is as high as the second dorsal fin, or higher 14 13. Anal fin is nearly as long as second dorsal fin Bluefish, p. 383 Anal fin is only about one half as long as second dorsal fin Rudderfish (Pompano family in part), p. 373 14. Body very thin through, flat sided, nearly two-thirds as deep as it is long to base of caudal fin; the back and also the ventral edge of the body are armed with bony plates; there is a finlet of three short spines in front of the anal fin John Dory, p. 297 Body stout, not more than one-third as deep as it is long; the sides are rounded; the back and lower surface are not armed with bony plates; there is no finlet in front of the anal fin 15 8 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Key E — Continued 15. First dorsal fin soft-rayed; second dorsal fin at least 5 times as long as first dorsal. refer to Cod family key, in part, p. 173 First dorsal fin noticeably spiny; second dorsal fin as long as first dorsal 16 16. Second dorsal fin not much longer than the anal fin refer to Sea bass family key, in part. p. 389 Second dorsal fin is about twice as long as the anal fin refer to Weakfish family key, p. 417 Key F Bony fishes with snouts of ordinary form; symmetrical tails; caudal fin distinct from the anal fin; neither with bar- bels on the top of the snout nor with canine tusks; and with only one well-developed dorsal fin; the latter is soft- rayed except that it may be preceded by a few short spines or by a series of hair-like rays without connecting membrane and that there may be an isolated spine on the top of the head (from No. 26, p. 6). 1. The rear parts of the dorsal fin and of the anal fin are broken up into series of almost separate finlets (fig. 191) 2 The rear parts of the dorsal and anal fins are not broken up into series of finlets 3 2. The forward parts of the dorsal and anal fins are very high and scythe-shaped; the pectorals are very long, reaching back considerably beyond the high part of the dorsal fin; there are no spines in front of the anal fin.. Sea bream, p. 361 The dorsal and anal fins are not very high and slope gradually rearward; the pectorals are small, their tips falling far short of the level of the front of the dorsal fin; the anal fin is preceded by two short stout spines. Leather jacket, p. 380 3. The mouth gapes back far beyond the eye refer to Anchovies key, p. 118 The mouth does not gape back much beyond the rear edges of the eyes, if that far 4 4. The whole of the anal fin is behind the rear end of the dorsal fin refer to Herring Tribe key, p. 85 Part or all of the anal fin is further forward than the rear end of the dorsal fin 5 5. There is a spine or a bristle-like rod on the top of the head over the eyes 6 There is no spine or bristle-like rod on the head over the eyes, but there may be a few short spines close in front of the dorsal fin 7 6. The spine on the top of the head is thick and very stiff and has no fleshy tab at its tip; mouth small; body stiff; fin rays slender, not fleshy refer to Filefish family, key, p. 521 The spine on the head is slender and flexible and has a fleshy tab or "bait" at its tip; body soft; mouth very large; fin rays thick and fleshy Deep-sea angler, p. 543 7. Form eel-like; snout sharp pointed Launce, p. 488 Form not eel-like; snout blunt 8 8. Dorsal fin originates on the head, about over the eyes Dolphin, p. 360 Dorsal fin originates far behind the eyes 9 9. Each ventral fin is represented by a single large stout spine refer to Stickleback key, p. 307 The ventral fins are of ordinary rayed type, or are lacking 10 10. The upper anterior profile of the head is conspicuously concave 11 The upper anterior profile of the head is more or less convex __12 11. The forward parts of the dorsal fin and of the anal fin are much higher than the rear parts, the first few rays of each being very much longer than the rays farther back Lookdown, p. 379 The dorsal and anal fins are only a little higher in front than rearward, the first few rays not being much longer than the rays farther to the rear Moonfish, p. 378 12. The forward rays of the dorsal and anal fins are very long and thread-like Thread-fin, p. 381 The forward rays of the dorsal and anal fins are not very long and thread-like 13 13. The entire body is armored with several rows of overlapping plates Alligator fish, p. 457 The body is not armored with overlapping plates 14 14. The skin is rough or prickly Refer to Puffers and Porcupine fishes, key, p. 526 The skin is smooth, though scaly 15 15. The front part of the dorsal fin is much higher than the rear part 16 The front part of the dorsal fin is not much higher than the rear part 17 16. The ventral fins are large and conspicuous Opah, p. 247 There are no ventral fins Refer to Butterfish and Harvest Fish, key, p. 363 17. The tail fin is conspicuously rounded 18 The tail fin is more or less deeply forked 19 18. The dorsal fin runs the whole length of the back from close behind the head to the caudal fin which it joins; there is a barbel on the chin disk, p. 238 The dorsal fin occupies only about one-third of the length of the back or less, and stands far to the rear; there is a considerable space between it and the caudal fin; there is no barbel on the chin. .Refer to Mummichog key, p. 162 FISHES OF THE GULF OF MAINE 9 Key F — Continued 19. The caudal peduncle is slender and has a conspicuous longitudinal keel on either side; the pectoral fins do not reach back as far as the point of origin of the dorsal fin Pilotfish, p. 372 The caudal peduncle is deep and has no longitudinal keel; the pectoral fins reach back farther than the point of origin of the dorsal fin 20 20. There are 6-8 short detached spines, each with a small triangular fin membrane, on the back in front of the dorsal fin Barrelfish, p. 369 There are no detached spines on the back in front of the dorsal fin 21 21. The ventral fins stand far behind the bases of the pectoral fins; the point of origin of the dorsal fin is little if any in advance of the anal fin; the pectoral fins (Gulf of Maine species) are very long, reaching back nearly to the base of the tail fin Flying fish, p. 172 The ventral fins stand about under the base of the pectoral fins; the point of origin of the dorsal fin is far in ad- vance of the anal fin; the pectoral fins are small, falling far short of the anal fin Black ruff, p. 370 KeyG Fishes as in Key F, except that at least the forward one-third of the single dorsal fin is spiny. There is no adipose fin behind the rayed dorsal nor fleshy flap in front of it (from No. 26, p. 6). 1 . The body (tip of snout to base of caudal fin) is at least as deep as it is long Boarfish, p. 438 The body is considerably longer than it is deep 2 2. The rear part of the dorsal fin is soft-rayed .3 The whole length of the dorsal fin is spiny 8 3. Sides of head bony, with knobs or spines 4 No knobs or spines on the sides of the head 5 4. Sides of head armed with conical spines; the spiny portion of the dorsal fin is at least as long as the soft part; the body is flattened sidewise Refer to Rosefish family, key, p. 430 Sides of head with low rounded knobs only; the spiny portion of the dorsal fin is considerably shorter than the soft part; body tadpole-shaped Arctic sculpin (Sculpin family in part), p. 453 5. The ventral fins are much larger than the pectorals; the eyes are very large Short big-eye, p. 410 The ventral fins are not larger than the pectorals; the eyes are not very large 6 6. The pectorals are sharply pointed; the body is much flattened sidewise Refer to Porgy family, key, p. 411 The pectorals are rounded; the body is not much flattened sidewise 7 7. The rear (soft) part of the dorsal fin is nearly as long as the front (spiny) part; the anal fin is much higher than long Seabass (Seabass family in part), p. 407 The rear (soft) part of the dorsal fin is less than half as long as the spiny (front) part; the anal fin is longer than high Refer to Cunner family, key, p. 473 8. The mouth is strongly oblique; there are no ventral fins Wrymouth, p. 500 The mouth is not strongly oblique; ventral fins are present (very small in one species) . .Refer to Blenny fishes, key, p. 491 THE CYCLOSTOMES. CLASS AGNATHA The lampreys are the most primitive of the appearance, but are easily distinguishable from true vertebrates, their skeletons being cartilagi- the true eels and, indeed, from most of the true nous without any true bone, and their skulls fishes, by their peculiar jawless sucking mouth hardly differentiated from the vertebral column situated at the tip of the snout, and, further, which forms a simple notochordal sheath. They from all Gulf of Maine eels by lacking pectoral have no true jaws, no ribs, no shoulder or pelvic fins, girdles, and no paired fins. They are eel-like in THE HAGFISHES AND LAMPREYS. FAMILIES MYXINIDAE AND PETROMYZONIDAE These two groups are easily distinguished, one nally, whereas the lampreys have no barbels, their from the other, by the fact that the hags have mouths are disc- or funnel-like, their eyes are well several barbels on the chin, that their mouths developed after the larval stage is past, and they are not disc- or funnel-like, that they have only have one or more dorsal fins separate from the one continuous fin fold on the back and around caudal fin. the tail, and that their eyes are not visible exter- 10 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Hagfish Myxine glutinosa Linnaeus 1758 Bigelow and Schroeder, 1948, p. 34. Description. — The hag, like the lamprey, lacks paired fins and fin rays. Its skeleton is wholly cartilaginous, without bones, its mouth is jawless; and its skin is scaleless. It is easily recognized by its eel-like form; by its single finfold (a fold of skin, not a true fin) running right around the tail and forward on the lower surface of the body with no division into dorsal, caudal, and anal fins; by the single gill pore on each side, just forward of the origin of the ventral finfold; by its lipless mouth, star-shaped in outline when closed; by the single nasal aperture at the tip of the snout; by its peculiar barbels or "tentacles," two flanking the mouth on either side and four surrounding the nostril; and by the evertible tongue studded with rows of horny rasplike "teeth." We might also mention the series of mucous sacs on either side of the abdomen, and point out that the dorsal finfold originates about two-thirds of the distance back from snout toward tip of tail, and the ven- tral fin fold one-third the way back, with the vent piercing it. Color. — Hags vary in color, perhaps to cor- respond with the color of the bottom, being gray- ish brown or reddish gray above, variously suf- fused, mottled, or piebald with darker or paler gray, with brown, or with bluish; they are whitish or pale gray below. Size. — Gulf of Maine hags grow commonly to a length of about 1% to 2 feet, with a maximum of 31 inches recorded off the coast of Maine. Habits. — The hag is found chiefly if not ex- clusively where the bottom is soft mud, where (to judge from its actions during the brief time it survives in aquaria) it spends its time lying em- bedded in the clay or mud with the tip of the snout projecting. And it is at home only in com- paratively low temperatures, cooler probably, than 50°, which confines it in summer to depths of 15 to 20 fathoms or more hi the Gulf of Maine. It is not a true parasite, as has sometimes been suggested, their being no reason to believe it ever attacks living, uninjured fish, but is a scavenger. Figure 2. — Hagfish {Myxine glutinosa). A, adult, Gulf of Maine, from Bigelow and Schroeder, drawing by E. N. Fischer. B, lower view of head of same; C, tongue-teeth of same as seen from above, about 3 times natural size; D, egg, after Dean, about 2 times natural size. FISHES OF THE GULF OF MAINE 11 Being blind, it doubtless finds its food by its greatly specialized olfactory apparatus. It feeds chiefly on fish, dead or disabled, though no doubt any other carrion would serve it equally well. And it is known to prey on marine annelid worms also, at least in Norwegian waters. It is best known for its troublesome habit of boring into the body cavities of hooked or gilled fishes, eat- ing out the intestines first and then the meat, and leaving nothing but a bag of skin and bones, inside of which the hag itself is often hauled aboard, or clinging to the sides of a fish it has just attacked. In fact, it is only in this way, or entangled on lines, that hags ordinarily are taken or seen. Being worthless itself, the hag is an unmitigated nuisance, and a particularly loathsome one owing to its habit of pouring out slime from its mucous sacs in quantity out of all proportion to its small size. One hag, it is said, can easily fill a 2-gallon bucket, nor do we think this any exaggeration. In American waters the commercial fishes most often damaged by it are haddock and the hakes (Urophycis) , these being the species most often fished for with long lines or with gill nets over the type of bottom the hag frequents. But it some- times damages cod also, and European authors describe it as attacking ling (Molva) and other members of the cod tribe, herring, mackerel, sturgeon, and even mackerel sharks under similar circumstances. Breeding habits. — -The hag was formerly believed to be a functional hermaphrodite, with its single sex organ first developing sperm in the posterior por- tion, eggs later in the anterior portion. However, recent detailed studies of the sex organ appear to show that such is not the case, but that either the male portion of the common sex organ matures in a given individual with the female portion remaining rudimentary, or vice versa." It has long been known that the eggs are few in number (only 19 to 30 having been counted in any one female) and large (up to 25 mm. in length), and the horny shell has a cluster of anchor-tipped filaments at each end that make the eggs easy of identification. Until 1900 none had been found that certainly had been laid naturally. In that year, however, hag eggs were reported from the western part of Georges Bank and from the south » Seo Blpelow and Schroeder, Fishes Western North Atlantic, Pt. 1, ch. 2, 1918. pp. 3.V-36, fur references. coast of Newfoundland by Dean (1900) ; " from the neighborhood of the Faroe Islands by Jensen; I5 from Norway by Hjort; I6 off Morocco bv Koe- foed.17 And they have been reported subsequently from the Bay of Fundy by Huntsman, from Frenchman Bay on the coast of Maine by Conel.18 The eggs are deposited on bottom, where they stick firmly to fixed objects of one sort or another by their terminal filaments and by threads of slime. The hag spawns throughout its range; also it spawns throughout the year, for females nearing ripeness and others nearly spent have been re- corded for winter and spring, as well as summer and autumn, in one part of its range or another. The few eggs so far reported have been from depths of 50 to 150 fathoms, most of them trawled on mud, clay, or sand bottom. We need only add that, to judge from their behavior in aquaria, the females cease to feed at the approach of sexual maturity, as many other fishes do. Newly hatched hags have never been seen, but inasmuch as the smallest yet described (about 2)i inches long), probably not long out of the egg, already resembled the adult in external appearance there is no reason to suppose that the hag passes through a larval stage greatly different from the adult. General range. — Arctic seas, and both coasts of the north Atlantic; Murman Coast and northern Norway south regularly to the Irish Sea, and to Morocco as a stray in the East; northern part of Davis Strait, south to the latitude of Cape Fear, N. C, in the west. It is represented in the cor- responding temperature-belt of the Southern I li misphere by a form (or forms) resembling it so closely that it is doubtful whether any sharp line can be drawn between them. Occurrence in the Gulf of Maine. — Apart from one record for the northern part of Davis Strait, the most northerly reports of the hag off the American coast are from southern Newfoundland and from the Grand Banks.19 But it is generally distributed along outer Nova Scotia at appropriate depths. And it is only too common in the Gulf » Mem. N. Y. Acad. Bel., vol. 2, Pt. 2, Art. 2, 1900. " Vlden. Meddel. Dansk nstui hist. Forenlng, 1900, p. 1. a Rept. Norwelglan Fishery and Mar. Invest., vol. 1, 1900, No. 1, ch. 4. p. 75. " Rept. Michael Sars North Atlantic Exped., Zool., vol. 4, No. 1,1927, p. 18 » Science, N. Ser., vol. 75, 1932, pp. 19-20. " It has not been reported for certain from West Greenland (so far as we can learn), from the outer coast of Labrador, or within the Gulf of St. Lawrence thouch It Is to be expected In the deeper parts of the latter. 12 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE of Maine; perhaps it is not absent there from any considerable area of smooth bottom. Thus, it is abundant off the north end cf Grand Manan; is reported from Passaamquoddy Bay and from various localities near Eastport; is to be found off-shore on muddy bottom all along the Maine coast; and is caught at times in considerable numbers on the Boon Island-Isles of Shoals fishing grounds and about Jeffreys Ledge, where we found it plentiful enough in the spring of 191.3 to have gutted 3 to 5 percent of all the haddock in the gill nets. Fishermen report it as equally numerous in the deeper parts of Massachusetts Bay. On the offshore banks the hag is well known, and it has been trawled at various localities along the outer edge of the Continental Shelf off New England at depths of from 100 to 200 fathoms, and deeper. We ourselves took 11 large ones in one set of a Monaco deep-sea trap in 260 fathoms off Nantucket on July 9, 1908, and it has been taken in from 300 to 500 fathoms off Marthas Vineyard; as deep as 524 fathoms on the southeast slope of Georges Bank. Sea lamprey Petromyzon marinus Linnaeus 1758 Lamprey; Spotted lamprey; Lamper; Eel- sucker Bigelow and Schroeder, 1948, p. 46. Description. — Lampreys are eel-like in ap- pearance, but have a soft, cartilaginous skeleton. They lack paired fins but have well developed dorsal and ventral finfolds. In the adult the jaws are so rudimentary that apparently they are wanting; the mouth is a longitudinal slit when closed, but forms an elliptical disk at the tip of the snout wben open, and is armed with many horny, hooked teeth arranged in numerous (11 to 12) rows, the innermost the largest. There are two dorsal finfolds, and there are seven open gill slits on each side, whereas the hag has only one gill pore on each side, and only one fin. The sea lamprey (the only member of its group known from our salt waters) can hardly be mistaken for any other fish, its eel-like appearance coupled with two dorsal fins and the jawless mouth placing it at a glance. Color. — Small specimens (whether on their way downstream or in salt water) are white below and uniformly colored above, usually described as blackish blue, or as lead colored, and more or less silvery. But large specimens usually are olive brown above, or of varying shades of yellow- brown, green, red, or blue, mottled with a darker shade of the same color, or sometimes nearly black if the dark patches are confluent. The lower surface is whitish, gray, or of a pale shade of the same hue as the ground color of the back. During breeding season, the landlocked form takes on more brilliant hues, with the ground tint turning bright yellow. Size. — The length at the time of transformation from the larval stage is about 4 to 8 inches (100- 200 mm.). Sexually mature individuals, taken ■?£+ **l 3& #"f >nyW? 4>'ii•*'I♦**,^, Figure 3. — Sea lamprey (Petromyzon marinus), about 18 inches long, Merriraac River; and open mouth disc of another Merrimac River specimen to show the arrangement of the horny teeth, about 0.9 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. FISHES OF THE GULF OF MAINE 13 in American rivers, average 2 to 2){ feet long, up to a maximum of about 3 feet. One of 33 inches weighed 2% pounds. Habits. — It has been known from early times that the sea lamprey breeds in fresh water. How- ever, it does not enter all the streams within its range indiscriminately. As an illustration, we may cite outer Nova Scotia and the Bay of Fundy, where lampreys run in the St. Marys, Sackville, Annapolis, Shubenacadie, Petit Codiac, and St. Johns Rivers, but not in the Moser or Apple Rivers, although these last also are "salmon" rivers. Their requirements are a gravelly bottom in rapid water for their spawning beds, with muddy or sandy bottom in quiet water nearby, for the larvae. In many small streams, and in larger ones if these are blocked by dams or high falls, they may spawn oidy a short distance upstream; even within the influence of the tide, although invariably in fresh water. But they are able to ascend fulls, if these are not too steep and high, by clinging to the rocks by their oral discs and resting. And they may run upstream for very long distances in large rivers, as they did formerly in the Moirimac and probably still do in the St. Johns River. They are still to be found 200 miles or more from the sea in the upper tributaries of the Delaware and Sus- quehanna systems. Since the breeding activities of the sea lamprey take place in fresh water, a brief account will suffice here. As the two sexes ripen, the males develop a strong ridge along the back, the females a crestlike fin between the anus and the caudal fin. Spawning, commencing when the tempera- ture of the water is about 50° F. (10° C.) is com- pleted by the time it has warmed to about 68°-70° (20°-21° C), and a sea lamprey has been found to contain 236,000 ova. Working in pairs, some- times with a second female assisting, they make depressions 2 to 3 feet in diameter and about 6 inches deep in the stream bed in stretches where the bottom is stony or pebbly, dragging the stones downstream in a pile with their suckerlike mouths. And they are able to move stones as large as one's fist. It is in these depressions that the eggs are deposited, not among the piles of discarded stones that have often been described as "nests." It seems that they all die after spawn- ing; not only have they often been found dead, but their intestines atrophy, they are attacked by fungus, and they become so debilitated that recovery seems out of the question. The larvae are different in appearance from the adults: blind, toothless, with mouths and fins of different shape. They continue in this state for a period estimated as 3 to 4 years, during most of which time they live in burrows in the mud or sand, or hide under stones. They are abundant in the mud of flats near the mouths of small tributary streams of river systems such as the Delaware and Susquehanna, where lampre3-s still breed in large numbers, and they subsist on minute organisms. At the end of this larval period, when they have grown to a length of 4 to 6 inches, they undergo transformation to the adult form and structure, an event occupying about two months, August to September or October. They run down to the sea in Xovember or December, to live and grow there for one or two years, so that large ones, not yet mature, are to be found in salt water all the year round. Little is known of the habits of the lampreys while they live in the sea further than that their mode of life centers around a fiercely predaceous nature. Judging from their land-locked relatives and from the occasions on which they have been found fastened to sea fish, they must be extremely destructive to the latter, which they attack by "sucking on" with their wonderfully effective mouths. The lamprey usually fastens to the side of its victim, where it rasps away until it tears through the skin or scales and is able to suck the blood. Its prey sucked dry, it abandons it for another. Probably lampreys are parasites and bloodsuckers pure and simple, for we cannot learn that anything but blood has been found in their stomachs, except fish eggs, of which lampreys are occasionally full.20 In salt water they have been found preying on mackerel, the various anadromous herrings, cod, haddock, American pollock (Pollachius) , salmon, basking sharks, swordfish, hake (Urophycis), sturgeons and eels. Sometimes as many as three or four are fast at one time to a single shad, and they are said to be exceedingly aggressive in their attacks on other fishes. Occasionally they are found fast to driftwood, even to boats. When not clinging to anything they are strong, vigorous swimmers, progressing by an undulating motion. » fioode. Fish. Ind. U. S., Sect. I, 18S4, p. 677. 14 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE General range. — Atlantic coasts of Europe and of North America; from the west coast of Greenland to Florida in the western side of the Atlantic ; from northern Norway to the Mediterranean in the eastern;21 running up fresh rivers to breed, and landlocked in certain American lakes. Occurrence in the Gulf of Maine. — No doubt the sea lamprey occurs along the whole coast line of the Gulf of Maine, for it is recorded in or at the mouths of numerous rivers and streams in Nova Scotia, New Brunswick, Maine, and Massachusetts; spe- cifically in the St John, Annapolis, Petit Codiac, and Shubenacadie Kivers and from the St. Andrews region in salt water in the Bay of Fundy; from Eastport, Bucksport, Casco Bay, and the Pre- sumpscott, Kennebec, and Penobscot Rivers in Maine; from the Merrimac River system; from the Exeter and Lamprey Rivers, tributaries of Great Bay, New Hampshire; and from the Parker River in northern Alassachusetts. Since lampreys never take the hook or are cap- tured in nets except on rare occasions they are sel- dom seen in salt water; only when running up our rivers are they familiar objects. But they have been taken as far offshore as the seaward slopes of Banquereau, Sable Island, and LaHave Banks off Nova Scotia; on Browns Bank; in the deep gully between the latter and Georges Bank, and over the continental slope off Nantucket and off Marthas Vineyard. Lampreys have long been known to run up New England rivers a little earlier in the spring than shad, perhaps beginning to work upstream as early as the beginning of April or even the end of March. In the rivers tributary to the Gulf of Maine the runs are at then peak during May and early June, with few, if any, entering later than that. The larvae have been reported by Doctor Huntsman as plentiful in the Shubenacadie (emptying into the Bay of Fundy) and no doubt they are to be found in the Merrimac system, in the Exeter River, and in other Gulf of Maine streams. Abundance. — The construction of impassible dams has sadly reduced the numbers of lampreys " Also reported from "West Africa" by Gunther, Cat. Fishes British Mu- seum, vol. 8, 1870, p. 502. in the larger rivers of New England. In the Mer- rimac, for example, once a famous lamprey river,22 so few now succeed in surmounting the succession of dams that a recent survey yielded no evidence of any now having access to the upper reaches. Some lampreys, however, are said to breed in the river below the Lowell dam;23 we have seen what resembled their "nests" in the Squannacook, a branch of the Nashua tributary to the Middle Mer- rimac, and they still continue numerous in some Gulf of Maine streams where they can reach suit- able spawning grounds without too great difficulty. We may quote catches of up to 119 recently in the Shubenacadie, where larvae also have recently been reported in abundance,24 and of more than 100 each on several occasions in the Exeter River,25 where they are familiar spectacles, as they gather at the falls at Exeter, N. H. But we ought perhaps to caution the reader that while lampreys, like other anadromous fishes, may seem plentiful when condensed between the narrow bounds of a river's banks, their numbers as a whole do not rival those of the more abundant of the salt-water fishes. Importance. — Lampreys were esteemed a great delicacy in Europe during the middle ages (histo- rians tell us Henry I of England died of a surfeit of them) and considerable numbers were captured of old in the rivers of New England for human food, particularly in the Connecticut and Merrimac Rivers. But the lamprey fishery has been scarcely more than a memory for 40 years past except lo- cally and in a small way for home consumption, or to supply the needs of biological laboratories. In the salt water of the Gulf of Maine the lamprey has never been of any commercial importance; the average fisherman might not see one in a lifetime, nor is there any sale for the few that are picked up by chance. But larvae are taken in considerable numbers for bait in the Susquehanna River, and perhaps elsewhere along the middle Atlantic coast. " For an account of the lamprey fishery in New England during the first half of the 19th century, see Goode, Fish, and Fishery Ind. U. S., Sect. 1, 1884, p. 680. « Bailey, Biol. Survey Merrimack Watershed, New Hampshire Fish and Game Dept., 1938, p. 155. « Information gathered for us by Dr. A. O. Huntsman. » Collected for the Biological Laboratory, Harvard University. FISHES OF THE GULF OF MAINE 15 CARTILAGINOUS FISHES. CLASS CHONDRICHTHYES The Shark and Skate Tribes, and the Chimaeroids These are fishlike vertebrates with well-devel- oped fins and teeth, and with 2 pairs of fins, one of them supported by the pectoral girdle, the other by the pelvic girdle. Their most distinctive char- acter, as contrasted with the bony fishes (p. 80) is that their entire skeleton, including the skull, is cartilaginous, without any true bone, though it is partly calcified, especially in the vertebrae; the skull is far simpler than it is among the bony fishes; the gill filaments are attached throughout their lengths to the partitions between the gill openings instead of being free; and the rear portion of the digestive tract is modified into the so-called "spiral valve" by the development of a special fold from its lining layer, which only a few bony fishes have. Fertilization is internal in all of them, and is effected by a pair of rodlike copulatory organs, each of which is developed from the inner edge of one of the two pelvic fins, and is supported by one or more cartilages. The sharks and rays are usually looked upon as more primitive than the bony fishes. SHARKS, TORPEDOES, SKATES, AND RAYS. SUBCLASS ELASMOBRANCHII The most obvious external character by which all the sharks, skates, and rays are distinguishable from all of the bony fishes is that they have five or more gill openings on either side of the head, in- stead of only one. They recall the lampreys in this respect, but it is a commonplace that their jaws and teeth are extremely well-developed. Their skins are tough, and are studded in most of them with denticles (placoid scales), which are not homologous with the scales of bony fishes, for both dermis and epidermis take part in their formation, instead of the dermis alone. The teeth of the sharks and rays represent placoid scales that are modified and are embedded in the gums alone, not in the jaws. The fins are supported at their bases by segmented cartilaginous rods, supplemented in all of the sharks, and in some of the rays by nu- merous slender horny fibers further out, instead of by rays or spines of the sorts that are seen in the bony fishes. All of their fins are covered with the same leathery skin that clothes the body. Among sharks the tail is uneven ("heterocercal"), with the vertebral column extending out into its upper lobe, but it is whip-like in most of the skates and rays, with no definite caudal fin. The torpedo is an exception to this rule. The modern representatives of the subclass may be grouped in two orders, the one (Selachii) to include all living sharks, the other (Batoidei) to include the sawfishes, the skates and the rays. They are separated one from the other by the following external differences, and there are skele- tal differences between them as well:26 1 . The gill openings are at least partly on the sides; the edges of the pectoral fins are not attached to the sides of the head in front of the gill openings; the upper edges of the orbits are free from the eyeballs, so that they form free eyelids Sharks, (p. 15). The gill openings are entirely on the lower surface; the edges of the pectoral fins are attached to the side of the head in front of the gill openings; the upper edges of the orbits are attached to the eyeballs so that they do not form free eyelids Sawfishes, skates and rays, (p. 57). *• For further discussion, s.t Blgelov and Schroeder, Fishes Western North Atlantic, Pt. 1, ch. :i, 1948. p M Sharks. Order Selachii Sharks always are objects of interest, not only to fishermen and mariners but to seaside visitors generally, because of their evil appearance, their ferocity, the large size to which some of them grow, the destruction they wreak on fishermen's nets and lines as well as on the smaller fishes on which they prey, and because of the bad reputation certain kinds have earned as maneaters. The Gulf of Maine is not particularly rich in sharks (very poor indeed compared with our southern coasts), for while the number of species actually recorded there is considerable (indeed any high-seas shark might stray thither) the little spiny dogfish alone is numerous in the sense in which this term is applied to the various com- mercial fishes. And only two of the larger species, 16 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE the mackerel shark {Lamna nasus), and the blue All Gulf of Maine sharks give birth to young shark (Prionace glauca), occur with us in numbers that are not only practically adult in structure sufficient for one to be fairly sure of seeing them but of relatively large size at birth, and there is a during a summer's boating off the coast north of placental connection between mother and embryo Cape Cod. in some, but not in others. Still other sharks lay With the larger sharks generally so scarce (the eggs; this is true of the chain dogfish (Scyliorhinus mackerel shark is harmless to anything larger than retifer, p. 34), which is common out on the conti- the fishes on which it feeds, and the blue nental shelf from the offing of Cape Cod, south- shark is also harmless, although better armed), ward, and of its immediate relatives; also of the the danger of attacks on bathers is negligible in heterodontids or Port Jackson sharks which are our Gulf. Indeed, not a single well-authenticated not represented in the Atlantic, instance of the sort is on record 27 for the past There is so little market for sharks in Gulf of 80 years for the coast north of Cape Cod, though Maine ports (attempts to introduce the dogfish as the beaches are crowded every summer with a food fisn having failed so far) that the amounts vacationists. But as long as the white shark or landed in Maine and Massachusetts were only man-eater (Carcharodon carckarias) does stray aD0Ut 240,000 pounds in 1947, and about 309,500 occasionally into the Gulf (p. 26), it is always poimds in 1949; they interest fishermen chiefly as remotely possible that we may be horrified some nuisances because of the damage they do to nets summer by the news of tragedies such as occurred and other gear, except that mackerel sharks are on the New Jersey coast in July 1916, when marketable. several persons were killed or injured, presumably Jfc ig posgible tQ y^ a]1 ^ gharks SQ faf by a small shark of this species that was captured kiwn from ^ Qulf (and ^ indudes aU fchftt nearby a few days later,-8 and near Mattapoisett, are likdy tQ Qcmr there fc g) by the on Buzzards Bay, Mass., on July 25 1936, when gizes ^ rektive locations of the ^ and by a swimmer was fatally mimed by a shark, species , , ,, , , , , ,_ . „ J ' * such tooth characters as mav be seen at a glance not determined. ,, ., -, ' t ,, • ,, ,, c at the open mouth or easily felt with the linger « In 1830 (an event often quoted) one Joseph Blaney, fishing from a small (qfter the shirk is dead') boat in Massachusetts Bay off Swampscott, Mass., was attacked by some ^ "■'" fish that was seen to overset and sink his boat and, presumably, devoured Wq have attempted ill the following descriptions him, for neighboring fishermen who hastened to his rescue found no trace of . _ . _ . him. Whether his attacker was a large shark or a killer whale is an open of the Several Species to include Ollly SUCh features question. „,,.,„,„ „„ „ , , ,„,„ x- as will tell what shark is at hand; for more minute "Murphy and Nichols (Brooklyn Mus. Quart., vol. 3, 191fi, No. 4, pp. t45-if;o) give a detailed account of this occurrence. particulars we refer the reader to our account of « See Gudger (Amer. Midland Natural., vol. 44, 1950, p. 714) for clinical . . , . , »T , , . . , 0, details of this case. tue sharks of the western North Atlantic (p. 2). Key to Gulf of Maine Sharks 1 . There is an anal fin 2 There is no anal fin 16 2. Head greatly expanded sidewise, at level of eyes, in hammer- or shovel-form 3 Head of ordinary shape, with rounded or pointed snout 4 3. Outline of front of head only slightly concave opposite nostrils if at all so; grooves (if any) from nostrils shorter than horizontal diameter of eyes; free tip of second dorsal fin is not longer than forward margin of the fin; rear margin of anal fin is only weakly concave; teeth near outer corners of mouth are rounded, without sharp cusps. Shovel head, p. 44 Outline of front of head is deeply indented opposite each nostril; grooves from nostrils are more than twice as long as horizontal diameter of eye; free tip of second dorsal fin is considerably longer than front margin of the fin; rear margin of anal fin deeply concave; teeth near corners of mouth are like those near center of mouth, with sharp eusps Hammerhead, p. 45 4. Caudal peduncle (root of tail) is not widely expanded sidewise as a lateral keel on either side; upper lobe of caudal fin is much longer than lower lobe 8 Caudal peduncle is widely expanded sidewise as a lateral keel on either side; lower lobe of caudal fin is nearly as long as upper lobe, suggesting the caudal fin of a mackerel or swordfish 5 FISHES OF THE GULF OF MAINE 17 5. Gill openings very large, the first pair nearly meeting below the throat; teeth tiny, many hundred in number; gill arches with numerous horny gill rakers directed inward-rearward Basking shark, p. 28 Gill openings, confined to sides of head; teeth large, few in number; gill arches do not have horny gill rakers 6 6. Upper teeth broadly triangular, with serrate edges; anal fin is entirely behind second dorsal fin White shark, maneater, p. 25 Upper teeth with smooth-edged cusp, with or without a denticle on either side, at the base; anal fin is not entirely behind second dorsal fin 7 7. First two teeth from center in each jaw are similar to the succeeding teeth; origin of first dorsal fin is over or in front of inner corner of pectoral fin when latter is laid back; forward part of caudal fin has a small secondary lateral keel on each side, below the primary keel formed by the lateral expansion of the caudal peduncle. Mackerel shark, p. 20 First two teeth from center in each jaw are noticeably more slender and more flexuous than the succeeding teeth; origin of first dorsal fin is behind inner corner of pectoral fin when latter is laid back; forward part of caudal fin does not have a secondary longitudinal keel Sharp-nosed mackerel shark, mako, p. 23 8. Upper lobe of caudal fin is nearly or quite as long as head and body combined Thresher, p. 32 Upper lobe of caudal is less than one-half as long as head body combined 9 9. Second dorsal fin is nearly as high vertically as first dorsal fin 10 Second dorsal fin is less than one-half as high vertically as first dorsal fin 12 10. First dorsal fin is wholly or mostly forward of the origin of the pelvic fins 11 First dorsal fin is wholly posterior to bases of pelvic fins Chain dogfish, p. 34 11. Teeth high, narrow, sharp pointed, not in mosaic arrangement; snout conical; fifth gill openings well in front of pectoral fins Sand shark, p. 18 Teeth small, low, rounded, in mosaic arrangement; snout flat, broadly rounded in front; fifth gill openings are behind origins of pectoral fins Smooth dogfish, p. 34 12. Origin of first dorsal fin far behind inner corner of pectoral fin; upper surface brilliant blue in life. Blue shark, p. 38 Origin of first dorsal fin is over or anterior to inner corners of pectorals; ground color of upper surface is gray, brownish or dusky in life, not bright blue, 13 13. Length of snout in front of mouth is not more than one-half as great as breadth of mouth: upper jaw has a furrow on either side extending from outer corner forward past level of eye: caudal peduncle with a low longitudinal keel on either side; upper and lower teeth are of shapes shown in figure 11; their margins coarsely serrate. Tiger shark, p. 37 Length of snout in front of mouth is more than two-thirds as great as breadth of mouth; furrows on upper jaw, if any, do not extend forward-inward as far as level of eye: caudal peduncle without longitudinal ridges: teeth are not of shape shown in figure 11, their margins either only very finely serrate or smooth 14 14. Outer corners of mouth have a short "labial furrow" extending inward-forward along each jaw; teeth are alike in the two jaws, directed sharply outward, margins of upper teeth smooth, as well as those of lower teeth. Sharp-nosed shark, p. 40 Outer corners of mouth have no labial furrow on lower jaw and upper labial furrow is so short as to be hardly notice- able; teeth directed only moderately outward, their margins only finely serrate; lowers noticeably more slender than uppers 15 15. Origin of first dorsal fin is about over inner corner of pectoral when latter is laid back; vertical height of first dorsal fin is less than distance from eye tn first gill opening Dusky shark, p. 41 Origin of first dorsal is about over axil (armpit) of pectoral, its vertical height (after birth) is at least as great as distance from eye to third gill opening Brown shark, p. 43 16. Trunk much flattened dorso-ventrally ; eyes on top of head; front margins of pectorals overlap the gill openings. Angel shark, note, p. 18 Trunk subcylindrical; eyes on side of head: front margins of pectorals do not overlap the gill openings 17 17. Each dorsal fin is preceded by a stout and conspicuous spine 18 Doisal fin-spines either lacking, or are so nearly concealed in the skin that their presence can be detected by touch only 20 18. Upper teeth with 5 erect cusps; lower teeth with only one cusp, the successive cusps directed outward, forming a nearly continuous horizontal cutting edge all along the jaw Etmopterus princeps, p. 47 Upper and lower teeth are alike in shape 19 19. Upper teeth quadrangular as well as lower teeth, with one cusp directed outward, forming a nearly continuous horizontal cutting edge along each jaw Spiny dogfish, p. 47 Upper and also lower teeth each have 3 to 5 erect, triangular cusps Black dogfish, p. 51 20. First dorsal fin well in advance of pelvic fins; upper teeth noticeably different in shape from lower teeth 21 First dorsal fin stands over posterior part of bases of pelvic fins; upper are teeth similar to lower teeth in shape. Bramble shark, d. 5& 18 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE 21. 22. Lower teeth erect, triangular, their edges serrate Dalatias licha, p. 55 Lower teeth quadrate, the cusp directed outward, forming a nearly continuous horizontal cutting edge; their outer margins deeply notched, the edges smooth ■ 22 Dermal denticles rounded, overlapping, scale-like, entirely concealing the skin (fig. 20) ; each dorsal fin is preceded by a short spine, embedded nearly to its tip in the skin, but recognizable by touch Portuguese shark, p. 52 Dermal denticles conical, only moderately close set, the skin visible between them; dorsal fins not preceded by spines Greenland shark, p . 53 Note. — Not yet known from the Gulf of Maine though reported from Marthas Vineyard. THE SAND SHARKS. FAMILY CARCHARIIDAE Outstanding characteristics of the sand sharks are that they have an anal fin ; the two dorsal fins are without spines and are nearly equal in size; the rear end of the hase of the first dorsal is over or in front of the origin of the pelvic fins; the anal fin is about as large as the dorsals ; the upper lobe of the caudal fin is much longer than the lower, but occupies not more than one-third of the total length of the fish; there are no lateral keels on the caudal peduncle ; the fifth gill openings are farther forward than the origins of the pectoral fins; and the teeth are slender and sharp-pointed. Sand shark Carcharias taurus Rafinesque 1810 Dogfish shark; Ground shark Bigelow and Schroeder, 1948, p. 100. Garman, 1913, pi. 6, figs. 1-3. Description. —The large size of the second dorsal fin, and of the anal as well (which is about equal to the first dorsal instead of much smaller) is of itself enough to distinguish this species from all other Gulf of Maine sharks. The fact that the first dorsal fin is located but little in front of the pelvics, and that the trunk seems crowded with fins of equal size, is a useful field mark. We may also point out that the pectoral fins are not much larger than the other fins — triangular rather than sickle-shaped; that the upper lobe of the tail is nearly one-third as long as head and body together and notched near its tip, with the lower lobe about one-third as long as the upper lobe; and that the head is flat above, the snout short, conical with rather sharp tip. The teeth also (alike in the two jaws) are diagnostic, being long, narrow, sharp-pointed, and smooth-edged, with one (rarely two) small spurs ("denticles") on either side near the base. Size. — Most of the sand sharks that are caught in the northern part of their American range, from Delaware Bay to Cape Cod, are immature, of perhaps 4 to 6 feet. But adults up to 8 or 9 feet long are reported there from time to time, espe- cially from the vicinity of Nantucket, where a commercial shark fishery yielded many of them in Figure 4. — Sand shark (Carcharias taurus), about 40 inches long, Cape Cod; and upper and lower teeth from front part of mouth of a larger specimen from New Jersey, about natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. FISHES OF THE GULF OF MAINE 19 the early 1920's. And large ones, alone, have been reported from North Carolina, southward. The greatest recorded length is 10 feet 5 inches, from southwestern Florida. And the sand shark does not mature sexually until perhaps 7 feet long, or more. A weight of 250 pounds is recorded for one 8 feet 10 inches long, showing how much lighter a fish this is, length for length, than various other sharks. Color. — Light gray-brown above, darkest along back, snout, and upper sides of pectorals, paling on the sides to grayish white on lower surface; sides of trunk rearward from pectorals variously marked with roundish to oval spots, of which there may be upwards of 100, varying in color from yellowish brown to ocher yellow. The rear margins of the fins are edged with black on some specimens, but not on others. Habits and food. — Despite its trim appearance and voracious appetite, this is a comparatively sluggish shark, living mostly on bottom or close to it; more active and taking a bait more freely at night than by day. During its summer visits to the New England coast it holds so close to the coast that it has never been reported from Georges Bank, or from the outer part of the Continental Shelf. Most of those caught are from depths not greater than 1 to 5 fathoms, occasionally perhaps as deep as 10 fathoms, and many come right in to tide line along the beaches. They may sometimes be seen moving slowly to and fro at the surface, over bars, with dorsal and caudal fins showing above the water; and they sometimes enter the mouths of rivers. They capture great numbers of small fish, which are their chief diet, particularly menhaden, dinners, mackerel, skates, silver hake, flounders, alewives, butterfish, and south of Cape Cod, scup, weakfish, and bonito. Sand sharks have been seen surrounding and harrying schools of bluefish ; they have even been known to attack nets full of bluefish, which gives a measure of their voracity. They also eat lobsters, crabs, and squid Breeding. — The eggs of the sand shark are hatched within the parent and are retained there until the resultant young are ready for independent existence, but there is no placental connection between mother and developing embryo. It has recently been discovered that while a ripe female contains a large number of eggs, only two embryos develop as a rule, one in each oviduct; they are nourished (at least largely) by swallowing the unfertilized eggs 30 with which the stomach of the embyro becomes greatly distended. Females with large embryos have so far been reported only from Florida and from Louisiana, whereas others taken near Woods Hole have contained eggs only, making it likely that the small specimens that are so common along southern New England have come from a more southerly birthplace. General range. — Coastal waters on both sides of the Atlantic; Maine to Florida and Brazil in the west; Mediterranean, tropical West Africa, Ca- naries, and Cape Verdes in the east; also South Africa; represented in Argentine waters and in the Indo-Pacific by close relatives. Occurrence in the Gulf of Maine. — The sand shark is by far the most common of its tribe, next to the smooth and spiny dogfishes, along southern New England and at the westerly entrance to the Gulf of Maine. It is plentiful at Woods Hole from June to November, to be found anywhere in that region in shoal waters, even coming up to the wharves. At Nantucket, too, it is so abundant that shark fishing, with the sand shark as the chief objective, is a popular sport. The facts that a catch of about 1,900 sharks by three boats on Horseshoe Shoal, in Nantucket Sound, June to September 1918, was mostly of this species, as was another catch of 350 sharks, taken near Nan- tucket in the early 1920's, illustrate their numbers there. Scattered sand sharks are also caught along the outer beaches of Cape Cod by surf anglers (published records are for Monomoy, Chatham, and Provincetown) and there are enough of them along this stretch of beach in some summers (1951 was a case in point) for them to be a nuisance to anglers casting for striped bass in the surf at night. In August 1947 we saw a large one at the surface pursuing a striped bass, that was being hauled aboard a fishing boat on a hand line, in the eastern side of Cape Cod Bay, where fishermen tell us that this is not an unusual happening. But this appears to be the northern boundary to their occurrence in any numbers, or with regularity. True, they are recorded at Cohasset, on the south- ern shore of Massachusetts Bay, where we caught one about 4 feet long, years ago in Boston Bay, and at Lynn, Mass. But so rarely does it stray north of Cape Ann that it has been reported only M For an account of the embryos, see Springer, Copeia, 1948, No. 3, pp. 153-1 5fi. 20 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE twice from Casco Bay, and once from St. Andrews, New Brunswick, near the mouth of the Bay of Fundy, its most northerly known outpost, where one was taken in a weir in 1913. In New England waters the sand shark occurs only as a summer visitor. The winter home of those that summer along the northeastern United States is not known, nor has any increase been noted in Florida waters (where they are taken at all times of year) coincident with their winter dis- appearance from the northern part of their range. Like various bony fishes they may move offshore, and perhaps southward, to escape winter chilling. Importance. — There were commercial fisheries for the sand shark around Nantucket during the first quarter of the present century, but these were short lived, reputedly because of exhaustion of the local stock. And the sand shark is of no commer- cial importance on the New England coast at present. Westward from Cape Cod it is of some interest to anglers, who catch considerable num- bers, both as objects of special pursuit, for it takes almost any natural bait readily, or incidentally while surf casting for better fish. But it is not plentiful enough in the Gulf of Maine to be worth fishing for. There is no record of attacks by sand sharks on human beings in North American waters, though bathers often come close to them. Our own experi- ence bears this out; in fact, it is looked upon as a harmless nuisance on the New England coast wherever it is plentiful enough to be familiar. But its relative (or relatives) of East Indian waters have a more sinister reputation. MACKEREL SHARKS. FAMILY ISURIDAE Sharks of this family are easily recognizable by the very firm half-moon-shaped (technically lunate) caudal fin, with lower lobe but little shorter than the upper, in combination with large awl-like or blade-shaped teeth, and with gill openings larger than any other Gulf of Maine shark except the basking shark. Their tail fins, in fact, recall the tails of such bony fishes as the mackerel tribe or the swordfish, in outline, likewise in firm tex- ture, hence their common name. The basking shark also has a caudal fin and peduncle of this same sort, but its teeth are minute and very numerous, and its gill openings are so long that those of the two sides nearly meet on the lower surface of the throat. Other diagnostic features are that they have an anal fin ; that their caudal peduncle is expanded as a prominent longitudinal keel on either side; that their dorsal fins are not preceded by spines; and that the inner margins of their gill arches do not have horny gill rakers. Mackerel shark Lamna nasus (Bonnaterre) 1788 Porbeagle; Blue dog (in Gulf of Maine) Bigelow and Sehroeder, 1948, p. 112. Garman, 1911, pi. 6, figs. 4-6 (as Isurus punclatus). This is a stout, heavy-shouldered shark, tapering in front to a pointed conical snout and behind to a very slim tail root. Its dorsal and pectoral fins are large ; the former, originating a little rearward of the armpits of the pectorals, is triangular and about as high as it is long; the pectoral fins are only half as broad as long. The second dorsal and anal fins are very small indeed, and the pelvics but little larger. The second dorsal fin stands over the anal. There is a conspicuous transverse furrow or pit on the upper surface of the root of the tail, also one on the lower surface close in front of the origin of the caudal fin. The lower lobe of the caudal fin is two-thirds to three-fourths as long as the upper lobe, and there is a small secondary keel on the base of the caudal fin on either side, below and behind the rear end of the primary keel formed by the sidewise expansion of the caudal peduncle. The teeth of the porbeagle are alike in the two jaws, slender, pointed, smooth-edged, and with a sharp denticle near the base on each side (young fish may not have these) which the mako lacks (p. 23). The only Gulf of Maine sharks with which the porbeagle might be confused are the maneater (p. 25), or the mako (p. 23). And it is easily told from the former by its slender, smooth-edged teeth, as well as by the position of its second dorsal fin directly over the anal; from the mako by the shape of its teeth (cf. fig. 5 with fig. 6), each usually with a small basal denticle on either side, which the mako lacks; also by its stouter body and by the presence of the secondary longitudinal keel on the anterior part of its caudal fin. FISHES OF THE GULF OF MAINE 21 Figure 5. — Mackerel shark (Lamna nasus), about 37 inches long, Xahant, Massachusetts. Upper and lower first to fifth teeth from center of jaw of a larger specimen from Platts Rank, about 0.7 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. Color. — Dark bluish gray to bluish black above, including the upper surfaces of the pectorals, changing abruptly, low down on the sides, to white below; lower surfaces of pectorals dusky to black on the outer one-half to one-third, more or less mottled while and dark toward their bases, and with the anterior and posterior edges narrowly rimmed with black; the anal is white or slightly dusky. Size. — The common run of mackerel sharks in the Gulf of Maine are from 4 to 6 feet long, with few heavier than 200 pounds; thus 18 recently landed at Portland and East port, Maine,'" averaged 4 feet 5 inches, the largest being about 8 feet long, the smallest 3 feet 7 inches. Specimens longer than 7 to 8 feet are not common; only two longer than 8 feet have been recorded previously from the Gulf of Maine, one of which was 10 feet,32 the largest recorded from either side of the North Atlantic. This shark has been said to reach a length of 12 feet. But the sizes of sharks often are overstated, uidess actually measured, point to point, not around the curve of the body. Information as to the relation- ship between length and weight is restricted to a report of 305 pounds at 8 feet 3 inches, and of about 400 pounds at about 9 feet. One 3 feet long that we measured weighed 20 pounds. » Scattergood, Copela, 1949, No. 1, pp. 71-72. « Hubbs, Copela, No. 1?3, 19?3, p. 101 Habits. — The whole mackerel-shark tribe lead a pelagic life, wandering about over the ocean in pursuit of the fishes on which they prey, and often uniting in small companies, though the, can hardly be called gregarious. Like swordfish they spend much time at the surface on calm days, when their triangular back fins, followed by the tip of the caudal fin (the bluntness of the former and the wavy track of the latter identify the shark as such) may often be seen cutting through the water. We have sailed close to sharks probably of this species again and again, only to see them sound, just out of harpoon range, plainly visible at first but soon fading from sight as they swim downward. The porbeagle has often been described as active and strong swimming. But it puts up only a very feeble resistance when hooked. We have never seen or heard of one jumping, as the mako often does (p. 24), nor is there any difficulty in landing one of 4 to 5 feet on an ordinary cod line. It is, in fact, as proverbial among fishermen for its sluggishness when hooked, as is the mako for its activity. While often seen "finning," many are caught close to the bottom, in depths down to 80 fathoms in the gill net fishery for ground fish that is carried on from Portland, Maine; some also on bottom on cod lines; how much deeper they may descend is not known. 22 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Food. — In the Gulf of Maine the porbeagle feeds chiefly on mackerel and on the herring tribe; on butterfish; on ground fish, as cod, hake, cusk, rosefish, flounders, or other kinds available; and on squid. It has also the annoying custom of foraging on the cod and other fish that have been hooked on long lines and biting off the snoods. It is also known to prey on the spiny dogfish in the eastern Atlantic; probably in the Gulf of Maine also. But we find no record of its eating crustaceans of any kind. Breeding. — The mackerel shark tribe are ovovi- viparous; that is, the eggs are hatched within the maternal oviducts, but there is no placental con- nection between mother and young. The embryos, like those of the sand shark (p. 19), are nourished chiefly by swallowing the unfertilized eggs that lie nearby in the "uterus," and their stomachs become enormously swollen by the masses of yolk that are eaten in this way. Another interesting feature of the porbeagle embryo is that the upper lobe of its caudal fin is much longer at first than the lower lobe, the latter increasing in relative length with growth. The embryos also are very large at birth; young of 18, 19, and 24 inches have, for example, been found in a five-foot mother. Corresponding to their large size, gravid females contain only one to four young (0-2 in each oviduct) . General range. — Continental waters in both sides of the North Atlantic; southern Scandinavia, Orkneys and North Sea southward to the Mediter- ranean and northwest Africa in the east; northern coast of Newfoundland,33 Newfoundland Banks and Gulf of St. Lawrence to New Jersey and per- haps to South Carolina in the west; represented in the northwest Pacific and in Australian-New Zealand waters by forms that are closely allied to it, but not identical. Occurrence in the Gulf of Maine. — It has been known from the days of the earliest settlement that stout-shouldered, surface-swimming sharks of moderate size, with "mackerel" tails and slen- der, smooth-edged teeth are tolerably common in the Gulf of Maine; they are universally referred to by the fishing popidation as "mackerel sharks." During the first half of the last century only one such shark species was recognized in our waters. And while more recent researches have proved " One reported at Raleigh, on the Newfoundland side of the Strait of Belle Isle, July 1929, by Dr. W. Q. Jeffers. that two actually occur within the limits of the Gulf (this and the next described) the present species is the more northerly of the pair, and much the more frequently taken in the Gulf. Hence it is probable that most of the mackerel sharks that fishermen often see swimming lazily on the surface, and often catch, off the shores of northern New England, belong here. Seemingly, the chief centers of population for the porbeagle in the western Atlantic are along outer Nova Scotia, and in the western side of the Gulf of Maine. Thus, while there are but two published records for it from the Newfoundland Banks, and one (besides verbal reports) in the Gulf of St. Lawrence, fishermen report it as the commonest large shark along the Atlantic coast of Nova Scotia. Apparently it tends to shun the cold waters of the Bay of Fundy, for it is recorded only twice from Passamaquoddy Bay, one in August 1900, the other on October 3, 1935.34 But it is so plentiful farther west in the Gulf that inci- dental catches are on record of 19 that were taken in one night by six men on hand lines, and of about 150 taken by one crew during three weeks' cod fishing near Monhegan Island, Maine. We have ourselves hooked or sighted about one per three or four days' fishing, on the cod grounds in general in the western side of the Gulf, the majority near Platts Bank off Cape Elizabeth, but some also on Nantucket Shoals.35 Certainly it is the most often seen of the larger sharks around the Isles of Shoals and near Cape Ann, and it has been characterized repeatedly as "common" in Massachusetts Bay.36 To the westward the porbeagle is described as not uncommon near Woods Hole (we have not seen it there). We saw a small one about 3 feet long taken in an otter trawl at 60 fathoms, off Marthas Vineyard, on February 20, 1950, by the Eugene H; and it has been reported on several occasions from Rhode Island waters. But it ap- pears only as a stray off New York and to the southward. Thus, the latitudinal range within which it occurs regularly off the American coast covers only something like 5°. And its on- and offshore range is correspondingly so narrow that no report » Reported by McGonigle and Smith, Proc. Nova Scotia Inst. Sci., vol. 19, 1936, p. 180. 85 Cod tagging cruises of the IT. S. Bureau of Fisheries. 18 Actually no sharks other than the spiny dogfish (p. 47) are "common" In the Gulf of Maine, In the sense that this term is applied to such fish as herring, cod, mackerel, and other species, but only as relative to other sharks of corresponding sizes. FISHES OF THE GULF OF MAINE 23 of it has come to hand from Georges or Browns Banks, only one from the Nova Scotia slope off Sable Island, and two from the Grand Banks, as just noted. On the other hand, few come in-shore close enough to be picked up in pound nets or weirs. All published records of mackerel sharks from the Gulf, and all that we have seen there, have been in the warm half of the year, and something like 70 percent of the landings of porbeagles on the coast of Maine are for August to November. But its presence in the Gulf in winter is proved by our receipt of a photograph of a porbeagle embryo, taken from a female caught in January, off Portland, Maine, in 1927. And it is also caught in winter as well as in summer in north European waters. Apparently it simply descends into deeper water during the winter to escape low surface temperatures, feeding little, else more of them would have been caught in the Gulf during the winter fishery with long lines for hake (Uro- phycis). In the Gulf of Maine, females containing em- bryos have been taken in August (near Monhcgan Island, Maine) ; in October (off Barnstable, Mass.) ; in November (off Portland, Maine); and in Jan- uary (off Portland, Maine). But the fact that the largest embryos have been found in European seas in summer suggests that most of the young are not born until then. Importance. — The liver oil of the porbeagle, mixed with other fish oils, was in demand for use in tanning leather during the first quarter of the 19th century. And it is interesting to read that as much as 1 1 gallons of oil has been obtained from the liver of a single shark 9 feet long. This demand had almost entirely died before 1850 and has never revived. But a new demand has developed of late years for porbeagle meat, which resembles swordfish in taste as well as in appearance, resulting in landings for this purpose of about 46,000 pounds in 1944 on the coast of Maine, and of 71,600 pounds in 1945. Assuming an average weight of, say, 50 pounds, this corre- sponds to a commercial catch of about 900 to 1,400 sharks. There is no special fishery for porbeagles at present in the Gulf of Maine, or for any other sharks for that matter. About four-fifths of those brought in are taken in gill nets set on bottom for ground fish, and most of the sharks caught in this way are landed in Portland, Maine. The re- mainder are taken by seines, traps, weirs, hook and line or harpoons. And most of the porbeagles taken in these ways are discarded at sea.37 The porbeagle is not "game" enough to be of any in- terest to sport-anglers. Sharp-nosed mackerel shark Isurus oxyrinchus Rafinesque 1810 Atlantic mako Bigelow and Schroeder, 1948, p. 124. Description. — This shark resembles the common mackerel shark so closely that we need merely point out the points of difference. Most obvious of these is that while the first dorsal originates about above the armpits of the pectorals in the common mackerel shark, it stands over or behind the inner corner of the pectoral in the mako, and that the second dorsal originates a short distance in front of the anal. The teeth, too, differ rather noticeably in appearance, for while of the same awl-like type, those of the mako lack the lateral spurs or denticles that are characteristic of all but the smallest porbeagles, and those in the front part of the mouth are conspicuously flexuous in form. The mako, too, is more slender bodied; its snout is more narrowly conical; its upper and lower caudal lobes are more nearly equal hi length; and the forward part of its caudal fin lacks the secondary lateral keels that are to be seen on the caudal fin of the porbeagle (c/. fig. 6 with fig. 5). Color. — Deep blue-gray above when fresh- caught, appearing cobalt or ultramarine in the water, with gradual transition along the sides to snow-white below; but turning dark slate gray above soon after death (especially if preserved), and to bluish white or pale dirty gray below and on the lower surfaces of the pectorals. Size. — The maximum length reported for a spec- imen of the Atlantic mako that was actually meas- ured is about 12 feet,38 though it has been said to grow to 13 feet. The largest western Atlantic specimen of which we find definite record, taken off St. Petersburg, Fla., was 10 feet 6 inches long, and one nearly as large (10 ft. 2 in.) was caught off New York Harbor many years ago. But the com- mon run caught off the middle Atlantic United 17 See Scattergood, Copela, 1949, p. 70, for further details as to landings In Maine and methods of capture. 11 3.7 meters ns calculated from the size of Its Jaws. 24 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 6. — Sharp-nosed mackerel shark, or Mako (Isurus oxyrinchus), about 64^2 inches long, Maryland. Below, teeth in front of mouth of a large specimen, Cape Cod. From Bigelow and Schroeder. Drawings by E. N. Fischer. States are perhaps 5 to 8 feet long. Males of about 6 feet are sexually mature (as indicated by the claspers). Kecorded weights at different lengths are about 135 pounds at 6 feet, 230 pounds at 7 feet 8 inches; and about 300 pounds at 8 feet. The heaviest Atlantic mako caught on rod and reel of which we have found record was one of 786 pounds taken off Bimini, Bahamas, by Ernest Hemingway in 1936; the largest Pacific mako one of 798 pounds, taken by E. White-Wickham off New Zealand.39 Habits. — This is one of the most active and swift swimming of the sharks. In seas where it is more common than it is m our Gulf, it is often seen swimming at the surface, and it is famous for its habit of leaping clear of the water, not only when hooked, but under natural conditions. Seemingly it preys chiefly on schools of smaller fishes of the mackerel and herring tribes. But it also attacks larger fishes. A 730-pound mako, for example, that was harpooned near Bimini in the Bahamas, contained a 120-pound swordfish (Xiphias gladius) almost entire, while one weighing about 800 pounds, harpooned off Montauk, Long Island, was seen attacking a swordfish, and was 39 A South African shark of 2,176 pounds, landed on rod and reel, and re- ported as a mako, Is proved by the photograph of Its teeth (London lUus. News, July 14, 1928, p. 53) to have been a maneater (Carcharodon) . found when landed to contain a large amount of its flesh.40 Young embryos of the mako, like those of the porbeagle (p. 22), have greatly dilated stomachs, being nourished on the unfertilized eggs that he near them in the oviducts, and they are very large at birth, relative to the size of the mother. General range. — This is an oceanic shark, of the tropical and warm-temperate belts of the Atlantic north and south, including the Mediterranean in the east and the Caribbean and Gulf of Mexico in the west. It is represented in the corresponding thermal belts of the Pacific and Indian Oceans by a close ally, the Pacific mako Isurus glaucus. Occurrence in the 6ulf of Maine. — The center of abundance for the mako lies in warmer seas to the southward of our Gulf. Considerable num- bers journey northward, however, in summer along the continental shelf, as far as to the offing of southern New England, and a few are caughl off Woods Hole. One of the earliest accounts of it in American waters was based partly on one from Cape Cod. During the past few summers we have heard repeatedly of makos seen jumping, or occasionally hooked near the northern end of «• See Farrington (Field and Stream, vol. 47, Feb. 1943) for these Instances of the mako attacking swordfish, and for other Interesting notes on this shark. FISHES OF THE GULF OF MAINE 25 Cape Cod, and in the summer of 1941 one about six feet long was landed on rod and reel in the southern side of Massachusetts Bay near Plym- outh.41 Thus stray individuals may be expected to visit the southern part of the Gulf in most summers, though we have never met it there ourselves. It has even been reported as far north as Seguin Island, Maine, but without convincing evidence that the shark in question was not a porbeagle.42 Importance. — The chief importance of the At- lantic mako, as of its Indo-Pacific relative, is as a game fish, because of its fast runs when hooked and of its habit of leaping. But it is not plentiful enough anywhere in the Gulf of Maine to be worth fishing for there especially. Maneater Carcharodon carcharias (Linnaeus) 1758 White shark Bigelow and Schroeder, 1948, p. 134. Garman, 1913, PI. 5, figs. 5-9. Description.— The maneater is of the general "mackerel shark" appearance, with firm lunate tail, the upper lobe only a little longer than the lower; and with triangular first dorsal of moderate size originating over the armpits of the pectorals, which are sickle shaped, and roughly twice as long as they are broad. The second dorsal and anal fins are very small, the former a little in advance of the latter; and the root of the tail 'I Information from Dr. W. J. Mlxter. *» Various early reports of It In the northern part of the Gulf seem to have referred, actually, to the porbeagle. bears a single well-marked keel on either side. The snout is conical, moderately pointed. Unfortunately, there is no obvious field mark to distinguish a small maneater from a large porbeagle or from a large mako when seen swim- ming at any distance. Once captured, however, no confusion could arise, for instead of the slim catlike teeth of the porbeagle and of the mako, we find the maneater one of the best armed of all sharks; its teeth large and triangular, and similar in shape in the two jaws, except broadest in the upper, with nearly straight cutting edges and strongly serrated margins. As a precaution, any large active shark, upwards of 10 or 12 feet long, with the tail not long, out of ordinary proportions, should be looked upon with suspicion, for it might prove to be a maneater. If it were sluggish, resting with the dorsal fin high out of water, it would be no doubt a harmless basking shark (p. 28). Color. — Maneaters up to 12 to 15 feet long are slaty brown or leaden gray above, sometimes almost black, shading more or less abruptly on the sides to dirty white below. There is a black spot in the armpit of each pectoral fin, and the lower surfaces of the pectorals are black toward their tips, usually with some black spots adjacent. The pelvics are white below, but olive along their anterior edges. Larger specimens (we have seen none) have been described as dun colored above or very pale leaden, and they may lack the black spot at the armpit of the pectoral fin.43 >' Information from Stewart Springer, from large Florida specimens. ' Figure 7. — -Maneater (Carcharodon carcharias), Massachusetts, about 7 feet long. A, first three upper and B, first three lower teeth, from center of jaw, from a specimen about 8% feet long, Woods Hole, about 0.6 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. 210941—53 8 26 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Size. — This is one of the largest of sharks. A gulf of Maine specimen about 3 feet long is the smallest, apart from embryos, that has been seen; one of about 5 feet the next smallest. So far as known it does not mature sexually until it has grown to a length of 12 to 14 feet. Among larger ones, from one place or another, the exact measure- ments for which have been reported, four have been between 14 and 16 feet long, two between 16 and 18 feet, and three between 19 and 21 feet. The largest on record was 36 K feet long; u the next largest about 30 feet, but perhaps not measured exactly. Maneaters of a given length may vary widely in weight, because of variations in their condition. Thus one specimen 8 feet 2 inches long weighed only 342 pounds, but another of 8 feet 3 inches, weighed 600 pounds. Five, weighing between 910 and 1,000 pounds ranged from 9 feet 8 inches in length to 12 feet 6 inches. Three, of 13 to 13% feet, weighed 1,291 to 1,344 pounds, but another, from South Africa of 13 feet 3 inches scaled 2,176 pounds, doubtless a very fat fish. A 15-foot 2-inch specimen weighed 1,720 pounds; and one of 21 feet, the largest that has been weighed so far, 7,100 pounds, its liver 1,005 pounds.46 Habits.- — So few maneaters are seen that little is known of their way of life, apart from their vorac- ity. Most of the records of them have been of specimens taken at or near the surface, and such specimens as visit our Gulf sometimes come very close inshore. Thus two specimens were seined close in, off Swampscott, at the northern entrance to Boston Harbor in 1939; one was harpooned in 1937 about 2 miles off Nantasket Beach, one of the most popular bathing resorts near Boston; another was harpooned about one-half mile off Cohasset, Mass., where the water is not over 20 feet deep; one in 10 feet of water in Provincetown Harbor, many years ago. Some have even been taken in fish traps close to the beach on Cape Cod and near Woods Hole; and in 1916 one was taken in the shallow water of Sandy Hook Bay, N. Y. On the other hand, the largest one that has been weighed yet was caught on a set line off the north coast of Cuba, at a depth of about 700 fathoms. Nothing is known of its breeding habits, beyond "This Australian specimen, the jaws of which are in the British Museum, is the basis for repeated statements that the maneater grows to 40 feet. *' For further details, see Blgelow and Schroeder, Fishes Western North Atlantic. Pt. 1, 1948, pp. 137-138. the bare facts that it is ovoviviparous like others of the mackerel shark tribe. The maneater is one of the most voracious of all the fish tribe, feeding indifferently on large prey and on small. Other sharks, 4 to 7 feet long and practically intact, have been found repeatedly in maneaters' stomachs; and a young sea lion of 100 pounds in one on the coast of California, while seals, sturgeons, and tuna have been found in maneaters no longer than 8 to 9 feet. In southern seas they are described as feeding regularly on sea turtles. But they also devour smaller fishes of whatever kinds are available, including small sharks and chimaeroids, also squids. When they come in on the fishing banks, they are known to take fish that they find hooked on long lines as porbeagles do (p. 22). Thus the mouth of one of 9 feet 8 inches, taken near Cohasset, Mass., and examined by us, carried several hooks with the snoods still attached, while its stomach contained a spiny dogfish (Squalus acanthias) that evidently had been torn off a hook. And a large Florida maneater, caught on a set line, contained 2 brown sharks (Carcharhinus milberti), 6 to 7 feet long, that had evidently been torn from hooks on the same set line on which the maneater was hooked. The maneater, like the Tiger shark, is not above feed- ing on slaughterhouse waste or other garbage. General range. — This is an oceanic shark, widespread in the tropical and warm temperate belts of all oceans, including the Mediterranean. In the western side of the Atlantic it has been recorded as far north as St. Pierre Bank south of Newfoundland, and as far south as Brazil." Occurrence in the Gulf of Maine. — The maneater is usually looked on as a warm water shark, doubtless correctly so. None the less, it has been reliably reported from the southwestern part of the Gulf of Maine more often than it has from any other coastal sector of comparable length on the Atlantic coast of North America. At least 10, for example, were actually captured or were harpooned and lost in Massachusetts Bay alone during the period 1935 to 1948. We ourselves examined three of these, one that was netted at Swampscott; a female of 9 feet 8 inches weighing 980 pounds that was harpooned within half a mile of the land off Cohassett, in August 1940; one of about 3 feet, that was harpooned in July 1948 *• For details and references, see Blgelow and Schroeder, Fishes Western North Atlantic. Pt. 1, 1948, pp. 140-141. FISHES OF THE GULF OF MAINE 27 near Boston Lightship, this last being the smallest that is on record to date (p. 26), and one about 14 feet long, weighing 1,050 pounds dressed, which sold for 10 cents a pound, was taken in a trap at North Truro on November 9, 1952. Carrying the record back to earlier years, a 15-foot shark, taken at Monomoy Point at the elbow of Cape Cod in the autumn of 1928, appears to have been a maneater, and one of about 16 feet, taken in a trap at East Brewster, October 16, 1923, and identified by Dr. Samuel Garman, certainly was, while one of 7 feet 2 inches, taken in Massachusetts Bay, about 1910, was the basis of Garman's (1913, pi. 5, fig. 5) beautiful illus- tration. Earlier still, a 13-footer, taken at Provincetown, Cape Cod, in June 1848, was described by Storer as a new species, atwoodi, while two small ones were mentioned by him as taken by Massachusetts fishermen between 1820 and 1850. And Capt. Atwood reported seeing four, caught in mackerel nets at Province I own man}7 years ago.47 Proceeding northward, we find scattered records from the vicinity of Portland, Maine, most re- cently, a 13-footer caught in a gill net off Casco Bay in November 1931; one from Eastport, Maine, many years ago; a very large one (esti- mated as about 26 feet long) taken in a wier at Campobello Island, November 23, 1932 *8 it was suggested locally that it may have been the same specimen that had attacked a fishing boat off Digby, Nova Scotia, the preceding July (p. 27); one from Deer Island, New Brunswick, taken in a herring weir, August 24, 1949;49 and one from Digby, on the Nova Scotian shore of the Bay of Fund}'', July 2, 1932. And there arc several re- liable records for St. Margaret Bay on the outer coast of Nova Scotia, perhaps also for Halifax. The most northerly positive record for it on tho Atlantic coast of North America is for St. Pierre Bank, south of Newfoundland, where one attacked a fisherman in a dory many years ago, leaving in the sides of tho boat pieces of its teeth, from which Dr. Garman was able to identify it.60 Westward and southward from the elbow of Cape Cod, we find nine or ten definite records for Nantucket and for the vicinity of Woods Hole «' Putnam. Bull. Essex Inst., vol. 6, 1874, p. 72. « Piers, Proc. Nova Scotian Inst. Set., vol. 18, 1934, p. 198. 48 A female 12 feet, 8 inches long, weighing 1,299 pounds, reported by Scatter- good, Trefethen, and Coffin, Copela, 1951, p. 298. » Putnam, Bull. Esses Inst., Salem, vol. 6, 1874, p. 72. (never more than two in any one year), with one of five feet (second smallest on record) netted at Sakonnet, Rhode Island, May 30, 1939. Maneat- ers are also reported occasionally near New York, notably one of about seven feet, taken in Sandy Hook Bay, July 1916, to which we recur below (p. 27). Relation to man. — So few man-eaters visit our Gulf that they would deserve only the briefest mention were this not the only shark that is ever likely to attack human beings there. Strong and active, equipped as it is with a most terribly effective set of cutting teeth, it has borne an un- savory reputation as a man-eater from the earliest times, and it is probable that the 7-foot specimen listed earlier from South Amboy, Sandy Hook Bay, was the cause of the shark fatalities along the New Jersey beach in July 1916 (p. 16). A fatal attack on a swimmer at Mattapoisett, on Buzzards Bay, on July 25, 1936, may also have been by a man-eater, though in this case the shark was driven away without being identified. This is also perhaps the only shark against which unprovoked attacks on small boats are proved by identification of their teeth, embedded in the wood. One such instance, from the Newfound- land Banks, was reported by Putnam 61 many years ago (p. 27). A recent local case is of a very large one that attacked a fishing boat in the Bay of Fundy off Digby Gut, Nova Scotia, July 2, 1932 and left in her keel or lower planking several of its teeth, by which it was identified.62 Storer M wrote of a case where one (apparently the 13-foot specimen that ho had described earlier as atwoodi) turned furiously on a boat, but was lanced to death anil brought into Provincetown. And a 15-foot shark, probably this species to judge from the il- lustration of it that was published,61 that was killed off Monomoy Point by two fishermen in November 192S, overturned their dory before it was subdued. And one of about 15 feet (similarly identified by teeth left in the planking) attacked a boat, from which it had been harpooned, in St. Margaret's Bay, Nova Scotia, on June 27, 1920.66 Hence, so long as maneaters wander within " Proc. Essex Inst. Salem, vol. 6, 1874, p. 72; teeth Identified by Dr. S. Garman. « Reported by Piers, Proc. Nova Scotia Inst. Scl., vol. 18, 1934, p. 198. « Fishes of Mass., 1867, p. 248. « Reported In Witman and Lee Co.'s Market Letter for Nov. 8, 1928; called to our attention by Dr. Lewis Radcllffe of the U. S. Bureau of Fisheries. •» For details of this occurrence, see Piers, Proc. Nova Scotia Inst. ScL, vol. 18, 1934, pp. 19G-19S. 28 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE our limits more often than had been realized pre- viously, the possibility is always open of attacks on bathers along the Massachusetts shores of the Gulf. Despite its ferocity, muscular strength and size, the man-eater does not put up so spectacular a re- sistance when hooked as does a mako, neither running so fast nor having the habit of jumping. Neither does it put up as strong a fight, pound for pound, as a tuna ordinarily does, or any of the swordfish tribe. Thus a 1,329-pound maneater was landed on rod and reel by an Australian angler in 53 minutes. One of 2,176 pounds, caught from the shore in South Africa, is the largest fish ever landed on rod and reel that has come to our notice.66 M London Illus. News, July 14, 1928, p. 53; photograph recorded as a mako but shown by its teeth to have been a maneater. BASKING SHARKS. FAMILY CETORHINIDAE Basking shark Cetorhinus maximus (Gunnerus) 1765 Bone shark Bigelow and Schroeder, 1948, p. 147. The basking shark resembles the mackerel sharks in the lunate shape of its caudal fin, with lower lobe nearly as long as upper; also in the presence of a noticeable lunate furrow above and one below on the root of the tail, and in the wide lateral expansion of the latter, forming a pro- nounced "fore and aft" keel on either side; also in the facts that the second dorsal fin and the anal fin are much smaller than the first dorsal, that its fifth gill opening is situated in front of the origin of the pectoral fin; in the position of the mouth on the under side of the head ; and in the wide separa- tion of the nostrils from the mouth. But the teeth of the basking shark are minute and very numer- ous (large and few in number in the mackerel sharks); its gill openings are so large that they extend right around the neck, with those of the first pair almost meeting below on the throat ; and the inner margin of each gill arch bears a great number of horny, bristle-like rakers, directed inward-forward, that correspond to the rakers of various bony fishes in their position and in their function (see p. 30). It was the fancied resem- blance of these rakers to the whalebone of the whalebone whales that suggested the vernacular name "bone shark" to the whalemen of olden times. Corresponding to its feeding habits, the mouth of the basking shark is very large and widely dis- tensible at the corners. The snout is short, conical, with rounded tip on large specimens. But it is much longer, relatively, on small ones, Figure 8. — Basking shark {Cetorhinus maximus), 26}4-foot female, Marthas Vineyard. A, side view of head of 12-foot Long Island specimen; B, a group of the teeth of same, about 1.2 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. FISHES OF THE GULF OF MAINE 29 projecting far beyond the mouth, obliquely trun- cate in front, terminating above in a sharp point, and with the head strongly compressed sideways abreast of the front of the mouth. This results in so bizarre an appearance that the young basking shark was thought at first to represent a separate species. A gradual transition takes place from the juvenile shape of head to the adult shape when a length of 12 to 16 feet has been reached. We need only note further that the triangular first dorsal fin stands midway between pectorals and pelvics; though not so high in proportion as that of the mackerel-shark tribe, it rises high in the air when a large basking shark lies awash on the sur- face, as is their habit, a convenient field mark (p. 29). Color. — Upper surface grayish brown, slaty gray, or even almost black. The lower surface has been described repeatedly as white. But the Menemsha specimen described by Allen w was of a somewhat lighter shade below than above, without white markings, as was a Massachusetts Bay specimen recently examined by us; while one 14 feet long captured at West Hampton, L. I., on June 29, 1915 68 had the belly as dark as the back, with a white patch underneath the snout in front of the mouth. Size.- — The basking shark rivals, though it does not equal, the whale shark of tropical seas in size. Reports that an occasional basking shark may reach a length of 50 feet probably aro not an exaggeration, for the catch on the coast of Norway, for the period 1884 to 1905, included one of about 45 feet and three of about 40 feet, with the six next longest ranging between 36 feet and 30 feet 3 inches. The three longest for which we find definite measurements for the western Atlantic were of 32 feet 2 inches, 32 feet, and 30 feet 3 inches. But others up to 35 feet long have been credibly reported as killed near Eastport, Maine, many years ago; and one captured at Musquash Harbor, New Brunswick, near the mouth of the Bay of Fundy in 1851 was said to have been about 40 feet long. It is probable that they are at least 5 to 6 feet long when born, the three smallest so far reported having been between 5 feet 5 inches and about 8 feet 6 inches long. Matthews 69 concluded from studies of basking sharks taken near the Isle of Skye that fish up to 10 feet are in their first year, those of 15 feet in their second year. Males mature sexually at about 18 to 20 feet as indicated by the lengths of their claspers, females at about 20 to 23 feet; i. e., when 3 years old or perhaps 4, according to Mathews' estimate. We find no exact weights for large basking sharks from the Atlantic. But 6,580 pounds for one of 28 feet, and 8,600 pounds for another of 30 feet, from Monterey, Calif., is doubtless a fair indication of what a fairly large one may be expected to weigh. Estimated weights for smaller ones, from the Pacific, are about 6,600 pounds at about 23 feet, 1,000 to 1,800 pounds at 13 to 15 feet, and 800 pounds at 8 feet 4 inches.60 A young one, 12 feet long, killed off Digby, Nova Scotia, August 16, 1939, weighed 359 pounds, after it had bled, 61 and one almost 20 feet long, taken off Portland, Maine, in 1936, weighed 550 pounds, dressed. Habits. — This is a sluggish, inoffensive fish, help- less of attack so far as its minute teeth are con- cerned. It spends much time sunning itself at the surface of the water, often lying with its back awash and dorsal fin high out of water, or on its side, or even on its back sunning its belly; some- times it loafs along with the snout out of water, the mouth open, gathering its provender of plank- ton. They pay so little attention to boats that it is easy to approach one of them within harpoon range, and excellent motion pictures have beea taken of them in Irish waters.62 But they have also been seen jumping, perhaps to shake off para- sites. Those seen in the Gulf of Maine are usually traveling singly. But they are known to congre- gate sometimes in loose schools which may include as many as 60 to 100 in the peak years of abun- dance for them in regions where they are more numerous than in the Gulf of Maine.63 It is chiefly during the warm half of the year that basking sharks are encountered off the northeast- ern United States and in the northern part of their range in the opposite side of the Atlantic. It is likely that those that summer in the inshore parts of the Gulf simply withdraw in the fall, to pass the " Bull. Boston Soc. Nat. Hist.. No. 24, March 1921, p. 5. » Described by nussakof, Copeia, No. 21, 1915, pp. 25-27. » Phllos. Trans. Roy. Soc. London, Ser. B., vol. 234, 1950, pp. 217-316. » For further details as to slics of basking sharks, see Bigelow and Schroeder, Fishes, Western North Atlantic, Pt. 1, 1948, pp. 151-152. •' Referred to by McKcnzte, Proc. Nova Scotia Hist. Set., vol. 20, 1940, p. 42. 81 Shown in the film "Men of Arran." « See Bigelow and Schroeder, Fishes Western North Atlantic, Pt. 1, 1948, pp. 153, 154, for details as to their centers of population and secular fluctuations in abundance in north European waters. 30 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE winter in deeper water where the temperature does not fall so low. Next to its vast bulk and its curiously sluggish habit, the most interesting peculiarity of the bask- ing shark is its diet, for it subsists wholly on tiny pelagic animals, which it sifts out of the water by means of its greatly developed gill rakers, exactly as plankton-feeders among fishes such as men- haden do, and whalebone whales with their baleen sieves. In several cases their stomachs have been found packed with minute Crustacea; this was true of the only western Atlantic specimen of which the stomach contents have been examined. And while digestion is so rapid that the food swallowed is soon reduced to a soupy mass, this usually is reddish, suggesting a crustacean origin. All that is known of the breeding of the basking shark is that the structure of the internal sex or- gans of the female accords with the nourishment of the embryo within the maternal oviduct, that the ovary of a female, with empty oviduct contained something like 6 million immature ova instead of the few that are usual in sharks that bear "living" young, and that an embryo about a foot long was said, long ago, to have been taken from its mother.64 Basking sharks reported as "sea serpents" or as other "monsters". — The remains of basking sharks have been reported as "sea serpents" on several occasions; nor is this astonishing. "As the carcass of the shark rots on the shore, or is buffeted against the rocks, the whole of the gristly skeleton of the jaws and gill arches ... as well as the pectoral and pelvic fins, is soon washed away,"65 leaving only the cranium and the long backbone, with larger or smaller amounts of muscle, so frayed out as to suggest a hairy or bristly mane. As a recent instance from the Gulf of Maine, we may cite the newspaper and radio publicity, that was given, as a supposed sea serpent, to a basking shark skeleton, about 25 feet long, that beached near Provincetown on the outer shore of Cape Cod, in January 1937, that we examined.66 A more spectacular instance of the fanciful in- terpretation that is likely to be placed on any large stranded carcass that has decayed partially, was the famous "Animal of Stronsa," that came « See Matthews, Phllos. Trans. Roy. Son. London, Ser. B, No. 612, vol. 234, 1950, pp. 347—366 for detailed account. u Norman and Fraser, Giant Fishes, Whales and Dolphins, 1937, p. 21. M For account and photograph, see Schroeder, New England Naturalist, No. 2, 1939, p. 1. ashore on the island of that name in the Orkneys, in September 1808. It was pictured by an eye- witness as having three pairs of limbs, but the published illustration of its cranium, vertebrae, and pelvic skeleton 67 show that it was only the remains of some very large shark, probably a basking shark. It has also been suggested repeat- edly that some of the stories of sea monsters of one sort or another may have been based on the dorsal and caudal fins of two or more basking sharks, swimming one behind another as they often do (we dare not touch further on the contro- versial subject of the "sea serpent"). General range. — This enormous fish, formerly thought to be an Arctic species, straying south- ward, is now known to be an inhabitant of the temperate-boreal zone of the North Atlantic.68 It is represented in the corresponding thermal belts of the South Atlantic and of the North and South Pacific by a similar great shark (or sharks), whose exact relationship to the basking shark of the North Atlantic is still an open question. The northern boundary of the normal range of the basking shark of the North Atlantic appears to follow the line of transition from waters of predominately Atlantic influence to those of Arctic origin. This, roughly, runs from the outer coast of Nova Scotia (1 record), and from southern Newfoundland (4 positive records) to western and southern Iceland, to the Orkney and Faroe Islands, and skirts the Norwegian coast to the North Cape, while basking sharks stray now and then to the Murman coast. To the southward, in the North Atlantic, they range as far as the Mediter- ranean and Morocco in the east, to North Carolina in the west. Occurrence in the Gulf of Maine. — -Before the coming of the white man this great shark seems to have been a regular inhabitant of the southern part of the Gulf of Maine. And tradition has it that large numbers were taken in Massachusetts waters, especially off the tip of Cape Cod, during the first half of the eighteenth century, for their liver oil which was then in demand for illuminating purposes. However, the local stock seems soon to have gone the same way as the local stock of the North Atlantic right whale; that is, into the try pot. And basking sharks seem never to have •' Barclay, Mem. Wernerlan Soc, Edinburgh, vol. 1, 1811, p. 418. •> It has long been realized that old tales of a tremendous whale-eating shark, on which Fabriclus based his statement that the basking shark occurs in Greenland waters, were fiction. FISHES OF THE GULF OF MAINE 31 visited the northeastern part of the Gulf in any numbers, there being only a few records for the vicinity of Eastport, Maine, and three from within the Bay of Fundy. At the present time the Gulf appears to harbor a sparse and fluctuating popu- lation, occasional members of which are encoun- tered from time to time, here or there, but whether as immigrants into the Gulf from the open ocean is not known. The list of specimens, the capture or stranding of which in the Gulf has come to our attention for the period 1908-1951 is as follows: 1908. One, 18 feet long, near Provineetown, taken in a fish trap; measured by J. Henry Blake. 1909. One, about 22 feet, in Provineetown Harbor. 1913. One, about 29 feet, Provineetown. 1925. One, about 29 feet, near Monhegan Island, Maine. 1931. Female, 12J4 feet long, York Harbor, Maine. 1934. One, 29 feet, Whale Cove, Grand Manan Island, and one, 28 feet, Back Bay, Bay of Fundy.69 1936. Two off Portland, Maine; the first about 20 feet long, weighing 550 pounds dressed, about May 1 ; the second, much larger (reported as of about 40 ft.), August 2. 1939. Skeleton of one of about 25 feet, examined by us, found on the beach near Provineetown in January. One of about 25 feet, Yarmouth, Nova Scotia. One of 12 feet, Bay of Fundy off Digby Gut.70 1947. Female, about 13 feet long, examined by us, har- pooned by W. T. Reid 3rd, near Boston Lightship, August 5th. 1949. A small one (size not recorded), near Rockport, Mass., September; identified from a good photograph by Miss D. E. Snyder of the Peabody Museum, Salem. 1951. One, 12 feet, near Bar Harbor, Maine, harpooned July 28.71 Occasional basking sharks also visit the shores of the southern coast of Massachusetts, westward from Cape Cod; one, for example, 12 to 14 feet long was taken at Alenemsha on Marthas Vine- yard, August 16, 1916; another of 20 feet 6 inches at that same locality on June 24, 1920;72 one 20 feet 2 inches long was stranded in Hadleys Harbor, Naushon Island, July 1937; and one of 8 feet (among the smallest on record) was taken in a fish trap near Woods Hole on June 15, 1948. ® McKenzie, Proc. Nova Scotia Inst. Scl., vol. 20, 1939, p. 14. '• McKcnzie, Proc. Nova Scotia Inst. Scl., vol. 20, 1939, p. 14. 'i Personal communication from J. W. Burger. 78 This specimen, mounted, In the New England Museum of Science and described by Allen (Bull., Boston Soc. Nat. Hist., No. 24, March 1921, pp. 3-10), served as chief basis for the Illustration (riven hereof the adult basking shark. Probably the basking shark is no more plentiful near shore in our Gulf in most years than the paucity of the recent records suggest, for popular interest in sharks is now so keen, as represented by newspaper publicity given to any unusual capture, that any well-grown one is apt to be seen in these frequented and hard-fished waters. We do not find evidence of any considerable incursion by them into coastal waters farther west since 1878, when 20, at least, were found dead in the fish traps near Woods Hole during the summer. And the only report that might be based on the basking shark on the offshore fishing banks that we have received from fishermen has been of a number of unusually large sharks of some sort, seen by Capt. Henry Klimm on the southeast part of Georges Bank during late June and early July 1947. Importance. — The clay of any regular fishery for the basking shark is long since past in New England waters, probably never to return. And no use is made there, nowadays, of the occasional specimens that are captured. But it may be of interest to point out that it was always hunted of old by the sperm whalers from New Bedford, for its liver oil was considered nearly or as good as sperm oil for illuminating purposes. Basking sharks are still the object of intermittent small vessel fisheries off the coast of Iceland, around the Orkneys, off western Ireland, and off southern Norway; also off Ecuador and Peru in the Pacific. And increasing numbers have been landed during the past few years in northern California, where they are considerably more plentiful than they are in the Gulf of Maine,73 for fish meal and for the liver oil. The yield of oil per fish varies from about 80 gallons to about 200, occasionally to 400 gallons, with as much as 600 gallons reported. The liver of a 30-foot fish weighing 6,5S0 pounds, taken off Monterey, Calif., had a fiver weighing 1,800 pounds, 60 percent of which was oil.74 But, sad to sa3r, it is very low in vitamin A. The fishery, wherever carried on, is by harpoon. And basking sharks are so sluggish and so un- suspicious of a boat, large or small, that it usually is a simple matter to harpoon one that is seen at » According to MacGlnltie (Science, N. Scr., vol. 73, 1931, p. 496), 21 basking sharks were landed in Monterey, Calif., between November 22, 1930 and February, 1931. '< MacQlnltie, Science, N. Ser., vol. 73, May 1931, p. 496. 32 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE the surface. Once struck, however, a large one is likely to put up an astonishingly active and enduring resistance. We read, for example, of one of 35 to 38 feet harpooned by Capt. N. E. Atwood off Provincetown, Mass., about 1863, that towed the fishing smack all night, and broke loose finally.76 '• Goode, Fish. Ind. U. S., 1884, Sect. 1, p. ( THRESHER SHARKS. FAMILY ALOPIIDAE The threshers (several species are known) are peculiar among sharks for their enormously long tail fin. Their closest affinities in other respects are with the mackerel sharks. Thresher Alopias vulpinus (Bonnaterre) 1758 Thraser; Swiveltail; Fox shark Bigelow and Schroeder, 1948, p. 167. Garman, 1913, pi. 7, figs. 1-3. Description. — The thresher is as easily distin- guished from all other Gulf of Maine sharks by its long tail as the hammerhead is by its head, the upper caudal lobe being a little longer than the head and body of the fish together, curved much like the blade of an ordinary scythe, and notched near the tip, whereas the lower lobe measured along the front margin is hardly longer than the pelvic fins. We need merely point out in addition that the first dorsal fin (of moderate size and about as high as it is long) stands about midway between pectoral and pelvic fins; that the second dorsal fin and the anal are very small; that the pectoral fin is long and sickle shaped; and that the thresher is a stout-bodied shark with short snout and blunt, rounded nose. Its teeth are small, subtriangular with a single sharp cusp and are smooth edged. Those near the center of mouth are nearly symmetrical, but the successive teeth are increasingly oblique outward, with their outer margins increasingly concave. Color. — Dark brown, blue-slate, slate gray, blue gray, leaden or even nearly black above, often with metallic luster, grading on the sides to white below, except that the snout and the lower surface of the pectorals are usually about as dark below as above, and that the sides near the pectorals may be more or less mottled with gray, the belly also. The iris is black or green. Size. — Threshers vary considerably in size at birth, for while free living specimens have been reported as small as 46 inches, with manjr of 48 to 60 inches (some with umbilical scars still showing), one unborn embryo was 61 inches long. The state of development of the claspers of males, with the lengths (14 ft. 6 in. and about 15K ft.) of females that have been found with embryos, makes it unlikely that they mature sexually until they are at least 14 feet long (tail included). Lengths up -•.---' D ^2 Figure 9. — Thresher (Alopias vulpinus), about 5 feet- long, Rhode Island, from Goode, drawing by H. L. Todd. A, upper second tooth; B, upper third tooth; C, upper fifth tooth; D, upper fifteenth tooth; E, lower second tooth; F, lower sixth tooth, counted from center of jaw; about 2 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. FISHES OF THE GULF OF MAINE 33 to 16 feet are usual;76 the maximum length (tail included) is about 20 feet. Threshers are so large- ly tail that they are much lighter than many other sharks, length for length. The few actually weighed have ranged from about 300 to 320 pounds at about 10 feet, and 375 to 400 pounds at about 13 feet, to about 500 pounds at about 14% feet. Perhaps 1,000 pounds is about the maxi- mum to be expected for a very large one. Habits. — The reports of threshers are mostly based on ones seen at the surface or caught either in nets set shoal, or in traps set close inshore. But a thresher has been hooked as deep as 35 fathoms in British waters.77 The thresher feeds chiefly if not exclusively on small schooling fishes; in American waters mostly on mackerel, menhaden, herring, and bluefish (Pomatomus) ; also on bonito and on squid. A pair of threshers often work in concert "herding" a school of fish, and it is to frighten its prey together that the enormously long, flail-like tail is em- ployed. Allen 78 gives an interesting eyewitness account of a thresher pursuing and striking a single small fish with its tail. The tale that the thresher leagues with the swordfish to attack whales is time honored, but has long since been relegated to the category of myth. And so weak toothed is this shark that the second part of the story (it makes a meal of its huge victim) is close to an impossibility. The thresher, we may add, does not harm human beings. In American waters it is probable that threshers are born throughout its range, very small free living specimens having been caught off New Eng- land on the one hand, and off Florida on the other. The embryos do not develop a placental attach- ment with the mother, and either 2 or 4 have been reported in gravid females. General range. — This is an oceanic shark of temperate and subtropical seas. In the Atlantic it is known from southern Ireland and the North Sea to Madeira and the Mediterranean in the east, and also from the Cape of Good Hope ; from Nova Scotia and the Gulf of St. Lawrence to Cuba and the northern part of the Gulf of Mexico in the '• Several of tint size have been I ikon In the traps at Woods Hole. " There Is another group of species or the genus, with very large eyes, that live at greater depths; for discussion of these, see Bigelow and Schroeder (Fish. Western North Atlantic, Pt. 1, 1948, pp. 162, 163). n Science, N. Ser., vol. 58, 19?3, pp. 31-32. west, and again from southern Brazil and northern Argentina. Seemingly it does not occur in the equatorial belt of the Atlantic. But it does in the Pacific, where it is known from Oregon to Panama and Chile. Threshers of this same type are also found in the central and western Pacific and in the Indian Ocean. Whether the thresher of the eastern side of the Pacific is identical with that of the Indian Ocean remains to be determined. Occurrence in the Gulf of Maine. — The thresher has often been seen off the southern coast of New England and in some numbers. Three about 16 feet long have been taken near Woods Hole, for example, in one trap in a single morning, and it has been classed as the commonest of the large sharks off Block Island. Scattered specimens also visit the Gulf of Maine in some years, though perhaps none in others. Thus two have been reported in print from Nantucket; we saw several large ones in Pollock Rip, off the southern angle of Cape Cod on August 4, 1913; it has been re- ported repeatedly on the coast of Massachusetts, as at Barnstable on Cape Cod Bay, where one about 10 feet long was taken in a trap on October 21, 1949, and from various localities in Massa- chusetts Bay (c. g. Boston Harbor and Nahant). Records for it along the coast of Maine include the vicinity of Monhcgan Island, east of Matinicus Island, the offing of Penobscot Bay where one weighing about 500 pounds (estimated) was caught in 1911 and another seen in 1911, in the vicinity of Eastport. It has also been taken in the cold waters of Passamaquoddy Bay; one for instance in a weir at Deer Island, August 28, 1936; 7° also in the Basin of Minas on the Nova Scotiau shore of the Bay of Fundy. Occasionally a thresher is netted or seen off the outer coast of Nova Scotia. The most northerly record for it from our side of the Atlantic is for the Bay of Chaleur in the southern side of the Gulf of St. Lawrence. It is to be expected in Gulf of Maine waters only during the warm half of the year, perhaps May to October (April to late autumn for Woods Hole); in the cold season it altogether deserts our northern coasts for warmer seas. Importance.— The thresher is not common enough in the Gulf of Maine to be of any impor- tance to fishermen one way or another, or to play '• Reported by McKenzle, Proc. Nova Scotia Inst. Sci., vol. 20, 1939, p. 11. 210941—53- 34 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE a practical role of any moment among the smaller fish. Further south, however, and wherever it is numerous in the Atlantic, it makes itself a pest, tangling and tearing mackerel nets as well as destroying and chasing away the more valuable fishes on which it feeds. CAT SHARKS Distinctive features of these little sharks are that they have five pahs of gill openings and an anal fin; that at least one-half of the base of the first dorsal fin is rearward of the point of origin of the pelvic fins; that the front margin of the nostrils does not bear a fleshy barbel; and that they lay eggs with horny shells and tendrils at the corners. Many species are known. The familiar spotted dogfishes of European seas (two species) fall in this group. And one species calls for mention here. Chain dogfish Scyliorhinus retifer (Garman) 1881 Description. — The chain-like pattern of narrow black stripes with which the reddish-brown back and sides of this little shark are marked are so distinctive that there is no likelihood of confusing it with any other shark. We need only add that its first dorsal fin stands wholly behind the rear ends of the bases of its pelvic fins; that its second FAMILY SCYLIORHINIDAE dorsal fin is about one-half as large in area as its first dorsal fin; that its tail fin is square-tipped and occupies only about one-fifth of the length of the fish; and that its teeth are similar in the two jaws, narrow-triangular with a small second- ary cusp on either side. Size. — The largest specimen measured so far was 17 inches long. General range and occurrence in the Gulf of Maine. — The range of the chain dogfish is con- fined to the 40-125 fathom zone between the offings of Cape Lookout, North Carolina, and of Nantucket. It seems to be the most plentiful off Virginia, in the general offing of Chesapeake Bay, where considerable numbers are taken during the winter trawl fishing. They are caught now and then as far as the offing of Marthas Vineyard, and Cap'n Bill II trawled one, in July 1952, south of Nantucket Lightship, Lat. 40°02' N; Long. 69°37' W, at 75-90 fathoms which brings it within the arbitrary boundary of the Gulf of Maine. Figure 9A. — Chain dogfish (Scyliorhinus retifer), male, about 17 inches long, New Jersey. After Bigelow and Schroeder. SMOOTH DOGFISHES. FAMILY TRIAKIDAE These are rather small sharks, with two dorsal fins without spines, the second dorsal (in Atlantic species) nearly as large as the first, and they have an anal fin. The tail fin is very strongly asym- metrical, its lower anterior corner forming a low but rather definite lobe in some, but not in others. The teeth are small, with several rows in function imultaneously, flat, and pavement-like in some, but with three or four definite cusps in others. The eye has no nictitating ("winking") mem- brane, but only a longitudinal fold along the lower eyelid. They resemble the requiem sharks (Family Carcharhinidae, p. 36), except for the teeth, and for the lack of a nictitating membrane. Only one species is known from the Gulf of Maine, or is ever likely to be found there. Smooth dogfish Mustelus canis (Mitchill) 1815 Smooth dog; Smooth hound; Grayfish Bigelow and Schroeder, 1948, p. 244. Garman, 1913, pi. 4, figs. 6-9, as Galeorhinus laevis. FISHES OF THE GULF OF MAINE 35 Figure 10. — Smooth dogfish (Mustelus cants), male, about 31 inches long, Woods Hole. A, tooth band of right-hand side of upper jaw, about 1.8 times natural size; B, teeth of another specimen, about 6 times natural size. From Bigelovv and Schroeder. Drawings by E. N. Fischer. Description. — The smooth dog is easily identi- fied among Gulf of Maine sharks by having two large spineless dorsal fins, the second only a little smaller than the first, combined with low, flat, pavement-like teeth. So different, indeed, are its teeth from the awl-like or blade-like teeth of all our other sharks that a glance at the mouth is enough to separate this species from the young of any larger Gulf of Maine shark. In form this little shark is slender, flattened below, with taper- ing but blunt snout. Its first dorsal fin originates nearly over the hind angle of the pectorals. The second dorsal fin is about twice as large as the anal, over which its stands. The tail is of typical "shark" shape, i. e. with upper lobe much longer than lower. The hind margin of the upper lobe of the caudal is deeply notched near the tip; the lower caudal lobe is very small. Color. — Upper surface grayish olive, slaty gray or brown, lower surface yellowish or grayish white. Newborn specimens have the upper part of the first dorsal fin edged with dusky gray; the apex of the second dorsal sooty edged or tipped, but with the rear edge white; the tail fin with a sooty blotch above near the tip, but white edged below. But these markings have mostly faded out by the time the little "dog" has grown to a length of two feet or so. Smooth dogs have a greater ability than most sharks to change shade to suit their surroundings, paling to a translucent pearly tint above white sand, but darkening on dark bottom.80 Size. — Smooth dogs range from about 11% inches to about 14% inches long when born. They mature sexually at about 3 feet, most of the ma- ture females with young are between about 3 feet 3 inches and 4 feet 4 inches long; and a few grow to a length of about 5 feet. Habits. — The smooth dog is most familiar as a shore fish and a bottom swimmer, commonly entering shoal harbors and ba3Ts, and even coming into fresh water. But fishermen also report them as far offshore as the "tile fish" grounds off southern New England and down to a depth of 80 to 90 fathoms. They reach the northern part of their range only as warm-season visitors; at Woods Hole they arrive sometime in May, to withdraw in late October or in November. Food of the smooth dogfish consists chiefly of the larger Crustacea, and it is perhaps the most relentless enemy of the lobster, which had been eaten by no less than 16 percent of the fish examined by Field. Large crabs are likewise an important article in its diet, as are the smaller fishes. It has been estimated that 10,000 smooth dogfish, in Buzzards Bay, might devour more than 60,000 lobsters yearly, and perhaps one-fifth » Experiments have shown that It requires only 1 lo 2 hours for one to darken, but as much as 2 days to pale to the extreme; see Parker (Biol. Bull., vol. 66, 1934, p. 31). 36 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE million crabs, besides a great number of small fish (menhaden and tautog are the species most often found in smooth dogfish stomachs). And these figures are based on a sufficient number of observations of the stomach contents to serve as a general indication of the destructiveness of the smooth dogfish. They also feed on squid, espe- cially in spring, and while they do not regularly take hard-shelled mollusks, razor clams have been found in the stomachs of several at Woods Hole. When kept in captivity they are constantly on the move, searching the bottom for food, which they find chiefly by the sense of smell though their sight is also keen.81 Any crab that may be offered is soon found, seized, shaken to and fro, and eaten. And with packs of these sea hounds hunting over every square foot of our southern bays and sounds it is a wonder any of the larger crustaceans escape where dogfish are abundant. Field 82 also made the interesting observation that the smooth dogfish never molested healthy and active menhaden but soon devoured any sick or injured fish that might be in the same tank with them. As this is not a characteristic Gulf of Maine fish, we need merely note that it is one of the sharks that develop a placental connection be- tween the embryos and the mother. In other words, it is truly viviparous. The period of gestation appears to be about 10 months; off southern New England the young are born be- tween early May and mid July. The number in a litter usually is between 10 and 20, but as few as 4 have been reported. A description of the un- born young is given by Fowler.83 General range. — Coastal waters of the western Atlantic, from Uruguay and southern Brazil, regularly to Cape Cod, and to Passamaquoddy Bay as a stray; also Bermuda.84 Occurrence in the Gulj of Maine. — The smooth dog is the second most numerous shark along the southern coast of New England, though falling far short of the spiny dogfish (p. 50). At Woods Hole, for example, pound-net catches varied during the summer of 1903 from 1 to 41, averaging about 7, and catches up to 100 have been reported from the vicinity at one time. Similarly, catches of 5 or 6 on a hand line are common in a few hours' fishing, with as many as 10 to 20 reported. But the elbow of Cape Cod and the region of Nan- tucket Shoals mark so definite a boundary to their dispersal eastward that while they have been reported from Provincetown, from various locali- ties within Massachusetts Bay, and even from as far north as St. Andrews in the Bay of Fimdy, where one was caught in July 1913, neither of us had ever seen one north of Cape Cod until Sep- tember 21, 1951, when an angler (Ellery Sidney) showed us a female about 3 feet long that he had caught at Cohasset, while casting with an eel skin, for striped bass. So far as known its occasional incursion- into the Gulf are sporadic, at least they have not been correlated with unusually warm summers or with the presence of other southern fishes. Neither has it been re- ported by fishermen from Georges or Browns Banks, nor was it detected there by the repre- sentatives of the Bureau of Fisheries during the trawling investigations of the years 1912 and 1913 (p. 60), or subsequently. REQUIEM SHARKS. FAMILY CARCHARHINIDAE This family, which includes a large number of species in tropical and temperate seas, is charac- terized by a head of normal shape, eye with a nictitating (winking) membrane, tail with the upper lobe considerably larger than the lower but not very long, 2 spineless dorsal fins, the first usually much larger than the second in most of 61 The senses of this shark have been studied by Parker (Bull., U. S. Bur. of Fish., vol. 29, 1911, pp. 43-57), and by Sheldon (Jour. Compar. Neurol, and Psychol, vol. 19, 1909, No. 3, p. 273). »» Rept. U. S. Comm. Fish., (1006), 1907, Spec. Pap. 6, pp. 14-16. 13 Occas. Pap. Mus. Zoo]., Univ. Mich., No. 56, 1918, p. 15. the Atlantic species,86 an anal fin, a caudal peduncle lacking lateral keels, and sharp, bladelike teeth with a single cusp. All bear "living" young; " Present indications are that several more or less isolated populations of this shark exist, with their areas of regular occurrence separated by wide gaps, where Ihere is little or no intermingling. One of the best known is along the Atlantic coast, Cape Cod to North Carolina; another centers in the Gulf of Mexico-Caribbean region; a third is along southern Brazil and Uruguay. For further details, see Bigelow and Schrocdcr, Fishes Western North Atlantic, Part 1, 1948, pp. 250-251. 85 The lemon shark (Negaprion brtpirosiris) of warmer waters, which has been known to stray to New Jersey, is an exception in this respect; its second dorsal is nearly as large as its first dorsal. FISHES OF THE GULF OF MAINE 37 some have a placental connection between mother and embryo, but others do not. Tiger shark Galeocerdo cuvier (LeSueur) 1822 Leopard shark Bigelow and Schroeder, 1948, p. 266. Description. — The tiger shark is characterized among the Atlantic members of its family by the forward position of its first dorsal fin (origin about over the arm pit of the pectorals), combined with a caudal peduncle with a low longitudinal ridge of skin on either side, besides a well-marked semilunar pit below as well as above; a very small second dorsal fin; a furrow, about as long as the snout along either side of the upper jaw; a very slender-tipped caudal fin with moderately large and pointed lower lobe; and large teeth alike in the two jaws, of very characteristic shape, with convex inner margins, deeply and conspicuously notched outer margins and strongly serrate edges (fig. ID- Young tiger sharks arc rather slender, but they become very heavy forward, with growth, though they continue tapering toward the tail. The first dorsal fin is high, triangular, and nearly as large as the pectorals, while the second dorsal is hardly one-third to one-fourth as high as the first and stands over the anal, which is of about equal size. The lower tail lobe is almost half as long as the upper, the rear margin of which is notched near the tip. The large size of the head, with very short, obtusely rounded front outline, and broad mouth occupying nearly four-fifths of the width of the head, with long grooves along the upper jaw, combined with the unique shape of its teeth, make the "tiger" easy to recognize among Gulf of Maine sharks. Color. — Gray, or grayish brown, darkest on the upper surface. Young "tigers" up to 5 or 6 feet long, are more or less conspicuously spotted or barred with darker brown on the back and along the upper parts of the sides. But these markings fade with advancing age until large specimens are plain colored, or nearly so. Size. — Tiger sharks are small at birth, corre- sponding to the large numbers in a litter, free living specimens having been reported only 18 to 19 inches long. By the time they mature they are among the larger sharks; but their size has often been overestimated. The majority of tigers caught in centers of abundance are less than 12 to 13 feet long, and the largest measured lately in the western Atlantic was one of about 18 feet, from Cuba. Repeated statements that the tiger grows to a maximum length of 30 feet have no reliable foundation, so far as we can discover. A 4-foot specimen from Woods Hole weighed 25% pounds when taken from the water. Larger tigers vary widely hi weight at given lengths depending on how fat they are and on the stage of development of the young in gravid females. Specimens from various localities have weighed 37 pounds at 5)2 feet; 168 pounds at 6 feet; 366 to 718 pounds at 10 to 11 feet; 450 to 825 pounds at 11 to 12 feet; 630 to 1,324 pounds at 12 to 13 Figure 11. — Tiger shark (Galeocerdo cuvier), young male, about 49 inches long, Rhode Island. A, upper tooth, and B, lower tooth of larger specimen, enlarged. From Bigelow and Schroeder. Drawings by E. N. Fischer. 38 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE feet; and 1,028 to 1,395 pounds at 13 to 14 feet.86 Habits. — This voracious shark, with wide jaws and powerful teeth, preys upon the large sea turtles, other sharks, fish, and occasionally on invertebrates such as horseshoe crabs, crabs, conchs, whelks. It is proverbial for its habit of feeding on slaughter-house wastes or any other carrion. Remnants of squeteague, mackerel, hake, scup, menhaden, goosefish, and dogfish all have been found in stomachs of tiger sharks taken at Woods Hole.87 There is no placental connec- tion between mother and young, and the broods are very large, as many as 82 having been counted in a large female; but other litters as small as 10 to 14. In the West Indies it is much dreaded, whether or not with good cause. General range. — Cosmopolitan in the warmer waters of all oceans; straying northward as far as Cape Cod on the American coast of the Atlantic. Occurrence in the Gvlj of Maine. — A few tiger sharks are taken in fish traps in the Woods Hole region every year, seldom before August or later than October although one was caught there July 20, 1951. 88 These specimens usually have been about 5 feet long, at most about 8 feet, and very rarely does a full-grown tiger shark stray so far from its tropical home. The tiger has not yet been recorded (on reliable evidence) from within the limits of the Gulf of Maine. It is included here *• For further details and references, see Bigelow and Schroeder, Fishes Western North Atlantic, Pt. 1, 1948, p. 269. " Bell and Nichols (Copeia, No. 92, March 1921, pp. 17-20) list the stomach contents of a number of tiger sharks caught off Morehead City, N. C. " This shark was 8 feet, 3 Inches long, taken in a pound net off Quisset Harbor, Buzzards Bay. because of the likelihood that a stray specimen may occasionally round the elbow of Cape Cod, or be encountered on the offshore Banks.89 Blue shark Prionace glauca (Linnaeus) 1758 Blue dog Bigelow and Schroeder, 1948, p. 2S2. Garman, 1913, pi. 3, figs. 1-3 (as Galeus glaucus). Description. — The blue shark is slender-bodied, thickest about its mid-length, and tapers toward head and tail (a shape usually named "fusiform"). Its snout is long with rounded tip. Its first dorsal fin is of moderate size, standing far back with the mid point of its base about midway between the inner corners of the pectorals (when these are laid back) and the points of origin of the pelvic fins. The second dorsal fin is less than one-half as high as the first, and is about equal in size to the anal over which it stands. The pectorals are narrow and very long, their tips reaching back nearly as far as the rear corner of the first dorsal. The lower lobe of the caudal fin (measured along its anterior edge) is about one-half as long as the upper lobe; the latter is conspicuously notched near the tip, and both of the lobes of the caudal fin are slender tipped. The teeth are large, sharp-pointed, with serrate edges, and distinctive in shape. The uppers are so closely spaced that the bases of adjacent teeth » The statement in the first edition that a tiger shark was once taken at Provincetown was an error. The original description of the specimen In question (Atwood, Proc. Boston Soc. Nat. Hist., vol. 10, 1S65, p. 81) suegests that it was a mako (hums oiyrinehus) . Figure 12.— Blue shark (Prionace glauca), male, about 7 feet 2 inches long, off Marthas Vineyard. A, third left-hand upper tooth, counted from mid-point of jaw; B, ninth left-hand upper tooth; C, third left-hand lower tooth; and D, eighth left-hand lower tooth; about 1.6 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. FISHES OF THE GULF OF MAINE 39 overlap. The median upper tooth is nearly symmetrical, but those along the sides of the mouth have strongly convex outer margins, and deeply concave inner margins, while their points curve sharply outward toward the respective corner of the mouth. The lower teeth are nar- rower, more nearly symmetrical, and nearly erect. Color. — Living specimens are dark indigo blue along the back, shading to a clear bright blue *° along the sides; but this beautiful hue changes to a slaty or sooty gray soon after death. The lower surface is snow-white, but with the tips of the pectorals dusky and the anal fin partly sooty. Size. — The usual length at birth seems to be between VA and 2 feet.91 Blue sharks do not ma- ture until they have grown to be 7 or 8 feet long, to judge from the sizes of the females that have been found with young; the longest we have handled was almost exactly 11 feet long. The fact that the greatest measured length so far re- liably reported was only 12 feet 7 mches (3.83 meters) suggests that repeated characterizations of the blue shark as commonly growing to 15 feet are an exaggeration. If any grow to 20 feet, as is rumored, they must be giants of their kind. Remarks. — The very long slender pectorals of the blue shark, combined with its long narrow snout, the position of its first dorsal fin far back, and its brilliant blue color, give it an aspect very different from that of the tiger shark (p. 37), of the sharp-nosed shark (p. 40), the dusky or brown sharks (pp. 41-43), or that of any other carcharhinid shark that might perhaps straggle to the Gulf of Maine. Habits. — The blue shark is "encountered indif- ferently far out at sea and in continental waters, its wanderings no doubt directed chiefly by the search for food, though it may drift with ocean currents. It is frequently seen at the surface, swimming lazily with first dorsal fin and tip of caudal out of water, or basking in the sun. There is no reason to suppose that it ever descends to any great depth." 92 They sometimes follow sail- ing ships for days on end, to pick up scraps, and their habit of gathering when a sperm whale was killed, to feed on the carcass, was proverbial dur- ing the days of the sperm whale fishery.93 But their normal diet is smaller fishes, of whatever kinds may be available. In northern waters herring, mackerel, spiny dogfish, and various others have been found in their stomachs. And we have several times seen a blue shark pick up a tagged cod, haddock or American pollock that we had put back in the water, on Georges Bank. The blue shark is viviparous, that is to say, the embryo has a well developed placenta attached to the mother. As many as 28 to 54 young have been reported in a litter in the Mediterranean. General range. — Cosmopolitan on the high seas in the warmer parts of all the oceans, including the Mediterranean; ranging northward to outer Nova Scotia and as a stray to the Banks of New- foundland in the western side of the Atlantic; to England and Scotland in the east, with stray specimens reaching the Orkneys and southern Norway. This, we think, is by far the most nu- merous of the large, oceanic sharks; it is the one witli which the sperm whalers were the most familiar; the one around which many of the super- stitions about sharks have developed; and the one with which we have had to do most often. Occurrence in the Gulj oj Maine and along Nova Scotia. — Only one blue shark had been reported definitely from the Gidf of Maine in scientific lit- erature, up to the time the first edition of this book was printed, though it was known to be rather com- mon along outer Nova Scotia. But we have learned since then that it is a regular summer visi- tor to the southern and western parts of the Gulf, appearing occasionally in July, more often in Aug- ust and September. In 1928, for example, we caught one on Stellwagen Bank on August 26, saw one over the northern end of Jeffreys Ledge on September 2, and caught four on Platts Bank on September 3, with others in sight from the vessel at nearly all times throughout the day. And many more have been seen or caught subsequently, on Platts Bank, in Massachusetts and Cape Cod Bays, where 1 8 were reported to us during the summer of 1935,9* on Georges Bank where blue sharks, swim- ming at the surface, are a familiar sight in summer; and on Browns Bank. Two have also been re- " "Sailor blue," as shown in Rldgeway's Color Standards and Color Nomenclature, 1912, p. 21. " Embryos have been reported as long as about 17H inches, and free-living specimens as small as 20-21 inches. •* Blgelow and Schroeder, Fishes Western North Atlantic, Pt. 1, 1948, p. 286. »» Nichols and Murphy (Brooklyn Mus. Scl. Bull., vol. 3, No. 1, 1916, p. 9) have given a graphic account of the blue shark as it was met with by whalers on the high seas. " By J. E. Lowes, an experienced shark fisherman. 40 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE ported to us recently from the coast of Maine, a few miles east of Casco Bay.96 We have never heard of a blue shark in the north- eastern corner of the Gulf, in the Bay of Fundy, nor along western Nova Scotia, whence they may be barred by colder surface waters. But fisher- men are familiar with them off the outer coast of Nova Scotia, both offshore, and also near the coast at the times when the warm surface water presses inshore. Blues were reported near Halifax, for instance, from time to time between August 15 and October 10, 1920, some coming close in to the entrance to the Harbor. And two specimens have been re- ported at Canso,96 but whether the "blue dogs" described by local fishermen as common on the neighboring banks actually are this shark, or per- haps the porbeagle, seems doubtful. It has also been recorded from the southwest part of the Grand Bank of Newfoundland.97 Following westward from Cape Cod, we find many records of blues from the traps near Woods Hole, and they are often seen (or harpooned) on the continental shelf in the offing. Twenty-eight were counted 4 to 10 miles off Block Island for ex- ample, during one hour, and something like 150 to 200 during the day (13 of them were harpooned) on August 22, 1943. Most of the blues that are seen or taken off our northern coast are medium sized or larger, though very small ones are taken from time to time.88 " By the late Walter H. Rich, who was long associated with the V. S. Bu- reau of Fisheries. « Cornish, Contr. Canadian Biol. (1902-1905) 1907, p. 81. « Rept. Newfoundland Fish. Res. Lab., 1935, p. 79. •! Robert Goffln reports one only 20 inches long, from Menemsha Bight, near Woods Hole, August 31, 1925; we have seen one of 21 Inches, taken a few miles off Block Island, August 22, 1943; and F. D. Firth reports one 34H inches long taken 65 miles southeast of Highland Light, Cape Cod, on October 23, 1930. And for some obscure reason all but two of the adults seen in our Gulf, for which we have the per- tinent information, have been males. Commercial importance. — This shark is of no commercial value. A few are caught by anglers, mostly on natural bait, and a Blue will sometimes take an artificial lure; we hooked one off Boone Island, Maine, on a feather jig, tipped with pork rind. We have never had blues put up much re- sistance on a heavy hand line until hauled in to the side of the vessel, when they thrash about vio- lently, but it is said that a large one will make long and powerful runs, if hooked on rod and reel. The blue shark has always been looked on with contempt by the sperm whalers, who were more familiar with it than anyone else. We find no well- authenticated case of one attacking a swimmer, sailors' yarns to the contrary notwithstanding. Sharp nosed shark Scoliodon terrae-nonas (Richardson) 1836 Bigelow and Schroeder, 1948, p. 295. Garman, 1913, pi. 2, figs. 1-4. Description. — This little shark is separable from any other carcharhinid that has yet been reported from the Gulf of Maine or that is likely to be, by its upper and lower teeth which are perfectly smooth along the edges from tip to base, combined with a so-called "labial furrow" of considerable length running forward along each side of each jaw from the corner of the mouth toward the nostril. This last character, while not conspicuous, is a precise one. The trunk is slender, highest about at the first dorsal fin, tapering both fore and aft. The snout varies rather widely in length and in bluntness at the tip. The point of origin of the first dorsal fin Figure 13. — Sharp-nosed shark (Scoliodon terrae-novae) , female, about 31 inches long, from the Bahamas. From Bigelow and Schroeder. Drawing by E. N. Fischer. FISHES OF THE GULF OF MAINE 41 is about over the inner corners of the pectorals when the latter are laid back; its height is about one-half as great as the distance from the tip of the snout to the level of the origin of the pectorals. The second dorsal is only about one-quarter as high as the first; its point of origin is about over the mid-point of the base of the anal fin; the anal is a little larger than the second dorsal. The tail fin occupies about one-quarter of the total length of the shark; its lower lobe (measured along the anterior edge) is a little less than one-half as long as the upper lobe, the rear edge of which is deeply notched near the tip. The pectoral fins are smaller relatively than in any other local species of this family, their length, armpit to tip, being only a little greater than the height of the first dorsal fin. The teeth are alike in shape in the two jaws, sharp-pointed and smooth edged; those in the center of the mouth are symmetrical and erect, but those along the sides have weakly concave inner margins, but deeply notched outer margins, and are increasingly oblique toward the corners of the mouth. Color. — Brown to olive gray above, with the dorsal and caudal fins more or less dark edged; white below and along the rear margins of the pectorals. Size. — Mature specimens are commonly between 26 and 30 inches long; a few grow to 36 inches. General range. — Both sides of the tropical-sub- tropical Atlantic; Morocco to Cameroon and the Cape Verde Islands in the east; Uruguay to North Carolina in the west; occasional to Woods Hole, and as a stray to the Bay of Fundy. Occurrence in the Gulf of Maine. — Our only reason for including this warm-water shark is that one was taken at Grand Manan Island," at the mouth of the Bay of Fundy, in 1857.1"2 Early reports of it from Newfoundland were based on a misconception. Sharks of the Genus Carcharhinus The members of the genus Carcharhinus are set apart from other Atlantic members of the family Carcharhinidae by the following combination of characters: The mid-point of base of the first dorsal fin is at least as near to the level of the axils of the pectorals as to the level of the origin of the •• This specimen, collected by A. E. Verrlll. Is In the Museum of Com- parative Zoology. '-' See Jordan and Evermann, Bull. 47, U. S. Nat. Mus., Pt. 1, 1896, p. 43, footnote. pelvics (separating them from the blue shark, p. 38) ; no labial furrows on lower jaw, and furrow on upper jaw reduced to a very short slit at the extreme corner of the mouth, directed outward (separating them from the tiger shark, p. 37, and from the sharp-nosed shark, p. 40) ; second dorsal fin much smaller than first dorsal (separating them from the lemon shark, p. 35, footnote 85) ; edges of upper teeth more or less finely serrate but without larger denticles near the base, and edges of lower teeth perfectly smooth, without lateral denticles (separating them from the tiger shark, p. 37, from the sharp-nosed shark, p. 40), and from Paragaleus pecloralis, a tropical shark that has been taken off southern New England.3 This is a warm-water group, fifteen species of which are known to inhabit the western side of the Atlantic, most of them resembling one another closely in general aspect. Only one of these (the dusky shark, described on p. 41) has yet been reported reliably from within the confines of the Gulf of Maine, while only one other (the brown shark, p. 43) is likely to be found there. If a stray Carcharhinus from offshore that does not agree with the following descriptions of one or other of these should be taken on Georges Bank, or on Nantucket Shoals cast of the longitude of Cape Cod, we hope that its captor can identify it by means of the keys and descriptions of the genus that we have given in Part 1 of the Fishes of the Western North Atlantic. Dusky shark Carcharhinus obscurus (LeSueur) 1818. Bigelow and Schroeder, 1948, p. 382. Description. — The combination of characters that place the dusky shark among the western Atlantic members of its genus are: Trunk about one-fifth as high at first dorsal fin as it is long to origin of the caudal fin, tapering both forward and rearward; snout broadly rounded in front, its length in front of the nostrils less than the distance between the nostrils; the front edge of the nostril is not expanded as a definite lobe; the midline of the back between the two dorsal fins has a low but definite ridge, a character which is very pre- cise, though seemingly minor; the first dorsal fin is considerably smaller than in the brown shark • For description, see Bigelow and Schroeder, Fishes of the Western North Atlantic, Pt. 1, 1948, p. 276. 42 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 14. — Above: Dusky shark (Carcharhinus obscurus), female about 39 inches long, Woods Hole. A, third upper tooth; B, fourth lower tooth; C, ninth upper tooth; D, tenth lower tooth; about 2.4 times natural size. Below: Brown shark (Carcharhinus milberti), female, about 4 feet 10 inches long, from Woods Hole. A, ninth upper tooth; B, eighth lower tooth; C, third lower tooth; about 1.4 times natuial size. From Bigelow and Schroeder. Drawings by E. N. Fischer. (p. 43), with more deeply concave rear margin, its point of origin about over the inner corner of the pectoral (over the armpit of the pectoral in the brown shark) ; its apex is narrowly rounded. The free rear corner of the second dorsal fin is less than twice as long as the vertical height of the fin. The anal fin is a little longer, along the base, than the second dorsal and stands about under the latter. The caudal fin occupies between one* quarter and one-third of the total length of the shark, the lower caudal lobe (measured along its anterior edge) is about two-fifths as long as the upper lobe; and the upper lobe is noticeably slender toward its tip. The pectorals are about as long (from origin to tip) as the distance from the tip of the snout to the level of the first pair of gill openings, usually narrower, relative^, than in the brown shark, and sometimes more definitely siclde-shaped. The upper teeth are broadly triangular; nearly erect toward the center of the mouth but wealdy oblique toward its corners; their inner margins are nearly straight, the outer margins increasingly concave outward along the jaw. The lower teeth are erect, symmetrical, with narrow cusp on a broadly expanded base. Both the upper teeth and the lower are serrate along the edges, the lower the more finely so. Color. — All the fresh caught specimens we have seen have been bluish or leaden gray on the back FISHES OF THE GULF OF MAINE 43 and upper part of the sides, including the pectorals, but this shark has also been described as pale gray above or even dirty white, perhaps over a white sand bottom. The trunk is white below, the pectorals grayish, darkening to sooty at then- tips ; the pelvics and anal fins grayish white. Size. — The usual length at birth is a little more than three feet.4 Adult dusky sharks so far measured have ranged from 10 feet 4 inches to 11 feet 8 inches in length, and they are said to grow to 14 feet, though perhaps not on very convincing evidence. General range. — Western Atlantic, north to southern New England and to Georges Bank, south to southern Brazil, at least by name. A shark very closely allied to obscurus has been reported under that name in the eastern Atlantic, from Spain to Table Bay, South Africa, including Madeira, the Canaries, the Cape Verdes, Ascen- sion Island, and St. Helena. But we have yet to learn its precise relationship to the obscurvs of the western Atlantic. Occurrence in the Gulf of Maine. — The dusky shark has been taken repeatedly off the coasts of New Jersey and of Long Island, N. Y.; also at Woods Hole, where we have handled 12 specimens during the past few summers, 6 of them in August 1944. But it so seldom strays to cooler waters farther east that only one shark has been recorded from Nantucket, and one from Georges Bank, that probably were of this species and not some other carcharhinid.6 Thus it has no real place in the fauna of the Gulf.6 Brown shark Carcharhinus milberti (Miiller and Henle) 1841 Sand bar shark Bigelow and Schroeder, 1948, p. 368. Garman, 1913, pi. 3, figs. 4-6 (as Carcharinus plalyo- dori). Description. — The brown shark differs from the dusky (only member of its genus that seems actually to have been taken within the Gulf) in the more forward position and larger size of its first dorsal fin, in its broader pectorals, and in its stouter trunk, heaviest forward (compare speci- mens in figure 14). Also, the anterior edge of its nostril is expanded as a low but definite triangular lobe, which is not the case hi the dusky shark. Other characters (in combination) that mark it off from other members of this genus that might stray to the Gulf are: Mid-line of the back with a low ridge between the two dorsal fins; snout forward of a line connecting the front margins of the nostrils, considerably shorter than the distance between the nostrils; point of origin of second dorsal fin about over origin of anal fin, its free rear corner only a little longer than the height of the fin; apex of first dorsal fin angular; length of pectorals along anterior margin about as great as distance from tip of snout to level of second pah of gill openings; distance from rear tips of pelvic fins to origin of anal fin as long as base of anal fin, or longer, fifth gill openings longer than horizontal diameter of eye. The teeth resemble closely those of the dusky shark (see figure 14). Color. — Upper surface slate gray to brown; lower surface a paler tint of the same hue, or white; fins without any conspicuous black mark- ings. When alive some of the dermal denticles are bright blue, at least on some specimens. Size. — Sexual maturity is reached at a length of about 6 feet; maximum length about eight feet.7 Generalrange. — Southern Brazil, Louisiana, both coasts of Florida, and northward along the Atlantic coast of the United States to southern New England; also the tropical-subtropical belt of the eastern Atlantic, and the Mediterranean, or represented there by an extremely close relative.8 Occurrence in the Gulf of Maine. — Next to the sand shark, this is the most numerous of the larger sharks along the coasts of New Jersey and of New York. Some visit the vicinity of Woods Hole, though so few that the number taken there in most summers probably is not greater than six or seven. It has not been reported as yet from « Embryos have been reported up to 38 in. long (965 mm.), and a free living specimen of only 39ln. (993 mm.); see Bigelow and Schroeder, Fishes Western North Atlantic. I>t. 1, 1948, p. 387. ■ Probably this species and not the brown shark because 11-12 feet long. « In the first edition of this book, the dusky shark was said to havo been taken at three localities within the Gulf. But one of these records, at least, was almost certainly based on a blue shark, and the others probably wcro (Bigelow and Schroeder, Fishes Western North Atlantic, Ft. 1, 1948, pp. 292, 368). » Seven feet 10 laches is the greatest measured length that we have found recorded, with convincing evidence that the specimen actually was one of this species. ! If the eastern Atlantic-Mediterranean form Is actually Identical with the American, as seems to be the case, the specific name milberti of Miiller and Henle, 1811, must be replaced by plumbeus proposed by Nardo In 1827 for the brown shark of the Adriatic. 44 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE within the limits of our Gulf, but is included here on the chance that a stray specimen may be taken, either on the outer coast of Cape Cod, on Nantucket Shoals, or on Georges Bank. THE HAMMER-HEADED SHARKS. FAMILY SPHYRNIDAE The peculiar hammer-shaped head, with eyes far apart, sufficiently characterizes the Gulf of Maine sharks of this family, which resembles the requiem sharks (p. 36) otherwise. Five species are known in the western Atlantic, all of them tropical-subtropical in nature. Two of these have been reported from our Gulf, but only as strays. Shovelhead Sphyrna tiburo (Linnaeus) 1758 Bonnet head shark Bigelow and Schroeder, 1948, p. 420. Garman, 1913, pi. 1, figs. 4-6 (as Ceslracion tiburo). Description. — The peculiar shovel-shaped head of this shark is enough to distinguish it readily from any other shark known from the Gulf of Maine, except for the hammerhead, from which it is readily distinguished by the fact that its head is considerably narrower, is more rounded in front, and is not deeply indented opposite each nostril; that the posterior margin of its anal fin is only weakly concave, and that the outermost four or five of its lower teeth next each outer corner of its mouth are low and rounded, not blade-like. The eyes of the shovel-head shark, like those of the hammerhead, stand at either edge of the expanded head; the first dorsal fin originates a little behind the "armpit" of the pectoral, is somewhat higher than the pectorals are long, and is higher than long; the very small second dorsal fin originates a little behind the origin of the anal fin; the upper lobe of the tail is notably long (about one-third as long as the body of the fish) and deeply notched near the tip, the lower lobe is about one-third as long as the upper lobe. The anal fin is larger than the second dorsal fin, its posterior margin is only slightly concave ; the pectorals are broadly triangu- lar, their anterior margins about as long as the distance from the level of their own points of origin to the front of the mouth. Color. — Gray or grayish brown above, and a. paler shade of the same below; some are marked with a few small dark, roundish spots along the sides. Size. — This shark is much smaller than the hammerhead, rarely exceeding 5 feet in length; it is said to reach 6 feet. General range. — Tropical-warm temperate At- lantic; from southern Brazil to North Carolina, in the west, and as a stray to southern New England and Massachusetts Bay; tropical West Africa in the east; also from southern California Figure 15. — Shovel head (Sphyrna tiburo), female, about 14% inches long, from Rio de Janeiro. A, under side of head; B, first to seventh upper teeth and first to sixth lower teeth counted from center of jaw, about 3.6 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. FISHES OF THE GULF OF MAINE 45 to Ecuador on the Pacific Coast of America, or represented there by a very close relative.' Occurrence in the Gulf oj Maine. — Our only reason for including the shovel-head here is that a stray specimen has been reported from Massa- chusetts Bay.10 It has also been taken once at Newport, R. I., and a commercial shark fishery that was carried on in Nantucket Sound in the summer of 1918 is said to have yielded six of them.11 Common hammerhead Sphyrna zygaena (Linnaeus) 1758 Bigelow and Schroeder, 1948, p. 436. • On this point, see Bigelow and Schroeder. Fishes of the Western North Atlantic, Pt. 1, 1948, p. 425, footnote 20. A shark has also been reported as tlburo from China and from the Philippines, but without convincing evidence as to its identity. >° By Garman, Mem. Mus. Comp. Zool., vol. 36, 1913, p. 181. Apparently the specimen is no longer In existence. » Persona] communication by R. U. Bodman, who operated this fishery. Description. — The very differently shaped head of the hammerhead, the shape of its anal fin with much more deeply concave posterior margin, and the fact that the outermost four or five of its lower teeth on each side are blade-like, like those nearer the center of its mouth, are ready field marks to separate the hammerhead from the shovelhead (ci. fig. 16 with fig. 15). The anal fin, too, is only about as large as the second dorsal in the hammer- head (considerably larger than the second dorsal in the shovelhead). Otherwise the positions and shapes of the fins and the size and shape of the tail are much alike in the two species. Color. — Leaden or brownish gray above, shading along the sides to pure or grayish white below; the tips and edges of the dorsal and caudal fins are more or less dusky; and the tips of the pectorals are black on some specimens. ■Figure 16. — Hammerhead (Sphyrna zygaena), female, about 27 inches long, from Nahant, Massachusetts. A, head from below, about one-third natural size; B, second upper tooth; C, ninth upper tooth; D, third lower tooth; E, ninth lower tooth; about 4 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. 46 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Size. — It appears that hammerheads are com- monly about 19 to 20 inches long when they are born; seemingly, they mature sexually at about 7 to 8 feet; they are often taken 9 to 11 feet long, and occasionally as long as 12 to 13 feet.12 Most of those that visit southern New England are less than 6 to 7 feet long, some very small indeed.13 In 1805, however, one of 11 feet was netted at Riverhead, L. I. And the fact that it contained parts of a man in its stomach has been chiefly responsible for the bad reputation of this species of hammerhead. Two other large sharks closely related to the common hammerhead, the tropical hammerhead (Sphyrna lewini Griffith, 1834)u and the great hammerhead (Sphyrna mokarran Riippell, 1835) 16 occur along the South Atlantic coast of the United States. The first of these, in particular, might stray as far as Cape Cod, as many tropical fishes do, for it has been recorded from the offing of Cape May, New Jersey. They resemble the com- mon hammerhead closely in general appearance, but both of them may be distinguished from the latter by the fact that the front outline of their head is scalloped in the midline, not evenly rounded there as it is in the common hammerhead. For further accounts of them, see Bigelow and Schroeder.16 Habits. — Since hammerheads are an accidental visitor to the Gulf, we need only remark that they are pelagic in habit, often swimming with dorsal and caudal fins out of water, and are to be met with indifferently out at sea or near land. They feed chiefly on fish, including smaller sharks (including their own kind), and sting rays, » The larger hammerheads that are sometimes reported probably are not this species, but the great hammerhead (Sihyrna mokarran, p. 46, note 16). » Dozens of little ones, of about 2H feet, have been seined on the outer shore of Long Island, N. Y., In August. '< The account of this species, In Bigelow and Schroeder, (Fishes of the Western North Atlantic, Pt. 1, 1948, p. 415) was as diplana Springer, 1941. But Frasor-Brunner (Rec. Austral. Mus., vol. 22, No. 3, 1950, pp. 213-214), has shown that it cannot be separated from the Indo-Pacific S. lewini of Griffith, 1S34, a much older name. 11 Tortonese has recently pointed out (Arm. Mag. Nat. Hist. Ser. 12, vol. 3, No. 36, 1950, p. 214) that the name tudes Valenciennes 1822 that has been applied commonly to the great hammerhead of the Atlantic actually belongs to a different species; consequently that the correct name of the great hammer- head Is mokarran Riippell, 1835, It being Identical with that Indo-Pacific species. M Fishes Western North Atlantic, Pt. 1, 1948, pp. 415, 428. the tail spines of which are sometimes found imbedded in their jaws. Like tiger sharks, they make themselves a pest in warmer latitudes where fisheries for sharks are carried on, by devouring those that they find entangled in the nets. As many as 30 to 37 embryos have been found in a gravid female, and the embryos do not develop any placental connection with the mother, so far as is known. General range. — Widespread in the tropical to warm temperate belts of the Atlantic, of the Pacific, and probably of the Indian Ocean as well; north commonly to southern New England, straying to Massachusetts Bay and as far as Halifax, Nova Scotia.17 Occurrence in the Gulf of Maine. — Hammer- heads (often in small schools) wander northward every summer, along the Atlantic seaboard; they are often to be seen basking at the surface (some harpooned) a few miles out, off Marthas Vine- yard and Nantucket; and one is occasionally taken in one or another of the fish traps near Woods Hole. But the longitude of Cape Cod so sharply bounds their yearly dispersal that the only records from the Gulf of Maine, or from Nova Scotia waters, are of stray specimens from Chatham and Provincetown on the outer shores of the Cape; of one about 27 inches long from Nahant, in the inner part of Massachusetts Bay;18 of two small ones recently from Casco Bay;19 of one taken many years ago, off Brier I., on the Nova Scotian side of the Bay of Fundy;20 of a 12-footer harpooned between Georges and Browns Banks hi August 1928 by the sword fishing schooner Doris M. Hawes; of a small one caught in Halifax Harbor, Nova Scotia, in September 1932 ;21 and of another about 21 inches long taken in a trap off Sambro Head, near Hali- fax, August 25, 1938.22 " For further details of distribution, see Bigelow and Schroeder, Fishes of the Western North Atlantic, Pt. 1, 1948, p. 442. i8 This specimen, obtained many years ago by Louis Agassiz, Is In the Museum of Comparative Zoology. » Seen In the fish market at Portland, Maine, by the late Walter H. Rich. » McKenzie, Proc. Nova Scotia Inst. Scl., vol. 20, 1939, p. 13. « Vladykov, Proc. Nova Scotia Inst. Scl., vol. 19, Pt. 1, 1935. p. 8. » McKenzie, Prov. Nova Scotia Inst. Scl., vol. 20, 1939, p. 13. FISHES OF THE GULF OF MAINE 47 THE SPINY DOGFISHES. FAMILY SQUALIDAE This group is characterized by the lack of an anal fin, combined with the presence of two dorsal fins, each of which is preceded by a fixed spine which is long and conspicuous in some, but so short in others that its presence can be detected only by touch. The teeth are alike in the two jaws in some, unlike in others. Spiny dogfish Squalus acanthias Linnaeus 1758 Dogfish; Piked dogfish; Grayfish Bigelow and Schroeder, 1948, p. 455. Garman, 1913, pi. 14, figs. 1-4. Description. — Any little gray or brownish shark, with a large sharp spine lying along the front margin of each dorsal fin, caught within the Gulf, or on the shoaler parts of the offshore fishing banks, is practically sure to be this "dog," of which there are thousands in the Gulf to every one shark of any other kind. One of its relatives, the black dogfish (p. 51), is a regular inhabitant of the deeper slopes of the offshore Banks that front the Gulf, where we also trawled more than 50 specimens of another relative Etmopterus princeps Collett 1904 during the summer of 1952. But there is no danger of confusing the common spiny -dog with either of these, for they are velvety black in color, the rear margins of their tail fins are indented near the tip, which is not the case in the spiny-dog, and each of their teeth, at least in the upper jaw (lower jaw as well in the black dog- fish) has 3 to 5 sharp points, but only one point in the spiny dog. This is a slender little shark, with flattened head and snout tapering to a blunt tip. Its first dorsal fin stands between pectorals and pelvics; its second dorsal fin is about two-thirds as large as the first; its pectorals form nearly an equilateral triangle; and its pelvics are well forward of its second dorsal fin. The dorsal fin spines lie close along the front margins of the two dorsals, the first not more tha.i one-half as long, and the second nearly as long as the front margin of their respec- tive fin, and they are very sharp. The spiny-dog has no anal fin, a lack separating it from all smooth-finned sharks known from the Gulf of Maine, except for the Greenland shark (p. 53), Dalatias (p. 55), and the bramble shark (p. 56). There is a low fold of skin on either side of the root of the tail back of the second dorsal fin, so small, however, that there is no danger of confusing it with the caudal keels of the mackerel-shark tribe. The teeth are small, their sharp points bent toward the outer corners of the mouth so that they form a nearly continuous cutting edge along each jaw. Color. — The upper surface is slate colored usu- ally, sometimes tinged brown, with a row of small Figure 17. — Spiny dogfish (Squalus acanthias), female, 27 Inches long; after Garman. A, upper and lower teeth, mid- point of mouth marked by the dotted line, about 3 times natural size. From Bigelow and Schroeder. Drawing by E. N. Fischer. 48 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE white spots on each side from the pectoral fin to abreast of the anal fin, and with a few other white spots in front of the first dorsal and behind it, also in front of the second dorsal fin. These spots are most conspicuous on small fish up to 12 or 14 inches long and they fade with growth until they disappear altogether in some specimens. The margins of the first and second dorsals, and of the caudal are more or less dusky at birth, but soon fade. The lower surface ranges from pale gray to pure white. Size. — The majority are between 8% and 13 inches long when born. Most of the adult males are from about 2 feet to a little less than 3 feet long; adult females are from a little less than 2% feet to almost V/i feet; maximum length about four feet. Mature females average 7 to 10 pounds, a few reach 15 pounds if very fat, and 20 pounds has been reported. Habits. — Much has been written of the habits of the spiny dogfish, but nothing to recommend it from the standpoint either of the fishermen or of its fellow creatures in the sea. It is one of the more gregarious of our fishes, swimming in schools or packs. Swedish fishermen assert that young dogs school separately from their parents, and it is certain that fish of a size continue to associate together as they grow, the result being that any given school runs very even, consisting as a rule either of the very large mature females, or of medium-sized fish (either mature males oi im- mature females), or of small immature fish of both sexes in about equal numbers. Apart from their general seasonal migratory movements, dogfish are governed by the move- ments of the fishes on which they prey. And re- cent marking experiments have shown that some of them cover long distances in their wanderings, for two tagged near St. Johns, Newfoundland, in mid -July 1942 were recaught off Cape Ann,23 one on November 23, 1943, the other on Decem- ber 4 of that year,24 while others from the same tagging experiment were caught within the Gulf of St. Lawrence.25 Fortunately they seldom stay long in one place, but there is seldom, if ever, a time during the summer when they are not com- mon on some part of the Gulf of Maine coast. So erratic are their appearances and disappearances 33 About 14 miles offshore. :* On Middle Ground about 25 miles off Cape Ann. "■> Tcmpleman, Fish. Res. Bull., Newfoundland Dept. Nat. Res., No. 15, 1944, pp. 67-69. that where one has good fishing today he may catch only dogfish tomorrow and nothing at all the day after, the better fish having fled these sea wolves and the latter departing in pursuit. The dogfish use their back spines for defense, curling around in a bow and striking, which makes them hard to handle on the hook. It is probable, too, that the spines are slightly poisonous, general report to this effect being corroborated by the fact that the concave surfaces are lined with a glandular tissue resembling the poison glands of the venomous "weever" (Trackinus draco) 26 of Europe. Voracious almost beyond belief, the dogfish entirely deserves its bad reputation. Not only does it harry and drive off mackerel, herring, and even fish as large as cod and haddock, but it destroys vast numbers of them. Again and again fishermen have described packs of dogs dashing among schools of mackerel, and even attacking them within the seines, biting through the net, and releasing such of the catch as escapes them. At one time or another they prey on practically all species of Gulf of Maine fish smaller than them- selves, and squid are also a regular article of diet whenever they are found. Dogfish arc also known to take worms, shrimps, and crabs. And when they first arrive at Woods Hole in May they are often found full of Ctenophores, being one of the few fish that eat these watery organisms. Often, too, they bite groundfish from the hooks of long lines, or take the baits and make it futile to fish with hook and line where they abound. Fishermen are familiar with the fact that the female spiny dog bears "living" young (this has been known since the days of Aristotle). The eggs are large, well stored with yolk, and during early stages those in each oviduct (so-called "uterus") are contained in a horny capsule that breaks down later, leaving the embryos free in the "uterus," to which they have no placental attach- ment. The number in a litter is commonly 4 to 6 ; sometimes as many as 8 to 11, or as few as 2. According to recent studies, the females carry their young for 18 to 22 months. Accordingly, the adult females caught in our Gulf contain either very early embryos, averaging only about three- fourths of an inch in length by September, or "■> Evans (rhilos. Trans. Royal Soc., London, Ser. B, vol. 212, 1923, pp. 8, 27) describes the spines and gives clinical records of the effects of wounds indicted by them. FISHES OF THE GULF OF MAINE 49 much larger ones, 7 to 11 inches long by that month; i. e., nearly ready for birth. Similarly, we have taken females with embryos 9 to lOK inches long in November, on the Cholera Bank near New York Harbor. And it now seems established that most of the young are born on the offshore wintering grounds.27 But dogfish so small as evidently to have been newborn are oc- casionally taken along southern New England and in the Gulf in early summer; also on Nantucket Shoals where the Albatross II trawled some of 10^ to 13 inches in August, showing that the season of production extends through the spring, or even into the summer as in 1905 when females taken off Gloucester in July gave birth to young on capture.28 General range. — Both sides of the North Atlantic, chiefly in the temperate and subarctic belt; also both sides of the northern Pacific; 29 and repre- sented in the corresponding thermal belt of the southern hemisphere by a relative (or relatives) so close that it is doubtful whether they differ in any recognizable way from the spiny-dog of the north. Occurrence in the Gulf of Maine. The spiny dog- fish ("dogfish" or "dog" in common parlance) makes up for the comparative rarity of other sharks in the Gulf of Maine by its obnoxious abundance. To mention all the localities from which it has been reported there woidd be simply to list every seaside villnse and fishing ground from Cape Cod to Cape Sable. It is as familiar, too, on the offshore banks as it is along the coast; also along outer Nova Scotia, in the Gulf of St. Lawrence, on the Grand Banks, and along the east coast of Newfoundland to southeastern Labrador. There is no record of it from the North American coast north of Hamil- ton Inlet, but stray specimens have been taken along the southwest coast of Greenland.30 To the southward, fishermen are familiar with it in season ** Females that we saw trawled oft Block Island in 60-65 fathoms In late January 1950, pave birth to young on the deck of the vessel. » Mclntiro, Kept. Comm. Fish. Game Massachusetts, (1905) 1906, p. 108. » We have found no consistent differences between North Atlantic and North Pacific specimens. For further discussion of this point, and further details as to the occurrence of the spiny-dog in the two sides of the North At- lantic, see Bigelow and Schroeder (Fishes of the Western North Atlantic, Pt. 1, 1948, pp. 453, 463). <• Jensen (Selachians of Greenland, Mindeskr. Japetus Steenstrup, Pt. 2, No. 30, 1914, p. 7) lists several definite records of this species at Sukkertoppen and near Holsteinborg, West Greenland. as far as Cape Lookout, N. C, and a few stray even to southern Florida and to Cuba.31 Dogfish are seasonal visitors on the coast, strik- ing in about as early along New Jersey (March), and even on Georges Bank (March-AprU) , as along North Carolina. In the inner parts of the Gulf of Maine the date of the first heavy run of dogfish varies widely from year to year and from place to place. We have not heard of them there before May. But the period of freedom may close as early as the last half of the month, in some years. In 1903, for example, they had appeared as far north as Penobscot Bay by the middle of May. And while it is not untU June that they usually arrive in numbers in the Massachusetts Bay re- gion, it is sometimes impossible to set gill or drift nets anywhere between Cape Cod and Cape Elizabeth after the first days of that month, so numerous are they. In 1913 the first heavy run of dogfish struck Ipswich Bay on June 14, and they appeared there at about the same date in 1905, but there is much local variation in this respect. In 1903, for example, they did not appear until early July at Provincetown, though swarming a month earlier in Massachusetts Bay, in Ipswich Bay, and off Penobscot Bay. But in 1920 they appeared at Provincetown by May 25 to 26 when one set of mackerel traps caught 23 barrels of them, and another 21 barrels. They usually strike in all along the northern Maine and west Nova Scotia coasts by the end of June; but few are seen until late in July in Passamoquoddy Bay. They have been recorded as early as July 1 near Raleigh, on the Newfoundland side of the Strait of Belle Isle, but they are not caught in any numbers in the inner parts of the Gulf of St. Lawrence until well into July, and they have not been reported from southeastern Labrador until early m Sep- tember.32 In the southern part of its range, from North Carolina to New York, the spiny dogfish is a spring and autumn transient only. West of Cape Cod (at Woods Hole, that is, and along Long Island) si Repeated reports of it as plentiful along eastern Florida seem to have referred to some other shark; the basis for similar reports from Cuba and Trinidad doubtless was the Cuban dogfish, Sguaius cudfnji* Rivero. >■ See Templeman (Res. Bull. 15, Newfoundland Dept. Nat. Res., 1941, pp. 56, 64) for dates of arrival around the coast of Newfoundland in different years. 50 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE they are transients mostly, passing north in spring and south in autumn, though some summer there; even considerable numbers in some years.33 And it seems that most of them withdraw from Mas- sachusetts Bay also during the warmest period, for few are taken there between June and Septem- ber. But they continue present all summer along outer Cape Cod, and here and there throughout the northern and eastern parts of the Gulf, in varying abundance. Most of the dogfish take their autumnal depar- ture from the inner parts of the Gulf during October, few being caught on the coast north of Massachusetts Bay after November 1. But they sometimes stay later, as in 1903 (a big dogfish year), and again in 1942, when they were abundant along the outer shore of Cape Cod as late as the first week of November. Ordinarily none are caught within the Gulf of Maine north of Georges Bank in winter, but this has its exceptions. In 1913, for example, a few were caught 20 miles off Cape Ann on November 19 to 24, many near Boon Island from December 5 to 13, and on Jeffreys Ledge on December 11 and 12. In 1882, schools were reported off Portsmouth, N. H., even as late as February, an exceptional event. Dogfish appear earlier in spring and linger later into the winter on Georges Bank (fig. 18) than in the inner parts of the Gulf. It is safe to say that there are few there in March, the earliest definite record (obtained during the investigations of 1913, only year of record, being of 25 fish caught on the "winter cod ground" east of the shoals (long, about 67°, lat. about 41°40') between the 20th and the 22nd, and of 46 from the same gen- eral region from the 27th to the 30th, while some are trawled there all summer. In 1913, a few were taken in November and in December; a few also on the southern part of the Bank (lat. about 41°, long, about 67°30') on January 20 to 22 in 1914. Apparently dogfish reach Browns Bank later than they do Georges, for none was taken there on April 14 in 1913, though they are only too plentiful there in summer. It is also likely that they depart earlier, although a few lingered as late as December 3 to 12 on Western Bank off Halifax in that year. - For details, see Blgelow and Schroeder, Fishes of the Western North Atlantic. Pt. 1, 1948, p. 464. 8000 — 7500 - 7000 - 6500 - 6000 - £ 5500 - S 5000 - £ 4500 - £ -40 0 0 - | 3500 ■ 2 3000 - A § Z500 - a= 2000 - 1500 - 1000 - 500 - ■', ' ', rL-L"-, M-. ^ 1, I I MAR. APRIL MAY JUNE JULY AUG. SEPT. OCT. _J_ J- N0V. DEC. Figure 18. — Numbers of spiny dogfish caught on certain otter trawling trips to Georges Bank, during the dif- ferent months of 1913. It now seems certain that the spiny dogfish winter chiefly in deeper water offshore, for con- siderable numbers have been trawled at that season on the outer part of the continental shelf off Block Island, in 50 to 65 fathoms, where we saw several hundred (200 in one haul) trawled during the last week of January 1950; off New York in November and January; 34 also in Febru- ary off the Middle Atlantic coast in 16 to 70 fathoms, south as far as the offing of Cape Hat- teras. On the other hand, the fact that numbers of them have been found washed on shore in January on the southwest coast of Newfoundland suggests that some of those that summer in that general region may survive the winter in the deep trough of the Gulf of St. Lawrence. They are usually so thin when they reappear on the coast in spring as to suggest that they feed but little during the winter. This is the only Gulf of Maine shark that even remotely rivals the important food fishes in num- bers. Unfortunately, the statistics of the com- mercial landings for American waters do not afford any information in this regard. But spiny dogs must be plentiful indeed in our waters when they can sometimes be caught as fast as they can " Mr. Thomas Quast informs us that many were taken from the schooner Victor, long-lining for tile fish, on the outer edge of the continental shelf, off New York, during the second week of January 1928. FISHES OF THE GULF OF MAINE 51 be hauled in; when a long line, with 1,500 hooks, has been known to bring in a dogfish on nearly every hook; and when an average trawl catch of 6,000 to 8,000 per trip was made on Georges Bank in 1913 during their season of abundance. At the time of the 1904 to 1905 peak it was estimated from recorded catches that at least 27,000,000 were being taken yearly off the coast of Massachu- setts.35 More precise information from waters farther north is that 10,391,000 pounds, or 2 to 3 million individual dogfish, were caught in 1938, in Pla- centia Bay, Newfoundland, with no apparent effect on their numbers.38 In short, they may be as plentiful in our Gulf as they are on the Cornish coast, where the record catch of 20,000 in a single haul was made many years ago. Spiny dogfish appear to have been more numer- ous in the Massachusetts Bay region during the last quarter of the past century and during the early nineteen hundreds than they had been pre- viously. At Woods Hole, on the contrary, they are said to have been much more plentiful before 1887 than they have been at any time since. To a certain extent, of course, reports of fluctuations in abundance from year to year must be discounted as reflecting the movements of the great schools that may visit one part of the coast one summer and another part the next, not a general altera- tion of the stock. But the many fishermen who reported to the Massachusetts Commissioners in 1905 were unanimously of the opinion that dogfish had multiplied steadily for 20 to 30 years past, and reports from British coasts were to the same effect. Perhaps the years 1904-1905 marked the apex of this wave of multiplication; at any rate dogfish were reported as distinctly less troublesome to the mackerel netters in 1913 than they had been previously. And little complaint has been made of them in late years. But it is not safe to conclude from this that the stock is at a low ebb at present, for it was the hand- and long-line fishermen that suffered most from them; and it is only as they increase the amounts of trash fish dumped overboard that the dogfish bother the otter-trawlers. Importance. — During the years when the ground fishery was chiefly by hook and line, fishing often was actually prevented by dogfish in Massachu- « Report, Comm. Fish and Game, Mass., (1906), 1907, p. 20. » Tompleman, Newfoundland Fish. Res. Bull., 15, 1944, p. 72. setts and Ipswich Bays, unless cockles (Polynices) were used for bait, for dogfish do not take these. The general replacement of hook and fine fishing by the otter trawl has put an end to widespread complaints on this score. But when schools of dogfish get into a net or seine, they so snarl the twine that disentanglement and repair may be the work of days. And it has been estimated that they may do some $400,000 worth of damage annually to fishing gear, and to fish caught by such gear, off the coast of Massachusetts alone, during their peaks of abundance there. With the dogfish so plentiful and destructive, it is no wonder that serious efforts have been made to make them a source of revenue instead of a dead loss. And the dog is a far better food fish when fresh than is generally appreciated, as is evident by the large amounts landed in the fishing ports of northwestern Europe. But it has never been in any demand for the table, on our coasts, though it would offer a large supply of cheap food were a satisfactory method found for canning it. During their more recent periods of plenty various efforts have been made to utilize them on a large scale for fertilizer, and for liver oil (it compares favorably with cod for vitamin A, though it is much poorer in vitamin D), on the Atlantic coasts of the United States and Canada; however such developments have been short-lived. And dogfish have not been of sufficient value up to the present to compensate for a hundredth part of the damage they do.37 Black dogfish Centroscyllium fabricii (Reinhardt) 1825 Bigelow and Schroeder, 1948, p. 482. Garman, 1913, pi. 10, figs. 5-8. Description. — The notched margin of the upper tail lobe distinguishes this shark at a glance from the spiny dogfish, with which it agrees in having a long pointed spine at the front edge of each dorsal fin. It differs further from the common dogfish in that its dorsal spines are deeply grooved along each side, whereas in the "dog" they are rounded; in the location of the pelvic fins, the rear axils of r For further discussion of the damage done by dogfish and of their com- mercial possibilities, see Ann. Rept., Comm. Fish. Game Mass. (1905), 1906, pp. 97-169; Rept. U. S. Comm. Fish. (1902) 1904, pp. 228-229; Field, Doc. 622, Rept. U. S. Comm. Fish. (1906) 1907, pp. 21-23; Field, Bull. TJ. S. Bur. Fish., vol. 28, 1910, pp. 243-257; Mayor, Contr. Canad. Biol. (1918-1920) 1921, pp. 125-135; and Templeman, Newfoundland Fish Res. Bull. 15, 1944 52 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 19 -Black dogfish (Cenlroscyllium fabricii) , female, about 25 inches long, from the southeast slope of Georges Bank A first three upper teeth counted from center of jaw; B, twentieth upper tooth; C, first three lower teeth; D lower sixteenth tooth; about 5 times natural size. From Bigelow and Schroeder. Drawings by E. N. 1'ischer. which stand almost directly under the front origin of the second dorsal fin instead of some distance in front of the latter; in its small pectorals of rounded outline; in the shapes of its teeth, each of which has 3 or 5 sharp points; in its broad rounded snout; and in its very dark color. Like the spiny dogfish, it lacks an anal fin. Size. — Adult specimens range from 2 to 3}i feet in length, that is, about the same size as the spiny dogfish. Color. — Uniform dark brown to black, below as well as above. Habits. — In West Greenland waters cephalopods, pelagic crustaceans, and medusae have been found in their stomachs, and females have been taken with embryos in February. Perhaps they are luminescent, for their skins bear minute deeply pigmented dots, suggesting the light organs of the brilliantly luminescent shark Isistius brasili- ensis. General range. — Northern North Atlantic; Faroe Bank, Faroe-Shetland Channel and Iceland in the east; West Greenland; Davis Strait; and outer slopes of the fishing banks in the west, southward to Georges Bank; chiefly deeper than 150 fathoms. Occurrence in the Gulf of Maine. In the years when a long line fishery for halibut was carried on regularly, black dogfish were often caught along the slopes of the offshore Banks, from Grand to Browns and to the eastern part of Georges, if sets were made down to 200 fathoms or deeper. And while they dropped out of sight with the general abandonment of that fishery, no doubt they are as plentiful now as formerly, for we trawled about 100 of them, 6 to 24)£ inches long, off southwestern Nova Scotia, at 290 to 580 fathoms, on the Caryn of the Woods Hole Oceanographic Institution, in June 1949. How far they range to the west and south, at the appropriate depths, is not known.38 Portuguese shark Centroscymnus coelolepis Bocage and Brito Capello, 1864 Bigelow and Schroeder, 1948, p. 494. Garman, 1913, pi. 14, figs. 5-8. Description.— This shark can be identified easily by the fact that while its general appearance (especially the absence of anal fin, the situation of its pelvics far back under the second dorsal, and its rather stout form and blunt snout) might lead a hasty observer to think he had caught a small Greenland shark; more careful examination, by touch if not by eye, would reveal a short spine close in front of each dorsal fin. The first dorsal fin is smaller than in any of our sharks except in the "Greenland," (p. 53), and in Dalatlas licha (p. 55), the second dorsal is a little larger than the first, and the pelvics are larger than either of the dorsals. The tail is noticeably short and broad and the rear edge of its upper lobe is notched. The teeth are different in the two jaws; narrow, pointed, and of the seizing type in the upper; broader, ob- long, with a notch on the outer side near the tip, and forming a continuous cutting edge in the lower. The dermal denticles are flat, scale-like, closely overlapping, and clothe the entire trunk. Color. — Dark chocolate brown, belly as well as back and fins. M Its range has been said to extend to New York, but without supporting evidence; and report of a young one from the Gulf of Mexico (Ooode and Bean, Smithsonian Contrib. Knowledge, vol. 30, 1895, p. 11), probably was based on some other shark. FISHES OF THE GULF OF MAINE 53 Figure 20. — Portuguese shark (Centroscymnus coelolepis), female about 42)4 inches long, off Banquereau Bank. A, upper teeth, and B, lower teeth from center of mouth, about 3.4 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. Size. — Adults measure from 3 to VA feet long, as they are caught. Garman records one 44 inches long taken off the coast of New England. About 9 inches is the smallest recorded.3' Habits. — Little is known of its habits beyond the fact that it is a deep-water species, and that it was caught regularly by Portuguese fishermen with hand lines, a fishery that Wright 40 described as follows: Some GOO fathoms of rope were let out, the first 30 or 40 fathoms of which had fastened to it at intervals of a fathom a series of small ropes, on each of which was a large hook baited with a codling. This fishing tackle remained below for about two hours, when they commenced to haul it in. When it arrived at the last few fathoms, they pulled in, one after another, five or six specimens from 3 to 4 feet long. The species was the Centroscymnus coelolepis Bocage and Capello. These sharks, as they were haulfd into the boat, fell down into it like so many dead pigs. Thirteen to 16 young have been found in fe- males caught off Portugal. General range. — This deep-water shark, origi- nally discovered off Portugal, has since been taken at various other eastern Atlantic localities.41 Defi- nite records of it for the western Atlantic arc from the slopes of the Nova Scotian Banks and of Georges, at depths of 180 to 250 fathoms, perhaps 15 to 20 specimens in all. But Goode and Bean's*2 old characterization of them as abundant on the Banks at 200 fathoms and deeper presents its local status more correctly, for fishermen long lining for halibut often caught one or two a trip in the deeper gullies between the offshore Banks. THE GURRY SHARKS. FAMILY DALATIIDAE The gurry sharks, like the spiny dogfishes, lack anal fins, but they have no spines in their dorsal fins. The teeth in the upper jaw are noticeably unlike those in the lower. Greenland shark Somniosus microcephalus (Bloch and Schneider) 1801 Sleeper shark; Gurry shark; Ground shark Bigelow and Schroeder, 1948, p. 516. Garman, 1913, pi. 15, figs. 4-6. Description. — The Greenland shark is notable n A male 228 ram. long, examined by us, in the U. S. National Museum from the continental edge south of Nantucket. <• Ann. Mag. Nat. Hist., Ser. 4, vol. 2, 1868, p. 426. for its small dorsal fins, without spines, the second dorsal being of about the same size as the first, and for small pectorals hardly larger than the pclvics, coupled with the absence of an anal fin and with a tail of more fish-like form than that of most other sharks except for the mackerel-shark tribe. Bearing these points in mind, particularly the absence of an anal fin and of dorsal spines, it cannot be confused with any shark common in our Gulf. And while it resembles the rare Portuguese shark in the sizes and relative situa- « Iceland; Faroe Bank; Madeira; Azores; Morocco; Cape Verde I.: For key to other species of the genus, see Bigelow and Schroeder, Fishes Western North Atlantic, P. 1, 1948, p. 494. « Smithsonian Contrib. Knowledge, vol. 30, 1895, p. 14. 54 FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE Figure 21. — Greenland shark (Somniosus microcephalus) , female, about 5 feet 9 inches long. Teeth at center of mouth; lower teeth from midway along the jaw of a speci- men about 11 feet long from the Gulf of Maine, about 1.8 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. tions of its dorsal and anal fins, in its general form, and in its teeth, it is easily separable from the "Portuguese," both by lacking any trace of spines in its dorsal fins, by its thorn-like and loosely spaced dermal denticles, and by its more lunate tail. It also grows much larger than the Portu- guese shark. We need only note, further, that while its upper teeth are narrow and awl-like, its lowers are broad, squarish, forming a nearly continuous cutting edge, with the single cusp directed sharply outward ; that its gill openings are short and located low down on the sides of the neck; that its eyes are very small; and that it is stout shouldered, with blunt rounded snout, as Scoresby pictured it more than a century ago.43 Color. — Blackish, coffee brown, or ashy-, pur- plish-, or slate gray, below as well as above; changing to bluish gray if the epidermis is rubbed off, as is apt to happen when one is caught; the back and sides are marked with many indistinct dark crossbars on some specimens. Size. — This is one of the larger sharks. It is said to grow to a length of 24 feet, but 21 feet is the largest of which we find definite record,44 and 16- to 18-footers are unusual. One of 16K feet was reported from the Grand Banks in 1934; one of « Arctic Regions, 1820, vol. 2, pi. 15, figs. 3 and 4. « Jenkins. Fishes British Isles, 1925, p. 325. 16 feet off Portland, Maine, in 1846; one of about 15 feet off Cape Ann in 1849; and another of about that same size was caught on a long line north of Cape Ann in February 1931. Perhaps 8 to 14 feet is a fair average for adults, that is not often ex- ceeded among the hundreds caught annually off West Greenland and around Iceland. The 21- foot British specimen mentioned above was said to weigh about 2,250 pounds; two Gulf of Maine specimens, each about 1 1 feet long, weighed about 600 and 650 pounds, respectively. Habits. — Off Greenland, and along the Labrador coast, the Greenland sharks tend to approach the surface in winter, often coming right up to the ice. But most of them withdraw in summer to 100 fathoms or deeper. And the few that visit our Gulf appear to hold rather closely to the bottoms of the deeper troughs, though a stray may come so close to the shore now and then, and into water so shoal as to blunder into a fish weir; one such event is on record for Passamaquoddy Bay. This is one of the most sluggish of sharks, offering no resistance whatever when hooked, and it is entirely inoffensive to man.45 But it is ex- '* Tales to the effect that it attacks Greenlanders in their kyaks are appar- ently mythical, and Doctor Porsild, Director of the biological station at Disko, said that the Eskimos do not fear it as they do the killer whale; nor is there any authentic instance on record of a shark attacking a human being near Iceland. FISHES OF THE GULF OF MAINE 55 tremely rapacious. It devours any carrion ea- gerly, such as whale meat, blubber from whaling operations, or the carcasses of young seals that are left on the ice off the Newfoundland-Labrador coasts. And its habit of gathering when there has been a big killing of narwhals in Greenland waters is proverbial. Apart from carrion (which cannot be available except on rare occasions), its diet includes a wide variety of fishes, large and small. Seals are a favorite food, and in view of its sluggishness, it is somewhat astonishing that it should be able to capture prey as active as seals, halibut, and salmon. The specimen from Cape Cod Bay, mentioned above, contained half a dozen flounders and a large piece (with hide and hair) that had been bitten out of the side of a seal. It is also known to eat crabs, large snails, even medusae. Objects as large as an entire reindeer (without horns), a whole seal, a 3-foot cod, and a 39-inch salmon, found in Greenland shark stomachs, give some measure of their appetite. In line with this, they will bite on any fish or meat bait, the more putrid and ill smelling the better. Large numbers of soft eggs, without horny cap- sules, ranging in size up to that of a goose egg, have been found repeatedly in female Greenland sharks, but never any embryos, suggesting that this may be an egg-laying species.48 General range. — Northern Atlantic, from Polar latitudes south to the North Sea and accidentally to the mouth of the Seine and perhaps to Portugal in the east; south regularly to Newfoundland and the northern part of the Gulf of St. Lawrence in the west, and less commonly to the Gulf of Maine. It is represented in the Mediterranean region, in the North Pacific, and in the sub-Antarctic by forms that appear to be distinct, though closely allied to it.47 Occurrence in the Gulf of Maine. — Although there is no reason to suppose that the Greenland shark ever appears in our Gulf save as a straggler from the north, its presence there has been signalized on a number of occasions. Two specimens, for example, were taken in the neighborhood of St. Andrews in 1915 (one caught in a weir and the other on a long line). It has been reported off Eastport; off Cape Elizabeth whence 6 were landed « The Mediterranean Somniosng rostratus, on the contrary, bears living young. <7 For recent discussion of the species of Rom-niosus, see Bigelow and Schroeder. Fishes Western North Atlantic, I't. 1, 1948, p. 515. at Portland between 1925 and 1948;48 on Jeffreys Ledge, where one of about 15 feet was caught on a long line, on February 16, 1931 ;49 near Cape Ann; off Marblehead and Nahant; in Massachu- setts Bay; off Barnstable in Cape Cod Bay; at Provincetown; and in Cape Cod Bay off the en- trance to the Cape Cod Canal, where one between 10 and 11 feet long was taken by a trawler in April 1924, landed in Boston and identified by us. Recorded captures in the Gulf include small specimens as well as large, and have been for all four seasons of the year, suggesting that when a Greenland shark does stray southward to the Gulf, it may survive there for years. The local records are distributed so widely as to show that an odd specimen is to be expected anywhere in the deeper parts of the Gulf. And rumor has it that they were more numerous in our waters in early colonial times when Atlantic right whales were still being killed in numbers off the Massa- chusetts coast.60 Commercial importance.' — This shark is not plen- tiful enough in our Gulf to be even of potential value. But it has long supported a fishery off northern Norway, around Iceland, and in West Greenland waters, chiefly for its liver oil.61 In Greenland the flesh is dried also for dog food, and to a small extent in Iceland for human consump- tion. But it produces an intoxicant poisoning if eaten fresh, though it is wholesome if dried.62 Dalatias licha (Bonnaterre) 1788 Bigelow and Schroeder, 1948, p. 502. Description. — This shark resembles the Portu- guese shark in the relative sizes and positions of its fins; also in its scales. But its dorsal fins do not have any trace of spines, while the serrate margins of its lower teeth, in combination with their triangular shape, mark it off from any other shark without an anal fin that is known yet from the North Atlantic. Its trunk is rather slender, its snout short and bluntly rounded, and the lower-anterior corner of its tail fin is not expanded as a definite lobe. Its upper teeth are slender, awl- " Reported to us by the late W. W. Rich. 4i This one was landed in Boston, where we saw it. 80 When they gather to feed on whale, narwhal, and seal carcasses in their northern home, they may linger for a long time in tbe vicinity. •' The annual catch off West Greenland was around 32,000 during the first decade of the present century. M For accounts, see Jensen, 1914 (Selachians of Greenland, Mindesk. Jap. Steenstrup, vol. 2, No. 30, 1914, p. 12); also Clark (Science, N. Scr., vol.41, 1915, p. 795). 56 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE A A Figure 22. — Dalatias (Dalatias licha), female, 58 inches long, from Georges Bank. A, upper teeth and B, lower teeth from central part of mouth, about 1.5 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. shaped, curving somewhat outward toward the corners of its mouth; but the lowers are erect, broadly triangular, with serrate edges. Color. — Dark chocolate, cinnamon, or violet brown below as well as above; the upper surface sometimes with poorly defined blackish spots; the dorsal and pectoral fins with pale or whitish edges, the tail tipped with black. Size. — Most of those caught are between 40 and 60 inches long; 72 inches is the longest re- corded so far. The Gulf of Maine specimen illus- trated in figure 22 was about 5 feet long and weighed 23 K pounds, gutted. General range. — Eastern Atlantic, from tropical West Africa to the Irish Atlantic slope; recorded once from the American coast. Occurrence in the Gulf of Maine. — Our only rea- son for mentioning this shark is that a female, about 5 feet long, was taken on the northern edge of Georges Bank on August 19, 1937 (fig. 22) .« THE BRAMBLE SHARKS. FAMILY ECHINORHINIDAE The only living representative of this family (it is represented among the tertiary sharks) re- sembles the Greenland shark and its allies in lacking both anal fin and dorsal spines, but its teeth are alike in the two jaws. Bramble shark Echinorhinus brucus (Bonnaterre) 1788 Spiny shark Bigelow and Schroeder, 1948, p. 527. Description. — The location of the first dorsal fin above the pelvics instead of about midway between the latter and the pectorals, and the very different shape of its tail fin (cf. fig. 23 with fig. 21), are the most conspicuous field marks separating this shark from the Greenland shark. Brucus also differs from the latter in that the teeth are alike in the two jaws, instead of unlike, and that the skin of its back and sides is sparsely strewn with large scales with either one or two sharp points. " Recorded by Nichols and Firth, Proc. Biol. Biol. Soc. Wash., vol. 52. 1939, p. 85. O t> « a V.3 m Figure 23. — Spiny shark (.Echinorhinus brucus), eastern Atlantic specimen about 3 feet long. From]Bigelow and Schroe- der. Drawing by W. P. C. Tenison. FISHES OF THE GULF OF MAINE 57 Color. — Described as dark gray, olive or brown above, with metallic reflections, and with or witbout darker blotches; as paler brown or gray to white below. The scales have been described as luminescent,64 but there are no special luminous organs. Size. — The largest of which we have found a record (a specimen from British waters) was 9 feet long. One 8 feet 4 inches long weighed about .300 pounds. General, range. — Eastern Atlantic (including the Mediterranean) from tropical West Africa to Ireland and the North Sea, and accidental in the western Atlantic; represented in South Africa; off California; in the Hawaiian, Japanese, and Australo-New Zealand regions, and in Arabian waters by forms that probably cannot be dis- tinguished from brucus of the Atlantic. Occurrena in the Gulf oj Maim. A single specimen of this little known shark came ashore at Provincetown in December 1878. This and one taken near Buenos Aires more recently 5S are the onlv records of it from the western Atlantic. Torpedoes, Skates, and Rays. Order Batoidei This tribe falls into four groups, so far as the Gulf of Maine fauna is concerned: first, the torpedoes (family Torpedinidae) , with large caudal fin, interesting because provided with electric organs capable of giving a strong shock; second, the skates (family Rajidae), with very thin bodies, comparatively short tails without tail spines, and only a trace of caudal fin; third, the sting rays (families Dasyatidae and Rhinopteridae), with long whiplike tails armed with a stiff saw-edged spine (or spines); and fourth, the devil rays •< Cornish, Zoologist, Sor. 2, vol. 10, 187S, p. 4501. (Mobulidae) with two ear-like fins extending forward from the front of the head. Most of our common species belong to the second group. Among torpedoes, skates, and rays, fertiliza- tion is internal as it is among sharks, and the modification of the posterior edges of die pelvic (ins into rodlike semitubular claspers (the copula- ton organs) distinguishes males and females at a glance. Some bear "living" young, ready for independent existence; others lay eg^s. » Berg, Com. tctlol, Comm. Mus N'ac. Buenos &irea, ral. I. No. 1. 1898, p. hi -!. KEY TO GULF OF MAINE SKATES AND RAYS The front, (if the head bears a pair of separate, ear-like fins, extending forward The front of head does not bear a pair of separate ear-like fins extend inn forward There is a large triangular caudal fin, as well as two well developed dorsal fins on the tail. There is no distinct caudal fin; the dorsal fins, if any, arc very small . . No long dorsal spine on tail - - There is a long saw-edged dorsal spine (or spines) on the tail The upper surface of the disc is marked with conspicuous black rosettes . . The markings on the upper surface of the disc are not in the form of black rosettes. Devil ray. p. 77 ...2 Torpedo, p. 58 ..3 Common skale< 4 ,.11 leopard skate, p. 66 ...5 There are no conspicuous thorns along the mid-dorsal zone of disc between the spiracle-; ami the base of tail; the lower surface of disc is marked with black dots or dashes, marking the openings of the mucous pores. Barndoor skate, medium sized and large specimens, p. 61 There are one or more rows of conspicuous thorns along the mid-dorsal zone of disc rearward from the spiracles; the lower surface of disc is not marked witli black dots or dashes — 6 There are no large thorns on the rear %-}', of tail . . - Smooth-tailed or Prickly skate, p. 70 There are one or more rows of large thorns along the rear part of tail a< well as fan her forwar I along it__ . 7 There are no large thorns on upper side of disc between the spiracles and I he level of axils of pectoral fins. Barndoor skate, very small specimens, p. 61 The uppersideofdi.se, rearward from spiracles, has more or fewer hour l horns . 8 The thorns of the midrow on the tail are much larger and more conspicuous than any other thorns on the tail, and not more than 9 or 10 in number Thorny skate, p. 72 No one row of thorns along the tail is much larger or more conspicuous than the other thorns on the tail; there are at least 15 thorns in each of the rows along tail — 'J 210941—53 5 58 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE 9. There is only one row of large thorns along the midzone of the disc from the nape to the level of the axils of the pectoral fins; the first and second dorsal fins are separated by a definite space or at least by 1 or 2 thorns; the forward angle of the disc is less than 110°; the upper surface of the disc is marked with short dark bars as well as with roundish spots. _ - Brier skate, p. 65 There are at least three rows of thorns along the midzone of the disc from the nape to the level of the axils of the pectoral fins; the first and second dorsal fins are not separated by a definite interspace or by a thorn or thorns; the forward angle of the disc is more than 125°; the upper surface is not marked with dark bars though it is variously spotted 10 10. Upper teeth in at least 72 series, most often 90-100; does not mature sexually until at least 26 inches long. Big skate, p. 63 Upper teeth in not more than 66 series and usually less then 54; matures when only 18-20 inches long. Little skate, p. 67 11. There is a small dorsal fin on the upper side of the tail, in front of the spine (or spines) ; the crown of the head is high- domed, with the eyes and spiracles on the sides; there are only 7-9 series of teeth in the form of large flat grinding plates _ Cow nosed ray, p. 76 There is no dorsal fin on the tail; the crown is low, flat, and with the eyes and spiracles on the upper surface; the teeth are in many series, in mosaic arrangement Sting ray, p. 74 THE TORPEDOES OR ELECTRIC RAYS. FAMILY TORPEDINIDAE The trunk of the electric rays has the form of a flattened, roundish or oval disc, fleshier toward the margins than it is in other Gulf of Maine skates or rays, and the body is softer. The tail, too, is broader and shorter; there are one or two relatively larger dorsal fins on the tail, and the latter ends in a well -developed caudal fin also. The most interesting feature of the electric rays is that they have two large electric organs, each of which occupies one side of the front part of the disc. In the only Gulf of Maine species the two organs together make up about one-sixth of the total weight of the fish. Torpedo Torpedo nobiliana Bonaparte 1835 Electric rat; Numbfish, Crampfish Bigelow and Schroeder, 1953, p. 96. Carman, 1913, pi. 25, fig. 2, as Narcacion nobilianus. Description. — No one would be apt to mistake a Figure 24. — Torpedo (Torpedo nobiliana), male, about 33 inches long, off Plymouth, Massachusetts. A, side view of caudal fin; B, teeth 3 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. FISHES OF THE GULF OF MAINE 59 torpedo for any other Gulf of Maine skate or ray, the rounded outline of the disk and the large caudal fin identifies it at a glance. Furthermore, its skin is soft and naked, without the spines or thorns so characteristic of all our common skates. The disk is roughly subcircular, truncate in front, and somewhat broader than long. The eyes are very small and set far forward. The two dorsal fins, of which the first is the larger, stand on the forward end of the tail, the first, indeed, partly above the bases of the pelvic fins, and they are separated by an interspace nearly as long as the second dorsal fin. The tail fin is of ordinary fish form, triangular and nearly as long as it is deep. The tail is shorter than in the skates for it occupies only about two-fifths the total length of the fish, measured from the cloaca. The teeth are small, with sharp curved points, and are in about 60 series, with up to 7 rows exposed and functioning at one time. Color. — Dark chocolate to purplish brown above, some with a few obscure darker spots; lower surface white except that the edges of disk, fins, and tail are of the same dark tint as the upper side. Size. — Adult torpedoes are usually 2 to 5 feet long or a little longer, and heavy for their size. Specimens taken at Woods Hole average about 30 pounds, while most of those taken anywhere on our Atlantic coast weigh less than 75 pounds. But we have seen one only about 4 feet long from Chesapeake Bay that weighed about 100 pounds; one of 144 pounds was brought from Nantucket to the U. S. Fisheries Station at Woods Hole many years ago; and the heaviest taken near Provincetown were estimated long ago by a fisherman of keen observation as 170 to 200 pounds. Habits. — The most interesting thing about the torpedo is its ability to give electric shocks of considerable strength to anyone touching it. The statement, even, has long been current that the shock from a large one in rested condition may be strong enough to throw a full grown man to the ground. And the story is told of a dog which was in the habit of wading on a Cape Cod beach in shoal water to catch flounders, but was so shocked by a torpedo that it ran away howling and could never be persuaded to go fishing again. In fact, this anecdote antedates the scientific nammg of the New England torpedo. But shocks of a strength even approaching what is suggested by such reports are to be expected only from torpedos of the largest size in rested condition. The voltage recorded recently was 170 to 220 for one that had been kept in a live well. And the most we have felt ourselves from medium-sized torpedos lying on the dock at Woods Hole has been a slight benumbing sensation. The torpedo, like others of its tribe, is a bottom fish. It is a fish eater. The stomach of one taken at Woods Hole contained a summer flounder (Paralichthys dentatus) about 14^ inches long. A 2-pound eel, a 1 -pound flounder, plaice (Pleuro- nectes platessa), red mullet (Mullus surmuletus), a salmon weighing 4 or 5 pounds, and the remains of spotted dogfish (genus Scyliorhinus) have been found in the stomachs of British specimens. The wide distensibility of its jaws allows it to swallow fishes much larger than might be considered possible from the breadth of the mouth when closed. And it is generally believed that it stuns its prey by its electric shocks. Otherwise it is difficult to conceive how so sluggish a fish could capture such active prey. It bears "living" young, but there is no placen- tal connection between embryo and mother. And it seems that the young are born offshore, for the smallest torpedo yet recorded from American inshore waters (from New Jersey) was about 2 feet (610 mm.) long. And we doubt if it succeeds in producing young in the colder waters of our Gulf. General Range. — Both sides of the North Atlan- tic M from southern Nova Scotia (La Have Bank), Bay of Fundy, and Georges Bank to North Carolina in the west;67 and from northern Scotland to the Mediterranean, Azores, Madeira, and tropical West Africa in the east. Occurrence in the Gulf of Maine. — The torpedo is more common south and west from Cape Cod than to the northward and eastward. But it strays past the elbow of the Cape often enough for it to be classed as a regular member of the Gulf of Maine fish fauna. The most northeasterly records for it are of one presumably of this species taken in St. Margarets Bay, Nova Scotia, some 30 years ago; one caught on a long line set for cod u Comparison of American specimens with one from the North Sea revealed no differ- « This torpedo is also reported from the Florida Keys and from Cuba, but on doubtful evidence. 60 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE on La Have Bank in 1890,68 and from Eastport, Maine, at the mouth of the Bay of Fundy. It has also been taken at Williamsport, Maine; off Seguin Island where one was examined in 1880; at the mouth of Casco Bay; at Wood Island near Cape Elizabeth (1, in a trap, in 1894); near Cape Ann; off Plymouth in the southern side of Massa- chusetts Bay; near Provincetown; and on the outer coast of Cape Cod, so it would be no sur- prise to find it anywhere along the shores of the Gulf. It has been caught occasionally on Georges Bank;69 there are records of long standing of torpedos off Nantucket and Marthas Vineyard, and they are caught yearly in Vineyard Sound and in Buzzards Bay. Most of the reports of torpedoes within the Gulf have been based on single specimens. But it has been known for a long time that torpedoes are caught in much larger numbers in some years than in others. Thus they are said to have been unusually common near Provincetown in 1819 and for the next 4 or 5 years, when 60 to 80 were taken there yearly. Again in 1845 about a dozen came ashore or were caught otherwise near Provincetown. Any fluctuation, however, that may have taken place from year to year thereafter seems to have attracted no attention until the summer of 1896, when Dr. W. C. Kendall, of the U. S. Fish Commission collected several along the coast of Maine. The Massachusetts Bay speci- men mentioned above, taken off Plymouth and now in the Harvard Museum of Comparative Zoology, is the only torpedo from the inner part of the Gulf of which we have heard since that time. But it is as likely to be found in the Gulf now as it ever was. Importance. — The torpedo is of no commercial value nowadays, but its liver oil was considered equal to the best sperm for illuminating purposes before the use of kerosene oil was general. There is an old tale that its oil was a good cure for cramps if rubbed on externally, for stomach trouble if taken internally. And when one is landed on the dock at Woods Hole it is an object of interest to the workers at the Biological Labora- tory because of its electric discharges. SKATES. FAMILY RAJIDAE Skates, with their disc -like outlines, thin as a shingle toward their outer edges, and with their rather long tails, are familiar objects along our shores. The outer edges of their pelvic fins are concave (convex in the sting rays) , they have two very small dorsal fins on the rear part of the tail, but no distinct tail fin, and they lack the large tail spine, that is so characteristic of the sting rays. The Gulf of Maine supports four species in abun- dance, while two others have been recorded on rare occasions. The common skates look so much alike that fishermen seldom distinguish between them. For this reason we know very little about the indi- vidual differences in habits among the several species. All live chiefly on the bottom or close to it, spending much of the time partially buried in the mud or sand. They move through the water by undulations of the flexible pectoral fins, steering themselves with the tail. All are decidedly om- nivorous, feeding largely on the larger Crustacea, 88 Reported by G. F. O. Hansen, then second mate and later master of the U. S. Fish Commission sehooner Grampus, who doubtless was acquainted with the torpedo at Woods Hole. M The most recent record is of one .18 inches long, trawled on the southwest part in December 1030. such as shrimps, crabs, lobsters; on mollusks, worms, and to a greater or less extent on fish. All the true skates lay large eggs with blackish or sea-green leathery shells, roughly oblong in outline, with a hollow tendril at each corner by which they become fastened to seaweeds or other obj ects. The empty eggshells, ' ' mermaids purses, " are familiar objects on our beaches among the flotsam along high water mark. While still in the egg the embryo skate develops temporary external gill filaments from the walls of the gill clefts, but these disappear completely before it hatches. Probably all our local skates spawn over a considerable part of the year, with incuba- tion periods of several months up to a year or more.60 To give some idea of their abundance on the offshore banks we may note that the average number of skates (all species together) taken on Georges Bank, per trip of 4 to 7 days, on 25 trips by several trawlers, January to December 1913, was about 800, the largest catch 4,520, the 1,0 Under aquarium conditions the incubation period for the little skate (R. ejinacea) was S to 6 months (p. 69); and it ranged from 4H to about 14?4 months for 6 common European skates; see Clarke, Jour. Marine Biol. Assoc. United Kingdom, vol. 12, No. 4, 1927, p. 587. FISHES OF THE GULF OF MAINE 61 poorest 82. Again, on a trip to the northeastern part of the bank, September 1929, on the otter trawler Kingfisher, 37 hauls yielded from 0 to 105 skates per haul (total 495) and 42 trawl hauls by the Eugene H, fishing from Nantucket Lightship to the south-central part of Georges Bank in late June 1951 caught an average of 146 skates per haul (total, 6,130 skates), which works out at about 9 to 10 skates per acre.61 Probably they are equally abundant on Browns Bank; certainly they are familiar enough there, but statistics are not available of the actual numbers caught. Skates are also plentiful inshore as appears from catches of about 1 skate to 33 fishes of all kinds on long lines, at various localities in the Gulf of Maine/'2 In the Gulf of Maine, skates are only a nuisance for they bite the hook readily and often are caught in great numbers in otter trawls, most of them to be thrown back into the sea, the market demand for them being so small that the total landings reported for New England (Massachusetts and Maine) in 1947 was only 28,200 pounds; and 59,100 pounds for 1948. But some are now being landed in Maine for fish meal.63 They are much more highly vabie( I in northwestern Europe for food with landings for the years just preceding World Win- TT, running around 90 to 100 million pounds. Barn-door skate Raja laevis Mitcliill i SI 7 Bigelow and Schroeder, 1953, p. 217. Garman, 1913, pi. 22, fi»i. 2. as K. stabulifon Description. — The barn-door skate is easily identified by its large size, its very pointed snout , and its smooth skin. The thorns along the mid- line of its back are comparatively small and run only from the hinder part of the disc back along the tail; the tail also has one or two rows of large, sharp spines (smaller on males than on females) along each side, besides the median row. There are small thorns on the snout also, sometimes below as well as above, and along the front edges of the pectoral fins. The male has a patch of erectile hooks on the outer part of each pectoral covering an area measuring 5 by iy4 inches on one side, and 4 '2 by 1 % inches on the other in a speci- fll Three mile hauls with the trawl sweeping a strip about 3.fj feet wide. ,; Examples are: 15 miles off Monhegarj I., Maine. June 24-25, 1913, total fish caught, 5,463; skates 17(1. Twenty mill s east of Cape Cod, Nov. 11, 1913; total fish caught 6,532, skates 202. Jeffreys Ledge, Dec. 11-12, 1913; total fish caught 3,996, skate, 62. •■ Bcattergood, Copela, IS50, p. 189. men 52 inches long; otherwise the pectorals are smooth for the most part. The front angle of the disc is sharper than in our other skates, being more acute than a right angle, but the tip of the snout is blunt. The outer corners of the pectorals are angular and the disc as a whole is diamond or lozenge-shaped. The two dorsal fins are separated by a short interspace, with one or more spines, and the tip of the tail extends farther beyond the second dorsal fin than it does in most skates. The teeth of the female are flat and pavement- like, but those of adult males are provided with sharp slender cusps. Thirty to forty series of teeth have been counted in the upper jaw, 28 to 38 series in the lower jaw. Color. — The barn-door skate like so many sea fish, varies in color. The upper surface is brown (as a rule usually of a distinctly reddish hue), variously marked with small scattered darker spots or blotches of varying size, and often witli pale marblings or waterings; usually there is a large oval spot on the base of each pectoral fin, in line with the outer angle. The lower surface is not as uniformly pale as it is in most skates, its gray or white ground being shaded with darker toward the snout, and speckled on one-third grown specimens and larger, with black or dusky dots or short streaks that mark the mucous pores, a conspicuous feature. Size. — The barn-door skate is our largest, growing to a length of :> feet; it has been said to reach 6 feet though there i< no definite record of one that large. One "I 58 inches was 42 inches wide with a tail 27 inches long, and a female of 50 inches, taken by us, was 33# inches wide, with a 22-inch tail. Barn-door skates weigh about 4 to 6 pounds when 28 to 30 inches long, about 10 to 11 pounds at 36 inches, and about 19 to 21 pounds at 45 to 46 inches. Very small specimens are seldom taken. Habits. — Barn-door skates are bottom lish. They prefer smooth to rocky ground, and we have caught them on very soft mud bottoms as well as on sand and gravel. The fact that the lower surface is more or less pigmented instead of white suggests that it hugs the bottom less closely than other skates, and it is a strong, active swimmer, as anyone will agree, who has landed a large one on a hand-line. Its usual depth range is from close to the tide line, down to about 100 fathoms. It is perhaps more plentiful at 25 62 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 25. — Barn-door skate (Raja laevis). A, dorsal view of female, about 47 inches long, Massachusetts; B, ventral view of one of about 26K inches to show the black markings; C, upper teeth from center of jaw of female 50 inches long; and D, upper teeth from center of jaw of male 52 inches long. B, C, and D from Nantucket Shoals. From Bigelowand Schroeder. Drawings by E. N. Fischer. to 35 fathoms on Georges Bank and on Nantucket Shoals than deeper, judging from average catches of 32 per haul at 26 to 35 fathoms, 13 per haul at 36 to 49 fathoms, and 6 per haul at 50 to 75 fathoms in 42 trawl hauls by the Eugene H, late June 1951, fishing from Nantucket Lightship to the south-central part of Georges Bank. But the Atlantis found it widespread (though not numer- ous), as deep as 100 fathoms both in the open trough of our gulf and in the bowl west of Jeffreys Ledge during experimental trawling, in August 1936; and it has been reported as deep as 235 fathoms off Nantucket. The temperature range of the barn-door skate is wider than that of the little skate (p. 67). They are found in the southern side of the Gulf of St. Lawrence in the icy-cold-bottom water on the banks, also, at lesser depths that warm in summer to 60° F. (16° C.) or more. In the Gulf of Maine, at one locality or depth or another, they are ex- posed to temperatures ranging from perhaps as low as 32° to as high as 64 to 6S° and the upper limit must be considerably higher in the southern part of their range. Garman has pointed out that the spines on the snout of this skate are usually worn smooth, as though used to dig in the mud or sand (very likely it thus obtains the bivalves that form part of its diet). It also feeds on worms, various crus- taceans, particularly on large rock crabs and lob- sters, shrimps, squid, and on fish. Probably it is more destructive to the latter than are any other of our skates thanks to its large size. Woods Hole records list spiny dogfish, alewives, herring, men- haden, butteriish, launce, cunners, tautog, scul- pins, silver hake, hake, and flatfish among its foods. No doubt cod, haddock, and other fish, suffer to some extent from this skate on the off- FISHES OF THE GULF OF MAINE G3 shore fishing grounds, for its European relative is a well-known enemy of the cod, and there is no rea- son to suppose that the barn-door skate is less voracious. It bites readily on almost any bait, and is often caught on hand and long lines as well as in otter trawls, and in weirs along shore. Little is known of the breeding habits. The yellowish or greenish brown egg cases are about 4% to 5% inches (124-132 mm.) long by 2% to 2% (68-72 mm.) inches broad, not counting the horns, and thus much larger than those of any other Gulf of Maine skate. Females containing fully formed egg capsules have been taken in December and January in Nova Scotia waters, evidence that the eggs are laid in whiter. However, it seems that the young are not hatched until late spring or early summer, for we have seen one, taken on Nantucket Shoals in July, so small (about 73« in. long) that it could not have been set free long be- fore its capture. General range. — Atlantic Coast of North America from the Banks of Newfoundland, Gulf of St. Lawrence and outer coast of Nova Scotia and the Nova Scotia Banks to North Carolina.64 It is replaced in European seas by a very close ally, the common skate, Raja batis. Occurrence in the Gulf oj Maine. — This is a com- mon fish in all parts of our Gulf, and any very large skate taken or reported there is almost certain to be a "barn-door." Following the coast around from east to west we find it reported as plentiful off the outer Nova Scotia shore; it is known from St. Mary Bay; is found very generally though not abundantly in the Bay of Fundy and in Passama- quoddy Bay; is reported from Eastport, Casco Bay, and generally along the coast of Maine; is known from various localities in Massachusetts Bay, where we have seen many caught; and its abundance on Georges Bank and on Nantucket shoals is illustrated by an average catch of about 21 per haul (about 14 percent of all the skates caught), in 42 trawl hauls by the Eugene H, fishing from Nantucket Lightship out into the south central part of Georges Bank in late June 1951. In short, it is to be expected anywhere within the limits of the Gulf. Like most other skates, it is often taken in shoal water in our Gulf in summer ; seldom or never in winter. Huntsman tells us that it comes to Passamaquoddy Bay from May to November. We once caught one nearly 5 feet ** Doubtfully reported from Florida. long at Cohasset in Massachusetts Bay in less than a fathom of water in midsummer; indeed, it is often stranded on the beach. This inshore migra- tion, however, does not involve the entire stock, witness its presence in 20 to 60 fathoms on Georges Bank and off Cape Cod throughout the year, and the fact that it is reported by fishermen and has been trawled by vessels of the former Bureau of Fisheries, also by the Atlantis, as deep as 100 fathoms in summer. In the warmer waters off the southern coast of New England it comes in- shore in spring and autumn, descending to some- what deeper water for the summer. Commercial value. — The barn-door skate is of no commercial value except as entering into the small landings of skates mentioned on page 61. Big skate Raja ocellala Mitchill 1815 Spotted skate; Winter skate; Eyed skate Bigelow and Schroeder, 1953, p. 240. Garman, 1913, p. 339, pi. 29, fig. 2, as Raia diaphanes. Description. — This skate looks very much like the little skate, but it is larger and has more numerous teeth. The front angle of the disc is much blunter than a right angle, bulging opposite the eyes, and the tip of the snout is rounded. The teeth are in from 72 to 110 series in cacb jaw instead of 66 series, or fewer as in erinacea, and they are sharper in males than in females. The backs of both sexes are rough with sharp spines on the head, around the eyes, along the anterior margins of the pectorals, over the shoul- ders, and on the sides of the tail. The midline of the back behind the shoulders is almost always free of spines in adults. But we have one speci- men, a female 18 inches long taken near Jeffreys Ledge, November 1, 1927, which bears a row of large spines along the midline of back and tail from the shoulder girdle to the first dorsal fin. Males, like those of other skates, have rows of retractile hooks on the outer parts of the pectorals. The two dorsal fms are close together; the outer corners of the pectorals are bluntly angular; the elaspers of adult males reach about halfway back along the tail, which occupies about half the total length of the fish. Color. — Light brown above with round dark brown spots. As a rule there is a large white eye spot with black center near the rear corner of the pectoral fin, and often two smaller ones 64 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE , 0% US ,'VJ * Figure 26.— Bis skate (Raja ocellata), male, about 36 inches long. From Jordan and Evermann. Drawing by H. L. Todd. close to it. And we have seen two large speci- mens from Georges Bank with several of these eye spots on each side of the disk. There is a translucent or white area on each side of the snout in front of the eyes and the lower surface is white. The eye spots, if present, serve to identify this skate at a glance; sometimes, however, they are lacking, in which case half-grown specimens so closely resemble the little skate that recourse must be had to the number of teeth to tell the one from the other. Size. — This skate does not mature until at least 25 to 26 inches long, and grows to about 3}i feet in length, commonly from 30 to 34 inches. Specimens 32 inches in length are about 20 inches wide. Habits. — Big skates feed on the same diet as little skates do (p. 69). Rock crabs and squid are favorite prey, but they also take annelid worms, amphipods, shrimps, and razor clams, and they eat whatever small fish are readily available, the list at Woods Hole including smaller skates, eels, herring, alewives, bluebacks, menhaden, smelt, launce, chub mackerel, butterfish, dinners, sculpins, silver hake, tomcod, and hake.65 66 From Vinal Edwards' and Linton's notes. It is caught right up to the wharves in the Gulf of St. Lawrence; often comes into very shoal water on sandy beaches, and we once caught an adult male in September in only 2 or 3 feet of water in Nauset Marsh on the outer coast of Cape Cod, but few are found shoaler in our Gulf than 2 to 4 fathoms. They are much more plenti- ful at 25 to 35 fathoms than deeper, on the offshore grounds, as appears from average catches, of 48 per haul at 26 to 35 fathoms, but only 11 per haul at 30 to 49 fathoms, and none at 50 to 75 fathoms, in 42 trawl hauls by the Eugene H, fishing from Nantucket Lightship to the south-central part of Georges Bank in late June 1951, and very few arc caught deeper than about 50 fathoms anywhere. In our Gulf they inhabit about the same range of temperature as the little skate does, i. e., from 68° or so, for those along the Massachusetts coast in summer, down to 34-36° in the coastal belt as a whole in winter, and to near 32° in the Buy of Fundy region, at least in some years. In the southern side of the Gulf of St. Lawrence they are found in the icy bottom water on the banks as well as shoaler, where temperatures rise to 61° (16° C.) or more in summer. Those living the shoalest in the southern part of their range FISHES OF THE GULF OF MAINE 65 must be exposed to temperatures as high, perhaps, as 68° to 70° at the warmest time of the year. Off the Atlantic Coast of Nova Scotia this skate deposits its eggs from summer into autumn, and probably through the same season in the Gulf of Maine for Scattergood 66 reports females with egg capsules in Maine waters in September. And it continues to do so into December and January off southern New England. Its egg cases are larger than those of the little skate, 2% to 2% inches by about l3/4 inches, not counting the horns. The length of the period of incubation is not known. General range. Atlantic Coast of North Amer- ica from northern North Carolina to the southern side of the Cull' of St. Lawrence, and to the southern part of the Newfoundland Banks. Occurrence in ll>< Gulj <>1 Maine. — Tins, the second in size of our skates, occurs commonly all around the Gulf of Maine from Xova Scotia to Cape Cod. There are many locality records for it for the Bay of Fundy as well as from the coasts of Maine and Massachusetts, but so closely does a half or two-thirds grown big skate resemble an adult little skate (p. 68) that it is often impossible to tell to which species published reports refer. It also makes up so considerable a proportion of the skate population on Georges Bank that about 14 percent of the catch of skates made on Georges Bank by one otter trawler in September 1929, and aboul I* percent (1,116) of the skates taken in 42 trawl hauls by the luiot /<< II. late .lime 1951, fish ing from \a tit ticket Lightship to the southwestern part of Georges Bank, were this species. But it has never been reported from the deeper troughs of the Gulf, nor have we taken it there. The name "winter skate" seems appropriate enough for it along the southern coast of Wn England, for it is only during the cold season that big skates come close inshore near Woods Hole. And they are said to be taken in larger numbers in winter than in summer in the Massachusetts Bay region (we cat t verify this). However, this is a misnomer in the cooler waters of the northern part of the Gulf of Maine, for it is common inshore in Passamaquoddy Bay from May to November, and this probably applies to the whole coastline east of Cape Elizabeth to judge from temperature. Big skates are taken on hook and line, in weirs, and in otter trawls, but they are of no commercial value, except as they form a. part of the general » Copeia 1951, No. 2, p. 160. 210941 53 6 landings of skates. And they are only a nuisance to anglers. Brier skate Raja eglanteria Bosc 1802 Bigelow and Schroeder, 1953, p. 165. Garman, 1913, pt. 23. Description. — In the brier skate, as in the thorny skate, the midline of the back and tail is armed with a continuous row of stout thorns from the shoulders to the first dorsal fin near the tip of the tail, usually with 1 or 2 in the gap between the 2 dorsal fins. But the thorns of this row are not much larger than those along the sides of the tail (they are in the thorny skate), and there are at least 16 thorns in the midrow along the tail (not more than 9 to 10 in the thorny skate). There also are groups of large thorns opposite and behind the eves, with 1 to ."> on each shoulder and 1 to 4 rows along either side of the tail. Elsewhere the upper surface of the disc bears only small sharp prickles (hence its name), most numerous on the forward parts of the pectoral-, over head and snout, and along the middle of the back and tail among the larger thorns. Thus it is a much smoother species than the thorny skate, and its snout is more acute, its outline being about a right angle with the mar- gins bulging less opposite the eyes than in any of the blunter-nosed skates. The outer corners of the pectorals are distinctly angular, and the dorsal fins are separated In a short gap. Color. Brownish to grayish above; the pec- torals variously marked with darker spots and blotches and with more elongate bars; this last is a characteristic feature; there is a translucent acb side of the snout ; it is white below. It is most readih recognized bj its color pattern, with short dark bars as well as spot-, which is not shared by any other Gulf of Maine skate. Size. The brier skate ordinarily grOWS to a length of about 2', feet. The largesl on record was about 37 inches long. (It n< ml range. — Off the eastern coast of the United States from Massachusetts Bay to both coasts of Florida. Occurrenct in thi Gulf oj Maine. — This is a southern species, uncommon even as far north as Woods Hole and decidedly rare in the Gulf of Maine. It has been recorded once from Glouces- ter, its most northerly outpost, and also from 66 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 27. — Brier skate (Raja eglanteria), female, about 29 inches long, Woods Hole, Massachusetts. Schroeder. Drawing by E. N. Fischer. From Bigelow and Province town. Two specimens were taken on Nantucket Shoals near Round Shoal buoy by the Halcyon, one in July, the other in September, in 1924. Leopard skate Raja qarmani "Whitley 1939 PiOSETTED SKATE Bigelow and Schroeder, 1953, p. 200. Garman, 1913, pi. 18, fig. 2. Description. — The conspicuous dark rosettes on the upper surface make this skate recognizable at a glance, since no other skate of the western Atlantic is marked in this way. And its tail is longer relatively than that of any other Gulf of Maine skate. The disc is considerably blunter in front than a right angle, with anterior margins bulging rather conspicuously a little anterior to the level of the eyes; the outer corners of the pectorals are very broadly rounded ; the tail measured from the center of the cloaca to the tip is about 1.5 times as long as the body from tip of snout to center of cloaca; and there is a definite gap with one or two thorns between the two dorsal fins.67 There are thorns along the ridge of the snout; a row around the inner and posterior margins of the eyes with a few in the space between the latter; a group on each shoulder; and one row along the midbelt of the back and tail in young specimens, increasing to 2 to 6 irregular rows in large ones. In young specimens the skin of the disc, as a whole, and of the tail, is also rough with small prickles, but, most of these are lost with growth, leaving large specimens mostly naked except for the thorns. The lower surface is smooth. There are 46 to 52 series of teeth in the upper jaw, a few less in the lower, and those of adult males are only a little sharper than those of females. Color. — The upper side is pale buff or brown, •' Carman's illustration is of an abnormal specimen with three dorsal fins. FISHES OF THE GULF OF MAINE 67 Figure 28. — Leopard skate (Raja garmani), female, 16 inches long, offing of Montauk Point, New York. From Bigelow and Schroeder. Drawing by E. N. Fischer. freckled with small spots, darker or lighter, and conspicuously marked with dark rosettes, each consisting of a group of 6 or more dark brown or black spots surrounding a dark central spot. The lower surface is white or pale yellow. Size. — This is one of the smaller skates, males maturing when only about 16 inches long. General range. — Outer part of the continental shelf and upper part of the continental slope from southern Florida to the offing of Nantucket, in depths of 30 to 300 fathoms. Occurrence in the Gulf of Maine. — Our only reason for mentioning this species is that one specimen was trawled by the Albatross III, May 14, 1950, at 52 fathoms southeast of Nantucket Lightship (lat. 40°05' N., long. 69°22' W.). And this is probably close to the eastern boundary of its range, for it has never been reported among the other skates that are trawled in abundance along the seaward slopes of Georges and of the Nova Scotia Banks. But it is one of the most plentiful of skates offshore to the westward, along southern New England. Little skate Raja erinacea Mitchill 1825 Common skate; Summeh skate; Hedgehog skate; Tobacco box Bigelow and Schroeder, 1953, p. 176 Garman, 1913, pi. 20. Description. — The most distinctive characters of grown specimens are their small size, absence of thorns along the midline of the back (except in the young) and blunt nose. The anterior angle of the disc is blunter than a right angle and the tip of the snout is rounded, with the margins bulging opposite the eyes. The teeth are in only about 38 to 66 series. Females have thorns scattered generally over the upper surface; these are especially prominent on head, snout, shoulders, and sides of tail. Ordinarily there are no spines on the midline, back of the shoulder girdle; but we found one fish, 13% inches long, among the many we have observed, with a median row of spines extending from the shoulder girdle to the first dorsal fin near the 68 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 29. — Little skate (Raja erinacea). A, male, 20 inches long, Boston Harbor; B, female, 17H inches long, Mystic Connecticut; C, side view, end of tail of same, about 0.6 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. end of the tail, and this is true of newly hatched specimens in general. Males are less spiny, but the spines on tail, shoulders, and along either side of the back ridge are noticeably strong in both sexes. The two dorsal fins are close together ; the tail is about half the total length. Large ones closely resemble small specimens of the big- skate (R. ocdlata, p. 63) that may chance to lack the ocellar spots with which that species usually is marked. A count of the teeth is then the only sure clue to the identity of the specimen in hand. Color. — Grayish to dark brown above, or clouded light and dark brown, paler at the edges of the pectoral fins; usually with many small round darker spots; white or grayish below. Size.- Ordinarily up to Hi to 20 inches long; the maximum recorded length is 21 inches (53 cm.); they weigh about % to f pound at 10 to 17 inches and anywhere from 1% to 2 pounds at 18 inches. Females mature sexually when 12V17 inches (32-43 cm.) long, males at about 14 to \7% inches.68 Habits. — It is common knowledge that this skate, like others, is most abundant on sandy or pebbly bottom; but they are likewise found on mud and we have seen them lying on ledges at times. The usual depth range is from close to tide line down to 7o fathoms or so. Many even follow uj) the shelving bottoms of our beaches until they •'Information supplied by Dr. Daniel Merriman, Dr. > . 11 Olsen, the m, g ii Wheatland and I.. B. Calhoun, who have made ;i detailed tuds of the little skate in southern N'e« England waters FISHES OF THE GULF OF MAINE 69 strand. And the bulk of the population hold to depths of less than 40 to 50 fathoms, as appears from average catches, per haul, of 100 at 26 to 35 fathoms, and 95 at 36 to 49 fathoms, but only 12 at 50 to 75 fathoms, in 42 hauls by the Eugene H, fishing from Nantucket Lightship to the southeentral part of Georges Bank, in late Juno 1951 . Fifty fathoms (off the Bay of Fundy I is, in fact, the deepest that positively identified specimens are known, in the inner parts of our Gulf; 80 fathoms off southern New England." The little skate tolerates a wide range of temperature, being found in water as warm as 68-70° in summer, while they are exposed to temperatures close to 32° in the Bay of Fundy in some winters, unless they move out, and deeper there than seems likely. In the southern side of the Gulf of St. Lawrence, writes Hunts- man,70 they are found in the intermediate zone between the icy cold waters of the banks and the surface stratum, which last warms to 61° (16° C.) or higher in summer. They have never been reported, to our knowl- edge, where the water is appreciably brackish. Hermit and other crabs, shrimps, worms. amphipods, ascidians ("sea squirts"), bivalve mollusks, squid, small fishes, and even such tm\ objects as copepods have been found in their stomachs. Probably crabs loom largest in their diet, for more than '-".) percent of the skates opened by Field at Woods Hole, contained them; 15 percent had bottom-dwelling shrimps {Crago and 6 percent had eaten squid. In Long Island Sound, however, amphipods (Leptoi are the dominant item in their diet, forming from one- third to one-half of the stomach contents at all seasons of the year.71 Launce, alewives. her- ring, dinners, silversides, tomcod, silver hake, have all been found in their stomachs, and the'. bite a baited hook readily, affording amusement to vacationists. The spawning habits of the little skate have aol been followed in the Gulf of Maine. Studies. at the Bingham < >ceanographic Laboratory, how ever, suggest that they ordinarily deposit their eggs in water not deeper than 15 fathoms and on a sandy bottom. It appears from anatomical examination of the sexual organs of the mature females that ii Seventeen that wc savi trawled on tl Al atrosi III. Mas 1950. :o Trar Roj i ■■' . er. 3, to!. 12, sec., i, 1918, p 63 '■Information from l>i Daniel Merriman, Dr. V. B. < Isen, nid the land and L. II Calhoun. copulation may take place at any time throughout the year, and frequently. Observations, too, on skates kept in aquaria have shown that the eggs are laid in pairs at intervals of from five days to several weeks; also that they are usually buried in sand, at least partially.72 The eggs have been taken off Southern New England, in fish traps and dredges in a few fathoms of water in abun- dance from July through September. Examination of large numbers of females has shown that eggs are laid there throughout the year. And there is no reason to doubt that this is the case to the north and east of Cape Cod as well. Aquarium experiments have also shown that eggs laid in the period, May-July, hatched between the end of November and the beginning of January, i. e., after 5 to (i months. But the incubation period is likely to be somewhat longer for spring-summer laid eggs in nature because of somewhat lower temperatures; and considerably longer for eggs laid in autumn and early winter. The eggs measure about 1 '-. to 1\ inches by aboul 2 . to 2% inches, nol counting the horns, and the great majority of the empty skate eggs that are washed up on the beaches of our Gulf belong to this species. The young skate, which emerges through a transverse opening at the edge of the egg case al I he end that has the longer pair of horns, is aboul 3 i to ! inches long at hatching; its abdomen is si ill swollen with yolk, and its tail terminates in a whiplash-like extension that disappears within a few days. Huntsman's ob- servations suggest that young hatched near the head of the Bay of Funds descend to deeper water the first winter, and this probably applies in the < rUlf of Maine as B H hole. It appears from information of various sorts thai a little skate S inches (I'D em.) long may be 1 to 1', years old; one of I 1% to 12 inches (30 cm.) 2 lo :; years; one of 15ni to Hi inches (40 cm.) 3 io 4 years; one of 19:'i to 20 inches 6 to 8 years old. And the mortality rate appears to be very high after five years, for very few of those taken are longer than about 18 to 19 inches." General range. Atlantic coast of America; southern side of the Gulf of St. Lawrence and n This summary of the ' i I nsive information supplied by Dr. Daniel Merriman, Dr. Y. II. Olsen, and the Misses S. B. Wheatland and h. 11. Calh >u :i, from Dr. an. Dr. Y. H. Olsrn, ani m. - i land nd L. II Call 70 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE northern Nova Scotia to Virginia, in coastal waters and on the shoaler of the offshore banks. Occurrence in the Gulf oj Maine.— This, the smallest of our skates, is the commonest and the most familiar from its habit of coming up into very shoal water in summer and of stranding on the beaches, where dried skate carcasses are often to be seen. It occurs all along the coast in the southern side of the Gulf of St. Lawrence and along outer Nova Scotia; is very abundant both on the New Brunswick and on the Nova Scotia sides of the Bay of Fundy, and is to be caught every- where and anywhere along the coasts of Maine and of Massachusetts; far more commonly, indeed, than one might suspect from the few definite records that have found their way into scientific literature. An average catch of about 88 per haul (about 60 percent of all the skates taken) in 42 trawl hauls by the Eugene H, in late June 1951, fishing eastward from Nantucket Lightship suggests that this is the most plentiful skate on the south- western part of Georges Bank and on the Nan- tucket grounds. But it seems to be far less numerous on the northeastern part of the Bank, if it is present there at all; at least we failed to find a single one, among 495 skates of other kinds caught there in 37 hauls by the otter trawler Kingfisher in September 1929. And we have never found it (nor has it been reported) in the deeper basins and troughs of our Gulf, probably because it is restricted in general to depths less than 40 to 50 fathoms (p. 69). In our Gulf many of the little skates appear to carry out an irregular migration into shoal water in April and May, where they remain throughout the summer, autumn, and early winter, to retire again to somewhat deeper, hence, warmer water in December or January. Its migration schedule appears to be more complex in Long Island Sound waters where summer temperatures are higher; i. e., inshore in spring, offshore in mid- or late summer, inshore again in late autumn and offshore again in midwinter.74 Doubtless little skates breed throughout the shoaler parts of the Gulf, and on the offshore banks. They are of no commercial importance in our Gulf except as they form a part of the landings of trash fish. " Information from Dr. Daniel Merriman, Dr. Y. H. Olsen, the Misses S. B. Wheatland and L. H. Calhoun of the Bingham Oceanographic Lab- oratory. Smooth-tailed or prickly skate Raja senta Garman 1885 Bigelow and Schroeder, 1953, p. 264. Garman, 1913, pi. 25, fig. 1. Description.— By the time this skate has grown to one-fourth its adult size it is made recognizable by the fact that the middorsal line of thorns runs back only to about the middle of the tail, where the thorns so dwindle in size that they are not dis- tinguishable from the tiny prickles with which the tail is clothed, generally. Newly hatched speci- mens in which this character is not yet established are separable from all other Gulf of Maine skates by the color pattern of the tail, which has two pale crossbars, each outlined in front and behind by a dark band or blotch. There is a single row of 16 or more medium- sized to large thorns along the midline of the back, spaced irregularly, and usually about 20 to 30 along the anterior one-half or so of the tail; about 10 to 13 around the inner ridge of each eye; and 3 to 5 on each shoulder. Immature specimens of both sexes are also closely and uniformly rough- ened with small prickles over the disc as a whole, on the pelvics and on the upper side of the tail. But irregular bare areas develop on the shoulders and around the outer parts of the pelvics of females as they approach maturity while mature males lose the pricldes from the central part of the disc as a whole, but develop a few thorns on the mid- ridge of the snout besides larger thorns over a roughly triangular area on either side of the head abreast of the eyes and farther forward. They also develop two rows of the usual curved sexual spines on either side on the pectorals, about 13 to 14 per row. The lower surface of the disc is smooth, except that a few pricldes develop, with growth, along the margins near the snout The lower surface of the tail as a whole is prickly on females and on immature males, but tends to become smooth on males by the time they mature sexually. The anterior angle of the snout is a little more obtuse than a right angle (about 110°); the tip of the snout is sharper than in either the big skate, the little skate, or the thorny skate. There are 38 to 40 series of teeth in the upper jaw, 36 to 38 series in the lower jaw; those of females are low, with only faintly indicated points, but those of mature males are longer, sharper, recurved, and FISHES OF THE GULF OF MAINE 71 Figure 30. — Smooth-tailed or prickly skate (Raja senta), male, about 20J4 inches long, Emerald Bank, Nova Scotia. From Bigelow and Schroeder. Drawing by E. X. Fiacher. spaced more loosely. There is no free space between the two dorsal fins. The disc is a little broader than long (1.2 to 1.3 times); the tail occupies about one-half of the total length. Color. — The upper side, including the tail, is pale brown, with many obscure darker spots. Newly hatched specimens are also marked on the tail with two pale cross bars, each outlined by a darker cross bar or blotch in front of it and one behind, but these bars disappear with growth. The lower surface is white, either plain or with a few dusky blotches. Sometimes the rear part of the tail is uniformly dark below. Size. — A male about 20 inches (515 mm.) long that we have seen seems to be sexually mature. The largest specimen of which we have record was 24 inches long. Habits. — This skate appears equally at home on the soft mud and clay bottoms of the deeper basins of the Gulf and on the sand, broken shells, gravel and pebbles of the offshore fishing banks. Nothing is known of its diet. Egg cases, appar- ently of this species, have been trawled in deep water (82-164 fathoms), in the estuary of the St. Lawrence Kiver in July and August ; probably thev are laid in summer in the Gulf of Maine, as well. General range. — Atlantic shelf of North America from the offing of Charleston, S. C, to the Nova Scotia Banks and Gulf of St. Lawrence, a few reaching the southern part of the Newfoundland Banks; mostly in depths greater than about 40 to 50 fathoms. The deepest record for it is 478 fathoms off South Carolina. Occurrence in the Gulf of Maine. — This skate, once considered a rare species, is now known to occur generally throughout the western side of the Gulf wherever the water is more than 45 to 50 fathoms deep, 25 fathoms being our shoalest record for it and on the offshore Banks as well. We have trawled it on several occasions in the deep trough west of Jeffreys Ledge; in deep water (80-100 fathoms) near Cashes Ledge; also in the basin east and southeast of Cape Cod. And, being known from the southeastern slope of Browns Bank, it is to be expected generally in the eastern side of the Gulf, as well as in the western, at the proper depth. It is widespread on Georges 72 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Bank also, but is far less plentiful there than other skates, to judge from the fact that trawl hauls in September 1929 brought in only 37 of them, and that we counted only 8, from 42 hauls on the Eugene H; in June 1951, fishing from Nantucket Lightship to the south central part of Georges. We have trawled it at 50 to 250 fathoms off southern New England. To the eastward and northward, it is recorded on La Have and Emerald Banks at 50 to 100 fathoms, and in the Gulf of St. Lawrence at 82 to 178 fathoms. Thorny skate Raja radiata Donovan 1807 Starry skate75 Bigelow and Schroeder, 1953, p. 255." Garman, 1913, pi. 21, fig. 2." Description. — The thorny skate can be identified at a glance among skates of the Gulf of Maine by the fact that the row of thorns with which the midline of back and tail is armed are not only large and conspicuous, but do not number more than 19 at most from the nape back along the tail. There are also 2 or 3 large thorns on each shoulder; and one in front of each eye and one behind it ; one close to the inner end of each spiracle; and other smaller thorns scattered on snout, pectoral fins, and tail. The bases of the thorns on the pectorals are star-shaped, a very distinctive character; those of the still larger thorns along the midline of the back are oval. Adult males have 2 rows of hooked, erectile thorns near the outer corners of the pectorals. The anterior angle of the disc is considerably more obtuse than a right angle (1 10-140°), and the tip of the snout is blunt with the margins bulging somewhat a little in front of the level of the eyes; the outer corners of the pectorals are less broadly rounded than in either the little skate or the big skate; and the two dorsal fins may either be joined at the base or be separated by a short space. There are 36 to 46 series of teeth in each jaw, those of females and of young males with low cusps that are worn nearly smooth along the 78 When the tirst edition of this book appeared, it was an open question whether the thorny skate of American waters (named /'. scabrata by Garman 1913) was identical with the thorny skate of northern Europe (R. radiata Donovan, 1807). Our subsequent comparison of American specimens with others from Greenland and Norway has convinced us that they all belong to the one species, which must be called by the older of the two scientific names. '• FiKiire lot Gam 's plate 2! is nol ofs thorny skate, as it is named in the accompanying caption, bul isol l small pecimen of the big katethat we lia\ e examined. older rows; those of mature males a little sharper and spaced a little more widely. Color. — Brown above, either uniform or slightly clouded, or spotted with darker, small specimens more conspicuously so than larger. Sometimes there is a white spot beside each eye, one on either side abreast of the nape, and another on each side on the rear part of the disc. The lower side is white, sometimes with irregular sooty or brownish blotches. Garman mentions a partial albino, white above with a few reddish brown and brown spots. Size. — The thorny skate is about 4 inches (100 mm.) long from snout to first dorsal fin at hatch- ing. The largest specimens so far recorded from American waters have been about 40 inches for the Nova Scotia Banks, 35% inches for Georges Bank, and about 31 inches for Massachusetts Bay. But some males may mature when only 21 to 22 inches long. One 32 inches long is about 23 inches wide. Habits. — The thorny skate is a cool water fish, at home in temperatures from about 50° or so down nearly to the freezing point of salt water. It is also restricted in general to depths greater than about 10 fathoms, even in the northernmost part of its range. In the Gulf of St. Lawrence it lives indifferently on the ice cold banks and in the warmer water on the bottom of the deep Lauren- tian Channel. Average catches of 1 per haul at 26 to 35 fathoms, 22 per haul at 36 to 49 fathoms, and 5 per haul at 50 to 75 fathoms, in 42 trawl hauls, by the Eugene H fishing from Nantucket Lightship, the central part of Georges Bank, June 1951, suggest a rather definite preference for the intermediate depth zone, perhaps because of the food supply. But thorny skates have been taken at many stations, also, down to 336 fathoms off the American coast, and as deep as 459 fathoms near Spitzbergen. The stomachs of thorny skates caught on Georges Bank contained shrimps, spider crabs, anemones, hydroids, and fish digested past identification. The egg cases vary considerably in size, prob- ably depending on the size of the parent fish. One from a fish 32 inches long, taken on Georges Bank, measured 3 by 2% inches exclusive of the horns. Others that have been measured from the Nova Scotia Banks ranged from 3 to 3% inches in length. They are flat on one side, strongly convex FISHES OF THE GULF OF MAINE 73 Figure 31. Thorny skate {Raja radiaia), female, about -i inches long. After Garman. on the other, and are rough with narrow cro ridges. A mass of delicate fibrils, matted to- gether, extends along each of the longer sides and partly over the surfaces also. And each horn ends in a slender fibril. General range.- The thorny skate is known on both sides of the northern Atlantic. In the east its range extends from the White Sea and Barents Sea to the North Sea, Dutch coast, and western part of the Baltic;77 in the west from West < Green- land, Hudson Bay, Atlantic coast of Labrador, cast and south coasts of Newfoundland, Grand Banks, Gulf of St. Lawrence and outer coast of Nova Scotia with the off-lying fishing grounds, to the Gulf of Maine, and thence westward and south- ward along the edge of the continental shelf to the :: Doubtfullj reported from Belgium and the Bay of Biscay. oiling of New York; and as a stray to the offing of Charleston, S. C.;s Occurrence in the Gulf oj Maim. -The thorny skate is not often seen close inshore along our coast, g restricted in general to moderately deep water (p. 72). Bui it is i.ow known to be generally dis- tributed in the deeper wafers of the Gulf. Thus il is frequently taken on the New Brunswick side of the Bay of Fundy in depths of 10 fathoms or deeper, in 20 to 30 fathoms in St. Mary Bay on the Nova Scotia side. It has been recorded from Casco Bay; from Ipswich Bay, off Gloucester, Salem and Nahant, and off Provincetown ; and we have taken it ourselves in numerous places in the Gulf at 14 fathoms and deeper, including the " One taken in lat.33 to' X.. long. 7i .1 fathoms, by the Albalrost 111 Is in the Museum of Compel 74 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE vicinity of Mount Desert; PlattsBank; and in the bottoms of the deep troughs. It has also been trawled at many stations on Georges Bank, like- wise along the upper part of the continental slope off southern New England, down to 336 fathoms. There is nothing in the available record to sug- gest that it carries out any regular migrations, whether in or offshore, or along the coast. And it is more catholic in respect to its choice of bottom than some other skates, for while it is most plenti- ful on the good fishing grounds of sand, gravel, and broken shells, we have taken it at many sta- tions in the Gulf on soft mud. And it is one of the most plentiful of Gulf of Maine skates at appro- priate depths. Thus 325 were caught in 37 trawl hauls on the northeastern part of Georges Bank on one trip in 1929; again, in June 1951, we counted 432, from 42 trawl hauls (7 percent of the total catch of skates), on the Eugene H fishing from Nantucket Lightship to the south central part of Georges. We once caught 12 in the western side of the Gulf in a beam trawl oidy 8 feet across the mouth in 30 minutes; and we have taken 1 to 100 of them in 26 hauls with larger trawls, between Mount Desert Island and Massachusetts Bay. Females containing eggs about ready to be laid, and deposited eggs in various stages of incubation, have been taken in Nova Scotian waters or in the Gidf of Maine, in April, June, July, and September, and they are to be expected there in winter as well, having been reported in January and February off Norway, and from February to June in Scottish waters. THE WHIP-TAILED STING RAYS. FAMILY DASYATIDAE The whip-tailed sting rays, like the skates, are disc-like in form, very thin toward the outer edges, with the anterior parts of the pectoral fins fused with the sides of the head, and with the eyes and spiracles on the upper surface. Their pelvic fins, however, have convex outer edges, not concave as are those of the skates. They have no dorsal fin. Their tails are long and whiplash-like to- ward the tip and armed, in most of them with one to several sawedged, venomous spines on the upper side. Their teeth are small and in many series, closely crowded in bands along the jaws. The upper surface of disc and tail is smooth in some of them, variously roughened with tubercles, thorns or prickles in others. They do not lay eggs as the skates do, but bear "living" young (p. 57). And the young resemble their parents closely when born. Four species are known along the Middle and South Atlantic States, but only one of them reaches the Gulf of Maine, and then only as a stray. Shoidd any long-tailed sting ray be picked up within the limits of the Gulf that does not fit the following description, its captor is referred to Bigelow and Schroeder, 1953, 79 for its identification. " Fishes Western North Atlantic, Pt. 2, Mem. 1, Sears Foundation, 1953. Sting ray Dasyatis centroura (Mitchill) 181580 Stingaree; Clam cracker Bigelow and Schroeder, 1953, p. 352. Garman, 1913, pi. 33, figs. 1, 2, as Dasybatus marinus. Description. — The most distinctive features of this sting ray, among other Gulf of Maine fishes, are its very long, whiplash-like tad without dorsal fins, and the long, sawedged spine or spines with which the upper side of its tad is armed. The disc is rhomboid, about \)i to l){ times as broad as it is long; the anterior angle is much blunter than a right angle (130-140°); and the tip of the snout projects very little if at all. The anterior margins of the disc are nearly straight, the posterior mar- gins are only slightly convex, and the posterior corners are abruptly rounded or even angular. The tad, measured from the center of the cloaca, is about 2% times as long as the body from cloaca to snout. The lower side of the tail has a narrow fold of skin extending rearward from below the origin of the tad spine for a distance about as 80 This ray was mentioned as Dasybatus marinus and as D. hastatns in the first edition of this book. But the specimens in question all belong to one species, the correct scientific name for which is Dasyatis centroura, proposed by Mitchill in 1S15, as Raja centroura. FISHES OF THE GULF OF MAINE 75 Figure 32. — Sting ray (Dasyatis cenlroura), male, about 55}i inches wide, Woods Hole, Massachusetts, and tubercle from tail, about 0.7 times natural size. From Bigelow and Schroeder. Drawings by E. N. Fischer. long as from its own origin to the cloaca; the upper side of the tail is rounded, except for a low ridge close behind the spine (or spines). The spear-pointed tail spines, of which there are 1, 2, or sometimes 3, are situated well back on the tail. One spine that we examined had about 40 saw teeth on each edge. Young ones are smooth skinned (except for the tad spines) . Larger ones develop 1 to 3 irregular rows of conical tubercles along the midline of the disc, with others on the shoulders as well as on the outer posterior part of the disc, and the tail be- comes very rough finally, with conspicuous thorns along its whole length on its upper side, and rearward from abreast of the tail spines on its lower side. The lower side of the disc is smooth. Large specimens are easily distinguishable from other sting rays of our Atlantic coast by their very thorny tails and by the large tubercles on the outer parts of their discs. Small ones on which the thorns and tubercles have not yet developed, are recognizable by the shape of the disc, com- bined with the presence of a skin fold on the lower side of the tad but none on the upper side. Size. — This appears to be the largest sting ray of the western North Atlantic. The greatest 76 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE measured width definitely recorded for it is 5 feet, the greatest measured length 10 feet 3 inches. But some certainly grow considerably larger, for a New Jersey specimen has been reported as nearly 7 feet across; the corresponding length would be 13 to 14 feet, if the tail were intact. Color. — Fresh caught specimens seen by us at Woods Hole, have been dark brownish above with the tail black from the spine rearward; white below. General range. — Coastal waters of the western Atlantic, from the latitude of Cape Hatteras to Cape Cod; most common from Delaware Bay to the Woods Hole region. Occurrence in the Gulf of Maine. — The only claim of this sting ray to mention here is that one was reported from Chatham on the outer shore of Cape Cod many years ago, and that it is said to have been seen on the shoaler parts of Georges Bank. It has no real status as a Gulf of Maine fish, appearing there only as a summer straggler from the south, tbough it is so common near Woods Hole that the fish traps there catch some 400 to 500 of them in ordinary summers.81 Beware of handling any skate-like fish with a long, whip-like tail, lest it prove to be a sting ray. The tail spine (brought into action as the tail is lashed to and fro) is a dangerous weapon; and the wounds made by it cause excruciating pain. THE COW-NOSED RAYS. FAMILY RHINOPTERIDAE The cow-nosed rays, like the whip-tailed rays, have a very long tail armed with one or more poisonous sawedged spines; a very flat broad disc; and pelvic fins with convex outer margins. But their pectoral fins are interrupted on each side of the head, so that the forward portions form a separate two-lobed fin extending forward from the lower side in front of the mouth and nostrils; their crowns are high-domed; their eyes and spiracles are on the sides of the head instead of on its upper surface; and they have a small dorsal fin on the upper side of the tail in front of the tail spines. Their teeth have the form of large, flat grinding plates, fitting close together in mosaic arrange- ment; and there are only 7 to 9 series of them in each jaw. Cow-nosed ray Rhinoptera bonasus (Mitchill) 1815 Bigelow and Schroeder, 1953, p. 469. Garman, 1913, pi. 37, as Rhinoptera quadnloba. Description. — The cow-nosed rays with all their close relatives82 have such characteristic out- lines, the shape of their heads is so peculiar with the eyes and spiracles on the sides, and their large, flat, plate-like teeth are so different from those of any other Gulf of Maine skates or rays that they are not apt to be mistaken for anything else. 61 This estimate is based on our own observations near Woods Hole. '•"The eagle lays, family Myliobatidae, and butterfly rays, family Gymmiridae, are close allies of the co id rays; none of them In is yet been encountered in our Quit The species in question is characterized among its confreres by the indented contour of the front of its head, and by the conspicuously bilobed outline of the short so-called "subrostral" fin that pro- jects forward from the lower side of the latter. The outer corners of the pectorals are pointed, and their posterior margins distinctly concave. The pelvic fins are small, reaching but a short distance hack of the posterior corners of the pectorals. The dorsal fin is rounded above, originating about opposite the rear ends of the bases of the pelvics. The tail measured from the center of the cloaca is about twice as long as the body from cloaca to front of head on adults if not damaged, nearly 3 times on small specimens. The tail spines (1 or 2) are close behind the rear limits of the pelvic fins, and thus much further forward on the tail than those of the sting rays (p. 74). There usually are 7 series of teeth in each jaw, with up to 11 to 13 rows exposed, and in function simultaneously. Size. — The cow-nosed ray has been said to grow to a breadth of 7 feet. But the largest specimen the width of which has either been actually measured or can be calculated from some other dimension, was only about 38 inches wide.83 Color. — Brownish above, white or yellowish white below. Some of them are marked both above and below with many narrow faint dark lines radiating out from the center of the disc. General range. — Western Atlantic coast from middle Brazil to southern New England. " Calculated from the dimensions <>f the head of one from Bio de Janeiro. FISHES OF THE GULF OF MAINE 77 Figure 33. — Cow-nosed ray (Rhinoplera bonasus), aboul 22 inches wide, Newport, Rhode Island. From Bigelow and Schroeder. Drawing by R. V Fischer. Occurrence in the Gulf of Maine. The cow-nosed ray has even le^s claim than the sting ray to be called a Gulf <>f Maine fish, for while schools of them appear occasionally near Woods Hole where ! 15 of them were taken m the lisli traps in one day on one occasion, and while it is recorded from Nantucket, il li.is never been seen, actually, east or in >ii h of the elbow of Cape ( !od. DEVIL RAYS. FAMILY MOBULIDAE The devil rays, like the sting rays (p. 74) and cow-nosed rays (p. 76) have the pectoral (ins interrupted along the sides of the head close behind the eyes. But they differ very noticeably from the others mentioned above in the shape of the anterior pails of the pectorals, for these are in the form of two separate narrow ear-like fins, set vertically and curving forward from the front of the head. They are further unique among skates and rays in the fact that I hey feed on small pelagic animals, which they sift, by a complex sieve-like modification of their gill arches, out of the water that is gulped in by the mouth and passed out via the "ill clefts. Some of them are the largest of the ravs and among the Largest of fishes. Being tropical-subtropical in nature they have no real place m the lisli fauna of our Gulf, hut Manta, the largest of them all, has been known to reach Georges Hank as a stray from warmer la til udes. Devil ray Manta biro&tris (Donndorff) 1798 Bigelow and Schroeder, 1953, p. 502. Description. — The so-called cephalic fins of the devil ray, pointing forward, give it so distinctive an appearance that it could not be confused with any other lisli, except for some other member of its own family. And it is marked off from all others of these that are known in the Atlantic by the 78 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 34. — Devil ray {Mania birostris), juvenile male, 11 feet 5}i inches wide, Bimini, Bahamas. Schroeder. Drawing by E. N. Fischer. From Bigelow and position of its mouth, which extends across the front of its head instead of being on the under side. Its cephalic fins are about one-half as broad at the base as they are long, with thin lower edges and thick fleshy upper edges and rounded tips, and each arises nearly vertical from the side of the head. When the owner is swimming they point directly forward, but when the ray is feeding, they can be curved inward, one toward the other until their tips nearly meet in front of the mouth. The disc (not counting the cephalic fins) is a little more than twice as broad as long, with tapering outer corners. The tail measured from the cloaca is at least as long as the body from cloaca to front of head and perhaps longer still if not damaged. And it bears a small rounded dorsal fin on its base. Some specimens have been described as having one or two small tail spines close behind the dorsal fins. However, those that we have seen have had none, but a rounded knob in its place, supported by a mass of bony tissue with a minute pointed spur on the upper side that can be felt but does not break the skin. The skin of disc, pelvic fin, and tail is roughened with small tubercles, below as well as above. The mouth is very wide, extending across a little more than one-half the whole breadth of the front of the head. And the teeth, the lower jaw only, are minute and very numerous; we counted about 270 series in about 12 to 18 rows or a total of about 4,500 in one specimen about 11}£ feet wide. The gill openings are noticeably long. Color. — The upper side varies from reddish or olive brown to bluish slate colored or black, either plain or with various white markings. The lower side is white toward the center of the disc but gray around the margins, and there may be various dark blotches in the region of the gills and on the abdomen. The rear part of the tail is gray. Size. — This giant ray matures when about 14 to 15 feet wide. They commonly grow to 18 feet or so, and there are recent records of measured specimens 19 feet 8 inches, 21 feet 2 inches, and 22 feet wide. One 14 feet wide weighed 1,686 pounds, one from the Galapagos Islands, 18 feet wide, 2,310 pounds; and one of 20 feet taken long ago off Venezuela weighed 3,502 pounds. General range. — Manta is known in the Atlantic from middle Brazil to the Carolinas and as a rare FISHES OF THE GULF OF MAINE 79 stray to southern New England and Georges Bank; from Bermuda; from Madeira; and from tropical West Africa. Mantas are also widespread in the tropical-subtropical belt of the Pacific and Indian Oceans, but it is not yet known whether they are identical with the Atlantic species or not. Occurrence in the Gulf oj Maine. — The only reason for mentioning this giant ray here is that a pair, judged to be 18 to 19 feet wide, were encountered on the southeast part of Georges Bank late in August 1949, by Capt. Henry W. Klimm, while out after swordfish, and so close at hand that their cephalic fins and purplish color were noted. The nearest record to the westward and southward is of one 19 feet wide, weighing 1,686 pounds, harpooned by a sword fisherman a few miles off Block Island and landed there in August 1921.84 Chimaeroids. Subclass Holocephali The chimaeroids, being cartilaginous fishes, are allied to the sharks, skates and rays, but are separated from them by many important ana- tomic characters. Most obvious of these externally are that they have no spiracle; that they have only one external gill opening on either side; that their tails are symmetrical; and that their gill filaments are free at the tips like those of bony fishes. The chimaeroids remotely suggest the grenadiers in general body form (p. 243), but arc easily separable from them at a glance; first of all by the softness of their bodies and by their naked skins, also by the location of the pelvic fins which are set far back under or behind the tips of the pectorals, and by the large size of the pectoral fins, to list only the most obvious differences. There is no danger of confusing them with any other Gulf of Maine fishes, so curious is their appearance. They lay eggs that are astonishingly large for the size of the parent fish, and enclosed in brown horny capsules which are elliptical, spindle-shaped or tadpole-shaped in different species. But fertilization is internal. The Chimaeras. Order Chimaerae FAMILY CHIMAERIDAE Chimaera Hydrolagus affinis (Brito Capello) 1868 Bigelow and Schroeder, 1953, p. 539 Description. — This species of chimaeroid, the o-dy one known from within the geographic limits of the Gulf, is deepest (one-sixth to one-seventh us deep as long) just behind the gills, tapers gradually backward to a weak slender tail, and is very soft-bodied. The head is short, its dorsal profile oblique, the snout conical with a blunt tip. The forehead of the male bears a curious cartilag- inous hook, armed with recurved prickles on its lower surface, which probablj- serves to clasp the female. The mouth, on the lower side of the head, is small, with thick fleshy lips; the upper jaw is armed with 4 flat plates in place of teeth, « Reported by Oudgcr (Science, N. Ser., vol. 55, 1922, p. 339). There are photographs of this specimen in the American Museum of Natural History in New York. Figure 35. — Chimaera (Hydrolagus affinis), female, about 31 Yt inches long, Banquereau Bank. From Bigelow and Schroeder. Drawing by E. N. Fischer. 80 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE the marginal pair set edgewise, the lower jaw with a pair of marginal plates set edgewise. The gill openings are vertical, set very low down on the sides of the neck, and each is covered with a flap of skin, paralleling the gill cover of bony fishes. There are two distinct dorsal fins. The first of these originates about over the gill openings, is triangular, about as high as long, and supported at its anterior margin by a stout spine that is free along the terminal part, with the rear surface of the free part double saw-edged. The second dorsal is separated from the first by a space vari- able in length, and is less than one-third as high as the first, with straight margin. The small caudal fin, marked off from the second dorsal by a deep notch, is lanceolate in shape, ending as a short, whiplike filament; and it extends a short distance forward along the ventral surface of the trunk, there being no separate anal fin. The pelvics and pectorals both have pointed tips, the latter being much the larger and reaching back nearly to the point of origin of the pelvics. The male has a trifid copulatory organ arising from the base of each pelvic fin on the inner side, and also a supplementary bladelike clasping organ close in front of each pelvic fin, its margin armed with 4 or 5 hooks, and lying in a pocket from which it can be protruded. The skin is smooth; the lateral- line system is well developed and ramifies over the head in several branches. This species is a close ally of the well known chimaera of north European seas (Chimaera monstrosa), but is distinguishable from it by the fact that it has no separate anal fin; that there is a considerable free space between its two dorsal fins; that the outline of the second dorsal fin is straight; that its caudal filament is much shorter; and that its pectorals hardly reach back to the pelvics. Color. — Lead color, tan-brown or dark sepia below as well as above, except paler on the throat and grayish on the snout. The margin of the first dorsal, the rear and inner margins of the pelvics, and the rear margins of the pectorals are dark. Size. — The largest specimen yet reported, taken 85 miles off Cape Sable, Nova Scotia, at a depth of between 400 and 500 fathoms, was 49 inches long and weighed 17K pounds dressed. General range. — Not uncommon on the conti- nental slope of North America from the latitude of Cape Cod northeastward, along the Nova Scotia Banks, to the Grand Banks, in 160 fathoms to more than 1,200 fathoms; also in the eastern side of the Atlantic off the coast of Portugal. Occurrence in flu Gulf of Maine. — Our only reason for mentioning this chimaera is that it is (or was) so plentiful along the offshore slopes of the Banks off the eastern part of the Gulf and off Nova Scotia that many were brought in for a few years subsequent to 1875, when fishermen long lining for halibut extended their operations down to 300 fathoms or so. Only one seems to have been reported during the past 25 years, caught off Browns Bank, 85 miles southwest of Cape Sable, between 400 and 500 fathoms on October 15, 1930.85 But perhaps it would be found no less plentiful now than of old, if sought at the proper depth. The shoalest capture of which we found record was at 1G0 fathoms. Nothing is known of its way of life nor have its egg cases been seen. THE BONY FISHES. CLASS OSTEICHTHYES THE STURGEONS. FAMILY ACIPENSERIDAE The sturgeons, like the sharks, have an uneven ("heteroccrcal") tail with the vertebral column extending out along the upper lobe. But there is no danger of mistaking a sturgeon for a shark for it has only one gill opening on each side, while the gills are enclosed by bony gill covers. And the combination of gills of this kind with sharklike t:ii] and with the fact that the head is covered by bony plates united by sutures, sets the sturgeons off from all other Gulf of Maine members of their own class. Two species of sturgeons are known from the Gulf, one of which once was rather common there; the other is extremely scarce everywhere. I' Reported by Firth, Dull. Boston Soc. Nat. Hist., 61, 1931, p. 9. It was 19 Inches long and weighed 17H pounds dressed FISHES OF THE GULF OF MAINE KEY TO GULF OF MAINE STURGEONS 81 1. The successive bucklers in the dorsal row touch each other or even overlap; the space between the dorsal row of buck- lers and the uppermost of the two lateral rows is thickly set with coarse prickles Sea sturgeon, p. 81. 2. The successive bucklers in the dorsal row are separated one from the next by spaces up to Yi as long as the bucklers; the space between the dorsal row of bucklers and the uppermost of the two lateral rows is only sparsely strewn with fine prickles Short-nosed sturgeon, p. 84. Sea sturgeon Acipenser sturio Linnaeus, 1758 86 Jordan and Evermann, 1896-1900, p. 105 Description. — The skin of the sturgeon is armored with a row of large bony shields or bucklers along the middle of its hack (the succes- sive bucklers touching or even overlapping) with a second row of smaller bucklers high up along each side of the body; and with a third row, also smaller, lower down, along the line of transition from side to belly. Each buckler has a longitu- dinal keel with a spur, which is so sharp on small fish that those arc hard to handle, lower and blunter on large. On the average there arc 1(1 or 11 (10 — 16) bucklers in the mid-dorsal row; 28 or 29 (26 — 34) in each upper lateral row; and 9 to 14 in each of the lower lateral rows. The dorsal row runs from above the gill covers hack to the dorsal fin, and each of the dorsal shields reaches to the next, shield or even overlaps it. The upper lateral rows run from the gill openings back to the root of the tail (in; the lower lateral rows from close behind the pectoral fin to the pelvic fin, also from the pelvic fin back as far as the anal (in. And each shield in each of the two lateral rows is separated from the next shield l>\ a space up to one-half as long as the shields. The body is rather slender and rendered more or less pentagonal in ('toss section by the five rows of shields, instead of rounded as it is in the majority of bony fishes. The snout is narrow in young sturgeons less than 2 to 2)i feet long, depressed below the level of the forehead, nearly Hal below, M It still is an open question, Hint we cannot answer, whethei thi sturgeon of eastern North America is identical with the European sea stur- geon, is a recognizable race of the latter, or is a separati ti e last, its scientific name i- \eipmset axyrinchut MitchiU, 1S15 and longer (from the eyes forward) than the dis- tance is from the eyes rearward to the upper corners of the gill openings. But it changes shape as the fish grows, becoming blunter, straight in dorsal profile, and considerably shorter rela- tively. The mouth, situated on the under side of the head, is small, toothless (except in larval stages), with protractile lobed lips, and there are four pointed barbels in a row across the lower side of the snout in front of the mouth. The single rather small triangular dorsal fin stands far back, with its rear edge over that of the still smaller anal fin. The ventral fins are likewise far back. The pectorals are set almost as low as the plane of the belly.87 t'ulcr. Olive greenish or bluish gray above, gradually fading on the sides and changing rather abruptly below the upper lateral rows of shields to the white of the belly. Size. — The sea sturgeon is a very large fish. In the Delaware River where sturgeon persisted until recently in larger numbers than in Xew England, ripe males tire up to about 6 to 7 feel in length, averaging 65 pounds in weight; the spawning females (which arc larger), up to about 10 feet and to about 250 pounds, ss with a larger one taken from time to time. And the general run was about the same in the Kennehec, to judge from an average weight of 120 pounds for males and females together, during the years when a fishery was carried on there. But some still grow considerably larger in Gulf of Maine waters. Thus !) weighing between 350 pounds ami 600 >' Vladykov ami Beaulicu (Natural. Canad., vol. : :. 1946, pp 14 giv< a detailed account of the characters that separate the sea sturgeon froiti the laki titer fuleeteen* Raflnesque, 1817). " Accordit tot Comm. (1899), 1! 00, p. 277. Figure 30. — Sturgeon (Acipenser sturio), Potomac River specimen. From Goode. Drawing by H. L. Todd. 82 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE pounds were landed in Portland, Maine, from the South Channel, Georges Bank, Browns Bank, and Western Bank off Nova Scotia during the period 1927-1935.89 About 12 feet is perhaps the greatest length to be expected today. But 18 feet, reported for New England many years ago, may not have been an exaggeration, for sturgeon as long as that have been reported from Europe also. The heaviest Gulf of Maine stur- geon reliably reported (to our knowledge) was one of 600 pounds, landed in Portland by the steam trawler Fabia from Georges Bank, Decem- ber 21, 1932. The following relationship between length and weight, for sea sturgeons up to 7% feet long, taken in the lower St. Lawrence River,90 would probably apply to Gulf of Maine, fish, equally: 7 to 9 pounds at 30 inches (to fork of tail); 15 to 18 pounds at 40 inches; about 35 pounds at 50 inches; 55 to 57 pounds at 5 feet; about 100 pounds at 6 feet; and about 190 pounds at 7% feet. Habits. — The sturgeon makes most of its growth in salt water but enters fresh-water rivers to spawn, as do the salmon, the shad, and the alewife. The large adult fish enter (or once entered) the Gulf of Maine rivers late in the spring, working their way slowly upstream beyond tidewater before deposit- ing their eggs. So far as known, spawning takes place in our rivers in May, June, and perhaps as late as July. It has been suggested that some may spawn in brackish water from the fact that females with large eggs have been taken near Woods Hole in June and July (i. e., in the spawning season). Spawning leaves the spent "cows" in very poor condition. In the Delaware, however, and pre- sumably in Gulf of Maine rivers, they "become again quite plump, acquiring considerable addi- tional weight" 91 before they go down stream again, which some of them do not do until September, according to observations in the Delaware. But we do not know how many years in succession a given fish may spawn. A single female may produce as many as 2,400,000 eggs which hatch in about a week after they are fertilized.92 Judging from European observations on artificially reared sea sturgeon, '• Records collected by the late Walter H. Rich of the U. S. Bureau of Fisheries. w According to measurements and weights of 1,592 sturgeons by Vladykov, Rapp. Gen. Ministr. Chasse, Pech., Quebec (1948-1949), 1949, pp. 43-54. " Ryder. Bull. U. S. Fish Comm., vol. 8, 1890, p. 266. '- Ryder (Bull. U. S. Fish Comm., vol. 8, 1890, p. 231) describes the spawn- ing and early development of the sturgeon in the Delaware River. the larvae may be expected to grow to 12 mm. in length within 5 days after hatching; to 16-17 mm. in 2 weeks; to 20 mm. in 4 weeks; and to 4-5% inches in 2 months. Some 3roung sturgeon may live several years in the lower tidal reaches of the rivers in which they are spawned, until they have grown to a length of 2% to 3 feet, as appears to be the case in the Hudson.93 And it seems that they pass their entire growth period in the salt estuary of the St. Lawrence River, for sturgeons are taken there of all sizes from a few inches long up to 7-8 feet or longer.94 But others may descend during their first year, for sperlets only 5 to 6 inches long have been found at the mouth of the Delaware River and of the Elbe in Europe.95 Some Gulf of Maine sturgeon have taken to the sea by the time they have grown to 3 feet or so, as proved by the capture of sturgeons of that size at various points around the coasts of the Gulf, and off southern New England. And recent ob- servations in the Hudson by Greeley make it likely that all the sturgeon that are spawned in rivers emptying into the Gulf of Maine go to sea sooner or later to complete their growth.96 Sturgeon grow rather slowly at first while still in their parent streams. Four, for example, that were tagged in the lower St. Lawrence when 29 to 33 inches long, and recaptured nearby 2 to 3% years later, had gamed only about 2 to 5 inches in length per year.97 Very slow growth is also indicated by ages of 5 to 6 years at 24 to 28 inches; 7 years at 25 to 31 inches; and 8 years at 32 to 34 inches, for sturgeon from the tidal waters of the lower Hudson, as estimated from the mark- ings on their otoliths.98 It also seems that sturgeon, like many other fish, make most of their growth during the warm season in such situations for one marked fish in the Elbe did not grow at all between November and the following February, whereas a second grew from 17 cm. (Q}i in.) to 38 cm. (15 in.) in length between January 17 and " See Greeley (Supp. 26 Ann. Rept. Conserv. Dept. New York, 1937, pp. 68, 78-82, 89) for a study of the sturgeon in the Hudson River. 94 A series of 1,592 sea sturgeons from the lower St. Lawrence River, studied by Vladykov (Rapp. Gen. Minstr. Chasse, FSch. Quebec (1948-1949) 1949, pp. 53-66) included a good representative of sizes from about 4 inches up to 90 inches. es Prince reports a 6-inch sturgeon from nudson Bay (Rept. Sixty-seventh Meeting, British Assoc. Adv. Sci., Toronto, 1897, p. 687). " Qreeley, Suppl. 26 Ann. Rept. Conserv. Dept. New York, 1937, p. 82. •» Vladykov (Rapp. Gen. Ministr., Chasse, Pech. Quebec, 1948-1949, pp. 61-63, 66. table 19). °s Greeley, Supp. 26 Ann. Rept. Conserv. Dept. New York, 1937, p. 68, table 10. FISHES OF THE GULF OF MAINE 83 the following April, and a third from 43% cm. (17% in.) to 64 cm. (25% in.) from April 9 to the following December. But sturgeon grow much more rapidly after they go to sea, if ages (esti- mated from otoliths) of 1 1 years for a 75-inch sturgeon, and 12 years for two others of 88 and 100 inches are anywhere near the truth." The sturgeon is a bottom feeder, rooting in the sand or mud with its snout like a pig (the barbels serving as organs of touch) as it noses up the worms and mollusks on which it feeds and which it sucks into its toothless mouth with considerable amounts of mud. It also consumes small fishes, particu- larly sand launce. Small ones, while living in estuaries and around river mouths, subsist largely on amphipod and isopod Crustacea. Sturgeon, like salmon, eat little or nothing while traveling up river to spawn. When at ease sturgeon swim slowly to and fro, seeming very sluggish. But they are capable of darting ahead like an arrow on occasion, and they often come to the surface to jump clear of the water. Though they usually offer no resist- ance when netted, large ones are very strong. General range, — Coastal waters from the St. Lawrence River to the Gulf of .Mexico, running up into rivers to spawn; reported from Hudson Bay, also Scandinavia to the Mediterranean, if the American and European sea sturgeons belong to the same species. Occurrence in the Gulf of Maine. — The sea sturgeon is (or was) well known in the St. John, Penobscot, Kennebec, and Merrimac Rivers, and has even been taken some distance from the mouths of streams no larger than the Charles River and the Parker River in Essex County, Mass.,1 where some arc still seen jumping in July and one is taken occasionally. In fact, sturgeon once entered practically every stream of any size emptying into the Gulf of Maine. Wood, writing of Massachusetts in 1634, u described them as "all over the country, but best catching of them be upon the shoales of Cape Code and in the river of Merrimacke, where much is taken, pickled and brought for England, some of these be 12, 14 and 18 foote long." In fact, an odd no Sop foe t note 98. 1 Two sturgeon 44 and 45H Inches long, netted in the Parks] River at Newbury, Mass., July 23, 1933, are (or were) in the collection of the Boston Society of Natural History, now the New England Science Museum (Bull. Boston Soc. Nat. Hist., 69, Oct. 1933, p. 8). ''New England's Prospect, 1634, p. 37. sturgeon still enters the mouth of the Merrimac, witness one of 230 pounds netted there on Sep- tember 14, 1938 and landed in Newburyport.2 Sturgeons may be expected anywhere off the coasts of the Gulf of Maine during their sojourn in salt water. There is definite record of them at sundry localities on both sides of the Bay of Fundy ; off Mt. Desert Island; ha Penobscot Bay; in Casco Bay; at the mouth of the Piscataqua River; on the Boars Head-Isles of Shoals fishing ground, where several 3 to 4 feet long were taken in gill nets dur- ing April and May 1913; at the mouths of the Essex and Ipswich Rivers, where jumping stur- geon have been reported recently in the daily press;3 at the mouth of Gloucester Harbor, where an angler reports catching one of about 12 pounds while fishing for tautog; inside and outside Boston Harbor; at Provincetown; off Truro, Cape Cod; and at Nantucket, as well as along the southern New England coast to the westward. Some also extend their wanderings to the offshore fishing banks as they grow. Thirty, for example, rang- ing in weight from 120 to 600 pounds were landed in Portland and Boston by otter trawlers from Nantucket Shoals, from South Channel, and from Georges and Browns Banks, during the years 1927-1936.4 Probably all of these were on bot- tom when caught, to judge from their diet (p. 83), and from the fact that sturgeon have been hooked on cod and haddock lines as deep as 25 fathoms in Scandinavian waters. Nothing beyond this is known of their movements in our Gulf. Importance. — It is only the scarcity of the sea sturgeon in the Gulf of Maine that limits its com- mercial importance there and in the tributary rivers. The few taken are picked up acciden- tally in traps or weirs, in drift nets, or by the otter trawlers. In former years, when our streams were less obstructed and sturgeons more plentiful, the catch was of considerable value in some of the larger rivers. It is interesting, for instance, to read that sturgeon, doubtless from the Kennebec River and cured near what is now Brunswick, Maine, were shipped to Europe as early as 1628; and that large quantities were also shipped to Europe from near Ipswich, Mass., in 1635. In the Kennebec, where an intermittent fishery had long been maintained. » Reported in the Boston Globe. Sept. 15, 1938. 3 The Boston Herald, June 1950. • Reports collected by the late Walter H. Rich, U. S. Bureau of Fisheries, and notices in the daily press. 84 FISHERY BULLETIN OK THE FISH AND WILDLIFE SERVICE the catch was about 250 fish in 1880, yielding 12,500 pounds of meat, and not much less Mil lit 38 inches from Waquoit, on the southern Bfaore oi Massacl 11 Hull. Essex Inst., vol. 11. 1879, p. 27. A sturgeon wa rep I as bre- Ij from Boston Hftrboi m ■". ■< u igo, bul nun is no v.:i> now of checking the I 5. (i. 7. KEY TO GULF OF MAINE HERRINGS AND TARPONS Last dorsal fin ray prolonged 2 Last dorsal ray not prolonged 3 Dorsal fin originates in advance of the ventrals; scales only i leratelj large Thread herring, p. 112 Dorsal fin originates behind the ventrals; scales very large- Tarpon, p. 87 Belly rounded 4 Belly sharp edged •> Scales very small; mouth very large with upper jaw-bone extending considerably beyond I Ik; rear edge of the i point of origin of dorsal fin about over 1 hat of (lie ventral fins _ Ten pounder; p. 86 Scales large; mouth small, with upper jaw-bone extending rearward only about as far as the fronl edge of the eye; point of origin of dorsal fin well in advance of that of the \ entral tins ... • Uud Herring, p. 87 Head (tip of snout to edge of gill cover) very large, occupying about one-third the total length "f the body to base of the central rays of the caudal tin; free edges of scales Suted, not rounded, - . . - Menhaden, p. 1 13 Head about one-fourth the total length of the body; free edges of the scales rounded .... 6 Distance from point of origin of dorsal fin to tip of lower jaw (mouth closed) about as long as from origin of dorsal fin to base of central rays of caudal fin; edge of belly hardly saw-toothed, though sharp; general form comparatively shallow; there is a cluster of teeth on the roof of the mouth Sea herring, p. 88 Distance from point of origin of dorsal fin to tip of lower jaw (mouth closed) considerably shorter, than from point of origin of dorsal fin to origin of central rays of caudal fin; edge of belly more or less strongly saw-toothed, especially in space between the ventral and anal fins; general form deep; there are no teeth on the roof of the mouth-- 7 The tip of the lower jaw extends noticeably beyond the upper when mouth is closed Hickory shad. p. 100 The tip of the jaw does not extend appreciably beyond the upper when mouth is closed 8 86 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE 9. The upper outline of the forward part of the lower jaw (visible if mouth is opened) is nearly straight, ana does not show a pronounced angle; the upper jaw extends back about level with the rear edge of the eye Shad, p. 108 The upper outline of the forward part of the lower jaw is concave with a pronounced angle; the upper jaw reaches back only about to the level of the center of the eye 9 Breadth of eye is greater than distance from front of eye to tip of snout; back distinctly grey green; lining of belly cavity pale grey Alewife, p. 101 Breadth of eye is only about as great as distance from front of eye to tip of snout; back distinctly blue green; lining of belly cavity sooty or black Blue back, p. 106 Ten pounder EIojjs saitrus Linnaeus 1766 Jordan and Evermann, 1896-1900, p. 410. Description. — The ten pounder is herring-like in the arrangement of its fins, with the single and soft-rayed dorsal fin originating about midway along its back; in having no adipose fin; in the position of its ventral fins about midway between tip of snout and fork of tail ; and in its forked-tail fin. But its scales are very much smaller relatively than those of any of our herrings, and its mouth is much larger, with the upper jawbone extending rearward considerably beyond the rear edge of the eye. Being about one-sixth as deep as it is long, it is a much more slender fish than any of our herrings except the round herring, and its belly is rounded like that of the latter. But its trunk is more flattened sidewise than that of the round herring, its dorsal fin-origin is over the ven- trals (well in advance of the ventrals in the round herring), and its tail fin is much wider relatively than that of any herring, and more deeply forked. A more important structural character is that its tliroat is stiffened between the branches of its lower jaw by a long bony plate, which it shares with the tarpon, but which no member of the her- ring tribe has. Its closest affinity among fishes yet known from our Gulf is with the tarpon. But its scales are very much smaller than those of the latter, nor does its dorsal fin have the prolonged ray characteristic of the tarpon. Color. — Silvery all over, with the back bluish, the lower parts of the sides and the lower surface yellowish; the dorsal and caudal fins dusky yellow- ish and silvery; the ventral and pectoral fins yellowish speckled and dusky. Size. — The ten pounder is said to grow to a length of 3 feet,12 but few of those caught are longer than about 20 inches. General range. — Atlantic coast of America, from Brazil northward; commonly to North Carolina, in small numbers and less regularly to southern New England, and perhaps straying around the elbow of Cape Cod on rare occasions. The ten pounder of our Atlantic coast is represented in tropical-warm temperate seas in various other parts of the world by relatives so close that they may all finally prove to represent only the one wide-ranging species.13 Our only reason for men- tioning this southern fish is that one reported as from Chatham, Mass., may have been taken on the Gulf of Maine shore of Cape Cod.H Ten pounders are taken from time to time near Woods Hole. u Jordan and Evermann. Bull. 47. U. S. Nat. Mas., Pt. 1, 1896, p. 410. » Smith (Sea Fishes Southern Africa, 1949, p. 86) considers this probable. » This specimen, taken on October 19, 1888, and reported by Bigelow and Schroeder (Copeia, 1940. p. 139) is in the Museum of Comparative Zoology. Figure 38. — Ten pounder (Elops saurus), Massachusetts. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 87 Tarpon Tarpon atlanticus (Cuvier and Valenci- ennes) 1846 Jordan and Evermann, 1896-1900, p. 409, fig. 177. Description. — The tarpon is herring-Eke in gen- eral form and appearance, but it is made easily recognizable by the fact that the last ray of the dorsal fin is greatly prolonged, its free portion being as long as the fin is high or longer, and by die presence of the bony plate on the throat men- tioned above (p. 85) in the characterization of the family to which it belongs. Furthermore, the anal fin of the tarpon is deeply falcate; that of all Gulf of Maine herring-fishes rhomboid in out- line. The ventral fins, which are situated under or behind the dorsal fin in herrings, alewives, shad, and menhaden, are considerably in front of the dorsal fin in the tarpon, while the lower jaw of the latter projects relatively further; its scales are relatively larger; and its caudal fin is relatively wider. < blor. — Bright silvery all over, the back darker than the belly. Size. — Tarpon grow to a length of 6 to 8 feet ; the longest recorded was 8 feet 2 inches; the heaviest taken on rod and reel weighed 247 pounds.15 General range. — Tropical and subtropical coasts of .America, from Brazil to Long Island, casually to Cape Cod, and to Nova Scotia, where it has been recorded off Isaacs Harbor and in Harrigan Cove.16 Its chief center of abundance is in the West Indies, about Florida, and in the Gulf of Mexico. Occurrence in tin Gvlj oj Maine. — A specimen 5% feet long, taken at Provincetown on July 25, 1915,17 is the only record of the tarpon in the Gulf of Maine, which it reaches only as an accidental straggler from the south. Round herring Etrumeus sadina (Mitchill) 1815 Jordan and Evermann, 1896-1900, p. 420. Description. — The most distinctive feature of this fish, among herrings, is that its belly is rounded, not sharp edged. It is, furthermore, the most slender of our herrings, its body being only " Taken on rod and reel in the Panuco River, Mexico, Mar. 24, 1938, by H. W. Sedgewick. ■» Halkett, Check List, Fishes Canada. Newfoundland, 1913, p. 45. lis, No. 28, 1918, p. 3. ; v 3 ' WW ': ^k Figuke 39. — Tarpon (Tarpon atlanticus). New Jersey. From Goode. Drawing by H. L. Todd. Figure 40. — Round herring (Etrumeus sadina). 88 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE one-sixth as deep as long, thus suggesting a smelt in its general outline. Its dorsal fin, too, stands wholly in front of the ventrals instead of over the latter, as in herring, alewives, and shad ; and there are fewer anal fin rays (only about 13, whereas the herring has about 17, the alewife about 19, and the shad about 21) than any of the latter. ( 'olor. — Olive green above with silvery sides and belly. Size. — Eight to ten inches long when adult. General range.- — Atlantic and Gulf of Mexico coasts of the United States; occasionally common as far north as Woods Hole; sometimes straying past Cape Cod, to the mouth of the Bay of Fundy. Occurrence in the Gulf of Maine. — This southern fish has been taken at Provincetown, Mass., whence the Museum of Comparative Zoology has two specimens; one was taken in the Yarmouth River which empties into Casco Bay, and one in the bay itself on September 15, 1924;18 it has been reported from Jonesport, Maine; also from East- port, Maine, in 1908. 19 And a number of them were taken at Campobello Island, at the mouth of Passamaquoddy Bay in September 1937.20 Herring Clupea harengus Linnaeus 1758 Sea herring; Labrador herring; Sardine; Sperling; Brit Jordan and Evermann, 1896-1900, p. 421. Description. — The sea herring is typical of its family in form, with body so flattened that it is much deeper than thick; moderately pointed nose; large mouth situated at the tip of the snout and >» Reported to us by the late Walter H. Rich of Hi': U. S. Bureau of Fish- eries. w Reported in the newspapers. !° Reported by Loira, I'rogr. Rept. 21. Atlantic Biol. Sta. Fish. Res. Bd. Canada, 1937, p. 5; and by McKenzie, Proc. Nova Scotia Inst. Sei., vol. 20, 1939, p. 15. lower jaw projecting a little beyond the upper when the mouth is closed; sharp-edged belly; and deeply forked tail. The dorsal fin stands over the much smaller ventrals, its origin about midway the length of the body. The scales are large, their rear margins rounded, and so loosely attached that they slip off at' a touch. There is no adipose fin, and its absence at once distinguishes all the her- rings from any of the salmon tribe. The chief ana- tomical character separating the sea herring from the shad and from the several alewives (genus Po- molobus) is that it has an oval patch of small teeth on the vomer bone in the center of the roof of the mouth. Conspicuous field marks separating her- ring from shad, hickory shad, and alewife are that the point of origin of its dorsal fin is about midway of the length of its trunk (considerably farther for- ward in the others) ; its body is not so deep, a differ- ence shown better in the illustrations; and the sharp midline of its belly is only very weakly saw- toothed but is usually strongly so in the others, especially along the space between ventral and anal fins. Color. — Deep steel blue or greenish blue on the back with green reflections; the sides and belly silvery; the change from dark belly to pale sides often marked by a greenish band. The gill covers sometimes glisten with a golden or brassy gloss; indeed, fish just out of the water are iridescent all over with different hues of blue, green, and violet; but these colors soon fade, leaving only the dark back and silvery sides. The ventral and anal fins are translucent white; the pectorals, however, are dark at the base and along the upper edge; the caudal and dorsal fins are dark grajush or shading into green or blue. Size. — Herring grow to a length of about 17 inches and to a weight of about 1 % pounds. Habits. — The herring is a fish of open waters, Figure 41. — Herring (Clupea harengus). From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 89 traveling as a rule in schools of hundreds or thou- sands; single fish are seldom seen, or even small companies of a few dozen. As a rule all the indi- vidual members of a school are about the same size, whether large or small. It is not known how long any given school may preserve its identity as such. Fridriksson and Aasen,21 it is true, found that herring tagged and released together might be recaptured from widely separated localities, sug- gesting that schools are more or less temporary formations. But this may not apply to schools that have assembled under natural conditions. When a school is at the surface, as often hap- pens on a calm day, its presence is betrayed by a fine rippling of the water, but we have never seen herring "finning" or lifting their noses above the surface as menhaden often do (p. 114). They come to the surface most often by night, when their presence is betrayed by their luminous trails, if the water is "firing," as we have often seen. A school is likely to be more or less stationary when feeding, its members swimming slowly to and fro and drifting as a whole with the current.22 But at other times schools are seen traveling with in- dividual fish swimming side by side, rank below rank, as far down in the water as the eye can see from a boat, all heading in one direction appar- ently with some purposeful intent. We have often watched schools of "sardine" size streaming close past a certain rocky headland in the southern side of Massachusetts Bay, seemingly in unending procession . As Dr. Huntsman points out, "There is no in- dication that herring swim against the current unless the water is somewhat turbulent." 23 If they do so under such conditions, it depends on the relationship between their rate of swimming and the strength of the current whether they ac- tually make headway against it or lose ground, tail first. We might also add that schools of herring, like schools of menhaden, are not so easily frightened by the approach of a boat, as mackerel often are, and striped bass. Herring do not jump unless frightened. But the smaller sizes are often seen jumping when pursued from below by larger pred- " Eept., Norwegian Fisb. Mar. Invest. Skriftcr, vol. 9, No. 11, Rept. 1, 1950, p. 22. "Huntsman (James Johnstone Memorial Vol., 1934, p. 83) gives an in- teresting account of the movements of herring schools in Passamaquoddy Bay. » James Johnstone Memorial Vol., 1934, p. 84. 210941—53 7 atory fishes, such as silver hake or striped bass, a common spectacle. Fridriksson and Aasen found that herring, held in live-nets, swam con- stantly at a rate of about 0.2 to 0.25 sea miles per hour (6-8 meters per minute) when not disturbed. And it is certain that they are capable of long journeys, for a number of herring tagged on the northeast coast of Iceland have been recaught in southern Norway, and some vice-versa.24 The activity of the herring is controlled in great part by the temperature of the water. In Pas- samaquoddy Bay, for example, they are "ob- served to move very sluggishly when the water is coldest in February and March," 25 and probably this applies all around the periphery of our Gulf, for the upper 20 fathoms ordinarily cools to about 33 to 36° F. during those months, with the sur- face often chilling to the freezing point of salt water in bays and harbors. The herring become active again when the water has warmed to about 40 to 43°. Food.— The herring is a plankton feeder. When first hatched, and before the disappearance of the yolk sac, the larvae (European) feed on larval snails and crustaceans, on diatoms, and on peridinians, but they soon begin taking copepods, and depend exclusively on these for a time after they get to be 12 mm. long, especially on the little Pseudocalanus elongatus.^ As they grow older they feed more and more on the larger copepods and amphipods, pelagic shrimps, and decapod crustacean larvae. Examination of 1,500 stomachs27 showed that adult herring near East- port were living solely on copepods and on pelagic euphausiid shrimps (.\ftganyctiphanes norwegica), fish less than 4 inches long depending on the former alone, while the larger herring were eating both. When feeding on euphausiids, we have often seen them pursuing the individual shrimps, which frequently leap clear of the water in their efforts to escape. Even in winter, when shrimp are rarely seen at the surface, Moore found them an important article in the diet of the Eastport herring. And it is likely that the local appear- » Fridriksson and Aasen, Rept. Norwegian Fish. Mar. Invest., Skrifter, vol. 9, No. 11, Rept. 1, 1950, pp. 26-27. " Huntsman. James Johnstone Memorial vol., 1934, p. 83. » The diet of herring, young and old, in the English Channel and in the North Sea has been described by Lebour in a series of papers (see especially Jour. Mar. Biol. Assoc. United Kingdom, vol. 12, 1921, pp. 458-467), by Hardy (British Fisheries Invest., Ser. 3. vol. 7, No. 3, 1924), and by Jesperson (Medd. Komm. Havund. Ser. Plankton, vol. 2, No. 2, 1928, Copenhagen). » Moore, Rept. U. S. Comm. Fish. (1896), 1898, p. 402. 90 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE ances and disappearances of schools of large fish in the open Gulf are connected with the presence or absence of euphausiid shrimp of one species or another. A few of the larger fish, however, as well as the smaller ones, will usually be found full of copepods, even when both shrimp and copepods abound, and copepods are the chief dependence of all our herring, large and small, in the absence of shrimp. The amphipod genus Euthemisto also is an important food for herring in European seas ; hence the absence of Euthemisto from the herring stomachs examined by Moore and by us has doubtless been due to the com- parative scarcity of this large active crustacean in the coastwise waters of the Gulf of Maine. The particular species of copepods on which Gulf of Maine and Woods Hole herring depend have not been identified, but we might guess that Calanus predominates, with Pseudocalanus, Acar- tia, and Centropages, and Temora also, at its times of abundance, while Euchaeta offers a rich food supply when the schools seek the deep waters of the basin frequented by these mammoth copepods. In default of an abundant supply of Crustacea, and sometimes even when these are plentiful, herring feed on whatever molluscan larvae, fish eggs, Sagittae, pteropods, annelids that the water contains, even on microscopic objects as small as tintinnids and Halosphaera. But the smaller microscopic plants, either diatom or peridinian, are never found in the stomachs of herring more than 15 to 20 mm. long, probably because their gill rakers are not fine enough to retain them. Although herring normally are not fish eaters, small launce, silversides, and the young of then- own species have been found in them at Woods Hole. And Templeman28 reports them as con- suming quantities of small capelin, in winter, in Newfoundland waters. Herring ordinarily pick up their food objects individually by a "definite act of capture" as Battle expresses it,29 while she found that herring in the aquarium at St. Andrews did not feed in complete darkness, though they did in faint liglil. But it seems that when feeding on very small objects they may strain these out with their branchial sieves as the manhaden does (p. 114), for Moore, a very accurate observer, described them as swimming open mouthed when feeding on " Bull. Newfoundland Government Lab., No. 17, 1948, p. 133. " Ann. Rept. Biol. Board Canada (1933), 1934, p. 14-15. minute crustaceans, crossing and recrossing on their tracks.30 Doubtless it is because of their feeding habits that herring seldom take a baited hook, if they ever do. But we think it likely that large ones when feeding on shrimp would take an artificial fly, as spent and hungry alewives will (p. 104) on their return to salt water, and as shad will on their way upstream (p. 109). Enemies. — The herring is the best of all bait fishes in our Gulf, where it is preyed upon by all kinds of predaceous fish, especially by cod, pollock, haddock, silver hake, striped bass, mack- erel, tuna, salmon, and dogfish, and by the mack- erel sharks. Silver hake, in particular, often drive schools of herring up on our beaches, where pursued and pursuers alike strand on the shoaling bottom. We once saw this happen at Cohasset in Massachusetts Bay many years ago, on an October morning, when hake and herring were so inter- mingled in shallow water at the height of the car- nage that we soon filled our dory with the two, with our bare hands. The finback whales also devour herring in great quantities. The short- finned squid (Ilex) likewise destroys multitudes of the young sardines. On one occasion near Provincetown, in June 1925, we watched packs of perhaps 10 to 50 squids circling around a school of 2- to 4-inch herring, bunching them into a compact mass. Individual squids then darted in, seized one or two herring, ate only a small part, then darted back for more. A silvery streak of fragments of dead herring remaining along the beach bore witness to the carnage. Breeding habits, development and groxoth. — Much attention has been devoted to the breeding habits and growth of the herring by European zoologists, by Moore, and by Huntsman in our own Gulf, and by Lea 31 in more northern Canadian waters. Herring may spawn in spring, in summer or autumn, according to locality, or both in spring and autumn (for further information on this matter, see p. 98). They do so chiefly on rocky, pebbly, or gravelly bottoms, on clay to some ex- tent, probably never on soft mud. Spawning in the Gulf of Maine (including the Bay of Fundy), takes place chiefly from 2 or 3 fathoms down to about 30 fathoms; perhaps never in the littoral »° Rept. U. S. Comm. Fish. (1896), 189S, p. 402. » Age and growth of the herrings in Canadian waters. 1911-15 (1919), pp. 75-164. Oanad. Fish. Exped., FISHES OF THE GULF OF MAINE 91 zone, nor has herring spawn ever been reported as cast up by the surf on the beaches of New England, a fate that often overtakes it in the Gulf of St. Lawrence. Occasionally they spawn as deep as 100 fathoms in Scandanavian waters, perhaps also in the eastern basin of our Gulf where the sea floor is hard, not soft and muddy as it is in the basins in the western side. During the act of reproduction as observed by Moore at Cross Island and at Machias, Maine, "the fish were darting rapidly about, and those who have opportunity to see the fish spawning in more shallow water where ob- servation is more favorable, state that both males and females are in constant motion, rubbing against one another and upon the bottom, appar- ently by pressure aiding in the discharge of the eggs and milt." 32 A female herring may deposit from 20,000 to upwards of 40,000 eggs, according to her age and size, averaging about 30,000. In sexually mature herrings, the genital organs are so large just before spawning commences that they make up about one-fifth the total weight of the fish. The eggs sink to the bottom, where they stick in layers or clumps to the sand or clay, to sea- weeds, or to stones, by means of their coating of mucus, or to any other objects on which they chance to settle. They are often found massed on net warps, anchors, and anchor ropes. The individual eggs are 1 to 1.4 mm. in diameter, de- pending on the size of the parent fish and also, perhaps, on the local race of fish involved. Tbe period of incubation is governed by temperature; European students tell us that it requires as long as 40 days at 38-39°, 15 days at 44-^6° and 11 days at 50-51° F.; while experiments on the Mas- sachusetts coast by the U. S. Bureau of Fisheries gave 10 to 12 days in the temperature prevailing there in autumn. And MacFarland 33 found that all of the eggs kept at Grand Manan at about 59° (15° C.) hatched, but that none hatched at 32-41° (0-5° C), and that all died that were warmed to 68° (20° C). Ten to fifteen days might be stated as an average incubation period for the Gulf of Maine, under existing temperatures. The larvae of the herring family are very slender and can easily be distinguished from all other young Gulf of Maine fish of similar form (e. g., launce, smelt, or rock eel) by the location of the " Moore, Kept. V. S. Comm. Fish. (1896), 1S98, p. 412. » Rept. Biol. Board Canada (1930), 1931, p. 23. Figure 42. — Eggs of the herriug (Clupea harengus), attached to seaweed (European). After Ehrenbaum. vent, which is so far back that it lies close to the base of the tail. But it requires critical examina- tion to distinguish our several clupeoids one from another in their early st;> The sea herring is about 5 to (i mm. long at hatching, with a small yolk sac that is absorbed by the time a length of about 10 mm. is reached. The dorsal fin is formed at 15 to 17 mm.; the anal at about 30 mm.; the ventrals are visible and the tail well forked at 30 to 35 mm.; and at about 40 mm. {V/2 in.), the little fish begins to look like a herring. According to Huntsman's observations, fry produced on the Grand Manan spawning grounds in late summer and early autumn grow to a length of 17 to 20 mm. by the end of November or first of December; they are 26 to 50 mm. (1-2 in.) long in March and April and 50 to 60 mm. (2-2% in.) long by June when fry of this size are abundant in the St. Andrews region. Tins is in line with our own observations that fry of 2 to 2% inches (50-65 mm.) predominate among the young her- ring at Provincetown at the end of June, and fry of 2){ to 4 inches (54 to 100 mm.) on Nantucket 92 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 43. — Larval stages of the herring (Clupea harengus) European. After Ehrenbaum. A, newly hatched, 7 mm.; B, 10 mm.; C, 19 mm.; D. 29 mm.; E, young fry, 41 mm. Shoals in mid-July. They grow to about 3K to near 5 inches (90-125 mm.) by the end of their first year of life; fish of that size, presumably of the previous autumn's hatch, are abundant in the fall in the Bay of Fundy, and at Boothbay, Maine. The growth rate is about the same at Woods Hole, where herring spawned in October and early No- vember are 3 to 5 inches (76-125 mm.) long by the following autumn. The Norwegian herring, also, average about 5 inches (125 mm.) long at the end of their first year, according to Hjort, and North Sea herring are about 4 inches (100 mm.) long then.34 Subsequent growth. — The herring has proved a particularly favorable object for growth studies based on the structure of the scales.35 Without pursuing this subject, which would lead us far afield, we may point out that herring not only grow at different rates at different times of year, with the contrast between the rapid growth of summer and the slow growth of the winter greater or less in different seas, but that they grow rapidly when young and slowly thereafter in some locali- ties, whereas they may grow slowly at first in other localities, but sustain a more even growth to old age. The Dogger Bank herring, for example, in the North Sea approximate 4 inches iD length at the end of the first year, 8% to 9 inches at the end of the third year, 10% at the end of the sixth, and 11% to 12 inches at the end of the ninth, though with considerable variation. The Norwegian herring, however, spawned in the year 1899, averaged only 7% inches when 3 years old, but were as large as the Dogger Bank fish of equal ages by their sixth year and subsequently.36 Newfoundland herring grow more slowly at first than those in the southern side of the Gulf of St. Lawrence, but catch up with them as they grow older. Huntsman credits the Bay of Fundy herring with about 10 inches at the end of their third year; i. e., when 4 years old, which agrees closely with an average growth of 9% inches at 4 years as cal- culated by Lea for Gulf of St. Lawrence fish. The average growth rate of the older Bay of Fundy fish probably falls between that of the Gulf of St. Lawrence fish and that of the herring of outer Nova Scotia which grow a little faster; i. e., to between 10% and 11% inches at 5 years; between 11 and 12% inches at 7 years; and betweeD 12% and 13% inches at 9 years.37 Bay of Fundy herring make most of their growth from May to Septem- ber. In the southern parts of our Gulf, where the growth period probably continues a month later into the fall, they may grow as fast as they do along outer Nova Scotia. When the little herring have reached an age of about 2 years and a length of 7% to 8 inches (190-200 mm.) they accumulate large amounts of fat among the body tissues and viscera during the warm months of the year when growing rapidly, but lose this fat in winter and also at the approach of sexual maturity. We can bear witness and the fact is well known to fishermen that this "fat" stage is as characteristic of American waters as of European, where "fat" herring are the objects of extensive fisheries. According to Moore, who examined thousands " Huntsman (Canad. Fish. Exped. (1914-1915), 1919, pp. 168-169) believed he could recognize spring as well as autumn-spawned herring fry in the Bay of Fundy, and credits the former with a length of about 90 mm. by the first, and 150 mm. by the second, winter. But this seems to call for confirmation, it being unlikely that any herring now spawn there in spring (p. 98). « See Lea (Canad. Fish. Exped. (1914-15), 1919, pp. 75-164) for an account of age determination by analysis of the scales, as applied to the herring. " Eapp. and Proc. Verb., Cons. Internat. Explor. Mer, vol. 20, 1944. •' As scaled from Lea's diagrams (Canad. Fish. Exped. 1914-1915 (1929), figs. 40 and 41). It has been found that the Norwegian herring grow from April to September only, remaining practically stationary in length from October until March; see Lea (Pub. de Circ, Cons. Perm. Internat. Explor. Mer, No. 61, 1911, pp. 35-57) and Hjort (Eapp. Proc. Verb., Cons. Perm. Internat. Explor. Mer, vol. 20, 1914). FISHES OF THE GULF OF MAINE 93 of fish about Eastport, herring rarely spawn when less than 9K inches long; usually not until they are 10 to 10K inches; and most of the spawners are 12 to 13 inches long. This means that some few spawn when only 3 years old, if the growth schedule outlined above is correct, but most of them not until 4 years or older, to continue spawning annually thereafter as long as they live. In Norwegian waters, too, a few spawn at 3 years, many at 4 years, and the majority at 5 years; some few not until 6 years old. Herring have been seen as old as 20 years, and they may live even longer. Success of reproduction. — The relative abund- ance of any species of sea fish from year to year depends less on how many individuals spawn in any locality than it does on how many of the resultant fry survive. And the many age analyses that have been made of herring in European waters have proved that while a very large crop of young may be produced in some years, hardly any are in others, even in favorable nurseries. Apparently this applies more to the northern breeding grounds than to the southern (to some extent, however, to all) the result being that the herring spawned in some one favorable breeding season may dominate the schools over large areas for many years, or until another successful breeding year comes, producing another large crop. In Norwegian waters, for example, the herring produced in 1904 was dominant in the catches for the next six years, at least; this is a classic instance. Lea found, similarly, that herring hatched that same year (1904) dominated the catches on the west coast of Newfoundland as long afterwards as 1914 and 1915. And while precise information is not avail- able for our Gulf, no doubt the same rule governs there. One case, at least, is well documented of a particular body of Bay of Fundy herring that received no important recruitment for something like 10 years, when the few still remaining seem to have disappeared, from old age (p. 99). Various explanations have been proposed to account for this, such as abundance or scarcity of microscopic plankton, favorable or unfavorable temperature, salinity, or other factors, all of which may enter in. And while it is during the first few weeks of life that the herring is most vulnerable, it is also possible that the conditions under which the parent fish lived for the year preceding spawning may influence the fate of the fry. Whatever the explanation, the fact that such fluctuations do occur from year to year, in the numbers of fry reared is of the greatest practical interest to all concerned with the sea fisheries, as evidence that variations existing in the stock of herring, and consequently in the catch, may be due more to the success or failure of reproduction than to any effect the fishery may have on the stock. General range. — Both sides of the North Atlan- tic. Off the European coast the herring ranges north to Norway, Iceland, Spitzbergen, and the White Sea; south to the Straits of Gibraltar. It is known on the American coast as far north as northern Labrador and the west coast of Green- land; regularly and commonly as far south as Cape Cod and Block Island ; and it is occasionally seen in small numbers as far south as Cape Hat- teras in winter. It is replaced by a close ally (C. pallasii) in the North Pacific. Occurrence in the Gulj oj Maine. — To list the localities where herring have been recorded would be to mention every hamlet along our coasts whence fishing boats put out, for more or less herring, large or small, appear at one season or another around the entire coast line of the Gulf, and on the offshore fishing banks as well. They also enter bays and estuaries freely, but they have never been reported in our Gulf from water that is appreciably brackish; perhaps 2.8 percent salinity 3S may be set at about their lower limit. The distribution of commercial catches, plotted by Needier (fig. 44) 3a shows that herring are far more plentiful from Casco Bay eastward along the coast of Maine, and especially in the Passama- quoddy Bay-Grand Manan region than they are along the western shores of the Gulf on the one hand, or up the Bay of Fundy on the other, or along western Nova Scotia. Thus the landings per unit length of coast averaged 3 times as great for the Passamaquoddy-Grand Manan region and for the coast of Maine to Mount Desert, as for the coast sector from Mount Desert past Penobscot Bay; about 4 times as great as for the Maine coast as a whole, westward and southward » Surface, in Bay or Fundy in May. >• A reliable index, for the herring is a valuable fish. 94 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE 70 68 46 HERRING FISHERY CATCHESOF THE YEAR 1919 400,000 POUND LOTS PASSAMAQUODDY PENOBSCOT BAY " _3 L—i ; 1 r'lil l^s I I I ' 1 1 * ft V\ *\ a .' CAPE C00*»\\ /""■-"" 42 68 66^ 64 Figure 44. — Catches of herring for the year 1919. Each dot represents 400,000 pounds. After Needier. from Penobscot Bay; and 13 times as great as for the coast of Massachusetts,40 for the years 1919, 1928, 1929, and 1930.41 Present day landings of upwards of 30 million pounds of sardines alone, for Charlotte County, New Brunswick, even in poor years, up to some- thing like 100 million pounds in good years, plus some 9-14 million pounds of larger herring, con- trasted with a maximum of only about 17 to 18 million pounds reported for 1947 for any sector of the Maine coast of comparable length,42 show that the Passamaquoddy-Grand Manan region has not lost its preeminence as a herring center. The abundance of little herring there is, in fact, the outstanding feature of the distribution of fishes in * Omitting the landings for Suffolk County, Mass., since these represent fish discharged at Boston by the vessel fisheries from offshore. " Graham, Jour. Biol. Board Canada, vol. 2, No. 2, 1936, p. 129, table 2. « Scattergood has given an interesting analysis, regional and seasonal, of the 1947 catch of herring for the coast of Maine. the Gulf of Maine. A catch of about 2,400,000 pounds for Massachusetts in 1947, contrasted with some 11,300,000 pounds for the Penobscot Bay region alone in that year, illustrates how much less rich in herring the southwestern side coast line of the Gulf is than the sector that happens to be the least productive part of the northern coast line of the Gulf. Fishermen tell us, too, that herring are much more regular in their occurrence from year to year in the Passamaquoddy-Grand Manan region than they are either off western Nova Scotia in the one direction, or along the coast of Maine in the other. And this is borne out by such statistics as are con- veniently available. Thus only one-fourth to one- fifth as many pounds of herring were caught in the Penobscot Bay region 43 in 1947 as either eastward « Scattergood's statistical areas 11-14. FISHES OF THE GULF OF MAINE 95 to Lubec Narrows on the one hand, or westward past Casco Bay to Cape Elizabeth on the other,44 whereas the catches for 1919 were rather evenly distributed along the northern and eastern Maine coast as a whole. We find herring even more and more sporadic in their appearances and disappearances, both from place to place, from week to week, and from year to year, passing southward around the western periphery of the Gulf. Very few, for example, are seen on the southern side of Massachusetts Bay in some years (as in 1950 and 1951) ; many schools in others. And herring are such wandering fish in general, here today and gone tomorrow even in their centers of abundance, that the successful location of the weirs depends largely on intimate local knowledge and on close observation of the movements of the schools. Herring appear, also, to be far less plentiful on the offshore banks and less regular in their occur- rences there than they are in their inshore center of abundance in the northeastern part of the Gulf. Trawlers, it is true, occasionally pick up schools on Georges Bank and on Browns Bank, as in 1931, when catches of 3,000 pounds were reported on the northern edge of Georges and of 2,800 pounds on the southwestern part in October. Schools, too, are occasionally reported as seen at the surface, by Albatross III for instance, in April-May 1950. Fishermen used sometimes to set drift-nets on Georges for herring for bait in the days of the long line fishery, and small numbers up to 130-160 per haul, were trawled by Albatross III, widespread on the western part in depths of 20 to 50 fathoms in May 1950, as well as off southern New England.45 But it is more usual for trawlers operating on Georges to pick up only odd fish or none. Thus the maximum catch on any trip during the otter trawl investigation of 1913 was only a dozen or two; 42 hauls by the Eugene H, in late June 1951, yielded only one herring, fishing from Nantucket Lightship out onto the south- central part of Georges; and the stomachs of cod caught on Georges seldom contain herring, if they ever do.46 The appearance of schools of large herring or of small is distinctly a seasonal event off most parts ** Coast sectors of comparable length. »' Average catches per haul about 56 fish at 22 to 40 fathoms, and 28 at 41 to 60 fathoms, but only 6 at 51 to CO fathoms. '* W. F. Clapp found no herring in many cod and haddock stomachs examined by him on Georges Bank. of our coast, and the picture is made still more complex by differences in the behavior of sardine- size, "fat," and spawning herring, the reasons for which are not yet well understood. The newly spawned fry, less than % of an inch (9-11 mm.) long, have been taken in September in the lower part of the Bay of Fundy, a product, doubtless, of the Grand Manan and West Nova Scotia spawning; also in October in Gloucester Harbor where one tow-net haul yielded us a great number on the 24th, in 1916. And they are to be expected wherever herring spawn in numbers in any particular year. It seems likely that most of them remain near their birth place during their first autumn and whiter, when the circulation of the Gulf is in its least active stage. But they become widely distributed during the spring (March-May), when 1% to 2 inches (30-50 mm.) long, both in the lower Bay of Fundy, around the entire periphery of the open Gulf, east as well as west, out over the basin, and on the northern and eastern parts of Georges Bank.47 Little seems to be known in detail about the movements of herring during their first year, but those that find their way into enclosed waters where mid-summer temperatures are high, such as Duxbury and Plymouth Bays and Provincetown Harbor, appear to move out during the early part of the summer, being reported as far less plentiful l line in June than they are in April and May. Sardine-size herring, 4 to 8 inches long including 1- and 2-year-olds, are to be expected in abundance all summer east of Penobscot Bay, and particularly in the Passamaquoddy Bay region, where thej support the sardine fishery for which the latter is famous, and where they are present throughout the year. It is probable, however, though not proved, that the 1- to 2-year-olds (fish in their second and third years) do not appear along the southwestern coasts of the Gulf until several months later in the season than the little fish of }{ to 2 inches do, that were hatched the preceding autumn. Thus it usually is not until late June, July, or August that "sperling" of 4 to 7 inches are reported in numbers off the Massachusetts coast, or that we *' During March and April 1920 we took them near Cashes Ledge, on the northern and eastern parts of Georges Bank, of! Seal Island; off Yarmouth, Nova Scotia; near Machias, Maine, and over the basin in the offing; near Boothbay; and near the Isles of Shoals. Graham (Jour. Biol. Board Canada, vol. 2, No. 2, 1936, p. 112, fig. 8) found them equally widespread in the open Gulf in May 1932, also in the lower Bay of Fundy (none, however, at the head of the Bay). 96 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE have seen them there. They are even more er- ratic, too, in their appearances and disappearances in Massachusetts Bay and along Cape Cod than they are to the eastward of Mount Desert. At Cohasset, for example, on the southern shore of the Bay where we have had many years' experi- ence, schools of sperling are here today in summer and early autumn, but gone tomorrow. It is also our impression that the sperling, like the larger herring, are not only far less concentrated in favorable localities around the southwestern shore of the Gulf than they are to the north and east, but far less numerous on the whole.48 These first two year classes (the fish in their first year having grown to a length of 3 or 4 inches by September; those in their second year to 7 to 9 inches) begin to thin out from the shore waters of the open Gulf after the middle of October as the water cools, and few "sardines" are taken there after early December. The corresponding ebb and flow, so to speak, for the sardine is suggested in a striking way by the average monthly catches of sardines by the weirs in Charlotte Co., New Brunswick (Passama- quoddy Bay, Campobello, and Grand Manan) for the year 1920, which are equally illustrative of conditions today: Month January February,. March Pounds Month Pounds 11,000 July 3,315,000 None August 6,475,000 56, 000 September 6, 730, 000 April. - . 1,049,000 October 6,012,000 May 3, 036, 000 November 1, 325, 000 June 2,542,000 December 147,000 Here, however, the seasonal variation (as Dr. Huntsman informs us) is simply a matter of local availability, for sardines remain in Passama- quoddy Bay all winter, but do not move about much then. Probably the sardines winter mostly on the bottom. And there is no reason to sup- pose that the bulk of them travel far in any part of the Gulf. Very little is known about the Gulf of Maine herring during their third summer, when they have passed the "sardine" or sperling stage and have not yet reached spawning age. In some years these "fat" herring, as they are often called, 48 No particular attention is paid to sperling around Massachusetts Bay, for they are too small to be in demand lor hait, and they are not plentiful enough (or not concentrated enough) to support a sardine fishery there. or "summer" herring, weighing up to about one pound (they are called "spawn" herring locally, but this is an error), are taken in the traps at Provincetown for a week or so about mid-April; they are taken at about the same time off Glouces- ter (in 1915 they were reported 8 to 15 miles off Cape Ann on the 17th), and they are said by the fishermen to "show" first off Seguin Island in May and June, off Mount Desert late in summer. Doubtless they form a large part (just what pro- portion is not known) of the catches of herring larger than sardines that are made in the Passa- maquoddy Bay region, also around Grand Manan. As a rule few of them are taken inside the inner islands elsewhere, though they came into the har- bor of Boothbay about May 14 in 1914. When a mackerel seiner picks up a school of herring out in the open Gulf in summer,49 or when an otter trawler makes a catch of herring on Georges Bank (p. 95), most of them are very fat and show no signs of approaching sexual matu- nty. Thus it seems that they tend to keep farther offshore than do either the younger herring or the still older mature herring. The peak season for herring larger than "sar- dines" inshore in the northeastern part of the Gulf is ordinarily from July through October; i. e., some 2 months less than that for the sardines (see p. 96). But a greater proportion of the larger fish continue available there through the cold months than of the younger fish, to judge from the fact that considerably larger catches are made of big herring in winter than of sardines, whereas the total local catch is much larger for the latter than for the former. A report m on the average monthly landings of large herring for Charlotte County, for the period 1920-1931, to the nearest 1,000 pounds, follows: Month Pounds Month Pounds January 132,000 July 1,065,000 February 164,000 August 4,334,000 March 275,000 September 7,098,000 April 312,000 October 2,817,000 May 306,000 November 646,000 June.. 284,000 December 268,000 Large herrings, yearly average 17, 701, 000 "Sardines", yearly average 30,698,000 « Many events of this sort have been reported. For example, a large catch of fat summer herring was made on Georges Bank and reported to the Massa- chusetts Commissioners in the mid-summer of 1901. » From Graham, Jour. Biol. Board Canada, vol. 2, No. 2, 1936, p. 130, table 3. FISHES OF THE GULF OF MAINE 97 Large herring (Dr. Hunstman tells us) are also present there throughout late winter and spring, though few find their way then into the weirs. In the southeastern part of the Gulf, as typified by Cape Cod Bay, large herring appear inshore in greatest numbers to about June and again in the autumn, with very few (and not many sardines) in June or July. This is illustrated by the largest and smallest catches made in 8 traps at North Truro for different months during the years 1946 to 1952. The following data are contributed by the Pond Village Cold Storage Company: Minimum Maximum Month (in pounds) (in pounds) April 0 117,375 May 221 523,550 June 0 88,057 July 0 0 August 0 1,000 September 0 57,287 October 0 9,526 November 0 170,435 The earliest catch of sardines there in those years, or in 1935, 1938, or 1943 was sometime in May, the latest November 16 to 17; the earliest catcb of large herring was made between April 20 and 30, the latest on December 10th. In most years the large herring vanish from the Massachusetts coast at some time in December. In 1950, for example, they vanished about De- cember 4th from Ipswich Bay, where considerable catches had been made for some time previous by about 15 boats.61 Nothing is known, definitely, as to their sea- sonal appearances and disappearances over the offshore banks. About all that is known of the movements of the large mature herring (in their fourth summer and older) is that they are encountered in num- bers only for the brief period before, during, and after the spawning season, when they are seen schooling at the surface, and are caught along shore. Fishermen report that they show about the off-lying islands some time before they make their way up the bays; two or three weeks earlier, for instance, at Grand Manan, Jonesport, and about Mount Desert Island than within Machias Bay. They are said to appear some time after the middle of July at Isle au Haut at the eastern entrance of Penobscot Bay, and at Castine within the Bay, though not until the end of that month » This happening was reported in the daily papers. 210941—53 8 or the first of August at Matinicus Island. Such of them as visit the Massachusetts Bay region are not expected there until the last week in Septem- ber. But they are in full force on all the spawning areas along the shores of the Gulf by October, from Grand Manan to Cape Cod ; they are equally widespread, if less abundant, inshore in November, and they are reported in December occasionally, and even later. It is probable that as the fish spawn out most of them move out promptly from the spawning grounds into deeper water, for fish recently spent are not often reported as taken in the weirs. Probably the spawners merely descend into deeper water to winter, as is the case in European waters. How deep the great body of them go is not known. But is has been proved that herring of all ages remain in the open Bay of Fundy throughout the cold season; also in the passages between the inner and outer divisions of Passama- quoddy Bay, even when water temperatures there are as low as 32° F.52 And the abundance of pelagic euphausiid shrimps (a favorite herring food) in the deeper water layers of the northeast corner of the Gulf suggests this as a rich wuntcr pasture for them. Studies carried out from the Atlantic Biological Station at St. Andrews, chiefly under Dr. A. G. Huntsman's S3 leadership, and by the International Passamaquoddy Fisheries Commission M during the early 1930's seem to us to have proved that the factor chiefly responsible for the great concen- tration of young herring in the Passamaquoddy region, and for their availability to the weir fishery there, is the differentia] circulation of the shoaler and deeper water layers that is set in motion by the inflow of fresh water from the tributary streams combined with superficial currents set up temporarily by the wind. In other words, the sardine-sized herring acts as does any planktonic animal such as the euphausiid shrimps and the copepod crustaceans, on which it feeds, as it swims to and fro, i. e., it drifts with the current. In technical language, it is "denatant." The case is not so clear for the larger herring, not because there is any reason to suppose they can direct their jourmys more intelligently, and because any directive swimming they may carry " Huntsman. James Johnstone Memorial Vol., 1934, p. S2. *' For summary, see Huntsman, James Johnstone Memorial Vol., 1934, pp. 95-96. «< See Graham, Jour. Biol. Board Canada, vol. 2, 1935, No. 2, pp. 95-140. 98 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE out is far more effective because more rapid; but because so little is known as to journeys any indi- vidual school actually makes as season follows season, whether of fat herring or of spawne.rs. Perhaps the most interesting question of all, and one as yet unsolved, is how and why the spawning fish seek their spawning grounds year after year, when their sex organs mature. Spawning grounds and season. — It appears that the most productive spawning ground for our Gulf formerly was and still is at the mouth of the Bay of Fundy, particularly on the shoals south- west of Grand Manan. The Trinity Ledges off western Nova Scotia are another important ground; and herring are reported as spawning commonly, though irregularly, in Machias Bay; about Jonesport; at Mount Desert; in French- mans Bay; among the islands at the mouth of Penobscot Bay (Swans, Isle au Haut, and Matin- icus); in Casco Bay; also about Wood Island a few miles south of Cape Elizabeth, which has long been known as the resort of breeding schools. Herring have also been found spawning off the beaches along the western shore of the Gulf, Ipswich Bay, for example; about Cape Ann; in Massachusetts Bay; about Provincetown; along outer Cape Cod ; in the Woods Hole region ; near No Mans Land; and about Block Island which is the southern breeding limit. But whatever spawning does take place either southward from the vicinity of Cape Elizabeth on the one hand, or in the inner parts of the Bay of Fundy on the other, is trifling as compared with the production along the eastern coast of Maine and in the Grand Manan region. Spawning takes place both along shore in our Gulf and on the various shoals and ledges that lie for 5 to 25 miles off the coast, a habit betrayed by the eggs that are found sticking to the anchor ropes of fishing vessels. But we find no definite record of herring spawning on Browns or Georges Banks, nor are young fry known there, a fact that was commented upon by Storer long ago. Spawning season. — Both spring - spawning schools and summer-fall spawning schools of herring were reported formerly, in the Bay of Fundy, the spring spawners visiting the south (Nova Scotia) side of the bay from Bier Island at the mouth in as far as Digby Gut, also the Parrsboro region on the New Brunswick shore near the head of the bay, spawning during April and May. But they seem never to have been very numerous, and it is not known whether any spawn now in the bay before summer. Spring- spawning as well as autumn-spawning herring have also been reported to us by fishermen along the west coast of Nova Scotia, though we have not been able to verify this. Other than this, spring spawners are neither recorded nor rumored anywhere in the Gulf of Maine. Around Grand Manan and in Machias Bay nearby, the heaviest runs of summer-autumn spawners usually come in July, August, and Sep- tember, the spawning season continuing until late in the fall in some years,55 but not commencing until early August, and ending by early Oc- tober in others.58 Passing westward we find the breeding period progressively later and shorter; mid-August for example until October around Petit Manan and near Mount Desert, while the few herring that spawn farther south do so chiefly during October in Ipswich57 and Massa- chusetts Bays; in late October and early Novem- ber in the vicinity of Woods Hole. So many observations have been taken in the Gulf from the vessels of the Bureau of Fisheries, and in the Bay of Fundy by the Biological Board of Canada, that it is possible to establish the temperatures rather closely at which herring spawn in our waters. Around Grand Manan and in the northern part of the Gulf generally, prac- tically all spawning is carried out in water of about 46-52° F. But such herring as spawn in the southern part of Massachusetts Bay and along the shores of Cape Cod, where autumnal cooling of the surface waters is not so rapid as it is farther north, may do so in slightly warmer water, say up to 53° or 55°. The Gulf of Maine herring spawn in rather low salinities (such characterize the coastal zone as a whole as compared with the North and Norwegian Seas), the most saline water in which it is known to spawn within our limits being not saltier than 33 per mille, the freshest probably about 31.9 per mille. They never spawn in brackish water within the limits of the Gulf, although known to do so at the mouths of certain European rivers in water that is nearly fresh. Destruction by natural causes. — The herring is a very "tender" fish, prone to wholesale destruction » So described by Moore, Rept. U. S. Comm. Fish. (1896), 1898, p. 408. » Dr. Huntsman informs us that this was the case in 1917. « Allen, Mem. Boston Soc. Nat. Hist., vol. 8, No. 2, 1916, p. 201. FISHES OF THE GULF OF MAINE 99 both by stranding on beaches during storms, and by pollution of the water. Many instances of this kind have been reported. Allen,68 for exam- ple, saw young herring in windrows for miles on the strand at Rye Beach, N. H., in August 1911. A slaughter of herring (still more instructive be- cause the exact course of events was followed) took place at Cohasset, on the south shore of Massa- chusetts Bay, in October 1920. On the 5th of that month a large school of "sperling," 4 to 5 inches in length, ran up the harbor (which is nearly landlocked), probably driven in by silver hake (at least so local fishermen said); were trapped there by the falling tide, and stranded on the mud. So numerous were they that the flats were entirely covered with them and it was esti- mated that 20,000 barrels of fish perished. Dur- ing the next few days the fish (alternately covered and uncovered by the tide) decayed, and despite the tidal circulation, so fouled the water that lobsters impounded in floating cars died. On the 10th there was a second smaller run of herring, and on the 15th a third run came as numerous as the first, the newcomers dying soon after they entered the harbor. Altogether, it was estimated that 50,000 barrels of fish perished, of which more than 90 percent were "sperling," 5 to 10 percent were large adults, and a few were small mackerel and silver hake, besides large numbers of smelt. The flats were silvery with herring scales at low tide by the last half of October, when we saw ihem, and the residents about the harbor found the stench almost unbearable. But the fish decom- posed and the water purified itself during the winter months. Mass destructions of young herring have also been reported in other Gulf of Maine harbors. Thus, Dr. Austin H. Clark reported that early in August 1925 the mud flats in Manchester Harbor, on the north side of Massachusetts Bay, were white with stranded herring 3 to 5 inches long, packed several deep at low tide along the sides of the little drains and hollows. Another such destruction took place in the same harbor in the summer of 1928. Vast quantities of herring spawn are likewise cast up on the beaches every year to perish in north European waters; this also happens to some extent in the Gulf of St. Lawrence. Numerical abundance and importance. — Moore « Mem. Boston Soo. Nat. Hist., vol. 8. No. 2, 1916, p. 202. (1898), who sifted many sources of information concluded (we believe rightly) that no general decrease had taken place in the abundance of young herring at the mouth of the Bay of Fundy up to that time. But it is common knowledge among fishermen that both the numbers visiting any given locality on our coast and the duration of their stay varies widely, not only from year to year, but over longer periods. Local spawning grounds, too, may be abandoned for a term of years — a common occurrence.59 The best documented case of local disappear- ances from a previously productive ground took place, as Dr. Huntsman writes us,60 from the shoals southwest of Grand Manan, whence large herring (previously very plentiful) withdrew in 1877, to reappear in 1881 on the Nova Scotia coast between Cape Sable and Digby. Dr. Huntsman has suggested that they had circled the Gulf offshore, for their exodus from the Grand Manan shoals was not accompanied by any coin- cident increase in the catch along the eastern part of the coast of Maine, but rather by the reverse.61 They persisted on the Nova Scotia shore until is'.li), when they gave out, probably from old age, for the large herring that remained in the Quoddy region also dwindled in numbers as shown by the collapse of the winter fishery there, evidence that this particular body of herring did not receive any significant recruitment after about 1880-1881. It remains to be seen whether large herring will ever reappear in their former plenty on the Grand Manan ground, as they did about 1857 hi Massa- chusetts Bay, where the stock had been at a low ebb since 1837; or whether the yearly drain on the population of young herring by the sardine fishery (well started by about 1880) is too great. The largest reported catch of herring for the Gulf as a whole for any year since 1928 for which statistics are readily available was 219,131,500 pounds taken in 1946, divided as follows: Massa- chusetts, 2,049,000 pounds; Maine, 80,107,400 pounds; and the Canadian shores of the Gulf, 136,975,100 pounds. The smallest catch was 70,519,886 pounds in 1932, divided 5,687,254 pounds, 31,988,132 pounds, and 32,844,500 pounds, » Moore, Kept. IT. S. Comm. Fish. (1890), 1898, p. 430. K Based on Canadian fishery statistics. «> Earll (Fisheries and Fish. Ind. U. S., sect. 6, vol. 1, 1887, pp. 423, 424) states that the fishery declined near Bois Bubert Island from 1875 to 1880, and that the catch was "considerably below average" at Matinicus during the 10 years previous to 1879. 100 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE respectively. In 1947, incidentally, the herring catch of the Gulf was topped only by the catches of haddock and of rosefish. It is not clear to what extent this range in the catch from year to year is due to fluctuations in the supply of fish; to differences in their availability; or to the sundry economic factors that enter in. What is certain is that with some 80 percent of the catch consisting of sardine-sized fish weighing only about one-half ounce, the toll taken cannot have been less than 1% billion fish in the poorest of recent years, 5 to 6 billion in the year when the yield was greatest, i. e., numbers far greater than that for any other Gulf of Maine fish. Come good year then or bad, Capt. John Smith's account of the herring of our Gulf thus applies equally well today: "The savages compare the store in the sea with the hair of their heads, and surely there are an incredible abundance upon this coast." 62 The sardine catch of the Bay of Fundy is made almost wholly in weirs, 347 of which were operated on the New Brunswick shore in 1947. On the Maine coast, as a whole, a little less than two- thirds the catch of herring, large and small, is made in weirs and in purse seines, combined, a little more than one-third nowadays in stop seines (about 44,500,000 pounds in 1947). These are used "to prevent the exit of the herring school from a cove or inlet. . . . The seine is stretched around the school with the ends of the net made fast to the shore." And stop seines are used mostly at night, when the presence of fish is betrayed by their luminous trails, if the water is firing, or by the « General Hlstorle of Virginia, New England, and the Summer Isles, 1616, reprinted In 1819 from London edition of 1629, p. 188. noise they make as they "flip" at the surface.63 And some are still caught in floating traps (about 2 million pounds in 1947) which we have often seen used in the harbors of Mount Desert. In 1947, seemingly a representative year, purse seines yielded the Maine fishermen nearly as much herring (about 36,100,000 pounds) as the weirs. Their presence is detected, Scattergood tells us, either by the firing of the water if by night, by echo sounding apparatus, or by the use of a thin wire suspended in the water, the vibrations of which indicate the presence of fish that strike it. In 1947 eleven purse seiners were active in the fall fishery for Maine herring. How many were engaged in the New Brunswick and Maine winter fishery is not known. Large catches of herring when on bottom also are made by special otter trawls in European wa- ters; and of the closely allied herring of British Columbia of late. But the possibility of develop- ing an otter-trawl fishery for herring in the Gulf of Maine has not yet been explored. Finally, we may remark that herring fresh from the water are among the most delicious of our fishes, especially the small sizes. Their only draw- back is that they do not keep well, being rich- meated and oily, and in the larger sized fish the many hair-like bones are troublesome. Hickory shad Pomolobus mediocris (Mitchill) 1815 Fall herring; Shad herring Jordan and Evermann, 1896-1900, p. 425. - See Scattergood, U. S. Fish and Wildlife Service, Sp. Sei. Ecpt. No. 67, 1949, p. 8, for further details. Figure 45. — Hickory shad (Pomolobus mediocris), Chesapeake Bay region specimen. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 101 Description. — The hickory shad differs rather noticeably from the sea herring in that the point of origin of its dorsal fin is considerably in front of the mid-length of its trunk; in its deep belly (a hickory shad 13'^ in. long is about 4 in. deep but a herring of that length is only 3 in. deep) ; in the fact that its outline tapers toward both snout and tail in side view (fig. 15); and in that its lower jaw projects farther beyond the upper when its mouth is closed; also, by the saw-toothed edge of its belly. Also, it lacks the cluster of teeth on the roof of the mouth that is characteristic of the herring. One is more likely to confuse a hickory shad with a shad or with the alewives, which it resembles in the position of its dorsal fin, in the great depth of its body, in its saw-toothed belly and in the lack of teeth on the roof of the mouth. But it is marked off from all of these by its projecting lower jaw. There is also a small difference in outline, its head tapering more to the snout, as seen in side view (fig. 45) . It has only about half as many gill rakers (19 to 21 on the lower limb of the first gill arch) as either the alewife or the blueback; and its upper jaw, reaching back only about as far as opposite the center of its eye, is shorter than that of the shad in which it reaches as far as the rear edge of the eye. Under favorable circumstances its color, also, is characteristic, for it is faintly marked on the sides with dusky longitudinal stripes, and the tip of its snout is dusky. Size. — This is the largest of our anadromous herrings next to the shad, growing to a length of 2 feet. A fish about 15 inches long weighs a pound, one of 18 inches, 2 pounds. Habits.— Nothing is known of the habits of the hickory shad in the sea to differentiate it from its close relatives of the herring tribe except that it is more of a fish eater. Launce, anchovies, dinners, herring, scup, silversides, and other small fish, squid, fish eggs, and even small crabs have been found in the stomachs of hickory shad at Woods Hole, as well as sundry pelagic Crustacea. It will strike a small spinner or other artifical lure, and it gives a good fight when hooked. In the southern parts of its range it is described as running up fresh streams, with the alewives in late winter and early spring to spawn.64 But it appears not to do so in the streams tributary to Chesapeake « Smith (N. C. Oeol. Econ. Surv; vol. 2, 1897, p. 121) describes it as doing so in the streams tributary to Pamlico Sound, N. C, where it is plentiful. Bay, though it is found in practically all of them. This opens the interesting possibilitj7 that the "green" fish found in Chesapeake Bay, leave the Bay, perhaps to spawn in salt water.65 General range. — Atlantic coast of North America from the Bay of Fundy to Florida. Occurrence in the Gulf oj Maine. — The hickory shad is a southern fish, with the Gulf of Maine as the extreme northern limit to its range. It is recorded in scientific literature only at North Truro; at Provincetown ; at Brewster; in Boston Harbor; off Portland; in Casco Bay; and from the mouth of the Bay of Fundy (Huntsman doubts this record), and it usually is so uncommon within our limits that we have seen none in the Gulf ourselves. But in 1932 anglers, trolling for striped bass and mackerel off the Merrimac River, met a run of hickory shad.66 It is much more plentiful west of Cape Cod, being common from spring throughout summer and early autumn at Woods Hole, where as many as 3,500 have been taken at a single lift of one trap. In 1919 the Massachusetts catch of hickory shad, practically all from the south coast, amounted to 12,800 pounds, and none are listed for Massa- chusetts for any subsequent year. Alewife Pomolobus pseudoharengus (Wilson) 1811 lapproximate date] Gaspereau; Sawbelly; Kyak; Branch herring; Fresh-water herring; Grayback Jordan and Evermann, 1896-1900, p. 426. Description. — The alewife is distinguishable at a glance from the sea herring by the greater depth of its body, which is three and one-third times as long as deep (an alewife of 13^ inches is about 4 inches deep; a herring that long has a depth of only 3 inches) also by the position of its dorsal fin, the point of origin of which is considerably nearer to the tip of the snout than to the point of origin of the central rays of the tail fin. Further- more, the alewife is much more heavily built forward than the herring, and the serrations on the midline of its belly are much stronger and sharper (hence the local name "sawbelly"), so much so that a practiced hand can separate «' Hildebrand and Schroeder, Bull., U.S. Bur. Fish., vol. 43, 1928, p. 84. w The Museum of Comparative Zoology received one from this run from Dr. J. C. Phillips, caught by him off the northern end of Plum Island, October 2, 1932. 102 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 46.— Alewife (Pomolobus pseudoharengus) , Chesapeake Bay region specimen. From Goode. Drawing by H. L. Todd. herring from alewives in the dark. The most useful distinctions between the alewife and the blueback are that in the former the eye is broader than the distance from its forward edge to the tip of its snout and the back grayish green, while in the latter the eye is only about as wide as the distance from front of eye to tip of snout, and the back is dark blue (p. 107). Also the lining of the abdominal cavity is pale grayish or pinkish white in the alewife, but is usually dusky or blackish in the blueback. But this distinction may not hold in all cases. Alewives are distinguishable from young shad by their smaller mouths with shorter upper jaws; also by the fact that the lower jaw of the alewife projects slightly beyond the upper when the mouth is closed, and by the outline of the edge of the lower jaw, the forward part of which is deeply concave in the alewife but nearly straight in the shad. The lack of teeth on the roof of the mouth distinguishes the alewife, with its brethren the hickory shad (p. 100) and blueback (p. 106) from the sea herring, anatomically. Color. — The alewife, like the herring, is grayish green above, darkest on the back, paler and silvery on sides and belly. Usually there is a dusky spot on either side just behind the margin of the gill cover (lacking in the herring) and the upper side may be faintly striped with dark longi- tudinal lines in large fish. The sides are iridescent in life, with shades of green and violet. The colors change, to some extent, in shade from darker to paler, or vice versa, to match the bottom below, as the fish run up stream in shallow water. Size. — The alewife grows to a length of about 15 inches, but adults average only about 10 to 11 inches long and about 8 to 9 ounces in weight; 16,400,000 fish taken in New England in 1898 weighed about 8,800,000 pounds. Habits. — The alewife, like the shad and the salmon makes its growth in the sea, but enters fresh water streams to spawn. This "anadro- mous" habit, as it is called, forced itself on the attention of the early settlers on our coasts. In the words of an eyewitness, "experience hath taught them at New Plymouth that in April there is a fish much like a herring that comes up into the small brooks to spawn, and when the water is not knee deep they will presse up through your hands, yea, thow you beat at them with cudgels, and in such abundance as is incredible." 67 And they are no less persevering in their struggles upstream today. Numbers of them are to be seen in many streams, any spring, alternately swimming ahead; resting in the eddy behind some irregularity of the bottom; then moving ahead again, between one's feet if one happens to be standing in midstream. And they are much more successful than the shad in surmounting fishways of suitable design. During the early runs some- times one sex predominates, sometimes the other, but the late runs consist chiefly of males, as a rule, and these are said to outnumber the females greatly on the spawning grounds. We have no firsthand observations to contribute on this score. Alewives are decidedly general in their choice • Capt. Charles Whltbome, In "The True Travels of Capt. John Smith," 1616, vol. 2, p. 250. FISHES OF THE GULF OF MAINE 103 of streams, running indifferently up rivers as large as the St. John, Merrimac and Potomac, or streams so small that one can almost leap across, and only a few inches deep. In large rivers they run far upstream — how far they may do so we do not know— or their journey may be one of only a few yards, as it is in the artificial cuts that are kept open through barrier beaches to allow the fish access to fresh water ponds behind the latter. The alewife spawns in ponds, including those back of barrier beaches (if there are openings to the sea, natural or artificial) and in sluggish stretches of streams, never in swift water, each female depositing from 60,000 to 100,000 eggs or more, according to her size.68 Spawning lasts only a few days for each group of fish. The spent fish run down stream again so soon after spawning that many of them pass others coming up, as we have often seen; fish on their return journey to salt water are familiar sights in every alewife stream. The adults, when entering streams to spawn, make the change from salt water to fresh within a short time without damage; this is equally true of the spent fish on their return to the estuaries. But Dr. Huntsman informs us that they appear unable to endure repeated changes between salt water and fresh, and that great numbers are killed in this way in the estuaries under certain conditions of tide. The strain of spawning leaves them very thin, but they recover rapidly after they reach salt water. We have seen spent alewives that had already put on considerable fat, taken from a trap at Provincetown as early in the season as July 16 (in 1915). Spawning ordinarily takes place at tempera- tures of about 55 to 60°. The eggs are about 0.05 inches in diameter, pink like those of the sea herring, and they stick to brush, stones, or any- thing else they may settle upon.89 Incubation occupies about 6 days at 60°. The young alewives, which are about 5 mm. long when hatched, growing to 15 mm. when a month old, soon begin to work their way downstream. They have been seen descending as early as June 15 in the more southerly of Gulf of Maine streams; «* The average number of eggs in 644 females taken In the Potomac was 102.800 (Smith, N. C. Qeol. and Eeon. Survey, vol. 2, 1907. p. 123). • The development of the eggs, larval stages, and young fry are described by Ryder (Report, U. S. Comm. of Fish. (1885), 1887, p. 505) and by Prince Contr. Canad. Diol. (1002-1005), 1907, p. 95). successive companies of fry move out of the pond and down with the current throughout the summer; and by autumn the young alewives have all found their way down to salt water when 2 to 4 inches long. We have seined young alewives as long as 4 to 4K inches (102-115 mm.) in salt water near Seguin Island, Maine, at the end of July, but others, only 3 to 3% inches long (78-92 mm.), near Mt. Desert Island as late as the first of October. Thenceforth the alewife lives in salt water until sexual maturity. Hildebrand and Schroeder70 found that little alewives in Chesapeake Bay had grown to about 4K to 5 inches long by the time they were 1 year old. The rate of growth of the older alewives, in salt water, has not been traced. But experiments in planting adult alewives in ponds in which there were none before, led, long ago, to the conclusion that they became sexually mature at 3 or 4 3rears of age, for none of their progeny returned until 3 or 4 years after the original plant. Specific instances, cited by Belding 71 are: (1) Three years after a large number of alewives were hatched in Keene's Pond, Maine, tributary to the Calais River, from a "plant" of mature fish, a run of adult fish entered Keene's Pond stream where none had ever been seen before ; this case was reported by the U. S. Bureau of Fisheries. (2) The establishment of a fishery, in the same way at Plymouth, Mass., in 4 years after restocking in 1865; and (3) G. M. Besse obtained results in 3 years in ponds in Wareham, Mass. The fact that alewives have been known to return, for spawning, to streams in which their parents had been planted, lends support to the "parent stream" theory; i. e., that alewives, like shad, tend to spawn in the stream system in which they were hatched. But a much more intensive study is needed of this interesting question before any categorical statement can be made, as to how generally this is true; and to what extent their return depends on their never having wandered far afield. Food. — The alewife is chiefly a plankton feeder like the herring; copepods, amphipods, shrimps, and appendicularians were the chief diet of speci- mens examined by Vinal Edwards and by Linton •• Bull. U. S. Bur. Fish., vol. 43, 1928, p. 91. " Rept. Alewife Fish. Mass., Mass. Dept. Conservation, Div. Fish, and Game, 1921, p. 18. 104 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE at Woods Hole. However, they also take small fish, such as herring, eels, launce, cunners, and their own species, as well as fish eggs. Unlike herring, alewives often contain diatoms even when adult. Alewives fast when they are running up- stream to spawn, but when the spent fish reach brackish water on their return they feed ravenously on the shrimp that abound in the tidal estuaries and which they can be seen pursuing. We have often hooked alewives on an artificial fly at such times. Movements at sea. — The alewife is as gregarious as the herring, fish of a size congregating in schools of thousands of individuals (we find record of 40,000 fish caught in one seine haul in Boston Harbor) and apparently a given school holds together during most of its sojourn in salt water. But they are sometimes caught mixed with men- haden, or with herring. Alewives, immature and adult, are often picked up in abundance in weirs here and there along the coast,72 and it is likely that the majority remain in the general vicnity of the fresh water influence of the stream-mouths and estuaries from which they have emerged, to judge from the success of attempts to strengthen or restore the runs of alewives in various streams, mentioned above. But it is certain that some of them wander far afield, for catches of up to 3,000 to 4,000 pounds per haul were made by otter trawlers some 80 miles offshore, off Emerald Bank, Nova Scotia (lat. about 43° 15' N., long, about 63° W.) at 60 to 80 fathoms, in March 1936.73 Odd alewives were reported from Georges Bank and the South Channel in March, June, August, and November of 1913. Some (up to 78 per haul) were trawled by Albatross III about 25 to 60 miles out off southern New England in May 1950; also 18 adults, 10 to 11 inches long, 70 odd miles off Barnegat, N. J., on March 5, 1931 ; and we saw 60 alewives trawled at the 25-fathom line off Marthas Vineyard 74 in late June, 1951 by the Eugene H. Where these wanderers come to shore to spawn, if they succeed in doing so at all, is an interesting question. It seems likely from various lines of evidence that alewives tend to keep near the surface for their first year or so in salt water, and while they " Huntsman (Contr. Canad. Biol., [1921) 1922, p. 58) reports it? young at Campobello Island, Bay of Fundy, in December and March. '» Reported by Vladykov, Copeia, 1936, No. 3, p. 168. One vessel brought in about 10,000 pounds. " At lat. 40° 58' N.; long. 70° 32' W. are inshore when older. But practically nothing is known as to the depths to which they may descend if (or when) they move offshore, there being no assurance that those taken by trawlers were not picked up, while the trawls were being lowered or hauled up again. General range. — Gulf of St. Lawrence and north- ern Nova Scotia south to North Carolina, running up into fresh water to spawn; landlocked races also exist in Lake Ontario, in the Finger Lakes of New York, and in certain other fresh-water lakes.78 Occurrence in the Gulf of Maine. — When the white man crossed the Atlantic probably there was no stream from Cape Sable to Cape Cod but saw its annual run of alewives unless they were barred by impassable falls near the mouth. And while its numbers have declined during the past two centuries and its range has been restricted, both by actual extirpation from certain streams by overfishing, by the pollution of the river waters by manufacturing wastes, and by the erection of dams that it cannot pass, the alewife is a familiar fish still, all along around our coast 76 and yields an abundant catch in many of our streams. Ale- wives are taken commonly about Yarmouth, Nova Scotia; in the Annapolis Basin; in Minas Channel; and farther still, up the Bay. Alewives still run in most of the streams tributary to the Bay of Fundy, many in the St. John. A few are taken in the weirs in Passamaquoddy Bay; while young ones have been taken around Campobello Island; as deep as 50 fathoms. They enter the large river systems all along the coasts of Maine and New Hampshire, likewise many small streams, the re- quirements being that these shaU lead to ponds or have deadwaters of sufficient extent along their courses, and no dams or falls that the alewives can not surmount. At Boothbay Harbor, for in- stance, a considerable number of alewives annually run, or did run, up to spawn in Campbell's Pond, a small body of water that is dammed off from the harbor, and reached by a fishway only 15 feet long. This is the shortest alewife stream of which we know. In 1896, when the alewife fishery was the sub- ject of inquiry by the Bureau of Fisheries,77 catches 75 Such a race has been reported in Cobbett Pond, Rockingham Co., N. H. by Kendall (Occ. Pap. Boston Soc. Nat. Hist., vol. 7, No. 8, 1908, p. 38) and by Bailey (Biological Survey Memmac Watershed, New Hampshire Fish and Game Dept., 1933, p. 162). " Belding (Rept. Alewife Fish. Massachusetts, Mass. Dept. Conserv., 1921) has given a very instructive report on the alewife in Massachusetts. " Smith, Rept. U. S. Comm. Fish. (1898) 1899, pp. 31-43. FISHES OF THE GULF OF MAINE 105 large enough to be worth special notice were re- ported from the mouths of the St. Croix, Dennys, Machias, Medomak, Penobscot, St. George, Pema- quid, Damariscotta, and Kennebec Rivers; from Casco Bay; and from sundry other shore localities in Maine; from the Piscataqua River system in New Hampshire; from the mouth of the Merrimac, and from Cape Cod Bay. Few alewives enter the Merrimac, now, so polluted is it, and so obstructed by dams.78 And Belding found them running in only about 9 or 10 streams on the Gulf of Maine coast of Massachusetts in 1920, out of 27 streams there that had formerly supported considerable alewife fisheries.79 At present, we learn from John B. Burns, of the Massachusetts Division of Marine Fisheries, only a few alewives manage to run up the Merrimac past the fish ladder at Lowell; there is a small but regular run in the Parker River; a few in the Ips- wich ; a good run in the Essex ; a few in the Saugus ; perhaps some in Weymouth Back River; M a small run in Wier River, Hingham (really a brook); a few in Bound Brook, Cohasset; a large run in Herring Brook, Pembroke (tributary to North River) yielding about 1,000 barrels yearly; in- creasing numbers in Jones River, Kingston, which had been restocked previous to 1938 when a fish ladder was installed; several thousand run yearly up Barnstable Mill Pond Brook; an improving run in Stony Brook, Brewster, where a ladder was built in 1945, and a good run in Herring River (really only a brook) in Wellfleet, Cape Cod. The first alewives ordinarily appear early in April in the few streams tributary to Massachu- setts Bay that they still frequent, and equally early (March or April) in the St. John River, New Brunswick, according to McKenzie;81 but their date of arrival varies considerably from stream to stream, according to local conditions. Thus few are seen in the streams of Maine until late April or early May; the first alewives appeared in 1915, for example, in Campbell's Creek, Booth Bay Harbor, on April 20. And the earliest good runs on the Nova Scotia shores of the open Gulf and of the Bay of Fundy may come as early as April (streams of Yarmouth, Annapolis, Hants, and Colchester *8 Fishways recently constructed now allow a few to ascend beyond Lowell' Massachusetts. »• See his report on the Alewife Fishery of Mass. (Mass. Dept. of Conser- vation, Div. Fish, and Game, 1921) which gives much information as to tho status of the alewife in Massachusetts streams. 80 Stocked with 28.000 adult fish in 1049, and fish ladders under construction. »' Kept. Biol. Board Canada (1931) 1932, p. 34. Counties), in May (Digby and King's County streams), or not until June (Cumberland County.82 Successive runs follow thereafter, all around the Gulf, until well into June, the later runs, going up, passing the earlier spawners coming down. In 1915, we saw this happening in Campbell's Creek, Boothbay, on May 20. And alewives have been seen, descending, as late as August 20, in Massa- chusetts streams. The extreme range of temperature within which eggs are spawned, in Gulf of Maine tributaries, is not known; probably the bulk of production takes place between about 55° and about 60°. Numerical abundance. — In 1896 S3 reported catches were 2,677,972 individual alewives (1,356,755 lb.) for Cape Cod Bay and for the Merrimac River combined; 526,500 (293,671 lb.) for New Hampshire streams; and 5,832,900 (3,388,326 lb.) from the rivers and streams and coast of Maine. The reported catch was 5,843,000 pounds 84 for the New Brunswick shore of the Bay of Fundy that year; 1,609,400 pounds for the Nova Scotia side and for the west coast of Nova Scotia, or about 10,510,000 and about 2,895,000 individual fish, respectively, assuming that the average weight was about the same as that for the alewives of Maine We thus arrive at a total catch for the Gulf of Maine of something like 22 million individual fish at that time and actually somewhat more, for the canvass certainly was not 100 percent complete. The run was much greater then in the St. John River system than in any other Gulf of Maine river and doubtless is still. The Damariscotta River, ranking second, was about one-third as productive as the St. John; the Merrimac, St. George, and Penobscot Rivers only something like one-tenth as productive each. Casco Bay yielded about one-sixth as many alewives as the St. John River, the shore line of Cape Cod Bay about one-fifth as many.85 And the catch of the St. John River system (including Kennebecasis Bay) still was about five times as great in 1931 as that for any of the other counties of New Bruns- « According to McKenzie, Rept. Biol. Board Canada (1931) 1932, p. 34. ■» A special study of the alewife fishery was made for that year, see Smith, Kept. V. S. Comm. of Fish. (1896) 1S99, pp. 33-43. 84 The Canadian catches for the year were reported in barrels; the conver- sion factor used is 200 pounds per barrel. " Reported catches for 1S96 wer.e about 4,234,000 pounds for the St. John River system; 1,390,612 pounds for the Damariscotta River, 3S5.804 pounds for the St. George River, 308,844 pounds for the Penobscot, 472,500 pounds for the Merrimac, 701,287 pounds for Casco Bay, and 8S1.255 pounds for Cape Cod Bay. 106 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE wick or of Nova Scotia that border on the Bay of Fundy or on the open Gulf.86 The alewife population of the Gulf is much smaller, today, than it was half a century ago. Thus the catch was only about one-half as great for the Bay of Fundy in 1945 and 1946 87 as it had been in 1896, and about one-third as great for Maine (1,224,600 lb.) while the Merrimac River, yielding 472,500 pounds in 1896, yielded less than 3,000 pounds in 1945.88 And though alewives may seem almost incredibly numerous when crowding into some stream, they made but a sparse population, even in their days of greatest plenty, when spread over the coastal waters of our Gulf, as compared to the sea herring. Importance. — Alewives are excellent food fish and they are marketed both fresh and salted, and are preferred by many to the sea herring. Thej- are good bait for cod, haddock, and pollock; and their scales commanded a high price for use in the manufacture of artificial pearls for a brief period during the first world war and for a few years afterward.89 By far the greater part of the catch of alewives is made in the lower reaches of the streams that they enter to spawn, in weirs, in dip nets or in haul seines according to locality. Most of those taken in outside waters (as in Casco and Cape Cod Bays) are either gill netted or are picked up in the fish traps. « MeKenzie, Kept. Biol. Board Canada (1931) 1932, p. 34. 87 5,051,100 pounds and 4,517,500 pounds, respectively. !s The reported catch for Essex County, Massachusetts, in that year was 2,700 pounds, only a part of which was from the region of the Merrimac. ** For details, see Report, Division of Fish and Game, Mass. (1920) J921, p. 140. Blueback Pomolobus aestivalis (Mitchill) 1815 Glut herring; Summer herring; Blackbelly; Kyack Jordan and Evermann, 1896-1900, p. 426. Description. — Bluebacks and alewives are diffi- cult to distinguish; experienced fishermen who recognize the existence of the two separate fish cannot always tell them apart, so closely do they resemble one another in general appearance. The most obvious external difference between them is that the back of the blueback is definitely blue green, that of the alewife gray green. But this applies only to fresh-caught fish; preserved speci- mens do not differ much in color, or fish that have been on ice for more than a short time. Another external difference is that the eye of the blueback is only about as broad as the distance from front of eye to tip of snout (or slightly broader), but is appreciably broader than that in the alewife; the blueback, too, with body about Z}{ times as long as deep, is a slightly more slender fish (on the average) than the alewife, and its fins are a little lower, but the two species probably intergrade in both these respects. The most dependable distinction between the two (though requiring the use of a knife) is that the lining of the belly cavity is sootj7 or blackish in the blueback, but pearl gray or pinkish gray in the alewife. We have yet to see a specimen that could not be named as the one or the other on this basis alone, unless so poorly preserved that the original shade of the cavity could no longer be determined. Figtjre 47. — Blueback (Pomolobus aestivalis), Chesapeake Bay region specimen. Todd. From Goode. Drawing by H. L. FISHES OF THE GULF OF MAINE 107 Color.- — Dark blue or bluish gray above, the sides and belly silvery, with coppery reflections at least in some waters; lining of the belly sooty or blackish. Size. — The blueback attains about the same size as the alewife, i. e., 15 inches; the adults average about 11 inches in length and about 7 ounces in weight. Habits.- — The blueback, like the alewife, makes its growth in salt water, but runs up into fresh water to spawn. And its breeding habits do not differ in any important particular from those of the alewife, except that it "runs" later in the season, does not run up as far above tidewater, and does not spawn until the water is much warmer, 70° to 75° instead of 55° to 60°.90 The eggs, about 1 mm. in diameter, sink like those of the alewife, and stick to anything they may chance to touch. Incubation occupies only about 50 hours at a temperature of 72°. The young are 30 to 50 mm. long within a month and already show most of the diagnostic characters of the adidt. Evidently they soon find their way down to the sea, for blue- backs of 50 mm. have been seined in abundance in Rhode Island waters late in July.91 Nothing whatever is known of their later rate of growth. The spent fish, return to sea shortly after spawning as do alewives. Practically nothing is known of their movements in the sea, except that they are schooling fishes. The fact, however, that 7 were trawled by the Albatross //on March 5, 1931 about 100 miles oil' Cape May, N. J., suggests that the blueback moves out from land and passes the cold season near the bottom. We need only note further that the blueback is as gregarious as the herring or alewife; that it is equally a plankton feeder, subsisting chiefly on copepods and pelagic shrimp, as well as on young launce and, no doubt, on other small fish fry. General range. — This is a more southern fish than the alewife, occurring along the American coast as far south as northern Florida; as far north as southern New England in abundance, perhaps less regularly in the Gulf of Maine though widespread « The early development and larval stages of the blueback are described by Kuntz and RadclifTe (Bull. U. S. Bur. Fish., vol. 35, 1918, pp. 87-134). 11 In Chesapeake Bay, nildebrand and Schroeder (Bull. U. S. Bur. Fish., vol. 43, 1928, p. 88) found that while most of the young bluebacks pass out to sea during the summer and fall, some remain in the deeper holes over the win- ter. By the following March when about a year old these are about 3M to 4 inches long; those in the sea may grow faster than this. there, and known definitely as far north as Cape Breton, Nova Scotia: 92 it spends most of its life in salt water but runs up into fresh water to spawn. Occurrence in the Gulf oj Maine. — Although fishermen have recognized the existence of two distinct species of alewives at least since 1816, it is difficult to arrive at a just idea of the status and migrations of this fish in our Gulf, because fish reported as "bluebacks" at sea sometimes turn out to be alewives, while the late runs of alewives are often referred to as "bluebacks." It seems, however, that schools of bluebacks are to be expected anywhere between Cape Sable and Cape Cod. Thus we have seen "gaspereau" fresh caught at Yarmouth, Nova Scotia, that appeared to be bluebacks.93 Huntsman had specimens from St. John Harbor and Shubenacadie River; they are reported, at least by name, from the St. Croix River; from Dennys River, Eastport; Bucksport; Casco Bay; Small Point; Freeport; and sundry other localities along the coast of Maine, as well as from the shores of Massachusetts Bay, including Cape Cod. L. W. Scattergood of the U. S. Fish and Wildlife Service has sent us about 40 typical bluebacks, about '■¥{ to 5t2 inches (92-124 mm.) long taken at Hodgdon Island, Sheepscot River, Maine, June 14, 1951 ; and we once saw thousands of fish taken from a trap near Gloucester, most of which we judged to be bluebacks from their color.94 A few fish were reported as "bluebacks" from Georges Bank during the investigation of 1913, and while there is no way, now, of checking whether these actually were bluebacks or alewives, the fact that we saw 10 bluebacks about 1 foot long, trawled by Albatross III at the 45 fathom line off southern New England, in mid-May, 1950, 95 shows that they may spread as far offshore as alewives. No definite information is at hand as to how regularly alewives run into our Gulf of Maine streams, for spawning; or what streams they enter at all. No distinction is made, commercially, on our coast between the blueback and the more abundant alewife; it is equally useful for bait and for food. u Dr. A. H. Leim has sent us four typical bluebacks about 12 inches lonf , taken at Cape Breton, Nova Scotia, in 1950. « We had no chance to examine them critically. » We did not then appreciate the desirability of positive identification. » Lat. 40° 06'; long. 71° 38' W. 108 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Shad Alosa sapidissima (Wilson) 1811 [Approxi- mate date] Jordan and Evermann, 1896-1900, p. 427. Description. — The shad is a typical member of the herring tribe in all respects with soft-rayed dorsal and anal fins of moderate size, the former situated above the ventrals and well forward of the middle of the body. It has a deeply forked tail and large scales that are loosened easily. Unlike the sea herring, the shad has no teeth on the roof of the mouth; adults have no teeth at aU, although young shad have small ones in the jaws which may persist until the fish is a foot or so long. The shad agrees with the hickory shad, alewife, and blueback, in its deep body and sharp saw-edged belly. But it differs rather noticeably from the hickory shad in its longer mouth, with upper jaw reaching to below the rear edge of the eye, and in the fact that the tip of its lower jaw is entirely enclosed within the tip of the upper when its mouth is closed. The most clear cut character distinguishing shad from alewife and blueback is that the upper outline of the shad's lower jaw is very slightly concave, without a sharp angle, the outline of theirs deeply concave with a pronounced angle. Furthermore the lining of the shad's belly is very pale. Color. — Dark bluish or greenish above, white and silvery low on sides and on belly, with a dusky spot close behind the rear edge of the gill cover, and usually with one or two longitudinal rows of indistinct dusky spots behind it. Size.— The shad is the largest of the herrings that regularly visit our Gulf, growing to a length of 2% feet. In the Bay of Fundy, according to Leim 96 shad weigh about % pound at 8 inches; about % pound at 12 inches; about 1% pounds at 15 to 16 inches; about 2% pounds at about 20 inches ; and about 4 K pounds at 23 to 24 inches, though with variations according to their condi- tion. Adult males weigh from 1% to 6 pounds; females from 2>}{ to 8 pounds. Shad are occasion- ally reported to 12 pounds, and the older writers mention shad of 14 poimds, but none so large has been credibly reported in the Gulf of late years. Habits. — The shad, like the alewife, spends most of its life at sea, and makes most of its growth there, but runs up into fresh rivers to spawn, the spent fish soon returning to salt water, and its fry soon running down also. During then- stay in the sea shad are schooling fish, often in thousands, and they never reenter fresh water until they return to spawn, though they sometimes do appear in brackish estuaries. Schools of shad are often seen at the surface in spring, summer, and autumn. In winter they disappear from sight. Probably the shad of the year winter near the mouths of their parent streams; the larger sizes somewhat farther out and deeper. The most direct evidence as to the depths to which they may descend is that shad have been trawled at about 50 fathoms off Nova Scotia in March (see footnote 22, p. 112), and at 26 to 68 fathoms off southern New England in May (footnote 23, p. 112). Food. — The shad, like other herrings, is pri- marily a plankton feeder. We have found shad taken in the Gulf of Maine in summer full of copepods (chiefly Calanus), and the stomach con- » Contrib. Canad. Biol.. N. Ser., vol. 2, 1924, p. 245, fig. 41. Figure 48.— Shad (Alosa sapidissima), Chesapeake Bay specimen. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 109 tents of fish from the Nova Scotia Coast of the Bay of Fundy examined by Willey 97 consisted chiefly of the copepod genera Arcatia and Temora with other smaller ones, of mysid shrimps and of the larval stages of barnacles; while Leim M found that the shad in the open Bay of Fundy feed chiefly on copepods and mysids. Shad are also known to feed as greedily on the pelagic euphausiid shrimps as herring do, on fish eggs, and even on bottom dwelling amphipods, showing that they forage near the ground at times. Occasionally they eat small fish, but these are only a minor item in their general diet.89 Shad, it appears, take little or no food just prior to spawn- ing. But they will often take an artificial fly, or a live minnow when running upstream to spawn.1 During the past few years, crowds of anglers have caught many on flies in the Connecticut River, and doubtless could in the few Gulf of Maine streams to which shad still repair (p. 110). Reproduction and growth.2 — The sexually mature fish enter the streams in spring or early summer when the river water has warmed to 50° to 55°. Consequently the shad run correspondingly later in the year passing from south to north along the coast, commencing in Georgia in January; in March in the waters tributary to Pamlico and Albemarle Sounds; in April in the Potomac; and in May and June in northern streams generally from the Delaware to Canada. In the Kennebec, according to Atkins,3 the first shad appear (or did) late in April, with the main run in May and June; the first ripe females are caught the last week in May and they begin to spawn about June 1, most of them doing so during that month, a few in July, and possibly an occasional fish as lato as August. Probably these dates applied equally to the Merri- mac in the good old days when shad were plentiful there, but the season is somewhat later in the St. John, also in the Shubenacadie as might be ex- pected; i.e., from mid-May until the end of June.4 » Contrib. Canad. Biol., N. Ser., vol. 1, 1923, p. 316. "Contrib. Canad. Biol.. N. Ser., vol. 2. 1921, p. 231. '• Leidy (Proc. Acad. Nat. Sci. Philadelphia, Ser., 2, 1868, p. 228) found 30 small sand eels (Ammodytes) in the stomach of a shad, probably caught in Delaware Bay. i Bean (Bull. 60, Zool., vol. 9, New York State Mus., 1903, p. 207) com- mented on this long ago. i Accounts of the breeding habits of tho shad have been given by Ryder, Eept. U. S. Comm. Fish. (1885) 1887, p. 523; by Prince (Supp. 6, Rept. Dept. Marine Fish. Canada, Fish. Branch. 1907, pp. 95-110; in the Manual of Fish Culture, published by the U. S. Bur. of Fish., 1887; and more recently by Leim (Contrib. Canadian Biol. N. Ser. vol. 2, 1924, pp. 184-202). > Fish. Ind. U. S., Sect. 5, vol. 1, 1887, pp. 683-684. < Leim, Contrib. Canad. Biol., N. Ser., vol. 2, No. 11, 1924, p. 182. In large rivers they run far upstream. In the St. John River, New Brunswick, they ascend about 200 miles to the grand falls even today according to Leim, and they still run up 300 miles (or did recently) in the Altamaha in Georgia; for 375 miles in the St. Johns River, Florida. But they could run up only about 35 miles at present in the Penobscot, where they formerly ascended some 90 miles, or 44 miles (to Augusta) in the Kennebec, which they formerly ascended 108 miles (to Carratunk Falls), though none enter either of these rivers now, so far as we know. And the dams at Lawrence, only 20-odd miles up- stream, now stop any stray shad that may still enter the Merrimac, which they formerly as- cended for 125 miles to Lake "Winnepesaukee.8 In the Shubenacadie, shad spawn mostly in temperatures higher than about 54°, and spawning is interrupted if the water chills below that, temporarily. The fish select sandy or pebbly shallows for spawning grounds, and deposit their eggs mostly between sundown and midnight. Females pro- duce about 30,000 eggs on the average, though as many as 156,000 have been estimated in very large fish. The spent fish, now very emaciated, begin their return journey to the sea immediately after spawning. In the Kennebec they were first seen on their way down about June 20 and constantly thereafter throughout July; in the St. John spent fish are running down in July and August. Ac- cording to Atkins they begin feeding before reach- ing salt water and recover a good deal of fat before moving out to sea. The eggs are transparent, pale pink or amber, and being semi-buoyant and not sticky like those of other river herrings they roll about on the bottom with the current. The eggs hatch in 12 to 15 days at 52° (12° C), in 6 to 8 days at 63° (17° C), which covers the range characteristic of Maine and Bay of Fundy rivers during the season of incubation. And Leim has made the interesting discovery that larval development is more successful in brackish than in pure fresh water, with about 7.5 parts of salt per thousand as about the most favorable salinity. The larvae are about 9 to 10 mm. long at the time of hatching, growing to about 20 mm., at 21 • Stevenson (Rept. U. S. Comm. Fish., (1898) 1899, p. Ill) has given a table of the distances to which shad ascended various rivers then, and formerly from the Penobscot in Maine to the St. Johns in Florida. 110 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE to 28 days. Shad larvae resemble alewife larvae, being extremely slender with the vent almost as far back as the base of the tail.6 The young shad remain in the rivers until fall, when they move down to salt water; they are now 1% to 4^ inches long, resembling their parents in appearance. According to Leim's investigation, based on scale studies and length frequencies, shad in the upper Bay of Fundy, average about 5 to 6 inches long when one year old; 9 to 10 inches long at 2 years; 13 to 14 inches at 3 years; 15 to 16 inches at 4 years; and 18 to 19 inches at 5 years. The two largest he examined, about 24 72 inches (62 cm. and 63 cm.) long, appeared to be 7 and 6 years old, respectively. They may grow somewhat faster in the open Gulf of Maine, to judge from the greater abundance of pelagic crustaceans on which they feed (p. 109). Most of the spawning fish are 5 years old in the Shubenacadie, and presumably in other Gulf of Maine rivers; the oldest 8 or 9 years old. General range. — Atlantic coast of North America from the southeastern coast of Newfoundland,7 which shad have been known to reach as strays, and the estuary of the St. Lawrence River, where there is a considerable population of them,8 to the St. Johns River in Florida; also represented in the Gulf of Mexico by a closely related species. The shad has been successfully introduced on the Pacific coast of the United States. It runs up rivers into fresh water to spawn. Occurrence in the Gulj of Maine. — When the first settlers arrived in New England they found seemingly inexhaustible multitudes of shad annu- ally running up all the larger rivers and many of the smaller streams, with the tributaries of the Gulf of Maine hardly less productive than the Hudson or Delaware. But one stream after another was rendered impassable by the construc- tion of dams near the mouth, for shad cannot or will not run up through fishways that are readily used by alewives. Indeed, they have been practically wiped out in the Merrimac River, as appears from the following compilation : 9 • Leim (Contr. Canad. Biol., N. Ser., vol. 2, No. 11, 1924, p. 196) gives a detailed comparison of shad with alewife larvae. 7 The most northerly record of a shad, on which we have chanced, is one taken in Bull's Bay, near St. Johns, Newfoundland. « See Vladykov (Contr. Dept. Fish., Quebec, No. 30, 1950, pp. 121-135, and Natural. Canad., vol. 77, 1950, pp. 121-135) for a study of the movements of the shad in the St. Lawrence estuary. • Fr<-m Stevenson, Rept. U. S. Comm. Fish. (1898) 1899, p. 262. Number of shad caught, reported, or estimated Year Number of shad caught, reported, or estimated 1789 830,000 1888 1805 540,000 1889 1835 365,000 1890-1892. 1865 50,000 1893 1871-1873 (aver- 1894 age) 1,942 1895 1880 2,139 1896 1885 130 None 18 None 2,020 2,750 94 7 The Gulf of Maine rivers to which shad are known to resort regularly today are the Annapolis, Petit Codiac, Shubenecadie, and St. John, tribu- tary to the Bay of Fimdy; perhaps the St. Croix; ,0 the only Maine rivers that see regular runs of a few shad are the Nonesuch and the Sheepscot.11 A few shad may enter other Gulf of Maine streams in some years if not yearly, and bright spots in the shad picture are that a considerable number of adult shad ran up the South River in Marshfield, Massachusetts, on the southern shore of Massachusetts Bay in 1950, and that there has been a run of something like 2,000 shad yearly in Mill Creek, Sandwich, Mass., for the past four years. 12 How successfully they may have spawned in either of these streams is not known. It appears that most of the shad hatched in the rivers tributary to the Bay of Fundy, and the spent fish from there, remain in or near the estu- aries where they take to salt water; and that most of the adults that survive the strain of spawning return to the parent stream to spawn again. Thus it is only in St. Marys Bay, in Annapolis Basin, in Cobequid Bay and Minas Basin, in Chignecto Bay and at the mouth of the St. John as well as for a few miles westward, that large Fundian shad are caught in any numbers.13 The fact, on which Leim I3a comments that "there is not a single record of a shad ever having been taken" at Grand Manan island, although this "lies almost directly in the path of any body of >o The St. Croix once had a large run of shad. None were seen there for 8 or 9 years prior to 1915, but they wero there in some numbers in 1915 and 1916, according to investigations by H. F. Taylor of the V. S. Bureau of Fisheries; their present status there is not known. They have been entirely extirpated from the Saco, where they were abundant formerly, probably from the Penobscot and Kennebec, and certainly from the Merrimac, as noted above. " Information from Dr. C. E. Atkinson, U. S. Fish and Wildlife Service. " Reported to us by John B. Bums of the Massachusetts Division of Fisheries and Game. U Leim (Contr. Canad. Biol. N. Ser., vol. 2, No. 11, 1924, fig. 2) gives a chart showing the location of shad catches for the Bay of Fundy. "» Contr. Canad. Biol. N. Ser., vol. 2, No. 11. 1924. p. 173. FISHES OF THE GULF OF MAINE 111 fish going in or out of the Bay of Fundy," is especially significant as emphasizing the localiza- tion of the St. John shad near the parent river. The behavior of the St. John river shad raises an interesting question, as to the source of the young fish that sometimes congregate in the Bays and among the islands along the coast of Maine (Casco Bay especially), for there seem to be too many of them, in some years, to be credited to the small runs that still frequent the rivers of Maine (unless runs may have been overlooked of late in other rivers there). Immature shad, up to 2 to 2% pounds in weight are observed more or less commonly in Cape Cod Bay near Provincetown in summer or autumn and in the inner parts of Massachusetts Bay (some- times taken in the traps at Beverly or Manchester), and off Cape Ann.14 Spent shad up to 10 pounds in weight (averaging about 5 pounds), are some- times reported by fishermen off the coast of Maine west of Penobscot Bay; near the Isles of Shoals; off York Beach, and off Cape Ann, in summer, autumn, and even in December.15 The few mature shad with ripening sexual organs that are picked up by the haddock netters between Cape Ann and Portland in April and May, most often about the Isles of Shoals and Boon Island,18 probably are headed for the rivers of Maine. Larger numbers of fish are seined in September and October, in the neighborhood of Mount Desert Island, where they have been the object of a frozen fish industry in some years.17 These, like the green fish mentioned above, seem far too numerous to be accounted for by the small production that still takes place in the rivers of Maine. Some few of them, it seems, are Bay of Fundy fish, for one of a batch tagged near Mount Desert Rock in August 1947, was recaptured in Kings County, New Brunswick (St. John River system) the following June, and a second in the Petitcodiac River that July, while a third, tagged farther west on the coast of Maine in August or September 1948 was " 502 barrels (about 100,400 lb.) were taken in one set of mackerel pounds at Provincetown in June 1910; the traps picked up numbers of shad of about 14 Inches from June 20 to July 6, 1921, at Magnolia and Beverly, where the catch was 10,300 pounds In 1945; and 14 shad 11 to 1554 Inches long were taken In one set of traps at Barnstable, on Cape Cod Bay, October 3, 1950. " 135,000 pounds of those large spent fish were caught near Gloucester In the autumn of 1915; 125 barrels of 2- to 5-pound shad, some spent, near Seguln Island, July 19, 1925. " A series of shad from that region, examined by the late W. W. Welsh In April and May 1913, averaged 5 pounds, all with well-developed sex organs. " About 250,000 pounds were brought In to the local freezers yearly in 1913, 1814. and 1915. recaptured in the St. John River in May 1950. But it seems established that most of the medium- sized shad and larger now found in our Gulf are immigrants from the south, growing and fattening on the rich supply of plankton they find there, but returning to the rivers west and south of Cape Cod to spawn. Direct evidence of this is that one tagged in Chesapeake Bay was recaught at Race Point, at the tip of Cape Cod, 39 days later; 18 one also was recaptured near Gloucester and another near Portland that had been tagged in the Hudson River, while 3 out of 1,380 tagged in New York Bay were recaptured in the Bay of Fund}' after 37 days, 75 days, and 85 days, respectively, and one tagged off Fire Island, N. Y., was recaught at St. John. New Brunswick, after 39 days.19 On the other hand, 18 shad, from a batch of 236 that were tagged ne;ir Mount Desert Rock in August 1947 were recaptured the next spring scattered along in different stream systems from the Connecticut to the Altamaha in Georgia. Others, from this same batch, were recaptured in the Connecticut, in the Hudson, on the coast of New Jersey, and in the Pamlico River, X. C, during the next two springs. And three others, from a batch of 431 tagged farther west along the coast of Maine in the sum- mer and autumn of 1948, were recaptured in the Hudson River; three in Chesapeake Bay, and one in I lie Pamlico River, N. C.20 The shad that take part in this intermigration must winter somewhere between their northern feeding grounds whence they have vanished wholly by mid-autumn, and their southern breed- ing streams near which they do not appear until spring. But it is not yet known where they pass the cold months, how deep down they go, bow far offshore, or how active they are then. Still other shad are known to make very long journeys that can hardly be fitted into any regular migratory pattern, and from which they may never find their way back. Thus one that was tagged in the lower St. Lawrence River was recaught on Brown's Bank 258 days later; a second, from that same batch, was recaught in Cumberland basin, near Amherst, Nova Scotia, at the head of the Bay of Fundy after 322 days; a third at Province- " Vladykov, Trans. Amer. Fish. Soc, vol. 67, 1938. p. 64. '■ Information supplied by C. E. Atkinson, U. S. Fish and Wildlife Service. "Information supplied by E. H. Hollls of the U. S. Fish and Wildlife Service. 112 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE town at the tip of Cape Cod, some 1,200 miles away from where it had been tagged 444 days previous.21 And one, from a batch of weir-caught fish tagged on the coast of Maine, August- September, 1948, was recaught in the Medway River, outer coast of Nova Scotia, a second, in the Miramichi River, tributary to the southern side of the Gulf of St. Lawrence in 1950, and a third, off Tor Bay, eastern Nova Scotia in 1951. To what extent the seasonal journeys of the sbad are passive with the dominant circulatory move- ments of the water, and to what extent (if any) they are self-directed is perhaps the most interest- ing question that now faces us in our studies of the shad of the Gulf of Maine. Shad have been trawled 50 to 60 miles out, off eastern Nova Scotia;22 have often been reported 40 to 50 miles out off the coast of Maine; also 25 to 90 miles out, off southern New England,23 and we saw one trawled by the Eugene H in late June, 1951, on the southern part of Georges Bank (lat. 40°52'N., long. 67°40'W.), about 110 miles from the nearest land. Evidently they may wander as far offshore as alewives do; perhaps even as far as herring. Shad reared in different regions may, perhaps, prove to differ enough hi racial characters for recognition when taken at sea, but this is a ques- tion for the future.24 Abundance. — The stock of shad in the Gulf is but a shadow in comparison with that of colonial days. In 1896, the only year for which detailed information is available as to the numbers taken in different streams, 290,122 shad were reported as caught in the Kennebec system, 9,000 in the Pleasant River, about 3,000 in the Harrington River, only 114 in the Penobscot and 12 in the St. Croix; 100 in the Piscataqua and 7 in the Merri- ll See Vladykov, Nat. C3nad., vol. 77, 1950, p. 121, for a detailed account of his tagging experiments on St. Lawrence River shad. 22 Vladykov, Copeia, 1930, No. 2, p. 108, reports between 25 and 30 shad of 4-0 pounds, taken per haul, by otter trawlers in March, 1935, southwest of Middle Ground, about lat. 44c25' N., long. 01°05' W., at about 50 fathoms. 22 Two shad were trawled by Albatross III on the eastern part of Nantucket Shoals at 08 fathoms, and 40 others at 9 stations distributed thence westward to the offing to Montauk Point (long. 71°52' W.) at 20-04 fathoms, May 11-18, 1950. 2< Vladykov and Wallace (Trans. Amer. Fish. Soc, vol. 07, 1937-1938, pp. 52-00) believe that Shubenacadie, Delaware River and Chesapeake Bay shad differ significantly in average number of vertebrae, of mid ventral scales and of pectoral fin rays. But Warfel and Olsen (Copeia, 1947, pp. 177-183) doubt whether any distinction can be drawn between shad in various streams along our North Atlantic coast, at least as far as average number of vertebrae goes. mac.25 In that same year the catch was about 1,059,000 pounds for the Nova Scotia shore of the open Gulf and for the Bay of Fundy;26 1,404,477 pounds for the rivers and coast of Maine; about 122,932 pounds (32,782 fish) for the Gulf of Maine coast of Massachusetts, or a total of about 2,586,400 pounds for the Gulf as a whole. With shad averaging about 3% pounds in weight,27 this corresponds to about 690,000 fish. But the yearly catch was only about one-third as great for the period 1916-1919 as it had been in 1896, whether for the United States shores of the Gulf or for the Canadian.28 And it was of about that same order of magnitude in 1931, i. e., 677,540 pounds for the Gulf as a whole (157,763 pounds for Maine, 147,277 pounds for Massachusetts, 237,200 pounds for the Bay of Fundy and West Nova Scotia region). Since that time, the catches have ranged between 10,400 pounds and 306,000 pounds for the Massa- chusetts coast of the Gulf and between 9,300 pounds and 1,106,800 pounds for Maine, a fluctuation so extreme (no regional correlation appearing) as to suggest that market conditions were the chief governing factor. On the other hand the catches for the Canadian shores of the Gulf increased rather consistently from 1931 to a total of 1,287,600 pounds in 1939 then declined to around 780,000 pounds for 1944 and 1946, a rise and fall regular enough to suggest a corre- sponding fluctuation in the actual abundance of the shad. The average yearly catch for the period 1944-1946 combined, was about 20,000 pounds for Massachusetts, about 224,050 pounds for Maine, and about 780,000 pounds for the Bay of Fundy and western Nova Scotia. Thread herring Opisthonema oglinum (LeSueur) 1817 Jordan and Evermann, 1896-1900, p. 432. Description. — The thread herring is distinguish- able at a glance from all the herrings that regularly inhabit the Gulf of Maine by the prolonged last ray (usually about as long as the body is deep) of its dorsal fin. It resembles the gizzard shad of 2» Stevenson, Rept. U. S. Comm. Fish (1898) 1899 pp. 205-209. 2' These catches were reported as "barrels" presumably of 200 pounds each. 2i Stevenson, Rept. U. S. Comm. Fish (1898) 1899, p. 121. 2» About 400,000 pounds for the United States coast, of the Gulf and about 374,000 pounds for the Bay of Fundy and in western Nova Scotia combined in 1910-17. FISHES OF THE GULF OF MAINE 113 Figure 49. — Thread herring (Opisthonema oglinum). Drawing by Louella E. Cable. fresh and brackish waters farther south in this respect, but the two differ rather conspicuously in various details. In the thread herring, the upper edge of the tail fin is about 1% times as long as the head (only about as long as the head in the gizzard shad); the point of origin of the dorsal fin is a little in front of the origin of the ventral fins (a little behind in the gizzard shad); the distance from the origin of the ventrals to the origin of the anal fin is at least 1% times as long as the base of the anal (only about % to % in the gizzard shad); and the anal fin is very low, with its first few rays a little shorter than the eye (about 1% times as long as the eye in the "gizzard"). There is no danger of confusing a thread herring with a young tarpon with which it shares the prolonged dorsal ray, for its dorsal fin originates in front of the ventrals, while the two fish are far apart in general appearance. This is a rather thin fish, its body about 2% to 3 times as long (to the base of the tail) as deep ; the belly is sharp and saw edged ; the tail deeply forked as in our other herrings. There are 18 to 19 rays in the dorsal fin, 22 to 24 in the anal. Color. — Bluish above, silvery on sides and belly. The scales along the back have dark centers, form- ing longitudinal streaks, and there is a faint dark spot just behind the upper margin of the gill cover; the dorsal and caudal fins have black tips. Size. — Maximum length about 12 inches. General range. — Atlantic coast of America in tropical and subtropical latitudes, south to Brazil, straying northward to Chesapeake Bay, and occasionally as far as southern Massachusetts. Occurrence in the Gulf of Maine. — A thread herring is caught off southern New England occasionally; they were even reported as rather common in Buzzards Bay and in Vineyard Sound during the summer of 1SS5. But there is only one record of it within the Gulf of Maine, a single specimen 7 inches long, taken off Monomoy Point, at the southern angle of Cape Cod, in August 1931.25 Being a tropical fish, it is not apt to reach the Gulf except as the rarest of strays. Menhaden Brevoortia tyrannus (Latrobe) 1802 Pogy; Mossbunker; Fat back Jordan and Evermann, 1896-1900, p. 433. Description. — This fish is universally called "pogy" in the Gulf of Maine but no less than 30 common names are in use south of Cape Cod. It is flattened sidewise like all our other herrings, has a sharp-edged belly, and is as deep proportion- ally as the shad (body about 3 times as deep as long), though the general form is altered when the fish are fat. The very large scaleless head, which occupies nearly one-third of the total length of the body, gives the menhaden an appearance so dis- tinctive that it is not apt to be mistaken for any other Gulf of Maine fish. It is likewise distin- guishable from all its local relatives by the fact that the rear margins of the scales are nearly vertical (not rounded), and are edged with long comblike teeth instead of being smooth. The dorsal fin originates over the ventrals or very slightly behind them. We need only point out further that the pogy is toothless, its tail deeply forked, its ventral fins very small, its dorsal and anal of moderate size, its mouth large and gaping back as far as the hind margin of the eye, and that the tip of its lower jaw projects beyond the upper. » Reported by Mactoy, Bull. Boston Soc. Nat. Hist., No. 61, 1931, p. 21. 114 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE \ ^ ^^^| Figure 50. — Menhaden {Brevoortia tyrannus). A, egg; B larva, newly hatched, 4.5 mm.; C, larva, 23 mm.; D young fry, 33 mm. A-D, after Kuntz, and Radcliffe. Color. — Dark blue, green, blue gray, or blue brown above, with silvery sides, belly, and fins, and with a strong yellow or brassy luster. There is a conspicuous dusky spot on each side close behind the gill opening, with a varying number of smaller dark spots farther back, arranged in irregular rows. Size. — Adidt menhaden average 12 to 15 inches in length, and from two-thirds to one pound in weight. One 18 inches long was taken at Woods Hole in 1876, and a fish 20 inches long has been reported. The heaviest of which we have heard was one of 1 pound 13 ounces, taken at Orient, N. Y. Habits. — The menhaden, like the herring, almost invariably travels in schools of hundreds or thou- sands of individuals, swimming closely side by side and tier above tier. In calm weather they often come to the surface where their identity can be recognized by the ripple they make, for pogies, like herring, make a much more compact disturbance than mackerel do, and "a much bluer and heavier commotion than herring, which hardly make more of a ripple than does a light breeze passing over the water," as W. F. Clapp has stated to us. Also, pogies as they feed frequently lift their snouts out of water, which we have never seen herring do, while they break the water with their dorsal fins, also with their tails. And the brassy hue of their sides catches the eye (as we have often seen), if one rows close to a school in calm weather. It is chiefly on warm, still, sunny days that the menhaden come to the surface, sinking in bad weather; and they are said to come up more often on the flood tide than on the ebb. It is also said (this we cannot vouch for) that the fish work inshore on the flood tide and offshore on the ebb. Food. — The menhaden, formerly thought to subsist on mud, is now known to feed chiefly on microscopic plants (particularly diatoms) and on the smallest Crustacea.30 It sifts these out of the water with a straining apparatus in the shape of successive layers of comb-like gill rakers as efficient as our finest tow nets. No other Gulf of Maine fish has a filtering apparatus comparable to that of the pogy, nor has it any rival in the *• For a detailed account of the food and of the branchial sieve of the men- haden, see Peck (Bull., U. S. Fish Comm., vol. 13, 1894, pp. 113-124, pis. 1-8. FISHES OF THE GULF OF MAINE 115 Gulf in its utilization of the planktonic vegetable pasture. Menhaden feed, as Peck described, by swimming with the mouth open and the gill openings spread. We have often seen specimens in the aquarium at Woods Hole doing this.31 And we have watched small ones in Chesapeake Bay, swimming downward as they feed, then turning upward, to break the surface with their snouts, still with open mouths. The mouth and pharyngeal sieve act exactly as a tow net, retaining whatever is large enough to enmesh, with no voluntary selection of particu- lar plankton units. The prey thus captured (as appears from the stomach contents) includes small annelid worms, various minute Crustacea, schizo- pod and decapod larvae, and rotifers, but these are greatly outnumbered as a rule by the sundry unicellular plants, particularly by diatoms and by peridinians. And the food eaten at a given locality parallels the general plankton content of the water, except that none of the larger animals appear in the stomachs of the fish on the one hand, nor the very smallest organisms (infusoria, and certain others such as the coccolithophorids) on the other. The menhaden, in short, parallels the whalebone whales, the basking shark, and the giant devil rays in its mode of feeding, except that its diet is finer because its filter is closer meshed. Peck has calculated from observations on the living fish that an adult menhaden is capable of filtering between 6 and 7 gallons (about 24 to 28 liters) of water per minute, and while the fish do not feed continuously this will give some measure of the tremendous amount of water sifted and of plankton required to maintain the hordes in whic'.i these fish congregate. The abundance of microscopic plants in the water of bays and estuaries, and along the coast has often been invoked to explain the concentration of menhaden close to shore. Enemies. — No wonder the fat oily menhaden, swimming in schools of closely ranked individuals, helpless to protect itself, is the prey of every pre- daceous animal. Whales and porpoises devour them in large numbers; sharks are often seen fol- lowing the pogy schools; pollock, cod, silver hake, and swordfish all take their toll in the Gulf of 31 Apparently Ehrenbaum (as quoted by Bullen, Jour., Mar. Biol. Assoc. United Kingdom, vol. 9, 1910-13, pp. 394-403) was not acquainted with the habits of menhaden when he wrote to the effect that no fish eat plankton indiscriminately, or swim about habitually with open mouth when feeding. Maine, as do weakfish south of Cape Cod. Tuna also kill great numbers. But the worst enemy of all is the bluefish, and this is true even in the Gulf of Maine during periods when both bluefish and menhaden are plentiful there (p. 384). Not only do these pirates devour millions of menhaden every summer, but they kill far more than they eat. Besides the toll taken by these natural enemies, menhaden often strand in myriads in shoal water, either in their attempt to escape then enemies or for other reasons, to perish and pollute the air for weeks with the stench of their decaying carcasses. Breeding and growth. — Very little is known about the breeding habits of the menhaden, except that it spawns at sea and that the chief production of eggs takes place south of our limits. According to observations at Woods Hole,32 the main body of the fish off southern New England spawn in June, continuing through July and August; even into October as in 1915, when the Grampus collected eggs and larvae in Nantucket Sound and westward from Martha's Vineyard in that month. And re- ports of spent fish in the Gulf of Maine in July and August, with others approaching maturity, BUggesI that the menhaden is a summer spawner there also. We have found no eggs in our tow- nel tings north of Cape Cod (young fry were taken in abundance in Casco Bay in October 1900), prob- jil.lv because our work there was carried on during a series of poor menhaden seasons. From Chesa- peake Hay southward the spawning season appears to be late in the autumn, and in early winter. Menhaden eggs are buoyant and resemble those of the European pilchard (Clupea pUchardus), bul are easily distinguished from the eggs of any other Gulf of Maine fish by their large size (1.5 to 1.8 mm. in diameter), broad perivitelline space, small oil globule (0.15 to 0.17 mm.), and very long em- bryo. Incubation is rapid (less than 48 hours), as Welsh found by experiment. The newly hatched larvae are 4.5 mm. in length, growing to 5.7 mm. in 4 days after hatching. The dorsal and caudal fins first become visible at a length of 9 mm.; at 23 mm. all the fins are well developed; scales are present at 33 mm.; and at 41 mm. the fry show most of the characters of the adult, except that their eyes are much larger, proportionately. The youngest larvae much resemble young herring, but the fins are formed, the tail becomes forked, and u By Kuntz and Radcliile. Bull. U. S. Bur. Fish., vol. 35, 1918, p. 119, who describe the eggs and larvae. 116 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE the body deepens at a much smaller size, a men- haden of 20 mm. being as far advanced in develop- ment as a herring of 35 mm., which makes it easy to distinguish the older larvae of the two fish. Welsh concluded from examination of great numbers of fry and from measurements and scale studies of fish of various ages that menhaden hatched in summer (which would apply to any fry that might be produced in the Gulf of Maine) are 2% to 3% inches (6 to 8 cm.) long by their first winter; and average about 6% inches (16 cm.) by their second winter; fall-hatched fish are 1% inches (3 cm.) and about 5 inches (about 13 cm.) long, in their first and second winters, with every gradation between the two depending on the precise season when the fish are spawned.33 Apparently sexual maturity is attained in the season following the third winter, and a few of the older fish that Welsh examined showed as many as 9 to 10 winter wings on their scales. General range. — Coastal waters along the At- lantic coast of America from Nova Scotia to eastern Florida; represented in the Gulf of Mexico, and southward to northern Argentina, by a series of named forms that differ from our northern menhaden in ways that would not be apparent to any one but to a trained student of fishes.3* Occurrence in the Oulj of Maine. — The Gulf of Maine is the northerly limit for the menhaden; St. Mary Bay on the west coast of Nova Scotia is its most easterly known outpost. Prior to about 1850 the pogy seems to have been common at the mouth of the Bay of Fundy; it was, indeed, reported by Perley as far up the bay as St. John, and fishermen spoke of it as abundant near Eastport up to 1845-1850. But it seems to have abandoned Fundian waters altogether 35 since then except for an occasional straggler, and very few menhaden have been noticed east of Mount Desert and Jonesport of late years. Perhaps the most interesting aspect of the oc- currence of the menhaden in the Gulf of Maine is that it fluctuates tremendously in abundance there from year to year, periods of great plenty al- ternating with periods of scarcity or entire absence " Young menhaden that we collected at Woods Hole on September 23, 1942, were 3J-S to 4 inches (91-99 mm.) long; others taken in Salt Pond, Fal- mouth. Mass.. on November 24, 1949, were 456 to 5 inches long. >' See Hildebrand (Smithsonian Misc. Coll., vol. 107, 1948. No. 18 for a revision of the genus Breiwrtm). One named species, P. Uevicaudala Goode 1878, is known only from Noank, Conn.; we doubt its validity. "According to Huntsman (Contr. Canad. Biol., (1921) 1922, p. 59) one was taken in St. John Harbor in August, 1919. from our waters. Thus they were extremely abundant off the coasts of Massachusetts and Maine, every summer, for some years prior to 1875, when a considerable fishery developed for them in Maine. Very few, however, were taken in the Gulf during the cold summer of 1877 until September and October, when they were reported as about as abundant as normal; practically none appeared north of Cape Cod in the year 1879; and they were so scarce along the coast of Maine for the next six years that it caused com- ment when an occasional one was caught. In 1883, for instance, a few were reported to the U. S. Fish Commission though no schools were seen and many people thought they had gone per- manently. But they were once more reported abundant off Maine and Massachusetts in 1886; they were so plentiful as far east as Frenchman Bay in 1888 that the menhaden fisheries were revived; they were as plentiful in Maine waters in 1889 as they had ever been (more than 10 million pounds taken there) and they were still so numerous in 1890 that four fertilizer factories were established, and nearly 90 million fish were taken during that season. But this period of abundance was short-lived, less than half as many fish being caught in Maine waters (about 41 mil- lion) in 1S91 as the year before, while few men- haden were taken or seen north of Cape Cod in 1892. They were plentiful enough, however, in 1894, for a single steamer to seine about a million fish off the Kennebec during that summer, while 582,131 fish were taken in Boston Harbor in 10 days' fishing during the last half of that August. Menhaden were scarce again in the Gulf during the period 1895-1897 but abundant again in 1898, when about 7 million pounds were taken along the Maine coast. They were scarce in 1902 (Maine catch about 300,000 lb.); reported as abundant again north of Cape Cod, in 1903, especially in Boston Harbor; rare north of Cape Cod from 1904 to 1921, when odd schools were seined along the Massachusetts and Maine coasts in some summers, while few or none were seen in others. Thej' reappeared, however, in such abundance again in the southwest part of the Gulf in the summer of 1922 that 18 steamers fished for them successfully for some weeks in Massachusetts Bay, when upwards of 1,500,000 pounds were landed by the larger fishing vessels, besides what the small boats brought in. And they were so plentiful at FISHES OF THE GULF OF MAINE 117 least as far north as Boothbay Harbor, that about 2,500 barrels were frozen there, though no large schools were reported east of that point. The appearance of menhaden in such abundance in the Gulf after so many years' absence prompted the Bureau of Fisheries to send the steamer Halcyon to Massachusetts Bay that August, and her towings indicated the presence of much greater quantities of diatoms than is usual at that season, evidence that the fish found a better pasture in Massachusetts Bay than in any summer since 1912. But we hesitate to assert that it was an unusually rich food supply that attracted them past Cape Cod. However this may have been, there were not enough menhaden in the Gulf to be- of any com- mercial importance from the middle 1920's to the middle 1940's. But so many visited Massachu- setts Bay, in 1946 and 1947 that local boards of health were forced to clean some of the bathing beaches of the fish that drifted ashore from schools netted for lobster bait. There were a good many in Maine waters in 1948 (reported catch 145,000 pounds);36 more still in 1949, when more than 5,000,000 pounds were taken there; and about 8,000,000 pounds off Gloucester,37 and when small fry, 2-3% inches (52-95 mm.) were taken in the Sheepscot River, December 5-11, suggesting that some had been reared in the Gulf that year. But this peak of abundance lasted no longer than the peak had in the early 1920's, for there seem to have been far fewer menhaden in Maine waters in 1950 than in 1949, as there certainly were in Massachu- setts Bay, where we did not chance to sight a single school, and very few were reported. In the years when menhaden come, they appear in Massachusetts Bay about mid-May; off the Maine coast during the last half of May or first part of June. They are most abundant during July, August, and early September, and most of them depart from the coast of Maine by the middle of October, from the Massachusetts Bay region by early November; and it is unusual to find a single menhaden along these shores after the middle of that month, although small ones have been taken in the Sheepscot River as late as the first third of December. The universal belief among fishermen, that the seasonal appearances and disappearances of men- « Reported by Scattergood, and Trefethen, Copcia, 1951, pp. 93-94. " Reported by Scattergood, Trefethen, and Coffin, Copeia, 1951, p. 298. haden in the Gulf of Maine result from a definite migration from the south around Cape Cod in the spring and a return journey in the autumn, probably is well founded. The brevity of the peaks of abundance, the fact that they come at such long intervals, and es- pecially the great local scarcity of young fish, are arguments against the possibility that menhaden are permanent inhabitants of our gulf, though a few fry may be produced there in favorable summers, as happened in 1949 (p. 117). Menhaden are warm water fish, and our studies of the temperatures of the Gulf of Maine cor- roborate earlier observations to the effect that they never appear in spring until the coastwise water has warmed to 50° or more, or in abundance until the temperature is several degrees higher, which is in accord with Bean's 38 experience that menhaden will not survive in an aquarium if the water chills below 50°. No doubt, it is the falling temperature of autumn that forces the menhaden to leave the coasts of northern New England. In menhaden years the fish occur all along the shores of the Gulf of Maine from Cape Cod to Penobscot Bay, even to Mount Desert. Their chief centers of abundance always lie in Massa- chusetts Bay within a mile or so of land, parlic, ularly off Barnstable and in the mouths of Boston and Salem Harbors; in Casco Bay; and among the islands, thence to Penobscot Bay. But we have never heard of them entering water that is appre- ciable brackish, and in some years they may con- gregate as much as 40 to 50 miles offshore, as happened in 1878, for instance. But we have heard no report of menhaden in the central part of the Gulf or on the off shore Banks. The men- haden are thin when they arrive on our coasts in spring, but they put on fat so rapidly that while the average yield of oil per thousand Gulf of Maine fish was about 12 gallons for the whole summer season of 1894, it rose to 14$ gallons for Boston Harbor fish in August, and to 16 or 18 gallons in September. It is generally accepted, furthermore, that fish taken on the New England coast, south or north, always average larger and fatter than those caught farther south. Commercial importance. — The menhaden is one of the most important, commercially, of the fishes of the Atlantic Coast of the United States, being used for the manufacture of oil, fertilizer and fish " Rept. Xew York State Mus., 60, Zool. 9, 1903, p. 213. 118 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE scrap.39 In 1946, when the catch for the Gulf of Maine was only about 20,000 pounds, the total catch for the Atlantic and Gulf States was 851,129, 000 pounds; the value of the catch to the fisher- men was $7,439,573; the value of the products made from menhaden was $18,196,573. Con- siderable numbers are used locally on the Middle Atlantic coast for bait. But the menhaden is so oily that it is unlikely to become popular as a food fish. Practically the entire catch of menhaden is taken by purse seines and in pound nets; they never bite a baited hook. THE ANCHOVIES. FAMILY ENGRAULIDAE <° The anchovies are small herring-like fishes; bu.t they are easily distinguishable from the herrings by the fact that then mouths are not only very much larger and gape much farther back, but are on the lower side of the head, and are overhung by the upper jaw, which projects like a short piglike snout in some species. Two anchovies are known to occur in the Gulf of Maine; both are stragglers from the south. KEY TO GULF OF MAINE ANCHOVIES 1. Anal fin originates under the front of the dorsal; the silvery lateral band is diffuse; 24 to 27 anal fin rays Anchovy, p. 118. Anal fin originates under the rear rays of the dorsal; silvery lateral band bright and well defined; 20 or 21 anal fin rays Striped anchovy, p. 119. Anchovy Anchoa mitchilli (Cuvier and Valenciennes) 1848 Whitebait Jordan and Evermann (Stolephorus mitchilli), 1896- 1900, p. 446. Description. — The only Gulf of Maine fishes with which one might confuse an anchovy are young herring, smelt, or silversides, but it is easily dis- tinguished from the former by the wide mouth, as just noted; by its much larger eye; by the relative positions of the fins with the dorsal wholly behind 11 For an account of the menhaden industry, see Harrison. Inv. Rept. No. 1, U. S. Bureau of Fisheries, 1931. the ventrals instead of over them and with the latter originating close behind the tips of the pectorals when these are laid back against the body; and by its much longer anal fin. The lack of an adipose fin behind the dorsal is sufficient to separate anchovy from smelt at a glance, while the silversides {Menidia) have two dorsal fins in- stead of one. The anchovy has large, thin, easily detached scales and a deeply forked tail. This species may be distinguished from the striped anchovy by the fact that its anal fin originates under the front of the dorsal ; by its more or less diffuse lateral band of silver; by its more numerous anal fin rays (24 to 27 contrasted with 20 or 21 for the striped anchovy), and by its relatively small size, for it seldom exceeds 3 inches in length. The body is about 4 to 5 times as long as deep in both anchovies. Color. — This is a whitish silvery, translucent little fish, its most characteristic marking being an ill-defined silvery band scarcely wider than the pupil of the eye, running from the gill opening back to the caudal fin. There are also many dark dots on body and fins. Size. — Seldom more than 3% inches long. General range.- — Coast of the United States from Maine to Texas, chiefly west and south of Cape Cod. *° For a recent review of the American anchovies see Hildebrand, Bull- Bingham Oceanographic Coll., vol. 8. art. 2, 1943. Figure 51. — Anchovy (Anchoa mitchilli). FISHES OF THE GULF OF MAINE 119 Occurrence in the Gulf of Maine. — -We mention the anchovy because it has been taken in Casco Bay and at Provincetown. It has no real place in the Gulf of Maine fauna, seldom straying past Cape Cod, though it is abundant about Woods Hole and thence westward and southward. Stragglers may be expected most often in the Gulf in midsummer for it appears from May to October in southern New England waters. Sandy beaches and the mouths of rivers are its chief resorts. An account of its embryology and larval development is given by Kuntz.41 Striped anchovy Anchoa hepsetus (Linnaeus) 1 758 Jordan and Evermann (Stolephorus bToumii), 1896-1900, p. 443. Description. — This anchovy resembles the pre- ceding species closely, but its anal fin is shorter (20 or 21 rays) and originates under the last rays of the dorsal, and it has a very bright and well de- fined silvery band along each side. It is a larger and more robust fish than the other anchovy, often more than 4 inches long. Color. — The bright silvery lateral band, already mentioned, is the most prominent marking on this fish. Fresh specimens are pale gray and irides- cent, the upper surface of the head with some green and yellow; and the back has dusky dots. The dorsal and caudal fins are more or less dusky on some specimens. Size. — Commonly 4 to 5 inches long, maximum length about 6 inches. General range. — Abundant from Chesapeake Bay to the West Indies, and south to Uruguay; north as a stray to Maine and to the outer coast of Nova Scotia;42 a more southerly fish than the other anchovy. Occurrence in the Gulf of Maine. — The claim of this species for mention in the Gulf of Maine is based on one record off the mouth of the Penobscot River, near Portland, October 8, 1930." One specimen was saved and identified, and the her- ring fishermen who brought it in stated that there were "lots of them" on that date. It is not likely that the striped anchovy is other than a straggler to the Gulf, else it would have been found there before this. As it is a gregarious fish, nearly always traveling in small schools, it is not aston- ishing that they may be found together in some numbers, on occasion, even out of their usual range. Figure 52. — Striped anchovy {Anchoa hepsetus), Somera Point, N. J., specimen 100 mm. long. THE SALMONS. FAMILY SALMONIDAE The salmons are soft-rayed fishes with no spines in any of the fins, with the ventrals situated on the abdomen far behind the pectorals, and with a fleshy rayless "adipose" fin on the back behind the rayed dorsal fin. The presence of this adipose fin, and its situation, separates them from all other Gulf of Maine fishes except for the smelt, capelin and the argentine, the pearlsides (p. 144), and some of the lantern, viper, and lancet fish » Bulletin. U. S. Bur. of Fish., vol. 33, WIS, p. 13. tribes (p. 141). 41 The blunt noses, stout bodies, and nearly square tails of the salmons distinguish them at a glance from the sharper-nosed, slender, forked-tailed smelts, their large mouths and smaller eyes from the argentine; the absence of lumi- nescent organs distinguishes them from the « Five were taken in Bedford Basin, Halifax, Nova Scotia, on September 29, 1931 (Vladykov, Proc. Nova Scotia Inst. Sci., vol. 19, 1935, p. 3). « Kendall, Bull. Boston Soc. Nat. Hist., No. 58, 1931. « Sundry other deep-sea fishes have adipose fins. 120 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE pcarlsides, while the lantern, viper, and lancet fishes are of different general aspect. Four salmons 45 occur in the Gulf of Maine, or have recently, one of which, the sea trout, resorts to tidal estuaries at the mouths of a few of our streams; a second and a third — the humpback salmon and the silver salmon — were introduced from the Pacific coast, leaving the Atlantic sal- mon as a characteristic inhabitant of the open waters of the Gulf of Maine. KEY TO GULF OF MAINE SALMONS 1. Scales so small that they are hardly visible; back with vermiculate markings; teeth on roof of mouth con- fined to a group in front Brook trout, p. 120 Scales large enough to be easily visible; back without vermiculate markings; a row of teeth runs back along the mid line of the roof of the mouth 2 2. Anal fin with only 8-10 rays Salmon, p. 121 Anal fin with 12 rays or more 3 3. Back and lower half of tail fin, as well as its upper half, conspicuously marked with large black spots Humpback salmon, p. 131 Back with very small black spots or none at all; no black spots on lower half of tail fin. Silver salmon, p. 133 Brook trout Salvelirms jontinalis (Mitchill) 1S15 Sea trout; Salter Jordan and Evermann, 1896-1900, p. 506. Description. — Although brook trout vary widely in general form in different streams, they are usu- ally salmon-like in shape when taken in salt water, that is, about one-fourth as deep as long, tapering gracefully to a small head. The nose of a trout, " A specimen of one of the whiteflshes (probably Coreqonus quadrilateralis Richardson) was taken in the mouth of the Sissibou River, St. Mary Bay. Nova Scotia, September 1919 [Huntsman, Contr. Canad. Biol., (1921) 1922, p. 59] straying down from fresh water. Whitefish have an adipose fin, like the true salmons, but have a very small mouth, and are flattened sidewise, and herring-like in appearance, rather than salmon-like. however, is blunter than that of a salmon, and its head is longer in proportion, the total length of the fish (not counting the caudal fin) being about four and one-half times that of the head,46 while its mouth (gaping back of the eye) is relatively larger. The general arrangement of the fins, in- cluding the "adipose," parallels that of the salmon, but the ventral fins stand under the middle of the dorsal, thus farther forward in relation to the latter than in its larger relative. All tbe fins, too, are relatively larger, particularly the ventrals; as a rule the anal has one less ray in the trout (usually 8) than the salmon, but the number of dorsal rays (about 11) is the same. The tail of the sea trout is less forked than that of a young salmon of equal size. Examination of the scales and of the teeth is the most positive means of distinguishing brook trout (in European terminology this is a "charr") from young salmon, for the teeth on the roof of the mouth of the trout are confined to a cluster near the front, instead of extending backward in a row along its midline as in the salmon; and the scales of the trout are so tiny as hardly to be visible whereas those of the salmon are large and easily seen. Color.- — Trout living in salt water almost wholly lack the yellow and red tints so conspicuous on their freshwater relatives. They are steel blue or bottle green on the back, with cheeks and sides silvery like a salmon and with a white belly. The sides above the lateral line are more or less dotted with pale yellow spots, but the dark vermiculate markings so characteristic of the fresh-water brook trout are rarely seen on the trunk of sea run fish, though evident as wavy crossbars on the dorsal " Some trout are longer headed. yfer U_d^_J" "" ' V '$iS^*Vj^ff-;; !. £jjjg»£ Figure 53. — Brook trout (Salvelinus fontinalis), about \5% inches long. FISHES OF THE GULF OF MAINE 121 fin and on the corners of the caudal fin. The sides and flanks below the level of the lateral line usually are strewn with small pale vermillion dots, but the ventral fins are often plain white; at most, the pink edging so conspicuous in trout caught in fresh water is faint on fish in salt water. General range. — Eastern North America, north to the outer coast of Labrador, west to Minnesota, and southward to Georgia along the Allegheny Mountains. Occurrence in the Gulf of Maine. — Brook trout are plentiful in many of the river systems and smaller streams that empty into tbe Gulf of Maine. Some of the trout in some of these seek salt water after the breeding season, to remain there over the winter. This applies particularly to the brooks that flow through the sands of Cape Cod, several of those on its southern slope being famous for their sea-trout fisbing. These, however, lie outside our present province, and only a couple of small streams on the Massachusetts Bay side of the Cape still support a race of trout that run down to the sea regularly. One or two small brooks tributary to Ipswich Bay, and the Merriland River, emptying between Wells and Kennebunk- port, Maine, are the only places between Cape Ann and Cape Elizabeth where we have heard of sea run trout. We cannot say how generally sea trout may now exist in the streams of eastern Maine, but accord- ing to Evermann 47 trout once inhabited the tidal portions of many of the brooks that empty into Casco Bay, and they still may. Some of good size are caught also in the Belfast River waters, tributary to upper Penobscot Bay.48 Huntsman found no definite evidence of trout in salt or brackish water on the New Brunswick side of the Bay of Fundy, but local inquiry has elicited the information that there are fish of this habit in a few streams (notably in Salmon River) on the north and west coasts of Nova Scotia, where many streams formerly held sea run trout that have been fished out long since. The "sea trout" are indistinguishable from the ordinary brook trout anatomically.49 They are simply fish that have the habit of running down to salt water, and most of the trout never leave •' Rcp't. U. S. Comm. Fish., (1901) 1905, p. 105. " Towne, Striped Bass Survey, Maine Development Comm. and Dept. Sea and Shore Fisheries. 1940. p. 21. '• There is another species of sea trout (Salvelinus alpinus) in northern Canadian waters which is very plentiful along the coast of northern Labrador. 210941—03 9 fresh water, even in streams offering free access to the sea, cold enough throughout their lengths, and harboring these "salters" (as they are called on Cape Cod). All who have given special atten- tion to our sea trout are agreed on this. It is still an open question whether the habit is hered- itary or whether it is acquired independently by each individual fish. We incline to the first view, chiefly because sea trout are slow in reestablishing themselves in any stream where they have been brought to a low ebb by hard fishing. The trout that follow this habit grow much more rapidly on the abundant rations the salt estuaries provide than do most of their relatives that remain in the brook. Sea fish weigh from 1 to 3 pounds in streams where few of the fresh-water trout exceed half a pound. On Cape Cod the sea trout go down to salt water in November immediately after spawning, to winter there. They begin to run again in April, and all of them are in brackish or fresh water by mid-May. But it is said that they do not appear until later in the Xova Scotia streams tributary to the Bay of Fund}- (we cannot vouch for this). While in salt water (at least along Cape Cod) the trout feed chiefly on shrimps or on gammarid Crustacea, on mummichogs (Fundulus), and on other small fish. Trout never stray far from the stream mouths; hence they have no place M in the fish fauna of the open Gulf. Salmon Salmo salar Linnaeus 1758 Atlantic salmon; Sea salmon; Silver salmon; Black salmon; Parr; Smolt; Grilse; Kelt Jordan and Evermann, 1896-1900, p. 486. I)i tcription. — The Atlantic salmon is a graceful fish, about one-fourth as deep as long, deepest below the dorsal fin, whence it tapers toward both head and tail; and oval in cross section. Its head is small (about one-fifth, or less of the fish's length, not counting the caudal fin), its nose is blunt, eye rather small, and its mouth gapes back to below the eye. The dorsal fin (about 11 rays) stands about midway between tip of snout and base of tail fin; the vcntrals are under the rear end of the dorsal. The anal is similar in form to the dorsal but has only about 9 rays (7 to 10 have » Trout arc taken about Woods nolc, occasionally, in winter. 122 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 54. — Salmon (Salmo salar). Drawing by H. L. Todd. been recorded), whereas the humpback has 14 anal rays or more. The tail is only very slightly emarginate in adults, and is almost square in large fish, but is more forked in fish that have been at sea for not more than one year ("smolts" and "grilse"). Color. — The salmon is silvery all over while in the sea, with brownish back and with numerous small black crosses and spots on head, body (chiefly above the lateral line), and fins. The young fish (or "parr") are conspicuously marked with 10 or 11 dark crossbars while in fresh water, alternating with bright red spots, much like young trout. Fish that have been at sea for only one year (grilse) are marked with a larger number of black spots than the older fish. Size. — The largest salmon we find mentioned was an English fish of S3 pounds. None even approaching this size is recorded from our side of the Atlantic, where a 50-pounder is unusual, though fish of 40 pounds are not uncommon in some of the larger rivers emptying into the Gulf of St. Lawrence. Very few fish reach 40 pounds in the Penobscot or St. John Rivers, and 30- pounders are unusual there, the usual run being 10 to 12 pounds. Taking one river with another, large and small, 10 pounds may be set as a fair average of the mature Gulf of Maine fish. A 2- foot fish will weigh about 6 pounds, one of 3 feet, 16 to 20 pounds, with allowance for individual and seasonal variation. Remarks. — The teeth and the scales afford the most certain distinction between small salmon and the New England sea trout (p. 120). In the salmon the roof of the mouth is armed both with a cluster in front and with a row of stout conical teeth running back along the mid-line, easily felt with the finger, whereas the sea trout has the forward group only. The scales of the salmon are so large that they are seen easily, whereas those of the trout are so minute that they are hardly visible. Old salmon sometimes lose the teeth on the roof of the mouth, but large size and large scales identify them at a glance. It should also be easy to tell an Atlantic salmon from a humpback (should any of the latter still exist in our Gulf) for the black spots on the upper part of the body of the humpback and on its tail fin are more close set and much larger and con- spicuous than the dark markings on a salmon. A more precise difference is that an Atlantic salmon never has more than 10 rays in its anal fin, whereas the humpback always has at least as many as 12, while most of them have 13 to 17. The danger will be greater of confusing smallish Atlantic salmon with silver salmon, if the attempts now in progress to establish the latter in our Gulf should succeed, for the two fish look much alike. A reliable criterion is, again, the number of rays in the anal fin, for the silver salmon always has as many as 13 of these, an Atlantic salmon never more than 10. Life history.61 — It is no wonder that the life of the salmon has been the subject of much scientific study and that a whole literature has grown up about it. As everybody knows, the salmon lives the greater part of its life in the sea and makes most of its growth there but spawms in fresh water. The salmon are silvery and very fat when they enter fresh-water on the spawning journey, but •i Huntsman (Bull. Biol. Board Canada, 21, 1831) has published an exten- sive study of the life history of the salmon of the Maiitime Provinces of Canada, from which we have drawn freely in the following account. See also Huntsman and others (Migration and Conserv. of Salmon, Pub. No. 8, Amer. Assoc. Adv. Sci., 1939) for discussions of the movements of the salmon in Canadian and Newfoundland waters; also Lindsay and Thompson (Rept. Newfoundland Fish. Res. Comm., vol. 1, No. 2, 1932) for an account of the biology of the salmon in the rivers and around the coasts of Newfoundland . FISHES OF THE GULF OF MAINE 123 they lose condition gradually as they work up- stream, for they feed very little in fresh water, if at all; they make no attempt, for example, to capture the parr they meet. Most anglers believe that they may occasionally snap up a small fish or other tempting morsel. Many are caught on artificial flies, while every salmon angler knows that they will sometimes take a hook baited with angleworms or with prawns. It has been suggested that salmon recover the feeding habits of their youth to some extent after they have spent some time in the river, for they often rise to floating insects. But the stomachs of salmon caught in fresh water never contain anything but a little yellowish green fluid. And the fact that they keep better with bellies intact than if opened and gutted suggests that the secretion of effective digestive juices has ceased. The maturing salmon of both sexes lose their silvery sheen in fresh water during the summer months, to take on a dull brownish or reddish hue, while the belly suffuses with some tint of red, large black spots develop, and the male not only becomes variously mottled and spotted with red or orange, but his jaws elongate, the lower becom- ing so hooked that only the tips come together. His body becomes slab-sided, his fins thicken, and his skin is covered with slime, until altogether he is but a caricature of the beautiful silvery creaturo that came in from the sea. In small streams salmon may spawn only a short distance above the head of tide; but they may run upstream for more than 200 miles in large rivers that are not obstructed, as they do in the St. John system in New Brunswick. In Gulf of Maine rivers they spawn in October and early November, on sandy or gravelly bottom, the females smoothing a shallow trough or redd and covering the eggs with gravel. As it is with the life of the salmon in the sea that we are concerned here, the reader is referred to Belding 62 and to Kendall M for recent accounts of the mating actions of the males and females. The spent fish, known as "kelts," "slinks," or "black salmon," are thin, weak, and so exhausted that many of them die. Most of those that survive in small rivers drop down at once to the sea after spawning. But many of them linger over the winter in large rivers, improving somewhat in " Trans. Amer. Fish. Soc, vol. 24, 1934, p. 211. « Mem. Boston Soc. Nat.. Hist., vol. 9, No. 1, 1935, p. 65-6S. condition and becoming more silvery, though they take little food. If they survive the winter (which many do not, for spawning leaves them thin and exhausted) they drop downstream to salt water the following spring.64 The large (6 to 7 mm.) thick-shelled eggs lie loose on the bottom and develop so slowly in the low temperature of winter that hatching does not take place until late in the following April or early in May. The newly hatched larvae are 15 to 18 mm. (0.6-0.7-inch) long, and carry a very large yolk sac for about 6 weeks, hiding among the pebbles of the spawning bed and taking no food. When the yolk sac is absorbed the little fish, now known as "parr," begin to swim and feed. Parr live in fresh water for longer or shorter periods according to locality or to other factors not well understood. In the St. John,65 and in the rivers of Minas Basin, most of them remain for 2 summers and 2 winters, running down to the sea the third summer. But Huntsman has found that some few stay in the Fundian rivers for 3 years. Most of the salmon of the Penobscot spend 2 years as parr, a few 3 years, according to Kendall. It is even possible that some may linger in Gulf of Maine rivers for 4, 5, or even 6 years, as is known to happen in Norway. And Dr. Huntsman informs us that some of the male parr in the rivers of the Chignecto Peninsula become sexually mature before visiting the sea. Parr may be moving downstream any time from late spring to autumn, but most of them probably make the journey in June and July in Gulf of Maine streams, when they are 5 to 6 inches long. They put off their barred and spotted pattern as they near tidewater, to assume the silvery coat worn by the salmon during his sojourn in the sea. They are now known as "smolts." Salmon, small or large, are voracious while in salt water, feeding altogether on live bait, chiefly on fish and on crustaceans. Among fishes avail- able to them in this side of the Atlantic, launce, herring, alewives, smelt, capelin, small mackerel, haddock, small sculpins, and even flatfish have all been reported as entering into their diet in one place or another. Salmon caught off Norway are sometimes packed full of herring, and a hook ** They are voracious now, and fly-fishing for these "black salmon" as they are called, is a favorite sport nowadays, especially in Miramichi waters tributary to the Gulf of St. Lawrence. •' Huntsman, Bull 21, Biol. Bd. Canada, 1931, p. 31. based on studies by Kerr and by Blair. 124 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE and line fishery is carried on for salmon in the Baltic, with herring for bait, while in British waters salmon are sometimes caught on hooks baited with launce and with pieces of mackerel. Launce and capelin had been the chief diet of thousands of salmon opened by Comeau66 in the northern side of the Gulf of St. Lawrence. And it is probable that the salmon of the Bay of Fundy and open Gulf of Maine feed chiefly on herring (herring up to 5 inches long have been found in salmon stomachs near Eastport) and on launce, taking alewives or any other small fish as occasion offers, including smelts and mummichogs (Fundu- lus), when they first enter the estuaries.67 Salmon also feed greedity on euphausiid shrimps (fish entering the Penobscot have been found full of "shrimp," probably euphausiids) ; to some extent on pelagic amphipods (Euthemisto) , while sand fleas (gammarid crustacean) are described as ranking with launce and herring as salmon food in the North and Baltic Seas. Salmon are also credited with eating crabs.68 Smolts, on the other hand, fall prey to any large predaceous fish (they have been found in the stomachs of pollock), but salmon are so heavy and strong after one or two years' sojourn in salt water that only fish as large as tuna, swordfish, or the larger sharks can menace them. Their worst enemy is the harbor seal, which is a com- mon inhabitant of the northeastern coasts of the Gulf of Maine and of the Bay of Fundy. The young smolts grow so rapidly on the abun- dant diet the sea affords that they usually reach a length of at least 16 inches and a weight of any- where from K to 7 pounds after one year at sea. They are now known as "grilse." And older sal- mon continue to put on length and weight very fast, as long as they remain in salt water. Thus, several St. John fish which were tagged and re- leased in the river in the autumn after spawning and which were recaptured the following summer after wintering in the sea had gained 2 to 8 pounds in weight, one of them more than 6 inches in length. Others which spent two uninterrupted years in the sea (as shown by their scales) averaged about " Life and Sport on the North Shore, 1909, Quebec. •' Kendall (Mem. Boston Soc. Nat. Hist., vol. 9, No. 1, 1935, p. 34) found smelts in Penobscot salmon, alewives in salmon from the St. John. •' See Kendall (Mem. Boston Soc. Nat. Hist., vol. 9, No. 1, 1935, pp. 33-34) for a recent survey of the diet of salmon in general; the Gulf of Maine fish in particular, with references. Eichelbaum (Cons. Perm. Internat. Explor. Mer, Rapports et Proc. Verb., vol. 21, 1916, p. 84) records the contents of many salmon from the Baltic and from the North Sea. 10 pounds heavier and 6 inches longer when re- captured.69 But they grow much less rapidly in winter than in summer. And they hardly grow at all during the years when they spawn if they enter the river early in season, though they con- tinue growing until later if the}7 enter late. Hence the size of a salmon depends more on the number of times it has spawned and on the date when it enters its river than on its age. Most of the exceptionally large fish of 40 to 50 pounds are virgin females entering fresh water for the first time, but some are fish that have already spawned once. An interesting case is that of a 45-pound 2-ounce fish, caught in the Moisie River, on the north shore of the Gidf of St. Lawrence, June 1950, by E. E. Steedman, the life history of which had been as follows:60 hatched spring 1942; went to sea June 1945; returned to river and spawned there in 1948; returned to sea autumn 1948; remained there until June 1950; then re- turned to the river, to be caught on a "Lady Am- herst" fry; age 8 years. Some salmon become "river mature" and return to spawn after only one year at sea; these, known as grilse, are distinguishable from the older fish by more forked tail, more slender body, thinner scales, and more numerous spots that are blue rather than black.61 Some spawn 2 or 3 years in succession, and hence never grow large; others spawn twice in alternate years; a few three times, very few oftener. It follows from this that large salmon are to be found in the sea throughout the year, though fewer of them in summer when the spawning fish are in the rivers, than in winter when the whole stock is in salt water except for the "parr," a few immature grilse (p. 129), and such of the spent fish as winter in the rivers. Some spawn only once after 3, 4, or even 5 years at sea, growing to a great size meantime. But very few salmon five to be more than 8 or 9 years old, including the time spent in fresh water as parr. Our ignorance of the way of life of the salmon in the sea has recently been characterized as abyssmal. Certainly they are swift swimmers, and the nature of the catches suggests that they " Huntsman (Bull. Biol. Board Canada, No. 21, 1931) gives an interesting account of these tagging experiments, from which this summary is drawn. •o As worked out from its scales by Dr. D. L. Belding, and reported in Field and Stream, August 1951. p. 10. •> It is commonly stated that this applies chiefly to the males. But. Hunts- man (Bull. Biol. Board Canada, No. 21, 1931, pp. 18-19) has found that grilso of both sexes spawn in the small rivers at the head of the Bay of Fundy. FISHES OF THE GULF OF MAINE 125 live scattered for the most part. But at least one case has come to our notice of a school seen, and some of them netted.62 While salmon often leap in the esturaries on their return journey and in the rivers, we have never heard of one doing so at sea. And they keep so constantly to the mid-depths that they are seldom seen at the surface, except in the estuaries. But this rule has its exceptions, for the school mentioned above was sighted at the surface, where they were mistaken for pollock. On the other hand, there is no reason to suppose that many of the Gulf of Maine salmon descend to any great depth, winter or summer. The weirs, gill nets, and other gear that yield so many in various regions, are all operated in rather shoal water (the Baltic hook- and-line-fishery is carried on at about 1 % fathoms) . Dr. Huntsman informs us that salmon are taken on hand lines in mid-winter in the Bay of Fundy. They are caught occasionally on long lines in the Gulf, and otter trawlers get stray salmon on the offshore Banks (p. 126), proof that at least some may go as deep as 50 fathoms or so, while diet (p. 124) proves that they sometimes feed near bottom if not actually on it. General range.- — Coastal waters of both sides of the North Atlantic, entering rivers to spawn. On the European side its range extends northward well within the Arctic Circle; southward to the Mifio River, at the boundary between Spain and Northern Portugal, perhaps with a few reaching the Duero River, midway of Portugal.63 It occurs in a few rivers in western Greenland.64 On the American side salmon ran up all suitable rivers, formerly, from northeastern Labrador to the Housatonic emptying into Long Island Sound; perhaps the Hudson also. The northern limit of the commercial fishery for it on the American side is only about latitude 54° N. (Indian Harbor, north shore of Hamilton Inlet). And while it is known to range to Hudson Strait,66 reports of it from stream mouths northward from Hamilton Inlet seem often to have been based * Kendall. Mem. Boston doc. Nat. Hist., vol. 9, No. 1, 1935, p. 32. •» This is the southern European limit given by Berg (Zoogeographies. vol.1, Ft. 2, 1932, p. 112. "Jensen, Fauna of Greenland, vol.1, Pt.3, Fishes, 192S, pp. 3 and 4, Copen- hagen. « Vladykov (Contrib. Canad. Biol., N. Scr., vol. 8, No. 2, 1933, p. 18, fig. 1) shows a locality record near Fort Chimo, and there are salmon in the rivers of the eastern part of Ungava Bay. on the sea run form of the Arctic charr Salvelin us alpinus, which also grows large in the sea.66 Occurrence in the Gulf of Maine. — When the white man first came to New England and to the Maritime Provinces, he found salmon in every large stream not barred by impassable falls, from Cape Sable to Cape Cod; i. e., in all the Nova Scotia and New Brunswick Rivers, tributary either to the open Gulf of Maine or to the Bay of Fundy, and in the following rivers in New England: St. Croix, Dennys, Orange, East Machias, Pleasant, Narraguagus, Union, Penob- scot, St. George, Medomak, Sheepscot, Andro- scoggin, Kennebec, Royal, Presumpscot, Saco, Mausam, Piscataqua, and Merrimac.67 One New England river, however, after another was so obstructed by dams after the beginning of the past century, that salmon regularly entered only the St. Croix, Dennys, East Machias, Machias, Penobscot, Sheepscot, Kennebec, and Andro- scoggin by the 1880's. The Kennebec was still an important salmon river as late as 1895. But by 1925 the Dennys and the Penobscot alone, of the rivers of Maine, saw regular runs, with a few fish in the St. Croix where pollution by sawdust was not as bad then as it had been, perhaps with an occasional fish in other streams. The fate of the salmon in the Merrimac "8 typi- fies its history in the rivers from which it is now barred. Salmon spawned plentifully in the upper tributaries, especially in the Pemigewasset, as late, as 1793 (in 1790 the run was so abundant in the lower river that 60 to 100 a day was the usual catch with a 90-yard seine near the mouth at Amesbury), but the completion of the dam at Lawrence in 1847 completely barred the upper reaches of the river. For some years thereafter salmon congregated below the Lawrence dam in spring and summer, vainly endeavoring to ascend, but there has been no run of salmon in the upper Merrimac since 1859 or 1860, when the last salmon hatched above the dam had lived its span of life, nor have any spawned there since then with the possible excep- tion of a few that have been lifted over the dam by hand. - Blair (Res. Bull. 12, Dept. Nat. Resources Newfoundland, 1913, pp. 5-17) gives a detailed account of the salmon rivers of the outer Labrador coast. Strait of Belle Isle to Hamilton Inlet. « Atkins (1887, Fish. Ind. U. S., Sect. 5, vol. 1, p. 679) has collected much information on the local history of salmon in northern New England. « Lyman and Reed, Rept. Comm. Fish. Massachusetts (1865) 1866, Senate Doc. 8, pp. 36-41. 126 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Enough salmon to yield a supply of eggs for artificial hatching continued to enter the lower Merrimac up to 1893 and there seems to have been what almost might be described as a run there in 1896, when salmon were seen leaping below the Lawrence dam nearly every day from June 10th to July 25th, often 10 or 20 at a time, and a few were lifted over. But we have not learned of a single sea-run salmon seen in the Merrimac since 1901, though watch has been kept for them by the wardens of the Massachusetts Division of Fish- eries and Game," and it is not likely that salmon would still run in the Penobscot were it not for the artificial propagation that is carried on there by the U. S. Fish and Wildlife Service. But the sal- mon situation now shows signs of improvement, for the run in the St. Croix has increased; salmon have reestablished themselves in the Narraguagus and provide sport there for many anglers since one obstructing dam has washed out and another opened. Enough salmon run regularly in the Dennys to attract anglers and a few also in the Machias and Pleasant Rivers. The Fisheries Com- mission of the State of Maine, and the U. S. Fish and Wildlife Service, are now concerned with the possibility of improving the runs in these streams, and of reestablishing runs of salmon in other Maine rivers. Along the Canadian shores of our Gulf a few salmon still run in the Tusket, Salmon, and An- napolis Rivers; many in the Shubenacadie River in Nova Scotia, some in the Petitcodiac, and great numbers in the St. John River in New Brunswick, which still is a famous salmon river. Movements in the Gulf.- — After the smolts reach salt water they are found for a time in the river mouths and about estuaries. No doubt the little salmon (too small to sell) that are caught in sum- mer and autumn in weirs at Matinicus Island have come from the Penobscot a month or two previous. They drop out of sight in winter, as do the older and larger salmon as well. But there is no reason to suppose that many of them go far out to sea in the Gulf. Odd salmon stray, it is true, as much as 90 to 100 miles sea- ward off the outer coast of Nova Scotia,70 while otter trawlers pick up odd salmon from time to time in the South Channel, and even on Georges Bank up to 160 miles or more at sea from Cape Cod.71 But the great majority of the salmon that are caught in the Gulf are taken within 25 miles of the land. The Gulf of Maine salmon also appear to con- tinue rather closely localized as a whole, not only within the coastal belt, but within the zone of fresh-water influence from the particular rivers or river systems from which they come. So few, for example, are caught near Cape Sable that there can be no general movement around the Cape by the fish that spawn in the rivers of the outer coast of Nova Scotia. Most of the fish that go to sea via Minas Channel from the Shubenacadie, and the few from smaller streams that discharge into Minas Basin, seem to remain along the Nova Scotia shore within a distance of 30 to 40 miles to the westward. And while tagging experiments have proved that some of them scatter more widely; i. e. to Cobequid Bay, to the estuary of the St. John River, to the Annapolis Basin, and to St. Mary's Bay, few of them leave the Bay of Fundy72 (for some that did, see p. 127). The much more numerous salmon from the St. John appear to hold rather closely to the tongue of low salinity that extends westerly from the mouth of the river, keeping out from the shore, for hardly any salmon are caught either on the New Bruns- wick shore to the eastward, except for a few near the head of the Bay (doubtless the product of the Chignecto Bay river system) or farther west than Point Lepreau, or around Grand Manan Island which stands directly in the route of any fish mov- ing westward out of the northern side of the Bay of Fundy. Thus it appears that a radius of, say, 40 to 50 miles would enclose the wanderings of most of the St. John River fish. The evident failure of salmon from the St. John to follow the myriads of sardine sized herring into Passamaquoddy Bay is especially interesting. The weirs there pick up a few salmon, the presence of which can be credited to the small run in the St. Croix River. And the numbers of salmon that are caught thence westward along the coast of Maine73 89 A few small "salmon" reported of late in the Merrimac probably were the landlocked form, running down from tributaries stocked with this fish. "> Three reports of salmon caught on Western Bank have appeared in the daily press since 1925 to our knowledge, and Kendall (Mem. Boston Soc. Nat. Hist., vol, 9, No. 1, 1935, p. 33) reports one caught on La Have Bank 100 miles from Halifax, and another 60 miles off Cape Sable. " Kendall (Mem. Boston Soc. Nat. Hist., vol. 9, No. 1, 1935, pp. 31-33) lists a number of such cases. » Huntsman, Ann. Rept. Fish. Res. Board Canada, (1947) 194S, p. 37, and unpublished notes. 1 The average was only 3,000 pounds (perhaps 300 fish) for the years 1939, 1940, 1943, 1944. Statistics are not readily available for 1941 and 1942. FISHES OF THE GULF OF MAINE 127 are not larger than can be credited to such of the Maine rivers as still have runs of salmon. It seems certain, also, that only odd salmon from the Penobscot and from the rivers farther east ordinarily disperse westward and southward be- yond Casco Bay, for while the average catch for the coast of Maine east of that point has averaged about 12,000 pounds (some 1,200 fish) for the 10 most recent years of record74 combined, the cor- responding 10-year average for the whole western side of the Gulf from Cape Elizabeth to the elbow of Cape Cod was only 600 to 700 pounds, or some 60 to 80 fish at most, with more than 100 pounds reported in only 5 of the 10 years and none in 3 of the years. Further evidence of a more gen- eral kind that Gulf of Maine salmon do not scatter far as a rule is that they appear about the river mouths in spring so soon after the ice goes out that they cannot have come from any great distance. A few do stray as far as Cape Cod Bay in most years; witness catches of one to 5 or 6 fish (10-55 pounds) in 14 out of 16 years by 8 traps, at North Truro, Cape Cod, during the period 1935 to 1950, in the months of May, June, July, September, and November.76 A year comes from time to time when a con- siderable number are taken off the coast of Massachusetts. The most recently recorded in- stance of this sort fell in 1937, when floating traps along the North Shore of Massachusetts Bay picked up 4,400 pounds of salmon. All of these were taken close inshore. But the 1,600 or so salmon (16,050 lb.) that were reported for Massa- chusetts in 1928 (the big year next previous) seemingly were farther out at sea, for all of them either hooked on long lines (10,134 lb.), or were taken in otter trawls. These must have come from as far as the Penobscot, if not from the Bay of Fundy, which is equally true of the salmon that are caught around Marthas Vineyard from time to time.78 One, however, of about 10 pounds, reported in the North River, Marshfield, in the summer of 1938, and a few seen jumping in the Parker River (also in Massachusetts) in the sum- mer of 1951, may have been the product of attempts to stock these streams. Occasional sal- mon that have been taken along the New Jersey " 1933, 1935, 1937, 1938, 1939, 1940, 1943, 1944, 1945, and 1940. " Information contributed by the Pond Village Cold Storage Co. '• In the spring of 1915 about 75 (including fish up to 35 lb.) were taken at Gay Head and in the neighborhood of Woods Hole coast and off Delaware 77 may have been the product of attempts to stock the Hudson. Salmon, also, of 25 to 50 pounds that are sometimes caught in Minas Channel at the head of the Bay of Fundy, must come from afar, as Dr. Huntsman points out,78 probably from the Gulf of St. Lawrence, there being no run of fish so heavy in any Bay of Fundy river or in any Maine river. It is not astonishing that some salmon should stray far afield in Gulf of Maine waters, for marked salmon have been known to make much longer journeys, elsewhere. Thus fish marked in the southern side of the Gulf of St. Lawrence have been recaught on the north shore of the Gulf; in Newfoundland; and in the Strait of Belle Isle.79 One marked at Bonavista on the east coast of Newfoundland was retaken 98 days later in the Margaree River, Cape Breton Island, Nova Scotia, 550 miles away 80 by the shortest possible route. One marked in Minas Channel at the head of the Bay of Fundy went out around Nova Scotia to Chedabucto Bay on the northeast, near the Gut of Canso, a journey of at least 440 miles.81 Five, tagged in the Annapolis River system, were recaught on the east coast of Newfoundland, a minimum distance of 900 miles, while a sixth, from the same lot, was taken at Ramah on the outer coast of Labrador, more than 1,000 miles still farther away to the northward.82 This last is the most spectacular case of wandering yet reported for any Gulf of Maine or Gulf of St. Lawrence salmon. What is chiefly interesting about the large catches that are sometimes made off Massachu- setts is their demonstration that so many fish may occasionally wander so far afield. And this ap- plies not only to large salmon but to smolts in their first year at sea, for salmon so small that they must have rim down to salt water but a few months previous have been taken in Cape Cod Bay in October. It is not likely that these wandering salmon return at all to their home rivers; probably they " Smith (Bull. U. S. Fish. Comm., vol. 14, 1895, p. 99) reports salmon seined among some mackerel oU Delaware in 1893. " Dull. 51. Biol. Board of Canada, 1936, p. 9. » See Huntsman, Pub. Amer. Assoc. Adv. Sci. 8, 1932, p. 35, for summary of these records. » Huntsman, Science, vol. 95, 1947, p. 381. " Huntsman, Ann. Rept. Fish. Res. Bd. Canada, (1947) 194S, p. 37. •> Huntsman, Science, vol. 85, 1937, p. 314; Pub. 8, Amer. Assoc. Adv. Sci., 1939. p. 35. 128 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE are lost permanently from the breeding popula- tion. But the much greater numbers that remain localized not very far from their parent streams are believed to follow about the same routes on their return journeys that they followed when they went to sea. Thus, only a few are caught on the Nova Scotia shore between the entrance to St. Mary's Bay and Digby Gut, but fish en route to the Shubenacadie River system are taken in some numbers as they follow the shore of An- napolis and Kings Counties (the Annapolis River also yields a few salmon in its lower course, and some are taken in the Annapolis Basin). Simi- larly, salmon approaching the St. John River strike the coast about Point Lepreau (about 23 miles to the west) and support an important fishery from there to the mouth of the river. A question closely bound to the movements of salmon to the sea is: what proportion of them return to spawn in the very rivers in which they were hatched? It seems demonstrated by a variety of evidence, especially by the recapture of tagged fish, that the majority do return. Huntsman, for example, reports83 an extraordinary instance, of a kelt taken from the Sackville River on the outer coast of Nova Scotia that was tagged and released in the Shubenacadie River system at the head of the Bay of Fundy, and then found its way out of the Bay, around the Nova Scotia coast, and back again to the Sackville, where it was recaptured. We can only speculate how it directed its course, and why it did not turn in to the mouth of any of the other salmon rivers it passed en route. On the other hand, marked fish are sometimes caught in strange rivers. Fish, for instance, that were tagged in Minas Channel have been caught later in the St. John River.84 And odd fish appear from time to time in rivers where no salmon have been hatched for many years (in the Merrimac for instance). In short, the parent-stream theory does not always hold. Probably the truth is that while most of the fish never stray far away and do return to the home stream, wanderers that chance, in the spring, to be in the physical state leading to maturity may enter any unpolluted stream they encounter, no matter how far from home. Dr. Huntsman's studies, carried on through many years, make it increasingly probable that «! Ann. Rept. Fish. Res. Bd. Canada (1947) 1948, p. 33. " Huntsman, Ann. Rept. Fish. Res. Bd. Canada (1948) 1949, p. 40. the journeyings of our salmon in salt water are not the result of purposeful swimming in a definite direction, but that they tend to drift with the current as herring do (p. 97), so that the direction in which they travel depends chiefly on the depth at which they happen to be, in relation to the dif- ferential circulation of the water at different levels. If so, the St. John River fish tend to drift out with the river water as they scatter. And most of them do appear to remain more or less concentrated in the mid-depths where the princi- pal mixing takes place between the river dis- charge and the water of the open Bay of Fundy, some 20 to 30 miles from St. John Harbor, living where they find an abundance of herring of various sizes as food. Here Dr. Huntsman 86 calculates the space for them is so great that no two of the approximately 50,000 fish that comprise the total yearly catch need be closer to each other than three-quarters of a mile in a layer of water 5 feet thick; so there is no crowding. But the tagging experiments have shown that the fish that go to sea from Minas Channel, where the outflow is not so definitely localized, scatter more widely, some of them drifting right around the Bay of Fundy with the anti-clockwise circulation.86 The situation is not so clear for the coast of Maine, partly because of the paucity of present- day information, partly because the several rivers there that once had runs of salmon are so closely spaced along the coast that it is not pos- sible to evaluate their individual contributions to the yearly catches. With the relationship between salmon journeys and water movements so extremely complex, all we dare say in this regard is that the inshore drift of the deeper layers (characteristic of circulation of the estuarine type) and the slackening of the offshore drift of the fresher surface water that is to be expected as the spring freshets diminish, may be the cause, at least in part, for bringing the salmon into the estuaries, and close inshore elsewhere, in spring. But the nature of the stim- ulus that impels a salmon to enter fresh water, and then fight his or her way upstream, remains a mystery. It is not known whether all the salmon move inshore in spring, or only those that are destined « Bulletin 21, Biol. Bd. Canada, 1931, p. 96. «• This was shown by Huntsman, Ann. Rep. Fish. Res. Bd. Canada (1947) 1948, p. 37. FISHES OF THE GULF OF MAINE 129 to spawn that year, plus a certain number of im- mature grilse that have passed 1 year at sea. And Dr. Huntsman87 has pointed out that the move- ment of the salmon riverward may be very slow; thus the salmon may take as much as a month to cover the 20-odd miles to the head of tide in the Petitcodiac River, while some of those that enter the estuary of the St. John River in autumn pass the winter there (probably in a lethargic state) before moving up to the head of tide 80 miles distant. In any case, only such fish as are ap- proaching sexual maturity (irrespective of age), and some immature female grilse, run far up into the rivers; all the others remain in salt water, or at most they do not run above the head of tide, as has often been remarked. The majority of the Gulf of Maine salmon be- come "river-mature" as it is called, long before the spawning season, for while none of them spawn before October, some of them enter fresh water as early as March and April. But the chief runs come later, varying in date, not only from river to river, but from year to year in a given river. In the Penobscot, some fish may enter in March ; they are to be expected in the lower reaches after the first week in April; more come in May, perhaps two- thirds in June, with a few fish entering later still. Available information is to the effect that few enter the Narraguagus and Dennys Rivers until well into May, the chief runs there coming in June, with some entering as late as September. We have not been able to obtain definite dates for the spring and early summer runs in the St. John River. But it seems that salmon continue to enter the latter until well into the autumn, judging from catches of fish so fat that they must have come in recently from the sea. Salmon enter other streams tribu- tary to the Bay of Fundy from May on. As a rule the large salmon come earliest, the grilse not until later, probably because it is not until later that the latter have reached the degree of fatness associated with river maturity. Accordingly, the heaviest runs in the Shubenacadie, mostly grilse (p. 130), are said to come from August until late in the autumn. Every salmon fisherman is familiar with the fact that salmon enter in "runs" that are spaced irreg- ularly in time, and that vary in date from year to year, depending on the height of water in the " Progress Report, Atlantic stations, Biol. Bd. Canada 8, 1933, p. 6; and unpublished notes. 210941 — 53 10 river and on the strength of the current. Freshets tend to bring them in; if the current becomes too strong they simply hold position, to breast it again as the flow slackens. The fish that are in the estuary remain there during the periods be- tween freshets, waiting, as it were, for the message from upstream that starts them on their way. And the salmon within the river are similarly quiescent during periods of low water and weak current. This is the chief reason why salmon angling is so uncertain a sport, even in the best of rivers A good deal of discussion has centered about the question whether the earliest fish stay in fresh water from then until spawning time (a matter of 6 months) or whether there is more or less move- ment in and out of the river mouths at the begin- ning of the season. The latter view may be cor- rect for the small streams, but it seems safe to say that after the run is well under way in late May or early June no fish return from fresh to salt water until autumn. Tagging experiments carried out in Canadian rivers have also yielded the very interesting information that no matter when a salmon runs upstream in one year, it may do so either early or late in the next.88 It is a matter of common knowledge that salmon average larger in some rivers than in others, and growth studies based on the scales have shown that these differences are due chiefly to the average ages of the fish that enter. In the St. John, as Huntsman has pointed out,89 there are three prin- cipal groups of salmon: (a) male grilse, averaging about 6 pounds, that are mature and fated to breed that same autumn; (b) the ordinary spawn- ers that have passed two years or more at sea; these average 10 to 15 pounds in weight and enter from May to August, the late comers running heavier than the early comers; most of them are virgins, but some of them have already spawned once or twice; (c) immature female grilse, averag- ing about 9 pounds, that enter from November to January. Few, however, return to spawn in the rivers of Maine until they have passed 2 years at sea; not more than 3 or 4 grilse to 70 adults were taken in the St. Croix, for example, when there still was a good run there, and not more than 1 '» Fifty-filth Annual Report of the Fisheries Branch, Department of Marine and Fisheries. Canada, (1921-22) 1922, p. 19. »■ Nature, vol. 141, 1938, p. 421; Pub. 8, Amer. Assoc. Adv. Sci , 1939, p. 34. 130 FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE grilse to 500 adults in the Dennys and Penobscot Kivers.90 The average weight of the salmon caught in the Penobscot was about UK pounds in 1905 (6,378 fish), 9 to 10 pounds in 1919 and 1920 (3,920 fish),91 or a little less than in the St. John. The heaviest Penobscot fish of which we found definite record of late years weighed a little more than 35 pounds.92 The fish in the rivers flowing into the head of the Bay of Fundy run much smaller, as Perley pointed out long ago, and recent studies show that most of them spawn first as grilse, i. e., after only one year at sea; a few, having spawned after one year at sea, return to spawn again a year later; and the percentage of larger and older fish is very small there. This, Huntsman points out,93 contrasts with the preva- lent 6-year-old fish in the Miramichi, which dis- charges into the southern side of the Gulf of St. Lawrence, and with 7- or 8-year-old fish in the Grand Cascapedia, tributary to the Bay of Chaleur. Various explanations have been ad- vanced to account for these differences from river to river, none of them convincing in our opinion. It also appears to be true (as often stated) that a larger proportion of the salmon are annual spawners in small streams, where most of the spent fish drop downstream again soon after spawning, than in large rivers where many of these "kelts" remain in fresh water over the winter. A plausible explanation is that kelts that return to the sea immediately after spawning have less opportunity to grow (though they recover condition sufficiently to spawn again the following summer) than such as await the spring to go downstream, and that spend a whole year at sea instead of one winter only between two suc- cesive spawnings. This, however, does not ac- count for the fact that it is almost invariably the large rivers that yield the very large maiden fish that have spent 4 years at sea, or more. Abundance. — The early extirpation of salmon from the Merrimac, Saco, Kennebec system, and various rivers to the eastward naturally resulted in a great decrease in the abundance of salmon » See Kendall (Mem. Boston Soc. Nat. Hist., vol. 0, No. J, 1935. pp. 58-60) for age determinations of Penobscot salmon. »i Radcliffe, Rept. U. S. Comm. Fish. (1921) 1922, p. 146. 'J Kendall. Mem. Boston Soc. Nat. Hist., vol. 9. No. 1, 1935, p. 32. •> Bull. 21. Biol. Board Canada. 1931. p. 19. in the open Gulf, clearly reflected in the catches. Data are not available for early years when all the rivers still offered free access. But the yearly catch had been reduced to about 100 to 500 fish in the St. Croix by about 1887; 200 to 1,000 each in the Dennys and in the Kennebec, and 5,000 to 15,000 in the Penobscot. The catch along the Maine coast, which had been a little more than 150,000 pounds in 1889 (more than seven-eighths of this in or about the approaches to the Penobscot), was only about 86,000 pounds in 1905 (of this 74,000 lb., or 6,378 fish from the Penobscot); was about 20,000 pounds in 1919 (13,557 lb. or 1,322 fish from the Penobscot); and was only 14,744 pounds (12,700 lb. or 1,221 fish from the Penobscot) in 1928. As 70 to 90 percent of the Maine catch comes from Penobscot River or Bay, the following table of salmon caught there in certain years from 1896 to 1928 is pertinent: Year Number offish Pounds Year Num ber offish Pouudi 1896- -.- 6, 404 80, 225 1918- ... 1,653 17, 212 1898- ... 3,225 42, 560 1919- --. 1,322 13, 557 1901. ... 6,821 86, 055 1920. ... 1,598 15, 135 1903_ ._- 4,859 67, 470 1928- -.. 1,221 12, 700 1905- --- 6,378 74, 158 The Maine catch then increased again to about 88,000 pounds in 1930 and to about 70,000 in 1931, suggesting a better run in the Penobscot, and var- ied between 16,000 and 40,000 pounds through the period 1932-1938.94 But the average reported catch for Maine for the period 1939 to 1947 9S was only about 3,600 pounds (maximum 9,300, min- imum 600), the average Massachusetts catch for the same period only about 100 pounds (maxi- mum 400, minimum 0). Thus the output of salmon from the rivers of Maine (none from the rivers of Massachusetts) has been only about one- fiftieth as great during the past few years as it was some 60 years ago. The numbers of salmon have held up much better in the Canadian waters of the Gulf, thanks to wise measures of conservation such as limiting netting at the mouths of the rivers, and keeping the streams free for access by fishways at the dams. The average yearly catches, from 1870 to 1946, " No data are available for 1934 or 1936. w No data for 1941. FISHES OF THE GULF OF MAINE 131 were as follows for the west coast of Nova Scotia and for the Bay of Fundy combined : Years Pounds 1870-1879 655,200 1880-1889 292,700 1890-1899 634,000 1900-1909 576,800 Years Pounds 1910-1919 540,000 1920-1929 470,300 1930-1939 424,000 1940-1946 278,000 The Canadian catch in the open Gulf and in the Bay of Fundy may be expected to run about 400,000 to 600,000 pounds at the present time, taking one year with another, or something like 40,000 to 60,000 fish, which is perhaps 100 times as great as that for the entire coastline of Maine and of Massachusetts. And the distribution of the catches shows that the St. John River contributes something like four-fifths of this, or a yearly aver- age of some 50,000 fish,8" contrasting with only a few hundred fish for the Penobscot in a poor year, and perhaps up to 8,000 in a good. Salmon anglers are only too familiar with the fact that the number of fish that enter even the best of salmon rivers is much smaller in some years than in others. During the 16-year period, 1931-1946, the commercial catches reported for St. John Harbor and St. John River (best salmon river tributary to the Gulf of Maine) were good in 1931 (164,000 lbs.); in 1935 (149,300 lbs.); in 1936 (148,600 lbs.); in 1937 (172,700 lbs.); and in 1943 (157,500 lbs.); but were poor in 1939 (48,500 lbs.); in 1945 (60,000 lbs.) and in 1946 (54,500 lbs.). The yearly average for this period was 116,000 pounds. '• Huntsman (Bull. 21, Biol. Board Canada, 1931) has made a very Inter- esting analysis of catches for the Bay of Fundy as a whole, as woll as for the St. John, for the Chignecto system, and for the Minas system, separately. In the Minas system the fishery produced as much as 383,800 pounds in 1907, 283,400 pounds in 1917, and 226,500 pounds in 1918; but since then, up to 1946, the best catches have been only 160,700 pounds in 1919, 165,100 pounds in 1923, and 143,300 pounds in 1925, while the poorest were 28,100 pounds in 1938 and 26,600 pounds in 1945. The average yearly catch from 1917 to 1930 was 133,000 pounds, and from 1931 to 1946, 48,000 pounds. The reader will notice at once that the big years have not been the same for these two bodies of salmon. It seems sufficiently established that yearly and regional differences, such as these, result in the main from corresponding differences in the numbers of smolts that reach salt water in any given year. And recent investigations in Canadian waters make it likely that the factor chiefly responsible is the height of the water from summer to summer, or over periods of several summers, which of course reflects the yearly vari- ations in rainfall. If the water is high the pan- arc protected from the birds that prey upon them and are more easily able to escape the trout, so that many survive to descend to the sea and to return one, two, or three years later. If the water in the river is low the parr are more at the mercy of kingfishers, megansers, and trout, so that fewer of them live to reach salt water, and there are fewer of them to return as grilse or as older fish. Humpback salmon Oncorhynchus gorbuscha (Walbaum) 1792 Jordan and Evermann, 1896-1900, p. 478. /'■■ "y^wrtfl =--^ ...i-.^-,^**.-' -- Figure 55. — Humpback salmon {Oncorhynchus gorbuscha). 132 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Description. — The humpback is of the familiar salmon outline while living in the sea, the body being deeper than thick, with rounded belly. The head is naked but the body is covered with scales large enough to be seen easily. The dorsal fin stands about midway of the body above the ven- trals, and the flaplikc adipose fin is over the rear end of the anal fin. It agrees so closely with the Atlantic salmon in all this that the one might easily be taken for the other, were it not that the anal fin of the humpback invariably has 14 rays or more, whereas that of the Atlantic sahnon has only about 9 rays. Also, the humpback is a stouter-bodied fish than the Atlantic salmon. The male humpback (like all the Pacific salmons, and the Atlantic salmon to a lesser degree) under- goes a very noticeable change in form in the spawning season, with the body deepening and developing a prominent hump in front of the dor- sal fin; the jaws elongating and becoming hooked at the tip and the teeth increasing in size. Color. — The back and tail of the humpback are bottle green with poorly defined black spots, while it is in the sea. These spots are particularly con- spicuous on the tail, where they are oval in outline and as much as a thud of an inch in longest diam- eter. These spots are one of the distinctive marks, whereby the humpback can be distinguished from all other salmons. The sides and belly are sil- very, with a faint pinkish tinge. Young hump- backs are unique among salmon in being of prac- tically adult coloration without "parr" marks (p. 122). Size. — The humpback is the smallest of the Pacific salmons and much smaller than the Atlan- tic salmon, adults averaging only about b){ pounds in weight and 20 to 25 inches in length. Males weigh to about 11 pounds and females to about 7% pounds. General range. — Pacific coast of North America and of northern Asia, from Oregon northward on the American side. This is the most abundant salmon in Alaska. It runs up fresh rivers to spawn, which it does but once and then dies. It has been introduced in the rivers of Maine. Occurrence in the Gulf of Maine. — The history of the introduction of this west coast salmon to New England waters is as follows: Humpback sahnon eggs seem first to have been planted in Maine rivers in 1906. In the autumn of 1913 a large consignment of humpback eggs was shipped to the Craig Brook and Green Lake (Maine) hatcheries, and the approximately 7,000,- 000 fingerlings hatched therefrom were distributed in the Penobscot, Androscoggin, Damariscotta, Dennys, Pleasant, Union, Medomak, Georges, and St. Croix Rivers. A year later some 5,000,000 young fish were liberated. A third plant was made in 1915; a fourth of 6,235,808 fingerlings in 1916; and a fifth of about 1,000,000 in the Dennys and Pembroke Rivers in 1917.97 The results of this attempt at acclimatization were first seen in the summer and fall of 1915 when fishermen reported large numbers of mature humpbacks along the Maine coast, and when humpbacks ran in the Dennys River (where many were caught) from August 15 until September 24, some probably spawning there, for the bodies of spent fish were seen drifting downstream. Hump- backs again entered the Pembroke and Dennys Rivers during August, September, and October of 1917 with a few reported from the Penobscot, St. Georges, Medomak, and St. Croix, the result of the plant of 1915. And at least 2,000 mature fish were seen that summer in the Dennys alone, where many were caught averaging about 5 pounds, and one as heavy as 10 pounds 9 ounces. Definite in- formation is lacking for 1918. But even larger numbers entered the Dennys and Pembroke Rivers in the autumn of 1919 than in 1917, with smaller runs in the Penobscot, Machias, St. Croix, and Medomak Rivers. Enough spawned that year in the Dennys and Pembroke Rivers for the fish- culturists of the Bureau of Fisheries to artificially fertilize half a million eggs there. And hump- backs were caught in the weirs in Passamoquoddy and Cobscook Bays during that season. Adult fish were taken again in the weirs in 1920,98 and one fish was caught in a weir as far from its native river as Lanesville, Mass. (near Cape Ann) " at some time during the summer of 1921. Large numbers of eggs were collected again from wild fish between 1922 and 1926, the resultant fry being returned to the Dennys and other rivers nearby. Artificial propagation was abandoned then, for it seemed that the species was estab- w More detailed accounts of these and successive plantings will he found in the annual reports of the U. S. Commissioner of Fisheries for the years 1914 to 19S8. " Reported catch, 'Washington County, Maine, 1920. 310 pounds. " It was forwarded to the Massachusetts Commissioners as reported by C. K. Grant of Olouccster. FISHES OF THE GULF OF MAINE 133 lished.1 But natural reproduction seems not to have been successful enough for the humpback to maintain itself in the few Maine rivers open to it, much less to increase in numbers, for very few have been reported since about 1926 or 1927, and none that we have heard of for some years past. Silver salmon Oncorhynchus kisutch (Walbaum) 1792 COHO SALMON Jordan and Evermann, 1896-1900, p. 480. Description. — The silver salmon resembles a rather stout Atlantic salmon closely in its general shape, also in the relative size and position of its fins, and in their shapes. But a safe morphologi- cal criterion for distinguishing the one from the other is that the silver always has at least 12 rays in its anal fin, and some of them have as many as 17, whereas most of the Atlantic salmons have only 8 or 9 anal rays, and never more than 10. The color is a help also, in this connection, for while a silver is silvery down its sides, like an Atlantic salmon, it is more closely sprinkled with small black spots along its back and on the upper part of its tail fin than is an Atlantic salmon. These spots, too, are always roundish or oval in a silver, never in the form of crosses. On the other hand, the black spots are much smaller and much less conspicuous on a silver salmon than oil a humpback, and the lower half of the tail fin, which is as conspicuously spotted as the upper half on a humpback, usually has no spots on a silver salmon. Size.- — -Up to 3 feet in length. General range, habits, and occurrence in the Gulj oj Maine. — The native range of the silver salmon is from northern California to northwestern Alaska, where it is an important food fish, and where anglers take many of them, both by trolling and by fly fishing. Like other Pacific salmons, it runs i Rept. of U. S. Comm. Fish. (1928), I't. 1, 1929. p. 379. up into fresh streams to spawn, dying thereafter. Most of the young remain about one year in fresh water, but a few do not move out to sea until they are in their third year. Most of them return to fresh water at the end of the third summer at sea, a few, however, by the end of the second summer in salt water, a few others not until the fourth summer. Our only reason for mentioning the silver salmon is that a plant of its fry and fingerlings that was made in the Duck Trap stream, tributary to the western side of Penobscot Bay, near Lincolnville, Maine, resulted in the return of 150 mature fish to Duck Trap stream in 1944, and perhaps of more of them. But nothing more was heard of them thereafter, and no returns have been re- ported up to this writing (Nov. 1, 1951) from other plants that were made in Maine waters 2 in 1948. THE SMELTS. FAMILY OSMERIDAE The smelts are small salmons in all essential respects, except that their stomach has few pyloric caecae, or none, whereas there are large numbers of such caecae in their larger relatives of the salmon family. However, it is not necessary to look so deeply to learn whether a fish be smelt or very young salmon, for the former all have pointed noses and are very slender, whereas the young of our four salt-water salmons -humpback, silver Atlantic, and sea trout — are much stouter, with blunt noses. In most cases, too, the shape of the tail would suffice of itself to separate smelt from salmon smolt, for it is never as deeply forked in the latter as in the smelts. Two smelt fishes occur in the Gulf of Maine: the smelt (very common), and the capelin (a sporadic visitor from the north). The argentine (p. 139) is so closely related to the smelts that it is included in the following key. >In Tunk stream, Duck Trap stream, Chandler River, and Bald Hill Cove Brook. 2. KEY TO GULF OF MAINE SMELTS AND ARGENTINES The dorsal fin originates over the tips of the pectorals; the mouth is very small Argentine, p. 139 The dorsal fin situated far behind the pectorals; the mouth is large 2 Upper jaw almost as long as lower; teeth large; there is a group of strong fangs on the tongue; the pectoral fins have 12 rays or fewer Smelt, p. 135 Lower jaw much longer than upper; teeth so small as hardly to be visible; no fangs on tongue; the pectoral fins have 15 to 20 rays Capelin, p. 134 134 FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE Capelin Mallotus villosus (M tiller) 1777 Jordan and Evermann, 1896-1900, p. 520. Description. — The capelin is an even slenderer fish than the smelt, its body being only about one- sixth to one-seventh as deep and about one- twelfth as thick as it is long, and of nearly uniform depth from gill cover to anal fin (except in the case of females when their abdomens are distended with spawn) , whereas the smelt is usually deepest about its mid-length (at least if the fish is fat), which gives the two species characteristically different aspects. The head of the capelin is pointed like that of the smelt, the mouth gaping back to below the center of the very large eye with the tip of the lower jaw projecting noticeably be- yond the upper. The scales are minute, much smaller than those of the smelt and more numerous (about 200 per row on the sides of the body) ; the teeth so small as to be hardly visible to the naked eye, and the tongue fangs, so characteristic of the smelt (p. 135), are lacking here. The outline of the adipose fin likewise helps separate capelin from smelt, for it is low in the former and about half as long as the anal, but short and high in the latter. The pectoral of the capelin is broader also, usually with 15 or more rays. The capelin exhibits a pronounced sexual dimorphism; the male has much the longer pectoral fins; and the base of his anal is elevated on a pro- nounced hump, whereas it follows the general out- line of the belly in the female. In males, too, the scales in one of the longitudinal rows immediately above the lateral line, and in another row along each side of the belly, are pointed, distinctly larger than the other scales, and become longer still at spawning time when each pushes up the skin as a finger-like process; they form four ridges that are very evident when the fish is held in the hand. Color. — The capelin is transparent olive to bottle green above, like a smelt, but its sides are uni- formly silvery below the lateral line and the scales are dotted at the margins with minute dusky specks (in the smelt there is a distinct silvery band on each side) ; the belly is white. Back and head darken at spawning time. Size. — Few capelin are more than 6% to 7K inches long. Habits} — Capelin are most in evidence during the spawning season, when they come inshore in multitudes along arctic-subarctic coasts. They spawn on gravel or pebbly bottom, chiefly close below tideline, many of them in the wash of the waves in the beach; many are stranded then on the beach between waves. But eggs have also been reported from as deep as 35 to 40 fathoms. Each female while spawning is accompanied by two males that crowd her between them; but she may have only one companion.4 Spawn- ing takes place chiefly at temperatures of 43° to 50° F. (6°-10° C.) and more actively by night than by day. The eggs are reddish, about 1/25-inch (1 mm.) in diameter, and so sticky that they cling to each other like herring eggs, and to the gravel and pebbles with which they are intermingled by the swash of the waves. They hatch in about 15 days at a temperature of 50° F. (10° C). And they will tolerate a salinity as low as 7 per mille, i Interesting accounts of the habits of the capelin and of its rate of growth in Newfoundland waters have been given recently by JefTers (Ann. Rept. Biol. Board Canada (1930), 1931, pp. 7-18); by Sleggs (Rept. Newfoundland Fish. Res. Comm., 1, No. 3, 1933); and by Templeman (Bull. Newfoundland Government Lab.. 17 (Research), 1948. » According to Lanman, Rept. U. S. Comm. Fish. (1S72-187.1.) 1874, p. 225. *&>& Figure 56. — Capelin (Mallotus villosus), Grosswater Bay specimen. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 135 as Dr. Jeffers writes us. The larvae, described as 5 to 7 mm. long at hatching, are very slender and resemble those of smelt, herring, and launce so closely that identification is a matter for the expert. In any case, capelin are encountered so seldom in our Gulf that their larvae are not apt to be seen there.6 Along the coasts of Newfoundland, capelin spawn chiefly in June and July, and we have found them doing so in multitudes along the outer Labrador coast in July. Probably any spawning that may take place within the limits of our Gulf would fall in May at latest, to judge from water temperatures. The capelin so seldom appears in the Gulf of Maine that we need only add that it is a fish of the high seas frequently encountered far out from land, coming inshore only to spawn and then as a rule moving out again; that it travels in vast schools at spawning time (when it often strands on the beach in countless multitudes). It is the chief bait fish of Arctic seas, preyed upon by whales and by every predaceous fish, particularly by cod, which are often seen pursuing the capelin at the surface in northern waters. Capelin themselves feed chiefly on small crustaceans, particularly on copepods, on euphausiid shrimps, and on am- phipods. It is also known to devour its own eggs. We can bear witness that the capelin is a delicious little fish on the table. General range. — Boreal-Arctic seas, south to the coast of Maine8 on the Atlantic coast of America. Occurence in the Gulf of Maine. — The capelin is a sub-Arctic fish that visits the Gulf of Maine occasionally; chiefly the eastern side as might be expected since it comes from the north. Dr. Hunstman writes: 7 In the Gulf of St. Lawrence it occurs abundantly in limited areas, which shift somewhat from year to year. It occurs periodically in similar limited areas farther south. The southeastern corner of Cape Breton is the center of such an area, where large quantities were taken in 1917. Halifax is the center of another area, where, however, it is more rare. In 1916 it was abundant at Sambro, near Halifax. The next area is in the Bay of Fundy, where they have, exceptionally, been taken in large quantities at long intervals. 'Templeman (Bull. Newfoundland Government Lab., 17, Res., 1948t figs. 18-20) cives a series of excellent illustrations of capelin larvae at different stages of growth. 8 According to Jordan and Evermarn the capelin finds its southern limit at Cape Cod, but we find no actual records of its occurrence farther *outh than is mentioned. ' Quoted from it letter. Apparently a period of this sort occurred about the middle of the past century, for Perley, writing in 1852, reported it from a number of points in the neighborhood of St. John, New Brunswick. It seems then to have disappeared from the Gulf of Maine, not to reappear until 1903 when it was common in the Bay of Fundy in May. A few were again taken off Passamaquoddy Bay in that same month of 1915.8 And this was the prelude to a period of local abundance, for capelin were noticed among the herring in the weirs of the Passamaquoddy Bay region in October 1916, becoming so plentiful by the end of November that one catch of 3,000 pounds of fish consisted of 2,000 pounds of capelin and only 1,000 of herring. They were also reported at various localities along the New Brunswick coast at that time. Probably they persisted locally in the Bay of Fundy through- out the winter of 1916-1917, for large numbers of capelin appeared in Minas Basin in the following May and June. We find no record of capelin within the limits of the bay in 1918, but they were taken again in 1919 in 50 fathoms of water off Passamaquoddy Bay in January, February, and March, and they appeared with smelts a month later as far west as the Penobscot River, pene- trating far inland. None, however, have been seen in the Gulf of Maine since then, so far as we have been able to learn. Smelt Osmerus mordax (Mit chill) 1815 Salt-water smelt Jordan and Evermann, 1896-1900, p. 523. Description. — The smelt is distinguishable from all other fish common in our waters by its slender form, combined with a long pointed head, large mouth, a small but evident adipose fin standing above the rear part of the anal, and a deeply forked tail. The location of its dorsal fin above the ventrals instead of in front of them, and its much larger mouth and small eye separate it from the argentine. The large, fang-like teeth on the smelt's tongue, its larger scales (of which there are about 75 along each row on the sides, all alike in the two sexes), its shorter adipose fin, its nar- rower pectoral fins, that its lower jaw projects only slightly beyond the upper and its scales slip off very easily, obviate any danger of confusing ' Huntsman (Contrib. Canadian Biol., (1921) 1922, p. 50) and Kendall (Copela, No. 42, 1917, pp. 28-30; and Copeia, No. 73, 1919, pp. 70-71) give details. 136 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE ■ wmm j^m*^ >■- Figure 57. — Smelt (Osmerus mordax), adult, Woods Hole. From Jordan and Evermann. Drawing by H. L. Todd. it with the capelin. The body of the smelt is only about one-fifth as deep as long (exclusive of caudal fin), with broadly rounded back but flat- tened enough sidewise to be egg-shaped in cross section. It is deepest about its mid-length, taper- ing thence toward the head and toward the tail (at least in fat fish), whereas the capelin is of nearly uniform depth from gill opening to anal fin (p. 134) . Its mouth gapes back of the eye. Printed accounts of the smelt usually credit it with a peculiar "cucumber" odor, and smelt fishermen often speak of a trace of this, but it is so faint that we have never noticed it though we have caught and handled many.9 Color.- — Transparent olive to bottle green above, the sides are of paler cast of the same hue but each with a broad longitudinal silvery band. The belly is silvery, while the fins and body are more or less flecked with tiny dusky dots. This color pattern is shared by another slender little fish, the silver- side (p. 302), but the latter has two large dorsal fins, so there is no danger of confusing the smelt with it. Size. — Smelt grow to a maximum length of about 13 or 14 inches. Few, however, are more than a foot long, and adults run only about 7 to 9 inches. Smelt weigh from 1 to 6 ounces accord- ing to size and fatness. Habits.10 — The smelt is an inshore fish, con- fined to so narrow a zone along the coast that none has ever been reported more than a mile or so out from the land, or more than two or three fathoms in depth, while many spend the whole year in estuarine situations. * The European smelt (0. eperlanus) smells so strong that It is not held In very high esteem as a food-fish. >° Kendall (Bull. U. 8. Bur. Fish., vol. 42, 1927, p. 244) has given a detailed account of the habits, distribution, and catches of the smelt of the New England coast, ako of the landlocked populations. Young smelts certainly, and old ones probably, travel in schools, which are mostly composed of fish of a size, hence probably are the product of one year's hatching, and they live pelagic, not on the bottom, though confined to shoal water. Most authorities describe the smelt as feeding on small crustaceans, which is correct as far as it goes, for shrimp (both decapods and mysids) and gammarids are probably its favorite food, and shrimp were long considered the best smelt bait. But it has been found that pieces of "sea worms" (Nereis) are more attractive to the larger smelt (at least in Massachusetts Bay). Small fish also form an important item in the diet. We have, for example, found smelts taken in the Sheepscot River in May packed full of young herring, and have caught many with small mummichogs (Fundulus) as bait; while cunners, anchovies, launce, sticklebacks, silversides, and alewives have been identified from smelt stomachs at Woods Hole. The Woods Hole diet list also includes shellfish, squid, annelid worms (Nereis), and crabs, but even as greedy a fish as the smelt ceases to feed during its spawning visits to fresh water. Young smelt depend chiefly on copepods and on other minute pelagic crustaceans. Smelt fisher- men are familiar with the fact that a smelt ap- proaches a bait slowly, then stops, and appears to suck it in.11 If the smelt take their living prey in this same way, it is somewhat of a mystery how they succeed in capturing animals as active as shrimps and small fish. Smelt, like alewives, shad, and salmon, make their growth in salt water, but run up into fresh water to spawn. The summer habitat of the smelt varies off n This method of feeding seems first to have been described In print by "Grif" (Forest and Stream, vol. 54, No. 8, Feb. 24, 1900, p. 151). FISHES OF THE GULF OF MAINE 137 different parts of the coast of the Gulf, depending on the summer temperature of the water and perhaps on the food supply. Most of them desert the harbors and estuaries of the Massachusetts Bay region and along the southern coast of Maine during the warmest season. But it is probable that they move out only far enough to find cooler water at a slightly greater depth, and a few may be found in harbors through the summer. Smelt, for instance, are caught then in Cohasset Harbor in some years, but not in others; and east of Penob- scot Bay, where the surface temperature does not rise so high as off Massachusetts, smelt are to be found in the harbors, bays, and river mouths all summer, and are sometimes taken in numbers then in the weirs.12 Adult smelt gather in harbors and brackNIi estuaries early in autumn, where smelt fishing with hook and line is in full swing by October. The schools then tend to move into the smaller harbors on the flood tide, and out again on the ebb, especially if the tidal current is strong, as it is in Cohasset, a locality with which we are famil- iar. But some smelt remain over the ebb in the deeper basins. And some of them have run as far as the head of tide by the tune the first ice forms in December. Most of them winter be- tween the harbor mouths and the brackish water farther up; the maturing fish commence their spawning migration into fresh water as early in the spring as the ice goes out of the streams and the water warms to the required degree. Temperature observations by the Massachusetts Commission show that the first smelt appear on the spawning beds in Weir River, a stream empty- ing into Boston Harbor, when the temperature of the water rises to about 40-42°. 13 This may take place as early as the first week in March or as late as the last, about Massachusetts Buy, depend- ing on the forwardness of the season and on the particular stream. The chief production of eggs takes place in temperatures of 50-57°, and spawn- ing is completed in Massachusetts waters by about the 10th or 15th of May, year in and year out. East of Portland, smelt seldom commence to run before April, and continue through May. In the colder streams on the southern shores of the Gulf of St. Lawrence they do not spawn until June. On the other hand, they may commence spawn- ing as early as February along the southern New England coast west of Cape Cod. As a rule smelt do not journey far upstream; many, indeed, go only a few hundred yards above tidewater, whether the stream be small or large. Thus Dr. Huntsman informs us that the smelt that enter the estuary of the Stewiacke River, Nova Scotia (a tributary of the lower Shubenacadie, near the head of the Bay of Fundy) spawn only in the tidal part. And some spawn in slightly brackish water in certain ponds back of barrier beaches (e. g., Straits Pond, Cohasset, Mass.). But flooding with salt water, wThich sometimes happens, kills the eggs. The adult smelts return to salt water immedi- ately after spawning to spend the summer either in the estuary into which the stream in which they spawn empties or in the sea close by. On the Massachusetts coast north of Cape Cod all the spent fish have left fresh water by the middle of May, earlier in some years. On the Maine coast, too, a good proportion of the spent fish are in salt water by the first weeks in May; thus we have seen a bushel of large smelt taken in a weir at Cutler (near the mouth of the Grand Manan Channel) as early as May 4. The eggs average about 1.2 mm. (0.05-inch) in diameter and they sink to the bottom, where they stick in clusters to pebbles, to each other, or to any stick, root, grass, or water weed they chance to touch. According to the Manual of Fish Culture a female weighing as little as 2 ounces will produce between 40,000 and 50,000 eggs;14 The eggs of the closely allied European smelt (Osmerus eperlanus) hatch in 8 to 27 days, accord- ing to temperature, and the incubation period of the American fish is the same, probably, for smelt eggs are reported as hatching in 13 days at the Palmer (Mass.) hatchery. 'i Atkins (Fish. Ind. U. S„ sect. 5, vol. 1, 1SS7, pp. 690-693) gives much information on the smelt in Maine. 1 Kendall (Dull. U. S. Bur. Fish., vol. 42, 1027, pp. 231-23!) summarizes these observations and cives additions] information for streams on the coast of Maine. Figure 58. — Smelt larva, 26 mm. The smelt has proved a favorable fish for artificial hatching and large numbers of fry are so produced yearly in Massachusetts, the eggs being M Kept. U. S. Fish Comm., 1897, p. 188. 138 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE taken in Weir River, just mentioned, and it has proved possible to re-establish smelt by intro- ducing the eggs or fry into streams from which it has been extirpated. For example, good smelt fishing was reported in "Poorhouse Brook," Saugus, a tributary of Boston Harbor, three years after the stream was stocked with eggs, and attempts have been similarly successful on Long Island, N. Y. Maintenance of the stock is a question either of providing accessible spawning grounds of sufficient extent, or of making up for lack of such by artificial propagation. The precise season when young smelt go down to the sea in the Gulf of Maine streams is yet to be learned; probably early in summer. We seined several hundred fry, 1% to 1% inches long, October 1, 1924, on a beach of Mount Desert Island, evidence that the rate of growth is about the same for our smelt during its first summer and autumn as for the European, i. e., to a length of 1% to 2% inches. Most of the smelt evidently do not spawn until they have passed an autumn, a winter, a summer, and a second winter in salt water. General range. — East coast of North America from eastern Labrador, Strait of Belle Isle, and the Gulf of St. Lawrence southward regularly to New Jersey, and reported to Virginia; running up streams and rivers to spawn. Smelt, also, are landlocked naturally in many lakes and ponds in New Hampshire and in Maine, also in Lake Champlain, and in various Canadian lakes.18 Occurrence in the Gulf of Maine. — The smelt is a familiar little fish around the entire coast of the Gulf of Maine, but varies greatly in abundance from place to place according to the accessibility of streams suitable for spawning, from which it seldom wanders far alongshore. Smelt are plenti- ful, still, all around the inner parts of Massachu- setts Bay and its tributary harbors, though many of the local streams are barred to them now; thence northward and eastward all along the coast of Maine; tolerably so in the region of Passama- quoddy Bay (catch for Charlotte County, New Brunswick, 7,400 pounds in 1945), and more so along the western shore of Nova Scotia (60,100 pounds for Yarmouth County in 1945). But they are less plentiful passing inward along the Nova Scotia shore of the Bay of Fundy, as illus- '■ The European smolt is landlocked in many lakes in northern Europe. trated by catches in 1945 of 20,100 pounds for Digby County, but only 7,600 pounds for Kings County, 2,000 pounds for Hants, and 1,800 for Colchester (covering the Minas Basin region). So few smelt exist along the New Brunswick side of the Bay, inward from the Passamaquoddy region, that none at all were reported for that stretch of coast in any year during the period 1939-1945. Doubtless this scarcity up the Bay is "due to absence of streams suitable for spawn- ing, and the general turbidity of the water," as Jeffers has remarked.16 Abundance. — Smelt once were so plentiful in the Back Bay at Boston (now mostly filled in) that "distinguished merchants of lower Beacon Street might be seen, at early hours, eagerly catching their breakfast from their back doors."17 Those happy days, however, are long since past, and smelt certainly are not so numerous as they were even 50 years ago,18 around the Massachusetts shoreline of our Gulf, where various streams either have been closed to them, or have been rendered uninhabitable by pollution. But enough still remain to provide sport for thousands of anglers,19 and we still hear of an occasional catch there of many dozens by some one lucky enough to hit a run of fish at the right time and tide. In 1938, when a special effort seems to have been made to gather smelt statistics, the reported catch for the inner part of Massachusetts Bay and northward to the New Hampshire line was 25,900 pounds, or some 300,000 fish, if they ran about a dozen to the pound. The yearly catch reported for the coast of Maine, added to that of the Passamaquoddy area (which form one faunal unit so far as the smelt is concerned) averaged about 644,000 pounds during the period 1937 to 1946,20 or perhaps some 8,000,000 fish; about 61,000 pounds for Digby and Yarmouth Counties, Nova Scotia, combined, which covers most of the catch for the Gulf, north and east of New Hamp- shire. The catches of smelt that are made along the coasts of Maine, New Brunswick, and Nova Scotia may seem impressive if taken by them- '• Ann. Rept. Biol. Board Canada, (1931) 1932. p. 27. >' Mass. Rept. for 1870, p. 23. » Kendall (Bull., U. S. Bur. Fish., vol. 42. 1927, pp. 244-249) gives many interesting details as to catches in Massachusetts. >* Smelt fishing has long been restricted to hook and line along this part of the coast. » Maximum 875,700 pounds in 1945, minimum 316,400 pounds in 1939. No data are available for Maine for the years 1941 or 1942. FISHES OF THE GULF OF MAINE 139 selves. But Miramichi Bay, alone, on the south- ern shore of the Gulf of St. Lawrence yields yearly between three and four times as much smelt as does the entire coastline of the Gulf of Maine.21 Catch records do not suggest any striking alteration in the abundance of smelts during the past 10 years or so for Maine or for the Canadian shores of the Gulf. But they seem to have been somewhat more plentiful along the Maine coast previous to the early 1900's, for catches of 1,125,268 to 1,279,550 pounds there in 1887, 1888, and 1902 have not been equaled since then, the nearest approach being 968,300 pounds in 1945. We are often asked what effect the disappear- ance of the eel grass (Zostera) from our coasts has had on the abundance of the smelt. Unfor- tunately, the statistics of the yearly catch do not yield any clear answer. Neither can we offer any convincing explanation for the violent fluctuations that take place from year to year in the abundance (or availability?) of smelts at one point or another. Fishermen report, for example, that they were far >< Average reported catch for Northumberland County, 1937-1946, was 2.268,030 pounds. less plentiful in Massachusetts Bay and in the Great Bay region, N. H., in 1950 than they were in either of the two previous years. The smelt also has a great recreational value, smelt fishing being a favorite pastime for home consumption. As many as 2,326 people, for in- stance, have been counted fishing at one time about Houghs Neck in Boston Harbor, and this same sort of thing is to be seen up and down the Massa- chusetts coast in harbors and stream mouths in autumn. Many smelt are caught in Great Bay, N. H., in good years, through the ice for the most part. And this applies equally to many localities along the coast of Maine. So plentiful are the fish on occasion and so greedily do they bite, especially on the flood tide, that it is usual to number the catch about Massachusetts Bay by the dozens rather than by the individual fish. Sea worms (Nereis) are generally considered the best bait, especially for the larger smelt, shrimp the second best, small minnows or clams a poor third. Smelt have also been taken with a small red artificial fly in the Gulf of St. Lawrence, and perhaps elsewhere. THE ARGENTINES FAMILY ARGENTINIDAE The argentincs resemble the smelts in most of their external characters. But their mouths are much smaller, with the upper jawbone reaching back only about even with the front of the eye, and the entire base of their rayed dorsal fin is in front of the ventral fins. Argentine Argentina situs Ascanius 1763 Herring smelt Jordan and Evermann, 1896-1900, p. 526. Description.- — The argentine has a pointed nose, deeply forked tail, and slender, compressed body, but it has much larger eyes than either smelt or capelin, a character no doubt associated with its deep-water home; its mouth is much smaller, not gaping back even as far as the eye; and its dorsal fin stands wholly in front of the ventrals, instead of above them as it does in both the smelt and the capelin.22 "The anatomy of Argentina situs Is described, and records along the American coast are given by Kendall and Crawford (Jour. Washington Acad. Scl., vol. 12, No. 1, January 1922, pp. 8-19). The body of the argentine (about one-fifth as deep as long) tapers toward both head and tail, but its sides are so flat, and its back and belly so broad, that it is nearly rectangular in cross section instead of oval. Its scales, too, are larger than those of the smelt, there being otdy 60 to 70 rows along the lateral line. Its adipose fin is very small and its jaws are toothless, though its palate and tongue are armed with small teeth. Color. — The color of the adult is variously de- scribed by different authors. All agree, however, that the back is brownish or olivaceous, the sides silvery or with iridescent golden or brassy luster, and the belly white. The adipose fin is yellowish. Size. — The argentine is a larger fish than the smelt or the capelin, growing to a length of about 18 inches. Habits. — Nothing is known of the fife of the herring smelt in our Gulf, and little enough is known of it in Scandinavian waters, where it is sometimes caught on deep set-lines baited with herring or mussels, and where it is occasionally 140 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 59. — Argentine (Argentina silus). A, adult, Biddeford Pool, Maine; from Goode and Bean, drawing by H. L. Todd; B, egg; C, larva, 28 mm.; D, larva, 45 mm. B-D, European, after Schmidt. swept up to the surface like other deep-sea fishes by some upwelling of the water, to drift there helplessly. Its eggs float chiefly in the deeper water layers, seldom rising to the surface, and they are among the largest of buoyant fish eggs (3 to 3.5 mm. in diameter), with flat oil globule (0.95 to 1.16 mm.) and vacuolated yolk. Newly hatched larvae are about 7.5 mm. long and have a large yolk sac, but this has been absorbed when they have grown to a length of 12 mm. and a line of spots has appeared along the belly. The fin rays are formed by the time the little fish has reached 45 mm., the anus has moved forward, and the forked outline of the tail is apparent, but the ventral fins do not appear until the larva is about 50 mm. long. General range. — North Atlantic, usually in water as deep as SO to 300 fathoms ; known from northern Norway south to the northern part of the North Sea on the European side, from the Nova Scotia Banks to the offing of southern New England on the American side.23 Occurrence in the Gulj oj Maine. — The argentine was considered rare in our waters until recent- ly. Some specimens have been brought in from H For recent records of argentines off Nova Scotia, see McKenzie and Homans. Proc. Nova Scotia Inst. Sci., vol. 19, No. 3, 1938, p. 277 and McKenzie, Proc. Nova Scotia Inst. Sci., 20, 1939, p. 15. widely scattered localities around the coast, name- ly, Belfast, Biddeford Pool, and Fletchers Neck, Maine; and from Hampton Beach, N. H. It has proved, with the development of otter trawling, that argentines are fairly common all around the edges of Georges Bank and off Cape Cod in mod- erately deep water. It is not unusual for one haul of the trawl to bring in from one to a dozen from depths of 30 to 100 fathoms, with much larger numbers taken occasionally; one vessel, for exam- ple, trawled 15,000 pounds on the northeastern edge of Georges Bank in about 100 fathoms during a week in mid-September 1929. Evidently there are at least a few argentines in the deep trough of the Gulf also. Firth'24 reports that ten were taken at 90 fathoms on the northwestern slope of Georges Bank on June 18; and the Albatross II trawled one at 1 15 fathoms off Mount Desert Rock. They spawn to some extent in the Gulf, for on April 17, 1920, a townet haul on the Albatross I from 109 fathoms in the southeastern part of the Gulf basin yielded 43 eggs, unmistakably of argentine par- entage, while we have taken a scattering of argen- tine fry at localities as widely separated as the offing of Mount Desert Rock and the northwestern edge of Browns Bank. « Firth, Bull. Boston Soc. Nat. Hist., fil, 1910, p. 10. FISHES OF THE GULF OF MAINE 141 LUMINESCENT FISHES FAMILIES MYCTOPHIDAE, MAUROLICIDAE, CHAULIODONTIDAE, GONOSTOMIDAE, STOMIATIDAE, AND STERNOPTYCHIDAE These families include a heterogeneous assem- blage of small oceanic fishes, that are primitive in some respects, but are highly specialized in others for existence in mid-depths, on the high seas. They all have light-producing organs, which no other Gulf of Maine fish has; this is the only reason why we group them together here. 6. 7. KEY TO GULF OF MAINE LUMINESCENT FISHES Trunk at least H as deep as it is long from tip of snout to base of tail fin; front part of rayed dorsal fin is a hard triangular plate, supported by 7 or 8 spines Hatchet fish, p. 149 Trunk less than % as deep as it is long from tip of snout to base of tail fin; rayed dorsal fin does not commence with a hard plate or hard spines 2 Mouth does not gape back as far as the eye Pearl sides, p. 144 Mouth gapes back beyond the eye 3 No barbel on the chin; the ventral fins are about mid- way of the trunk; origin of rayed dorsal fin either in front of mid- length of trunk or at least not much behind it 4 There is a long fleshy barbel on the chin; the ventral fins are considerably behind the mid-length of the trunk; the rayed dorsal fin is far back, close to the tail fin 7 The rayed dorsal fin is far in advance of the ventrals; the jaws are armed with long and conspicuous fan gs Viperfish , p. 145 The rayed dorsal fin is about over the ventral fins (it may be a little in front of them or a little behind) ; the teeth are small 5 Eyes very small; no adipose fin behind the rayed dorsal fin; anal fin reaches nearly to the base of the caudal Cy clothone, p. 146 Eyes very large; there is an adipose fin behind the rayed dorsal; there is a considerable interspace between the rear end of the anal fi n and the origin of the tail fin 6 There are 3 or 4 separate luminescent dots at the base of the caudal fin; the Gulf of Maine species has a large lumines- cent patch on the snout Headlight fish, p. 142 There are only 2 separate luminous dots at the base of the caudal fin; the snout does not have a large luminescent patch Lantemfish, p. 143 The point of origin of the anal fin is in advance of the origin of the rayed dorsal fin by a distance about as long as the diameter of the eye; the tip of the chin barbel is distinctly swollen as well as bearing several filaments Stomioides, p. 147 The point of origin of the anal fin is not in advance of the rayed dorsal fin , the tip of the chin barbel is not swollen.. 8 Each side has only about 68 luminescent spots; then' is a large luminescent patch crossing the top of the cheek, behind the eye; the point of origin of rayed dorsal fin is in advance of origin of anal fin by a distance about as long as the diameter of the eye; the tip of the lower jaw does not enclose the tip of the upper jaw when the mouth is closed.. Trigonolampa, p. 148 Each side has about 85 luminescent spots; the side of the cheek behind the eye does not have a large luminescent patch; the point of origin of rayed dorsal fin is about over origin of anal fin; the tip of the lower jaw encloses the tip of the upper jaw when the mouth is closed Stomias, p. 147 LANTERN FISHES. FAMILY MYCTOPHIDAE The most distinctive external characters of the lanternfishes are their large eyes (situated close to the tip of the blunt snout), wide mouths gaping back beyond the eye, one soft-rayed dorsal fin, a deeply forked tail, and the. presence of a series of luminous organs as conspicuous pale spots along the sides. Some of them have an adipose fin on the back behind the dorsal fin, but others lack this. When present, this fin is so small and fragile that it is apt to be destroyed by the rough treatment the fish receive in the tow net in which they are taken. They most nearly resemble the anchovy (p. 118), the pearlsides (p. 144), and the cyclothone (p. 146) among Gulf of Maine fishes; but they are readily distinguished from the first of these by the presence of luminous organs and by the fact that the snout does not project beyond the mouth; from the second by their much wider mouths; and from the third by their much larger eyes. 142 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE They are among the most numerous fishes on the high seas, where they live at a considerable depth by day but often rise to the surface at night. Only two species of the group, representing as many genera {Diaphus and Myctophum), have been recorded within the Gulf of Maine. But each of these genera includes a considerable number of species that are common along the continental slope abreast of the Gulf, hence are as likely to stray into the latter as are the two that have actually been found there. And this applies equally to various other genera of lantern- fishes. The species of Diaphus and of Myctophum all resemble one another in general appearance, in having a short dorsal fin, with an adipose fin behiod it; a deeply forked tail; large eyes; wide, oblique mouth; and numerous luminous organs along the sides; all, too, are blackishsilvery in color. The members of each genus are separable only by differences in the arrangement of the luminous organs. Hence, positive identi- fication of a given specimen calls for the services of a specialist in the group. Should a lanternfish be taken in the Gulf in which the arrangement of luminous organs does not agree precisely with the two described here, we suggest that it be sub- mitted to the U. S. Fish and Wildlife Service to be named.25 *> Parr (Bull. Bingham Oceanng. Coll., vol. 3, art. 3, 1928), and Tanlng (Vidensk. Meddel., Dansk Naturhlst. Forening, vol. 86, p. 49, 1928) have recently published critical synopses of the lanternfishes. Headlight fish Diaphus effulgens (Goode and Bean) 1895 Jordan and Evermann, 1896-1900, p. 566. Description. — This curious little fish is separable from the lanternfish (p. 143) and from the pearlsides (p. 144) at a glance, by the large and very noticeable luminescent patch that covers the entire tip of its snout (including the anterior margin of the orbit) and that extends down over the edge of the upper jaw, a structure that has no parallel in any other fish regularly inhabiting the Gulf of Maine. It also differs from the pearlsides in its much more deeply cleft mouth, its even larger eyes, in the more convex dorsal profile of its head, and in lack- ing the regular horizontal row of luminescent spots along each side about at the level of the pectoral fin, that are conspicuous on the pearl- sides.26 The four separate luminescent spots at the base of the tail (besides the organ on its snout) separate it from its close relatives of the genus Myctophum (p. 143). The arrangement of the fins (all of which are soft, the dorsal with about 15 rays, the anal with about 16), is essentially the same as in the latter, and in the pearlsides; the caudal fin is more deeply forked than in the pearlsides, the adipose fin proportionately shorter. Color. — The color has not been described. Probably it is black, overlaid more or less with » The structures along the lateral line shown here on the Illustration of the headlight fish are large scales, not luminescent organs. Figure 60.— Headlight fish (Diaphus effulgens), Browns Bank. From Goode and Bean. Drawing by A. H. Baldwin. FISHES OF THE GULF OF MAINE 143 silver, with the luminescent organs pale blue or green. Size. — The specimens from which this species was originally described seem to have been about 7 inches long.27 General range and occurrence in the Gulf of Maine. — This oceanic species is only a stray within the limits of the Gulf. One specimen has been found in the stomach of a cod caught on Browns Bank,28 and another, also from a cod stomach, has been reported on Western Bank off the outer coast of Nova Scotia.29 Lanternfish Myctophum affine (Liitken) 1892 Jordan and Evermann, 1896-1900, p. 570. Description. — The most noticeable features of this little oceanic fish are its silvery black color, the luminous dots along its sides, its enormous eye situated close to the tip of the snout, its very deep oblique mouth, and its deeply forked tail. The anal fin is mostly or wholly behind the short, soft dorsal, and there is an adipose fin behind the latter, as in the headlightfish (p. 142). The longer snout and smaller mouth of Myctophum, with the fact that the luminous organs on its snout are in the form of small dots instead of a large patch covering the entire tip of the snout, are the readiest field marks to distinguish it from the latter. The dorsal profile of the head is much arched, the body moderately flattened sidewise, tapering gently backward to the rather deep caudal pe- duncle. The location of the luminescent spots is shown in the drawing (fig. 61). Color. — This lanternfish is silvery when alive, the silver underlain on the back with deep brown- ish black, the sides below the lateral line, and the belly varying (below the silver) from dark brown to dusky gray, or even to white finely dotted with gray. The luminescent organs are pale green or blue. Size. — All members of the genus Myctophum are small; a little more than %}% inches (89 mm.) is the maximum length recorded for this particular species. General range. — All the species of this genus are oceanic, occurring only as strays inside the edge of the continent. Occurrence in the Gulf of Maine. — Goode and Bean30 report the capture of this lanternfish over the southeast slope of Browns Bank (lat. 42° 21' N., long. 65° 07' W.) at 104 fathoms, which still remains the only record for it in the Gulf of Maine,31 or for any Myctophum for that matter. Figure 61. — Lanternfish (Myctophum affine). After Parr. PEARLSIDES. FAMILY MAUROLICIDAE The Pearlsides resembles the lanternfishes (p. 141) in shape of body, but it has a shorter rayed * The Illustration (Goode and Bean, Smithsonian Contrlb. Knowl., vol. 31, 1895, fig. 103) , about 6 Inches long, Is characterized In the legend as "slightly reduced." » Reported by Goode and Bean (Smithsonian Contrlb. Knowl., vol. 30, 1895, p. 88) as Atthoprora effulgens. » Vladykov, Proc. Nova Scotia Inst. Scl., vol. 19, 1935, p. 2. dorsal fin, a longer adipose fin, a longer anal, and a much smaller mouth. » Smithsonian Contrlb. Knowl. (vol. 30, 1895, p. 72) as M. opalinum. » It Is likely that Myctophum alacialt will be found In the Gulf of Maine sooner or later, judging from Its widespread distribution In the boreal belt of the Atlantic and from the fact that It has often been caught at the surface. It resembles M. afiint very closely In appearance, and In the general arrange- ment of the luminous organs, but differs from It In that one of the luminescent spots above the base of the ventral On Is elevated above the others. 144 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Pearlsides Maurolicus pennanti (Walbaum) 1792 Pearlfish Jordan and Everman, 1896-1900, P. 577. Description. — The presence of an adipose fin be- tween the dorsal and caudal fins, together with luminous organs, distinguishes the pearlsides from all other fishes that occur regularly in the Gulf of Maine. It agrees hi both these respects with the lanternfish (p. 143) and with the headlightfish (p. 142), but it has a much smaller mouth and a longer adipose fin than the first of these, and it lacks the large luminous patch on the snout that is so strik- ing a feature of the second. Also, the pearlsides, with its herring-like coloration (p. 88) differs strik- ingly from the lanternfish, which has a black back overlaid with silver; and probably the headlight fish as well. Figure 62. — Pearlsides {Maurolicus pennanti). After Smitt. The pearlsides is a flat-sided, large-headed little fish, its body (about one-fifth as deep as long, ex- cluding caudal fin) deepest forward of the ventral an d dorsal fins ; its eye very large ; its lower j aw pro- jecting; its mouth oblique; and both its jaws armed with minute teeth. The dorsal fin (about 11 or 12 rays) stands above the space between the ventrals and the anal; the anal is longer than the dorsal. The adipose fin (both of Woods Hole 32 and of Norwegian 33 examples) is low and long, much as it is in the capelin.34 The caudal fin is broad and slightly forked. The pearlsides has been described as without scales, but this is not correct, for both Scandinav- ian and Woods Hole specimens have been found to be clothed with large but extremely thin trans- parent scales. There is no definite lateral line. » Sumner, Osbum, and Cole, Bull. U. S. Bur. Fish., vol. 31, Ft. 2, 1913, p. 743. »» Smitt, Scandinavian Fishes, vol. 1, 1892, p. 933, pi. 44, flg. 3. « Ooode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895 p. 96) describe It as "very small," but probably their specimens were damaged. The most interesting and diagnostic feature of the pearlsides is the presence of a series of lumi- nescent dots situated as follows: 35 First, 12 pairs along the belly between the pectoral and the ven- tral fins, followed by 5 or 6 from the ventral fins to the anal fin, and, after a gap, by 24 or 25 between the center of the anal fin and the base of the caudal fin ; all these together form a practically continuous row on each side of the belly from throat to tail. Second, there is a row of larger spots a little higher up on each side, 6 from chin to pectoral fin, and 9 thence backward to the ventrals. Third, there is a group of 6 low down on each side of the cheek and throat ; there is likewise a spot in front of the base of each pectoral fin and 2 on the chin. Color. — -The pearlsides is colored much like a herring, with dark bluish or greenish back and lus- trous silvery-white sides and belly. The lumines- cent spots are described as black rimmed, their centers as pale blue in life but turning yellow in alcohol; and there is a narrow black band along the base of the anal fin and from there to the base of the caudal, the latter being barred with a similar black band. Size. — Only 1 to 2% inches long. Habits. — The relatives of the pearlsides are oce- anic, living in the mid-depths mostly below 150 fathoms, but the pearlsides itself has been found so often in the stomachs of cod and of herring (fish that do not descend to any great depth) that there is no reason to regard it as a "deep-sea" stray, nor has it ever been taken far from land so far as we can learn. It probably spawns in early spring, females with large eggs having been taken in Scottish waters in winter. General range. — The pearlsides (there are several other species closely allied to it) ranges widely in the open Atlantic, occurring at times in shoals on the coasts of Norway and in British waters. It is especially common off the coast of Scotland, but has not been recorded often on the American side of the Atlantic. Occurrence in the Gulf oj Maine. — The known occurrences of the pearlsides in the Gulf have been few. Storer36 (1S67) records one found alive on the beach at Nahant, Mass., in December, 1837; another taken from the stomach of a cod at »» This account is based chiefly on Smitt's description and plate, the speci- mens we have seen being in poor condition. » Fishes of Mass., 18(57, p. 160, as Scoprtus hvmboldtil. FISHES OF THE GULF OF MAINE 145 Provincetown ; a third picked up alive there in July, 1865 (pictured by Storer on pi. 25, fig. 5); and five others found on the Provincetown beach soon afterward. We have seen one specimen 41 mm. long taken from the stomach of a cod, on Platts Bank, July 27, 1924; one 43 mm. long, also from a cod's stomach, on Cashes Ledge, August 16, 1928; and four, 32 to 39 mm. long, -taken from the stomachs of two pollock that we caught in 20 fathoms, 7 miles southeast of Bakers Island, Mount Desert, Maine, July 24, 1930. It has been found twice at Grand Manan,37 and speci- mens were picked up on the beach at Campobello Island at the mouth of the Bay of Fundy in July 1914,38 while others were taken from the stomach of a pollock caught near by. It has also been recorded twice near Woods Hole. These locality records are distributed widely enough to show that it is to be expected anywhere in our Gulf. And we suspect that the pearlsides is not as scarce there as the paucity of actual records for it might suggest (in fact, Storer tells us that a Nahant fisherman reported finding them repeatedly in the stomachs of haddock many years ago), but that it keeps out of sight, being an inhabitant of the deeper water layers as its luminescent organs would suggest, coming up to the surface chiefly at night. VIPER FISHES. FAMILY CHAULIODONTIDAE The viper fishes have slender bodies, bulldog- like faces with long fangs; the first dorsal very far forward, the anal far back; and no barbel on the chin. Viperfish Cftauliodus sloani Bloch and Schneider 1801 Jordan and Evermann, 1896-1900, p. 585. Description. — The viperfish not only has lum- inescent organs, but it is very different in general appearance from all the fishes that are regular inhabitants of the Gulf of Maine. Most obvious of its characteristics is its bulldog-like mouth. It shares this with its fellow strays, Stomias (p. 147), Stomioides (p. 147) and Trigonolampa (p. 148) and the general form is much alike in the three. But there is no danger of confusing it with any one of thes,e if one looks closely, for the viperfish has an » Cos (Bull. Nat. Hist. Soc. New Brunswick, 14, 1890, append., p. 56) reported one found dead there, on the shore. !' Huntsman (Contrlb. Canadian Biol., (1921) 1922, p. 61.) adipose fin and its rayed dorsal fin is far forward, whereas Stomias, Stomioides, and Trigonolampa have no adipose fin and their rayed dorsal fin stands far rearward. In the viperfish the lower jaw is longer than the upper, the upper is armed with four long fangs on each side, while the lower has a series of pointed teeth set far apart, those in front very elongate and all of them so long that they project when the mouth is closed. Furthermore, the snout is so short that the very wide mouth gapes far back of the eve. The body is about seven times as long as deep, flattened sidewise, deepest close behind the head, and tapering evenly to the tail. The very short dorsal fin (6 or 7 raj-s) stands far forward and its first ray is separate, very slender, and about half as long as the fish when not broken off, as it usually is. The ventrals are about midway between the snout and the origin of the anal fin, variously pictured as either larger or smaller than the dorsal. The small anal is close to the caudal, with the adipose fin over it. The Figure 63. — Viperfish (Chauliodus sloani), \ southern slope of Browns Bank. After Goode and Bean. 146 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE body is clothed with large but very thin scales. There are several longitudinal rows of small luminescent spots on the ventral surface, running from throat to tail; several more such spots on each side of the head; and many tiny unpigmented dots scattered over the trunk.39 Color. — Greenish above, the sides with metallic gloss; blackish below. Size. — -Up to about one foot long. Habits. — Nothing is known of its habits except that it is an inhabitant of the mid-depths of the Atlantic Basin and that it probably does not rise closer to the surface than 150 or 200 fathoms except, perhaps, during its larval stages. Its teeth suggest a rapacious habit but there is no actual record of its diet. Occurrence in the Gulf of Maine. — The only definite Gulf of Maine records are of one specimen found in the stomach of a cod caught on Georges Bank in 1874, and of a second found in the stomach of a swordfish that was harpooned in the gully between Browns and Georges Banks in 1931.40 But the viperfish may be expected on the offshore banks as a stray at any time, for several have been taken off the continental slope abreast of southern New England 41 in deep water. THE STOMIATIDS. FAMILIES GONOSTOMIDAE AND STOMIATIDAE The stomiatids include many soft-rayed fishes of the mid-depths, of most diverse appearance, all of them with well developed luminescent organs, with large eyes, large mouths, and teeth in both jaws. Some have and others lack the adipose fin, but the ventrals are inserted more than one-third of the way back on the abdomen in all of them. They differ from the herrings and salmons in the structure of the skull. Four species have been taken in our Gulf, as strays from offshore. Cyclothone Cyclothone signata Garman 1899 Garman, Mem. Mus. Compar. Zoology, vol. 24. 1899, p. 246, pi. J, fig. 3. Description. — The general aspect of cyclothone is extremely characteristic, the somewhat com- pressed body beiDg deepest at the gill opening with the upper surface of the head concave in profile, the mouth so large that it gapes back of the eye, the lower jaw projecting, the eye very small, and the gill openings very long. The dorsal fin stands over the anal (the latter is much «• Brauer, Ttefsee-Flscho WIssensch. Ergeb. Deutschen TIefsee-Eiped., (1898-1899) 1906, vol. 15, Pt. I, p. 40. the longer of the two), both originating close behind the middle of the body. The caudal fin is deeply forked and there is no adipose fin. The luminescent spots are arranged as follows: One on the head; 1 close below the eye and in front of it; 2 on each gill cover; 9 or 10 between the branchiostegal rays; 2 longitudinal rows along each side of the body, a lower row of 13 from throat to ventral fins, 4 from ventrals to anal fin, and 13 from anal to caudal, and an upper row of 7 reaching about as far back as the ventrals. Color. — Cyclothone signata is colorless or pale gray, except that the blackish, dark silvery lining of the abdominal cavity shows through, that the luminous organs are black rimmed and silver cen- tered, and that there are the following black markings: a Y-shaped mark on the forehead; a series of spots or short transverse stripes on the flank; spots between the bases of the dorsal and anal fin rays; one or two transverse streaks across the bases of the caudal fin rays; and a number of « Reported to us by Walter H. Rich. «' Goode and Bean (Smithsonian Contrlb. Knowl., 1895, p. 97) list these captures. o o o o c ■Q Qnn rJT^TTh o Figure 64. — Cyclothone (Cyclothone signata). After Brauer. FISHES OF THE GULF OF MAINE 147 irregular flecks and dots along the back and on the gill covers.42 General range.- — This is an oceanic fish, very- abundant in temperate latitudes in the Atlantic where it lives pelagic from about 100 fathoms down to 250 fathoms; hundreds have often been taken in a single haul. It is also known from the Pacific. Occurrence in the Gulf of Maine. — Cyclothone appears within our limits only as a stray from the Atlantic Basin; one 23 mm. long that we took in a haul from 30 fathoms on Browns Bank, June 24, 1915, and a second mutilated specimen probably of this species from the Fundy Deep (haul from 90 fathoms), March 22, 1920, are the only definite records of it within our limits. Stomias Stomias ferox Reinhardt 1842 Jordan and Evermann, 1896-1900, p. 588. Description. — The members of this genus (there are several), resemble the viperfish in their bull- dog-shaped heads, with large mouth and long, fanglike teeth. But they do not have an adipose fin; the dorsal fin and the anal fin both stand far rearward close to the tail fin; the dorsal fin is even with the anal fin; and the first dorsal fin ray is not prolonged as it is in the viperfish. The chin bears a fleshy barbel nearly as long as the head and ending in a group of about three simple fila- ments. The sides of the body are clothed with about 6 rows of large, thin, somewhat irregular, hexagonal scales, and there is one row of lumines- cent spots low down along each side and two rows along the belly; also one small, circular light organ below each eye. The tip of the lower jaw overlaps and encloses the tip of the upper jaw when the mouth is closed in the only member of the genus that has been reported from the Gulf of Maine (or is likely to be found there) ; the slender body is about 17 times 18 For detailed accounts and colored Illustrations see Oarman (Mem. Mus. Comp. Zool., vol. 24, 1899, p. 24R, pi. J, fig. 3), Brauer (Wissensch. Ergeb. Deutschen Tlefsee-Eiped. (1898-1899), 1906, vol. 15, Pt. 1, p. 77. pi. 6, fig. 6), Murray and HJort (Depths of the Ocean, 1912, pi. I). as long as it is high; the ventral fins are only about as long as the head; the dorsal fin is of about the same size and shape as the anal fin, over which it stands; and there are about 85-86 light organs in each of the ventral rows, about 60 light organs in each of the lateral rows. Color. — Black below as well as above, the sides with metallic iridescence. General range and occurrence in the Gulf of Maine. — This oceanic fish is so widespread in the northern North Atlantic that it was taken at almost all the stations that the Michael Sars occupied there in 1910,43 mostly between the 75 to 80 fathom level and the 410 fathom (750 meter) level, most plentifully at about 275 fathoms (500 meters). The early cruises of the Blake and Albatross I took it at many localities also, along the continental slope of North America between the southeastern slope of the Newfoundland Banks and the Bahama Channel.44 Our only reason for mentioning it is that one specimen about 12 inches long (tip of snout to base of tail fin) was taken by a trawler on the northeastern part of Georges Bank (lat. 42°10' N., long. 67°05' W.), at about 100 fathoms, on January 20, 1936.4' Stomioides nicholsi Parr 1933 Parr, Copeia, 1933, No. 4, p. 177. Description. — The chief anatomical character separating Stomioides from Stomias is the struc- ture of the chin barbel. In Stomias this terminates in three simple filaments. But in Stomioides it not only has these barbels, but the main trunk is swollen at the tip and has two additional filaments on one side a little inward from its tip. Another difference is that the point of origin of the anal fin is in advance of the origin of the dorsal fin by a distance about as great as the diameter of the eye in Stomioides, whereas the point of origin of the anal fin is about even with that of the dorsal in )i inches in length, appear along our shores in spring. As yet we have few data on the exact date of their arrival on the Gidf of Maine coast. They appear as early as March at Woods Hole; by mid- or late April both in Narragansett Bay and in Passa- maquoddy Bay at the mouth of the Bay of Fundy, while Welsh encountered a tremendous run in Little River, near Gloucester, on May 5, 1913, suggesting that they may be expected in the mouths of most Gulf of Maine streams during that month. And they are found ascending streams in the Bay of Fundy region during the summer. A run may last for a month or more in one stream, only for a few days in another. And there is a noticeable segregation even at this early stage, some of the elvers remaining in tidal marshes, in harbors, in bays back of barrier beaches, and in other similar situations, some even along the open coast, especially where there are beds of eel grass (Zostera); while others go into fresh water, some of them ascending the larger rivers for tremendous distances.65 It is now generally believed that most of the eels that are caught in fresh water are females. But some of the females remain in salt marshes and harbors, to judge from the large size of many of the eels that are caught there. And nothing is known as to what preference the males of the American eel may show in thig respect. It is no wonder that the ability of the elvers to surmount obstacles as they run upstream is proverbial, for they clamber over falls, dams, and other obstructions, even working their way up over damp rocks as Welsh saw them doing in Little River, where they were so plentiful on May 5 and 7, 1913, that he caught 1,500 in one scoop '* The life history of the eel Is presented in more detail than is possible here by Schmidt (Philos. Trans. Roy. Soc. London, Series B, vol. 211 (1922) 1923, pp. 179-208, summarized in Nature, vol. 110, 1922, p. 716), and by Cun- ningham (Nature, vol. 113, 1924, p. 199). See also Schmidt (Rapp. et Proc- Verb. Cons. Perm. Internat. Explor. Mer, vol. 5, No. 4, 1906, pp. 137-204, pis. 7-13); for a popular account see Smith (Nat. Geog. Mag., vol. 24, No. 10, October 1913, p. 1140). " Eels are native in Lake Ontario which they reach by way of the St. Lawrence River; and up the Mississippi drainage systems even as far as North Dakota, Wisconsin, Ohio, and western Pennsylvania. of a small dip net and 545 with a few grabs of his bare hand. Elvers in equal multitudes have often been described in other streams, American as well as European. Eels can live out of water so long as to give rise to the story that they often travel overland. There is no positive evidence for this. But Sella 56 has proved, by experiments with European eels marked so as to be recognizable if recaught, that they can earn7 out journeys as long as 31 miles (50 kilometers) along underground waterways. Doubtless it is this ability that explains the presence of eels in certain ponds that have no visible outlet nor inlet, a fact often attested. It is true in a general way that eels seek muddy bottom and still water, as has been said so com- monly. But this is not always so whether in salt water or in fresh. Thus the rocky pool at the outer end of the outlet from Little Harbor, Cohasset, on the south shore of Massachusetts Bay, is a good place to catch eels; and large ones are only too common in swiftflowing, sandy trout streams on Cape Cod ; we have had one follow and nibble at the trout we were dragging behind us on a line. The fact is, they can live and thrive wherever food is to be had, which applies to them in estuarine situations and in fresh water. No animal food, living or dead is refused, and the diet of the eels in any locality depends less on choice than on what is available. Small fish of many varieties, shrimps, crabs, lobsters, and smaller Crustacea, together with refuse of any kind (for they are scavengers) make up the bulk of the diet in salt, estuarine, and brackish water. Being very greedy, any bait will do to catch an eel. They are chiefly nocturnal in habit, as every fisherman knows, usually lying buried in the mud by day to venture abroad by night. But eels, large and small, are so often seen swimming about, and so often bite b}r day that this cannot be laid down as a general rule. Eels tolerate a wide range of temperature. But it is common knowledge that those inhabiting the salt marshes and estuaries of our Gulf, and its tributary streams, mostly lie inactive in the mud during the winter. Eels grow slowly. Hildebrand and Schroeder 57 concluded from a series of measurements taken at different seasons in lower Chesapeake Bay that those M Mem. R. Cornit. Talassogr. Dal., vol. 158, 1929. " Bull. U. S. Bur. Fish., vol. 43, 1928, p. 114. FISHES OF THE GULF OF MAINE 153 2}2 inches long in April are about 5 inches long a year later, or about 2 years after their transforma- tion. The winter rings on the scales have shown that full grown adults of the European species are from 5 to 20 years old, depending on food supply, and other conditions; this is corroborated for the American species by the fact that Dr. Hugh M. Smith, former Commissioner of the United States Bureau of Fisheries, found that a female, on the way down the Potomac, was in her twelfth year. At the approach of sexual maturity, which takes place in the fall, the eels that are in fresh water drop downstream, traveling mostly at night. They now cease feeding, as do those that have been living in the river mouths, bays, and estuaries; the color of the back changes from olive to almost black, the ventral side turns silvery, and the eyes of the males grow to twice their previous size. Both males and females then move out to sea, and it is not until after they reach salt water that the ovaries mature. In fact, no perfectly ripe female eel has ever been seen, and only one ripe male (of the European species). So little is the life history of the eel understood by our fishermen that we again emphasize the undoubted fact that no eel ever spawns in fresh water. The eels drop wholly out of sight when onco they leave the shore;68 no one knows how deep they swim, but they certainly journey out beyond the continental slope into the oceanic basin before depositing their eggs. Schmidt has been able to outline the chief spawning center of the American species (from the captures of its youngest larvae) as between latitudes 20° and 30° N. and between longitudes 60° and 78° W.; i. e., east of Florida and of the Bahamas south of Bermuda. But it may also spawn (always in deep water) farther north as well.69 The American eel spawns in midwinter, thus occupying one to two months in its journey from the coast to the spawning ground, for Schmidt found very young larvae (7 to 8 mm.) in February. Eels, like Pacific salmon, die after spawning, the evidence of this being that no spent eels have ever been seen and that large eels have never been •' Large eels, on their seaward journey, have occasionally been caught by otter trawlers in the western part ot the British Channel, but we know of no such occurrence on this side ot the Atlantic. » See Schmidt (Ann. Rep. Smithsonian Inst., (1924) 1925, pp. 279-314) tor a readable account ot the investigations which enabled him to chart the breeding places and seasons of the American and European eels. 210941—53 11 known to run upstream again. Smith suggests that they probably '"jellify" and disintegrate, as the conger does. Eels (European) are among the most prolific fish, ordinary females averaging 5 to 10 million eggs and the largest ones certainly 15 to 20 million. It is doubtful whether eggs laid by the American eel have been seen, or of the European either, for that matter.60 But it is generally supposed that they float in the upper or inter- mediate water layers until hatching. The larval, so-called "leptocephalus" stage, like that of all the true eels, is very different in appearance from the adult, being ribbon-like and perfectly trans- parent, with small pointed head; and it has very large teeth, though it is generally believed to take no food until the time of its metamorphosis. These leptocephali of our eel, living near the surface, have been found off our coasts as far north as the Grand Banks, but never east of longitude 50' W. Inasmuch as the breeding areas of the American and European eels overlap, not the least inter- esting phase of the lives of the two is that the larvae of the American species should work so consistently to the western side of the Atlantic, and those of the European to the eastern side that no specimen of the former has ever been taken in Europe or of the latter in America. The American eel takes only about one-third as long as the European to pass through its larval stage; i. e., hardly a year, as against 2 to 3 years. The leptocephali reach their full length of 60 to 65 mm. by December or January, when meta- morphosis takes place to the "elver"; the most obvious changes being a shrinkage in the depth and length of the body but an increase in its thickness to cylindrical form, loss of the larval teeth, and total alteration in the aspect of head and jaws, while the digestive tract becomes functional. It is not until they approach our shores, how- ever, that the adult pigmentation develops or that the elver begins to feed, a change that is accompanied by a second decrease in size. How such feeble swimmers as the leptocephali find " Four eggs taken on the Arcturus expedition near Bermuda in 1925 wer« provisionally identified as those of the American eel by Fish who has pictured them and the larvae hatched from one of them (Zoologica, New York Zool. Soc, vol. 28, 1927, pp. 290-293, figs. 103-107). But the date at which they were taken (July 15-17) makes it more likely that they belonged to some other member of the eel tribe. 154 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE their way into the neighborhood of the land remains a mystery. It seems certain, however, that all the young eels bound for the Gulf of Maine complete the major part of their meta- morphosis while they still are far offsbore. Thus we have never taken one in the leptocephalus stage in the Gulf of Maine in all our tow-nettings, whereas (more significant) the Albatross towed three young eels in the so-called "glass-eel" stage, 54 to 59 mm. long, of practically adult form but still transparent, during her spring cruise in 1920, one of them on Georges Bank, March 11 ; a second on Browns Bank, April 16 ; and one in the western basin of the Gulf off Cape Ann, February 23. Evidently they were intercepted on their way in to land. And since all three were on the surface, we may take it that glass eels, like leptocephali, keep to the uppermost water layers during their journey. General range. — Coasts and streams of West Greenland,61 eastern Newfoundland,62 Strait of Belle Isle, and northern side of the Gulf of St. Lawrence south to the Gulf of Mexico, Panama, West Indies, and (rarely) to the northern coast of South America; also Bermuda; running up into fresh water but going out to sea to spawn p. 153. Occurrence in the Gulf oj Maine.- — The occur- rence of the eel around the periphery of our Gulf can be described in the one word "universal." There is, we believe, no harbor, stream mouth, muddy estuary, or tidal marsh from Cape Sable on the east to the elbow of Cape Cod on the west but supports eels in some numbers, and they run up every Gulf of Maine stream, large or small, from which they eventually find their way into the ponds at the headwaters unless barred by insurmountable barriers such as very high falls. Examples of long journeys by eels upstream, in New England rivers, are to the Connecticut Lakes, New Hampshire, at the head of the Connecticut River; to the Rangeley Lakes at the head of the Androscoggin, and to Matagamon Lake, at the head of the East Branch of the Penobscot. Eels are even caught in certain ponds without outlets, as noted above (p. 152). On the other hand, we have seen a few (and some large ones) along the open coast, at Cohasset, for example, but always •' Jensen (Invest, of the Dana In West Greenland Waters, 1925, Eitr. Papp. et Proc. -Verb Cons. Internal. Expl. Mer, vol. 39, 1926, p. 101) records the American eel as one of the four fresh-water fishes known from the west coast of Greenland. •> Reported by Dr. G. W. Jefiers as common. close in to the shore line and in only a few feet of water, where flounder fishermen catch them from time to time. Importance. — Schmidt has suggested that the American eel is not as plentiful in actual numbers as the European, arguing from the facts that its larvae have not proven so common on the high seas, and that the American catch of eels (about 2,000 tons yearly) was but a fraction as large as the European catch (about 10,000 tons annually). But it is not safe to draw any conclusions from the statistics because the American catch is limited more by the fact that eels are not much in demand, than by the available supply. And the local demand is less for them today than it was 30 years ago, as is reflected in a decrease in the re- ported landings from about 305,000 pounds for Maine and about 240,000 pounds for Massachu- setts in 1919 to about 19,000 pounds for Maine and about 32,000 pounds for Massachusetts in 1947. The yearly landings of eels along the Cana- dian shore of our Gulf and from the tributary fresh waters are 30,000^0,000 pounds nowadays. Practically the entire coastwise catch is made in salt marshes, estuaries and stream mouths; the numbers captured up stream are negligible of recent years, except in New Brunswick where 16,000 pounds were caught in the lower sections of the St. John River System in 1950.63 In Germany, however, where the demand for eels is much greater, the yearly catch is nearly four times as great for rivers and other fresh waters as it is for the coast. And many millions of elvers were transplanted, during the 1930's, from British rivers (the Severn in particular) to landlocked bodies of water in Central Europe which the young eels could not reach naturally. The greater part of the catch is made in nets and eelpots; and some are speared, mostly in late autumn and winter, often through the ice. American conger Conger oceanica (Mitchill) 1818 64 Sea eel Jordan and Evermann, Leptocephalus conger (Linnaeus 1758), 1896-1900, p. 354. " Information from A. H. Leiin. u The American conger had long been considered Identical with the Euro- pean. But Schmidt (Nature, vol. 128, 1931, p. 602) has recently shown that it is a distinct species, characterized by having fewer vertebrae; a rela- tionship paralleling that between the American and European eels of the genus Anguilla. FISHES OF THE GULF OF MAINE 155 Figure 70.- — American conger (Conger oceanica). A, adult, Connecticut; from Goode, drawing by H. L. Todd; B, "Leptocephalus" stage, 84 mm., Chesapeake Bay. Description. —The readiest characters by which to distinguish the conger from other eels are noted in the key (p. 150); notably the origin of the dorsal fin above or only very slightly behind the tip of the pectoral when the latter is laid back, the rathe? long-pointed snout, the large mouth cleft back at least as far as the middle of the eye, and the scaleless skin. The conger has many more verte- brae than the common eel and there are other skeletal differences.66 The conformation of the tip of the snout likewise helps to identify the conger, for its upper jaw usually projects beyond the lower, whereas in the common eel the reverse is true, or at least the lower equals the upper. Fur- thermore, the eyes of the conger are oval and larger than the round eyes of the common eel. To give an idea of the proportions of the conger, we need only add that the distance from tip of snout to dorsal fin is about one-fifth of the total length; the length of the snout is one-fourth that of the head; the length of the pectorals is equal to one-third to one-fourth of the distance from dorsal fin to tip of snout; and that the body is of the snake-like form characteristic of eels in general. Color. — Bluish gray or grayish brown above, sometimes of a reddish tinge, sometimes almost black; paler on the sides; dingy white below. Size. - This is a much larger fish than the com- mon eel. The larger ones taken off southern New England and New Jersey are said to measure 4 feet up to 7 feet in length. The general run of those caught weigh 4 to 12 pounds, the heaviest we have seen weighed about 22 pounds. But the North American species never attains the enormous size reached by the European species; the largest European conger reliably reported, of which we have read, was 9 feet long, and weighed 160 pounds.66 Habits. — The depth range of the conger is from close to the coastline (they arc caught from the dock at Woods Hole) out to the edge of the continental shelf, the deepest record for it being for one that we trawled at 142 fathoms off southern New England, on the Albatross III, in May 1950. It feeds chiefly on fish: butterfish, herring, and eels have been found in their stomachs at Woods Hole. They also prey on shrimps and small mollusks at times. And we have caught them (and have seen them caught) on crabs, on soft clams (Mya), on sea clams (Mactra) and on cut fish bait. It is nowr well established that the European species (hence no doubt the American also) breeds but once during its life and then perishes like the common eel. Ripe congers are never caught on hook and line, for they cease to feed, hence to bite, for some time previous. But the males of the European species, kept in aquaria, •* For an account of these, see Smltt (Scandinavian Fishes, vol. 2, 1895 pp. 1016-1017, 103". "Jenkins, Fishes of the British Isles, 1925, p. 275; see also Day, Fishes of Great Britain, vol. 2 ,1884, p. 253, for large European congers. 156 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE have repeatedly been known to become fully ripe, females nearly so,67 then invariably dying. The ripening of the sexual products is accompanied by changes in the shape of the head; in the loss of the teeth; and in a jellification of the bones, while the eyes of the males become enormous and the females become much distended by the ovaries. It is probable that the American conger ripens off the coast of southern New England in summer; European congers in captivity have been known to do so every month in the year except October and November. It seems that the conger, like the common eel, moves out from the coast to spawn, for its young larvae have never been taken inshore, and Dr. Johannes Schmidt's 68 discovery of very young larvae in the West Indian region, but nowhere else, points to this as the chief spawning ground of the American conger, if not the only one. The congers are extremely prolific fish, the number of eggs a European female may produce having been estimated as high as 3 to 6 millions. American conger eggs have never been identified, for although eggs taken over the tilefish grounds 30 miles south of Nantucket lightship in July 1900 69 have been credited to this species, there is no certainty that this was their true parentage. It has long been known that the congers, like the common eels, pass through a peculiar ribbon- like larval stage (the so-called "leptocephalus") very broad and thin and perfectly transparent, with a very small head.70 In fact the first lepto- cephalus ever seen (about 1763) was the larval European conger. But its identity was not estab- lished definitely until 1886, when the famous French zoologist, Delage,71 reared one through its metamorphosis at the biological station at Roscoff. The leptocephalus stage of the conger is rela- tively more slender than that of the common eel, it grows larger (to a length of 150-160 mm.), and its vertebrae and muscle segments are far more numerous (140-149 in the American conger, " Cunningham (Jour. Mar. Biol. Assoc. United Kingdom, N. Ser., vol. 2, 1891-92, pp. 16-12) gives an interesting account of this and other phases of the life history of the conger. » See Nature, vol. 128, 1931, p. 602, for a discussion of this question by Dr. Schmidt. •• Eigenmann, Bull. U. S. Bur. Fish., vol. 21, 1902, p. 37. 70 For photographs of the leptocephalus stage of the European conger, see Schmidt, Rapp. et Proc. Verb. Cons. Perm. Internat. Explor. Mer, vol. 5, No. 4, 1906, pi. 9, figs. 8, 9; and Meddelelser Komm. Havundersjigelser, Ser. Flskerl, vol. 3, No. 6, pi. 1, figs. 1-3. " Conptes Rendus Acad. Scl. Paris, vol. 103, 1888, p. 698. 154-163 in the European) than in the common eels (about 107 in the American eel and about 114 in the European). But the number of body segments (visible only under a lens) is not of itself a safe clue to identity, for there are as many or more in the long-nosed eel (p. 158) which has been reported in the Gulf; also in the morays, and in various other members of the eel tribe.72 The duration of the larval period of the conger is not known. The process of metamorphosis consists essentially in a thickening and narrowing of the body, an enlargement of the head, the for- mation of the swim bladder and permanent teeth, and the development of pigment in the skin, a change that occupied about two months (May to July) in the case of Delage's European specimen. His young conger was 9.3 centimeters (3.6 inches) long at its completion.73 General range. — Continental shelf of eastern America: adults are known north to the tip of Cape Cod; larval stages to eastern Maine. Its southern boundary cannot be stated until the congers of the coasts of North and of South America have been critically compared. It is rep- resented by a closely allied species (Conger conger) in the eastern North Atlantic. Occurrence in the Gulf of Maine. — The only records for adult congers within the limits set here for the Gulf of Maine are of one taken at North Truro, Cape Cod; a second specimen trawled close to Provincetown in Cape Cod Bay, July 5, 1951, by Capt. Herman Tasha; and a third, trawled south of Nantucket shoals by Albatross III, in mid May 1950.74 But the conger must be much more plentiful at times off the shoals than the foregoing would suggest for Capt. Henry Klim of the dragger Eugene H reports trawling 1 ,400 pounds of them there, at 76 fathoms, March 25-30, 1951.75 And its curious band-like "lepto- cephalus" larvae have been found within the Gulf on several occasions. Thus, half a dozen speci- mens were picked up on the beach at Cherryfield and Old Orchard, Maine, and at Nahant, Mass., » Fish (Zoologlca, New York Zool. Soc, vol. 8, 1927, pp. 307-308) gives a table of the numbers of body segmentsfor various eels and for "leptocephalus" larvae of known and unknown parentage, " Schmidtlein (Mittleil, Zool. Stat. Neapel, vol. I, 1879, p. 135) speaks of young "congers" at Naples In April as hardly one-third as long as this, a discrepancy suggesting that these may actually have belonged to one of the Muraenold eels. N Local reports of congers do not necessarily relate to the true conger, for the eel pout (p. 510), which is common in the Gulf, Is often misnamed thus. » At lat. 40° N., long. 69° 50' VV. FISHES OF THE GULF OF MAINE 157 more than a half century ago. Two specimens, also picked up on the beach, were sent up from Newburyport, Mass., in November 1929; and A. H. Clark, of the U. S. National Museum, informs us that he has found many larvae of the leptocephalus tjrpe at Manchester, Mass., which probably were congers to judge from their size. The conger occurs regularly and commonly to the west and south of Cape Cod, being taken near Woods Hole from July into the autumn, and about Block Island from August until November. Very little is knowTi about their movements. But we suspect that they shift offshore into deeper and warmer water for the winter, judging from their absence then in shoal water, contrasted with the large offshore catch in March mentioned above (p. 156) and with the fact that we saw several trawled at 50 to 142 fathoms off southern New England on the Albatross III, in May in 1950. Slime eel Simenchelys parasiticus Gill 1879 Snub-nosed eel Jordan and Evermann, 1896-1900, p. 349. Description. — The most distinctive characters of the slime eel, its eel-like form, snub nose, long dorsal fin, and soft and slimy body, have been mentioned already (p. 150). It is stouter and more sway-bellied than the common eel, very soft, and with a more tapering tail. The dorsal fin origi- nates a short distance behind the tips of the pectorals when the latter are laid back against the body, and the anal runs forward on the lower surface almost to the vent, which is situated about midway of the body. The head is much shorter than in either the common eel or the conger; the mouth is small, gaping back only about half way to the forward edge of the eye, with upper and lower jaws of equal length and each armed with a single series of small, close-set cutting teeth. The gill openings are small, and instead of being vertical and on the sides of the neck as they are in the common eel, they are longitudinal and lower down on the throat. Color. — Dark brown, with the belly only a little paler than the back, though usually more or less silvery. Size. — About 2 feet long. Habits. — It is partly parasitic in habit, burrow- ing into the bodies of halibut and other large fish, circumstances under which a considerable number of specimens have been brought in by fishermen. Very likely it was common inshore in the old days when halibut were plentiful there. It also lives independently on the bottom. Nothing is known of its manner of life beyond this, nor of its breeding habits. We may add from experience that it is as slimy as a hag and drips with sheets of mucus when drawn out of the water. General range. — The continental slope, and the slopes of the offshore banks, from abreast of the eastern end of Long Island to the Newfoundland Banks, in depths ranging from 200 to more than 900 fathoms; also in deep water about the Azores, and represented in Japanese waters by an ex- tremely close relative, if, indeed, it is separable at all from the Atlantic slime eel.76 Occurrence in the Gulf oj Maine. — There is no definite record of the snub-nosed eel actually with- in the southern rim of the Gulf so far as we can learn, and our only first-hand experience with it was on the slope south of Nantucket lightship, where we captured 21 in a Monaco deep-sea trap " The Japanese slime eel, described first as a distinct species (leptosomut) by Tanak-a in 1908, has been classed more recently by him (Fishes of Japan, vol. 42, 1928, p. 810, pi. 173, fig, 470) as identical with the Atlantic paratiticut. <%Z0&2%k Figure 71. — Slime eel (Simenchelys parasiticus), off Sable Island Bank. From Goode and Bean. Drawing by H. L. Todd. 158 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE in 455 fathoms, on the Grampus in July 1908. It must be extremely abundant along that zone, how- ever, for so many to find their way into the trap in as short a set as two hours. And it has been recorded so often in water as shoal as 200 fathoms that it may be expected in the bottom of the Eastern Channel and in the southeastern deeps of the Gulf of Maine. Long-nosed eel Synaphobranchus pinnatus (Gronow) 1854 Jordan and Evermann, 1896-1900, p. 351. Description. — This deep-sea species, a typical eel in general appearance, is readily identifiable among its tribe by the fact that while its dorsal fin originates about as far back as in the common eel (p. 151), relative to the length of the fish, its point of origin is considerably behind the vent instead of in front of the latter, and that its anal fin originates considerably in front of the dorsal fin instead of behind it as is the case in all other Gulf of Maine eels. Furthermore, its mouth is much wider, gaping far back of the eye, and its snout is pointed. The most interesting anatomic characteristic of this eel is that its gill openings, opening longitudinally on the lower side of the throat, join together in front, apparently as a single V-shaped aperture, though actually they are separate within. Color. — Grayish, darkest below, with the ver- tical fins darker behind but pale-edged in front, and with the inside of the mouth blue black. Size. — The largest of 89 specimens measured by Goode and Bean was nearly 22 inches (545 mm.) long, the smallest about 9 inches (221 mm.) The largest we trawled on the Caryn, in June 1949, was 24 inches (605 mm.) long. Collett 77 mentions one 26% inches (675 mm.) long from the Azores. " Result, des Camp. Sci. Prince de Monaco, Ft. 10, 1896, p. 154. Habits. — Nothing is known of its habits except that it is a ground fish; that the readiness with which it bites a baited hook proves it predaceous; and that specimens in spawning condition have been taken in summer.78 On June 17, 1949 hi lat. 42° 38' N., long. 64° 04' W., in 400-460 fathoms, we trawled many on the Caryn, both males and females, 18% to 24 inches (470-605 mm.) long that had well developed gonads, one female having already spawned. The ripe eggs are orange in color and about 1 mm. in diameter. In its development it passes through a lepto- cephalus stage even more slender than that of the American conger (p. 156), and its body segments (144-149) overlap those of the American conger (140-149) in number. General range. — This deep-water species has a wide distribution. In the western side of the North Atlantic it has been taken at many local- ities along the continental slope from the offing of South Carolina to the Grand Banks ; it is known in the east from the Cape Verdes; off Morocco; from the Canaries; from the Azores; near Madeira; also from the Faroe Bank and Faroe-Shetland Channel. And its leptocephalan larvae have been taken in such numbers from north of Spain to south of Iceland that it must be one of the most plentiful of deep-water fishes there.79 It is also recorded off Brazil in the South Atlantic; likewise in the Arabian Sea; about the Philippines; and in Japanese waters, or is represented there by a very close relative. Most of the captures have been from depths of 300 to about 2,000 fathoms, but it has been taken as shoal as 129 fathoms. » The "leptocephalus" larvae of the long-nosed eel are described, with photographs by Schmidt (Rapp. et Proc. Verb. Cons. Perm. Internat. Explor. Mer, vol. 5, No. 4, 1906, p. 191, pi. 9, figs. 4-6; and Meddel. Komm- Havunders0ge]ser, Ser, Flskerl, vol. 3, No. 6, 1909, p. 7). " This fact is commented on by Schmidt (Rapp. Cons. Perm. Internat. Esplor. Mer, vol. 5, No. 4, 1906, p. 191). For further details as to Its distri- bution see Koefoed, Rept. Michael Sars North Atlantic Esped., (1910), vol. 4, Pt. 1, 1927, pp. 11, 14. Figure 72. — Long-nosed eel (Synaphobranchus pinnatus), La Have Bank. H. L. Todd. From Goode and Bean. Drawing by FISHES OF THE GULF OF MAINE 159 Occurrence in the Gulf of Maine. — This eel has not actually been reported within the geographic limits of the Gulf. But it is to be expected in the eastern channel and possibly above 150 fathoms along the slopes of Georges Bank, for fishermen have caught them in water as shallow as that off La Have Bank, while they have been trawled in 168 fathoms and 129 fathoms off southern New England by the Fish Hawk and Albatross. So many of them have been brought in by fishermen 80 from deep water off the fishing banks to the east- ward of longitude 65°, and so many have been trawled along the continental slope thence west- ward,81 that this eel must be one of the commonest of fishes below 150 to 200 fathoms, all the way from the Grand Banks to abreast of New York. Snake eel Omochelys cruentifer (Goode and Bean) 1895 Jordan and Evermann, 1896-1900, p. 377, Pisodonophis cruentifer. Description.- — The most striking feature of the snake eel and one that distinguishes it from all other Gulf of Maine eels is that the tip of its tail is hard and pointed. Other distinctive features are that it is only about one thirty-seventh to one thirty-eighth as deep as it is long; that its dorsal fin originates oidy a short distance behind the tips of the pectorals when these are laid back; that its anal fin originates far behind its dorsal fin; that its snout is bluntly pointed; and that its mouth gapes rearward considerably beyond its eyes (but not so far back as in the long-nosed eel, p. 158). The dorsal and anal fins end a little in front of the tip of the tail. The gill openings M Many such Instances are listed In the Rept. U. S. Comm. Fish. (1879) 1882, p. 787. " Goodo and Bean, Smithsonian Contrlb. Know]., vol. 30, 1895, pd. 143-144. are short new-moon-shaped slits, close in front of the bases of the pectoral fins. Its "peculiar and savage physiognomy" was stressed by its describers.82 Color. — Originally described as uniform brown- ish yellow. But those that we have seen have been uniform light brown below as well as above, large ones darker than small ones. A young one about 2}{ inches (6% cm.) long was pale with dark speckles. Size. — The largest yet seen was 16% inches long. Habits. — The original account of the snake eel includes the information that specimens had been received that had been taken from the bodies of other fish, evidence that it is a parasitic-boring form. Nothing else is known of its habits. General range. — Western side of the Gulf of Maine to the offing of Cape Henry, Va. Occurrence in the Gulf of Maine. — The snake eel was originally described in 1895 from 6 speci- mens trawled off Nantucket by the Fish Hawk, and a number have been taken thence southward to the latitude of Cape Henry, Va., by the Alba- tross II, in depths of 24 to 245 fathoms. The only report of it within the Gulf of Maine is by its describers of specimens taken by fishermen on Jeffreys Bank many years ago. Snipe eel Nemichthys scolopaceus Richardson 1848 Jordan and Evermann, 1896-1900, p. 369. Description. — The snipe eel is easily recognizable by its extremely slender body (the fish may be 75 times as long as deep), with its tail tapering to a thread, and by its elongate, slender, bill-like jaws, one as long as the other, the upper one curving upward, but the lower more nearly >' Ooode and Bean, Smithsonian Contrlb. Knowl., vol. 30, 1895, p. 147. ; Figure 73. — Snake eel (Omochelys cruentifer), continental slope south of Nantucket Shoals. From Goode and Bean. Drawing by H. L. Todd. 160 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 74. — Snipe eel (Nemichthys scolopaceus) . Drawing by E. N. Fischer, from Bigelow and Welsh. straight. The head is much deeper than the neck, with large eyes. The dorsal fin originates in front of the pectorals, the anal about abreast of the tip of the pectorals, and both dorsal and anal run back to the tip of the tail. There has been some confusion in the published accounts and illustrations as to the dorsal and anal fins, for while Vaillant 83 shows both as about as high throughout their length as the fish is deep, Goode and Bean84 picture the dorsal as much bigher than the anal (the artist evidently having transposed the two fins) , whereas Brauer 86 repre- sents the anal as approximately twice as high as the dorsal and the latter as soft rayed in its an- terior and posterior portions but composed of short thorn-like spines along its central third. The fins of two specimens taken off New England, now in the collections of the Museum of Comparative Zoology are as follows: Dorsal, soft-rayed and nearly as high as the body is deep for its first half; back of that it con- sists of a series of very short, stiff rays that extend to the tip of the tail. Anal, soft-rayed throughout its length and about as high as the body is deep, tapering to almost no tiling on the tail. The confusion has been due in part to the rather fragmentary state in which these deep-water fish usually arrive on board, but at the same time it is probable that two distinct species have been con- fused under the name scolopaceus, as Brauer suspected. Color. — Described as pale to dark brown above with the belly and anal fin blackish after preserva- tion. Judging from experience with other deep- sea fishes and from Brauer's plate (which, however, may be another species), we suspect that it is chocolate brown above in life and velvety black below. Size. — Maximum length about 3 feet. Habits. — Although commonly spoken of as a "deep-sea" fish, this species is undoubtedly an inhabitant of the mid depths, not of the bottom, and judging from the occurrence of other black fishes it probably finds its upper limit at 100 to 200 fathoms. Nothing further is known of its habits, but Mowbray's 86 capture near Bermuda of a snipe eel clinging by its jaws to the tail of a large red snapper has suggested that such may be a regular habit of this curious species. General range. — The snipe eel has been taken in deep water at many stations off the east coast of North America between latitudes 31° and 42°N., longitudes 65° and 75°W. ; also in the South Atlan- tic; near the Azores; near Madeira; off the Cape Verde Islands; off West Africa; and in the Pacific of New Guinea. Occurrence in the Gulf of Maine. — One specimen taken from the stomach of a codfish caught on Georges Bank in 45 fathoms is the only Gulf of Maine record, but several have been taken in depths of from 300 to 2,000 fathoms on the sea- ward slope of the bank. THE LANCET FISHES. FAMILY ALEPISAURIDAE The lancet fishes have one long and very high dorsal fin, soft-rayed from end to end; a small » Poissons. Exped. Sci. TravaitteuT et Talisman, 1S88, pi. 7, figs. 2 and 2a. » Smithsonian Contrlb. Know]., vol. 31, 1895, pi. 46, fig. 170. " Tiefsee-Fische, Wlss. Ergcb. Deutsch. Tiefsee-Exped. 0898-1899), vol. 15, Pt. I, 1900, p. 126, pi. 9, fig. 1. adipose fin behind the dorsal fin, like that of a salmon or smelt; a deeply forked caudal fin; a short anal, most of which is behind the rear end of the dorsal; large pointed pectorals and ven- "Copeia.No. 108, 1922. p. 49. FISHES OF THE GULF OF MAINE 161 trals; and a very wide mouth with large teeth. Several species are known, all belonging to deep water; only one has been taken within the province covered by this report. Their closest affinities seem to lie with the lanternfishes (p. 141). Lancetfish Alepisaurus ferox Lowe 1833 Handsawfish Jordan and Evermann, 1896-1900, p. 595. Description. — The combination of a long and very high dorsal fin, soft-rayed from end to end, with the presence of an adipose fin behind it, distinguishes the lancetfish from all other Gulf of Maine fishes. The body is slender, somewhat flattened sidewise, deepest at the gill covers, and tapers back to a slender caudal peduncle. The snout is long and pointed, the mouth wide, gaping back of the eye, and each jaw has two or three large fangs, besides smaller teeth. The dorsal fin (41 to 44 rays) originates on the nape and occupies the greater length of the back, is rounded in outline, about twice as high as the fish is deep, and can be depressed in a groove along the back. The adipose fin recalls that of the smelt in form and location. The caudal is very deeply forked; its upper lobe is prolonged as a long filament, and although most of the speci- mens so far seen have lost this we have an ex- cellent photograph showing it. The anal fin originates under the last dorsal ray, and is deeply concave in outline. The ventrals are about halfway between the anal and the tip of the snout, while the pectorals are considerably longer than the body is deep and are situated very low down on the sides. There are no scales and the fins are exceedingly fragile. Color. — Sides described as metallic silvery. We have not seen a newly taken specimen. Size. — The collection of the Boston Society of Natural History contains the cast of a specimen about 6 feet long that was taken off Nova Scotia in August 1910, and this is probably about the maximum size. Habits. — This is an oceanic species, of the mid-depths, appearing only as a stray shoaler than 200 fathoms. Nothing is known of its habits. A Block Island specimen had eaten a small spiny dogfish. General range. — -Widely distributed in the deep waters of the Atlantic, also reported from the northeastern Pacific.87 Occurence in the Gulj oj Maine. — A specimen brought in by a fisherman from Georges Bank 88 about 1878 or 1879 is its only claim to mention here. Goode and Bean and Yladykov and McKenzie 89 have reported other captures of lan- cetfishes from La Have Bank, from southeast of Emerald Bank and Banquereau. Another speci- men 5Yi feet long was caught alive in the surf on Block Island, R. I., March 12, 1928, and reported by Mrs. Elizabeth Dickins who sent us a photo- graph of it. " Crawford (Copela, No. 164, 1927, p. 60) reports several .4. feroz from the halibut banks off the northwestern coast of British Columbia. 11 No definite information Is available as to this specimen. - Proc. Nova Scotia Inst. Scl., vol. 19, 1935, p. 63. Figure 75. — Lancetfish (Alepisaurus ferox) . New York market specimen. From Jordan and Evermann. Drawing by H. L. Todd. 210941—53 12 162 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE THE MUMMICHOGS OR KILLIFISHES. FAMILY POECILIIDAE The mummichogs are small fishes made rec- ognizable by having only one short soft-rayed dorsal fin situated far back, and ventrals situ- ated on the abdomen, combined with a small mouth at the tip of the snout, a very thick caudal peduncle, and a rounded tail fin. The family is represented in the Gulf of Maine by three species, two of Fundulus and one of Cyprinodon, the former slender and the latter deep in outline, a dif- ference in body form sufficient to distinguish the one genus from the other at a glance. The teeth are likewise different in the two genera, those of Fundulus being sharp-pointed, whereas they are wedge-shaped in Cyprinodon and in- cisorlike. The two local species of Fundulus are separable by their markings, majalis of all ages being barred or streaked with black while the adult heteroclitus is not. Common mummichog Fundulus heteroclitus (Linnaeus) 1766 Killifish; Salt-water minnow; Chub; Mummy Jordan and Evermann, 1896-1900, p. 640. Description. — This is a stout-bodied little fish, about one-fourth as deep as long, its body thickest just back of the pectoral fins, whence it tapers to the tail. Both its back and its belly are rounded, but the top of the head is flat between the ej^es. The snout, as seen from above, is blunt. The mouth is at the tip of the snout and is so small that it does not gape back to the eye. Perhaps the most striking feature of Fundulus is its very deep caudal peduncle and rounded caudal fin. The fins are of moderate size, the dorsal situated behind the middle of the body above the anal, the pectorals broad and rounded. Both head and body are covered with large rounded scales. On males in breeding condition the scales on the sides of the head and those on the flanks below and behind the dorsal fin develop fingerlike processes on their free edges, called "contact organs." The mummichog shows a striking sexual dimor- phism in the dorsal and anal fins, which are not only larger in the male than in the female, and the anals of a different shape,90 but are more muscular and are used as claspers in the act of spawning. Color. — Males and females differ in color as well as in the sizes of the fins. Out of breeding season the males are dark greenish or steel blue above, with white and yellow spots, and marked on the sides with narrow irregular silvery bars or mot- tlings made up a series of dots. The belly is white, pale yellow, or orange; the dorsal, anal, and caudal fins are dark green or dusky with pale mottlings; the front edges of the anal and of the ventrals are yellow. Sometimes there is a dark- edged, pale-centered eyespot on the rear part of the dorsal fin. At spawning time the pigmenta- tion of the male is generally intensified, the back and upper sides darkening almost to black, while the yellow of the belly becomes more brilliant and the body generally takes on steel-blue reflections. The females (much paler than the males) are uni- x> A detailed account of the sexual differences is given by Newman (Biol. Bull., vol. 12, No. 5, 1907, pp. 314-348). <« ^^palso, as is the number of its vertebrae. But these last two characters are matters for the specialist. Color. — Dark blue gray on the back and on the upper part of the sides, silvery lower down on the sides, and below; the dorsal fin is plain gray, the rear margin of the pectorals with a very narrow pale edging. Size. — The species heterurus is one of the larger flying fishes, commonly growing to a length of about 1 foot (to the base of the tail fin). Occurrence in the Gulf oj Maine. — A flying fish, about 9% inches long to the fork of the tail, seem- ingly of this species but not in good enough con- dition for certain identification, was taken in a trap of the Pond Village Cold Storage Co. at North Truro, on the Massachusetts Bay shore of Cape Cod, on August 4, 1952. This is the only record of one of its tribe, from our Gulf. And the only record of a flying fish from Nova Scotian coastal waters is by Jones, of one taken at Sable Island, in 1859. Flying fishes are taken now and then at Woods Hole, the species heterurus perhaps more often than any other, according to published report, but several of the kinds to be expected in the Gulf Stream off our coast resemble one another very closely indeed. So we suggest that if a flying fish should be taken in our Gulf that does not seem to fit the accompanying illustration (fig. 83A) it be forwarded either to the Fisheries Laboratory of the U. S. Fish and Wildlife Service, Woods Hole, Massachusetts; to the Department of Fishes, U. S. National Museum, Washington, D. C; or to the Department of Fishes, Museum of Com- parative Zoology, Cambridge, Massachusetts, to be named. Figure 83A. — Flyingfish (Cypselurus heterurus). After Bruun and a specimen from North Truro, Mass. Drawing by Jessie Sawyer. FISHES OF THE GULF OF MAINE 173 THE SILVER HAKE AND COD FAMILIES— FAMILIES MERLUCCIIDAE AND GADIDAE The silver hakes and the cods are so closely allied that many European ichthyoloigsts group them in a single family. American practice, how- ever, is to separate them because of certain dif- ferences in the structure of the skull and ribs. They are soft-finned fishes, lacking true spines at any stage in development (though in one local species, the silver hake, the basal parts of the dorsal and anal fin rays are so stiff as to feel like spines to the touch), but they are distinguishable from all other soft-rayed Gulf of Maine fishes by the fact that their large ventral fins are situated under the pectorals or in front of them, and not behind them, as they are in the herrings and salmons. They and their relatives, the grenadiers (p. 243), are separated from most of the typical spiny-rayed fishes by the structure of the skull.21 Key To Gulf Of Maine Hakes, Cods, And Other Species 1. There are three separate dorsal fins and two anal fins 2 There are two separate and well developed dorsal fins 5 There is only one well developed dorsal fin 11 2. The lateral line is black; there is a black blotch on each shoulder Haddock, p. 199 The lateral line is pale; there is no shoulder blotch 3 3. The lower jaw projects beyond the upper; the chin barbel is very small, if there is one Pollock, p. 213 The upper jaw projects beyond the lower; the chin barbel is large 4 4. The ventral fins are narrow, and prolonged as filamentous feelers that are as long as the rest of the fin; the eyes are small Tomcod, p. 196 The ventral fins are broad, and their filamentous tips are less than one-third as long as the remainder of the fin; the eyes are large Cod, p. 182 5. The anal fin originates considerably in front of the point of origin of the second dorsal fin Hakeling, p. 233 The anal fin originates under the point of origin of the second dorsal fin or behind it 6 6. The ventral fins are short and of ordinary form Silver hake, p. 173 The ventral fins are very long and feeler-like 7 7. The first dorsal fin is hardly higher than the second dorsal, and none of its rays are prolonged or filamentous Spotted hake, p. 230 The first dorsal fin is much higher than the second dorsal, with one or two long filamentous rays 8 8. The ventral fins reach nearly or quite as far back as the rear end of the anal fin Long-finned hake, p. 232 The ventral fins do not reach back to the middle of the anal fin 9 9. The anal fin is so deeply notched about midway of its length as to suggest two separate fins Blue hake, p. 233 The anal fin is of about equal height from end to end 10 10. There are about 140 rows of scales along the lateral line from gill opening to base of caudal fin; the upper jaw bone reaches back to below the rear edge of the eye White hake, p. 221 There are only about 1 10 rows of scales along the lateral line; the upper jaw bone reaches back only as far as the rear edge of the pupil --- Squirrel hake, p. 223 11. There are no isolated rays in front of the dorsal fin, nor barbels on the top of the snout Cusk, p. 238 The dorsal fin is preceded by a fringe of short rays and one long ray; the top of the snout bears barbels as well as the chin 12 12. There are three barbels on the top of the nose -- Four-bearded rockling, p. 234 There are only two barbels on the top of the nose Three-bearded rockling, p. 237 Silver hake Merluccius bilinearis (Mitchill) 1814 Whiting; New England hake Jordan and Evermann, 1896-1900, p. 2530. Description. — The presence of two separate and well developed dorsal fins, both of them soft-rayed, the second much longer than the first, combined with the location of the ventrals on the chest, is sufficient field mark to distinguish the silver hake from all other Gulf of Maine fishes except for the true hakes (genus Urophycis, p. 221). And there is no danger of confusing it with any of the latter, for it lacks the chin barbels so characteristic of them, and its ventrals are of the ordinary finlike form, whereas those of the true hakes are altered into long feelers. It is a rather slender fish, about five to six times as long as it is deep, its body rounded in front of the vent but flattened sidewise behind it, with large flat-topped head occupying " The hypercoracoid bone lacks an aperture (technically a "foramen"). 174 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE ■ ■■'%■ .fzwm*,.^,. ■"*■■' Hm aBswww^^^^A ^W Figure 84. — Silver hake {Merluccius bilinearis) . A, adult, Nova Scotia, from Goode, drawing by H. L. Todd; B, egg; C, larva, 6.5 mm.; D, larva, 11 mm.; E., young fry, 23 mm. B-E, after Kuntz and Radeliffe. about one-fourth of the total length; large eyes; and wide mouth armed with two or more rows of sharp recurved teeth, and with the lower jaw projecting beyond the upper. The first dorsal fin (11 to 14 rays) originates close behind the gill openings, is roughly an equi- lateral triangle in shape, and is separated by a short space from the second dorsal. The second dorsal (38 to 41 rays) is about four times as long as the first dorsal, but hardly more than half as high, and is of distinctive outline, being deeply emarginate two-thirds of the way back, with the rear section the higher of the two. The anal fin (38 to 41 rays) corresponds in height and in shape to the second dorsal, under which it stands. The caudal fin is square tipped when widespread, but its rear margin is weakly concave, otherwise. The pectorals are rather narrow, their tips slightly rounded, and they reach back far enough to over- lap the second dorsal a little. The ventral fins, situated slightly in front of the pectorals, are perceptibly shorter than the latter, with about half as many rays (7) . Color. — The silver hake is dark gray above of brownish cast; but silvery -iridescent, as its name implies, or with golden reflections. The lower part of its sides and its belly are silvery. The inside of its mouth is dusky, the lining of its belly blackish. The fish is brightly iridescent when taken from the water, but fades soon after death. Size. — Maximum size about 2% feet long and about 5 pounds in weight, but adults average only about 14 inches long. Habits. — Silver hake are strong swift swimmers, well armed and extremely voracious. They prey on herring and on any other of the smaller school- ing fish, such as young mackerel, menhaden, ale- wives, and silversides. Probably a complete diet list would include the young of practically all the common Gulf of Maine fishes, for Yinal Edwards FISHES OF THE GULF OF MAINE 175 recorded the following from silver hake taken at Woods Hole: alewife, butterfish, cunner, herring, mackerel, menhaden, launce, scup, silversides, smelt, also the young of its own species. A 23 %- inch silver hake, taken at Orient, N. Y., had 75 herring, 3 inches long, in its stomach.22 And it is probable that the silver hake that frequent Georges Bank feed chiefly on young haddock. They eat squid when occasion offers. The small ones in particular prey regularly on large shrimp (Pandalus) in the deep troughs in our Gulf, where experimental trawlings by the Atlantis in the summer of 1936 took about four times as many silver hake at stations where these shrimps were abundant as at stations where shrimp were scarce.23 They sometimes take crabs, and bite freely on almost any bait, such as clams or cut fish. Though silver hake do not school in definite bodies, multitudes of them often swim together, and such bands sometimes drive herring ashore, and strand themselvas, in the pursuit. Events of this sort are oftenest reported in early autumn when the spent fish are feeding ravenously after the effort of spawning, but this may also happen at any time during the summer. Thus, Prof. A. E. Gross saw the beach at Sandy Neck, Barnstable, Mass., covered with them on several occasions in June and July 1920. 2i Doctor Huntsman informs us that spent fish frequently strand on the beaches on both sides of the Bay of Fundy in September. Wo once saw an army of silver hake harrying a school of small herring on a shelving beach at Cohasset, Mass. We half filled our canoe with pursuers and pursued, with our bare hands. It is said that European silver hake rest on the bottom by day and hunt by night, and it is usually at night that the American fish run up into the shallows and enter the traps. But strandings also take place by day. Silver hake, like many other rapacious species, are wanderers, independent of depth within wide limits, and of the sea floor. Sometimes they swim close to the bottom, some- times in the upper levels of the water, their vertical movements being governed chiefly by their pursuit of prey. Their upper limit is the tide line; at the other extreme they have been trawled repeatedly as deep as 150 to 400 fathoms on the continental slope off southern New England, and as deep as 296 fathoms off North Carolina.25 When they are on bottom they are caught in- differently on sandy or pebbly ground, or on mud (as in the deep trough west of Jeffreys Ledge, p. 175); seldom around rocks. The lowest temperatures in which we have known of silver hake being taken have been between 38° and 40° F. (probably), in the bottom of the deep trough west of Jeffreys Ledge, August 1936, 26 about 40° F. (4.4° C.) at 28 fathoms off New York, February 28, 1929, and about 39.5° F. (4.2° C.) at 19 fathoms in the same general region, February 5, 1930.27 And most of the winter and early spring records for it have been where the bottom temperature was warmer than about 43° F. (6° C.).28 At the other extreme, we have never heard of them in any numbers where the water was warmer than about 64° F. (18° C.) ; the monthly catches made in Cape Cod Bay (see p. 180) are especially instructive in tins regard. Breeding habits. — The silver hake is the most important summer spawner among Gulf of Maine fishes that are important commercially, just as the haddock is for spring and the pollock for autumn. The Gulf is probably its most prolific nursery, too, and it spawns over the outer part of the Nova Scotia Banks also, as far east as Sable Island, Dannevig 20 having recorded large egg catches in the offing of Halifax. But this is probably its eastern breeding limit, for the Canadian Fisheries Expedition found no silver hake eggs or fry on Banquereau or Misaine Banks; in the Laurentian Channel; or on the Newfoundland Banks. In the opposite direction, eggs in fair numbers have been taken in the tow nets off Woods Hole in July and August ; the Albatross II has found them and the resultant larvae near shore off Long Island in June and July, with eggs as far south as the offing of Cape May; and the young fry have been caught off New York 30 from spring to autumn. We have no evidence that silver hake commence to spawn before June, north of Cape Cod, our earliest egg record having been for the 1 1th of that " Nichols and Breder, Zoologica, N. Y. Zool. Soc, vol. 9, 1927, p. 163. » For details, sec Bigelow and Sehroeder, Biol. Bull., vol. 76, 1939, p. 315. « The Auk, vol. 40, 1923, p. 19. « Goode and Bean, Smithsonian Contrib. Knowl., vol. 30, 1895, p. 387. » No temperature was taken, but 38.6" F. (3.66° C.) was recorded there at 85 fathoms, on August 15, 1914, and 39.8° F. (4.33° C.) at 72 fathoms on August 15, 1913. 37 Specimens trawled by Albatross II. 18 Albatross II trawled a considerable number at stations scattered along the continental slope, from the offing of southern New England to the offing of Chesapeake Bay, in February 1929 and 1930, and in April 1930. » Canad. Fish. Exped. (1914-1915), 1919, p. 27. » Nichols and Breder, Zoologica, New York Zool. Soc, vol. 9, 1927, p. 163. 176 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE month; in our Gulf, egg production (as evidenced by the numbers of eggs taken in our tow nets) is at its height in July and August and continues through September, though less freely, with October 22 as our latest date. Similarly, the Canadian Fisheries Expedition found no silver hake eggs in Nova Scotia waters east of Cape Sable in May, but many in July. It is impossible to establish the exact tempera- ture at wbich silver hake are spawning at any particular station without knowing at what level ripe fish are in the water, which may be anywhere between the surface and the bottom with this species. It may be definitely stated, however, that they never spawn in as cold water as cod and haddock usually do in the western Atlantic. In 1915, for example (a representative season), it was not until the entire column of water was slightly warmer than 41° F. at the locality in question that we found the first silver hake eggs in our Gulf. And if the parent fish were in the upper water layers, as they may have been, all the rich spawnings we encountered in the Gulf during that year, and during the next, took place in tem- peratures considerably higher still. Similarly, the silver hake eggs towed off Halifax by the Canadian Fisheries Expedition in July 1915, and off Shel- burne, Nova Scotia, by the Grampus on September 6 of that same year may have been spawned in water warmer than 50° F., there being no need to assume that the parent fish were lying in the colder bottom stratum. As the spawning season draws to its close, in September and October, the minimum temperatures for most of our egg stations have been higher than 46°, with one (our latest record for the season) as warm as 57° F. at all depths. These data point to 41° to 45° F. as the lowest temperature limit for the spawning of the silver hake, with most of the eggs produced at 45° to 55° F. In the case of any fish producing buoyant eggs the tendency of the latter to rise (unless counter- acted by active vertical circulation of the water) insures that their development shall take place at the temperature of the upper stratum of water, not at that of the deeper levels where they were spawned. And the silver hake is no exception to this rule. While we have towed its eggs in June, when the surface was still only about 42° F., most of the egg records, and all our rich catches, were all made where the upper 5 fathoms or so were warmer than 50° and usually warmer than 55° F., with the temperature of the immediate surface 60° or higher in most cases. Similarly, silver hake eggs taken off Halifax by the Canadian Fisheries Expedition in July 1915, and off Shel- burne, Nova Scotia, by the Grampus on September 6 of that year, may well have been in water at least as warm as 53° F., there being no reason to suppose they were far below the surface.3' All this suggests that incubation does not proceed normally in water cooler than about 50°, and that it is most successful in temperatures as high as 55° to 60° F. This evidence that while the eggs of the silver hake may be spawned in low tem- peratures, a comparatively warm surface layer is necessary for their later development, offers a reasonable explanation for the failure of this fish to breed successfully along the New Brunswick shore of the Bay of Fundy, where active vertical circulation maintains surface temperatures as low as 50° to 55° F. throughout the summer, at least in most years. At the other extreme, the failure of the eggs that had been fertilized artificially to develop in the hatchery at Woods Hole in August temperatures points to 65° to 70° F. as the upper limit to successful incubation. According to Kuntz and Kadcliffe 32 only part of the eggs mature at one time, but we know of no estimate of the number of eggs a single female may produce. The eggs are buoyant, transparent, about 0.88 to 0.95 mm. in diameter, with a single yellowish or brownish oil globule of 0.19 to 0.25 mm. Incubation is rapid; Kuntz and Kadcliffe assumed a duration of 48 hours at Woods Hole, but it has not been determined for the cooler waters in the Gulf of Maine. The larvae are about 2.8 mm. long at hatching, slender, with small yolk sac, and they are made recognizable by the fact that the vent is located on one side, near the base of the larval fin fold, as is the general rule in the cod family, not at its margin as in most larval fishes, and that the trunk behind the vent is marked with two black and yellow cross bars. The dorsal and anal fins and the caudal fin have all assumed their definite outlines by the time the little fish is 10 to 11 mm. long, and fry of 20 to 25 mm. begin to resemble their parents in general appearance. 31 These catches were all made either at the surface or in oblique hauls with open nets. >» Kuntz and Radcliffe (Bull. U. S. Bur. Fish., vol. 35, 1918, p. 109) describe the spawning and early development. FISHES OF THE GULF OF MAINE 177 Evidently the young silver hake take to the deeper water layers toward the end of their first summer or that autumn, when about 1 to 3 inches long, for fry as small as this have been trawled in good numbers off southern New England at 150 fathoms and deeper at that season during the early explorations of the U. S. Fish Commission,33 by the Albatross II off Rockaway and off Long Beach, N. Y., in November 1928. By February they may be anywhere from 1% to 5 inches long, de- pending on how early they are hatched, on when they take to bottom, and on the feeding conditions they find there.34 They may be anywhere from about 2 inches long to aboit 6% inches long by April.35 The sizes of the many small silver hake that we have collected at different times of the year, both within the Gulf of Maine and southward as far as the offing of Chesapeake Bay, suggest that they reach an average length of 5% to 7% inches when 1 year old, and of about 9^ to 11 inches at 2 years of age, i. e., in their thud summer.36 Fish of 1 1 to 14 inches that dominated the pound-net catches made near Provincetown, August 1939,37 were three-year-olds, probably. The rate of growth has not been traced for the older fish, nor can it be deduced from that of the European silver hake for the latter grows to a considerably greater length, averaging as much as 30 inches at 8 years in the extreme northerly part of its range (Iceland) and considerably larger still, in the southern part (Gulf of Gascony and off Morocco).38 But it is reason- able to assume that the growth of the American fish varies similarly with the latitude (i. e., that it is most rapid in high temperatures) and that the American female, like the European, grows faster than the male. The European Merluccius ma- tures at 2 years, which is probably true of the American species as well. General range. — Continental shelf of eastern North America, northward to the Newfoundland « Goode, Fish, and Fish. Ind. U. S., Sect. 1, 1S84, p. 242. " Fry taken In February of different years by Albatross II have ranged from 1M inches (31 mm.) long to 4H inches (120 mm.). » In April 1930 Albatross II trawled young fry ranging in length from 2 inches (64 mm.) to 6% Inches (163 mm.) long at a number of stations from the offing of Rhode Island to the offingof Chesapeake Bay, at 14 to 85 fathoms. » For further details, see Bigelow and Schroeder (Biol. Bull., vol. 76, 1939, pp. 319-320, fig. 8). " Information supplied by Wm. A. Ellison, Jr. " Belloc, Notes et Memoires No. 21, Office Seientifique et Technique des Peches Maritimes, France, 1923. Banks, southward to the offing of South Carolina;39 most abundant between Cape Sable and New York. It is represented farther offshore and in the Gulf of Mexico by forms, the relationship of which to the Merluccius of our northeastern coast has not yet been determined. The silver hake is represented in Europe by a close relative, the European hake (Merlvccius merluccius), an excel- lent account of the natural history and migrations of which is given by Le Danois.40 Occurrence in the Gulf of Maine. — Silver hake are familiar fish all around the coasts of the Gulf of Maine from Cape Cod to the Bay of Fundy and to the west coast of Nova Scotia. But it has long been a matter of common knowledge that their chief center of abundance is in the southwestern part of the Gulf. Thus in 1945 (most recent year for which detailed regional statistics are available), the reported landings were between 46 and 47 million pounds " from off eastern Massachusetts in general, including the shores of Cape Cod out to the western slope of the so-called South Channel, contrasting with only about 4 million pounds for the western and central coasts of Maine, and with only about 6,500 pounds for eastern Maine. Silver hake, it is true, are said to be common in the Passamaquoddy region (more so in some years than in others), also around Grand Manan at the mouth of the Bay of Fundy. And they are re- ported at various localities along the Nova Scotia side of the Bay and along western Nova Scotia. But they are not mentioned in the statistics of the Canadian catches for these waters, hence cannot be very plentiful there. Silver hake are numerous over the west-central deeps of the Gulf also; in fact we found this the most plentiful fish at 70 to 90 fathoms in the basins off Cape Cod in the southwestern part of the Gulf and off Mount Desert in the northeastern, in August 1936; also in the trough west of Jeffreys Ledge, where the catches of them averaged 292 fish (maximum S40, minimum 1) as reduced to the common standard of one hour's trawling with an 82-foot shrimp trawl. And it is interesting that the catch there averaged about four times as great » The silver hake has been said, repeatedly, to range southward to the Bahamas, in deep water, following Jordan and Evennann (Bull. 47, U. S. Nat. Mus., Pt. 3, 1898, p. 2630). But the most southerly positive record we have found for it is off Charleston, S. C. (Blake Sta. 313, lat. 32° 32' N., long. 78° 45' W.; Goode and Bean, Smithsonian Contrib. Knowl., vol. 30, 1895, p. 387). « Notes et Mem., 2, Off. Sci. Tech. PGchps Maritimes, France, 1920. " "Round" and dressed fish combined. 178 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE (494 fish) at the stations where shrimp (Pandalus) were plentiful as at the stations where these were scarce (114 fish), evidence that silver hake con- gregate where feeding conditions are good.42 Reported landings throw little light on the numbers of silver hake that frequent the offshore rim of our Gulf, both because the otter trawls used there are so large-meshed that many pass through, and because most of those that are caught on Georges and Browns Banks are thrown overboard when the price is low.43 Experimental trawling, however, on Georges Bank, April to September 1913, yielded about one-seventh as many silver hake on the average (about 1,800 fish) as haddock (about 14,000 fish) per trip, and the Albatross III caught an average of about 150 silver hake, running about one-half pound in weight, per trawl haul, in 250 hauls on various parts of Georges Bank, July, August, and Septem- ber of 1948, 1949, and 1950. Thus they are moderately plentiful at least over Georges Bank as a whole, and there is no reason to doubt that this applies to Browns Bank equally. These catches do not suggest any definite con- centration on any one part of the bank, at least for summer, except that the largest that were made on its northern part were in hauls from shoaler than 30 fathoms, whereas the largest catches on the southern part were in hauls from deeper than 60 fathoms, a difference which may well have been a matter of the food supply.44 In April, however, of 1950, the silver hake were not only more plentiful along the northern edge of the bank (average 305 per haul) than on the southern part (average 77 per haul) but so strictly confined to the deeper levels that the total yield of 66 trawl hauls at shoaler than 60 fathoms was only 11 fish, contrasting with an average catch of 232 fish per haul at 60 fathoms and deeper (25 hauls).45 Silver hake spawn along the entire coastal zone from Cape Cod to Grand Manan, as proved by the locations of the egg catches (fig. 85). The sloping « For further details, see Bigelow and Schroeder (Biol. Bull., vol. 76, 1939, p. 308, table 1; p. 314, table 5. <> Reported landings, 1945-1947, ranged between 3,000 and about 33,000 pounds for Georges Bank, between 0 and 6,000 pounds for Browns. ** The average catch per haul was 262 fish from shoaler than 30 fathoms and 151 fish from deeper than 00 fathoms on the northern part of the bank; 90 fish per haul from shoaler than 30 fathoms and 285 fish per haul from deeper than 60 fathoms on the southern part. *• Twenty-one trawl hauls at 60 fathoms and shoaler yielded none at all in March; but no hauls were made in that month deeper than 60 fathoms, where the silver hake doubtless were. sandy bottom around the northern extremity of Cape Cod and off the eastern slope of the Cape evidently is an important center of reproduction. Thus we found an abundance of eggs off Race Point on July 7, 1915; our tow nets yielded many eggs at two stations off the outer shore of the Cape on July 22 of the following year, when a 1 5-minute tow there at 20 fathoms, with a net one meter in diameter, produced approximately 25,000 larvae of 3 to 7 mm., the richest haul of young fish we have ever made in our Gulf. And the fish were still spawning there a month later, as proved by the presence of eggs. Other occasions when we have taken silver-hake eggs in large numbers have been off Duck Island near Mount Desert on July 19 and on August 18, 1915; near Monhegan Island, August 4, 1915; off Wooden Ball Island near the mouth of Penobscot Bay on August 6, 1915; and off Rye, N. H., on July 23 of that same year. But we have never found them in any number in Massachusetts Bay though some eggs have been taken there on several occasions (fig. 85). Unfortunately, no quantitative hauls were made at any of the more productive egg stations, hence the number of silver-hake eggs present in the water cannot be approximated. But the vertical net yielded about 190 eggs per square meter of sea surface at one station in the eastern basin. Apparently the silver hake does not breed suc- cessfully in the northern side of the Bay of Fundy for neither its eggs nor its fry have ever been found there. But the capture of a few eggs in Petit Passage in our tow nets on June 10, 1915, suggests that it may spawn on the southern side of the bay as the cunner does (p. 478). And it may be expected to do so along the west coast of Nova Scotia, for the Canadian Fisheries Expedition found eggs at several stations off outer Nova Scotia, eastward to the longitude of Canso. The presence of silver hake on Georges Bank throughout the summer is presumptive evidence of local spawning, though we have taken no silver hake eggs or larvae there. The locations where we have found its eggs suggest that the silver hake, in the Gulf of Maine, spawns chiefly in water shoaler than 50 fathoms. But we have made one rich haul of its eggs in the center of the eastern basin. And the discovery of its eggs over the continental slope off Nova Scotia FISHES OF THE GULF OF MAINE 179 70° 60° Figure 85. — Localities where eggs (•), and larvae (O) of silver hake, or both (O) have been taken. by the Canadian Fisheries Expedition,46 with the presence of ripe fish as well as of spent, in depths as great as 150 fathoms and more off southern New England 47 proves that it spawns over deep water also. The European silver hake usually spawns in 50 to 100 fathoms. All our records for the free-drift ing larvae of the silver hake in the Gulf, unlike those for its eggs, have been in the southwestern part. And we have towed along the coast of Maine so often in August, September, and October (when the larvae spawned from June to August might have been expected) that our failure to find them east of Cape Elizabeth seems sufficient evidence that they actually are limited, in their regular occur- rence, to the southwestern part of the Gulf (they parallel the pelagic stages of the cod, the haddock, and the flatfishes in this) and to the waters westward from Cape Cod. Dannevig, too, has called attention to the absence of larvae of the silver hake in Nova Scotia waters, con- trasted with the presence of their eggs there.48 One possible explanation for this contrast between larvae and eggs is that it may mirror the relative percentage of eggs that hatch in the regions in question. A more likely explanation we think, when taken with other similar facts of distribution, is that it results from a peripheral drift around the shores of the Gulf from north- east to southwest, in which the eggs take part first and then the resultant larvae. This type of circulation, in fact, has been established so well for our Gulf by hydrograpliic evidence, that some such involuntary migration is inevitable, not only for various buoyant fish eggs and larvae that are produced near the coast line, but likewise for the drifting communities of invertebrates, and of plants. It is now known that large numbers of the silver hake that descend to the deeper water layers in the southwestern part of the Gulf during their first autumn remain there during the following " Dannevig, Canadian Fish. Exped., (1914-15) 1919, p. 28. «' Goode, Fish. Ind. U. S., Sect. I, 1884, p. 242. " Canad. Fish. Exped. (1914-1915) 1919, p. 28. 180 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE year, some of them still longer. Thus our ex- perimental trawlings, in August 1936, yielded large numbers of the 1-year-olds at 70 to 90 fathoms in the deep basins off Cape Cod and west of Jefferys Ledge; also off Mount Desert.49 And nearly all of the silver hake that come close enough inshore to enter the traps in the south- western part of the Gulf, or to strand on the beaches there, are good sized individuals of 9 inches and larger. In fact, the only instance that has come to our notice of any considerable inshore catch of one-year-old fish (about 6 to 8 inches long) in the Cape Cod Bay region was near Provincetown, August 1939, when about 1,900 of them were taken during a 14-day period.60 Huntsman, however, reports all sizes near shore from yearlings on, in the Passamaquoddy region to the northeast, and in the neighboring parts of the Bay of Fundy. The silver hake 3 years old and older, that provide the commercial catches, sometimes ap- pear in the Cape Cod Bay-northern Massachusetts region as early as the last week in March, regularly by May. Welsh saw some fish, for example, in Ipswich Bay in March and April in 1913 (a fairly representative season), considerable numbers in May, and an abundance in June. And this may be taken as typical for the whole coast line of the Gulf south of Portland; also for Georges Bank, where the first silver hake were taken by the otter trawlers from April 27 to 29 in 1913, and on almost every trip thereafter. We have not been able to learn how early silver hake ap- pear on the coast of Maine cast of Portland, or off western Nova Scotia, where it is only within the past few years that any attention has been paid them. Around Cape Cod Bay, silver hake are usually the most plentiful in June ; disappear more or less during August and September; and reappear in numbers in October, though far fewer then than in June, as is illustrated by the average monthly catches made by a set of eight traps at North Truro, for the years 1946-1947 and 1950: 61 June, 185,200 pounds; July, 36,700 pounds; August, 1,206 pounds; September, 1,780 pounds; and October, 10,852 pounds. <> For further details, see Bigelow and Sehroeder, Biol. Bull. vol. 70, 1939, pp. 308, 319-320, fig. 8. M Information supplied by William A. Ellison, Jr. "Information supplied by the Pond Village Cold Storage Co., North Truro, Mass. Whether their withdrawal thence in summer is a matter of food, or whether they move deeper to escape the heat of summer is a question for the future. Farther offshore in the western side of the Gulf, and to the northward, silver hake are about as plentiful in July and August as they are in June, as indicated by the vessel landings at Boston and Gloucester; somewhat less so in Sep- tember and October. And what little information we have suggests that summer is the season of greatest plenty for them in the Bay of Fundy region, though there are far fewer of them there. The great majority of the silver hake vanish from the inshore waters of the Gulf during the late autumn, November seeing the last of them in Massachusetts and Cape Cod Bays, according both to our own observations and to general report. The latest catches made on Georges Bank during the experimental trawlings of 1913 were on De- cember 3 and 12. And though a few are brought in from the grounds off Massachusetts and Cape Cod during January, February, and March, the catches average less than Vno as great for those months as for the period May through October, as illustrated by the monthly landings by trawlers at Boston and Gloucester for 1947 :62 January 1.400 July 4,444,000 February 2,255 August 4,879,000 March 1,700 September 1,974,000 April 7,540 October 2,381,000 May 860, 000 November 438, 000 June 1,158,000 December 207,000 It is probable that the fish of the year and those that are only 1 year old winter in the deeper de- pressions near where they first took to the bottom. It is unlikely that fish as small fish as those we have trawled in these situations, in August, can travel far. The wintering ground of the Gulf of Maine pop- ulation of larger silver hake is not known. Many of them may winter near the sea floor in the deep open troughs of the Gulf,63 where the bottom water at 75 to 100 fathoms and deeper continues warmer than 39° F. (4° C.) even at the coldest time of year. Evidence in this direction is that it is only deeper than 60 fathoms that good April catches have been reported on Georges Bank (p. 180). It is also possible that part of them move out to the shelf off southern New England to winter, or B Pounds of round fish and dressed fish combined. " Practically no trawling is done in winter In the deepest parts of the Gulf. FISHES OF THE GULF OF MAINE 181 even to the continental slope as the European silver hake do. Scattered catches, in fact, of half- grown silver hake and larger are made by otter trawlers off southern New England, and off New York in January and February.64 But it seems more likely that these are fish that either remain there throughout the year or that visit the coasts of New York and of southern New England at other times of year, than that they come from the Gulf of Maine. Fluctuations in abundance in the Gulf of Maine. — Every shore fisherman in the Massachusetts Bay- Cape Cod region knows that silver hake vary widely in abundance from year to year. Catches by one set of six pound-nets at North Truro on Cape Cod yielded about 60,000 pounds in 1946; 237,000 pounds in 1948; 232,000 pounds in 1949; and only about 10,000 in 1944; but about 458,000 pounds in 1950. Yearly fluctuations of this sort are to be expected at any given locality, in the case of any predaceous wanderer. And there is nothing in the available record to suggest that a major alteration has taken place in the numbers of silver hake in its center of abundance in the Gulf, whether upward or downward, since it has been an important fish on the market. Occurrence to the westward and eastward of the Gulf of Maine. — Silver hake are described as abun- dant from October to December as far westward as New York, sometimes in May also, though few are seen there in summer. And yearly catches of some 2 to 5 million pounds of "whiting" by pound nets 65 suggest that the beaches of New Jersey rival those of the Cape Cod Bay region in the seasonal abundance of silver hake. But we have not heard of any great numbers of them close in shore beyond Cape May, though pound nets do take a few as far south as the mouth of Chesapeake Bay. Farther out on the shelf, silver hake of all sizes are to be found at all times of the year, from the offing of southern New England, westward and southward, in numbers large enough for otter trawlers to land 3 to 5 million pounds yearly in New York and New Jersey,66 and smaller amounts in Delaware. Eastward from our limits we find the silver hake described as abundant57 in outer Nova Scotian waters generally. But we have no clue as to their actual numbers there, relative to the Gulf of Maine, for they are not yet important enough commercially to be included in the Canadian fisheries statistics. The experimental cruises of the Newfoundland Fisheries Research Commission took them on Banquereau and Misaine Banks; in the northern side of Cabot Straits; on the southern part of the Grand Banks; and at Bay Bulls on the east coast of the Avalon Peninsula, which is the most northern record for them of which we chance to know. But it seems they are not known anywhere in the Gulf of St. Lawrence.68 Importance. — Silver hake are as sweet a fish as one could ask, if eaten fresh or if slack salted over night and used for breakfast next morning. But they soften so fast that there was no regular market demand for them of old, and most of those that were caught incidentally were thrown over- board. In fact, we can remember seeing them used locally for fertilizer. Thus only some 37,000 pounds were saved in Maine and Massachusetts combined, even as recently as 1895. But improved methods of freezing fish were followed by landings of about two million pounds by 1902; of between four and five million pounds in 1905, rising through the years of the first world war to more than 14 million pounds in 1919.69 The yearly landings then fell off, for some reason, to only about 6 million to 9 million pounds for the period 1924 to 1933, which was far less than the potential catch. But the landings then increased again, as frozen whit- ing became more popular in the Middle West, to about 15 million pounds in 1935, to about 40 mil- lion pounds by 1940, with from 46 million to 74 million pounds during the 6-year period 1942 to 1947.60 All but a small part of the Maine and Massa- chusetts landings, recorded in the following table, are from within the limits of the Gulf of Maine. The silver hake now ranks fourth or fifth among Gulf of Maine fishes in amount landed. But it " Albatross II trawled 8 fish, 7 to 9 Inches long, off New York, February 28, 1929, at 28 fathoms; and the dragger Eugene H., Capt. Henry Klimm, picked up 115 of market size In a week's trip, about 80 miles off Martha's Vineyard, at 47 to 67 fathoms, January-February 1950. « 1912, 5,313,300 pounds; 1945, 5,842,900 pounds; 1947, 1,784,500 pounds. " Otter trawl landings of "whiting," for New York and New Jersey com- bined, were 3,408,200 pounds In 1942; 5,243,700 pounds in 1945; and 7,498,600 pounds In 1947. Delaware trawlers reported 203,500 pounds for 1947. »' VTadykov and McKenzIc, Proc. Nova Scotia Inst. Sci., vol. 19, 1935, p. 72. » According to Dr. Huntsman all ostensible reports of their presence In the Gulf of St. Lawrence have been based In reality on the other hakes of the genus Urophycis (p. 221). And It is these that aro meant when "hake" are mentioned In the early publications of the U. S. Fish Commission, such as Balrd's (Rept. U. S. Comm. Fish. (1886) 1889, app. A.) report on the fisheries of eastern North America. B Landings for Maine and Massachusetts combined. " Maine and Massachusetts combined. 182 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE ranks only about seventh in value. In 1945, the year when the catch was largest, its value was $1,736,200. Its rank is low as a sportman's fish, for while it bites greedily, it puts up only a feeble resistance when hooked. Year Catch to nearest 1,000 pounds Year Catch to nearest 1,000 pounds 1919 14, 607, 000 6, 377, 000 7, S75, 000 7, 943, 000 6, 936, 000 6, 379, 000 8, 678, 000 15,420,000 21,038,000 1938 24,851,000 1924 1939 — 27, 539, 000 1929 1940 39. 990, 000 1930 1942 45, 900, 000 1931 1943 1944 48, 460, 000 1932 - — 47, 373, 000 1933 . 1945 73, 866, 000 1935 1946- 48, 844, 000 1937 - 1947 58, 936, 000 Cod Gadus callarias Linnaeus 1758 61 Rock cod Jordan and Evermann, 1896-1900, p. 2541. Description. — The most noticeable external char- acteristics of the cod, emphasized above in the gen- eral survey of the cod family (p. 173), are its three dorsal fins and two anal fins; its lack of fin spines; the location of its ventral fins forward of its pectoral fins, and the fact that its upper jaw pro- trudes beyond the lower; that its tail is usually nearly square, and that its lateral line is pale, not black. The cod is a heavy-bodied fish, only slightly flattened sidewise, its body deepest under the first dorsal fin (cod neither very fat nor very lean are about one-fourth to one-fifth as deep as they are long) , tapering to a moderately slender caudal peduncle, and with a head so large that it takes up about one-fourth of the total length of the fish. The nose is conical and blunt at the tip; the mouth wade, with the angle of the jaw reaching back as far as the anterior part of the eye; and there are many very small teeth in both jaws. The first dorsal fin usually (if not always) originates well in front of the midlength of the pectoral fins; it is the highest of the three dorsals, triangular, with rounded apex and convex margin. The second dorsal fin is nearly twice as long as the first dorsal and about twice as long as it is high, decreasing in height from front to rear with slightly convex margin. The third dorsal fin is a little longer than the first dorsal, and is similar to the second dorsal in shape. The caudal fin is about as broad as the third dorsal fin is long (rather small for the size of the fish) and broom-shaped. The two anal fins stand below the second and third dorsals, to which they correspond in height, in length, and in shape. The number of fin rays was as follows, in a large series of Gulf of Maine cod, 23 to 37 inches long, examined by Welsh. " Jordan, Evermann, and Clark (Rept. U. S. Comm. Fish. [1928], Pt. 2, 1930, p. 210) use the species name morrhua Linnaeus 1758. But the use of callarias accords better with modern practice, because it preceded morrhua on the same page of the Systema Naturae. Number of finrays Dorsal Anal First Second Third First Second 13 15 16 19 21 24 18 19 21 20 22 24 17 IS 22 Figure 86. — Cod (Gadus callarias), Eastport, Maine. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 183 As few as 12 rays have occasionally been re- corded for the first dorsal, 16 for the second, 17 for the third, 17 for the first anal and 16 for the second. The pectoral fins, set high up on the sides, reach back as far as the rear end of the first dorsal. The ventral fins are nearly as long as the pectorals in young cod but are shorter, relatively in large fish, with the second ray extending beyond the general outline as a filament for a distance almost one-fourth as long as the entire fin. Both the head and the body are clothed with small scales. Young cod are easily distinguished from large tomcod by their relatively broad ventral fins with slender filaments, by the location of the first dorsal fin, and by their larger eyes, as explained in the description of that species (p. 196) . The pale lat- eral line readily distinguishes the cod from the haddock; and the square-tipped tail, projecting upper jaw, and spotted color pattern of a cod give it an aspect quite different from that of the pollock. Color. — Cod vary so widely in color that sundry of its color phases have been named, but all of them fall into two main groups, the gray-green and the red. The back and upper sides of the former range from almost black through dark sooty or brownish gray, olive gray, olive brown, sepia brown, mouse gray, ashy gray, clay colored, and greenish to pale pearly (darker on the back than on the sides) ; the fins are of the general body tint, and the belly is whitish, usually tinged with the general ground color. The red or "rock" cod vary from dull reddish brown to orange or brick red, with white belly tinged with reddish, and with red, olive, or gray fins. In most cod the upper surface of the body, the sides of the head, and the fins and tail (but not the nose or belly) are thickly speckled with small, round, vague- edged spots. On the "gray" fish these are of a brownish or yellowish cast, darker than the gen- eral body color, while they are usually reddish brown or sometimes yellowish on the "red" fish. Occasionally one sees a spotless cod, but these arc unusual. The lateral line is invariably paler than the general body tint, pearly gray or reddish ac- cording to the hue of the particular fish in ques- tion, and it stands out against the darker sides. Size. — Cod sometimes grow to a tremendous A huge one of 21 1 % pounds and more than 6 chusetts coast in May 1895; 62 one that weighed 138 pounds dressed (hence must have weighed ISO pounds or more alive) was brought in from Georges Bank in 1838; and Goode 63 mentions several others of 100 to 160 pounds as caught off Massa- chusetts. But cod of a hundred pounds are exceptional, the largest New England cod of which we have heard recently being one of 90 pounds, that was taken off the coast of Maine early in July 1922. Even a 75-pound fish is a rarity, but 50 to 60 pounders are not unusual. The so-called "large" fish that are caught near shore run about 35 pounds; and "large" ones taken on Georges Bank about 25 pounds. But the shore fish, large and small together, average only be- tween 6 and 12 pounds in weight. The relationship between length and weight is usually about as follows for fish caught on the in- shore grounds between Cape Ann and Portland, though this varies with the condition of the fish and with their state of sexual development.64 Females Males Inches Pounds Inches Founds 19 to 20 . 2W-3 3Hr- 4 *V>- 7 5-7 7-9 7^-10 9 -13 12^-17^ 16 -23 18 -22 16 -32 29H-32 31 -51 50 54 20 to 21 3 - 3'i 23 to 24 4 - by, 23 to 24 25 to 26 6H- 8 25 to 26 27 to 28 7 - 8*4 27 in 28 30 to 31 7 -11 30 to 31. . 32 to 33 7 -13 32 to 33. . 34 to 35 12 -17 34 to 35 . 36to37 12H-U 36 to 36^ - 38 to 39 17 -21 38 to 39 40 to 41 19 -25 40 to 42 43 to 45 25H-29 43 !■> li 46 43 48H to 50^ 52 ... 57H A 99,'2-pound fish recorded by Earll was 62 inches long, and one of 100 pounds caught off Wood Island, Maine, on April 9, 1883, measured 65 inches, its head 17% inches. Any fish of 5% to 6 feet will weigh 100 pounds or more. Habits. — Cod in one place or another range from the surface down to 250 fathoms at least. During the first year after the young cod take to bottom (p. 186) many of them five in very shoal water, even along the littoral zone, and many young fry have been taken at Gloucester and else- where along the shores of New England, while size. feet long, was caught on a long line off the Massa- •> Jordan and Evermann, American Food and Oame Fishes. 1902, p. 514. " Fish Ind. U. S.. Sec. 1, 1884, p. 220. « Based chiefly on measurements given by Earll (Rept. V. S. Comm. Fish. |1878|, 1880, p. 734), and on a large series of cod measured fresh from the nets by Welsh during the spring of 1913. 184 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE many small cod are caught about the rocks only a fathom or two deep even in summer. But it is certain that many cod fry take to bottom on the offshore banks also, for we have trawled young fry at many localities between Nantucket Shoals and Browns Bank. As a rule, the large cod lie deeper than 7 or 8 fathoms in summer in our latitudes. But the fishing is often good in only 3 to 5 fathoms of water in wintertime, especially in Ipswich Bay. At the other extreme, comparatively few cod are caught much deeper than 100 fathoms within the Gulf of Maine. And although fisher- men sometimes do well at much greater depths on the slopes of the offshore banks, the 5- and 75- fathom contours probably include the great majority of all the cod living in the Gulf, summer or winter. The largest catches of cod are made on rocky and pebbly grounds; on gravel; on sand, and on a particularly gritty type of clay with broken shells. They also frequent the deeper slopes of ledges along shore, where they forage among the Irish moss (Chondrus crispus) and among sea- weeds of other kinds. Young red ones are espe- cially common in these situations, while one some- times catches a large rock cod as these dark brown or red fish are called. And the bottoms where cod and hake are found are so distinct that a long line set from a hard patch out over the soft sur- rounding ground will often catch cod at the one end, hake at the other. But fair catches are sometimes taken on mud, as off Mount Desert, where large- and medium-sized cod are regularly caught on soft ground in winter. And a few very large cod (35-60 lb.) have also been brought in from the mud bottom of the deep basin to the westward of Jeffreys Ledge (about 90 fathoms). The cod, as appears from the foregoing, is typi- cally a ground fish; except on some journey (a subject to be discussed later) or when following its prey, it usually lies within a fathom or so of the bottom. And large ones keep closer to the ground than small ones as a rule, so that the closer one fishes to bottom the larger the cod are likely to run. But even the large ones sometimes follow herring up to the surface; we have known of large cod gaffed from a vessel's side in Northeast Harbor, Mount Desert Island, in September, while they were chasing sardines. And they come to the surface more commonly on the Grand Banks and along the eastern coast of Labrador, when they are following capelin. Cod even strand on the Labrador beaches while harrying schools of capelin, but we have never known cod to strand anywhere around the coasts of the Gulf of Maine, as silver hake so often do (p. 175). The adult cod is at home in any temperature from 32° to 50°-55° F. ; in all but the superficial layers of the Gulf of Maine, that is, at all seasons. But experience at the Woods Hole hatchery, proves that freezing may be fatal by the forma- tion of anchor ice. On the other hand, while large cod tend to avoid water warmer than about 50° F., except that they are abundant at times in temperatures as high as 58°-59° F. on Nan- tucket Shoals (the most southerly year-round cod-ground in the Atlantic). Small cod are somewhat less sensitive to heat than large, a fact reflected in the presence of greater numbers of them in shoal water in summer than of larger fish. The relationship of the spawning of the cod to temperature is discussed below (p. 194). Food. — When the larval cod first breaks from the egg it subsists on the yolk with which its abdomen is distended (fig. 88), as do most other sea fishes. But this source of nutriment is com- pletely absorbed by the sixth day after hatching, and the future existence of the little fish depends as much on finding a plentiful supply of food as on escaping the enemies by which it is encom- passed. So far as known, the larval and post- larval cod subsist almost exclusively on copepods and on other minute Crustacea, during the several months while they are drifting in the upper layers of water.65 And this same diet, varied with amphipods, barnacle larvae, and other small crustaceans, as well as with small worms, is the chief dependence of the little cod when they first seek the bottom 68 but as they grow larger they consume invertebrates in great variety and in enormous amount. Mollusks, collectively, are probably the largest item in the cod's diet in the Gulf of Maine; any shellfish that a cod encounters is gobbled up, so that their stomachs are mines of information for students of mollusks. Large sea clams (Mactra), •» Bumpus, Science., N. Ser„ vol. 7, 1898, p. 485. •* For further details on the diet of cod larvae and fry, see Brook (5 ann. Rept., Fish. Board Scotland (1SS6) 1887, p. 327), Mcintosh and Masterman (British Marine food flsbes, 1897, p. 242), Kendall (Rept. U. S. Comm. Fish. (1896) 1898. p. 179), Bumpus (Science, N. Ser., vol. 7, 1898 p. 485), and Good- child, Graham and Carruthers (British Mlnist. Agric. Fish., Fish. Inv., Ser. 2, vol. 8, No. 6, [1925) 1926. FISHES OF THE GULF OF MAINE 185 the empty shells of which are often found neatly nested in cod stomachs: cockles (Polynices) ; and sea mussels (Modiolus) are staples, all of which they swallow whole. Cod also eat crabs, hermit crabs, lobsters (large and small) , shrimps, brittle stars (of which they are sometimes crammed full) , sea urchins, sea cucumbers, and sea worms {Nereis). Brittle stars and small crabs, for example, had been the chief diet of the cod examined by Welsh on the Isles of Shoals-Boon Island ground in April 1913, while Wilcox 67 states that a number of 17-pound fish caught in Ipswich Bay were full of large red prawns 2 to 4 inches long (evidently the northern edible shrimp Pandalus). And we have found crabs {Cancer; Libinia) the chief food of the cod on Nantucket shoals. Tunicates (sea squirts) also bulk large in their diet. Occasionally they eat hydroids, bryozoans, and algae, perhaps taking these for the amphipods that are hidden among them. And in late summer cod frequently feed on ctenophores {Pleurobrachia pileus). But while its diet list would probably prove almost as extensive as that of the haddock (p. 202), the cod shows so decided a preference for large shells rather than for small ones that the stomach contents of cod and haddock taken side by side differ noticeably. Nor is it likely that cod root the bottom as haddock do (p. 202), for worms. Cod pursue and gorge on squid at every oppor- tunity, and on various small fish, particularly on herring, on launce, and (in the north) on capelin; also on shad, mackerel, menhaden, silversides, alewives, silver hake, young haddock, and even on their own young, rising into the upper waters for this purpose when necessary (p. 1S4). They also pick up flounders, cunners, rock eels {Pholis), blennics, sculpins, sea ravens, small hake and skates from the bottom. In fact, they take any fish small enough to swallow, including the hard slim alligatorfish (p. 457) and even the sea horse (p. 315). And Welsh noted that many cod taken near the Isles of Shoals on May 1, 1913 spat up small rosefish from 4 to 6 inches long. The eggs of the longhorn sculpin 6S and of the eelpout {Macro zoarces) 69 also have been found in cod « Bull. U. S. Fish. Comm., vol. 6, 1887, p. 95. « Warfel and Merrlman, Copela, 1944, p. 198. '• Olsen and Merrlman, Bull. Bingham Oceanogr. Coll., vol. 9, art. 4, 1946, p. 77. 210941—53 13 stomachs. Adult cod as well as small are also known to feed on pelagic shrimps in the waters around Iceland,70 but we have never heard of them doing so in the Gulf of Maine. Even a wild duck does not escape from a large cod now and then. Thus we have heard of several scoters found in the stomachs of large fish caught off Muskeget Island in 1897; and though sea fowl are not a normal article in their diet, the flesh of the greater shearwater (hagdon) has long been considered excellent cod bait. Objects as indigestible as pieces of wood and rope, fragments of clothing, old boots, jewelry, and other odds and ends have repeatedly been found in cod stomachs. And they often swallow stones; but probably for the anemones, hydroids, and other animals growing thereon, and not to take on ballast for a journey as the old story has it. Although cod are so rapacious they fast gener- ally while they are spawning; the stomachs of nearly all the ripe fish examined by Earll, and recently by Welsh, were empty. Experiments performed on the cod in captiv- ity,71 combined with the general experience of fishermen, suggest that they capture moving objects by sight. But apparently cod (and for that matter other fish), can see clearly only for a few feet, and their greediness in snapping up the naked meat of clams and cockles (foods which they never find in that condition in nature), added to the fact that they bite as readily by night as by day, seems sufficient evidence that they depend largely on smell. Enemies. — In the Gulf of Maine, large sharks and the spiny dogfish are the worst enemy of the adult cod. Formidable enemies of young cod fry are the small pollock which infest our harbors. These are so fierce that a single pollock 7 or 8 inches long will disperse a school of hundreds of cod fry, driving them to shelter among the weeds and rocks, while Earll remarks that in the aquar- ium a cod so fears a pollock of equal size that it will invariably hide if possible. Young cod, up to 7 to 8 inches, are also devoured in large numbers by the larger cod. to Schmidt (Skrlft. Komm. Havunderstfgelser, No. 1, 1904, p. 70) and Paulsen (Meddclel. Kommis. Havunders0gelser, Serle Plankton, vol. 1, No. 8, 1909, p. 39). " Bateson, Jour. Mar. Biol. Assoc. United Kingdom, N. Ser., vol. 1, 1889-90, p. 241. 186 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Migrations and wanderings. — It has long been known that cod carry out extensive migrations in some regions, but that they are more nearly stationary in others. European (especially the Scandinavian) biologists have succeeded in tracing the major outlines of their movements for North European seas, and enough evidence has accumu- lated to show that their travels fall into the same categories in the one side of the Atlantic as in the other. These categories are: (a) involuntary drifts by the eggs and by the larvae before they take to the bottom; (b) the various journeyings by the older cod in search of food; (c) journeys associated with the concentrations of cod on particular spawning grounds; and (d) regular seasonal migrations (with return movement) between different regions that are suitable for cod during different parts of the year. To begin with, the eggs, larvae, and young fry of the cod, like those of so many other sea fishes, drift helplessly with the current from the time they are spawned until they seek the bottom (a fact established by European observations too numerous to list).72 The length of this period (varying in duration in different seas) depends partly on whether the fry are near land or are far out at sea, and partly on whether they are floating over deep water or over shoal. It is not likely to last for more than two months for fish that are hatched on the inshore spawning grounds in the Gulf of Maine, where the bottom is within easy reach. Even so, it is extremely unlikely that any cod fry take to the bottom near where they were spawned. This matter is discussed further in relation to the occurrence of the cod in our Gulf (p. 190). The journeyings of the cod that are associated with their spawning are especially extensive along the Norwegian coast, where they have been the subject of much study, leading (among other things) to the very interesting probability that their journeys up and down the coast of Norway are chiefly involuntary, for the ripe fish drifting north become so fat that they tend to be suspended in the water near the surface, whereas the spent fish become so thin that they are deeper down in the water.73 But there is no reason to 71 In European seas young cod often live under the disks of tho large red Jellyfish (Cyanca), but they have not yet been found in this situation in the Oulf of Maine. " See especially Hjort, Journal du Consell, Cons. Perm. Internat. Explor. Mer, vol. 1, No. 1, 1926, p. 9. suppose that any of our Gulf of Maine cod need travel far to reach the localities where they spawn. In the extreme northern and southern fringes of their geographic range cod are regularly "migra- tory" in the common understanding of the term. Thus it is only in summer and early autumn that they visit the waters of the polar current along the eastern coast of Labrador, from which they withdraw again later in the autumn, to pass the winter and spring either to the southward or in deep water. On the other hand, it is only during autumn, winter, and early spring that cod are caught off the coasts of southern New England, of New York, of New Jersey, or further south. The fish that winter along this westerly and southerly extension of the cod's geographic range appear off southern Massachusetts in mid-October; off western Long Island and off the coast of New Jersey in November; they go back eastward again by the first part of May. And the numbers involved are large enough to support a profitable autumn- winter and early spring fishery from Nantucket to New Jersey. Tagging experiments carried out by the U. S. Bureau of Fisheries, first at Woods Hole in the winters of 1898-1 90 1,74 and in various parts of our Gulf on a much larger scale from 1 923-1 930,76 have shown that most of the fish that take part in this westerly movement pass the summers in the Nantucket Shoals region. But it is clear that a large part of the cod stock that summers on the Shoals fails to join this westerly mass movement in autumn, for fish tagged there in summer have been recaptured there the next winter, while many others have been recaught there the following spring. And it is established now that the great majority of the cod that live off our coasts from Cape Cod to northern Nova Scotia, in the south- ern part of the Gulf of St. Lawrence, and on the southern part of the Grand Banks, can fairly be termed "nonmigratory" in a broad sense. Breeding habits. — The cod is one of the more prolific fishes. A female 39 or 40 inches long may be expected to produce about 3,000,000 eggs yearly, one of 41 inches at least 4,000,000. And Earll estimated the number in a 52%-inch fish weighing 51 pounds at 8,989,094, with 9,100,000 in '< Smith, Rept. U. S. Comm. Fish (1901) 1902, pp. 193-208. " 22,884 fish tagged in the region of Nantucket Shoals, and about 30,000 hi other parts of the Oulf of Maine, including the offshore Banks, 308 flsh recaptured westward from Marthas Vineyard,. For further details, see Schroeder (Bull. U. S. Bureau of Fisheries, vol. 46, 1930, pp. 1-136). FISHES OF THE GULF OF MAINE 187 a 75-pounder. But the average production of eggs is perhaps not more than 1,000,000 for the general run of Gulf of Maine fish. The eggs are buoyant, transparent, without oil globule, and 1.10 to 1.82 mm. in diameter. Gulf of Maine eggs, artificially fertilized and measured by Welsh, averaged about 1.46 mm. in diameter, but the size varies somewhat with the temperature of the water, being larger in cold than in warm.79 The period of incubation for cod eggs depends on temperature. According to experience at the hatcheries, hatching may be expected in 10 or 11 days at 47° F., in 14 or 15 days at 43° F., in 20 to 23 days at 38° to 39° F. and not for 40 days or more if the water is as cold as 32° F. Fertilization can take place and development commence in tem- peratures even lower than this, as proved by ex- periments by Krogh and Johansen.77 But their observation that the mortality is great among eggs incubated at 32° F. (although full development can take place) corroborates the experience of the 11 Fish (Bull. U. S. Bur. Fish., vol. 43, 1929, p. 292) found cod eggs taken in the tow net in Massachusetts Bay to average about 1.63 mm. in February, smaller (1.46 to 1.49 mm.) in December and in May. " Dannevig, Canadian Fisheries Exped. (1914-1915)., 1919, p. 44. Figure 87. — Egg. After Heincke and Ehrenbaum. Figure 89. — Larva, 4.5 mm. After Schmidt. hatcheries, where it has proved impossible to hatch more than 25 to 50 percent of the eggs in water as cold as that. And the relative strength of the larvae that are hatched at different temperatures points to 41° to 47° F. as most favorable for in- cubation. All this suggests that extreme cold prevents the successful reproduction of the cod, not by interfering with spawning (for this can take place in the lowest temperatures to be found any- where in the open sea, p. 195), but by its effect on the developing eggs. And it is interesting that cod in the tank at Woods Hole produced eggs in Febru- ary, when the water may have cooled to 30° F. (and quite normally to judge from the fact that the eggs incubated successfully in the warmer water of the hatchery), for these same fish would have spawned naturally in temperatures at least as high as 36°-38° F. if they had been left at liberty. Newly spawned cod eggs are indistinguishable from those of the haddock, with which they inter- grade in size. But shortly before hatching, the pigment of the cod gathers in 4 or 5 distinct patches: one over the region of the pectoral fins, one above the vent, and the others equally spaced behind the latter (fig. 87) ; whereas in the haddock Figure 88.- — Larva, just hatched, 4 mm. After Masterman. Figure 90. — Larva, 9 mm. After Schmidt. Figure 91. — Fry, 20 mm. After Schmidt. Figure 92.— Young, 40 mm. After Schmidt. Cod {Gadus callarias), developmental stages, European. 188 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE the pigment cells are arranged in a row along the ventral side of the trunk (p. 203). There is also danger of confusing newly spawned cod eggs with those of the witch flounder (p. 287), which they over- lap in size; but the black pigment of the cod eggs identifies them as gadoid as soon as this appears, for the embryonic pigment of the witch is yellow. (See also haddock on p. 203.) Figure 93.- — Diagram of the pigmentation of the young larvae of the cod, A, and of the pollock, B. After Schmidt. The larvae are about 4 mm. long at hatching with the vent (which is close behind the yolk sac) located at the base of the ventral fin fold on one side instead of at its margin, so that the intestine seems to end blindly, as is also the case with haddock and pollock larvae. At this stage young cod much resemble the latter, but are separable from them by the fact that the pigment is in two dorsal and three (rarely two) ventral bars, with the dorsal bars shorter than the ventral bars opposite them, whereas the dorsal bars are longer than the opposing ventral bars (fig. 93) in pollock larvae up to 10 mm. long. Neither is there any danger of confusing cod larvae with haddock even at this early stage, for the latter are not barred but have a continuous row of pigment cells along the ventral margin of the trunk behind the vent, besides other patches on the nape and in the lining of the abdomen. The young cod float helplessly, when first hatched, yolk uppermost. But they assume the normal position in about 2 days; the yolk being absorbed and the mouth formed in 6 to 12 days, according to temperature, when the larvae are about 4.5 mm. long. As the little cod grows the pigment bars gradually fuse, and at 8 to 10 mm. a median band forms. Cod 10 to 20 mm. long may easily be distinguished from pollock by the fact that the pigment extends to the tail, whereas it ends abruptly some distance in front of the tail in the pollock. Haddock of this size show much less pigment (p. 203). Cod fry of 15 to 30 mm. are made recognizeable by the location of the vent under the second dorsal fin, combined with dense pigmentation. At 20 mm. the dorsal and anal fin rays have attained their final number and the separate fins are outlined, while at 30 mm. the fry begin to show the spotted color pattern so characteristic of the cod. Rate of growth. — In 1898 a large number of newly hatched larvae were released in December at Woods Hole in the "eel pond" (a lagoon freely communicating with the harbor and with a temper- ature about paralleling that of the outside water), where they grew to an average length of 50 to 100 mm. by the following June.78 The experiment was repeated in the winter of 1899 79 with similar results, as appears from the following table show- ing the growth of approximately 2 million freshly hatched larvae that were placed in the eel pond on January 1 1 . Date Extreme lengths Average length Date Extreme lengths Average length Apr. 8 Apr. 25 May 13 _ mm. 29 to 38.... 34 to 49.... 35 to 51.... mm. 32.9 40 42.8 May 25 June 6 . June 20 mm. 28 to 68.... 71 to 76.... 73 to 77.... mm. 64 75.5 75 Captures of young fry 1% to 3 inches long in the neighborhood of Cape Ann late in June (Earll 1880), and subsequently around Woods Hole and on Nantucket Shoals, show that cod hatched from January to March in the Gulf of Maine grow at about this same rate. But fish that are hatched in the rising temperatures of spring might be expected to grow faster during their first few months. European experience80 is to the effect that young cod are 4% to 8 inches long by the end of then first autumn, which probably applies equally to the Gulf of Maine. In later life cod grow at varying rates in different seas, and even fish that are caught in the same haul n Bumpus, Science N. Ser., vol. 8, 1898, p. 852. '• Smith, Bull. U. S. Fish Comm., vol. 19, 1901, p. 307. M Damas (Rapp. et Proces-Verb., Cons. Perm. Intern. Explor. Mer, vol.10, No. 3, 1909) gives an account of the European in .estigations on the life history of the cod, up to that date. FISHES OF THE GULF OF MAINE 189 may have grown at very different rates, as shown by the structure of their scales. Consequently, the length of a fish older than a yearling is no criterion to its age within 2 or 3 years. Wode- house's 81 studies on cod caught at the mouth of the Bay of Fundy and the Bureau of Fisheries investigations on Nantucket Shoals, suggest that cod grow more rapidly in the Gulf of Maine than in European waters, as follows: Age, in years Average length, in inches, Nan- tucket Shoals Average length, in inches. Bay of Fundy European (approx- imate average) 1 2 7-8 14-17 19-22 23-26 27-29 30-32 33-34 6 14 20 26 32 36 39 45 49 5 8 3 12 4 15 5 19 6 7 8 21 24 27 9 29 The fact that cod run much larger in the Gulf of Maine than in either the North Sea or the Norwegian Sea, and that those of 75 pounds and heavier, such as are brought in every year from our coastal waters are unusual on the other side of the Atlantic, tends to corroborate the American age estimates, but the desirability of further investigation along this line is self-evident. Judging from the foregoing table the general run of mature shore cod caught in the Gulf of Maine (5 to 20 pounds) are 3 to 8 years old, but whether the very large fish have grown excep- tionally rapidly or are many years old, remains to be learned. The smallest ripe male recorded for American waters weighed about %){ pounds; the smallest ripe female 4 pounds,82 that is, they were in their fourth winter. Probably a considerable propor- tion of our cod mature when they are 5 to 6 years old ; and practically all of them do so by the time they are 9 years old, as Thompson found for the cod of Newfoundland.83 General range. — Both sides of the North Atlan- tic, north to West Greenland, Davis Strait, Reso- lution Island, Hudson Strait in the west,83" south " Contrib. Canadian Biol. (1914-15), 1916, p. 103. •» Earll. Rept. 0. S. Comm. Fish. (1878) 1880, p. 717. " Research Bull. No. 14, Newfoundland Dept. Nat. Resources. 1943, p. 87. >!■ Dunbar (Kennedy, Natural History, Amer. Mus. Nat. Hist., vol. 62, No. 2, 1953, p. 78) has recently reported cod landlocked in southern Baffin Land in a so-called ' ' lake" where the surface is fresh but the deeper water salt. nearly if not quite to Cape Hatteras on the Ameri- can coast; abundant from northern Labrador to Nantucket Shoals, and to New York and New Jersey in winter, when a few are annually caught as far south as the northern part of the North Carolina coast. The continental slope marks the offshore boundary for the cod off the North American coast. The range of the cod in the eastern Atlantic extends from Nova Zembla, Spitzbergen, and Bear Island in the north to the northern part of the Bay of Biscay in the south, and up the Baltic to Finland. The North Pacific cod, with smaller air bladder (G. macrocephalus) cannot be separated from the Altantic cod by external appearance. Occurrence in the Gulf of Maine. — The cod ranks with the herring, mackerel, rosefish, had- dock, pollock, and silver hake as one of the most plentiful of the important food fishes in the Gulf of Maine. Cod were the mainstay of its com- mercial fisheries from earliest colonial times and until the market began to welcome the haddock. We fancy there is no patch of hard bottom, rock, gravel, or sand with broken shells, from Cape Sable in the east to Cape Cod on the west, but supports more or less cod at one time or another. Cod are even caught on soft mud bottoms, though they are not common there. And while the cod are essentially fish of the open sea, they appear regularly in various river mouths in Maine and Massachusetts during the late autumn and winter. One is taken in brackish water occasionally. The eastern half of Georges Bank has always been a most productive cod ground and one of the most famous south of the Grand Banks of New- foundland. The next largest Gulf of Maine fares are brought in from the South Channel- Nantucket Shoals region in the southwestern part of the Gulf, and from Browns Bank in the eastern part, the latter being especially productive in winter. The broken bottom off Seal Island, Nova Scotia, the ground near Lurcher Shoal, and Grand Manan Bank are all famous cod grounds. Other well-known inshore grounds are certain hard patches off Chatham (Cape Cod) ; between Provincetown and Plymouth and off the latter port; Jeffreys Ledge, Ipswich Bay, Cashes Ledge, Platts Bank, and Fippenies. Small vessels like- wise make good catches on the succession of hard and rocky patches that border the coast 190 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE from the Isles of Shoals to the mouth of Casco Bay; on "Seguin" and "Kettle" bottoms off Seguin Island; on the "Matinicus ground" off Matinicus Island; on the "Grumpy" off Isle au Haut; in the neighborhood of Mount Desert Rock and of Mount Desert Island; and on sundry small ridges thence eastward to the mouth of the Bay of Fundy. Rich,84 in fact, lists no less than 175 cod grounds around the inner parts of the Gulf, and many other smaller spots all up and down the coast yield a few cod to the small-boat fishermen. The following summary of the landings of fresh cod from several of the more important Gulf of Maine grounds for 1935 85 illustrates their relative productivity at that time, and there is no reason to suppose that the situation has altered signifi- cantly since then, so far as the numbers of cod are concerned. Locality Georges Bank Browns Bank South Channel.. - Cashes Ledge Stellwagen Bank. Fippenies Bank.. Jeffreys Ledge Nantucket Shoals Platts Bank Pounds 21,598,594 9, 288, 806 2, 993, 580 602, 901 284, 265 48, 865 42, 430 26, 075 20, 060 Percentage of cod in total catch of ground fish 26 30 18 18 37 19 21 14 18 Cod, for some reason not yet explained, become scarcer passing up the Bay of Fundy, and very few are caught near the head, though there are plenty about the mouth of the Bay. Movements of cod in the Gulf of Maine. — The young cod that are hatched within our Gulf tend to follow around the general coastline from north- east to southwest, during the period while they are adrift, as has been shown by Fish 86 very clearly for the Cape Ann — Massachusetts Bay spawning grounds. Our few captures of pelagic cod fry have, in fact, all been in the southwestern part of the Gulf, in which they agree with those of haddock, silver hake, and most of the common flatfishes. As Fish 87 pointed out, the fry from eggs that are spawned north of Cape Ann and on the Massachusetts Bay grounds have ample time to become distributed over the offshore banks " Rept. U. S. Comm. Fish. (1929), 1930, App. 3, table 2, pp. 85-86; table 3, p. 96. » Most recent year for which catches for the smaller inshore grounds are listed separately in the published catch statistics. '• Bull. U. S. Bur. Fish., vol. 43, 1929, pp. 266-290. " Bull. U. S. Bur. Fish., vol. 43, 1929, p. 289. before they seek the bottom (with 14 to 30 days' drift as eggs, and two months or more as pelagic larvae). They might even circle around to the coast of western Nova Scotia and so to the eastern Maine coast. And fry from the Georges Bank spawning grounds would have ample time to do this in years when they are neither held over the Bank by the local circulation nor carried out over the continental slope, to be lost, as happens in the case of the haddock in some years (p. 212). Our Gulf may also receive contributions of cod larvae and fry drifting past Cape Sable, from outer Nova Scotia waters farther east. On the other hand, the cod fry that are taken at Woods Hole in spring may have come from Nantucket Shoals. But those that we found as far south as the Capes of the Chesapeake in April 1930, probably were the product of the spawning that has long been known to take place in winter off New York and off New Jersey. Little is known of the wanderings of the cod in the Gulf of Maine from the time they first seek the bottom when 1% inches long or so, until they are large enough to be caught on hook and line, say 10 or 11 inches long, or l}{ to 2 years old. Young fry, however, from 2 to 4 or 5 inches long and upwards, have been trawled often enough offshore as well as inshore, and they have been found in the stomachs of older cod often enough to show that they soon become distributed all around the Gulf, including the outer part of the Bay of Fundy where it seems that none are hatched (p. 193). But they usually are much more plenti- ful on the rough inshore bottoms than on the smoother offshore banks. A reasonable explana- tion is that if young cod take to the bottom on rough, iocky groimds, or among algae, they have a fair chance of escaping their various enemies, but that thejT find no hiding places on the smooth bottoms that characterize extensive areas on Georges Bank and on Nantucket shoals, hence, are soon decimated. Some of the larger Gulf of Maine cod probably travel very little out of the spawning season, except as they gradually exhaust the food supply in one spot and are therefore driven to move on over the bottom to fresh foraging grounds. Such fish usually are dark and dull colored, with large heads, a sign of scanty diet. Thus tagging experiments, involving many thousands of fish, have shown that a large percentage of the rather FISHES OF THE GULF OF MAINE 191 small cod that make up most of the population along the coast of Maine shift ground but little from season to season. The red fish that haunt the rocks also belong to this category, and red "rock" fish are sometimes caught as large as 10 or 20 pounds. Other cod (and these compose the greater part of the Gulf of Maine stock), are always on the move over the bottoms of their chosen banks. Though cod can hardly be described as schooling in the same sense as herring or mackerel school, these traveling cod often hold together so closely that it is common enough for one-half of a long line to come in loaded with cod, but the other half to come in empty, and these bodies of fish often run very even in size, color, and shape, suggesting that they may hold together for con- siderable periods. But fishermen report them mixed as to sex, sometimes males predominating, sometimes females. It is these "school" fish, as they are called, that most often prey on fish and on squid, though they feed chiefly on shellfish as all cod do. They run slenderer and lighter colored than ground cod and have smaller heads, but it is probable that such differences are only temporary reflections of the surroundings of the individual fish, and that a cod that is a ground fish this month, may start on its travels next, turning brighter and becoming more shapely as it goes, either from a change of diet, from a change of surroundings, or from more active exercise. Furthermore, cod may flee a given locality if harassed too much by the spiny dogfish (p. 48), and no doubt other enemies drive them at times. When cod are on their travels they often rise to the middepths (a fact proved by the levels at which they are caught in nets) ; netted fish are so often empty, whereas those caught on hook and fine are full of food, that they are popularly (and perhaps rightly) believed to fast while they are on a journey. It is probable that the wanderings of these schools of fish are confined to rather small areas, in most instances. Very few cod, for example, that have been tagged on one of the major Gulf of Maine grounds north or east of Cape Cod have been recaught on any other ground. But the experience of fishermen makes it probable that a certain amount of intermingling does take place between Browns Bank and Georges; also between the latter and Nantucket Shoals. An interesting fact in this connection, and one for which we see no explanation, is that the majoritj^ of such cod as stray afield from the coast of Maine tend to travel to the eastward as a rule, as shown by tagging experiments. Thus 50 out of 76 cod that were marked near Mount Desert, and that are known to have journeyed more than a few miles afield went eastward to Petit Man an (5); to Grand Manan (6); to the west coast of Nova Scotia (20); to the outer coast of Nova Scotia as far as Scatari, Cape Breton (16); to Browns Bank (1); to La Have Bank (1); and to Sable Island Bank (1). But only 26 of them were recaptured to the southward and westward; i. e., Penobscot Bay to Cape Ann, including Cashes and Jeffreys Ledges (20); inner part of Massa- chusetts Bay (1); off Provincetown (1); South Channel (1); Nantucket Shoals (1) and Georges Bank (2).88 Canadian tagging experiments have shown a similar state for Nova Scotian cod, most of them remaining nearly stationary for long periods, some straying eastward, very few moving westward." And Thompson's very extensive tagging experi- ments have shown that the movements of most of the cod of Newfoundland waters are confined similarly within regions where physical conditions are comparatively uniform. Some of the cod there make long journeys, discussions of which would carry us too far afield.80 And in two different winters, (1877-1878 and 1892-1893) hooks of a kind that are used by French fishermen on the Grand Banks of New- foundland have been found in cod that were caught near Cape Ann,91 evidence that cod some- times carry out journeys from north and east to south and west along the American coast, com- parable in length to the seasonal migrations that cod have long been known to make along the Norwegian coast, and between Iceland and the West Greenland Banks.92 " About 12,000 cod were tagged by us near Mount Desert, on the U. S. Bureau of Fisheries vessels Halcyon and Albatross II and from other craft, from 1924 to 1931. Recaptures nearby totaled 1,764. " For details as to tagging experiments in Nova Scotian waters, see McKenzie, Contrib. Canadian Biol, and Fisheries, N. Ser., vol. 8, No. 31, 1934. » See Thompson (Research Bull. 14, Newfoundland Dept. Nat. Resources, 1943. pp. 20-45, charts 1-8) for detailed discussion in relation to spawning and to racial subdivisions of the local stock. •i Earll, Rept. U. S. Comm. Fish. (1878) 1880, p. 706. Kendall, Rept. V. S. Comm. Fish (1896), 1898, p. 178. •• See especially HJort, Journal du Conseil. Cons. Perm. Internat. Explor. Mer., vol. I, No. 1, p. 9, 1926; also Schmidt, Rapp. Proc-Verb. Conseil Perm. Intern. Explor. Mer., vol. 72, p. 37, 1931. 192 FISHERY BULLETIN OF THE FISH AND "WILDLIFE SERVICE The only regular seasonal migrations that the cod within our Gulf are known to carry out are: (a) their concentrations on their spawning grounds, followed by their dispersal therce after they are spawned out; and (b) a tendency of the fish living closest in shore and shoalest to shift depth with the season, according to the temperature of the water. Thus the cod tend to work in shore, and shoaler, around Massachusetts Bay in autumn, to work out into deeper (herce cooler) water again for the summer. On the other hand, local fishermen report that the cod abandon the shoalest (7-10-fathom) parts of Nantucket Shoals, after the water there has been chilled by the first heavy snows, to congregate from January until April in the deeper (12-20-fathom) channels (warmer in this case.). Spawning grounds and season. — Thanks to Earll's painstaking studies, and to the large scale on which the Bureau of Fisheries subsequently collected and hatched cod eggs at the Gloucester and Woods Hole hatcheries, the spawning season and the major spawning grounds of the cod are fairly well known for the coastal waters between Nantucket Shoals and the Bay of Fundy. According to the reports of fishermen and to W. F. Clapp's first-hand experience, large bodies of cod spawn on the eastern part of Georges Bank east of Georges shoal, centering at about latitude 41°21' to 41°31', longitude 66°50', to 67° F. in about 35 fathoms of water. Vague rumors are our only indication as to where and when cod spawn on other parts of Georges; they may do so there, wherever the water is shoaler than 35 to 40 fathoms. And there is every reason to suppose that they spawn regularly on Brown's Bank, though we have no definite record of it. The broken bottom of Nantucket Shoals, east and south of Nantucket Island (fig. 94), has long been known as a center of abundance for ripe cod fish in late autumn and early winter. So far as we can learn few cod, if any, spawn on the sandy bottom along the outer shores of Cape Cod. But great numbers of ripe fish congregate in Massachusetts Bay on well-defined grounds 3 to 10 miles offshore, extending from abreast of Sandwich (some 12 miles south of Plymouth) to Minots Light off Cohasset. Years ago many cod also spawned over a small area off Boston Lighthouse and thence northward toward Bakers Island. Few breeding fish have been reported there of late, however, probably because this general locality has been used as the dumping ground for the refuse from Boston, but a few still spawn on various small rocky patches off Gloucester. Figure 94.- — Chief spawning grounds of cod in the western side of the Gulf of Maine. The Ipswich Bay region, where large schools of ripe cod gather in winter and spring, is probably the most important center of production for the inner part of the Gulf of Maine north of Cape Ann, but this ground, like the Massachusetts Bay spawning ground, is limited to a rather small and well defined area extending only from a few miles south of the Isles of Shoals to abreast of the mouth of the Merrimac River and (less produc- tively) to Cape Ann, chiefly within 4 to 6 miles of land. A glance at the chart (fig. 94) will show how limited the more important breeding grounds of the southwestern part of the Gulf of Maine are in extent (not more than 300 square miles in all) FISHES OF THE GULF OF MAINE 193 compared to the whole peripheral zone of this part of the Gulf within the 50-fathom curve. And ripe fish are seldom found even close by, though the fishing for green or spent fish may be good there. One consequence of the limited extent of these spawning grounds is that the cod congregate on them at the spawning season, in great numbers. During the spring of 1879, for example, when fishing was less intensive than it is at present, and when the cod may have been correspondingly more plentiful, more than 11,000,000 pounds of cod, mostly spawning fish, were taken on the Ipswich Bay ground alone by local fishermen. Spawning cod are caught only in small numbers, and at scattered localities in the coastal zone north and east of the Isles of Shoals, the more productive of these minor grounds being near Cape Elizabeth; off Casco Bay; off the Sheepscott River; off Boothbay; and in the neighborhood of Mount Desert Island. Very few ripe cod are reported along the Maine coast farther east. And the egg-collecting activities of the several hatcher- ies have been carried on over so many years that important centers of production there could hardly have been missed. Cod eggs have been taken in the Bay of Fundy but the larvae are unknown there. Neither has any definite evidence been obtained that cod breed in any abundance off the west coast of Nova Scotia. And we should em- phasize that the small ledges in the western part of the Gulf, e. g., Jeffreys and Platts, are not breeding centers though they are important feeding grounds. We cannot speak for Grand Manan Bank or for German Bank. Cod, in short, are quite as local in their choice of spawning grounds in the Gulf of Maine as they are in Nor- wegian waters.93 Cod spawn at least as far south and west as New Jersey,94 and captures, in 1930, of a considerable number of fry 1% to 2% inches long off New Jersey and off Virginia in April are evidence that spawning is successful at least as far south as the offing of Chesapeake Bay. But the fate of these southern-spawned cod is yet to be learned. Following the cod eastward and northward, we learn that eggs are produced in profusion as far north as the Gulf of St. Lawrence and the Grand Banks. But it is not known how much spawning takes place along the eastern coast of Labrador, although eggs have been taken in some numbers along the west coast of Greenland as far north as latitude 66°56' N.95 Cod spawn in shoal er water than haddock on the whole. In fact, we can find no record of ripe cod deeper than 50 fathoms in our waters, and most of the Gulf of Maine spawning takes place on considerably shoaler bottoms. The Georges Bank ground, for example, is about 25 to 35 fathoms deep ; the Nantucket grounds are hardly anywhere deeper than 20 fathoms, and as shoal as 7 fathoms in places; the Massachusetts Bay grounds are about 12 to 25 fathoms; and the Ipswich Bay ground is only 5 to 25 fathoms deep according to the precise locality. It has long been known that while cod spawn chiefly in winter, both in American and in Euro- pean waters, the breeding season lasts much longer and is less definitely limited at either end for cod than it is for the haddock or for the pollock. And experience has shown that the season when the production of eggs is most active differs widely even within the comparatively small area now under discussion. On Nantucket Shoals, ripening fish are caught from late October on, with the cod spawning there in early Novem- ber to mid-February, and occasionally until April. Corresponding to tins, the brood fish taken off Nantucket that were formerly brought in to the Woods Hole pool spawned there from about the first of December until well into February and occasionally as late as March, with the major pro- duction usually from December 20 to January 7.96 And the spawning season is about the same as this off Plymouth in Massachusetts Bay, where ripe cod of both sexes are common from November until as late as April.97 On the north side of Cape Ann, however, only 50 miles distant, ripe fish seldom appear in any numbers until January and not until February in some years, though odd ones may be expected from November on. Earll, for example, found that not one female in ten had commenced to throw her eggs by Feb- ruary, in Ipswich Bay, though spawning wTas then " See Hjort (Rapp. Proc.-Verb., Cons. Perm. Internat. Explor. Mer.,vol. 20. 1914). '< Smith, Rept. U. S. Comm. Fish. (1901) 1902, p. 208; Schroeder, Bull. T. S. Bur. Fish; vol 46, 1930. p. 70. 210941—53 14 "Jensen (Rapp. et Proc. Verb., Conseil Internat. Explor. Mer., vol.39, 1926, p. 85. '• Information from W. II. Thomas, former superintendent of the Woods Hole hatchery. " Information from C. O. Corliss, former superintendent of the Qloucester hatchery. 194 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE at its height in Massachusetts Bay, nor were as many as 50 percent of the Ipswich Bay fish ripe before mid-March. Commencing to spawn later there and near Cape Ann than they do off Ply- mouth, they also continue later, i. e., until the end of April or even into the first part of May, as appears from the following table of cod-egg col- lections supplied by the Gloucester hatchery: Season Collecting field Number of eggs taken Spawning season 1911-12 67, 032, 000 170, 840, 000 91, 980. 000 82. 460. 000 145,630.000 92, 540. 000 119,020,000 249, 510. 000 570, 740, 000 210, 040, 000 No v. 24 to Jan . 3. 1912-13 1913 14 Ofl Rockport (Ipswich Bay). Feb. 16 to Apr. 7. Feb. 1 to Apr. 15. 1914-15 1916-16 1916-17 In Ipswich Bay and off the New Hampshire coast. Feb. 9 to Apr. 13. Feb. 27 to Apr. 13. 1917-18 Feb. 25 to Apr. 27. 1918-19 1919-20 do .. do . Feb. 27 to Apr. 30. Dec. 28 to Apr. 30. 1920-21 ___do Jan. 15 to Apr. 29. Off the western coast of Maine, according to Capt. E. E. Hahn, former superintendent of the Boothbay Harbor hatchery, cod spawn from late February or early March until the last of May, with the production of eggs at its peak in March; they spawn from March through May off the eastern Maine coast, and cod eggs (and hence spawning cod) have been recorded in spring in the Bay of Fundy. On Georges Bank cod spawn in abundance in February,98 March, and April. The records of the hatcheries just summarized tell when eggs are produced in maximum abund- ance, but they throw little light on the limits of the spawning season, for it was oidy during the period when there were enough ripe fish to warrant the effort and expense that spawn taking was carried on. And occasional ripe cod of both sexes are seen long before the bulk of the fish breed, and long after. Thus Earll " reports the first ripe female as taken near Cape Ann on September 2 during the season of 1S7S-79, while we have taken cod eggs, far enough advanced in incubation for positive identification as such, off Shelbui-ne (Nova Scotia) on September 6; near Mount Desert on September 15; and off Penob- scot Bay on October 6 (all in 1915). 93 This fact has long been common knowledge, and W. F. Clapp, formerly of the Museum of Comparative Zoology, has seen many cod with eggs running, caught on Georges Bank in February and March. » Rept. U. S. Comm. Fish. (1878) 1880, p. 713. On the other hand Earll saw ripe fish about Cape Ann as late as June. And our tow-nettings make it likely that some may even spawn in midsummer in the coastal zone east of Cape Elizabeth, for we have occasionally found eggs identifiable as either cod or haddock by their black pigment, and probably the former, near Mount Desert Island on July 19; near Wooden Bell Island at the mouth of Penobscot Bay on August 6 and near Cape Elizabeth on Septem- ber 30. This sporadic summer breeding of cod in our Gulf is hardly comparable to the so-called "after- spawning" that has been observed off the north coast of Iceland by Schmidt,1 in the North Sea, and in the Baltic.2 But it is not unusual for cod to breed in summer off the outer coast of Nova Scotia where ripe fish are reported by local fishermen in June and July. Similarly, spawning cod were caught from the deck of the Grampus (Capt. E. E. Hahn in command) on Bradelle Bank in the Gulf of St. Lawrence late in August many years ago, while gadoid eggs (probably cod) were towed at various localities there during June, July, and August of 1915 by the Canadian Fisheries Expedition.3 Cod spawn chiefly if not altogether in summer on the Grand Banks where Arctic temperatures prevaU during the spring. Corresponding to the prolonged period of repro- duction, spawning takes place over rather a wide range both of temperature and of salinity in our Gulf. On the Ipswich Bay grounds, for example, some are spawning late in November when the bottom water at the depth in question (p. 193) is at its warmest for the year (near 48°) ; they ripen regularly in temperatures of 41°-43° F. (January) ; spawning is at its height in the minimum temper- atures of the year (35°-37.5°), and some spawning continues until the bottom water has once more warmed to 38°-41° (mid-May). On the Massachusetts Bay ground, spawning fish appear in numbers (late November) when the bottom water is still as warm as 47°-48°; the chief production taking place in temperatures of 36°-42° (December through January), hence in warmer water than in Ipswich Bay. And the peak of the ' Rapp. et Proc. Verb., Cons. Perm. Internat. Explor. Mer., vol. 10, 1909, pp. 21, 123. » Ehrenbaum (Nordisches Plankton, vol. 1, 1905-1909, p. 225) and Fulton (Cons. Perm. 1'Explor. Mer, Pub. de Circonstance, No. 8, 1904). ' Dannevig, Canadian Fish. Exped. (19H-15) 1919, p. 22. FISHES OF THE GULF OF MAINE 195 spawning season has passed before the tempera- ture drops to its winter minimum, although some cod spawn there through the coldest season (mini- mum temperature 33°-370). The temperature range through which the cod breed on the offshore grounds cannot be stated so precisely, for want of data for autumn and for early winter. In the Gulf of St. Lawrence, cod are known to spawn in water as cold as 32° F. or even slightly colder,4 though the eggs develop at higher tem- peratures for they rise to the upper water layers. Around Newfoundland, the cod appear to seek temperatures of 35°^0°F. (1.5-4.4° C.) for spawn- ing, with the chief production of eggs taking place at 37°-41°F. (3-5° C).6 Cod spawn in rather colder water on the whole in the Gulf of Maine (still more so in the Gulf of St. Lawrence and on the Newfoundland Banks) than they do in the other side of the North At- lantic, or about Iceland, where the chief production of eggs takes place at temperatures of 40°-45° F. Probably no cod spawn in water fresher than about 32 per mille nor saltier than about 32.8 per mule, either on the Ipswich Bay grounds or on the Massachusetts Bay grounds. And our records (as far as they go) point to a salinity of about 32.6 per mille as typical for the spawning of the cod on Georges Bank. This is water much less saline than ripe cod seek in European seas, and necessarily so, the Gulf of Maine being decidedly fresher at all times of the year than the Norwegian Sea or the waters around Iceland. On the Massachusetts Bay spawning ground the specific gravity of the water is high enough to insure that the eggs shall float throughout the breeding season, but in Ipswich Bay the spring freshets often so lighten the surface that late- spawned cod eggs and haddock eggs may fail to rise to the uppermost water layers, a phenomenon which hinders the operations of the hatchery but which does not militate against the successful incubation of the eggs in nature, since the eggs merely float suspended at some deeper level. This subject . is discussed at greater length in connection with the haddock (p. 208). We have yet to learn what proportion of the cod larvae that are hatched in the Gulf of Maine (doubtless a very small one) survive to grow to ' Hjort, Canadian Fish. Exped. (1914-1915) 1919, p. XXVII. * Thompson, Research Bull. 14, Newfoundland Dept. Nat. Resources, 1943, p. 89. market size. And what few bits of evidence we have in this regard are contradictory.6 Importance.- — In 1945, the most recent year for which detailed statistics of the catch are available for the coastlines of Massachusetts and Maine, as well as for the offshore Banks, the Gulf of Maine yielded about 62,500,000 pounds of cod to United States fishermen;7 some 8,000,000- 9,000,000 to Canadian fishermen; 8 or a grand total of some 70-71 million pounds, plus an inde- terminate amount landed in small Nova Scotian harbors between the Yarmouth County line and Cape Sable. This is about the same amount as the Gulf had yielded in 1919 (about 67,000,000 pounds) ; nor is there anything in the catches of intervening years to suggest that any very pro- nounced fluctuations had taken place meantime in the abundance of cod within our Gulf. A representative yield, in round numbers, broken down into the statistical areas now em- ployed by the U. S. Fish and Wildlife Service, would be about 7,000,000 pounds along the west- ern coast of Nova Scotia and along the lower Nova Scotian shore of the Bay of Fundy; about 380,000 pounds for the upper Nova Scotian shore of the Bay; about 1,600,000 pounds for the New Brunswick shore of the Bay near its mouth; * about 500,000 pounds for eastern Maine; about 4,500,000 pounds for central Maine; about 3,350,000 pounds along western Maine; about 600,000 pounds from the small fishing grounds in the inner-central part of the Gulf; about 5,000,000 pounds off eastern Massachusetts; a little less than 5,000,000 pounds for the grounds from Cape Cod out to the so-called South Channel; about 17,000,000 pounds for Georges Bank as a whole; about 2,000,000 pounds for the western part of Browns Bank; and about 2,200,000 pounds for X a ut ticket Shoals. During the early days of the fishery, the entire Gulf of Maine catch of cod was made on hook and line; on hand lines at first, but with long or • Fish (Bull. U. S. Bur. Fish., vol. 43, 1929, p. 266) caught no cod larvae in Massachusetts Bay, though eggs were abundant there, but the Albatross II towed several hundred little cod (4 to 9H mm.) ofl the tip of Cape Cod near- by, on May 28, 1927. The paucity of our other catches of cod larvae (80 to 90 all told) for other part? of the Gulf of Maine may have been accidental. ' Total landings in New England ports were about 139,700,000 pounds, but something over 77,000,000 of this was taken on the grounds along outer Nova Scotia. ' About 9,259.900 pounds in 194-1, about 8,226,000 pounds in 1945, and about 8,174.800 pounds in 1946. ' No cod are mentioned for the head of the Bay on the New Brunswick side in the Canadian statistics of late years. 196 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE trawl lines coming iDto general use about the middle of the 19th century. And it is not aston- ishing that a fish so nearly omnivorous as the cod should be caught on various baits. Those most in use in the Gulf of Maine are clams (Mya arenaria), cockles (Polynices), herring (fresh, frozen, or salt), and squid. General experience suggests that there is little to choose between the first two of these, while the razor clam (Ensis direcius) is equally attractive though limited by the small supply. And tests made in the Gulf of St. Lawrence 10 proved that fresh herring and fresh squid are about as good as clams, but that frozen and salt herring are less attractive. Other kinds of fish are also used as cod bait in other parts of the world; capelin, especially, in more northern seas, and launce. The earliest important addition to fishing methods came during the winter of 1880-1881, when gill nets, based on the Norwegian system, were introduced in the Ipswich Bay region, yield- ing unexpectedly large catches.11 Since about 1908, when otter trawls came into general use in our waters, an increasing proportion of the catch has been taken by this method. Today about 80 to 85 percent of the Gulf of Maine catch is made in otter trawls ; only about 10 percent on long lines; about 1 percent in gill nets; less than 1 percent in pound nets, and less than 1 percent on hand lines. Cod still bite as greedily, however, as they ever did on clams, cocldes (Polynices), or on pieces » Knight, Contrib. to Canad. Biol. (1906-1910) 1912, pp. 23-32. » For account of cod Ashing methods in North American waters before the introduction of the otter trawl, see Goode and Collins, Fish. Industries U. S., Sect. 5, vol. 1, 1887, pp. 123-198. of squid or herring. We have even caught fair- sized cod on a pickerel spinner tipped with a bit of pork rind, over ledges in shallow water; we have heard of small cod caught on bucktail lures, also on tin-clad lures cast in the surf. And anglers fishing from small craft for pleasure or for home use catch large numbers all along the coast, though these are mostly of the smaller sizes. So far as we can learn, cod have never been jigged successfully in the Gulf of Maine, as they are in abundance in northern Labrador waters. Tomcod Microgadus tomcod (Walbaum) 1792 Frostfish Jordan and Evermann, 1896-1900, p. 2540. Description. — The tomcod resembles a small cod so closely in its fins, in the projection of its upper jaw beyond the lower, in the presence of a barbel on its chin; and in its pale lateral line, that the one might easily be taken for the other. But the outlines of the ventral fins offer a field mark by which the two fish may be separated, for while their second rays are filamentous at the tip in both species, the ventrals of the cod are moder- ately broad, rounded, and with the filament occupying less than one-fourth the total length of the fin, whereas the ventrals of a tomcod are so narrow, so tapering, and with so long a filament (as long as the rest of the fin) that the whole suggests a feeler rather than a conventional fin. Furthermore, the margin of the caudal fin of a tomcod is noticeably rounded, while that of the cod is square or slightly concave; the eye of the Figure 95.— Tomcod (Microgadus tomcod), Woods Hole. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 197 tomcod is decidedly smaller than that of a cod (about one-fifth to one-sixth as long as the head in the tomcod, about one-fourth in the cod, in fish 7 to 10 inches long) ; and the general form of its body is more slender. A less obvious difference is that the first dorsal fin of the tomcod originates over the middle of the pectoral fins or farther back still, farther forward in the cod; and the pectoral fins reach back only a little beyond the middle of the first dorsal fin in the tomcod, but nearly to the rear end of the first dorsal on a cod. Unfortunately, the number of fin rays varies so widely in both these fish that it is not diagnostic, there being from 11 to 15 in the first dorsal, 15 to 19 in the second dorsal, and 16 to 21 in the third dorsal of the tomcod: 12 to 21 in its first anal fin and 16 to 20 in its second anal fin. Most of the recent accounts list the position of the vent as the chief external distinction between tomcod and cod, describing it as in front of the origin of the second dorsal fin in the former and back of it in the latter. But we must caution the reader that it is only for adults of the two species (which no one could confuse in any case, cod being so very much the larger) that this distinction holds; cod as small as tomcod (that is, up to a foot long) often have the vent well in front of the second dorsal, while it may hardly be further forward than that in adult tomcod in breeding condition. Color.- — Tomcod are not so variable in color as cod. Those we have seen (a considerable number) have been olive or muddy green above, with a yellowish tinge, darkest on the back, paling on the sides, and mottled with indefinite dark spots or blotches. The lower parts of the sides usually show a decided yellowish cast in large fish; the belly is grayish or yellowish white; the dorsal and caudal fins are of the same color as the back; the anals are pale at the base but olive at the margin ; and all of the fins are more or less dark mottled. The tomcod has often been described (following Storer) as thickly speckled with black dots, but we have never seen one marked in that way. Size. — The maximum size is about 15 inches and VA pounds, but few of them are more than 9 to 12 inches long. Habits. — The tomcod is strictly an inshore fish; probably few ever descend more than two or three fathoms, or stray as much as a mile outside the outer headlands. In our Gulf they chiefly fre- quent the mouths of streams and the estuaries into which these empty, as well as shoal, muddy harbors like Duxbury Bay. As often as not they are in brackish water, and they run up into fresh water in winter. Dr. Huntsman, for example, writes us that they are caught in the Petit Codiac River 12 miles above the head of tide. Tomcod are less plentiful in harbors where there is no stream drain- age, but now and then they are caught off open shores, off Nahant, for instance, and such fish are usually large ones. South of Cape Cod, most of them move out from the shore into slightly deeper (hence cooler) water in spring, coming in again in autumn to winter in the estuaries. But a year comes from time to time (such as 1925) when they are plentiful close inshore all summer, as far south even as New York.12 And they do not carry out any inshore-offshore migrations of a regular sort in the cooler Gulf of Maine, so far as is known. Indeed, they are so resistant to cold that we find no record of them killed by winter chilling, a fate that sometimes overtakes other fishes that live in shoal water. And they are equally hardy toward sudden changes of salinity. Tomcod feed chiefly on small crustaceans, es- pecially on shrimps and amphipods, a great variety of which have been found in their stomachs; also on worms; small mollusks; squids; and fish fry, such as alewives, anchovies, cunners, mummichogs, herring, menhaden, launce, sculpins, silversides, smelt, and sticklebacks. According to Herrick 13 tomcod arc not so keen- sighted as pollock nor so active as hake, but spend most of their time quietly on the bottom in the aquarium. His experiments also proved that they are able to recognize concealed baits by the sense of smell if they chance to swim near and that they search the bottom by dragging the chin barbel and the sensitive tips of the ventral fins as they swim to and fro, either for food, or to stir up shrimps and other food items. Tomcod spawn in the shoal waters of estuaries, in stream mouths and such places, either in salt water or in brackish, and their eggs have been hatched artificially in fresh water. The season lasts from November to February, inclusive, with the height of production in January. The eggs are about 1.5 mm. in diameter with a conspicuous oil ii Nichols and Breder (Zoologies, N. Y. Zool. Soc, vol. 9, 192", p. 166) state that tomcod up to 1094 inches long were common throughout that summer In Sundy Hook Bay. " Bull. U. S. Fish Comm., vol. 22, 1904, p. 262. 198 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE globule, and (unlike those of its larger relative) they sink to the bottom where they stick together in masses, or to seaweeds, stones, or any available support. Incubation occupies about 24 days at an average temperature of 43°; 30 days at 40°. The larvae are not only somewhat larger (5 mm.) at hatching than those of the cod, but are farther ad- vanced in development, the mouth being formed. And they differ from all other Gulf of Maine ga- doids at a corresponding stage by the presence of the oil globule and by the fact that the vent opens at the margin of the ventral fin fold and not at its base at one side.14 Although great numbers of tomcod have been hatched artificially by the State of New York, its later larval stages have not been described, nor have we seen them ourselves. The fry, which are said to remain through their first summer in the waters where they are hatched, grow to a length of 2%-3 inches by the following autumn. But nothing is known of the rate of growth of older fish. General range. — North American coastal waters from the Gulf of St. Lawrence and northern New- foundland to Virginia, running up into fresh water. Occurrence in the Gulj oj Maine. — The tomcod is locally common around the entire coastline of the Gulf. It is reported at Pubnico and inSt. Mary Bay, for example, on the west coast of Nova Scotia; at various localities on both shores of the Bay of Fundy (e. g., Annapolis Basin and River, Minas Basin, St. John Harbor, and the St. Andrews region) ; at Eastport; from almost every river mouth along the Maine coast; in the vicinity of Boothbay Harbor; at sundry stations in Casco Bay; and in Portland Harbor in Maine. And it is to be found in practically every estuary around the Massa- chusetts Bay region. Tomcod are caught from docks and bridges and in salt creeks in mid-summer as well as in winter. Tomcod are in the inner parts of Dux- bury bay, for example, in midsummer; there are also plenty of them in a certain salt marsh creek at Cohasset at all seasons; and this applies to many similar locations all up and down the coast, including the Bay of Fundy, where tomcod are in and near the estuaries the year round, as Huntsman !6 remarks. Westward and southward from Cape Cod, the tomcod is plentiful in suitable situations all along the coast to New Jersey, where Abbott 18 described them many years ago as a "very common" little fish, and we have often caught them while fishing from docks in lower New York Harbor. In the opposite direction, they are common along the outer shores of Nova Scotia. They are plentiful enough around the shores of the Gulf of St. Lawrence for catches of 684,000 pounds to be reported from the New Brunswick coastline of the Gulf in 1947, 20,400 pounds from the southern shore of the estuary of the St. Lawrence River, 152,900 pounds from the north shore of the estuary and Gulf, while Jeffers " reports them as taken in considerable numbers through the ice in winter, on the Newfoundland side of the Strait of Belle Isle. And they are to be expected along the southern and eastern coasts of Newfoundland, though they seem not to have been reported there as yet. Importance. — The tomcod is a delicious little fish. But it seems to have been more higldy con- sidered a century ago, when between 5,000 and 10,000 pounds were caught annually in the Charles River tributary to Boston Harbor; today, it is unusual to see any for sale in a Massachusetts fish market. And, in any case, tomcod are not plentiful enough anywhere around our Gulf to support a regular commercial fishery of any mag- nitude. In 1929 the reported catch was about 6,000 pounds for Massachusetts, about 16,500 pounds for Maine, and about 6,100 pounds for the Canadian shores of the Gulf. In 1942,18 27,500 pounds were reported for Maine, none for Massa- chusetts, about 10,000 pounds for the Nova Scotian shore of the Bay of Fundy. Since that time a few thousand pounds have been reported yearly from the Nova Scotia shores of the open Gulf and of the Bay of Fundy; " none at all, however, from its New Brunswick shore. Most of the tomcod marketed in Maine (also most of those formerly marketed in New Bruns- wick) are taken in bag nets or in pocket nets set » Ryder (Rept. U. S. Comm. Fish., (1885) 1887, p. 523, pi. 13, flg. 67) de- scribes and pictures the newly hatched larva of the tomcod. i» Contrib. Canadian Biol., (1921) 1922, p. 67. « Oeol. New Jersey, 1868, p. 818. O Contrib. Canadian Biol., N. Ser., vol. 7, No. 16 (Ser. A, general, No. 13), 1932, p. 7. 18 Most recent year when tomcod were mentioned in the United States catch statistics for the Gulf of Maine coast. ■• 35,000 pounds of tomcod were reported for Digby County in 1944, bul this amount is so much larger than for preceding years, or for 1946, as to suggest some error. FISHES OF THE GULF OF MAINE 199 in the courses of the larger rivers, a few in weirs. In the days when the commercial catch for Massa- chusetts was large enough to be worth reporting, most of it was taken on hook and line north of Plymouth, in weirs and traps south of Plymouth. Besides the fish reported in catch statistics, a considerable number are caught in autumn on hook and line by smelt fishermen and by anglers fishing especially for "frost fish," all along the shores of northern New England and used for home consumption. Hence they are not re- ported or included in the fishery statistics. Tomcod bite any bait greedily. Clams, shrimp, sea worms, or cut fish will serve, and they afford amusement to a larger number of anglers in harbors and stream mouths than the meager commercial catch might suggest. Haddock Melanogrammus aeglefinus (Linnaeus) 1758 Jordan and Evermann, 1896-1900, p. 2542. Description. — The most obvious ways in which the haddock differs from the cod are in its black Figure 96. — Haddock (Melanogrammus aeglefinus). A, adult, Eastport, Maine, from Goode, drawing by H. L. Todd; B, egg (European); C, larva (European) just hatched; D, larva (European), 4.2 mm.; E, larva (European), 15 mm.; F, young fry (European), 25 mm. B and C, after Heincke and Ehrenbaum; D, after Ehrenbaum; E and F, after Schmidt. 200 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE lateral line (that of cod and of pollock is paler than the general ground tint) and in the presence of a dusky blotch on each side over the middle of the pectoral fin, and close below the lateral line. Furthermore the first dorsal fin of a haddock (higher than that of a cod, relatively) is con- siderably higher than either the second or third dorsal, more acutely triangular in outline, and with slightly concave margin. The margin of the haddock's tail is more concave than that of the cod ; and its second and third dorsal fins are more angular than is usuaUy the case with the cod, though they are similarly rhomboidal in outline. The haddock's mouth is relatively the smaller, not gaping back to below the eye, and the lower profile of its face is straight, with the upper profile only slightly rounded, giving the nose a charac- teristic wedge-shaped outline in side view. The upper jaw projects further beyond the lower in the haddock than in the cod, and the snout is usually more pointed and the body more flattened sidewise. But the general arrangement of the fins is the same; there are about the same number of dorsal fin rays in haddock as in cod (14 to 17, 20 to 24, and 19 to 22, in the first, second, and third fins, respectively) ; and while the anal fins average one or two more rays each (21 to 25 and 20 to 24), individual cod may have more anal rays than individual haddock. Finally, the haddock is a slimmer fish than the cod and al- though its scales (which clothe it from nose to tail) are of about the same size relatively (about 160 rows along the side), they are scarcely visible through the mucus with which the skin is coated.20 Color. — When a live haddock is first taken from the water, the top of its head, back, and sides down to the lateral line are dark purplish gray, paling below the lateral line to a beautiful silvery gray with pinkish reflections, and with the black lateral line and the sooty shoulder patch (just mentioned) standing out vividly. This patch, the "devil's mark," is indefinitely outlined and varies in size and in distinctness, but oidy very rarely does a haddock fail to show it. The belly and lower sides of the head are white. The dorsal, pectoral, and caudal fins are dark gray; the anal fins pale like the lower part of the sides and black specked at the base; and the ventrals are white, more or less dotted with black. Haddock usually » Vladykov (Canadian Field Natural., vol. 49, No. 4, 1935, p. 64) describes a haddock with 3 eyes, and includes a photograph of it. run very uniform in color, but occasionally one shows from one to four dark transverse bars or splotches in addition to the black shoulder blotch. Several of these serially striped haddock have been taken in Passamaquoddy Bay 2I and we have seen such near Mount Desert. Occasionally a haddock may be decidedly golden on the back and sides, with the lateral line golden, and such fish may lack the dark blotches. Size. — The haddock is a smaller fish than the cod, the largest on record having been only 44 inches long, weighing about 37 pounds.22 One of 30 pounds, caught on La Have Bank in the autumn of 1949 23 is said to have been the heaviest ever landed at the Boston Fish Pier. The largest among 1,300 fish that were measured and weighed by Welsh near Gloucester during the spring of 1913 was 35K inches long, weighing about 16K pounds. Only 4 or 5 out of the more than ten thousand haddock that we have helped to tag were as long as 32 to 34 inches. And the great majority of the fish that are brought in measure from 14 to 23 inches long, and weigh from 1% to 4% pounds. The largest among 627,996 fish meas- ured during the period 1931-1948 was 34% inches long.24 The relationship between length and weight averages as follows, according to Shuck ; 2S 10 inches, 7 ounces; 12 inches, 12 ounces; 14 inches, 1 pound 2 ounces; 16 inches, 1 pound 11 ounces; 18 inches, 2 pounds, 6 ounces; 20 inches, 3 pounds 3 ounces; 22 inches, 4 pounds 3 ounces; 24 baches, 5 pounds 5 ounces; 26 inches, 6 pounds 9 ounces; 28 inches, 8 pounds 3 ounces; 30 inches, 9 pounds 15 ounces. Habits. — Haddock live deeper than cod on the whole; few are caught in less than 5 to 10 fathoms of water and most of them in 25 to 75 fathoms. In fact, they so seldom come into shoal water where young cod are so plentiful that the pound nets of Massachusetts reported only about 5,000 pounds of haddock in 1919, as compared with almost 300,000 pounds of cod. Neither do we remember hearing of a haddock of any size in any of the shoal harbors where little pollock so abound. And the difference in habitat between these closely related species holds from the time the young fry =i Prince, Contrib. Canadian Biol., (1915-1916) 1917, p. 86. " This giant was an Icelandic fish, reported by Thompson (Rapp. et Proc. Verbaux, Conseil Internat. Perm. Explor. Mer, vol. 57, 1929, p. 29). » Received by O'Hara Bros., and reported by Moore, Boston Herald, Nov. 29, 1949. »* Information from Howard W. Schuck. » Fishery leaflet No. 198, U. S. Fish and Wildlife Service, 1947. FISHES OF THE GULF OF MAINE 201 first seek bottom, for haddock usually do so in 20 to 50 fathoms or deeper, seldom close to the shore, and perhaps never in the littoral zone.26 On the other hand, comparatively few haddock, are caught deeper than 100 fathoms in American waters,27 though they have been taken as deep as 120 fathoms (220 m.) on the slopes of the Faroe Bank, and as deep as 164 fathoms (300 m.) off Iceland.28 WII6HT lbs lb 14 13 IZ II 10 / / / / / / t / / / / / f / / f s / S 8 7 . s s s • Figure 97. — Average weight of ripe haddock of different lengths; male ( — ) and female (..) at Gloucester, Mass., March to May 1913. The haddock, like the cod, is a cold-water fish, though it is not at home in temperatures quite as low. Thus it is almost wholly absent off New- foundland, in the Gulf of St. Lawrence, and off Nova Scotia when the bottom water is as cold as 32° F.; few are caught there, generally speaking, where the bottom water is colder than about 35-36° F. (2° C.) though good catches are some- times made in temperatures as low as 34°. At the opposite extreme, haddock appear to avoid water warmer than about 50-52° F. Thus Vlady- kov 29 reports that young haddock withdraw from Halifax Harbor if the temperature near the bot- tom rises above about 52°, though they can sur- :fl The fact that haddock fry less than 1 year old have never been reported in shoal water in the Gulf or at Woods Hole corroborates European fishing experiments summarized by Damas (Rapp. et Proc.-Verb., Cons. Intcmat. Explor. Mer, vol. 10, 1909) and by Schmidt (ibid.). » Thompson, Research Bull. No. 6, Newfoundland Dept. Nat. Res., 1939, p. 9. » Ooode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, p. 354) list a haddock from 499 fathoms but with suspicion as to the accuracy of its label. » Contrib. Canadian Biol., N. Ser., vol. 8, No. 29, 1934, p. 418. vive considerably higher temperatures for limited periods.30 It is evident from this that the entire Gulf of Maine, at the depths frequented by the haddock, is suitable for them so far as tempera- ture is concerned, but that the upermost stratum may be too warm from late summer through early autumn, and too cold from late winter through early spring. In exceptional years, too, such as 1926, the whole column of water may chill to a temperature too low for their comfort in the Bay of Fundy (p. 210). The salinities at the localities and depths where haddock live in our Gulf range from about 31.5 per mille inshore to a maximum of about 34.5 per mille on the offshore edge of Georges Bank, with most of the catch made in water more saline than about 32 per mille. And while they enter the bays and reaches between the islands along the coast of Maine in some numbers (p. 210), they never run up estuaries into brackish water. Thus, haddock seem to require somewhat higher salinities than cod, which are sometimes caught in considerable numbers where the water is below 31 per mille (as in the Bras d'Or Lakes, Nova Scotia).31 Id general, the haddock live in rather cooler and less saline waters in the American side of the Atlantic than in the European, as Thompson 32 has emphasized. The haddock is more exclusively a ground fish than the cod and though they sometimes pursue herring and other small fish, as cod do more often, we have never heard ol haddock com- ing to the surface when so engaged, events by no means unusual with cod, and a characteristic phase in the life of the American pollock (p. 214). Haddock are more selective than cod in the type of bottom they frequent, being rarely caught over ledges, rocks, or kelp (where cod are so plentiful), or on the soft oozy mud to which hake resort. They are chiefiV taken on broken ground, gravel, pebbles, clay, smooth hard sand, sticky sand of gritty consistency, and where there are broken shells; they are especially partial to the smooth areas between rocky patches. Food. — During their first few months, while living pelagic near the surface, haddock fry probably depend on copepods as cod do. After » At the St. Andrews Laboratory, haddock kept at a temperature varying between about 57° and about 0S° F. survived for 3 to 4 months. •' Needier, Contrib. Canadian Biol., N . Ser., vol. 4, No. 20, 1929, p. 10. •» Research Bull., No. 6, Newfoundland Dept. Nat. Resources, 1939, p. 12. 202 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE they take to the bottom they become bottom feeders like cod, devouring all kinds of inverte- brates so indiscriminately that, as Baird 33 re- marked long ago, "a complete list of the animals devoured by the haddock would doubtless include nearly all the species belonging to the fauna" of the particular ground on which the fish in question were living. And they begin to depend on this adult diet when they are small. Thus we have found 7- to 9-inch fish full ot brittle stars, bivalve mollusks, small worms, and amphipods. The larger Crustacea, such as hermit, spider, and common crabs, shrimps, and amphipods, with gas- tropods and bivalve mollusks in great variety, worms, starfish, sea urchins, sand dollars, brittle stars, and sea cucumbers all enter regularly into the dietary of the haddock, according to locality. W. F. Clapp, for instance, listed no less than 68 species of mollusks, both bivalves and gastro- pods, from 1,500 haddock that were caught on the northwest part of Georges Bank in 40 to 60 fathoms, and he has called our attention to the fact that haddock usually contain smaller shells than do cod, and never the very large sea clams (Mactra) which are so important a constituent of the diet of the latter. Neither do haddock eat crabs larger than about 2 inches across, as cod so greedily do. On the other hand, haddock depend more on worms than cod do, and they are often packed full of worm tubes when they are caught on bottoms covered with the latter (the "spaghetti bottom") as in the locality known as "Cove Clark" on the northwest face of Georges Bank (about lat. 41° 08', long. 68° 40'). Haddock caught near Eastport, Maine, contained 8 species of annelid worms, and they must root out much of their food from the mud and sand of the sea bottom; in no other way could they obtain the burrowing worms and mollusks that their stomachs contain so often. Haddock take squid when opportunity offers; they are said to prey on herring in Norwegian waters; on launce around Iceland; on fish, mostly launce, on the Nova Scotian banks;34 on young eels off Cape Breton, Nova Scotia;35 on herring near Woods Hole and, in 1931, we received reports of haddock having eaten small mackerel on Georges Bank in January. And many baby had- " Rept. U. S. Comm. Fish (1886) 1889, p. 37. 81 See Homans and Needier (Proc. Nova Scotian Inst. Sci., vol. 21, 1946, pp. 15-49) for a study of the haddock. » Needier, Copeia, No. 171, 1929, p. 41. dock about 8 inches (20 cm.) long, trawled on the southwest part of Georges Bank, August 13, 1945, were not only seen by John R. Clark of the U. S. Fish and Wildlife Service, to disgorge large num- bers of small fish (apparently young silver hake) on the deck of the vessel, but had been feeding chiefly on them. They have also been accused of feeding greedily on herring spawn, perhaps without much justice. But fish ordinarily form so small a part of the diet of the haddock of our Gulf that none of those examined by Welsh near Cape Ann in 1913, nor the Georges Bank haddock opened by Clapp (about 5,000 altogether), and only two of the many that we have ourselves opened, con- tained fish of any kind, nor have any of the fisher- men of whom we have inquired (and their practical experience is of course vastly wider than ours) described Gulf of Maine haddock as feeding to any great extent on fish. And none of the East- port haddock that were opened by Doctor Kendall had risen to take the large pelagic shrimps (euphausiids) that are so abundant there and which are the chief food of the local pollock. Welsh's experience with the haddock near Cape Ann during April 1913 was that they are apt to fast at spawning time; more than 95 percent of the hundreds of fish caught there in the gill nets were totally empty, while long lines set nearby were bringing in very few haddock though they were taking hake in fair numbers. But spawning haddock elsewhere "both male and female, have been found with well filled stomachs, and many spawners have been observed in the catches of line fishermen,"36 so the rule is not universal. It also seems that they feed less actively, or at least they take the hook less freely, at temperatures lower than about 36°, as it is in the coldest parts of the Gulf in winter, and the best hook and line catches are made at about 45°-50° F. The haddock, like the cod, is a prolific fish for its size. Earll 37 estimated the number of eggs in a female weighing 2% pounds and 19}{ inches long at 169,050; 634,380 in one of 4% pounds and 24 inches long; 1,839,581 in one 9 pounds 9 ounces and 28 K inches long. Incubation occupies 15 days at a temperature of 37°; 13 days at 41°, a fair average for the eggs that are spawned in the Gulf of Maine. The eggs are buoyant, without oil » Needier, Contrib. Canadian Biol, and Fish., N. Ser., vol. 0, 1930, No. 10 p. 7. « Rept. U. S. Comm. Fish. (1878) 1880, p. 733. FISHES OF THE GULF OF MAINE 203 globule, and from 1.19 to 1.72 mm. in diameter; eggs taken at Gloucester in March 1913 averaged 1.57 mm., varying from 1.47 to 1.72 mm. Thus they average slightly larger than those of the cod. The haddock egg cannot be distinguished from that of the cod in early stages in its development, hence the term "cod-haddock," and when they are newly spawned there is even danger of confusing them with the eggs of one of our commonest flounders, the "witch" (p. 287), whose breeding season immediately follows that of the haddock. But the formation of black pigment soon identifies the cod-haddock egg as such (the embryonic pigment of the "witch" is yellow). The newly hatched larva is about 4 mm. long, with the vent close behind the yolk sac and at the base of the ventral fin fold, not at the margin, so that it seems to end blind. It resembles a cod so closely that the two would be indistinguishable one from the other, were it not that the post-anal pigment granules of the haddock are arranged in a row along the ventral surface of the trunk from vent to tip of tail, and not in bands as they are in the cod (p. 188) and in the pollock (p. 216), while the dorsal wall of the body cavity of the haddock is densely pigmented. In water of 41° F. the yolk sac is absorbed in about 10 days when the little fish is about 5.5 mm. long; the dorsal and anal fins are fully formed at 16 to 20 mm.; and the young haddock begin to take on the general aspect of the adult by the time it is 30 to 40 mm. long. The arrangement of the larval pigment serves to differentiate the little haddock until it is about 12 mm. long. Larger fry are distinguish- able from both cod and pollock by their pale pigmentation, and by the greater height of their first dorsal fin. Gulf of Maine haddock average about 6 inches long (extremes, 5 to 7 inches) at the end of their first year, and investigations show that the rela- tionship between length and age averages about as follows for larger haddock in different seas: Age, years Gulf of Maine North Sea Norway Length, inches Length, inches Length, inches 2_ 12 17.5 19 21 22.5 24 25 10 12 15 17.5 20 22 24.5 10.6 3.. 13 4. 15.5 8 17.6 6 19.5 7 21.5 8 23 Thus, American haddock grow more rapidly on the whole than European haddock while they are young, but more slowly when older, so that had- dock on both sides of the Atlantic appear to be of about the same size by the time they reach 7 or 8 years of age. Needier 38 has found too, that had- dock also differ considerably in their rate of growth in different parts of the Gulf of Maine, St. Andrews fish growing faster than those of Browns Bank, with Nantucket Shoals fish intermediate in this respect, as is illustrated in the following table: Age, years 8K Average length, inches Nantucket Browns St. Andrews Shoals Bank is'. UM 16?i 20H 20 X 18K Zlj-i 22 1»H 24 23H 20»i 25« 25 21 H 26Ji 25H 22H Eastern Nova Scotia 16Ji lWi 21 22.4 24 25 X According to Thompson 38 haddock on the Grand Banks grow more slowly than the Nova Scotian fish, averaging about 23 to 26 inches when 8 to 10 years old, while in the vicinity of Halifax Vladykov 40 gave about 12/4 inches as the length of 2 + -year-old haddock and 13M inches for 3+- year-old, a rate of growth slower than for other parts of the western Atlantic and perhaps not typical for all years. But individual fish grow at such different rates (probably due to food supply) that a haddock of a given length may differ by 1 or 2 years in age, or even by 3 years in the case of the larger fish. Thus a Gulf of Maine haddock, 14 inches long, may be 2 to 2)i years old; one of 20 inches, 3 to 4 years; one of 28 inches, 8, 9, or 10 years old. An illustration of this variability is that 6 out of 10 fish that were tagged by the vessels of the U. S. Bureau of Fisheries and were recaptured later had gained ){- to K-inch in 2 months though another had not grown at all in that period; one grew 2 inches in 9 months, but two others grew only K- to %-inch in 1 1 months.41 And Vladykov's » Contrib. Canadian Biol, and Fish., N. Ser., vol. 4, No. 20, 1929, pp. 11-20, 275-284; N. Ser., vol. 6, No. 10, 1930, p. 64 [295], fig. 17, p. 55 [296]. >• Research Bull. No. 6, Newfoundland Dept. Nat. Resources, 1939, p. 15, fig. 3 and table 3. <° Vladykov (Contrib. Canad. Biol., vol. 8 (29), 1934, p. 7) gave his lengths to the last vertebra, but we have converted these into total lengths to middle of caudal fin. >' Schroeder, Jour. Marine Res., vol. 5, No. 19, 1942, p. 16. 204 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE studies of the age-length relationship among young haddock of different sizes near Halifax, Nova Scotia, have shown, similarly, that their average rate of growth may differ considerably within short distances in Nova Scotia waters.42 The oldest haddock noted by Needier, one about 28% inches (72 cm.) long, taken off Ingonish, Nova Scotia, was in its 14th year. But the largest, about 30% inches (78 cm.) long, taken off Campo- bello Island at the mouth of the Bay of Fundy, was in its tenth year, only. In general, Gulf of Maine haddock grow most rapidly in late summer and early autumn, when the temperature of the water is highest at the depths in which they live, but there is much varia- tion in this respect from place to place and from year to year, as various authors have noted. Shuck 43 describes the haddock of New England waters as maturing sexually at 3 or 4 years, when they weigh 2 or 3 pounds. And the smallest sexually active specimens found by Welsh among 1,300 haddock were 2 females of about 20 inches long each; i. e., about 4 years old. Most of the Nova Scotia haddock also spawn first in their fourth or fifth year, according to Needier, as some do in Icelandic waters, also. This supports Duff's M view that the slackening of the rate of growth at 4 or 5 years of age, which she observed, reflects the first ripening of the sexual organs. In the eastern Atlantic, mature haddock have been reported as small as 9 inches. And almost all the fish spawn there by the end of their third year. General range. — Both sides of the North Atlantic. On the American coast haddock are the most abundant from the southern part of the Grand Bank and from the more easterly of the Nova Scotian Banks to Cape Cod. In winter they are taken southward to New York and New Jersey, and they have been recorded in deep water as far southward as the latitude of Cape Hatteras. But the species as a whole is so much more closely confined to waters east of Marthas Vineyard than is the cod, that in 1947, for example, only 158,992 pounds of haddock were caught off New York and New Jersey, contrasting with 2,962,559 pounds of cod for that part of the coast.45 Neither « Contrib. Canadian Biol. Fish., N. Ser., vol. 8, No. 29, 1934, p. 415, fig. 2. *3 Unpublished manuscript. " Contr. Canadian Biol. (1914-1915) 1916, p. 39. 48 This is exclusive of 4,110,508 pounds of haddock and 739,759 pounds of cod landed at New York City, most if not all of which were caught in waters to the east of Marthas Vineyard. does the range of the haddock extend as far north as that of the cod. Small catches are made in the southern side of the Gulf of St. Lawrence; also along its north shore both in the St. Lawrence estuary and nearing the Strait of Belle Isle, and a scattering are taken among the cod along the west coast of Newfoundland.46 And while the experimental trawling campaigns of the Newfound- land Fishery Research Laboratory have shown that there is a distinct and extensive stock of haddock on the southern part of the Grand Banks region47 very few are caught farther north along Newfoundland, though some fish have been re- ported from the Strait of Belle Isle, likewise from West Greenland.48 And haddock are unknown in the icy waters along the outer coast of Labrador, where great quantities of cod are caught every summer. Occurrence in the Gulf of Maine.- — Haddock are very plentiful all around the open Gulf, as well as on all the offshore banks, especially on Georges where they greatly out-number the cod. This is, in fact, one of the two species that now rank at the top among Gulf of Maine fishes, from the com- mercial standpoint; the rosefish is the other (p. 430). Good haddock grounds, it is true, are less extensive close inshore and more scattered there than good cod grounds, haddock being confined for the most part to depths greater than 5 to lOfathoms (p. 200), and being more selective in types of bottoms they frequent (p. 201). But the number of individual haddock that inhabit the coastal belt of the Gulf within 15 to 20 miles of the land may be as great as the number of individual cod, for while the yield of the inshore small boat fisheries has run only one-third to one-half as great in pounds for haddock as for cod, in Maine and Massachusetts, in years for which data are readily available,49 and one-half to three-fourths as great for haddock as for cod in the Bay of Fundy,50 this discrepancy may <> For locations, see Needier, Contrib. Canadian Biol., N. Ser., vol. 6, No. 10, 1930, p. 5 [245], fig. 1. " Thompson, Research Bull. No. 6, Dept. Nat. Resources Newfoundland, 1939, p. 7. <• Jensen and Hansen (UndersjSgelser over den Gr0nlandske Torsk, p. 52, 1930). *• Between 14 and 15 million pounds of cod and about 5 million pounds of haddock in 1919; between 6 and 7 million pounds of cod and about 3 million pounds of haddock in 1924, these being the only two recent years when the yield of the small boat inshore fishery was listed separately in the published statistics of the catch. 10 Bay of Fundy catch, about 7 million pounds of cod and about 5 million pounds of haddock in 1919; about 6 million pounds of cod and about 4 million pounds of haddock in 1946, years that seem to have been fairly representative. The inshore catches for western Nova Scotia are not separated from the off- shore catches in the published statistics. FISHES OF THE GULF OF MAINE 205 not be greater than can be accounted for by the considerably greater weights of individual cod than of individual haddock. And haddock certainly are far more numerous than cod on Georges Bank as a whole, especially on its western half. Haddock, for example, large and small, made up 60 to 70 percent by number of all the fish caught on various parts of the bank, spring to autumn, by certain otter trawlers in 1913, cod less than 10 percent; similarly, in 1948, 1949, and 1950 haddock formed about 21 percent by number, cod less than 1 percent of the fish trawled there by the Albatross III." In 1945 (most recent year for which detailed statistics are avadable both for the New England fishery and for the Canadian), the landings were as follows, for different parts of the Gulf, to the nearest 100,000 pounds: western part of Browns Bank, 6,000,000; grounds along the Nova Scotian shore of the open Gulf, 1,000,000; Nova Scotian side of the Bay of Fundy, 3,400,000; New Bruns- wick side of the Bay of Fundy near the mouth, 1,100,000; 52 off eastern Maine, 200,000; off central Maine, 2,100,000; off western Maine, 900,000; off eastern Massachusetts, 5,400,000; small grounds in the inner central part of the Gulf, 400,000 to 500,000; northern part of the Gulf, not classified, 1,700,000; Cape Cod out to the so-called South Channel, 3,900,000; Nan- tucket Shoals, 2,200,000; Georges Bank as a whole, 53,200,000. If this proportional relation- ship is roughly representative, as seems likely on various grounds, the Georges Bank-South Channel area as a whole harbors perhaps two-thirds to three-fourths of the total haddock population of our Gulf, with an average yearly yield of about 94,000,000 pounds, for the period 1931-1948, equivalent to something like 37 million fish.63 This indeed, is perhaps the greatest haddock ground for its size in the world, or has been in the past.64 According to the combined landings for the years 1942-1947, the northwestern "-northern parts of the Bank, and its central-southeastern 11 Information contributed by Clyde C. Taylor of the U. S. Fish and Wildlife Service. ,J Few haddock are landed near the head of the bay on the Nova Scotian side; none there on the New Brunswick side. " Estimate by Howard W. Schuck, from Fish. Bull. 66, 1951. '< Hen-ington (Fishery Circular No. 23, U. S. Bur. Fish., 1936) so classed it. '• During recent years this part of the Bank has been classified as "eastern side South Channel" in the catch statistics published by the U. S. Fish and Wildlife Service. part, are two to three times as productive each, as is the southwestern part, which agrees with fishermen's reports in general.66 Browns Bank, much smaller in area than Georges, is perhaps equally densely populated. The following table shows the percentages of the total catch of haddock taken on Georges Bank in each of the major statistical areas, in different years: Year Northwest- ern part Northern edge Central and southeast- ern part Southwest- ern part 1942. 19 17 20 31 26 19 39 27 37 24 35 40 36 45 35 24 29 33 6 1943 11 1944... 9 1945 22 1946 11 1917 9 22 34 34 11 Proceeding next to a more detailed survey of the inshore grounds we find that considerable numbers of haddock are caught on German Bank, and on the broken grounds off Lurcher Shoal. And while haddock are less plentiful than other ground fish on Grand Manan Bank at the mouth of the Bay of Fundy, perhaps because of the type of bottom, yearly landings of something like 3 million pounds along Digby Neck, Nova Scotia,67 reflect a rich center of population at the mouth of the Bay of Fundy on the Nova Scotia side.68 Haddock, like cod, diminish in numbers inward into the Bay, so much so that the counties at its head (Hants, Colchester, Cumberland, Westmoreland) report a few hundred pounds, at most, in some years, none at all in others. But they are plentiful enough on the New Brunswick side of the Bay near its mouth and within Passamaquoddy Bay to yield yearly catches about one-third as great as on the Nova Scotia side. The most productive of the small grounds in the western side of the Gulf 69 are Cashes Ledge, " Needler's chart of haddock catches, 1917-1925 (Contrib. Canadian Biol., N. Ser., vol. 6, No. 10, 1930, p. 5 [245], fig. 1) would suggest that haddock were concentrated on the western edge of the Bank chiefly and on the neigh- boring parts of Nantucket Shoals. But it is probable, as he points out, that "an exaggerated impression is given of the abundance on the grounds nearest Boston, which is the most important market center." " Classified in Canadian Fisheries statistics as "Digby County, from Sissiboo River to Annapolis County line." *' This appears clearly on Needler's (Contrib. Canadian Biol., N. Ser., vol. 6, No. 10, 1930, p. 5, fig. 1) chart of the distribution of the haddock catch, 1917- 1925. » Rich (Rept. U. S. Fish Comm. (1929) 1930, App. 3, pp. 51-117) gives a detailed account of the fishing grounds of the Gulf of Maine. In table 2, pp. 85-86, and table 3, p. 96, he lists 130 grounds in the inner parts of the Gull where haddock are taken regularly. 206 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Jeffreys Ledge north of Cape Ann, Stellwagen Bank at the mouth of Massachusetts Bay, and the several areas of "haddock bottom" off Chatham, Cape Cod. Small isolated rocky banks, such as Cashes and Platts, usually yield fewer haddock than cod, but in recent years of intensive fishing, haddock have been taken in numbers even on these so-called "cod grounds," as appears from the following table (landings to the nearest 1,000 pounds) : Locality 1919 1929 1934 1935 Platts Bank 68, 000 34,000 '320 1,094,000 736, 000 1,373,000 193, 000 83,000 494, 000 1,705,000 790, 000 1, 044, 000 75, 000 85, 000 423, 000 226, 000 682, 000 678, 000 18,000 26, 000 384, 000 27,000 236, 000 339, 000 i The reported landings from Cashes Ledge for 1919 were so small as to suggest some error. Spawning grounds. — One part or another of Georges Bank appears to be the most productive spawning ground for haddock off the American coast, one of the most productive anywhere, for that matter. And Walford's detailed studies60 have shown that haddock may spawn anywhere on the Bank eastward from Nantucket Shoals, except on Georges Shoals where the water is not deep enough. In most years there is a definite spawning center on the northeastern part of the bank, just east of Georges Shoals; Walford found this to be the case in 1931 and in 1932, corroborating our experiences on the Albatross I in 1920, when we found haddock eggs in great abundance 61 over an area there of at least 1,600 square miles. In 1932, there was a second spawning center in the so- called South Channel, where there seems to have been little spawning the year before. That Browns Bank, also, is a productive spawning center is proved both by Walford's studies, and by the fact that a fair proportion of the many gadoid eggs we towed there on the Albatross I in Aprd 1920 were far enough advanced in development to show a haddock parentage. Our own egg records, added to reports from the hatcheries and from local fishermen, show that haddock also spawn here and there, along the coastal belt from the entrance to the Bay of Fundy to Cape Cod, though in much smaller numbers than on Georges and Browns. » Bull. U. S. Bur. Fish., vol. 49, Bull. 29, 1938, pp. 3-12. •• Captures of ripe fish, male and female, in the trawl established the identity of these eggs as haddock, not cod. The more productive of the inshore spawning grounds which are neither as sharply circumscribed as those of the cod, nor as regularly occupied, are along the outer (eastern) and northern slopes of Stellwagen Bank, whence many eggs have been obtained for the Gloucester hatchery; the coastal belt between Cape Ann and Cape Elizabeth, especially off Ipswich Bay; the vicinity of the Isles of Shoals; about Boon Island; and off Wood Island, Maine. Breeding haddock are plentiful east of Cape Elizabeth in some years and scarce or altogether absent there in other years, or for terms of years. Thus, Captain Hahn, former superintendent of the Boothbay hatchery, has informed us that spawning haddock came into Boothbay Harbor in abundance and into Linekin Bay in April and May of 1912, while gill-netters made large catches in the general vicinity, but that spawning haddock did not approach this part of the coast at any time during the next 12 years in numbers large enough either to support any extensive fishery there, or to provide the hatchery with more than a few eggs. Spawning haddock have also been reported to us from the neighborhood of Mount Desert Island and off Cutler, Maine, while we found a few cod- haddock eggs near Petit Manan Island on April 12, 1920.62 But there is no reason to suppose that any considerable body of haddock spawn along the Maine coast east of Mount Desert, nor on the northern side of the Bay of Fundy, where neither eggs, larvae, nor young fry have ever been seen. However, our capture of a few haddock eggs63 and others in the younger "cod-haddock" stage (p. 203) in Petit Passage on June 10, 1915, proves that some spawn on the Nova Scotian side of the bay near its entrance; a few do so on the coastal banks along the western shores of Nova Scotia southward to Cape Sable according to general report, and we have taken a few cod or haddock eggs on German Bank in our tow nets in May. Turning, now, southward and westward, we learn that gill-netters sometimes get good fares of ripe fish off Boston Harbor, though no great body spawns in the inner part of Massachusetts Bay, and few if any on the cod-spawning grounds off « In a previous report (Bulletin, Museum of Comparative Zoology at Har- vard College, vol. 59, 1917, p. 258) we recorded eggs taken along this part of the coast in June as "cod-haddock", but fresh examination of the material shows that they might equally have belonged to the witch tiounder, none being sufficiently advanced in incubation to show the pigment. •' Far enough advanced to show the pigment in its distinctive arrangement. FISHES OF THE GULF OF MAINE 207 Plymouth (p. 192). Some ripe haddock are caught on the shelving-sandy bottom along Cape Cod as far south as Nauset; spawning fish, too, are caught off southern New England every winter. Nearly 800 baby haddock less than 1 year old were taken off Fire Island Inlet, Long Island, and 10 miles off Ambrose Lightship, in November 1948.6'1 But then" presence there does not necessarily mean that they were spawned so far west, as Dr. Howard A. Shuck of the Fish and Wildlife Service has pointed out to us. Haddock may at times deposit their eggs within a couple of fathoms of the surface in our Gulf, as, for instance, in Boothbay Harbor on the occasion just noted (p. 206). But this is most unusual, 15 to 20 fathoms being the upper limit to regular spawning with the depths of the more pro- ductive Gulf of Maine spawning grounds as fol- lows: Browns Bank, 30 to 50 fathoms and prob- ably deeper ; Georges Bank, from about 30 fathoms; Cape Cod grounds, about 40 to 70 fathoms; Stell- wagen ground, 20 to 40 fathoms; grounds between Cape Ann and Cape Elizabeth, 20 to 65 fathoms. The presence of newly spawned eggs out to the 100-fa thorn contour on the southeastern slope of Georges Bank at the height of the breeding season (late March 193 1)05 is evidence that the fish were spawning down nearly or to that depth. But about 100 fathoms appears to be the lower limit to any regular spawning. When eggs are found over greater depths they have drifted from shallower regions, as Walford has emphasized. The few eggs, for example, that we found over the deep basin of the Gulf, and in the Eastern Channel, in April 1920, were flotsam from the neighboring slopes or banks. The haddock spawn rather shoaler in the Gulf of Maine on the whole than they do in the North Sea region, where the maximum production of eggs takes place at 50 to 100 fathoms. Consequently, there is less difference in this respect between had- dock and cod in the western North Atlantic than in the eastern. Neither do haddock confine their spawning so definitely to smooth bottom in Amer- ican seas as they do in European waters. Welsh found ripe fish chiefly on broken ground "wherever sand, gravel, mud and rocks alternate — if any- thing, more are taken on the mud in such local- ities," between Cape Ann and Cape Elizabeth. The Gulf of Maine haddock spawn chiefly from late February until May and the following record, supplied by C. G. Corliss, former superintendent of the local hatchery, illustrates how brief the peak period of reproduction is near Cape Ann: Year First eggs taken Last eggs taken Period of greatest abundance Total eggs collected 1917 Apr. 16 Mar. 22 Feb. 12 Jan. 20 Jan. 22 May 3 Apr. 24 Apr. 30 Apr. 29 Apr. 25 10.820.000 1918---. .- Apr. 9 to Apr. 23- Feb. 20 to Apr. 23 Mar. 25 to Apr. 25 Jan. 27 to Apr. 14 32.380.000 1919 332. 740. 000 1920 303, 380, 000 1921 629, 130, 000 It appears from the hatchery records, cor- roborated by Welsh's experience in 1913, that the commencement of spawning varies considerably in date from year to year, with the fish breeding freely as early as the end of January in early seasons, but not until the end of March or even until the first part of April in late. But most of them are spawned out invariably by the middle or end of May at the latest. In normal years the spawning season is about the same on Georges Bank as it is near Cape Ann. In 1920, for example, we found cod-haddock eggs in moderate numbers across its western end late in February; great numbers of them (and took ripe haddock in the trawl) on the eastern end of the Bank on March 11 and 12; and they were still plentiful there on April 16 and 17, but we found none on the western part of the bank on May 17. Similarly, Douthart, of the Bureau of Fisheries, towed haddock eggs over the north-central portion of the bank on April 14 and again on the 26 and 27th in 1913, while Walford found that spawning commenced in February, was at its peak in March and April, and had about come to an end by late May in 1931. Spawning is likewise at its height in mid-April on Browns Bank (large egg catches were observed in our tow-nets April 16, 1920). Occasional haddock, however, may spawn long after the majority are spawned out. Thus we have towed eggs off Petit Passage, Nova Scotia, on June 10, and have caught a ripe female and a ripe male on Nantucket shoals on June 13 (in 1927). Ripe haddock have even been taken as late as the first part of July near Gloucester,66 but this is exceptional. The spawning season continues well into the summer in the colder water along the outer shores » As reported by Arnold, Copeia, 1949, p. 239. » Walford, Bull. U. S. Bur. Fisb., vol. 49, Bull. 29, 1938, p. 16, fig. 7. " Earll, Rept. TJ. S. Comm. Fish., (1878) 1880, p. 730. 208 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE of Nova Scotia and south of Newfoundland. Thus we took several unmistakable haddock eggs among numerous newly spawned cod or haddock eggs a few miles off Shelburne on June 23, 1915, while Dannevig 67 records occasional haddock larvae off Halifax on July 23; near Sable Island on July 25 and 26 ; and on St. Pierre Bank off Newfoundland on July 27 and 28 for that same summer. The breeding season is about the same in European as in American seas, that is, end of January until June, with the peak of production falling as early as March and April in the North Sea region but not until June around Iceland.63 The Georges and Browns Bank haddock spawn in temperatures ranging from about 36.5° to about 42°-43° F., and spawning is likewise completed on the coastwise grounds between Cape Cod and Cape Elizabeth before the stratum of water in which the fish are living has warmed more than a few degrees from its coldest for the year; i. e., in tem- peratures of about 35° to 40°-42°. Allowing for annual variations, this gives an extreme range of from about 35° to about 44° F. for the most active spawning over the Gulf of Maine as a whole, tem- peratures averaging considerably lower than those in which haddock spawn the most freely in Euro- pean waters (41° to 50°). The Gulf of Maine haddock likewise spawn in less saline water than does its European congener; and necessarily so, for the more important Gulf of Maine spawning grounds are considerably less saline at all depths and seasons (about 31.5 to 33.5 per mille, mostly). The specific gravity of the water at the tempera- ture in situ (the factor that determines whether buoyant fish eggs float suspended, and develop, or sink to the bottom and die) is usually between 1.0255 and 1.0270 in our Gulf in spawning season, at the depths where the fish spawn, both along shore and on the offshore Banks. Experiments by us and by Walford have shown that these values are high enough for the notation of tbe eggs. And while the water at the surface often is so light, near shore, as to interfere with the operation of the hatcheries, this layer of low specific gravity is so thin there is no reason to suppose that 17 Canadian Fish. Exped. (1914-15) 1919, p. 21. M Damas. Eapp. et Proc.-Vcrb. Cons. Internat. Explor. Mer, vol. 10, 1909; Schmidt, ibid. any of the haddock eggs produced in the Gulf fail to rise from the bottom.89 Populations and migrations within the Gulf of Maine. — Needler's 70 analysis of the results of tagging experiments, and of the differences in rate of growth between fish caught in different regions, and Vladykov's 71 studies of the number of verte- brae, confirmed by comparison between the growth rates of the haddock of Georges Bank and of Browns Bank by Schuck and Arnold,72 have shown that the haddock of North American waters include three more or less self-contained popula- tions; one (Needler's "New England population") inhabiting the Georges Bank-Nantucket shoals region and the inner waters of our Gulf from Cape Cod around to the New Brunswick shore of the Bay of Fundy; a second (Needler's "Nova Scot- ian") in the Nova Scotian side of the Bay of Fundy, and around Nova Scotia (including Browns Bank) to the Laurentian Channel; and a third in New- foundland waters. The geographic ranges of the New England and Nova Scotian populations are separated by the deep so-called "Eastern Channel" between Georges Bank and Browns, which extends inward as the "Fundian Channel" more than 100 fathoms deep, to the mouth of the Bay of Fundy. And it is probable that the depth is an actual barrier in this case, there being no evidence that haddock normally cross channels that are deeper than about 100 fathoms (at least in American waters), once they have taken to the bottom. Only within the Bay of Fundy, where there is no intervening water as deep as 100 fathoms, have tagging experiments given any evidence of a mixture between these two adult populations.73 And the still greater depth of the Laurentian Channel probably makes it an even more effective barrier between the Nova Scotian and the Newfoundland populations. The movements of individual fish within each of these populations fall in three groups: (a) those of the eggs and larvae while they are still adrift in the intermediate and upper water layers; (b) those of the young fry from the time they take H For a discussion of the relationship between flotation of haddock eggs and the specific gravity of the water, with references to European studies, see Walford, Bull. U. S. Bureau of Fisheries, vol. 49, Bull. 29, 1938, pp. 13-15. n Contrib. Canadian Biol, and Fish., N. Ser., vol. 6, No. 10, 1930. " Progress Rept. Atlantic Biol. Sta. Biol. Board, Canada, No. 14, 1935. '- Fish. Bull. No. 67, U. S. Fish and Wildlife Service, 1951. 73 One fish that was tagged by us near Mount Desert Island was recaptured in the Nova Scotian side of the Bay of Fundy oft" Digby. FISHES OF THE GULF OF MAINE 209 to bottom until they are large enough to figure in the commercial catches, and (c) those of the larger fish. It may be assumed that the pelagic life of the haddock lasts about as long in American waters as in European; i. e., for three months or so (we have no first-hand information) before the fry seek the bottom. Meantime the eggs and larvae, like those of many other fishes may drift for con- siderable distances from where they were spawned. And these involuntary drifts may be greatly ex- tended by a habit that the very young haddock have (like those of other gadoids) of living under the bells of the larger kinds of jellyfishes. Welsh, for instance found many small haddock of 2% to 3 inches (60 to 77 mm.) in company with the common red jellyfish (Cyanea) on Georges Bank and off Nantucket Island, in late July of 1916, while Willey and Huntsman 74 found young haddock about 2 inches long under Cyanea in the Bay of Fundy. In fact, it is in company with Cyanea that young haddock in the late larval stage have been taken most often in the other side of the Atlantic. Our few records for the pelagic larvae in the inner parts of the Gulf all have been in the south- western part. Thus the coastal zone east of Cape Elizabeth, and the whole deep basin of the Gulf, seems to be as barren of larval haddock (so far as our catches go) as of larval cod, of larval silver hake, of larval flatfishes, and, in fact, of most other larval fishes except rosefish (p. 433) and herring. It appears from Walford's studies that in normal years, as represented by 1931, the haddock population of Georges Bank is recruited by a good supply of larvae hatched from eggs that have been spawned on the bank itself. But a large proportion of the Georges Bank eggs and larvae drift off the bank in other years, as in 1932, either to the westward and southward past Nantucket Shoals, where their mortality is too great for them to support a population of any importance, or southward out over the continental slope, to even more certain destruction, 76 with results disastrous to the ensuing brood of young fry (p. 212). It is during their pelagic stage (whether drift- ing independently or with Medusae) that inter- mingling is the most likely to take place in signifi- cant amount between the New England and the Nova Scotian populations of haddock. All that is known in this respect is that Georges Bank seems not to have received any important re- cruitment from elsewhere, either in 1931 or in 1932. In any case, hosts of young fry settle on the bottom on the offshore banks generally. Thus we have repeatedly found 10 or more little haddock 3 or 4 inches long, in the stomachs of pollock caught on Georges, while we have trawled numbers of equally small ones there as well as on the other offshore grounds. And 1- to 2- year -old fish, 6 to 12 inches long (too small to market) sometimes make up as much as 35 to 40 percent of the total catch of haddock on Georges as well as in the South Channel, while many more of them doubt- less escape through the meshes of the trawls. On the other hand, very young haddock are seldom seen inshore for they are too small to be caught either on long lines or in gill nets. But it is probable that they are plentiful there, also, for yearlings are reported in the Bay of Fundy, by Huntsman. Notliing is known about the movements of the young haddock during the first year or two after they take to the bottom. But our fisher- men have long realized that the larger haddock, like the larger cod, are so constantly on the move in search of food that the fishing may be poor tomorrow where it was good today, or vice versa. And analysis of the catches that we made on Nantucket Shoals during the tagging campaigns of the U. S. Bureau of Fisheries, 1923-1931, shows that considerable changes took place in the abundance of fish within periods of a few days or weeks at the spots fished, also with occasional brief periods of unusual abundance that are most reasonably interpreted as reflecting the passage of large bodies of fish from else- where.79 The extensive tagging experiments that we have made within the Gulf of Maine on vessels of the U. S. Bureau of Fisheries, 77 and that have been made in Nova Scotian waters78 by the Biological Board of Canada have now proved " Canadian Field Natural, vol. 35, 1921, p. 2. "For further details we refer the reader to Walford's very interesting study (Bull. U. S. Bur. Fish., vol. 49, Bull. No. 29, 1938). " For details, see Schroeder, Jour. Marine Research, vol. 5, No. 1, 1942, p. 9, table 2. " Schroeder, Jour. Marine Research, vol. 5, No. 1, 1942. " Needier, Contrib. Canadian Biol, and Fish., vol. 6, No. 10, 1930. 210 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE (as was previously suspected) that most of the wanderings of the Gulf of Maine haddock are of short extent. Thus 34 fish that had been tagged on Nantucket Shoals were recaptured nearby, 16 after periods longer than 200 days; only 10 were captured at a distance. And the pre- ponderance of relatively stationary fish is even more impressive for the vicinity of Mount Des- ert Island, where 114 tagged fish were recaptured within a few miles of the tagging stations after an average period of 224 days, contrasting with recaptures of only 25 of them at a distance. The haddock of the coasts of Massachusetts and of western Maine, with the offlying banks, may be less stationary, for only two of the fish that were tagged on Stellwage.n Bank and be- tween Boone Island and Boothbay were recap- tured locally; 13 of them far afield. The tagging experiments do not suggest that such of the Gulf of Maine haddock as do wander follow any regular migratory routes. Thus some of the few Nantucket Shoals fish that are known to have strayed were recaptured to the eastward (eastern part of Georges Bank, 2) ; some of them to the northward (western side of Gulf and Platts Bank, 6); and some to the northeastward (nor- thern entrance to Bay of Fundy, 2). Conversely, it was in the opposite direction, i. e., to Platts Bank, to the coasts of western Maine and of Mas- sachusetts, to the South Channel, and to Georges Bank that wanderers are known to have strayed from the Mount Desert tagging ground. And the few fish that were recaught from those tagged at localities intermediate between Nantucket Shoals and Mount Desert, have fanned out in various directions. An obvious reason why haddock of the New England population, that commence their adult journeyings in the northeastern part of the Gulf, should tend to stray southwestward, southward, and perhaps then eastward along Georges Bank, whereas others, commencing in the southwest should tend either eastward, or northward and then northeastward, is that these are the only routes left wide open to them within the Gulf, between the coastline on the one side and the barrier that is set for them by the 100-fathom depth line on the other side. How effective is this barrier is emphasized by the fact that only one fish, among 9,416 that we tagged off the coasts of Massachusetts and of Maine was recaptured in Nova Scotian waters (it had gone from Mount Desert to the southern side of the Bay of Fundy) ; and that none of the haddock that were tagged in Nova Scotian waters by the Biological Board were recaptured west of the Fundian Channel. Very little is known as to the shifts in location and in depth that haddock may make between winter and summer, the difficulty lying in the in- terpretation of the differences from season to season in the amounts of haddock that are caught on neighboring grounds in the inner parts of the Gulf.76 In general, it appears that when the temperature of the upper 15-20 fathoms of water rises above about 50° to 52° F., as happens along the coasts of Maine and Massachusetts in July or August, the haddock tend to withdraw from the shallower grounds where they are plentiful in spring and early summer. But certain bodies of fish may linger all summer in the deeper channels among the islands of Maine, on patches of suitable bot- tom. In 1923, for instance, haddock were caught throughout July, August, and September, be- tween Suttons Island and Bear Island, near Mount Desert Island, as well as at other inshore localities near by. Fishermen report them as working inshore again in autumn or early winter, as the water cools, but those that come closest inshore then are supposed to work out again, in mid and late winter, to avoid extreme chilling. Thus few or none are caught at that season in the Bay of Fundy, where the temperature may fall as low as 32° in occasional winters,80 though it does not drop below 34° to 36° in most years. We must caution the reader, however, that these supposed disappearances in winter from inshore localities are based on failure to catch haddock then on hook and line, which may actu- ally result more from a reluctance on their part to bite at low temperatures (p. 202) than from sea- sonal scarcity of fish. Experimental trawlings at different seasons are needed to clarify this matter. At any rate, the temperatures of the open Gulf of Maine at the depths where haddock are the most plentiful never fall too low for their comfort in the winter, nor rise too high in the summer. '» Rich (Rept. V. S. Fish Comm. for 1929, 1930, App. 3) gives information in this respect. *> As happened in 1926 (Needier, Contrib. Canadian Biol, and Fish., N. Ser., vol. 6, N'o. 10, 1930, p. 19 [259]). FISHES OF THE GULF OF MAINE 211 Accordingly, haddock are caught on all the major grounds the year around. Except for shifts in depth, apparently asso- ciated with temperature, the haddock as a whole are year-round residents as far east as the offing of southeastern Nova Scotia (Roseway, La Have, and Sambro Banks) ; many of them as far east as Halifax and Sable Island Bank. But they appear only as late spring to early winter visitors farther to the east and north where they are wholly absent (as indicated by the catches) in late winter and early spring. Large catches, for example, are made in traps near Ingonish on the northeastern coast of Cape Breton Island in late May and in June. The first haddock are caught within the Gulf of St. Lawrence in June, whether on the Cape Breton shore, or westward, the largest catches are made there from July through the late autumn, and very few are taken as late as December. But catches are made again near Ingonish in December and January, of haddock, seemingly en route out of the Gulf of St. Lawrence. And it now seems established that these visitors to the Gulf of St. Lawrence pass the late winter and early spring on Sable Island Bank and farther to the westward in Nova Scotian waters.81 Part of the haddock in the southern part of the Grand Banks region, which form a distinct popu- lation separated from that of Nova Scotia by the deep Laurent ian Channel, are described by Thompson 82 as making a summer inshore migra- tion to the southwest coast of Newfoundland, and as clearly avoiding regions where the bottom water is colder than about 34° F. Abundance. — The haddock and the rosefish rank next after herring in numbers, among the fishes of our Gulf that are important commercially. In good years it has not been unusual for a trawler to take 10,000 to 20,000 haddock in 5 or 6 days' fishing on the Georges Bank and South Channel grounds; a catch of 240,000 pounds of haddock (something like 70,000 fish) brought in by the trawler Fabia in March 1926 is one of the largest of which we have chanced to hear. One must remember, too, that this represents only the fish that are large enough to be worth saving, and that multitudes of baby haddock too small to be marketable, caught on Georges, are thrown back dying or dead; in 1947, for example, the number so wasted was estimated at almost 17 million on Georges Bank alone.83 Howard W. Schuck in- forms us that the average weight of the haddock landed from Georges Bank during 1928 was about 3% pounds. Fishermen have long been aware that the had- dock vary widely in abundance from year to year and over periods of years, on one ground or an- other, independent of any effects the fishery may have had on the numbers of fish. It has been amply proved by investigations both in Europe and in North America, that these fluctuations re- sult chiefly from differences, from year to year, in the number of young that survive and take to the bottom on the grounds in question; the Gulf of Maine is no exception. The production, for ex- ample, of young haddock at the mouth of the Bay of Fundy, on the New Brunswick side was low from 1915 to 1919, very much higher in 1920, but somewhat lower again in 1921-1923 though some- what better than it had been in the 5 years pre- ceding 1920.84 But a larger number of haddock (by report) were produced near Digby, on the Nova Scotian side of the Bay in 1921 than had been in 1920.85 Similarly, two exceptionally successful year classes that were spawned in the Georges Bank- South Channel region during the period 1921-1924 were followed by poor year classes from 1925- 1928, but then by an abundant class that was spawned in 1929.86 Since then Georges Bank has been abundantly recruited with haddock fry in 1936, 1937, 1939, 1940, and 1945.87 On the other hand, the crop, so to speak, was unusually scanty on the Bank in 1930, 1931, 1932, 1942, and 1947. Perhaps a good crop comes a little more often for the Nova Scotia population, and every 3 years or so in the North Sea, "where the fry have a much better chance of being retained in the axea owing to the prevailing currents."88 11 For further discussion, see A. W. H. Needier, Contrib. Canadian Biol, and Fish., vol. 6, No. 10, 1930, and A. B. Needier, Bull. 25, Biol. Bd. Canada, 1931. 8! Research Bull. No. 6, Newfoundland Dept. Nat. Resources. 1939, p. 7. »i Schuck, Commercial Fish. Review, vol. 10, No. 10, October 1948, p. 6. " Huntsman and Needier, Contrib. Canadian Biol, and Fish., N. Ser., vol. 3, No. 18, 1927, see summary on p. 14 [4361. " Needier, Contrib. Canadian Biol, and Fish., N. Ser., vol. 6, 1930, No. 10, p. 44 [2841. "• The data for 1921-1929 are summarized in the Proc. No. 2, for 1931- 1933, N. Am. Council on Fishery Investigation, Ottawa, 1935, p. 13. " From dat3 supplied by Howard A. Schuck of the U. S. Fish and Wild- life Service. << Thompson, Res. Bull. No. 6, Newfoundland Dept. Nat. Resources, 1939, p. 22. 212 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Analyses from year to year of the relative pro- portions of fish of different ages in the catch 89 show, too, that our offshore banks may receive as much as 30 times as many fry in a good year as in the average for a run of years, and as much as 60 times as many as in the poorest years. One essential for a good year class of haddock, perhaps the chief essential, is that large numbers of larvae shall not only be hatched and survive until old enough to take to bottom, but shall remain in the area in general, to take to bottom there, as happened in 1931, and not drift else- where. Conversely, a poor brood automatically ensues if the circulation of the water is such that a large proportion of the larvae drifts away, as happened in 1932, when so many of them drifted off Georges Bank altogether, to be lost perma- nently to the local population, that the success of that year class was seriously affected.90 Herrington has also suggested that in years when large fish are the most plentiful the resulting competition for the supply of available food makes conditions difficult for the surviva' and growth of the young fry. Evidence is that the "largest spawning stocks have almost invariably yielded the leanest year classes 3 years later, and the poorer spawning stocks have done much better."91 No doubt a combination of various other factors helps to determine whether any particular year class shall be plentiful or the reverse. But the relative im- portance of these factors has not yet been evalu- ated for our haddock. The incidence of a good brood in any particular year, or the reverse, shows up in the commercial catch 2 years later; i. e., when the young fish first reach market size in significant numbers. And it is now well established, for both sides of the Atlantic,92 that the differences in the numbers of fry reared in different years are the chief cause for the short term fluctuations in the catches that are so characteristic of the haddock fishery. Our reason for emphasizing the qualification "short term" in this connection is that the situa- tion is complicated by the unhappy fact that the haddock populations of Georges and Browns Banks have been seriously reduced by the fishery. Commercial importance and effects oj the fishery. — The haddock was once much less in favor than the cod. But the expansion of the fresh-fish trade93 brought an increasing acceptance of haddock on the market because of their good keeping qualities and convenient size for the table. In 1919 the Gulf of Maine, inshore and offshore combined, yielded something like 85 million pounds of haddock to United States and Canadian fishermen. And the development of the filleting and packaging of fresh and frozen haddock soon brought so great an increase, both in the demand and in the intensity of the fishery, that some 206 million pounds were caught in 1929 from the New England population, with some 17 million pounds more from the Nova Scotian popu- lation on Browns Bank, off western Nova Scotia, and in the Nova Scotian side of the Bay of Fundy, making a total of at least 223-224 million pounds from the Gulf of Maine as a whole, corresponding to perhaps 60 to 70 million indi- vidual fish. This, however, was the high point, for trawlers working on Georges during the five years, 1930- 1934, "averaged scarcely one-third as much had- dock per day as during the previous five years," 9* while the Gulf of Maine catch as a whole had fallen by 1934 and 1935 to only about one-quarter of what it had been in 1929.95 Since then, down to 1947 (most recent market year for which we have seen the returns), the yearly yield of market-size haddock from the New England population has varied between about one-third to one-half as great, and about two-thirds as great as it was in 1929, to judge from the landings in the major New England ports, which form at least 90 percent of the total take from this population.96 A recent estimate is that there were only about one third as many haddock on Georges Bank in 81 From unpublished data for Georges Bank and the South Channel area supplied by Howard A. Schuck. » For details, see Walford's (Bull. U. S. Bur. Fish., vol. 49, Bull. 29, 1938) very interesting study of the drift of the Georges Bank eggs and larvae in these two years. 11 Tians. 9th North American Wildlife Conference, 1944, p. 260. M See especially Thompson's studies for Iceland (Fisheries Scotland, Sci. Invest. [1928], No. 5, 1929), and Raitt's for the North Sea (Journal du Conseil, Cons. Internat. Explor. Mer, vol. 11, No. 2, 1936, p. 211). 03 Fish that are iced at sea, not salted. » Herrington, Fishery Circular No. 23, IT. S. Bur. Fish., 1936, p. 9. " About 78 million to 80 million pounds in 1934, judging from the landings at Portland, Boston, and Gloucester from within the Gulf (which usually run about 9i *-S of the total catch in the Gulf by United States and Canadian vessels combined) plus perhaps 4 million to 5 million pounds taken by Canadian fishermen off western Nova Scotia and in the Bay of Fundy. 11 For tabulations of the total catches of haddock in the western Atlantic by Canadian and United States vessels, 1880-1927, see Needier, Contrib. No. 2, North American Council on Fish. Investigations, Ottawa, 1929, 13 pp., also Kept. U. S. Comm. Fish. (1930) 1930, App. 2, pp. 27-40. FISHES OF THE GULF OF MAINE 213 1931 as there had been there a year or two earlier.97 This conclusion is based on the assumption that yearly changes in the average yearly catches, per day's fishing of a standard group of the large otter trawlers, fishing consistently for haddock, over the period in question, have been propor- tional to the relative changes in the number of haddock on the banks. In 1939-1947 the catch statistics suggest that the total population on the banks had, on the average, increased somewhat from the relatively small population of 193 1.98 Landings of haddock Landings by United States vessels in the major New England ports, to nearest 100,000 pounds Canadian landings to nearest 100,000 pounds Total Oulf Year Georges Bank, Nan- tucket shoals, and inner parts of Gulf of Maine Browns Bank and off western Nova Scotia Western Nova Scotia and southern side of Bay of Fundy of Maine 1929 174, 700, 000 45, 300. 000 66, 200, 000 78, 500, 000 87, 500. 000 83, 200, 000 95, 600. 000 88, 800, 000 116,400,000 101, 400, 000 89, 700. 000 86, 800, 000 72, 500, 000 99, 300, 000 107, 400. 000 8,200,000 14,800.000 18, 000, 000 13, 600, 000 14, 900, 000 22, 500. 000 11,300.000 8,200.000 6, 100. 000 6,100.000 2, 800. 000 4, 400, 000 6, 000, 000 5, 200. 000 4,900,000 11, 500. 000 6, 500. 000 6,500,000 5, 100. 000 4, 700, 000 8,200.000 7.200.000 7, 100, 000 6,600.000 6. 900, 000 4. 600. 000 6, 200, 000 6, 400. 000 5. 200, 000 1934 1935 1936... 89,700,000 97, 200. 000 1937 1938 1939 1940 114,100.000 1941 128. 100.000 112.4110.000 1942... . 1943 1944 96,41X1, IKK) 83,900.000 109, 700, 000 1945 1946 1947 The yield from Browns Bank and the Nova Scotian side of the Gulf has also been significantly smaller since 1939 than it was during the few years previous, when American vessels began to fish Browns Bank more intensively than they had previously. The persistence of poorer catches through so long a term of years in the face of sustained demand, added to continued improvement in the gear and in the general efficiency of the fishing fleet, is only too clear evidence of overfishing. The decrease in the yield of haddock from within the Gulf of Maine has been partially offset by increased catches from the Banks along outer Nova Scotia eastward to Banquereau Bank. The landings, for example, were about 8 times as great, from east of Cape Sable in 1947 (about "Herrington, Trans. 9th North American Wildlife Conf., 1944, p. 259. Schuck, Commercial Fish. Rev., vol. 10, Oct. 1948, p. 1. " See Schuck (Biometrics, Amer. Statistical Assoc, vol. 6, No. 3, 1949, p. 215, table 1, and p. 216, fig. 2). 26,400,000 pounds) as had been the case back in 1929 (about 3,300,000 pounds). Further dis- cussion, however, of the fishery aspects of the matter would lead us too far from our main theme. Previous to the general adoption of the otter trawl in American waters, haddock were caught mostly on hand lines or on long lines; some in gill nets, especially in spawning time inshore between Cape Ann and southern Maine. Today all but a very small part of the catch is made in otter trawls. In 1947, for example, nearly 97 percent of the haddock that were landed in Maine and Massachusetts had been taken in otter trawls; only 3 percent of them on long lines; and only a small fraction of 1 percent on hand lines and in gill nets. While the haddock is of primary interest from the commercial standpoint, it deserves a word from the angler's viewpoint also, for it bites as freely as the cod does, on almost any bait, and, being a much more active fish, a haddock of fair size is likely to prove an astonishment to anybody who is lucky enough to hook one while fishing with a light sinker. A new-caught haddock is also a very beautiful object. American pollock Pollachius virens (Linnaeus) 1758 Pollock; Boston bluefish; Coalfish (in Great Britain); Green cod (in Great Britain) Jordan and Evermann, 1896-1900, p. 2534. Description. — The American pollock 99 has a deep, plump body (about four and one-fourth times as long as it is deep) tapering to a pointed nose and to a slender caudal peduncle. Its mouth is of moderate size. Its projecting lower jaw (giving it an undershot facial aspect); its forked, sharp-cornered tail, small ventral fins, small chin barbel (as a rule the latter is lacking altogether in large fish), and its beautiful olive green color, are ready field marks when it is caught with cod and haddock. Its first dorsal fin (13 or 14 rays), originating slightly behind the pectoral, is triangular, and is a little the highest of the three dorsals. The second dorsal, also triangular, is the longest of the " This is the "coalfish, green cod, or saithe" of British, Scotch, and Irish fishermen. The European "pollack" is a different species (Qadus pollachiua) 214 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Length, in Weight, in Length, in Weight in inches pounds inches pounds 24H 4-5H 30 8H-9H 26 4 31 10 27 V-4 31 H 10 Z?H 8H 32 10-12 28H 8 33 12 29 8-9 35 14 29'A 8H-9 Figure 98. — American pollock (Pollachius virens), Eastport, Maine. From Goode. Drawing by H. L. Todd. three (21 or 22 rays) and is separated by a con- siderable space from the third dorsal fin (19 or 20 rays) which is more rhomboid in outline. The second anal fin (20 or 21 rays) corresponds in shape and size to the third dorsal, under which it stands, but the first anal (24 to 28 rays) is consid- erably longer than the second dorsal though similar to the latter in shape. The ventral fins are a little in front of the pectorals, and are only about half as long as the latter. The pectorals are set high on the sides, and are longer than the first dorsal, but shorter than the second dorsal; they have rounded lower corners and bluntly pointed tips. The caudal fin is noticeably forked, with angular corners, unless it is spread to its widest when its margin becomes nearly straight. Color. — Pollock are always of a greenish hue, usually deep rich olive green or brownish green above, paling to yellowish or to smoky gray on the sides below the lateral line, and to silvery gray on the belly. The lateral line is white or very pale gray, contrasting strongly with the dark sides. The dorsal, caudal, pectoral, and anal fins are olive, the latter pale at the base. The ventral fins are white with a reddish tinge. Young fish are darker than large ones, and many of them are more tinged with yellow on their sides. Size. — Pollock reach a maximum length of about 3% feet and a weight of about 35 pounds. But fish of this size are exceptional, few growing larger than 40 inches or 30 pounds, with about 2 to 3 feet and 4 to 15 pounds as the average for adults. The proportion of length to weight was as follows among fat fish measured by Welsh off Boon Island on April 22 to 25, 1913: Large pollock, however, of a given length vary widely in weight; for example, we have found 40- inch fish to weigh from 25 to 35 pounds; 35-inch fish, from 14 pounds to 21 pounds. Habits. — The pollock is an active fish, living at any level between bottom and surface according to the food supply and on the season, often school- ing, and sometimes gathering in bodies so large that it is on record that a purse seiner once took 60,000 fish from one school at a single set. In our Gulf their depth range is from the surface down to 100 fathoms at least,1 while they may descend somewhat deeper in the deepest troughs. And it is the local presence or absence of prey that gov- erns the movements of the larger pollock. Pollock feed chiefly on small fish, and on pelagic crustaceans; among the latter most often on the large pelagic shrimp-like euphausiids. It is com- monplace that pollock destroy great quantities of small herring, launce, young cod, young haddock, young hake, silver hake, and other small fish in the Gulf of Maine just as they do on the other side of the Atlantic. Pollock chasing schools of herring are a familiar sight;2 pollock of 1 to \){ pounds commonly run up estuaries in pursuit of smelt in autumn; and newly hatched haddock or other ' We have seen them trawled as deep as this on the northern slopes of Georges Bank. ' Sars (Kept. U. S. Comm. Fish., (1877) 1879, p. 619-620) has given a graphic account of pollock rounding up schools of launce and of young cod in Nor- wegian waters. FISHES OF THE GULF OF MAINE 215 larvae that are liberated in harbors from the hatcheries are always in danger of being snapped up by the young pollock that are plentiful in such situations. When a pollock only 9 inches long is capable of eating 77 herring up to 2% inches long at one meal,3 "ravenous" is only mildly descriptive. However, pollock so seldom strand in pursuit of prey that we have never seen one on the beach though schools often come close in and are caught in the traps. In the Gulf of Maine, pollock depend perhaps as much on pelagic shrimps as on fish. At East- port, for example, where these shrimps (genera Meganyctiphanes and Thysanoessa) are very abun- dant all summer, Kendall 4 reports pollock of all sizes not only fattening on them but so evidently preferring them to young herring that he did not find a single "sardine" in a pollock stomach, though these were plentiful enough at the time. He adds that "if at any time the crustaceans disappeared from a place the large pollock disappeared also." And pollock, breaking the surface in pursuit of shrimp are familiar sights there, as we can bear witness with many others. Similarly, Welsh found large pollock in schools feeding on the surface on shrimp (Thysanoessa raschii) off the Isles of Shoals and off Boon Island in April 1913, remarking in his field notes for the 25th that "in the last few days pollock have begun to appear in small schools of 400 to 500 fish with the appearance of large schools of feed (shrimp, 'all eyes'), the feed (shrimp) breaking water trying to get away from the pollock which are after them." He described the fish themselves as "rising and sinking at intervals ; when at the surface swimming like porpoises, leaping up and over with open mouths, the feed being in dense streaks 6 inches to 1 foot down." These feeding fish were "very sluggish and tame on this feed and easily taken in the purse seines." All were "stuffed to capacity" with shrimps, and only a few contained herring. Large pollock take morsels as small as copepods. Willey 5 for example, speaks of a fish caught near Campobello Island which contained proportion- ately as many of these as of euphausiid shrimps, and it is probable that the little pollock depend chiefly on copepods. Glass worms (Sagitta), too, » Smitt, Scandinavian Fishes, vol. 1, 1892, p. 503. * Rcpt. U. S. Comm. Fish., (1896) 1898, p. 180. • Proc. Amer. Acad. Arts, Sci., vol. 56, 1921, p. 192. have been found in pollock stomachs. Sometimes they consume considerable quantities of cteno- phores; we found many pollock full of them on Cashes Ledge and on Platts Bank in August 1928; one had 105 of these watery organisms in its stomach. They also feed to a small extent on bottom-dwelling crustaceans on both sides of the Atlantic, thus crabs, and bottom-dwelling shrimp have been found in fish caught at Woods Hole and in the Gulf of Maine. They have also been reported as gorging themselves on herring spawn. They never take shelled mollusks, so far as we are aware. But the}7 bite on clams as greedily as on fish baits. And fishermen speak of them as one of the few species that will bite, that is, feed, during the spawning period. Experiments on fish kept in captivity at Woods Hole 6 have shown that the pollock captures its food more by its keen sight than by scent. The pollock is a cool-wa ter fish. We have never seen any large ones caught at the surface when the temperature there was higher than about 52° F., though there may be plenty of them a few fathoms deeper down where the water was cooler. Even the little "harbor pollock" of 8 inches or so do not appear in any great numbers at times or places where the water is warmer than perhaps 60° F. At the other extreme, pollock of all sizes from the 1 year-old fish upward must experience temperatures as low as 32° F. on the fishing grounds in the southern side of the Gulf of St. Lawrence, and on the more easterly of the Nova Scotian banks during the late winter or early spring, unless they descend then to considerably greater depths, a possible shift of which we have no direct evidence. But it is probable that the pollock's need of water as warm as about 38° F. for the incubation of its eggs (p. 216) and perhaps of temperatures a little higher than that for the maturation of its sex organs (p. 216) is the factor that sets the northern boundary to the mainte- nance of a permanent resident population (p. 2181. The pollock is a late autumn and early winter spawncr, and the shortness of the spawning season (p. 220), with the fact that the vertical temperature gradient covers a range no greater than 3° to 5° F. down to 50 fathoms at that season, makes it easy to establish the physical conditions under which the eggs are produced • Herrick, Bull. U. S. Comm. Fish., vol. 22, 1904, p. 258. 216 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE and in which they develop. On the Massa- chusetts Bay grounds breeding commences when the whole column of water has cooled to about 47° to 49°, and is at its climax (late in December) in temperatures of 40° to 43°, while the major production of eggs takes place long before the water has cooled to its winter minimum of 35° to 36° F. at the level at which the fish lie. Thus tbe pollock spawns on a falling temperature, with most of the eggs produced within a comparatively narrow range and in water several degrees wanner than that in which haddock spawn most actively (p. 208). This agrees closely with the European pollock which spawns only in temperatures near 44.5°, so far as is known. As the successful propagation of any fish depends as much upon the incubation of its eggs as on its spawning, we should note that incubation proceeds normally, and that the resultant larvae are strong and active over the whole range of temperature just outlined, that is, from about 38° to about 48° as proved by experience in the Gloucester hatchery. The Massachusetts Bay spawning takes place in salinities ranging from as low as 32 per mille to as high as 32.8 per mille, according to precise locality, depth, and season, salinities much lower than those in which pollock breed on the other side of the Atlantic (35.14 to 35.26 per mille). The number of eggs produced by a female pol- lock averages about 225,000, but more than 4 million eggs were reported in one fish of 23% pounds. The egg is buoyant, has no oil globule, and averages about 1.15 mm. in diameter. Thus it is decidedly smaller than the egg of the cod or of the haddock. Incubation occupies 9 days at a temperature of 43°; 6 days at 49°. The larvae are about 3.4 to 3.8 mm. long at hatching, slender, with large yolk sac, and with the vent situated on one side of the body at the base of the ventral fin fold as it is in other larval gadoids; they are sprinkled with black pigment cells. About 5 days' time is required for the entire absorption of the yolk sac and for the formation of the mouth; meantime the pigment of the post-anal section of the trunk becomes grouped in longitudinal bars, two dorsal and two ventral, the former longer than the latter. At this stage pollock closely resemble cod of the same size, but the ventral bars are longer than the dorsal bars opposite them in the cod, and usually three in number in the cod instead of two as in the pollock. These bars persist until the pollock grows to a length of about 15 mm., when the pigment becomes more scattered. The caudal fin rays appear at about 9 mm., all the dorsal and anal rays and the ventral fin rays at about 15 mm., the dorsal fins are separate from one another and also the anal fins at 20 mm. (at about 2 months), and fry of 25 to 30 mm. show most of the characters of the adult. In European seas the young pollock lives near the surface for its first 3 months. The young fry have been taken similarly, in the tow nets near the surface at Woods Hole from Jan- uary to May, and they are to be expected in Massachusetts Bay then, though we have no actual record of them there. Rate of growth. — Thanks to the shortness of its breeding season and to the readiness with which its scales can be "read" European students 7 have found it easy to trace the rate of growth of these "saithe" or "coal fish"; and this has been done for the American pollock by Mavor,8 also by us. Judging from scale studies and from the sizes of the fry that are caught near Woods Hole in the spring, pollock hatched in mid-winter are about 1 to 2 inches long by the following spring, growing to 3-5 inches by late summer; to 5-7 inches in their first winter, when a year old; to 12-13 inches at two years of age; to 17-18 inches at three years; to 21-22 inches at four years; to an average of about 23 inches at 4% years; of 25 inches at 5% years; and of about 27 inches at 6K years. Thus the 11-12 inch and 15-16 inch pollock that appear in such numbers along the New England coast late in summer are about 1% and 2% years old, respectively. The annual rate of growth thus is about 5 or 6 inches for the first three years, 2-4 inches for the next three years, and 1-2 inches for the next few years, after which they grow still more slowly. These sizes are somewhat larger than the aver- ages given by Damas for European fish of corre- sponding ages, but the difference is so small that it is safe to apply the European figures to older Gulf of Maine fish, for which we have no data. On this basis we may expect the American pollock to average about 28 inches at 7)i years; about 29 ' For resume see Damas (Rapp. et Proc. Verb., ConseU Perm. Internat. Explor. Mer, vol. 10. No. 8, 1900, p. 167). » Contr. Canad. Biol., (1917-1918) 1918, No. 6. FISHES OF THE GULF OF MAINE 217 Figure 101. — Larva (European), 6.75 ram. AfterSchmidt. Figure 99. — Egg (European). After Mcintosh. Figure 102. — Larva (European), 12.5 mm. AfterSchmidt. Figure 100. — Larva (European), 5 days old, 4.3 mm After Mcintosh. FIGURE 103. — Fry (European), 23 mm. After Schmidt. American I'ollock (Pollachinn virens). inches at 8% years, and about 30 inches at 9}i years. Fish of 3 feet and upward are therefore of considerable age. The oldest recorded b}r Damas among the thousands he examined was in its nineteenth year. In European seas pollock grow faster in the southern part of their range than in the northern, but we have yet to learn whether this applies to the American fish. The age at which Gulf of Maine pollock lirsi mature is not known, but this is probably at a somewhat greater size than in Norwegian waters, where most of them mature by the time they are 1% feet long; i. e., 3 years old. All of them that are 2 feet long, or longer, in summer have spawned at least once. Oeneral range. — Continental waters on both sides of the North Atlantic in cool temperate and boreal latitudes; regularly in the west from the southeastern part of the Gulf of St. Lawrence 9 ami northeastern Nova Scotia to New Jersej ; southward occasionally to Chesapeake Bay and to Cape Lookout,10 N. C, and northward in small numbers to the southern part of the Grand Banks, to the southeastern coast of Newfoundland, and 9 Pollock appear not to be known anywhere farther within the Gulf or in its northern side. 10 Reported from Chesapeake Bay by Hildebrand and Schroeder (Bull. U. S. Bur. Fish., vol. 43, pt. 1, 1928, p. 156) and from Cape Lookout by Colos (Copeia, No 151, 1926, p [105)). to Sandwich Bay on the southeastern coast of Atlantic Labrador; " West Greenland; Spitz- bergen; Iceland; and the coasts of northwestern Europe south to the North Sea, English Channel, and Brittany coast of the Bay of Biscay in the eastern Atlantic; occasionally to the Gulf of Gascony (Arcachon). Occurrence in the Qvlj oj Maim . — In our side of the Atlantic the pollock lias its chief center of abundance in the Gulf of Maine, where it is caught in large numbers both on the offshore hanks, and all around the coast line, from Nant ticket Shoals and Cape Cod to Cape Sable. The only regional exception is in the inner part of the Bay of Fundy along the New Brunswick shore, where so few pollock are taken that they do not appear at all in the landings reported thence (Albert County). The following statistics of the United States catch for 1945, combined with the Canadian catches for 1944 and 1946, '- give a general idea of the regional abundance of pollock, on a broad scale, also of how universal they are, with the one exception just noted. Browns Bank, about 965,000 pounds; western coast of Nova Scotia to the Annapolis County H The pollock is listed in the Reports of the Newfoundland Fisheries Research Commission for 2 stations on the southern edge of the Grand Bank, from Bay Bulls, Newfoundland, and from Sandwich Bay, Labrador. ii We have not yet seen the Canadian statistics for 1945. 210941—53- -15 218 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE line, at least 7,300,000 pounds; I3 Nova Scotian shore of Bay of Fundy, about 500,000 to 1,000,000 pounds; M inner part of Bay of Fundy on New Brunswick side (Albert County), 0; mouth of the Bay of Fundy on New Brunswick side, about 2,000,000 to 3,500,000 pounds;15 off eastern Maine, about 1,045,000 pounds; off central Maine, about 2,573,000 pounds; small fishing banks in west-central part of the Gulf, about 516,000 pounds; off western Maine, about 1,861.000 pounds ; off eastern Massachusetts and off northern Cape Cod, about 7,347,000 pounds; Cape Cod out to the so-called South Channel, about 1,518,000 pounds; Georges Bank as a whole, about 3,184,000 pounds. In general, pollock are more abundant around the coastal belt of the Gulf, out about to the 75 to 80 fathom line, on the isolated fishing grounds enclosed within that depth limit, and over the offshore banks than they are over the deeper central basin of the Gulf; though some are taken there also. And this has always been one of the principal fishes to be caught with hook and line on the various small banks and ledges in the inner part of the Gulf; near Lurcher Shoal for instance; on Grand Manan Bank; on Jeffreys Ledge, and on Stellwagen Bank at the entrance to Massa- chusetts Bay, while the neighborhoods of Boon Island and of the Isles of Shoals long have been famous pollock grounds. Pollock are decidedly less plentiful on the Nantucket grounds in general (only about 56,000 pounds landed thence in 1947) and west of Cape Cod than they are either farther within the Gulf to the northward or on Georges Bank to the east- ward. But commercial quantities are caught yearly (in season) along southern New England and New York. The landings for Rhode Island, Connecticut, and New York, combined, ranged, for example, between 787,000 pounds and 883,000 pounds for the years 1930 to 1933. And though the landings ran less than one-half as great (be- tween 135,000 pounds and 452,000 pounds) for 1935 to 1947 we suspect that this decrease re- sulted from market conditions, rather than from any decrease in the numbers of pollock that are available there. A few hundreds or thousands of pounds of pollock are landed yearly 19 in New Jersey ports also. But this is the extreme southern limit for the pollock as a market fish. To the eastward and northward, we find pol- lock caught in abundance all along the outer Nova Scotian coast and banks. In 1946, for example, 840,000 pounds were landed in Guysborough County, Nova Scotia, and 277,200 pounds along the Atlantic coast of Cape Breton Island, east of the Gut of Canso.17 This, however, is the north- eastern limit of our pollock as a market fish of any importance. True, a few thousand pounds were reported yearly from the southeastern side of the Gulf of St. Lawrence near the Gut of Canso during the early nineteen hundreds.18 But the catch is so small that pollock have not been mentioned in the catch statistics for more recent years, nor any- where else within the Gulf of St. Lawrence. Small pollock, 8 to 10 inches long and weighing less than half a pound (1 or 2 years old) swarm inshore after early April, when we have seen thou- sands of them taken from the traps at Gloucester and Magnolia. In the southern part of Massa- chusetts Bay these "harbor pollock," as they are called locally, move out in June, probably to avoid the rising temperature, to return again in autumn. But they continue abundant all summer and au- umn in the harbors and bays and among the islands along the coast northward from Cape Ann and eastward to Nova Scotia. Most of them seek slightly deeper water in winter, however, probably to avoid the cold. The larger fish tend to keep farther offshore than the small ones; they live deeper on the whole, ex- cept when they are pursuing some particular feed (p. 214), and they are caught in more definite local- ities, not everywhere and anywhere along the coast as are the little fish. In the southwestern part of the Gulf, as exemplified by Massachusetts Bay and by the belt from Cape Ann to the Isles ■' U. S. catch, 492,400 pounds, 1945; Canadian catches 7,017,000 pounds in 1944 and 0,642,000 pounds iu 1946, besides an Indeterminate- amount landed along this part of the Shelburne County coast line. '< 1944, 513,000 pounds; 1946, 983.000 pounds. I' Charlotte and St. Johns Counties, about 2,000,000 pounds in 1944, about 3,507,000 in 1946. u Maximum, 10,700 pounds, minimum 600 pounds for the years 1930-1937 and 1939-1947, 101 ,200 pounds were credited to New Jersey in 1938; an amount so much larger than usual as to suggest that it was because of economic reasons that tho flsh were landed in New Jersey rather than in New York. " Richmond County, Nova Scotia, 223,600 pounds; Cape Breton County, 53,600 pounds. " Yearly catch, 1,600-4,000 pounds; for 1902 to 1906 and 1909 to 1915-1916, 61,500 pounds were credited to Inverness County in 1901, but this amount is so much larger than usual as to suggest some error. FISHES OF THE GULF OF MAINE 219 of Shoals, large pollock are taken in greatest num- ber in late autumn and early winter when the gill- net fishery taps the spawning fish (fig. 220). Few are caught there later in the winter after they finish spawning, showing that the spent fish do not winter on particular grounds, but scatter to and fro in search of food. Along this part of the coast they often reappear in abundance at the surface near land during April and May and even into June. In 1951, for ex- ample, we heard of schools of large pollock at various points off the tip of Cape Cod, in the north- ern side of Massachusetts Bay, and off the Merri- mac River dining the first week of that month. They tend to move out again, and deeper, as the surface warms with the advance of the season, and very few large ones are taken inshore in the Massachusetts Bay region during July and August. But it is not likely that they travel far, or sink very deep, for good fares of fish 2 to 3 feet long are brought in by line fishermen from Jeffreys Ledge throughout the summer, most of them caught some distance above bottom. North of the Isles of Shoals, pollock arc more commonly seen on the surface during the hot months. Thus, we remember one year (1922) when small boats from Cape Porpoise and from neighboring ports were doing well trolling during July and early August; in 1951 schools were re- ported off Baileys Island, Casco Bay, during the first week of July. And great numbers of good sized pollock are caught all summer in the tide rips at the mouth of the Bay of Fundy; in Passa- maquoddy Bay; in Digby Gut on the Nova Sco- tian side of the Bay of Fundy; and along outer Nova Scotia.18 No information is available as to the relative frequency with which pollock appear at the surface over Georges Bank and other offshore fishing grounds, though they are caught all through the year at deeper levels, with no greater seasonal fluctuation in the landings than might result from the various vicissitudes of fishermen's luck, the weather, and the market. Pollock spawn in great numbers at the mouth of Massachusetts Bay, especially on the broken bot- tom southeast of Gloucester and along the seaward (eastern) slope of Stellwagen Bank, where most of the eggs were taken during the years when pollock were hatched in great numbers at the Gloucester hatchery of the U. S. Bureau of Fisheries.20 In some years many pollock spawn (and large quantities of their eggs have been collected lor the hatchery) right up to Boston Lightship in the inner part of Massachusetts Bay, though this is not a regular annual event. And gill-netters also catch an abundance of ripe fish between Cape Ann and the Isles of Shoals, where breeding pollock congregate in such abundance that they have sup- ported a lucrative fishery in some years. This in general seems to cover the most produc- tive spawning area so far as the inner parts of the Gulf are concerned. Few spawning pollock are caught in the Gulf south of the Massachusetts Bay region, while we find no report of them as breeding anywhere west of Cape Cod, although fry of the winter's hatch appear at Woods Hole in spring (p. 220). On the other hand only a fewripe fish are seen along the coast of Maine, though the Boothbay hatchery has made diligent search for them east of Casco Bay; neither have we found pollock eggs anywhere north of the Isles of Shoals in our autumn or winter bowings. And it seems that very few larvae are hatched at the mouth of the Bay of Fundy on the New Brunswick side, for none of their young fry have been found in the Passamaquoddy-Grand Manan region, though yearlings, half-grown fish, and adults are there in great numbers. Thus it is safe to say that no production of any importance takes place any- where in the Gulf of Maine east of Cape Elizabeth. We cannot offer any explanation for this regional contrast in pollock productivity. Temperature seems not the cause, for this differs by only a couple of degrees between Massachusetts Bay and Passamaquoddy Bay at the commencement of the spawning season. And while the coastal water as a whole is slightly cooler east than west of Cape Elizabeth at the height of the season, the differ- ences from station to station have been small; and all the readings we have taken there during late December and early January have fallen well within the range at which pollock spawn freely in 19 Near Canso good-sized fish are caught on hook and line at the sur- face from June to December, according to Cornish, Contributions Canadian Biology (1902-1905) 1907, p. 189. " Information supplied by C. G. Corliss, former Superintendent of the Gloucester Hatchery. 220 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Massachusetts Bay, as appears in the following table : Water temperatures, Massachusetts Day to Lurcher Shoal, 1920-1921 Off Off Cape Off Mount Desert I., Jan. 1, sta- tion 10497 Off Fundy Off Depth Glouces- Elizabeth Machias, Deep, Lurcher in ter, Dec. Dec. 30, Jan. 4, Jan. 4, Shoal, fathoms 29, station station station station Jan. 4, sta- 10489 10494 10498 10499 tion 10500 0 42 42 40.5 42 42 42.5 10 43.7 42.5 41.4 42 42.4 42.7 20 44.4 43.1 41.8 42.1 42.6 43.1 40 44.4 44.9 42.3 42.1 42.9 43.9 75 44.6 43.5 Presumably the pollock of Georges and Browns Banks and of outer Nova Scotian waters to the eastward reproduce themselves there. But we have no definite information in this regard. A few ripe fish are caught in the Massachusetts Bay region as early as the last week in October, and the first of November to the middle of Jan- uaiy covers the most active production there, as illustrated by the following table supplied by C. G. Corliss, former Superintendent of the hatchery, where many millions of pollock eggs were once hatched yearly. Year First eggs taken Last eggs taken Eggs most plentiful Total eggs collected 1911-12 Nov. 10- Nov. l._ ..do ..do do . Jan. 22. . 499. 875, 000 1912-13 Jan. 31 856, 680, 000 1913-14 Feb. 6 974, 240, 000 1914-15 Feb. 9 855, 020, 000 1915-16 Feb. 17 1,713,730,000 1916-17 . Nov. 7. - Nov. 6.. Nov. 10. Nov. 15. Jan. 27- - Jan. 23. Jan. 16-. Jan. 21.. Nov. 16 to Jan. 20 Nov. 20 to Jan. 8 Nov. 17 to Jan. 16 Nov. 21 to Jan. 16 2, 081, 400, 000 1918-19 1,110,470,000 1019-20... 1920-21. 954. 800, 000 650, 850, 000 The first week of March is the latest that the gill netters have reported any spawning fish. The pollock spawns considerably earlier in the Gulf of Maine than in European waters, where spawning does not begin until January, is at its height in March, and continues into April. The Gulf of Maine pollock, like the cod and had- dock, spawn in comparatively shoal waters. Thus we have towed a considerable number of pollock eggs over Stellwagen Bank where the water was only 16 fathoms deep (on November 8, 1916) and most of the ripe fish that supplied the Gloucester hatchery with eggs were netted in depths of 25 to 50 fathoms. Probably few spawn deeper than 50 to 60 fathoms, and there is no evidence in egg records, in captures of ripe fish, or in fishermen's re- ports, that any pollock eggs are produced in the deep basins of the Gulf. In European waters, however, this fish is described as breeding only in depths greater than 75 fathoms. The gill netters have described it to us as spawn- ing over hard bottom chiefly, though the pollock is not a ground fish at other seasons. The migrations of the young pollock in our Gulf, from hatching until they appear on the coast as yearlings, are of special interest because of the probability that the great majority of all the pollock that frequent the eastern coast of Maine and the Bay of Fundy region are produced elsewhere. Some of them may come from spawn- ing grounds (as yet unmapped) off southern or western Nova Scotia; our own observations throw no direct light on this point. But what is known of the general circulation of the Gulf in spring and early summer suggests, rather, that the bulk of them come from the spawning grounds on the western side, south of Cape Elizabeth, having circled around first southward, then eastward and northeastward, and so finally to the Bay of Fundy and to the east part of the Maine coast. Others, hugging the coast more closely in their involuntary journeyings, may follow past Cape Cod and so westward, evidence of which is the presence of an abundance of pollock fry in spring at Woods Hole, for pollock are not known to spawn in quantity anywhere west of the Cape (p. 219). Strangely enough, we have caught no pollock less than 8 or 9 inches long on the offshore banks either on hook and line or in our tow nets, nor have we seen any that had been trawled there. Wh ether this is because the young are too nimble to be taken in trawls, whether because they live well off bottom, or whether because they are scarce offshore, is not known. The larger pollock of our Gulf seem to wander but little, for many that have been tagged by the U. S. Bureau of Fisheries have been recaptured within short distances of the localities where they were marked, and after long periods of time. And while a few of the marked fish are known to have made considerable journeys eastward, (one, for example, from Jeffreys Ledge to Sable Island), instances of this sort have not been numerous enough to suggest any mass movements. Pollock appear to be similarly stationary all along the outer Nova Scotian coast, for they are caught there throughout the fishing season. But FISHES OF THE GULF OF MAINE 221 we think it likely that the few pollock that are caught within the Gulf of St. Lawrence wander in via the Gut of Canso. On the other hand, pollock are seldom caizght west of Rhode Island after June,21 and it is chiefly as cold season visitors that they appear off the coasts of Connecticut, New York, or New Jersey; the commercial catches reported thence are made mostly in winter and in early spring. Importance. — At the time the first edition of this book appeared (1925) our Gulf was yielding about 35 million to 40 million pounds yearly. In 1946, most recent year for which we have seen the Canadian catch statistics as well as those for the United States, the total catch for the Gulf was close to 48 million pounds,22 say 5 million to 7 million fish. A quarter of a century ago, the gill net was regarded as the most effective apparatus for catching pollock; hand and long lines ranking next; otter trawls yielding only a few,23 while schools that were seen at the surface were often seined, especially the smaller sizes. But the relative proportions have been reversed with the great development of the otter trawl fishery. In 1946, for example, a representative year, a little less than % of the pollock landed from the Gulf of Maine by United States fishermen were caught in otter trawls; a little less than l{ in gill nets; with hand and long lines, traps of one sort or another, and purse seines accounting for the remainder in the order named.21 Some of our readers will be more interested in the fact that pollock will take an artificial lure and put up a strong resistance. Small ones up to 4 or 5 pounds will take a bright artificial fly freely (silver body with white wings of hackle or hair is good, especially with a touch of red). We have caught many fly casting from the rocks in autumn when smallish pollock are inshore after smelt or other small fish. And a pollock rises so fiercely to the fly and makes so long and strong a run when it is hooked that a small one gives fully as good sport as a trout caught on a light fly rod; a medium-sized pollock provides nearly as good sport as a salmon of equal weight. When the larger pollock are schooling at the surface near shore in May and June, many of them are taken by anglers trolling with spoons or with feather lures of one kind or another, from party boats out of Plymouth, Gloucester, Ipswich, Newburyport, Hampton, York, Casco Bay, and various other places along our coasts; also off Gay Head, Marthas Vineyard, and still farther to the westward. And pollock of all sizes bite eagerly on clams, minnows, or on bait of cut fish. WHITE HAKE AND SQUIRREL HAKE We are forced to discuss these two hakes to- gether, for they are so hard to tell apart that they are often confused, while they agree so closely in habits and distribution that what is said of one applies equally to the other, except as noted below. White hake Urophycis tenuis (Mitchill) 1815 25 Boston hake; Black hake; Mud hake; Hake; Ling Jordan and Evermann, 1896-1900, p. 2555. Description.- — Although the hakes of the genus Urophycis (true hakes in general parlance on this « Two pollock tagged on Nantuckeil Shoals in June and October were caught off Block Island in the following May and July respectively. 32 47,670,776 pounds, plus an indeterminate amount for Shelburne County, Nova Scotia, that may have been caught on the Gulf of Maine side of Cape Sable. " Bigelow and Welsh, Bull. U. S. Bureau of Fisheries, vol. 40, rt. 1, 1925, p. 406. " No statistics are available as to what proportions of the Canadian catch in the Gulf are made with the different kinds of pear. " The European hake is a Merluccim (p. 177). side of the Atlantic) are close relatives of the cod and of the haddock, they are not at all codlike in appearance, being more slender and softer- bodied fish, tapering backward from the shoulders to a slim caudal peduncle and to a small weak tail, witli much larger eyes than the cod hut with smaller chin barbels. Furthermore, they have only two dorsal fins, the second much longer than the first, and only one anal (in instead of the three dorsals and the two anals of the pollock, cod, and haddock. The ventral fins are long, narrow, and fcelerlike. The body of the white hake is rounded in front of the vent, flattened sidewise behind the vent, and is about five and one-half times as long as it is deep. The mouth is so large that it gapes back to below the eyes, the upper jaw projects beyond the lower, and the chin bears a small barbel. The first dorsal fin (9 to 10 rays) orig- inates close behind the pectorals, and is shorter 222 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 104. — White hake (Urophycis tenuis), Halifax, Nova Scotia. From Goode. Drawing by H. L. Todd. than the latter, triangular, with the third ray prolonged as a filament that is longer than the fin proper is high. The second dorsal fin (about 54 to 57 rays) runs the whole length of the trunk from close behind the first dorsal fin to the caudal peduncle, is of about equal height from end to end, with rounded corners, and is only about half as high as the first dorsal. The anal fin is similar in outline to the second dorsal but is shorter (about 48 to 50 rays). The pectorals are rounded when spread; the ventral fins are situated considerably in front of the pectorals, and each is reduced to two very much prolonged rays (apparently a single branched ray), with the lower (longer) ray of the two falling slightly short of the vent. The length of the ventral fins has often been given as an alternative character separating the white hake from the squirrel hake (p. 223), in which the ventrals are usually described as reaching beyond the vent. This distinction is not to be relied on, however, for we have seen squirrel hakes in which the ventrals lacked something of reaching the vent. The scales on both head and body are. smaller than those of the closely allied squirrel hake, and their number is the most reliable distinction be- tween the two species, there being about 140 oblique rows of scales along the lateral line between gill opening and base of tail fin in the white hake, but seldom, if ever, more than 110 rows of scales in the squirrel hake. Another difference, which seems equally depend- able, though it is less obvious, is that the upper jaw (maxillary) bone reaches as far back as the rear edge of the eye in the white hake, but only as far as the rear edge of the pupil in the squirrel hake. Color. — Like most bottom fish, white hake vary in color. As a rule they are muddy or purple brown above, sometimes almost slaty (we saw one of this shade caught in Northeast Harbor, Maine) , the sides sometimes bronzed, and the belly dirty white or yellowish white peppered with tiny black dots. The dorsal fins are of the same color as the back, the anal fin the same as the belly, and both the dorsals and the anal are edged with black. The ventral fins are pale, like the belly, but usually they are more tinged with yellow. Figure 105. — Side view of heads of white hake, A, and of squirrel hake, B, to show the difference between the two in length of the upper jaw bone. Size. — The maximum length is about 4 feet, the maximum weight about 40 pounds. But most FISHES OF THE GULF OF MAINE 223 of the fish that are caught weigh between 1 and 20 pounds, averaging no more than 8 pounds. A hake 28 inches long will weigh about 8% pounds if it is in good condition; one of 30 inches, about 9 pounds; 36 to 38 inches, 13 to 16 pounds; and about 18 pounds at 40 inches, according to Welsh's experience. Squirrel hake Urophycis chuss (Walbaum) 1792 Red hake; Ling Jordan and Evermann, 1896-1900, p. 2555. Description.- — The squirrel hake resembles its larger relative, the white hake (p. 221) so closely that the one is often taken for the other. The number of scales affords the most reliable means of identification, those of the squirrel being much larger relatively than those of the white, and arranged in only about 100 to 110 oblique cross rows along the side from gill opening to base of caudal fin, and in about 9 longitudinal rows on the upper part of the sides between lateral line and dorsal fin, as against about 140 transverse rows and about 12 longitudinal rows in the white hake (p. 222). Also, the upper jaw (maxillary bone) reaches back only as far as the rear edges of the pupil in the squirrel hake, but as far as the rear edge of the eye in the white hake (p. 222), and this difference can be relied upon, even for very small fish. The ventral fins of the squirrel hake overlap the vent as a rule, whereas those of the white hake fall short of it, but this is not invariably the case, as already remarked (p. 222), for we have seen squirrel hakes in which the ventrals did not reach to the vent. Furthermore, the filamentous part of the third ray of the first dorsal fin is much longer (if undamaged) in the squirrel than in the white hake, i. e., three to five times as long as the rest of the fin, and the nose is blunter. The color, too, is of some value in identifying these species, for while the squirrel hake is almost always reddish brown, the white hake has a decidedly purplish lustre when fresh caught. Color.- — The squirrel hake is reddish, muddy, or olive brown on sides and back, darkest above; sometimes almost black, sometimes more or less mottled, and sometimes plain, with pale lateral line. The lower part of its sides usually are washed with yellowish, and sometimes marked with dusky dots. Its belly and the lower parts of the sides of its head are pure white, grayish, or yellowish; its dorsal, caudal, and anal fins are of the same color as the back except that the anal is pale at the base. The ventral fins are very pale pinkish or yellowish. Size.- — The squirrel hake does not grow so large as the white hake, seldom reaching a greater length than 30 inches (the largest of 780 Bay of Fundy fish measured by Craigie was about 27 inches long) , or a greater weight than 6 to 7 pounds, and the average of those caught will not run above 1 to 3 pounds. In fact, a fish as heavy as 5 pounds is exceptional. Females are both longer and heavier than males of the same age (p. 226). Habits. — These two hakes,28 like many other sea fishes, spend their first months drifting at or near the surface, and fry of K to 4 inches (among which both species are no doubt represented) are often taken in summer under floating eelgrass or rockweed. On calm days we have seen them darting to and fro on the surface on many oc- casions (p. 224). And it is evident that the dura- tion of this pelagic stage varies, for we have towed fry as long as 4 inches on the surface although others seek the bottom while they still are only " The youngest stages of the two species are so much alike that in most cases we have been forced to list them simply as "hake," awaiting more critical examination than we have been able to give them. ^^^^y^^-^:^^^ Figure 106. — Squirrel hake {Urophycis chuss), off Marthas Vineyard. From Goode Drawing by H. L. Todd. 224 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE about 2 inches long. But it is not known how far they may journey while they are at the mercy of currents. After they have taken to the bottom, they are ground fish for the remainder of their lives, only rising into the upper layers in pursuit of food. They are rather sluggish swimmers, as their body form suggests, and even a large one makes only a very feeble resistance when it is hooked. When hake first take to bottom many of them do so in very shallow water, fry 2 to 6 inches long being common close below the tide mark among the eelgrass (Zostera); and fish a little larger are often caught by flounder fishermen in the harbors all around the Gulf of Maine. Others, however, seek the ground in somewhat deeper water, where they have an interesting habit of hiding within the living shells of the giant scallop (Pecten magellanicus) . This has often been observed on the outer part of the Continental Shelf off south- ern New England; Nichols and Breder 27 have found little hake hiding in the mantle cavities of scallops in 20 fathoms off New York, and scallop fishermen have informed us that they often find little hake in the scallops that they dredge off the coast of Maine. Both of the common species of hake are known to use this curious refuge (they do not feed on the scallops but merely use their shells as hiding places) , but most of the specimens so taken have proved to be squirrel hake. And the latter adopts this form of commensalism so commonly that Welsh records as many as 27 taken from 59 scallops in one haul of a scallop dredge, and 11 hake from 9 scallops in another haul, besides many others not counted off southern New England, New York, and New Jersey during the summer and autumn of 1913. Slightly larger hake of both species, up to 8 to 12 inches long, are not only plentiful offshore, but are rather common close inshore in a fathom or two of water, in harbors, and even well up estu- aries. The larger fish usually keep to deeper water, especially in summer, when hake of market- able sizes are most plentiful below 20 fathoms, and when only a few large ones are caught in less than 10 fathoms of water. But this rule, like most others, has its exceptions. For instance, we once saw a white hake of about 8 pounds caught from a float in Northeast Harbor, Maine, in about 27 Zoologies, N. Y. Zool. Roc, vol. 9, 1927, p. 172. 10 feet of water, in July (in 1922). On the other hand, hake of both the species in question are to be caught in the deepest parts of the Gulf, and white hake have been taken down to 545 fathoms at least, on the offshore slope of Georges Bank. Both of these hake haunt soft bottom chiefly, few being caught on the gravelly or shelly grounds that are so prolific of cod and haddock, or on rocky grounds. And it has been our experience that the whites are the more strictly mud fish of the pair. The temperatures in which hakes of different ages are found cover the entire range proper to the Gulf except perhaps the very lowest. At the one extreme many of the youngest fry that are seen swimming at the surface in the west central part of the Gulf in summer are in water as warm as 68° to 70° F., while young hake are in still higher temperatures west and south from Cape Cod if they are at the surface. And the somewhat larger fry found on our beaches a little below tide mark may be in water as warm as 60° locally. But the great majority of the hakes living deeper are in water at least as cool as 50° throughout their later lives, most of them in temperatures lower than 45° F. At the other extreme, all of the hakes living around the inner slopes of the Gulf at depths less than 50 fathoms experience temperatures as low as 35° to 37° F. in late winter and early spring; as low as 33° to 34° locally if they are living as shoal as 20 fathoms, which many of them do. But the fact that the bottom temperatures at the particular stations on the Grand Banks (all on the southern part) where white hake have been reported by the Newfoundland Fisheries Eesearch Commission have all been between about 42° and about 33° F. (5.5° C. and 0.6 C), and that they were not taken on other parts of the Bank where the bottom is colder, suggests that they tend to avoid regions where the temperature is as low as 32° F. or lower. And this finds some corrobora- tion in the report (see p. 228) that hake tend to withdraw in autumn from Passamaquoddy Bay, where the water chills at least as low as 32° at some time during some winters. Food. — Less is known of the diet of the hakes than of the cod, the haddock or the pollock. However, it is certain that they are not shell eaters to any extent, for it is seldom that their stomachs contain even the smaller univalve or FISHES OF THE GULF OF MAINE 225 bivalve mollusks, and neither large mollusks nor echinoderms have ever been found in a hake, so far as we know. The stomach contents so far recorded M show that shrimps (Pandalus), amphi- pods, and other small Crustacea which they find on the bottom are their chief dependence at most times and in most places. They also feed as greedily on squid as others of the cod tribe do, and a variety of small fish have been foimd ia hake stomachs at Woods Hole,29 such as alewives, butterfish, dinners, eels, flatfishes, tautog, her- ring, mackerel, menhaden, launce, silversides, silver hake, sculpins, sea robins, smelt, and tomcod. Small white hake trawled some 75 miles south of Martha's Vineyard, in 56 fathoms, January 29, 1950, by the dragger Eugene H had fed on small squid, crabs (Cancer) and small butterfish (Poronotus); others trawled off Chesapeake Bay (lat. 38°13' N., long. 73°49' W.) hi 52 fathoms by the Albatross II, March 2, 1931, had small mack- erel, flounders, crabs, and squid in their stomachs. And we have seen squirrel hake caught ofl' north- ern New Jersey with their bellies distended with launce, and with launce hanging from their mouths. Hake of both species bite on fish bait such as herring readily; in fact, most of those that are caught on long lines (p. 230) are hooked on pieces of herring. But they also take clams on the hook greedily enough. In the northeastern part of the Gulf of Maine hake feed far enough off bottom to capture the pelagic euphausiid sin-imps (Meganyc- tiphanes and Thysanoessa) that are so plentiful there, while the general character of their diet is sufficient evidence that the)' do not root in the ground like haddock. Ever since 1616, when Capt. John Smith 30 wrote "Hake you may have when the cod fades in summer, if you will fish in the night," it has been common knowledge that they bite best after dark, from which it is fail- to assume they do most of their foraging between sunset and sunrise. 3«Cioode, (Fish. Ind. U. S., Sect. 1. 1884. p. 235); Kendall, (Kept. U. S. Coram. Fish., (1896) 1898, p. 180); Linton, (Bull. U. S. Fish Comra., vol. 19, 1901, p. 478); Hanson, (Proc. U. 8. Nat. Mus., vol. 48, 1915, p. 94); Bredcr (Zoologica, X. Y. Zool. Soc., vol. 2. No. 15, 1922, p. 350); and Vinal Edwards' notes. 39 A large white hake taken at Woods Hole in May 1908 had a fish (Lepo- phidium) encysted in the wall of its body cavity, having no doubt penetrated the hake's stomach after it had been swallowed. (Sumner, Osburn, and Oole, Bull. U. S. Bur. of Fish., vol. 31, pt. 2, 1913, p. 768). 30 General Historie of Virginia, New England, and the Summer Isles, 1616, ed. of 1819, vol. 2, p. 188. 210941—63 16 Herrick 31 has given an interesting account of the perceptions of squirrel hake kept in a tank at Woods Hole, where they proved to have keen sight (though less so than pollock) and usually caught bits of meat before these had sunk. But it seems that it was only while food was in motion that the fish recognized it by sight, and that they depend chiefly on the sense of touch for their livelihood. They exercised this by swimming close to bottom with the sensitive tips of the ventral fins dragging the ground. When a hake touched a fragment of clam in this way it immediately snapped it up, but not otherwise. And they paid no attention whatever to live clams in their shells, though they often brushed over them. These observations, applied to the conditions under which hake actually live, suggests that they rec- ognize shrimps, crabs, and other foods by their ventral feelers, and that they snap up their victims as these dart ahead, when the feelers drag over them. No doubt the eggs of the white hake are bouyant like those of the squirrel hake (p. 225), but few wholly ripe females, no eggs naturally spawned, or young larvae have been seen yet. We were equally ignorant of the spawning and early stages of the squirrel hake up to the summer of 1912. But we trawled squirrel hake with running spawn and milt in Ipswich Bay in that July, fertilized the eggs on board the Grampus, and thus identified the eggs. Since then large numbers of squirrel-hake eggs have been hatched arti- ficiallv at the Gloucester hatchery. Figure 107. — Squirrel hake (Uropkycis chuss), eggs, after 1 hour's incubation, A; and after 74 hours' incuba- tion, B. The eggs are buoyant, spherical, transparent, and 0.72 to 0.76 mm. in diameter. When first spawned they have variable numbers of small » Bull. U. S. Fish Comm. .vol.22 ,1904. p. 258. 226 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE colorless oil globules 0.02 to 0.07 mm. in diameter, scattered over the yolk, but most of these globules unite shortly after fertilization into one large one of 0.15 to 0.17 mm., which is sometimes single but which usually has two or three tiny ones close beside it. The embryo extends half way around the yolk sphere within two days after fertilization (at a temperature of 60° F.), and pigment has appeared, one of the most characteristic features of this species being the development of black chromatophores not only on the embryo, but over the yolk, and finally on the oil globule as well. In late stages of incubation this feature, combined with the small size of the egg and (usually) with a multiple oil globule, distinguishes eggs of the squirrel hake from all other buoyant fish eggs of known parentage that have been found in the Gulf, except for any rockling eggs that may have pigmented oil globules (p. 236). There is also some danger of confusing newly spawned eggs of the squirrel hake with those of the butterfish (of about the same diameter) for these sometimes have two oil globules (p. 364). The newly hatched larvae have not been de- scribed. Older fry (identity established either as white hake or squirrel hake by comparison with young fry that have been reared in the hatchery by Louella E. Cable) already show the long, slender ventral fins, the short first dorsal but long second dorsal, and the tapering body form, char- acteristic of the adults. These little hakes, green- ish blue on the back, with silvery sides, are sep- arable from rockling fry by their more slender form, and by their scattered pigment. Older stages are separable from rocklings by their two well developed dorsal fins, while their silvery sides mark them at a glance from the dull colored fry of the cusk.32 Rate of growth. — The rate of growth during the first few months cannot be stated until many more young fry have been measured and identified as the one species or as the other. It is probable that two year classes are represented among the fry that are caught along shore in summer. Some of the smaller ones (2 to 3 inches long) may be from the earliest spawned eggs of that same season, but other squirrel hake of 2% to 4% inches (60-1 10 mm.) 31 Fry figured by A. Agassiz (Proc. Amer. Acad. Arts, Sci., N. Ser., vol. 9, 1882, pi. 7, fig. 6; pi. 8, figs. 1-3), as Molella argentata undoubtedly were either white hake or squirrel hake. But the simple post anal pigment band, short, stocky bodies, and fan-like ventrals of the younger stages pictured by him under this same name (pi. 7, figs. 1-4) suggest that they were rockling. that were seined at Provincetown, on June 26, 1925, must have been about one year old, as they were taken too early in the season to have been the product of that year's spawning. And the larger ones of 6 or 7 mches are yearlings. D Figure 108. — Young stages of either white hake or squirrel hake. A, larva, 2.2 mm.; B, larva, 6.2 mm.; C, larva, 9 mm. ; D, young fry, 40 mm. silvery still, and living at the surface of the water. Specimens collected off Woods Hole. The growth of older squirrel hake in the Bay of Fundy has been studied by Craigie; his conclusions from scale studies,33 combined with the relative abundance of fish of different size groups, being as follows. : Average length, inches Age Male Female 1 year-old 8 8 2 years-old 13 14 3 years-old 16 19 The indicated rate of growth is so uniform during the first three years of life as to suggest that spawning (an event so exhausting that it « Contrib. Canadian Biol., (1914-1915) 1916, p. 87. Unfortunately, hake scales do not show the yearly growth zones as clearly as cod and haddock scales do. FISHES OF THE GULF OF MAINE 227 slows the growth of any fish) probably does not take place until they have passed their third birthday. Nothing definite is known of the rate of growth of the white hake, but it is fair to assume that it grows faster than the squirrel, to attain its greater length and weight. General range. — Both the white hake and the squirrel hake are exclusively American, occurring in continental waters from the Gulf of St. Lawrence and the southern part of the Grand Bank of Newfoundland southward to the Middle Atlantic States. The squirrel, though common as far south as Chesapeake Bay, has not been reported from farther south than Virginia. But the white hake is known off North Carolina (we have seen a 30 inch specimen that was trawled off Bodic I., North Carolina, lat. 35°52' N., long. 74°51' W. in 70 fathoms by the Albatross II, Feb. 24, 1931). And very likely the "squirrel" actually ranges as far south as the "white" does. Both of them occur from near tide mark, the squirrel down to about 175 fathoms, the white down to about 545 fathoms. Occurrence in the Gulf of Maine. — Hake are very common fish in our Gulf, where the two species, white and squirrel, are caught side by side regularly. In the Bay of Fundy there are so few toward the head that stragglers are caught, or none at all, but they are plentiful enough toward the mouth where, for example, about 6,400,000 pounds were landed on the Nova Scotian side by Canadian fishermen in 1944, and about 8,200,000 pounds in 1946, while the yearly catch on the New Brunswick side is about 500,000 to 600,000 pounds. Other centers of abundance for them inshore are along the coast of Maine between Machias and Mount Desert Island, in Frenchman's Bay (formerly the site of an important hake fishery), the ground known locally as the "Grumpy" near Isle au Haut, and off Penobscot Bay. Sundry small grounds outside the islands from Penobscot Bay to Cape Elizabeth and all along the western side of the Gulf, also yield good numbers of hakes, especially near Boon Island; the vicinity of the Isles of Shoals, a famous hake ground for small boat fishermen; Ipswich Bay; the lower slopes of Jeffreys and Stellwagen Banks; also the deeper parts of Massachusetts Bay, which yielded 750,000 pounds in 1919 when the demand for hake was better than it is now. Hake, indeed, are so widespread on the lower slopes of all the banks and ledges in the inner parts of the Gulf, as well as on the mud floors between them, that Rich34 listed 119 named grounds in the western side of the Gulf as good haking bottoms. Hake, with flounders, rosefish, and silver hake are practically the only commer- cially valuable fish one is likely to catch on the floors of the deep basins and channels of the Gulf; and a catch of 2,880 of them with 580 cusk, but no cod or haddock, by long-line fishing 15 miles southeast of Monhegan on June 24 to 25, 1913, will illustrate how completely they may monop- olize suitable bottoms. Hake are plentiful in the so-called South Chan- nel also, and on the northwest slope of Georges Bank, whence about 2,000,000 pounds were landed in 1919, about 1,500,000 pounds in 1947. And it has long been known that there is an abundance of hake at depths greater than 60 to 70 fathoms all along the southern slope of Georges Bank. Long-line fishermen, too,' have told us that while it was unusual to hook a hake on the shoaler parts of Georges, man}' were caught wherever the line was run off into deeper water on the northwest face of the bank; i. e., onto soft bottom. And this is borne out by the statistics of the catches, for the good trawling grounds on Georges Bank yield far fewer hake of marketable size than the inner parts of the Gulf do, if the year 1945 can be taken as representative.85 It has only been since 1944 that the landings of white hake and of red (i. e., squirrel) hake have been reported separately. Taken at their face value, these would point to the white hake as by far the more plentiful member of the pair throughout the inner parts of the Gulf as a whole, and on Georges Bank. In 1945, for example,38 the reported landings were some 14 times as great for white as for red hake, for every one of the subdivisions into which the inner part of the Gulf is divided for statistical purposes, while only a few thousand pounds of red hake were reported from off eastern or central Maine, or from the northwestern part of Georges Bank; and none from the eastern or southwestern part of the " Rept. U. S. Comm., Fish. (1929) 1930, App. 3, pp. 85-86, 96. « Landings of hake In 1945 were about 414,000 pounds lor Georges Bank; about 12,700,000 pounds for the Inner parts of the Gulf by United States fishermen and about 9,140,000 pounds by Canadian fishermen. >• The only year when the landings of the two have been reported by counties for Maine and Massachusetts, besides the landings at the major ports. 228 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Bank. But this would give a wholly false pic- ture of the actual situation, because most of the red hake that are caught on these grounds are thrown overboard because they are too small to be worth gutting and icing under present market conditions. Actually, we trawled 34 squirrel hake and only two other hake37 in Ipswich Bay, in 22 fathoms on one occasion in July 1912; Welsh counted 5,450 squirrel hake to 652 white hake caught in otter trawling on the northwest slope of Georges Bank in June 1912; we counted 2,457 red hake to only 196 white hake from 29 trawl hauls at 22 to 150 fathoms on the southwestern part of Georges Bank and south of Nantucket Shoals on the Albatross III, May 11-18, 1950; and we saw 2,040 squirrel hake taken in 42 hauls by the Eugene H on the southwest part of Georges Bank, in late June 1951, but only 51 white hake. Re- ported landings also, in pounds, for 1945, were about 100 times as great for red as for white hake from the Nantucket grounds, whence all the little hake are brought in for the trash fish industry. And the discrepancy is greater still in numbers, for the white hake are much the heavier of the two, individually. Red hake also predominate over white among the hake landed in New York and to the southward, as is illustrated by the catch statistics for 1947. 38 Landings, for 1947, to nearest 100,000 pounds: New York New Jersey Delaware Red hake 1,200,000 5,600,000 200,000 White hake 1,000,000 200,000 100 On the other hand, inquiries of fishermen, corroborated by our own experience, point to the white hake as the more plentiful of the two in the basin of our Gidf at depths greater than 40 to 50 fathoms. The Atlantis, for example, trawled about 700 white hake in the deep basins off Cape Cod, west of Jeffreys Ledge and off Mount Desert, in August 1936, but only a scattering of squirrel hake. Tl»is appears to apply equally to the deeper holes in Massachusetts Bay at depths greater than 30 fathoms or so (both Storer and Goode and Bean spoke of the "white" as the more common of the two there), also to the Bay of 87 The latter were listed by Welsh as V. rtgius, but probably they were white hake. B About 13,000 pounds of white hake were reported [rom Maryland In 1947, about 65,000 pounds from Virginia, and about 4,000 pounds from North Carolina, with no reds. But we suspect that reds were actually Included as well as whites, and spotted hake also. Fundy region in general, Including Passama- quoddy Bay, according to Huntsman. And nearly all of the hakes that have been listed by name from the more easterly of the Nova Scotian Banks, or from the southern part of the Grand Banks in the annual reports of the Newfoundland Department of Natural Resources, have been the white (tenuis). Tenuis, also, is the only member of the pair that was reported by Cornish 39 from Canso, but chuss alone is recorded from the Cape Breton shore of the Gulf of St. Lawrence by Cox,40 who also records one from 205 fathoms on the Nova Scotian side of Cabot Strait. The situation is made more confusing by Corn- ish's report of hake with 123 rows of scales from Prince Edward Island, and with 130 rows of scales from Canso, fish intermediate, that is, be- tween chuss and tenuis in this regard, though favoring the latter. Perhaps the separation between the two species in number of scales, and also in other features, may not be so sharp toward the northern boundaries of their geographic ranges as we have found it farther south. A more or less regular inshore movement of hakes of one or the other species, or of both of them, is said to take place in autumn, especially in the northeastern part of the Umlf , made evident by capture of considerable numbers in winter in the deeper, muddy harbors and bays east of Penobscot Bay, including St. John Harbor, and Kennebecasis Bay, which they reach by running up the St. John River, and where they are caught all winter through the ice. They also carry out correspond- ing movements in and offshore off southern New England, with goodly numbers appearing in shoal water at Woods Hole in autumn. But it is only in the spring and autumn that they are found close inshore off New York and off New Jersey. On the other hand, they are said to enter Passama- quoddy Bay in early summer, to withdraw in autumn. Probably the explanation is that the adults, being cool water fish, are barred from the shallows in summer by high temperature along the coasts of Massachusetts and of west-central Maine, but that the low summer temperature of Passama- quoddy Bay allows large hake to summer there, as well as small. Their reported withdrawal from » Contrib. Canadian Biol. (1902-1905), 1907, p. 89. " Contrib. Canadian Biol. (1918-1920), 1921, p. 113. FISHES OF THE GULF OF MAINE 229 Passamaquoddy Bay in autumn may be in avoid- ance of extreme winter chilling. But we should remind the reader that failure to catch fish on hook and line in the cold season of the 3Tear (it is in this way that hakes are caught in the Passamaquoddy region) does not necessarily mean that they have departed. The hake may have stopped biting, as every fisherman knows by experience. The evidence of otter trawl catches is much more reliable in this respect, for ground fishes in general. Except for in and offshore movements, hake are resident throughout the year in the open Gulf of Maine wherever they are found, once they have taken to the bottom. And they appear to be much more stationary than either cod or haddock. The localities where we have found eggs, pro- visionally identified as squirrel hake (fig. 109), show that it spawns all around the Gulf from Cape Cod to Nova Scotia. And despite its rather deep- water habitat and preference for soft bottom, most of these egg stations have been in shoal water near the coast; a haul in the eastern basin which yielded both squirrel hake and silver hake eggs (p. 178) has been the only exception. This, of course, points to a movement from the basins into shoaler water for spawning. It seems that the white hake spawns from late winter through spring to late summer, for we saw a 71* 70* ee' 68' «7* 66* . ., . . , . -^- — . ... vjtl/l- i . V) /V-f^ ^>a (/^7 44 /bay>-» ^3r /"/nova /> ? ^> a • y^Cy \ SCOTIA _y*A $f"V r3^ (Yarmouth Portland C ▲ J " '-'■-• v ! \l\ CAPE \ Ca ' A a JA V^° 4 V ♦/ The U. S. National Museum has specimens taken ofl Charleston, S. C, at 87 and 124 fathoms. »' Nichols and Breder, Zoologica, N. Y. Zool. Soc., vol. 9, 1927, p. 169. " Fauna boreali Americana, vol. 3, 1836, p. 253. Richardson's wood cut of the specimen in question, from a sketch by Lt. Col. Hamilton Smith, shows the low first dorsal with black apex that is characteristic of the species regius. » Report. Newfoundland Fishery Res. Comm., vol. 1, No. 4, 1932, p. 109. « This species was also listed from Ipswich Bay, from Casco Bay, and off ofMonhegen Island in the Crampus collections of 1912 (Bull. Mus. Comp. Zool. vol. 58, No. 2, 1914, p. 113), but it is probable that these specimens were white hake in reality. 232 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE the southwestern part of the Gulf every year, for 49 spotted hake were taken south of Block Island, in 47 to 67 fathoms, January 27 to Feb- ruary 3, 1950, by the dragger Eugene H. Long-finned hake Urophycis chesteri (Goode and Bean) 1878 Jordan and Evermann, 1896-1900, p. 2556. Description. — The most distinctive character of the long-fumed hake is its very long ventral fins, the longest of the rays of which reach back nearly to the rear end of the anal fin (about to its for- tieth ray), with the next longest ventral ray considerably overlapping the origin of the anal. The filamentous dorsal ray is longer also, than in the other Gulf of Maine hakes, reaching back to about the middle of the second dorsal fin or beyond. Furthermore, there are only about 90 rows of scales from gill opening to caudal fin along the lateral line, and the scales are rela- tively larger than in either the white hake or the squirrel hake; the eye is larger in the long- finned hake; the anal fin rays are more numerous (average about 56) , the rear corners of the dorsal and anal fins are more rounded. The outline of the anal is slightly concave instead of straight (fig. Ill); the pectoral fins are more slender and more pointed, and the caudal fin is narrower with more strongly convex margin; these differences are more clearly shown in the illustrations than verbally. The skin of the long-finned hake is curiously loose, like that of many deep-sea fishes. Color. — Freshly preserved specimens are olive above and on the sides, with a silvery white belly. The fins are olive, with dusky markings on the dorsal filament, on the outer edge of the dorsal fins, on the caudal fin, and on the ventrals. Size.— Specimens 14 to 15 inches (36 to 38 cm.) long, trawled by Albatross III, on the south- western slope of Georges Bank and off Nantucket Shoals, in 105 to 240 fathoms, May 11-18, 1950, are the largest yet recorded. Habite. — The long-finned hake is a bottom fish, living chiefly between 100 and 500 fathoms, the deepest record for it is from 538 fathoms. It is a summer and autumn spawner, judging from the fact that Goode and Bean saw specimens in breeding condition at that season. We have taken pelagic young of 8 to 35 mm. in our tows off Marthas Vineyard during the last week of August.65 And captures of 3 fry, about 2% to 2% inches (57-71 mm.) long on April 26 (1931) and of 16 fish of about 3 to 4% inches (74-1 10 mm.) late that July suggests that a length of 4 to 5 inches is reached at 1 year of age. General range. — This is a deep-water fish, occurring in great abundance on the continental slope off North America from the Laurentian Channel in Cabot Strait to abreast of Cape Lookout, N. C. Occurrence in the Gulf oj Maine. — This hake is plentiful all along the seaward slopes of Browns Bank, of Georges Bank and of Nantucket Shoals at depths greater than 100 fathoms, where it has been trawled at many stations.46 The Albatross III, for example, caught 861 in 63 half-hour trawl hauls, at 105 to 240 fathoms on the slope between the south-central part of Georges Bank (long. 67°14' W.) and the offing of the eastern end of Long Island, New York (long. 71° 57' W.) in May 1950. Up to 1931 the only » Bigelow, Bull. Mus. Comp. Zool., vol. 59, No. 8, 1917, p. 275. " For early locality records see Ooode and Bean, Smithsonian Contrib. Knowl., vol.30, p. 361. Figure 111.— Long-finned hake (Urophycis chesteri), off Cape Ann. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 233 definite records of them from the inner parts of the Gulf had been of three specimens taken off Cape Ann," in 1 10 to 140 fathoms, in 1878; and of a few others that were trawled on the northern edge of Georges Bank by the Kingfisher, hi September 1929, in 85 to 100 fathoms. But captures of a number to the westward along the Bank and in the central basin of the Gulf in the summer of 1931, by the Albatross II 68 at depths of 70 to 140 fath- oms, show that long-finned hakes are more numer- ous in the deeper parts of the Gulf than had been suspected previously. Blue hake Antimora rostrata Giinther 1878 Jordan and Kvermann, 1896-1900, p. 2544, as A. viola Goode and Bean. Description. — This species resembles the white, squirrel and spotted hakes in the form of its body and hi having two separate dorsal fins, the first very short and the second very long; but it is readily distinguished from them by the fact that its anal fin is so deeply notched about midway of its length that it almost seems to have two sepa- rate anals, and that each of its ventral fins is 6-rayed, with the second ray prolonged and fila- mentous. The shape of the snout, which is flattened above, keeled at the sides, and blunted at the tip hi some but forming an acute angle in others is distinctive, likewise its vent is situated much farther back than hi the true hakes (genus Urophycis), and its body, in life, is deep violet, blackish brown, or blue black, below as well as above. 87 These were the basis of Goode and Bean's original descrlptlOD Of the species (Proc. U. 8. Nat. Mus., vol. 1, 1878, p. 256). '» Reported by Bigelow and Schroeder, Bull. U. S. Bur. Pish., vol. 48, 1936, p. 339. Size. — The longest yet measured was one of about 2V/2 inches (545 mm.). Range. — The blue hake was reported at so many localities along the continental slope from the early cruises of the U. S. Bureau of Fisheries Ma on the Nova Scotian slope ; off southern New England ; and southward to the offing of Cape Hatteras, North Carolina, at 350 to 1,000 fathoms that it must be one of the more plentiful of fishes there. We have recently trawled a few at 220 to 460 fathoms, on the southeastern Nova Scotian slope, on the Caryn of the Woods Hole Oceanographic Institution, between longitudes 64° W. and 65° 56' W., and halibut fishermen have occasionally brought them in. But the blue hake has not been taken within the limits of the Gulf of Maine, and it is hardly to be expected there; the shoalest cap- ture recorded for it so far is from 220 fathoms. We mention it on the chance that vessels trawling on the slope may occasionally work deep enough to pick up a few. The known range of this deep sea hake includes the North Atlantic from Denmark Strait to the offing of Gibralter in the east and from the New- foundland Banks to the offing of Cape Hatteras in the west; Uruguay; the eastern Pacific, British Columbia to Panama, and the southern Indian Ocean. It has been taken as deep as 1,456 fathoms. m Hakeling Physiculus Jvlvus Bean 1884 Jordan and Evermann, 1896-1900, p. 2547. Description. — This fish is hakelikc in its general appearance, also in the general arrangement of its "•See Goode and Bean (Smithsonian Oontrlb. Know!., vol. 30, 1895, pp. 374-375) for list of stations. '• For a recent summary, see Schroeder, Copeia, 1940, No. 4, pp. 236-237. t***^r^?*^ Figure 112. — Blue hake (Antimora rostrata). La Have Bank. From Goode. Drawing by H. L. Todd. 234 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Wi ■f^^^^^^^ *&*£mwfiuijiiizt,- Figure 113. — Hakeling (Physiculus fulvus), outer edge of Continental Shelf off Nantucket. From Goode and Bean. Drawing by H. L. Todd. fins, for it has two dorsals, the first (10 rays) triangular and much shorter than the second (about 49 rays) which is of nearly uniform height from end to end ; one long anal fin (about 54 rays) which is similar to the second dorsal in shape; and ventral fins situated in front of the pectorals. It is separable from the white, squirrel, and long- finned hakes (genus Urophycis, pp. 221 and 232) in that its anal fin originates in front of the origin of the second dorsal fin instead of considerably behind the latter and that its ventral fins have 5 rays each instead of 2 and are much shorter than those of the true hakes, with the longest ray (the second, which is filamentous at the tip) hardly reaching back as far as the middle of the pectoral fins. Furthermore, the snout of the hakeling is blunter than that of any true hake; its caudal fin much smaller; its body tapers more abruptly; and none of the rays of its first dorsal fin are pro- longed. Color. — Described as light yellowish brown with the lower surface of the head, the abdomen, and the margins of the dorsal and anal fins very dark brown, and with a dark brown blotch on each cheek (on the subopercular bone). We have not seen it fresh from the water. Size. — The maximum size is not known. Habits. — Nothing is known of the habits of the hakeling except that it is a deep-water fish, having been taken from 79 fathoms down to 955 fathoms, where it lives on or near the bottom, to judge from its general structure. General range and occurrence in the Gulf of Maine. — This hakeling has been taken at several localities in the Gulf of Mexico and on the con- tinental slope off the eastern United States. The most northerly record for it is off Nantucket flat. 40° 01' N., long. 69° 56' W.) in 79 fathoms, and it is on this record that the hakeling is men- tioned here.60 Four-bearded rockling Enchelyopus cimbrius (Linnaeus) 1766 Rockling Jordan and Evermann, 1896-1900, p. 2560. Description. — The rocklings, of which this is the only common local representative, differ from their near relatives, tUe hakes (genus Urophycis), in the facts that their ventral fins are short, with 5 to 7 rays, and that the first section of their dorsal fin consists of only one ray, which is nearly as long as the head, and which stands over the upper corners of the gill openings, followed by a series of about 50 very short, separate, hairlike rays without connecting membrane, which can be laid down in a groove on tbe back. Thus there is only one well-developed dorsal fin. Rocklings differ fur- ther from all other gadoids in the presence of long barbels on the top of the nose as well as on the chin, the number of these being the most obvious specific character among the several species of rocklings. In the present species there are a pair of these barbels close in front of the nostrils, a third and somewhat shorter barbel standing alone on the tip of the snout, and there is a fourth barbel hanging from the chin. Rocklings remind one of young hake in their slender bodies tapering back from the shoulders; *> Another small hakeling (Lotella maiillaris) has been taken orl Marthas Vineyard. It is separable from the hakeling described above by the fact that its anal fin originates behind the origin of the second dorsal fin, and by its larger teeth. FISHES OF THE GULF OF MAINE 235 Figure 114. — Rockling (Enchelyopus cimbrius). A, adult, Bay of Chaleur, from Jordan and Evermann, drawing by H. L. Todd; B, egg; C, larva (European), 3.6 mm.; D, larva (European), 5.3 mm.; E, larva (European), 13.6 mm.; F, silvery fry (European), 17.5 mm. B, after Battle; C, after Ehrenbaum and Strodtman; D, after Brook; E, after Ehrenbaum; F, after Brook. and (hakelike) they are rounded in front of the vent but flattened sidewise behind it. Their upper jaw is longer than the lower and their teeth are smaller than in the hakes, while their noses are shorter and blunter; their eyes are smaller, and the dorsal profile of their heads is more rounded than it is in any of the hakes. The pectorals are rounded and the narrow pointed ventrals are sit- uated well in front of the latter. The second dorsal fin (45 to 53 rays) originates over the mid length of the pectorals, runs back nearly to the base of the caudal fin, and is equally high from end to end with a rounded rear corner. The anal fin is similar to the second dorsal in shape, but it is shorter (39 to 43 rays).61 The caudal fin is oval when it is spread. Color. — The color of this rockling is compara- tively constant by all accounts and this is cor- roborated by our own experience. Its back is •i Storer credits It with 48 rays, but subsequent students have not found so many. dark yellowish olive or dusky brown, its sides are paler, and its belly is white dotted with brown. On some individuals the sides behind the vent are more or less clouded with a darker shade of the general body hue. The first dorsal ray, the posterior edges of the second dorsal fin and of the anal fin, the lower half of the caudal fin, and the pectoral fins are sooty or bluish black. Other- wise the vertical fins are grayish or bluish brown. The ventral fins are pale, and the lining of the mouth is dark purplish or bluish. Size. — This rockling has been described as growing to a length of 16% inches in Scandinavian waters, but about 12 inches is the longest recorded from the Gulf of Maine, where they average only about 6 to 10 inches. Habits. — Rocklings are bottom fish, like hake. Occasionally they have been found in very shallow water, on Nahant Beach in Massachusetts Bay, for example; in water only a few feet deep at Woods Hole; in 6 to 7 fathoms, both in St. Mary's 236 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Bay, Nova Scotia, and in Buzzards Bay on the south coast of Massachusetts. But they appear to be more plentiful in depths of 25 to 30 fathoms or more; there are rocklings in the deep gully off Halifax, and also in the deep trough of the Gulf of St. Lawrence.62 They have been taken on the continental slope off southern New England to a depth of 724 fathoms.63 And there is no reason to suppose that the adult fish ever rise far above the bottom, unless by accident. The occasional appearance of adult rockling in very shallow water in winter near Woods Hole M suggests that some may work inshore and into shoal water in autumn, to work offshore again and deeper in spring, for the summer. Beyond this they seem to be year-round residents wherever they are found. The name "rockling" is a misnomer for this fish for it is found most often on soft bottom in the Bay of Fundy, while those that we have trawled in Massachusetts Bay and in Ipswich Bay from the Grampus were on smooth muddy sand be- tween the hard patches. And most of the rock- ling living in the deep sinks and channels in the western side of our Gulf, and on the continental slope, are on soft smooth ground. Judging from the stomach contents of Scandi- navian and British fish (their stomach contents have not been examined on this side of the water so far as we know) they feed chiefly on shrimps, isopods, and other small crustaceans, less often on fish fry. On the other hand, rockling have been found in cod stomachs in Massachusetts Bay, and no doubt all fish of prey devour them on occasion. The eggs are buoyant, described (we have never seen them) as 0.66 to 0.98 mm. in diameter. When newly spawned the oil is in small droplets, most of which soon coalesce into one globule of 0.14 to 0.25 mm., often with one or two smaller ones close to it. The danger of confusing them with squirrel-hake eggs is discussed in the account of that fish (p. 226). And Battle has found that « Huntsman (Trans. Roy. Soc. Canada, 3, vol. 12, Sect. 4, 1918, p. 631 and further information contributed by him. '» Qoode and Bean, (Smithsonian Contrib. Knowl., vol. 30, 1895, pp. 384-385) give a long list of locality records for the rockling on the shelf and slope between the offings of eastern Nova Scotia and of North Carolina (lat. 35°40' N.). « Sumner, Osburn, and Cole, Bull. U. S. Bur. Fish., vol. 31, Pt. 2. 1913, p. 771. they develop normally at temperatures ranging from 55° to 660.66 Newly hatched larvae are a little more than 2 mm. long. The yolk is absorbed at about 3.6 mm. and the later larval stages, up to about 10 mm. long, are characterized by the very large black ventral fins shown in the illustrations (fig. 114); by the presence of oidy one post anal band of black pigment; and by the short stocky body- form. Young hake are more slender and have scattered pigment; young eusk have two post anal bands; and all other Gulf of Maine gadoids have short ventral fins. After the rockling is 17 to 20 mm. long the structure of the first dorsal fin serves to identify it. These larger fry are silvery, awaiting their descent to bottom before assuming the dull colors of the adult. In British waters they are some- times called "mackerel midges" because they sug- gest little mackerel remotely, in their general appearance. In European waters, where there are more plentiful populations of the silvery fry of one or the other species of rockling they are often cast ashore. And one such instance is described for our Gulf by Storer 66 who writes that many were picked up on Nahant Beach during one tide in the summer of 1860; and others found in the surf at West Beach, Beverly.67 Rockling fry, like those of other gadoids, drift at the surface for then first few months. How long they do so in our waters is not known, but analogy with cod, haddock, and other species suggests three months at most. And it may be assumed they seek the bottom at a length of about 2 inches for our largest pelagic fiy were 40 to 45 mm. long. During this pelagic stage they drift with the current like any other fish fry, and are at the mercy of mackerel and other fish. But they are not plentiful enough in the Gulf of Maine to be as important an article in the diet of the mackerel as the fry of the far commoner European ■' Battle (Contrib. Canadian Biol., X. Ser., vol. 5, No. 6, 1930) has made a careful study of the effects of extreme temperatures and salinities on the development of the eggs of the rockling. •« Fishes of Massachusetts, 1867, p. 279. •' These fry, and one recorded at Nahant earlier by Gill (Proc. Acad. Nat . Sci., Philadelphia, (1863)1864, p. 241) were reported as an Arctic 3-bearded species (QaidrnpsaTus argentatus Reinhardt) which was described originally from Greenland and which has been found widely distributed in Denmark Strait; on the north coast of Iceland; and in the Norwegian Sea from the Faroes north to Bear Island. But there is no reason to suppose that the Nahant specimens were anything other than the fry of our common four bearded rockling. For a recent account and discussion of the species awn- talvs, with excellent illustrations, see Jensen, Spolia Zool. Mas. TTauniensis, Copenhagen, vol. 9. 1948, pp. 167-173, pi. 4. fig. 4. FISHES OF THE GULF OF MAINE 237 rocklings are, on the other side of the Atlantic. Nothing is known of their subsequent rate of growth. General range. — Both sides of the North Atlantic. The American range is from the northern part of the Gulf of St. Lawrence and the northeastern coast of Newfoundland (perhaps even farther north) to Narragansett Bay and Long Island Sound in coastal waters, and to the latitude of Cape Fear (N. C.) in deep water along the con- tinental slope.68 The Arctic three-bearded rock- ling (Gaidropsarus ensis Reinhardt), ottierwise known only from Greenland, has been trawled on the lower part of the continental slope in the offings of southeastern Nova Scotia, of Cape Cod, of Martha's Vineyard, of New York and of New Jersey at depths of 858 to 1106 fathoms, by the Fish Hawk and Albatross I, but this is not shoal enough to bring it within our limits.69 There are several other species of rockling in north European waters, but none of them have been recorded from our side of the Atlantic. Occurrence in the Gulf of Maine. — The little rockling is of no commercial value, and it seldom comes up into very shallow water where it would force itself on the notice of seaside visitors. But it is a common bottom fish in the deeper parts of Massachusetts Bay as Goode and Bean 70 remarked long ago, while our experience, corroborated by Huntsman for the Bay of Fundy, is that (his ap- plies to the entire Gulf. Definite Gulf of Maine records for adult rocklings are from St. Mary Bay (Nova Scotia) ; various localities in the Bay of Fundy including Passamaquoddy Bay; Jones- port; off Mount Desert; off Pemaquid; near Seguin Island; mouth of Casco Bay; the deep gully to the westward of Jeffreys Ledge; Ipswich Bay; Gloucester; Nahant; various stations in the deeper parts of Massachusetts Bay; Prov- incetown; the deep open basins of the Gulf;71 and Georges Bank. And we have taken its young fry rather frequently in our tow nets in season. 89 A specimen trawled by the Albatross 11 in 12 fathoms off the mouth of Chesapeake Bay on February 10, 1030, is the only one recorded in shallow water so far southward. •• Goode and Bean (Smithsonian Contrib. Know]., vol. 30, 1895, p. 381) give a list of these localities. For a recent account of O. ensis. with illus- trations, and list of Greenland localities, see Jensen, Spolia Zool., Mus. Hauniensis, Copenhagen, vol. 9, 1948, p. 167, pi. 4, fig. a. '« Bull. Essex Inst., vol. 11, 1879, p. 9. " The Atlantis trawled it both In the Jeffrey bowl, and in the open basin of the Gulf, August 1936; and we trawled it In the central basin in July 1931. Huntsman72 and Battle " have found the eggs of this rockling in Passamaquoddy Bay through- out the summer, commencing in May and most abundantly at the time the bottom water warms to 9° or 10° C. And its breeding season probably continues from spring to early autumn in the western Atlantic as it does in the eastern,74 for Dannevig 75 (1919) records rockling eggs (prob- ably this species) as earhr as the end of May near Halifax, while we have taken rockling larvae only 5.5 mm. long as late as September and October in our tow nets in Massachusetts Bay. It is probable that the rockling spawns all around the peripheral belt of the Gulf, with Massachusetts Bay as an important nursery, to judge from our repeated captures of its larvae there. And we have taken the pelagic fry in our tow nets at the various localities marked on the accompanying chart (fig. 109) from the first week in July until October; seldom, however, more than half a dozen in any one haul (the largest catch was 18 specimens). Huntsman, similarly, describes the fry as common in the center of the Bay of Fundy, and they have been taken in the tow nets at Woods Hole in April. But we have taken neither the eggs, the larvae, nor the pelagic fry in any of our tow nettings in the central parts of the Gulf, which perhaps justifies the assumption that the spawning grounds of the rockling within our Gulf are limited mostly to depths less than 75 fathoms, though it may spawn much deeper than that on the continental slope. To the west of Cape Cod, the rockling is now known to occur in coastal waters as far as Nar- ragansett Bay, and in Long Island Sound, where it was found generally in 5% to 9 fathoms, and abundantly at 21 fathoms by the Fish Hav)k in the summer of 1914.76 And it has been trawled by the Fish Hawk and by the Albatross I at many stations in deeper water offshore along the shelf and slope, southward to the offing of Cape Hat- teras (lat. 35° 40' N.).77 " Contrib. Canadian Biol. (1921) 1922, p. 69. " Contrib. Canadian Biol., Fish., N. Ser., vol. 5, No. 6, 1930, p. 13 [119]. " It spawns from the end of January until August in the Baltic. '• Canadian Fisheries Exped., (1914-1915) 1919, p. 53, table 1C. » Nichols and Breder, Zoologlca, N. Y. Zool. Soc, vol. 9, 1927, p. 172. " For list of early stations, see Goode and Bean (Smithsonian Contrib. Know]., vol. 30, 1895, pp. 384-385). They also report a specimen apparently of this species from the offlng of Cape Fear, N. C. (lat. 34° 01' N., long. 76° 11' W.). But it was in poor condition, hence of doubtful identity. 238 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Eastward and northward from our limits, the rockling is said to be rather common in Nova Scotia waters in general, coastwise as well as on the fishing banks. The Albatross trawled it at three stations along the continental edge between the offing of southwestern Nova Scotia and of Sable Island, at 93 to 134 fathoms; and while Huntsman 7S describes it as characteristic of the deep channels of the Gulf of St. Lawrence, Danne- vig 79 points out that the stations within the Gulf of St. Lawrence where the Canadian Fisheries Expedition took rockling eggs and larvae in any number, rather generally distributed in the south- ern part, a few in the northeastern part, were all "close to land or above the more shallow banks." Pelagic rockling fry are listed under this name in the Reports of the Newfoundland Fishery Re- search Commission also, from many stations in the Grand Banks region, and around the coast of Newfoundland to the Northern Peninsula on the east and to the inner end of the Strait of Belle Isle on the west. But it would not be astonishing if the fry of the three-bearded rockling (p. 237) should prove to be represented in these collections, together with those of our four-bearded species. Dannevig, indeed, has suggested that part of the rockling eggs taken by the Canadian Fisheries Expedition in Nova Scotian waters and south of the Grand Banks in May and June belonged to some species other than cimbrius. Importance. — The rockling is neither large enough nor plentiful enough to be of importance commercially, or of interest to the angler. Cusk Brosme brosme (Miiller) 1776 Tusk; Torsk Jordan and Evermann, 1896-1900, p. 2561. » Trans. Roy. Soc. Canada, Ser. 3, vol. 12, Sect. 4, 1918, p. 63. '< Canadian Fisheries Exped. (1941-1915) 1919, p. 27: charts figs. 18, 19. Description. — The cusk is separable from all its Gulf of Maine relatives at a glance by the fact that it has only one dorsal fin. The relationship of the anal and dorsal fins to the caudal and the outline of the latter are distinctive also, for both the dorsal and the anal are continuous with the caudal at the base but are separated from it by notches so deep that they are obviously distinct. And the caudal is evenly rounded. The cusk is a more slender fish than the hakes, being only about one-fifth to one-sixth as deep as it is long, round- bodied in front of the vent but flattened sidewise behind the vent, and tapering evenly backward to the base of the caudal fin. The mouth is large, gaping back to opposite the rear third of eye, is set slightly oblique, and is armed with small, sharp, curved teeth. The snout is blunt at the tip. The upper jaw encloses the lower when the mouth is closed; the eye is of moderate size; the chin bears one barbel; and the entire head and trunk are clad with small scales. The dorsal fin (85 to 105 rays) runs the whole length of the back from the nape of the neck, and is of uniform and moderate height from end to end with rounded corners. The anal fin is similar to it in outline but is only a little more than half as long (71 to 76 rays). The pectoral fins are rounded, and about half as long as the head. The ventral fins are about as long as the pectorals, with their 5 rays free at the tips, and are situated a little (but ob- viously) in front of the pectorals. All the fins are so thick and fleshy at their bases that it is only near their margins that the rays are to be seen. Color. — The cusk varies in color, no doubt con- forming to the bottoms on which it lives. Its upper parts range from dark slaty to dull reddish brown or to pale yellowish, paling to grayish on the lower part of the sides and to dirty white on the belly. Old fish are plain colored, the sides of small ones, however, are often cross-barred Figure 115. — Cusk (Brosme brosme). Boston market. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 239 with about half a dozen yellowish bands. The pectoral and ventral fins are of the same color as the sides, and the ventral fins are sooty at their tips. The most characteristic color mark is that all three of the vertical fins (dorsal, caudal, and anal), which are of the general body tint at their bases, are black at the margin, and they are nar- rowly edged with white, except that the anal may lack the white edging on some individuals. Size. — Cusk grow to a maximum length of about 3}i feet; one 40 inches long, weighing 27 pounds, trawled by the Albatross II in the central part of the Gulf of Maine, in 120 fathoms, was the largest that has been recorded definitely from our waters. But those caught in the Gulf of Maine average only IK to 2/i feet long, and from 5 to 10 pounds in weight. The relationship of weight to length, in fish we have handled recently, was as follows: 26 inches, about 5}i pounds; 33K inches, about 14% pounds; 36 inches, about 20 pounds. The size at which cusk first mature sexually seems not to have been recorded. Habits. — Once the young fry have taken to the bottom they are ground fish so exclusively that we have never heard of one swimming up to the upper waters, as cod so often do, and even hake. They are sluggish, too, and weak swimmers, but powerful of body; when a cusk is hooked it is likely to twine itself around one's line in a bother- some way. They are more or less solitary, not so abundant anywhere as cod, haddock, or hake are, as may be illustrated by the following catches counted as they came from the water by representatives of the Bureau of Fisheries in 1913: Twenty miles east of Cape Cod Light, November 16 and 17, 1913, long line, 460 cusk to 2,150 haddock and 1,22S cod; 15 miles southeast of Monhegan Island, June 24 and 25, 1913, long lines, 580 cusk to 2,880 hake; Jefln^ys Ledge, December 11 and 12, 1913, long line, 230 cusk to 470 haddock and 475 cod; northwest part of Georges Bank, October 10 to 13, 1913, otter trawl, 4 cusk and 12,473 haddock; 6 miles east of Boon Island, March 30, 1913, gill net, 5 cusk, 1,055 haddock; 51 cod, 20 pollock, and 76 dabs (Hippoglossoides) . It also seems that cusk move little from bank to bank. Thus the "Massachusetts fishermen tell me," wrote Goode m "that these fish are usually found in considerable abundance on newly dis- covered ledges, and that great numbers may be taken for a year or two, but that they are soon all caught. Sometimes, after a lapse of years, they may be found again abundant on a recently de- serted ground." "Neither is there any definite evi- dence that the cusk performs in or offshore migra- tions with the seasons, at least in our Gulf. The cusk is so purely a fish of at least moderately deep water that we have never heard of one taken in less than 10 to 15 fathoms of water within our Gulf. On the other hand, it is safe to say that there are few cusk living below 100 fathoms or so in the deep basins of the Gulf. But they range down to 250-300 fathoms on the continental slope off southern New England, according to Goode and Bean.81 And they have been caught down to 530 fathoms in the Faroe Channel. Cusk are decidedly fastidious, too, in their choice of bottoms, being found chiefly on hard ground, especially where the sea floor is rough with rocks or boulders; on gravelly or pebbly grounds; occasionally on mud with hake, but seldom on smooth clean sand. In Norwegian waters they often lurk among gorgonian corals, and they may have this same habit on the parts of our offshore banks where these are plentiful. The cusk is a fish of cool water, but not of the coldest. In the Gulf of Maine (once the fry have deserted the surface for the bottoms at their chosen depths), cusk spend their lives in water which does not warm above about 48° to 50° at the warmest season, nor cool below about 33° to 34° at the coldest. And it is probable that temper- atures of 32° F. or lower are the factor that limit their American range in the north (p. 242). Food. — Little is known of the diet of the cusk. European students describe the stomachs as usu- ally containing crustaceans, sometimes mollusks. And crabs, with occasional mollusks, that we found in the stomachs of several cusk caught on Platts Bank in the summer of 1924, are the only record of its food of which we know, for this side of the Atlantic. But the cusk is not fastidious as to bait, accepting clams, cockles, and herring readily. Cusk spawn in spring and early summer in both sides of the Atlantic. In European waters the season lasts only from April until June; but » Fish. Iuci. U. S., Sect. 1, 1884, p. 233. " Smithsonian Contrib. Knowl., vol. 30, 1895, p. 385. 240 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE throughout July in the Gulf of Maine, for we have caught several nearly ripe females on Platts Bank and around Boon Island at the end of that month, though we have seen no perfectly ripe fish. In the eastern Atlantic cusk spawn chiefly deeper than 100 fathoms, to judge from the distribution of the eggs at the surface. But the chief production of eggs probably takes place in shallower water in the Gulf of Maine, since most of the stock lives in lesser depths there. And some must spawn close inshore, for we have taken cusk larvae only 6 to 13.8 mm. long off Cape Cod; in Provincetown Harbor; and near the Isles of Shoals.82 We owe what is known of the eggs and larvae to European students. The cusk is among the more prolific of fishes, more than 2 million eggs having been estimated in a female of medium size. Their eggs are buoyant like those of other gadoids; 1.29 to 1.51 mm. in diameter, with one oil globule of 0.23 to 0.3 mm.; and they may be recognized by the brownish or pinkish color of the oil globule, together with the fact that the entire surface of the egg is finely pitted. The larvae are about 4 mm. long when they hatch. The vent is situated at the base of the ventral finfold as it is in other gadoids, but they are separable from all other gadoid larvae that occur in the Gulf of Maine by the pinkish oil globule at the posterior end of the yolk. The yolk is absorbed in about a week after hatching, when the larvae are about 5 mm. long. The ventral fins of the little cusk elongate as it grows, like those of young hake and of young rockling, besides becoming heavily pigmented with black. But cusk larvae are separable from those of hake and of rockling by the fact that their ventral fin rays are separate one from another, and by the presence of three patches of black pigment: one on the top of the head; a second over the gut; and a third at the tip of the tail, besides two vertical black bands which divide the trunk behind the head into three nearly equal sections. The rockling has only one band of pigment behind the vent, and neither of the hakes that are common in the Gulf of Maine has a definite cross-band of pigment. The first traces of the vertical fin rays of the young cusk are visible at about 12.5 mm.; the dorsal and anal fins are differentiated at about 28 mm.; and it is at this stage that the ventrals are at their longest, relatively. Fry of 40 mm. and upward show most of the characters of the adult. And the relationship of their dorsal and anal fins to the caudal, and the presence of only one dorsal fin and one anal fin is sufficient to identify them from this stage on. Figure 116. — Egg (European). After Schmidt. Figure 117.— Larva, 6.8 mm. (European). After Schmidt. Figure 118. — Larva, 9.25 mm., off northern Cape Cod. Cusk (Brosme brosme). The older cusk fry, while still living at the sur- face, are described by Schmidt 83 as greenish yellow with blue eyes, not silvery-sided. The young cusk drifts near the surface, as other gadoids do, until it is 2 inches long or more, and there is reason to believe that in European seas they first seek the bottom in considerable depths. But we have nothing to offer on this point for the Gulf of Maine. 8* The records arc July 22, 1912, 1 specimen; July 20, 1916, 4 specimens; and July 22, 1916, 1 specimen. •3 Meddel. Kommis. for Havundersftgelser, Serie Fiskeri, vol. 1, No. 8, 1905, p. 7. He also describes the larval stages of the cusk. FISHES OF THE GULF OF MAINE 241 The rate of growth of the cusk has not been The landings, 1931-1935, in Boston, Gloucester, studied, so far as we know. and Portland (to nearest 1,000 pounds) follow: General range. — Both sides of the North Atlan- Locality mi mi ms mi ms tic, chiefly in moderately deep water and on hard <;«>»... ..225.000 w.000 ,73.000 012.000 1.023.000 1 J . Fippemes 19,000 69.000 35.000 47,000 61.000 bottoms; north on the American coast to the piatts 7,000 6.000 105.000 84,000 45.000 xt e Ji i t> 1 I a ix. CM -a _* T>„11„ Jeffrey Ledge 301.000 143,000 148,000 122,000 53,000 Newfoundland Banks, and to the Strait of Belle stellwagen Bank ..... 65i000 ,53.000 ^wo 259,00o 78.000 Isle, south regularly to Cape Cod, rarely to .... ... , , . , ... ., ,T t-i 1 1 1 -u vr >> p are mclmed to believe that the wide dif- southern New England, and occasionally to JSew , . , , , _ , ' , _ . . , T . _. ferences from year to year, m the catches on these Jersey; northern coasts of the Bntish Isles, Den- ^^ ground; reflect the Dumber of vessds that mark (Jutland), northern part of the North Sea, fished there) rather than the number of cusk and Kattegat off Bohuslan, Sweden, to Iceland waiting there to be caught. and the Murman coast in the eastern Atlantic. Cusk are said to be plentiful on the rather in- It reaches east and west Greenland only as a rare definite ground off Penobscot Bay that is known stray from the south. as Jeffreys Bank (not Ledge) or "Matinieus Sou'- Occurrence in the Gulj of Maine— -The cusk is sou'west." In 1921, for example, 43,545 pounds distributed very generally in the Gulf in water werc reported thence, and considerable numbers deeper than 10 to 15 fathoms, its presence or are taken, in the aggregate, on the patches of hard 1 1 ,• ,1 ■ . <• 1 i* ™ bottom that skirt the coast of Maine, as appears absence depending on the precise type of bottom. . , , , • „ . ., , . „ . , , „„.N from the approximate amounts landed m the Because of its preference in this respect (p. 239), „ / „. . ., ..,_ . ,r ■ i- . . smaller ports S5 m the dill erent Mame counties in it varies greatly in abundance m different parts of mQ and m ig45. York> QfiQQ pounds anJ 2fiQQ the Gulf, and the grounds occupied by it are pounds; Cumberland (exclusive of vessel landings much less extensive than those haunted by cod, at Portland), 79,000 pounds and 182,000 pounds; by haddock, by pollock, or by the hakes. Thus Sagadahoc, 15,000 pounds and 44,000 pounds; cusk are rarely taken in Cape Cod Bay or in the Lincoln, 27,000 pounds and 3,000 pounds; Knox, deeper holes in Massachusetts Bay, and we have 52,000 pounds and 109,000 pounds; Hancock, taken none on the soft mud of the deep bowl west 12,000 pounds and 22,000 pounds; Washington, of Jeffreys Ledge. But considerable numbers are 4,000 pounds and 500 pounds, respectively. caught on the ledges off Chatham, Cape Cod, on Some cusk are caught at the mouth of the Bay Stellwagen Bank, and on the broken grounds of Fundy also, especially about Grand Manan between the latter and Cape Ann, while they are on the N«w Brunswick side, and off Brier Island plentiful off Cape Ann and on Jeffreys Ledge, the on thc Nova Scotian side' as Doctor Huntsman 1 ,. i • . ,1 __ . , „j„ .• . informs us, though none are reported toward the latter being one of the most productive cusk ' 6 J 1 /• /-1 i* mi 11 e r^ 1 head of the Bay. Small rocky patches along the grounds of our Gulf. 1 he rocky slopes of C ashes , ,T „ ; . , . J „ ~ , T , f , ° , . , . , , , T west Nova Scotian shore and oil Seal Island also Ledge, also have long been famous for cusk. In ^ gome cugk; ^ ^ &rc ^^ regulaily 0Q past years when more fishing was done there (as Grand Manan Ban]. German Bank and the in 1902 and 1905) this ground was the chief source fishing grounds off Lurcher Shoal are less pro- of supply for the cusk landed in New England. ductive of cusk, perhaps because they are floored, In 1935, similarly, about 30 percent of all the mostly, with patches of gravel and pebbles and cusk landed in Portland, Gloucester, and Boston small stones alternating with sand and clay, came from Cashes. And we have caught more But large catches are taken on Browns Bank, and cusk there than anywhere else. As might be fair numbers on the rougher spots on Georges expected, cusk are also caught on Fippenies and Bank, though its smoother expanses yield only Piatts Banks by the few vessels that fish there as an occasional cusk. is illustrated by the catches reported from these The only important exceptions in our Gulf to inshore grounds for the 5-year period 1931-35" ,llL> rule fchat cusk hold to rock-v -round are that they are at least tolerably plentiful in the co-called 14 1935 is the most recent year when landings were reported from these grounds, separately. "B -Mostly by small boat fishermen. 242 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE South Channel, where the bottom is mostly smooth (see regional summary of 1945 catches, p. 242) ; that some are caught with hake off the coast of Maine on broken or even muddy bottom; and that we have trawled a few, on the Atlantis, in depths greater than 78 fathoms off Cape Cod, where the bottom is mostly a sticky sand. One striking accompaniment of the preference of cusk for rough or stony grounds in moderately deep water, is that many more are caught around the peripheral belt of the Gulf, between, say, the 15-fathom and the 75-fathom con torn- lines, than are on the off-shore rim formed by Nantucket Shoals, Georges Bank, and Browns Bank. The one not- able exception is that there are so few cusk, if any, in the inner parts of the Bay of Fundy that they are not mentioned in the fishery returns for the Bay, except for a few thousand pounds taken near its mouth on the Nova Scotian side. This regional contrast is illustrated by landings by United States fishermen (1945) 86 and Canadian fishermen (1944, 1946) combined, of between 215,000 and 250,000 pounds off western Nova Scotia; 87 1,000 to 15,000 88 pounds at the mouth of the Bay of Fundy, Nova Scotian side; about 63,000 pounds off eastern Maine; about 333,000 pounds off central Maine; about 255,000 pounds off western Maine; about 419,000 pounds off east- ern Massachusetts; about 338,000 pounds from the small grounds in west central part of the Gulf; about 68,000 pounds from the South Channel; a few hundred pounds only from Nantucket Shoals; about 25,000 pounds from the northwest part of Georges Bank ; none reported from the southwest- ern part of Georges; about 17,000 pounds from the eastern central and northeastern parts of Georges Bank; and about 18,000 pounds from Browns Bank. Following the cusk eastward and northward, we find that considerable quantities are caught all along the Nova Scotian Banks, from Browns to Banquereau and to the Canso grounds off Cape Breton Island (catch, in 1946, about 542,000 pounds by United States and Canadian vessels combined). Cusk were also reported from the Newfoundland Banks many years ago by Goode,89 but there cannot be many of them there, for they are not included among the fishes reported thence from cruises of the Newfoundland Fishery Ke- search Commission.90 And the only report we have found of cusk anywhere in the Gulf of St. Lawrence is at Cheticamp, on the Cape Breton shore.91 In fact, the only definite record we have found of cusk on the American coast farther north than Cabot Strait is of one that was caught in the Strait of Belle Isle at 80 fathoms many years ago.92 And while the cusk has been credited repeatedly to Greenland, it is a rare stray there from the south, only 7 specimens having been reported there dur- ing the period 1936-46, 5 of them on the west coast, 2 on the east.93 Westward from Cape Cod, the cusk is said to have been "not uncommon" formerly in Vineyard Sound, but it is so rare there now (if it ever occurs there) that we have not heard of one caught any- where in the Woods Hole region of late years. But one was caught off Newport, Rhode Island, in November 1898,94 and two were reported from Cape May, New Jersey, many years ago.95 Importance. — The cusk is a good food fish and there is a ready market for all that are brought in. The landings from the Gulf of Maine by United States fishermen ranged between about 1,600,000 pounds and about 2,200,000 pounds for the years 1945-47; between about 100,000 pounds and about 200,000 pounds by Canadian fishermen for 1944 and 1946, which contrasts with 2 to 7 million pounds yearly by United States fishermen alone for the few years that pre- ceded the publication of the first edition of this book (in 1925). We attribute this decrease to the evolution that has taken place in the fishery from long lining to otter trawling chiefly, and to the ■• Most recent year for which landings have been published by counties, for Maine and Massachusetts, in addition to the landings at Portland, Gloucester, Boston, and New Bedford. »' Off western Nova Scotia, by United States fishermen, 1945, about 108,000 pounds; Yarmouth County landings, Nova Scotia, about 140,500 pounds in 1944, about 106,000 pounds in 1946. « 1944, 15,000 pounds; 1946, 700 pounds. •• Fish. Ind. U. S., Sect. 1, 1884, p. 232. « Frost (Service Bull. 8, Newfoundland Dept. Nat. Resources, 1938, p. 29) states that there is no definite record of cusk on the Newfoundland fishing grounds. •' Recorded by Cornish (Contrib. Canadian Biol. (1918-1920) 1921, p. 114) from fishermen's reports. W. R. Martin of the Fisheries Research Board of Canada, writes us that any fisheries reports of cusk for the Gulf of St. Lawrence actually refer to hake. " Weitz, Proceedings, Boston Soc. Nat. Hist., vol. 10, 1866, p. 274; Packard, Labrador Coast, 1891, p. 819. n For further details and discussion of the status of the cusk as a Greenland fish, see Jensen (Spolia Zool., Mus. Hauniensis, Copenhagen, vol. 11, 1948, p. 175). •' Tracy, 40 Ann. Rept. Commiss. Inland Fish. Rhode Island, 1910, p. 159. 11 Abbott, Geol. New Jersey, 1868, p. 819. FISHES OF THE GULF OF MAINE 243 fact that the cusk (frequenting rough bottom) is are 70 to 80 times as great by otter trawls as by not a good trawl fish. And 80 to 90 percent as long lines.96 much cusk (pounds) are caught on long lines as A few cusk are caught from party boats by are caught in otter trawls even today (as illus- sportsmen hand lining for ground fish in general, trated by 1947), although the yearly landings of but most of the cusk live too deep to be of any fish of all kinds in Maine and Massachusetts now particular interest to anglers. THE GRENADIERS. FAMILY MACROURIDAE The grenadiers are characterized externally by of all the species known from the western North having large heads, projecting snouts, and slender Atlantic and from central American seas, bodies that taper to whiplike tails, with no defi- _ nitely demarked caudal fin. They have two KEY TO GULF OF MAINE GRENADIERS dorsal fins, the first high, the second very low l. The dorsal spine is perfectly smooth but occupying the greater part of the back. The Long-nosed grenadier, p. 246 anal fin is nearly as long as the second dorsal, or The dorsal sPine is crated, with teeth which can be , felt if not seen 2 longer. The grenadiers are allied to the cod familv, in 2. The vent is considerably in front of the origin of the classification, by the structure of their skull,' but anal Gn> the skin surrounding the vent is naked and ,, ,.~ . ., , . ., . , black; the dorsal fin spine is strongly serrated they differ from the cod tribe in having one stout Common grenadier, p. 243 spine in the first dorsal fin. They are deep-sea The vent is close to the or;g;n 0f the anal fin; the skin fishes, living on the bottom, loose in texture and around the vent is scaly and pale colored; the serra- weak swimmers. Many species are known, but tions on dorsal fin spine are so fine that they are only three of them have ever been taken within hard'y visible> thoush they can be fdt--- --- ,, n , ,, „ „ ••»» . Rough-headed grenadier, p. 245 the confines of the Gulf of Maine. Besides the species described below, three Common grenadier Macrourus bairdii others, C'oryphaenoides rupestris, C. carapinus and Goode and Bean 1877 ' Nematonurus armatus,97 have been taken on the continental slope abreast of the Gulf and off Rat-tail; Marlin-spike southern New England often enough to show Jordan and Evermann, 1896-1900, p. 2583. that they are common there below 350 fathoms. They are typical inhabitants of the deep-sea floor, Description— This grenadier could hardly be never likely to rise shoal enough to come within mistaken for any other fish except for one of its the limits of the Gulf of Maine.98 But fish have own tribe, so characteristic is its slender body a way of straying, and if any grenadier should be (flattened sidewise behind the vent and tapering picked up in the Gulf that proves difficult to to a whiplike tail with no definite caudal fin), in identify, we recommend forwarding it either to combination with a pointed snout that overhangs the Laboratory of the Fish and Wildlife Service the mouth; very large eyes; and high first dorsal at Woods Hole, to the U. S. National Museum, fin with one large spine; but very low second dorsal Washington, D. C., or to the Museum of Compar- An. And it has a chin barbel like a cod (not shown ative Zoology, Cambridge, Mass., to be named. in the illustration). As noted above, the second Parr " has recently published a detailed synopsis ray of the first dorsal fin is a true spine, serrated along its front edge with about 15 sharp and very » Otter trawlers landed about 499 million pounds of flsh of all sorts in noticeable teeth pointing Upwards. Maine and Massachusetts in 1947; long liners about 7 million pounds. rp. fi t J,™,,! f,n (o „tjfp _„„„ fl.„ fi_o+ vprv "According to Parr (Bull. Bingham Oceanogr. Coll., vol. 10, art. 1,1946, itXe llrSt °-orsal Im ^ Stin rays, Hie nrsi very p. 54) this is the correct name of the grenadier that was reported by Ooode short, and 1 1 Softer rays) is triangular, about twice "l(fi t^Z"*' Kn°WL' V01' 3°' 1895' P' 4°7) " IIV™' as hiSh as ifc is lonS; and [t originates over the » For descriptions and lists of localities where they were taken during the pectorals, close behind the gill Openings. The early cruises by vessels of the U 8 Fish Commission see Ooode and Bean between the two dorsal fins is about as long (Smithson. Contnb. Knowl., vol. 30, 1895). In June 1949 we trawled about ^ «*~v ~ o 200 rupestris on the slope off southern Nova Scotia and off the southeastern face of Georges Bank, at 290-120 fathoms, from Caryn of the Woods Hole ' Parr (Bull., Bingham Oceanogr. Coll., vol. 10, art. I, 1946, p. 37) places Oceanographic Institution. this grenadier in the genus Nezumia of Jordan, 1904. But it seems wiser to ■ Bull. Bingham Oceanographic Coll., vol. 10, art. 1, 1946. follow the older and more familiar usage here. 244 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE ---:;-v5^^ ^ Figure 119. — Common grenadier (Macrourus bairdii), off Cape Ann. From Goode and Bean. Drawing by H. L. Todd. as the height of the first dorsal fin. The second dorsal fin (about 137 rays) extends back to the tip of the tail, is so low that its membrane is hardly visible, and tapers to practically nothing at the rear end. The anal fin is considerably longer than the second dorsal (only about 120 rays, however) and more than twice as high as the second dorsal. The pectoral fins are rounded at the tip. The ventral fins, which stand under the pectorals or a little behind the latter, are triangular, with the first ray prolonged as a threadlike filament. The exposed parts of the scales on the body, including the head and shoulders, are rough with minute sharp spines closely crowded together. The jaws are armed with several bands of small recurved teeth. The vent is situated a consider- able distance in front of the point of origin of the anal fin, and the skin immediately surrounding it is scaleless and black. Color. — The many we have seen have been uni- form gray above and below. Also described as light brownish gray above, silvery below, with dark bluish or blackish belly. The lower surface of the snout is pink, the throat is deep violet, the first dorsal is pink with blackish spines, and the eyes are dark blue. Size. — Usually about 1 loot long. The largest we have seen was 16 inches long. Habits. — Grenadiers are bottom fish, usually found on soft mud, and they are very feeble swim- mers. They usually live in at least 80 to 90 fathoms of water, and down to 1,000 to 1,200 fathoms (deepest record 1,255 fathoms). But one was trawled in 9 iathoms in Vineyard Sound by the Fish Hawk many years ago; a second was found floating near the surface at Eastport, Maine, by Dr. W. C. Kendall; and a third was taken in a weir at Lubec, Maine, as reported by Huntsman. Hansen 2 reports pelagic euphausiid shrimps (Thy- sanoessa longicaudata) in a grenadier stomach, while several examined by us from 100 fathoms on the edge of Georges Bank contained amphipods chiefly, together with an occasional worm and euphausiid shrimp. It is probable that grenadiers spawn in summer and autumn, for the spermaries of a specimen taken in the western basin of the Gulf on August 19 were nearly ripe, while a fully ripe male has been reported from South Channel in the last week of September. The eggs of this fish have not been seen, but it is probable that they resemble other macrourid eggs described by European authors3 in being buoyant at least for the first part of the period of incubation, with a large oil globule, wide perivitelline space, and with the surface sculptured into concave hexagonal facets. The larvae have not been seen yet. Those of other species of grenadiers have the rays of the first dorsal and ventral fins greatly prolonged. General range. — This (normally) deep-water fish has been found at many localities along the continental slope from the West Indies northward and eastward to the Grand Banks of Newfound- land,4 and rarely in the Gulf of St. Lawrence. It is also known from the mouth of the Laurentian Channel, on the Scotian Banks, in the Gulf of Maine, and even in Vineyard Sound. It has also been reported from the Azores. Occurrence in the Gulf of Maine. — The common grenadier was formerly regarded as a rare stray in the inner parts of the Gulf of Maine for oidy two had been recorded there aside from the East- port and Lubec specimens mentioned above, the ' Proc. U. S. Nat. Mus., vol. 48, 1915, p. 99. " Ehrenbaum CN'ordisches Plankton, vol. I, 1905-1909) summarizes what little is known of tho eggs and young of this group of fishes. < Rept. Newfoundland Fish. Res. Comm., (1933) 1934, p. 116. FISHES OF THE GULF OF MAINE 245 one from the western basin in 1G0 fathoms, the other from off Gloucester, both of them taken many years ago. But they must be rather com- mon on the muddy bottoms ot the deeper parts of the Gulf in 85 to 125 fathoms, for we have caught more than 100 of them at various localities on recent trawling trips. No doubt it is because few vessels ever fish on these grounds, which are not productive either of cod or of haddock, that the presence of grenadiers there has been overlooked. A grenadier, too, was reported from the slope of Jeffreys Ledge, in about 50 fathoms, during March 1934. Grenadiers, together with the long-finned hake (p. 232), are the most abundant fish on the con- tinental slope abreast of the Gulf below 100 fathoms.' Rough headed grenadier Macrourm berglax Lacepede 1802 e Rat-tail; Onion-eye Jordan and Evermann, 1896-1900, p. 2583. Description. — This species resembles the com- mon grenadier (p. 243) so c'osely in general appear- ance that we need only indicate the points of difference. Most obvious of these are that its snout is shorter and blunter, with more highly arched dorsal profile; that it has 4, 5, or 0 distinct ridges on the top of its head; that its head is • For a list of captures on the continental slot* during the early cruises "I the U. S. Fish Commission vessels, see Ooode and Bean (Smithsonian Con- tiib. Knowl., vol. 30, 1895, p. 394). • Authorities disagree as to the correct scientific name of this grenadier, for while it has usually been referred to as berglax Lacepede. 18(12, Jensen (Spolia Zool., Mus. Hauniensis, Copenhagen, vol. 9, 1948, p. 178) prefers il)'1 species- name /aorfcii that was applied to it more recently by Sundevall, 1840. on the ground that the grenadier to which Lacepede gave the name berglax was another fish, Coryphaenoidcs rupettris Gunnerus, 176.1, which is eommonly termed "berglax" on the west coast of Norway. relatively larger (about one-fourth to one-fifth the length of the fish, only one-sixth to one-seventh in the common grenadier); that its trunk is rela- tively stouter (about six times as long as it is deep) ; that its vent is close to the point of origin of the anal fin with the skin scaly around it, and no darker colored than on the back; and that the serrations on the large spine in the first dorsal fin are so fine that they are hardly visible. Furthermore, there are fewer (about 124) rays in the second dorsal fin, but more rays (about 148) in the anal than in the common grenadier, and its first dorsal fin is of rather different outline. The second dorsal fin, too, is relatively higher than in the common grenadier and with its membrane more developed (compare fig. 120 with fig. 119), while the filamentous prolongation of the outer ray of the ventral tins is not so long in berglax as it is in bairdii. The structure of the scales, too (visible to the naked eye), is diagnostic, for those on the head and shoulders of berglax are armed with either one longitudinal row of spines (10-12 rows of spines on each scale), or with up to 3 or 4 radiating ridges of spines while those further back each have a single row of spines, which together form conspicuous longitudinal ridges along each side of the rear part of the body. Color. — The only newly caught specimens we have seen were ash gray below as well as above; with the chest a little darker; with the rear edges of the scales on the rear part of the body still darker; with the anal fin narrowly dark edged; with the first dorsal fin and the pectoral fins sooty; and with the ventral fins sooty, except that the outermost rays are white after preservation in alcohol. Size. — This fish is larger than the other grena- diers (p. 243). It is credited with a maximum IfflBBBiBfiiri isSsSwW**''^^ ^P ; Figure 120.— Rough-headed grenadier (Macrourus berglax), Banquereau Bank. From Goode and Bean. Drawing by H. L. Todd. 246 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE length of 3 feet and a weight of 4 or 5 pounds but the largest we have seen is only 29 inches long. General range. — This is a deep-water fish like its relative, but is more northerly in its distribu- tion, being known off northern Norway, Spits- bergen, Iceland, southern Greenland, in Davis Strait, and southward along the continental slope of North America as far as Georges Bank. One has even been found floating dead on the surface, off New York Harbor, but it may have been thrown overboard from a fishing boat returning from the off shore banks. Occurrence in the Gulf of Maine. — Three quarters of a century ago, when halibut were more plenti- ful in the Gulf of Maine than they are today, and when vessels, long-lining from Gloucester, still resorted regularly to the deep channel between Georges Bank and Browns Bank as well as to the deep gullies that interrupt the Nova Scotian banks, large grenadiers were often hooked. Fishermen described them as common enough to be a nuis- ance, for they stole the baits meant for other fish and were of no commercial value themselves. It was on the strength of such reports that Goode7 characterized them as "exceedingly abundant on all of our offshore banks." A few were brought in "from off the coast of New England."8 And our re-examination of three specimens, one taken on the outer edge of either La Have Bank or Banquereau in 1878 a second taken "off New England" in 1880, the third (probably from the Grand Banks) obtained in Boston Market by Prof. G. H. Parker in 1903 8 has proved that ear- lier identifications of them as berglax were correct. ' Fish. Ind. U. S., Sect. 1, 1884, p. 244. > Bean, Proc. U. S. Nat. Mus., -vol. 3, 1881, p. 80. ■ These three specimens, the largest 29 inches long, are in the Museum of Comparative Zoology. We have not heard of one, either from Nova Scotian waters or from the Eastern Channel since 1903 ;10 not because they have vanished thence, but simply because very little long-line fishing is now done deep enough off our coasts. And there is always the chance that some vessel, fishing down the slopes of Sable Island Bank, La Have Bank, or southeastern Georges, may pick a few rough headed grenadiers at any time when least expected. One hundred fathoms may be set as about their upper limit ; most of those caught have been from 100 to 300 fathoms on both sides of the Atlantic; and they have been taken as deep as 677 fathoms by the Albatross off the southeast slope of Georges Bank. They are supposed to feed on small fish and on Crustacea but we find no definite record of the contents of then- stomachs. Females with the roe nearly ripe have been taken off northern Norway in May, suggesting that this is a spring spawner, but nothing definite is known of its breeding habits. Long-nosed grenadier Coelorhynchus carminatus (Goode) 1880 Jordan and Evermann, 1896-1900, p. 2588. Description. — This species resembles the com- mon grenadier (p. 243) so closely in its general appearance that there is danger of mistaking it for the latter; but it is identifiable by the facts that its dorsal spine is perfectly smooth and that its first dorsal fin is rounded instead of triangular; and that its snout not only overhangs the mouth slightly farther, but is thinner tipped.11 10 The most recent record with which we are acquainted is of one 16 inches long that we trawled on the southeastern slope of Georges Bank, at 500 fathoms, June 1949, on Caryn of the Woods Hole Oceanographic Institution. 11 Commonly described as "sturgeon-like," but this characterization applies better to other members of the genus which have still longer snouts. Figure 121.— Long-nosed grenadier (Coelorhynchus carminatus), continental slope off Marthas Vineyard. From Goode and Dean. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 247 Color. — Described as silvery gray. Size. — About 10 inches long. General range and occurrence in the Gulf of Maine. — This deep-water ground fish has been taken at many localities off the American coast from the West Indies and Gulf of Mexico north- ward along the continental slope to abreast of southern Nova Scotia, in depths of 104 to 464 fathoms. It is included here because it has been recorded once off Nantucket in 148 fathoms. THE OPAHS. FAMILY LAMPRIDAE For the characteristics of this family, see the following description of its unique representative, the opah. Opah Lampris regius (Bonnaterre) 1788 Moonfish; Jerusalem haddock Jordan and Evermann, 1896-1900, p. 954, as Lampris luna (Gmelin) 1789. Figure 122. — Opali {Lampris regius). and Bean. After Goode Description. — The thin, deep form of the opah (trunk less than twice as long as it is deep) with moderately slender caudal peduncle, which does not have longitudinal keels, and the rather pointed snout, might suggest an enormous butter- fish, were it not provided with very long falcate ventral fins, whereas the butterfish has no ventrals. The ventrals, also, of the opah have 14 to 17 rays; none of the mackerel or pompano tribes has more than 8. The forward part of the single dorsal fin (53 to 55 rays) is high, its outlines strongly falcate. The anal (38-41 rays) is shorter than the dorsal, and it is about equal in height to the low part of the dorsal throughout its length. Both anal and dorsal fins extend back close to the base of the caudal fin, and each of them is depressible in a groove. The tail fin is emarginate, the pectorals are conspicuously pointed, with their bases hori- zontal instead of vertical. The mouth is small and toothless, the scales are minute, and the lat- eral line is strongly arched upward above the pec- toral fin, then downward toward the rear. Color. — We have never seen this fish alive, but it is described as of a beautiful dark steel blue above, shading into green with silver, purple, gold, and lilac luster down the sides, and as rosy on the belly, with vermilion fins, while the whole body is speckled with silvery and milk-white spots. Size. — The opah grows to a length of 3 to 6 feet; most of them are 3 to 4 feet long. Habits.— The opah is usually spoken of as a deep-sea fish, but this is a misnomer, for it is caught on hook and line no deeper than 50 to 100 fathoms off Madeira, where it is taken in some numbers. Being so very rare off our coast, we need merely note that it feeds chiefly on squid, isopods, and small fish, as well as on seaweeds; that it is an excellent food fish; and that nothing is known of its breeding habits. General range. — Open waters of the Atlantic and Pacific Oceans; recorded off Madeira, Scan- dinavia, the British Isles, Norway, Iceland, New- foundland, Nova Scotia, Maine, Cape Cod, and Cuba in the North Atlantic; also in the Gulf of Mexico off the west coast of Florida. Occurrence in the Gulf of Maine. — Two speci- mens, only, of this oceanic wanderer have been reported definitely within the limits of our Gulf, one caught on a long line on Browns Bank in the spring of 1932, 12 the other, weighing 165 pounds fresh, was taken in an otter trawl on the northeastern part of Georges Bank, in August 1947.13 One also was reported from Maine by Goode and Bean,14 but this may have been based on a letter to D. S. Jordan from Everett Smith, July 19, 18S8, reporting that a "Sun Fish," identified '» Reported by Vladykov, Proc. Nova Scotia Inst. Sci., vol. 19, 1935, p. 8. »' This specimen Is in the Museum of Comparative Zoology. M Goode and Bean. Smithsonian Contrib. Knowl.. vol. 30. 1895, p. 223. 248 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE as an opah by the included description and sketch, had been landed in Portland, Maine, from the Grand Banks.16 One was caught off Sable Island, Nova Scotia, about 1856, and another off La Have Bank many years ago; 16 a specimen about 3 feet long was taken in July 1925 on Western Bank, southwest of Sable Island, by the schooner Falmouth; 17 and another of the same size stranded on the beach at Hyannis, Mass., on September 17, 1928. 18 THE FLOUNDERS AND SOLES. FAMILIES HIPPOGLOSSIDAE, PARALICHTHYIDAE, PLEURONECTIDAE, BOTHIDAE, AND ACHIRIDAE The flatfishes are a very homogenous tribe, so different from all other fishes that no one is likely to mistake any one of them for any other sort of fish. What strikes one first is their flatness ; less obvious is the fact that they do not lie on the belly but on one side, right or left. And their skull twists in the course of development so that the eye which was originally on the side that is fated to be underneath, migrates around the head, until both the eyes finally come to lie close to- gether, on the side that is uppermost as the fish lies on bottom. But the mouth retains its origi- nal position more nearly, so that it is often de- scribed as opening sidewise. The larval flounder swims on edge like any other fish; the migration of the eye takes place shortly before the fry take to the bottom. All of the flatfishes have a single long fin on each edge, one the dorsal and the other the anal; they also have well-developed ventral fins (at least on the eyed side) which are either on the right-hand edge or on the left-hand edge as the fish lies. Most of the Gulf of Maine species also have pectoral fins, one on the upper side as the fish lies on the bottom, the other on the lower side. The ventral fins are in front of the pectorals or in line with them; the abdominal cavity is very short, and some species are armed with a stout anal spine. Our several flatfishes look much alike; indeed, they are often confused. But it is not difficult to tell one from another, for the distinctive char- acters are rather precise, even if not obvious at first glance. Huntsman 19 has published a very useful key to the eastern Canadian species, which is expanded here to cover the Gulf of Maine. KEY TO GULF OF MAINE FLATFISHES 1 . Eyes on the left-hand side, and guts at left-hand edge, as the fish lies on bottom 2 Eyes on the right-hand side, and guts at right-hand edge 5 2. The lateral line is straight Citharichthys arclifrons, p. 294 The lateral line is arched over the pectoral fin 20 3 3. The two ventrals are not alike, the left (upper) being continuous with the anal fin, the right (lower) separate from it Sand flounder, p. 290 The two ventral fins are alike 4 4. The upper side is marked with four large oblong black eve-spots; there are fewer than 82 rays in the long right-hand (dorsal) fin Four-spotted flounder, p. 270 The upper side is marked with many small spots; there are more than 84 rays in the long right-hand (dorsal) fin Summer flounder, p. 267 5. There is a well-developed pectoral fin on the eyed side fi There are no pectoral fins Hog choker, p. 296 6. Mouth large, gaping back as far as the eye; jaws and teeth nearly equally developed on both sides . 7 Mouth small, not gaping back as far as the eye; the jaws are nearly straight on the upper side, but curved on the lower side 9 7. Margin of tail fin rounded American dab, or plaice, p. 259 Margin of tail fin slightly concave, with angular corners 8 8. Lateral line arched close behind the gill opening Halibut, p. 249 Lateral line nearly straight Greenland halibut, p. 258 »5 We are indebted to Norman J. Wilimovsky for showing us a copy of this letter. " Vladykov, Proc. Nova Scotia Inst. Sci., vol. 19, 1935, p. 6. " Radcllffe, Copeia, No. 151, 1926, p. 112. »■ Reported by Robert Coffin of the Bureau of Fisheries station at Woods Role. Mass. '• Our Eastern Flat Fishes, Canadian Fisherman, vol. 5, No. 6, 1918, pp. 788-790. a In all the flounders of this type so far recorded from the Gulf of Maine both of the pectoral fins are well developed. Should one be taken with no pectoral fin on the blind side it would probably be the deep-water Mcmoltnc gts.iUicaiida. FISHES OF THE GULF OF MAINE 249 9. Lower side of head with large open mucous pits; 100 or more rays in the long left-hand (dorsal) fin.. Witch, p. 285 Lower side of head lacks open mucous pits; fewer than 90 rays in the long left-hand (dorsal) fin 10 10. Lateral line arched behind the gill opening Yellow-tail, p. 271 Lateral line nearly straight 11 11. Top of the head between the eyes rough with scales— Winter flounder (including the Georges Bank flounder) p. 276 Top of the head between the eyes naked and smooth Smooth flounder, p. 283 Atlantic halibut Hippoglossus hippoglossus fish but is hidden by the skin in old fish. The (Linnaeus) 1758 two pectoral fins are of different shapes, the one ,0„„ „„„ „„„ on the upper (eyed) side of the fish being obliquely Jordan and Everrnann, 1896-1900, p. 2661. • . , , ., \, ~ ,, , • , ? , , pointed while tne fin on the lower side is rounded. Description. — This is not only the largest of The rather small ventral fins, which are situated flatfishes, but is one of the best characterized; 'n lront of the pectorals and are separated from its most obvious distinctive characters, apart from the anal by a considerable space, are alike. Hab- its size, being the fact that it lies on the left side; 21 but, like other flatfishes are scaly on the whole that its mouth gapes back as far as the eyes, and bead and body and they are very slimy with mucus, is armed with sharp curved teeth; that the rear Color. — The halibut is chocolate to olive or edge of its tail fin is concave, not rounded; that its slaty brown on the eyed (upper) side. Young two ventral fins are alike; and that its lateral line nsn arc paler, and are more or less mottled, while is arched abreast of the pectoral fin. Furthermore larg° ones are more uniform and darker, some- it is a narrower fish, relatively, than most of our times almost black. The blind (lower) side usu- fiatfishes (only about one-third as broad as it is aHy is P^e white in small fish, but large ones are long) but is very thick through, and its eyes are often more or less blotched or clouded below with farther apart than they are in most of the other gray (known by fishermen as "grays"). Occa- flounders. sionally a halibut is taken the blind side of which The dorsal (long) fin (98 to 105 rays) commences is marked with patches of the same color as the abreast of the eye and runs back the whole length eyed side. And we have seen one medium-sized of the fish, broadening but slightly for the first fish in which the rear third of the lower surface third of its length and then abruptly, to narrow was uniform dark brown. again toward the caudal peduncle. The anal fin Size. — Only swordfish, tuna, and some of the is similar to the dorsal fin in shape but is shorter larger sharks reach a greater size than the halibut, (73 to79rays), originates close behind the pectorals, among (iulf of Maine fishes for while reports of and is preceded by a sharp spinelike extension of specimens as large as 600 to 700 pounds have the post-abdominal bone, which projects in young usually l>een looked on as exaggerations we are glad to be able to give at least one record of a « Left-handed halibut have been caught, occasionally. Gulf of Maine halibut ill this Weight class. The ..,- ... Figure 123. — Halibut {Hippoglossus hippoglossus) , Eastport, Maine. From Goode. Drawing by H. L. Todd. 210941—53 17 250 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE fish in question was taken in June 1917, by Capt. A. S. Ree, about 50 miles eastnortheast of Cape Ann, and since it weighed 615 pounds, eviscerated with the head still attached, when brought in to the Boston fish pier, it must have been as heavy as 700 pounds while alive.22 Another halibut of 602 pounds is said to have been taken near Isle au Haut in 1902, but we cannot vouch for this one. Halibut of 500 to 600 pounds are rumored al- most every year, but the next largest of which we have definite knowledge was one of about 450 pounds caught on a hand line in the deep water between Browns and Georges Banks in 1908 by W. F. Clapp. Goode 23 likewise had records of a dozen fish of 350 to 400 pounds caught off the New England coast; the heaviest was one of 401 pounds taken near Race Point, Cape Cod, in July 1849. But a 410-pound halibut that was brought in to the Boston fish pier by the Dawn, March 27, 1941, 2i was spoken of as the largest that had been landed there in a "score of years," and it seems that halibut heavier than 300 pounds always were rarities anywhere in the North Atlantic. Full-grown females average about 100 to 150 pounds. Males run smaller, and most of the "large" fish landed in New England ports weigh from 50 to 200 pounds. The largest we have caught, taken on Browns Bank, weighed exactly 100 pounds and was 5 feet long. Halibut be- tween 7 and 8 feet long usually weigh 300 to 350 pounds, and the following table based on Ice- landic fish measured by Jesperson,25 and others from the Gulf of Maine, give the relationship of length to weight for the smaller sizes. Iceland Gulf ot Maine Length in Weight in Length in Weight in inches pounds inches pounds 74 215 63 1120 70 168 42H 30 61 107 42 33 54 to 56 60H iVA 27H 40 to 42 29 31 12 36 11 to 12 20 2% 30 9K 27 6H 24 5M — ' This flsh weighed 98 pounds dressed, the intestines accounting for 15 pounds and the ovaries (with immature eggs) for 7 pounds. Habits. — The halibut, like all the flatfish tribe, is normally a ground fish, once the young fry have taken to bottom. But it comes to the surface on occasion (p. 257), and it is a very powerful fish, when hooked. Halibut caught in shallow water are very active, usually starting off at great speed when they are hauled up from the bottom, often spinning the dory around in then- attempts to escape.26 They are usually found on sand, gravel, or clay, not on soft mud or on rock bottom; 400-500 fathoms may be set as the lower boundary to their existence in any numbers,27 but their absolute depth limit is not known. The young halibut, like the young of so many other ground fishes, drift helplessly with the cur- rent for some months after hatching (just how long is not known) ; not at the surface, howTever, but in the mid-depths (p. 253). During this period they tend both to rise in the water as they grow, and to be carried inshore, so that when they finally take to the bottom they do so in quite shallow water (p. 254). But the fry as a whole tend to work offshore again thereafter, and deeper, so that halibut caught in deep water are larger than those caught in shallow water. This fact was noticed early on Georges Bank, where most of the fish taken on the bank in depths of 30 to 40 fathoms or less ran from 125 to 180 pounds, whereas much larger ones were caught on the deeper slope to the southeast. Fishermen have also reported catching smaller fish on the inner ends of long lines set from shoal water out into deep, and larger fish on the outer ends.i8 And this rule holds equally for the other side of the Atlantic. The halibut is a boreal, not an arctic fish, in its relationship to temperature. Thus, large catches are (and were) made only at times and places where the water is at least as warm as 36°-38° F. (about 3° C). In the Grand Banks region, for instance, halibut are mostly caught either far enough down the slope to be below the icy touch of the Labrador Current, or at times and places where the latter does not reach bottom, if the fish are on the bank. But the lower limit to the temperature range of the halibut is not sharp-cut. We do, in fact, find record of at least one halibut " An account of this flsh was published In the Boston Globe, June 12» 1917. It was bought by the Shore Fish Co. » Fish. Ind. U. S., Sect. 1, 1884, p. 194. « Reported In the Boston Herald, March 28, 1941. « Meddelel. Kommls. Havunderstfgelser, Ser-Flskerl, vol. 5, No. 5, 1917. » Goode and Collins (Fish. Ind. U. S., Sect. 5, vol. 1, 1887, p. 17) give a readable account. " Capt. Baldersheim (Rapp. Cons. Intemat. Explor. Mer, vol. 56, 1929, p. 25) reports good catches at that depth In Davis Strait off west Greenland. » Goode, Fish. Ind. TJ. S., Sect. 1, 1884, p. 195. FISHES OF THE GULF OF MAINE 251 trawled on the southern part of the Newfoundland Bank in bottom water as cold as 33° F. (0.6° C), while others reported from Bay Bulls, on the east coast of Newfoundland and from Cut-throat Harbor on the outer coast of Labrador (p. 254) doubtless were in water equally cold. At the opposite extreme, it has been found that only a few halibut are taken in the parts of the North Sea where the bottom water is warmer than 46°^47° (go c )( none at all where it is warmer than 59° F. (15° C). And there is no reason to suppose that halibut ever were plentiful anywhere in the western side of the Atlantic in temperatures much higher than about 46°-47°, for while the bottom water warms locally to 50°-52° on Georges Bank in summer, and to 52°-59° on Nantucket Shoals, it was only during the winter and spring that there ever was any regular fishing for halibut on either of these grounds. On the other hand, the halibut that summer on banks where the bottom chills below about 36° in winter have been described repeatedly as with- drawing to deeper (i. e., to warmer) water for the coldest part of the year. Perhaps the best known example is off west Greenland. Here the halibut work in over the banks regularly in summer, from the deeper waters of Davis Strait, as the temperature rises, but work out again, and deeper, in autumn, as the water cools again.29 Thus it was only deeper than 350 fathoms that long liners, fishing there in 1926-28 30 found halibut in paying quan- tities at the beginning of June, when the bottom temperature on the banks was about 33°-37°. But good catches were made as shoal as 200 fathoms by the middle of the month when the temperature had risen to 35°-38°. And there was good fishing as shoal as 70 fathoms by mid-July, when the banks had warmed to 37°-39°, though many of the halibut were in deeper water still. Halibut have been described as shifting ground in the same way in the coastal belt of the Gulf of Maine (p. 257) from season to season. On the other hand, we suspect that halibut finding them- selves in water shoaler than 30 fathoms or so in the southernmost part of the range of the species, on the American side, at the onset of summer may withdraw to slightly deeper water for the time being, but definite information is lacking. The seasonal movement of halibut in onto the Greenland Banks as early in the summer as temperature allows seems to be in search of food, as Jensen points out, for a much richer supply of small fish is available to them on these shoaler bottoms than deeper down the Davis Strait slope, where they must depend chiefly on large shrimps (p. 252). And we suspect that the food supply is equally important in influencing the seasonal movements of halibut in our Gulf.31 If the prevalent view is correct, the Atlantic halibut resort to rather definite and circumscribed ground to spawn, much as the Pacific halibut do. Halibut have also been credited with extensive wanderings from bank to bank, for no evident reason. And recent tagging experiments carried out off Nova Scotia by the Fisheries Research Board of Canada,32 have proved that some of them certainly do so, in American waters. Thus fish that were marked on German and Browns Banks have been recaught as far to the eastward as Western Bank and in the general vicinity of Sable Island, while one that was tagged at Anti- costi was recaught at Seven Islands more than 100 miles to the westward. But most of the recaptures were* made within a few miles of the places where the fish had been tagged. And available evidence as to halibut migrations in the Gulf of Maine and in Nova Scotian waters is so contradictory, and so greatly complicated by the local effects of hard fishing, that it is not worth while to attempt any further discussion here. Food. — The halibut is very voracious, preying chiefly on other fishes, a long list of which have been reported from their stomachs, including cod, cusk, haddock, rosefish, sculpins, grenadiers, silver hake, herring, launce on which they often gorge in northern seas,33 capelin, flounders of various sorts (these seem to be their main dependence), skates, wolffish, and mackerel. Halibut are also known to eat crabs, lobsters, clams, and mussels; "Jensen (Meddelelser, Dansk Komm. Havunders., Ser. Flskert vol. T, No. 7, 1925, pp. 17-18) seems to have been the first to bring this to scientific attention. » Baldersheim, Rapp. Proc. Verb. Consell Internat. Explor. Mer, vol. 86, 1K29, pp. 25-28. « For a further discussion of the range and movements of the halibut m relation to temperature, with references, see Thompson and VanCleve, Rept. Internat. Fish. Comm. No. 9, 1936, pp. 22-38. « Martin and McCracken, Fish. Res. Board Canada, Progress Rept., Atlantic Coast Station, No. 50, 1950, pp. 3-8. 35 Capt. Baldersheim described halibut off west Greenland as sometimes in schools, preying on launce (Rapp. Proc. Verb. Consell Internat. Explor. Mir., vol. 5fi, 1929, p. 25). 252 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE even sea birds have been found in them.34 Fish- ermen have reported finding in halibut the heads and backbones of cod thrown overboard, and a variety of indigestible objects such as pieces of wood or iron, and even fragments of drift ice. The diet of the halibut in any particular locality depends chiefly on what other ground fish are most easily available. Thus they are reported as feeding chiefly on flatfish on Georges Bank, but on cod, haddock, cusk, and sculpins on other groimds. Halibut, like other flounders, must be nearly invisible as they lie on bottom, capturing any fish that passes within reach by a sudden rush. On one occasion a halibut of about 70 pounds was seen at the surface trying to kill a small cod with blows of its tail. "We hove out a dory and two men went in her taking with them a pair of gaff hooks. They soon returned bringing not only the halibut but the cod." 36 And halibut are very destructive to smaller fish. We read, indeed, of half a bushel of flatfish taken from one halibut. And fishermen said the appearance of a school of halibut soon drove away the cod and haddock, in the days when halibut were still plentiful on the shoaler banks. It appears that halibut do not eat many invertebrates at least in the Gulf of Maine, or in Nova Scotian waters. But a case is on record when 6 lobsters, 6 inches long, were found in the stomach of one. And Jensen found that halibut caught in deep water off west Greenland had fed chiefly on large shrimps (Pandalus borealis) .3C According to fishermen who have watched them in clear shallow water, "The halibut will advance to the bait . . . then retreat 4 or 5 feet from it . . . after repeating this performance several times — generally three or four — the fish seems to make up its mind to eat the bait, and, suddenly darting toward it, swallows it down at a gulp." 37 Halibut, in their own turn, fall prey to seals, and especially to the Greenland shark, for which they are a staple article of diet. » Smitt (Scandinavian Fishes, vol. 1, 1892, p. 414) speaks of a halibut that had eaten a razor-billed auk; Good and Collins (Fish. Ind. U. S., Sect. 5, vol. 1, 1887, p. 35) record an "lee bird" (probably a dovekie) as taken from a halibut caught on Georges Bank; and Scudder (Fish. Ind. U. S., Sect. 5, vol. 1, 1887, p. 110) reports finding the skeleton of a gull in the stomach of another. »» Ooode, Fish. Ind. U. S., Sect. 1, 1884. p. 196. >' Meddelelser, Dansk. Komm. Havundcrs., Scr. Fisk., vol. 7, No. 7, 1925, p. 18. » G oode and Collins, Fish. Ind. U. S., Sec. 5, vol. 1, 1887, p.. 17. Large halibut are very prolific, the ovaries of an Atlantic female of about 200 pounds having been estimated as containing 2,182,773 eggs, while a female of the Pacific form of "140 pounds may have as many as 2,700,000." 38 Very little is known about the breeding of the Atlantic halibut. In the eastern Atlantic halibut spawn chiefly in March, April, and May with the chief production of eggs in April, while a few fe- males may ripen as early as the end of January, and some not until June.39 Off west Greenland they spawn late in spring.40 Off the American coast it seems that the spawning season continues through the summer, for fishermen have reported ripe fish, both male and female, in April, May, June, July, August, and early September at var- ious localities from Georges Bank to the Grand Banks; 41 while the report that part of the eggs in the ovaries of a fish examined on Banquereau by representatives of the Bureau of Fisheries on Sep- tember 13, 1878, were ripe, but others immature, is evidence (if correct) that individual halibut may spawn over a considerable period. Presumably they spawn on bottom, like other flat-fishes, but definite information is lacking. The Pacific halibut is known to spawn at depths of 150 fathoms to about 225 fathoms: 42 and European students, generally, have believed that the Atlantic fish spawns deeper still, perhaps even outside the 400-500 fathom line; evidence is that naturally spawned eggs have been taken only where the depth was greater than about 550 fathoms (1000 meters), the drifting larvae less than 19 mm. long only over depths greater than about 220 fathoms (400 meters).43 On the other hand, halibut spawn regularly in the aquarium at Trondhjem, Norway, where the eggs have been fertilized artificially and hatched successfully.44 This, with fishermen reports of ripe fish, both fe- males and males, on the slopes of all the offshore Banks east of Cape Cod and, with Cox's report of >! Clemens and Wilby, Bull. No. 68, Fisheries Res. Board Canada, 1946, p. 312. " For a general survey of available information, see Taning, Meddelelser Komm. Danmarks Fisk. Havunders., Ser. Fisk., vol. 10, No. 4, 1939, p. 14. " A larva 14.75 mm. long taken on June 19 and another 21 mm. long taken on July 7 is recorded by Jensen (Rapp. Proc. Verb. Cons. Intern. Explor. Mer, vol. 39, p. 96, 1926). <> Goode (Fish and Fishery Industr. U. S., Sec. 1, 1884, pp. 196-197), mentions reports to this effect. <2 Clemens and Wilby, Bull. 68, Fish. Res. Bd. Canada, 1946, p. 312. « Tantng, Meddel. Komm. Danmarks Fisk. Havunders., Ser. Flskeri, vol. 10, No. 4, 1936, p. 8. " Rollefsen, Kgl. Norske Vidensk. Selsk. Trondhjem, Forhand., vol. 7, No. 7, 1934. FISHES OF THE GULF OF MAINE 253 two larval halibut, 20 and 21.5 mm. long, taken close in to the southern coast of Nova Scotia in shoal water,45 suggests that the American fish may spawn at least as shoal as the Pacific fish does, and perhaps even shoaler. The eggs are buoyant, drifting suspended in the water at depths greater than 30 to 50 fathoms, not at the surface. Usually they are 3 to 3.8 mm. in diameter, and the}' do not have any oil globule.46 The only other buoyant fish eggs equally large that are likely to be found in the Gulf of Maine are those of the Argentine (p. 140), but these have a large oil globule, so there is no danger of mistaking them for halibut eggs. The buoyant eggs of the Greenland halibut (p. 258) are larger still.47 In the Trondhjem aquarium the incubation of artificially fertilized eggs occupied 16 days at a temperature of about 43° (6° C.) . The larvae were 6.5 to 7 mm. long at hatching, with very large yolk sac and no pigment, growing to about 8.5 mm. by the sixth day, and developing pigment by the 10th day.48 The smallest naturally hatched Atlantic halibut yet seen 4Q was 13.5 mm. long, with the vertical fin rays appearing. The dorsal and anal fins are developed and the ventral fins are visible at about 22 mm. (fig. 125), by which time the left eye has moved upward until its margin is just visible above the contour of the head, forecasting that the fish is to be a right-handed flatfish. Fish of this size also show the large mouth characteristic of the species. Up to this stage there is little pigment. About one-fourth of the eye appears above the profile when the little halibut is about 27 mm. long, but even at 34 mm. (the largest pelagic stage yet found) the eye has not entirely completed its migration (fig. 126), though the pigmentation is « Contrib. Canadian Biol., N. Ser., vol. 1, No. 21, 1924, pp. 109-412. " For description of eggs artificially fertilized in the Trondhjem aquarium, see Rollefsen, Kg]. Norske Vidensk. Selsk. Forh., vol. 7, N'o. 7, 1934, p. 20-23; for descriptions of naturally spawned eggs taken in tow nets in Icelandic waters, see Taning, Meddel. Komm. Danmarks Fisk. Havunders. Ser. Flskeri, vol. 10, No. 4, 1936, p. 5; for description of the eggs and larvae of the closely allied Pacific halibut, see Thompson and Van Cleve, Kept. No. 9, Internat. Fisheries Comm., 1936. " 4 to 4.5 mm. in diameter according to Jensen, Kgl. Dansk Vidensk. Selsk. Skr. Nat., aid. 9. R . 6, 1935, p. 4. " For illustration of these artificially fertilized eggs In Incubation stages, and of the larvae hatched from them, see Rollefsen, Kgl. Norske Vidensk. Selsk. Forhand., vol. 7, No. 7, 1934. " What little we know of the early stages of the halibut is due to European students, chiefly to Schmidt (Meddel. Komm. Havunder-sS inches long, was reported to us by R. H. Backus as found dead in the water, in Cutthroat Harbor, August 5, 1950, by the Blue Dolphin. » Fish. Ind. U. S., Sect. 1, 1884, p. 190. « Moreau, Hist. Nat. Poissons France, vol. 3, 1881, p. 288. » 6,614 pounds, for example, were landed from the English Channel In 1932. For further details as to landings from the various statistical areas In the eastern Atlantic, see Thompson and Van Cleve, Rept. 9, International Fish- eries Comm., 1936, p. 21. M New Englands Prospect, 1634, p. 37. FISHES OF THE GULF OF MAINE 255 these of little esteem, except the head and finnes, which stewed or baked is very good; these halli- buts be little set by while basse is in season." They seem to have maintained their numbers there down to the first quarter of the nineteenth century, when contemporary writers described them as extremely numerous in Massachusetts Bay and along Cape Cod, in fact around the whole coast line of the Gulf of Maine. And they were dis- covered in abundance on Nantucket Shoals, on Georges Bank, on Browns Bank, and on the Seal Island ground as soon as fishing was regularly undertaken offshore. The cod fishermen of those days looked upon them as a nuisance, seldom worth bringing to market. And "It was the practice of the fisher- men when halibut were troublesome to string them on a line and hang them over the stem of the vessel."60 But a demand for halibut developed in the Boston market sometime between 1820 and 1825, and they have been pursued relentlessly ever since then, first inshore and then farther and farther afield. The Massachusetts Bay — Cape Cod region yielded large numbers of these great fish during the early years of the fishery. Four men, for instance, are reported as having caught 400 in two days off Marblehead in 1837, while a party of equal size is said to have landed 13,000 pounds off Cape Cod in three weeks. And it was discovered some time prior to 1840 that halibut congregated in winter in the 25-30 fathom gully between the tip of Cape Cod and Stellwagen Bank. However, a shrinkage in the supply had been noticed along shore even before 1839, for we find halibut de- scribed in that year (in the Gloucester Telegraph) as "formerly" caught along Cape Cod and in Barn- stable Bay. And they had been so nearly fished out in the Massachusetts Bay region by about 1850 that it no longer paid small boats to go there especially for them. Halibut held out better in the northeastern corner of the Gulf where there was not as ready a market for them as there was in Boston; Perley wrote of them as plentiful enough to be a plague to the local fishermen off Brier Island as recently as 1852. But it was not long thereafter before their numbers were greatly reduced there also. 1 Ooode and Collins, and Fish. Ind. U. S., Sect. 5, vol. 1, 1887, p. 29-30. The offshore fishery for halibut began about 1830, when cod fishermen brought word to Gloucester of a great abundance of them on Georges Bank,61 and they were caught there for a few years thereafter in numbers that seem almost unbelievable today. Thus we read of 250 caught in three hours; of vessels loaded in a couple of days; and of a single smack landing 20,000 pounds in a day. They were taken in great plenty on Nantucket Shoals, also, during this same period. But the supply seems to have dwindled suddenly, in the shoal waters both of Georges Bank and of Nantucket Shoals, and so permanently that few vessels went thither especially for halibut after 1850. Now forced to go further afield, the fishing fleet found that halibut were plentiful on the Seal Island ground; on Browns Bank; and in the Eastern Channel or gully that separates the latter from Georges Bank (localities which supplied the New York and Boston markets for the next decade). And in 1875 halibut fishing was extended to deeper water (100 to 200 fathoms) on the south- east slope of Georges Bank. But it was not long before all these grounds were fished out to the point where it was seldom possible to make paying trips to them for halibut alone. And for many years now, what few halibut have been caught in the Gulf of Maine have been taken incidentally. The history, in short, of the halibut fishery leaves no doubt that this species shows the effect of hard fishing sooner than most sea fish, it being possible to catch the majority of the stock on any limited area in a few years. Long liners and otter trawlers search all the good ground-fish bottoms of the Guff of Maine and its banks so thoroughly and con- stantly that the halibut never have a chance to reestablish themselves in any abundance on the shoaler grounds. They maintain their numbers better on the deeper slopes chiefly because they are subject to less intensive fishing there. It was fortunate for the fishing industry that the depletion of the Gulf of Maine of halibut was counterbalanced by the discovery of halibut in abundance along the deeper slopes of the banks to the north and east. And halibut fisherman sailing from Gloucester had begun resorting to the Grand Banks region by 1864-1866; to the west Greenland Banks by 1866; to the Magdalens by •i Ooode and Collins (Fish. Ind. U. S., Sect. 5, vol. 1, 1887, p. 3) have cot lected data on the Georges Bank halibut fishery and the former abundance of the fish there. 256 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE 1873; to the northern part of the Gulf of St. Lawrence near Anticosti by 1881 ; to Iceland by 1884.62 And the Gloucester vessels continued sailing to the Greenland Banks for halibut until the early 1880's. But by 1889 practically all the salt halibut that was landed in Gloucester, was being brought from Iceland. With salt fish in less and less demand, it became unprofitable, next, to sail so far afield. And it is many years, now, since any halibut fisherman from Gloucester has outfitted for Iceland. Long liners, out of New England ports, fished especially for halibut in the northern part of the Gulf of St. Lawrence until 1938; on the Grand Banks grounds until 1940 or 1941, when competi- tion with frozen halibut from the northwest coast had become severe, while the majority of fisher- men preferred to ship on otter trawlers, for com- fort and safety. Vessels continued long lining for halibut down the slopes of the Nova Scotian banks, and in the deep gullies between these, until the middle of the 1940's, but we have not heard of a vessel making a special trip from any New England port for these great fish, during the past few years. Although there is not one halibut hi our Gulf today, where there were hundreds or even thou- sands of them during the first quarter of the nine- teenth century, the geographical range of this noble fish is as extensive there as it ever was. Thus a few halibut are still caught along Cape Cod; in Massachusetts Bay (a few "chickens," 10 pounds and upward, are brought in every summer with some larger fish) ; along the Maine coast; and on all the offshore grounds. Rich, writing in 1929,63 listed 25 named inshore grounds off the coasts of Massachusetts and of Maine as still yielding a few halibut. We have enjoyed the acquaintance of several fishermen, especially in- terested in halibut, who treasure to themselves a hard-gained knowledge of particular spots, not too far offshore, where they are likely to catch one, in a day's pleasure fishing. And small groups of halibut accumulate occasionally on suitable patches of bottom; soon to be decimated, however, when then- presence is accidentally dis- covered. Thus, we knew of some 25 or 30 halibut, ranging from 40 to 110 pounds in weight, being caught within 1 to 3 miles of land, near Mount Desert Island, in 1930, in 10 to 15 fathoms of water. And one of 54 pounds was caught off Boston Harbor, from the steamer Westport, on June 24, 1951. M A catch of 9,500 pounds, or perhaps about 135 fish (assuming an average weight of 70 pounds) was reported off the coast of Maine in 1947 on hand lines, while eleven fish (largest 125 pounds) had been caught inshore, off Casco Bay, by local fishermen, up to the last week in May 1951.65 And many other instances of this sort might be quoted, no doubt, were our knowledge sufficient. Halibut are also caught fairly regularly still, about Grand Man an (4,700 pounds reported thence in 1947), but only occasionally about Campobello and near St. Andrews, and not at all along the north (New Brunswick) shore of the Bay of Fundy east of St. John. Small numbers occur, however, right up to the head of the bay on the Nova Scotia side.66 And there are enough of them off Brier Island at its mouth and on the fishing grounds along western Nova Scotia to have brought the landings for Digby County and for Yarmouth Coimty to 108,300 pounds in the year in question. The largest catches of halibut now made within the limits of the Gulf of Maine come from the Cape Sable-Browns Bank ground, from the deeper slopes of Browns Bank, from the deep gully that separates Browns from Georges, and from the eastern part and the deeper slopes of Georges, where otter trawlers are likely to pick up anywhere from 1 to 75 fish per trip. But not many are caught now on Nantucket Shoals where they were once so plentiful. In 1945 (most recent year for which detailed information is readily available for the coasts of Maine and Massachusetts), landings for the different parts of the Gulf, by United States and Canadian fishermen, were about as follows: off eastern Massachusetts, about 31,000 pounds; off western Maine, about 800 pounds; off central Maine, about 10,000 pounds; small banks in the inner west central part of the Gulf (Cashes, Fippenies, Platts), about 2,500 pounds; off eastern « See Collins, Fish. Ind. U. S., Sect. 5, vol. 1, 1887, pp. 43-89, and Scudder, Fish. Ind. I". S., Sect. 5, vol. 1, 1887, pp. 90-119, for historical surveys of the fresh and salt halibut fisheries from New England ports. " Rept. V. S. Comm. Fish. (1929) 1930, pp. 85-86, 90. M Reported in the Boston Clobe, June 25, 1951. « Reported in Salt Water Sportsman for May 29, 1951. M 11.300 pounds reported for Annapolis and King's Counties, Nova Scotia, in 1947. FISHES OF THE GULF OF MAINE 257 Maine about. 2,000 pounds; mouth of Bay of Fundy on New Brunswick side, about 700 pounds; Nova Scotian side of Bay of Fundy, about 45,000 pounds; off western Nova Scotia and Browns Bank (Canadian and United States vessels combined), about 73,000 pounds plus an indeter- minate part of the landings for Shelburne County, Nova Scotia, that may have come from Browns Bank; Georges Bank, about 65,000 pounds; South Channel, about 4,000 pounds; Nantucket Shoals, about 1,400 pounds; or a total of about 235,000 pounds that can be credited definitely to the Gulf. For some unknown reason, 1945 was a poor year; the Georges catch alone, for example, was about 110,000 pounds in 1946, about 211,000 pounds in 1947.67 And the yearly catch for the Gulf as a whole, by United States and Canadian fishermen combined, averaged about 316,000 pounds for the 0-year period 1941-1946, plus what fish may have been landed in Shelburne, Xova Scotia, from Browns Bank. Even so, the Gulf yields only about one-tenth as much halibut by weight today as it did, say, 30 years ago.68 We dare not guess in what degree this continued decrease has been a result of the progressive replacement of long-line fishing by otter trawling, of market conditions, or of a continuing decrease in the numbers of halibut. Halibut may have maintained their numbers somewhat better on the outer Nova Scotian Banks and slopes, which yielded about 3,400,000 pounds in 1934 (with Browns Bank); about 1,350,000 pounds in 1946.60 In the early days of the fishery, halibut were common hi the Gulf of Maine in water no deeper than they were farther north; near Anticosti in the Gulf of St. Lawrence for example, or near Miquelon, south of Newfoundland, where many were caught in 5 to 10 fathoms.70 A case is on record, for example, of a catch of 5 halibut, made in 1849, on one set of a long line with only 37 •' The otter trawlers that carried on investigations (or the Bureau of Fish- eries in 1913 took halibut on more than half their trips to Georges. Contrast this with a catch of 570 halibut by a long-liner on a patch of rocky bottom there in one day. in the early years of the Georges fishery! M The Gulf of Maine catch was nearly 3 million pounds in 1919. « Catch, Cape Sable to Cape Breton in 1940, about 50,000 pounds by United States vessels, about 1,300,000 pounds by Canadian vessels. For a general survey of the catches of halibut in both sides of the Atlantic, for 1934, see Thompson and Van Cleve, Rept. No. 9, International Fisheries Commission, 1930, p. 21. ™ Goode and Collins, Fish. Ind. U. S., Sec. 5, vol. 1, 1887, p. 17. hooks, in 7 fathoms, just off the mouth of Glouces- ter Harbor. A good many, too, were caught in those days on the southeastern part of Stellwagen Bank, where the depth (on the fishing grounds) ranges from 15 fathoms to about 30 fathoms. And many were reported as wintering in the gullies west of Stellwagen and between the latter and the tip of Cape Cod, in depths of 30 to 50 fathoms. Similarly, the early fishery also on Georges was on the shoaler parts of the bank in depths of 15 to 30 fathoms. And the early visitors to this ground describe the halibut, not only as schooling at the surface in pursuit of herring and launce (not an uncommon event in the Gulf of St. Lawrence and off Newfoundland when they are chasing capelin), but as often following their hooked companions up to the top of the water, so that more than one vessel made a good part of her fare by gaffing them alongside. The Nantucket Shoals halibut of old were like- wise in less than 30 fathoms depth, and when the licet first repaired to Browns Bank and to the Seal Island grounds they found halibut very plentiful in water but little deeper than that. In fact, it was not until 1874 or 1875 that the presence of this fish was suspected in the deeper gullies or on the offshore slopes of the banks below 100 fathoms. But it did not require many years of hard fishing to catch most of the halibut that were living in very shallow water, and so thoroughly that very few are now taken shoaler than 25 to 40 fathoms in our Gulf, while most of the halibut that are caught still on the offshore banks are from water deeper than 75 fathoms. All that has come down to us as to any general movements of the halibut in the Gulf of Maine during the days of their plenty there, beyond the prevailing tendency of the larger fish to work down deeper than the smaller (p. 250), is that some of them (though not all) worked inshore into shoaler waters for the winter, to work offshore again and deeper for the summer. But this offshore move- ment in whiter may not have extended far, or very deep, if it was to avoid low temperatures, for hali- but (or any other fish for that matter), that sum- mer inshore in shoal water where they would be most subject to winter chilling, need never move out for more than 60 miles or so off any part of the coast line of the open Gulf, nor descend deeper than about 70 to 75 fathoms, to find water per- manently warmer than 38°, except in the Bay of 210941—53- -IS 258 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Fundy in the coldest winters, or in the submarine embayment between Jeffreys Ledge and the coast. We have nothing to contribute as to present-day spawning of halibut within the Gulf of Maine. Of old, ripe females were reported on Georges in May and June, and have been, repeatedly, on the deeper slopes of the Nova Scotian banks generally, to the eastward, as well as off the Grand Banks and in the Gulf of St. Lawrence. But halibut smaller than a couple of pounds are seldom caught in the inner parts of the Gulf though "chickens" of 10 to 20 pounds are not uncommon there, there being a sharp contrast in this respect between the Gulf of Maine and the waters around Iceland, where Jespersen found an abundance of little fish of 8 to 10 inches. This, added to the fact that the inshore grounds were fished out so soon with little apparent tend- ency to recover when the fishery slackened, and that depletion by overfishing has not been ac- companied by any corresponding decrease in the average size of the fish that are caught, suggests that the halibut population of the inner parts of our Gulf always depended more on immigration from east and north of Cape Sable for its main- tenance than it did on local production. Fry may have been produced in greater numbers over the offshore slope of Georges Bank, where the Alba- tross III trawled two little halibut about 6 inches long, at 175 to 195 fathoms, on May 16, 1950. Importance. — The halibut, because of its pres- ent-day scarcity, is of only minor importance commercially in our Gulf; in 1947 the landings in New England, including what halibut were brought in from the Nova Scotian banks eastward from Cape Sable, amounted to only about 586,000 pounds, valued at $144,680. But the demand is always so good that all that are brought in are readily salable, and (being so large) each one that is caught is well worth saving. In the year in question (representative of present-day condi- tions), about one-fifth of the total New England landings, were caught on long lines 7l three- fourths by otter trawlers. The small remainder (10,000 pounds) were taken on hand lines, mostly by small-boat fishermen off the coast of Maine. We can only regret that there are not enough halibut inshore in our Gulf today to be of any general concern to anglers, for this is a very "sporting" fish as well as welcome on the table. Greenland halibut Reinhardtius hippoglossoides (Walbaum) 1792 Greenland turbot; Newfoundland turbot Jordan and Evermann, 1896-1900, p. 2611. Description. — This is a right-handed, large- mouthed flatfish (that is, it lies on its left side, with its eyes on its right side, and its abdomen at its right edge), with slightly concave tail, and symmetrical ventral fins like a halibut. In fact it so closely resembles the halibut that it might easily be taken for one were it not that its lateral line is nearly straight abreast of the pectoral fin, (arched in the halibut) and that its long fins (dorsal and anal) are of rather different shape (compare fig. 127 with fig. 123), though with about the same number of rays (about 100 dorsal and " Goode and Collins (Fish. Ind. U. S., Sec. 5, vol. 1 1887, pp. 10-18) have given a readablo account of the long-line fishery. •:' -^ Figure 127. — Greenland halibut (Reinhardtius hippoglossoides). From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 259 75 anal). Its mouth, furthermore, is larger, its eyes smaller relatively and its jaw teeth stronger, though the differences in these respects are not great enough to serve as useful field marks. It is yellowish or grayish brown, paler below than above but not white. Size. — This is one of the largest of the North Atlantic flatfishes, next to the halibut, growing to a length of about 40 inches and to a weight of 20 to 25 pounds. But fish caught about the Grand Banks weigh only from about 5 to 10 pounds. General range and occurrence in the Gulf of Maine. — This is a fish of the Arctic and subarctic Atlantic. It is taken from northern Norway and northern Iceland to the Faroe ridge, and to south- west of Iceland 72 as a stray. It supports a fishery off west Greenland that is important for the Eskimos.73 In the west considerable numbers are taken off the south coast of Newfoundland,74 also on the Grand Banks, hence it is to be expected along outer Labrador, though it has not been re- ported thence as yet. Odd specimens are to be expected here and there in the Gulf of St. Law- rence too, for it has been taken near the Biological Station at Trois Pistoles.75 " Norman (Ann. Mag. N'at. Hist., ser. 9, vol. 13, p. 539) reports a single specimen taken southwest of Iceland. " See Jensen (Meddel. Dansk. Komm. Havundersdgelser, Ser. Fiskerl, vol. 7, No. 7, 1927) for a general account of the Greenland halibut off west Greenland. '< According to Goode (Fish. Ind. U.S., Sect. 1, 1884, p. 198) long liners have reported it as "very abundant" in and of! Fortune Bay, in 60 to 300 fathoms. " Vladykov and Tremblay, Natural. Canad., vol. 62 (Scr. 3, vol. 6), 1935 p. 82. It is described as "not uncommon" off Canso, Nova Scotia,76 and as occasionally brought in from the more northerly of the Nova Scotian fishing banks long ago.77 Our only reason for mentioning it here is Goode and Bean's78 statement that "fishermen take them frequently in the gully between La Have and Georges Bank at depths greater than 200 fathoms." This has been corroborated by the capture of 16 specimens, at 300 to 530 fathoms, along the slope of La Have Bank to the southwestern slope of Georges Bank by the Cap'n Bill II, in July 1952. It has no real place in the Gulf of Maine fish fauna. American dab Hippoglossoides platessoides (Fab- ricius) 1780 79 Canadian plaice; Long rough dab Jordan and Evermann, 1896-1900, p. 2614. Description. — The most obvious distinctive char- acters of the American dab are that it is right- handed and large-mouthed like the halibut, but with a rounded tail instead of concave, and with the lateral line nearly straight instead of arched; it is the only Gulf of Maine flounder in which these characters are combined. Our only other large-mouthed flat-fishes with rounded tails (the » Cornish, Contrib. Canadian Biol. (1902-1905) 1907, p. 90. " Jones, Proc. Trans. Nova Scotia Inst., Sci., vol. 5, Pt. 1, 1879, p. 92. " Amer. Jour. Sci. Arts, Scr. 3, vol. 17, 1879, p. 40. n Various other common names are applied to this fish In different seas. It is usually termed "Long rough dab" In England and is so listed in British fishery statistics. It is not the "plaice," or the "dab" ol Europe. Figure 128. — Canadian plaice, or Dab {Hippoglossoides plattessoides) , La Have Bank. From Goode. Drawing by H. L. Todd. 260 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE sand-, summer- and four-spotted flounders, pp. 290, 267, and 270) are left-handed, and the wide- gaping jaws readily distinguish the American dab from the various small-mouthed flounders. It is a comparatively broad (really deep) flounder (about two and one-half times as long to base of caudal fin as it is broad) , more roimded in outline than the halibut, with pointed nose, mouth gaping back to abreast of the middle of the eyes, and with one irregular row of sharp conical teeth in each jaw. The free edges of the scales on the entire eyed (upper) side of its body and of its head are serrated with sharp teeth, which give the fish a characteristic rough feeling when handled, but the scales of the blind (lower) side are smooth-edged except on the rear part of the body and along the bases of the fins. The dorsal fin (76 to 96 rays) originates in front of the middle of the left eye and the anal fin (64 to 77 rajrs) arises slightly in advance of the base of the pectorals. Both of these long fins taper toward the head and toward the tail, and there is a short, sharp, spine (the prolongation of the post abdominal bone) pointing forward close in front of the anal fin. The pec- toral fin on the eyed side usually (not always) has one or two more rays than its fellow fin on the blind side, and is longer and more rounded, but the two ventral fins, which are close in front of the anal fin though entirely distinct from it, are alike in size, shape, and location. The margin of the caudal fin is always convex, either rounded or with its middle rays so much the longest as to form a blunt angle. The lateral line on the eyed side is more clearly evident on the dab than on most of our flatfishes, and it is straight from end to end, except for a slight arch over the pectoral fin. Color. — Dabs run more uniform in color than most of our smaller flatfish, ranging from reddish to greyish brown (darker or paler) above and pure or bluish white below. The tips of the rays of the two long (dorsal and anal) fins are white. On one specimen we saw the right edge of the eyed side was white (like the blind side) from the gill opening to the rearmost ray of the ventral fin but this is unusual. Small fish are usually marked with three to five dark spots along each edge of the bod}r; large ones are occasionally, though they are plain colored as a ride. Size. — Adults measured by Welsh off Cape Ann ran from about 12 inches to 24 inches, and few of those that are caught in our Gulf are longer than 2 feet. Nova Scotian fish measured by Hunts- man 80 ran from 12 to 24 inches in length, while fish caught in the colder waters off Newfoundland averaged 18 inches.81 The largest dab recorded from American waters, taken near Sable Island, May 1939, was 32 K inches long and weighed 14 pounds.82 The next largest, taken in 90 fathoms on the northern edge of Georges Bank, November 1951, was 29 inches long.83 According to Huntsman, Nova Scotian fish average about half a pound at 12 inches, l){ pounds at 16 inches, 1% pounds at 18 inches, 2% pounds at 20 inches, 4 pounds at 22 inches, and 6 pounds at 24 inches. Massachusetts Bay fish are about equally heavy at corresponding lengths. And a 16-inch fish from Georges Bank that we measured weighed 1 pound 5 ounces; two fish of 18% inches weighed 1 pound 13 ounces, and 2 pounds, respec- tively; one of 19/2 inches weighed 2 pounds 8 ounces, and one of 29 inches weighed 9 pounds 6 ounces. This flatfish tends to differentiate into local races in different seas. Thus the fin rays are more numerous on the average in fish from high latitudes than in those from low latitudes, while the body is relatively wider in fish caught off Greenland and off America than in those from Scandinavia and from the North Sea. But these characters vary so -widely even in limited areas that the Arctic-American and European species (plates- soides and limandoides) have been united by common consent long since, and we doubt whether the corresponding "varieties" still recognized by several recent authors will stand the test of time. Huntsman's statement that the dorsal rays aver- age more numerous in dabs from Bay of Islands, Newfoundland, than in those caught on the New Brunswick shore of the Gulf of St. Lawrence, with Welsh's note of a variation of 7 in the number of dorsal rays and of 6 in the anal rays in one lot of fish caught off Gloucester, illustrates this vari- ability. Notwithstanding the low latitude of the locality of capture (about 42° 30' N.), this same lot contained a specimen with the largest number of fin rays yet reported (96 dorsal and 77 anal). All we dare say until many more specimens are examined is that hereditary local races may perhaps exist off different parts of the American » Bull. No. 1, Biol. Board Canada, 1918. p. 10. *' Frost, Res. Bull. 4, Newfoundland Dept. Nat. Resources, 1938, p. 8. « McKenzie, Proc. Nova Scotian Inst. Sci., vol. 20, 1940, pp. 43-44. " We measured this dab, taken by Capt. Arthur Nelson of the Eugene H. who also caught several others, 27 to 28 inches long on this same trip. FISHES OF THE GULF OF MAINE 261 shore line, and that the growth marks on the scales, in relation to the length of the fish, may give a clue to the local origin of a given specimen, for it seems that the rate of growth is governed by the temperature of the water (p. 263). Habits. — Dabs are bottom fish like other flat- fishes. But the}' must rise some distance from the ground on occasion, and move about to a con- siderable extent to account for the capture of so many in gill nets (p. 264). We once caught one a foot long in a tow net at least 5 to 10 fathoms above the bottom off Ipswich Bay, where the water was about 50 fathoms deep. Like some other flatfishes, they avoid rocky or hard bottom, preferring a fine, sticky but gritty mixture of sand and mud, such as floors much of the Gulf between the hard patches, from the 20- fathom contour out to the 100 fathom contour. And they are also to be caught in numbers on the soft oozy mud of the deeper basins in the western side of the Gulf, as pointed out below (p. 264). In one part of their range or another, they are found from tide line down to as deep as about 390 fathoms (700 meters). This is an arctic-boreal species in its tempera- ture relations, reaching its highest development in water of 35° to 45° F.; able to live, however, in the lowest polar temperatures (29° to 30°); and finding the upper temperature limit to its regular occurrence at about 50° to 55° F. In different seas it lives through a wide range of salinity, from 30 per mille or lower in the Baltic to upwards of 34 per mille in the open Atlantic. So far as we are aware, it is never found in water which could be described as brackish along the coasts of New England or of the Maritime Prov- inces. But R. II. Backus informs us that the Blue Dolphin found it in brackish water (salinity 23 per mille) at the west end of Lake Melville, Labrador. Huntsman's84 statement that it feeds on minute planktonic plants (diatoms) at first, but on cope- pods as it grows larger and more active is our only information as to the diet of the young fry in American waters, while they are drifting near the surface. When they first take to the bottom they eat small shrimps and other Crustacea of various sorts. But they turn (as they grow) to a diet consisting chiefly of sea urchins, sand dollars, and brittle stars, as proved by the contents of their stomachs, though they also take various shrimps, hermit and spider crabs and other crus- taceans, mollusks, worms and ascidians (sea squirts), in fact, practically any bottom living animals that are small enough for them to devour. Occasionally they catch small fish. They do not bite a baited hook as readily as various other ground fishes, partly, no doubt, because they are sluggish fish, but partly, we believe, because the clams, cockles, and herring that are usually used for bait are not their favorite food. Still, considerable numbers are caught on hand and long lines. All the large predaceous fish that feed near bottom probably prey more or less upon them, and halibut no doubt destroyed great numbers of them in the Gulf of Maine formerly. But the adults can have no serious enemy in our Gulf today except large cod and perhaps the spiny dogfish. In more northern seas Greenland sharks prey regularly on them. Smitt and Huntsman both speak of the numbers of round worms to be found in the intestines and body cavity of the dab, and its gills are sometimes attacked by parasitic copepods. While the young are drifting near the surface (p. 262), they share in the same involuntary journeyings as other fish fry do, that are spawned at the same place and time. But it is one of the more stationary fishes from the time it seeks bot- tom. It has been said to work inshore more or less in winter, though not on very definite evidence, and it may congregate on definite grounds for spawn- ing, though this is yet to be proved. But it is certain that they are to be caught at any season of the year wherever they are plentiful. And Huntsman,85 who has paid special attention to this fish, believes that it "remains pretty much in the same; place from season to season and year to year. Perhaps in the course of years it may shift a few miles." Individual females produce 30,000 to 60,000 eggs, according to size. The eggs are buoyant and have no oil globule, but they have a trans- parent (perivitelline) space around the yolk so broad that they are not likely to be confused with those of any other Gulf of Maine fish. »< Bull. Biol. Board Canada, No. 1. 1918, p. 15. " Bull. Biol. Board Canada, No. 1, 1918, p. 18. 262 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 131. — Larva (European), 9 mm. After Ehrenbaum. Figure 129. — Egg (European). After Cunningham. Figure 132. — Larva, 14.5 mm., off Massachusetts Bay. Figure 130. — Larva, just hatched, 4 mm. (European) After Mcintosh. Figure 133. — Larva, 22.5 mm., off Cape Cod. Canadian plaice, or Dab {Hippoglossoides plaltessoides) . This space is formed by the entrance of water between the egg proper and its covering mem- brane, after the eggs are shed, and it about doubles the total diameter of the egg. The eggs we have taken in the Gulf of Maine have averaged about 2.5 mm. in diameter, but they have been reported as small as 1.38 and as large as 3.2 nun. in other seas, depending on the breadth of the perivitelline space. Incubation occupies 11 to 14 days at a tempera- ture of 39° F., and it seems that the eggs gain weight as development proceeds, for Huntsman found, in the Gulf of St. Lawrence, that the newly spawned eggs floated at the surface, but that eggs nearly ready to hatch drifted suspended at a depth of some 10 fathoms. We have no first-hand information to offer on this point. During the development of the egg, minute black and yellow pigment cells are scattered over the embryo, not aggregated into any diag- nostic clusters. But the pigment gathers in five definite groups very soon after hatching (which takes place when the larvae are 4 to 6 mm. long) ; one on the gastric region, one about the vent, and three behind the vent; a pattern similar to that of the larval witch flounder (p. 287) . The yolk is absorbed about 5 days after hatch- ing, when the larva has grown to 6.2 to 7.5 mm. in length. The caudal rays appear shortly after this, the dorsal and anal rays at about 11 to 12 mm., and the three vertical fins are differentiated at about 15 to 18 mm. By this stage the body has begun to assume the deep but very thin form characteristic of all young flounders, while the jaws have developed sufficiently to show that the little fish belongs to one of the large-mouthed species. The left eye may commence its migra- tion when the larva is about 20 mm. long, while Welsh found it visible above the outline of the snout in Gulf of Maine specimens of 24 mm., and almost at the dorsal edge at 34 mm. But larvae as long as 35 mm. may still be symmetrical in other seas. The only other Gulf of Maine species for which the larval dab might be mistaken (except in its very earliest stages) are the witch flounder and the halibut; but the witch is longer at corresponding stages of development, but ■with the distance from snout to vent proportionately much shorter, and the outlines of throat and abdomen are suf- ficiently different to distinguish the dab from the halibut (p. 253). The young dab drifts freely up to the time of its metamorphosis, as the young of most sea fishes do; close to the surface at first but sinking deeper as it grows, until it seeks the bottom finally. FISHES OF THE GULF OF MAINE 263 Welsh's observations suggest that this takes place, in our Gulf, when the little fish are about 1% to 1% inches long, with their metamorphosis already complete, their body scaly, and their eyed side densely pigmented. But there is wide variation in this respect. And European authors report that the fry may take to the bottom even before the left eye has completed its migration around the head. The period occupied in larval growth and in metamorphosis varies with temperature. Proba- bly it covers three to four months in the Gulf of Maine, where we have taken the pelagic larvae as early in the season as May 26 and as late as August 2. The little fish grow to a length of 2 to 3 inches by their first winter, with their exact size then de- pending upon how early in the season they are hatched, and probably on the temperature in which they live. And they average about 3 inches long 86 when they are one year old. Thus it may be assumed that bottom stages 2% to Z% inches (69-80 mm.) long that we have trawled off Cape Cod, on May 1, were about one year old; others of Z% to 4% inches (85-118 mm.) that we have trawled in July and August off Mount Desert, in the deep gully to the westward of Jef- freys Ledge, on Cashes Ledge, and on the edge of Stellwagen Bank were between 1% and 1% years old; and that those of 8 to 10 inches were 2% to 2% years old. Subsequent growth is more rapid in higher temperatures than in lower, throughout the temperature range favorable to this particular flatfish. Huntsman,87 for example, has found that it takes only 3 to 5 years for dabs to grow to a length of 12 inches in Passamaquoddy Bay, where the bottom water at 15 to IS fathoms warms to about 49° to 51° F. in August, but that it requires 4 to 6 years in the open Bay of Fundy, where the bottom temperature in summer is somewhat lower (45°-48°); 6 to 9 years in the cooler water (about 38°) of Chedabucto Bay, eastern Nova Scotia; and upwards of S years in the still lower temperatures (colder than 35°) of the Gulf of St. Lawrence. On this basis, dabs living on the shoaler parts of Georges Bank, and as shoal as 15 fathoms or so in coastwise waters from Cape Cod to Cape Eliza- " Huntsman, Bull. Biol. Board Canada, No. 1, 1918. " Bull. Biol. Board Canada, No. 1, 1918, p. 23. beth, probably grow about as fast as the Passa- maquoddy Bay fish, i. e., they may reach a length of 15 inches in 5 years or even sooner, gaining something like 4 ounces in weight yearly. Those in the eastern side of the open Gulf of Maine may be expected to grow about as fast as those in the Bay of Fundy, but somewhat more slowly there if they are living as deep as 50 fathoms, though not so slowly as in the Gulf of St. Lawrence. Some individuals may become sexually mature when only 6 inches long, probably all of them do so by their third year; and they are known to live to an age of 24-30 years, perhaps longer, at least in the Gulf of St. Lawrence. In general, females grow faster than males. Huntsman has also found, in the Gulf of St. Lawrence, that a majority of the fish of 3 years and younger were males, but that females out- numbered the males among the older fish, while all of those 14 years old and older that he saw were females. We have no explanation to offer for this (apparently) higher mortality rate for the males than for the females among the older fish. General range. — This is a very common fish on both sides of the North Atlantic, where its range parallels that of the cod, except that it does not c\l end as far south and west along the American seaboard. It is found in abundance along the outer coast of Labrador, southward from Hamilton Inlet, where (Frost88 writes) they are so abundant locally that a 5-minute haul with a torn trawl yielded 50 (at lat. about 54°) in Newfoundland waters in general; on the Grand Banks, including the eastern edge; " in the Gulf of St. Lawrence as a whole, and thence westward and southward to Cape Cod, from close inshore out to the 100- f a thorn contour. Westward from Cape Cod, a few are caught in the Woods Hole region; off Marthas Vineyard; and off Narragansett Bay which marks their western limit in general. The most southerly and westerly record with which we are acquainted is of one 15% inches long that was caught off Montauk Point, N. Y., in 112 fathoms, February 6, 1930.90 81 Res. Bull. 4, Newfoundland Dept. Nat. Resources, 1938, p. 8. R. H. Backus informs us also that the Blue Dolphin collected them at various local- ities as far north as the northern shore of Hamilton Inlet (lat. 54°30' N.), but did not take any farther north, in spite of extensive collecting. " Reported in abundance down the eastern edge of the Grand Banks, in the 20th Rept. Dept. Fish. Canada (1949-1950) 1951, p. 36. 90 We find no other credible records from New York or from New Jersey, those mentioned by DeKay being market fish which might have oome from anywhere to the eastward. 264 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE It is common in west Greenland waters, as far north as Upernavik near the Arctic Circle, in lat- itude about 72° N.91 And it ranges in European waters from Iceland and Spitzbergen southward to the North Sea, where it is an important commercial fish, and to the west Baltic; the English Channel is the southern boundary to its regular occurrence. Occurrence in the Oulj of Maine.92 — This is not as familiar a fish as are the winter and smooth flounders (pp. 276 and 283), for it is not common in water shallower than 15 to 20 fathoms. But it is probably the most abundant of all Gulf of Maine flatfishes at depths greater than 30 to 50 fathoms, except, perhaps, the witch (p. 285). Thus they are recorded from Provincetown ; from Massachusetts Bay; off Cape Ann; on Stellwagen Bank, where we have hand-lined a number of them in 25 fathoms; in Ipswich Bay; near Boon Island; off Cape Porpoise; off Casco Bay; on Cashes Ledge, where we have trawled both young and adults; off Seguin; south of Monhegan (we trawled them at the last four localities on the Grampus) ; close in to Little Duck Island, off Mount Desert; in Passa- maquoddy Bay; in St. Mary Bay; and right up to the head of the Bay of Fundy. In fact, they are to be caught all around the inner parts of the Gulf wherever the water is more than 15 fathoms deep or so, and where the bottom is smooth. Trawlings, too, by the Albatross II and by the Atlantis have shown that they are generally dis- tributed throughout the basin of the Gulf down to 120 fathoms. This, indeed, was the only flatfish, other than the witch (p. 288), that was taken by the Atlantis on the soft mud bottoms off Cape Cod, west of Jeffreys Ledge, or off Moimt Desert, at 66 to 105 fathoms during her experi- mental trawlings for the edible shrimp (Pandalus) in August 1936.93 Dabs are widespread on Georges Bank also, for they were reported at many localities there by representatives of the Bureau of Fisheries in 1913, while we have seen catches of up to 100 per trawl haul on the northern edge of Georges, in 60 to 100 fathoms of water. They are so plentiful along the 50-100 fathom zone on the northern edge of Georges Bank that draggers fishing there during 1951-1952 were making catches averaging about 5,000 pounds s' For an account of it in west Greenland waters, see Jensen (Meddel. Dansk Komm. Harandersdgelser, vol. 7, No. 7, 1925, p. 24). « Huntsman (Bull. 1, Biol. Board Canada, 1918) gives an interesting account of this fish in Canadian waters. « Bigclow and Schroedcr, Biol. Bull., vol. 76, 1939, p. 308. per day. A good example of their numbers there is furnished by the dragger Eugene H of Woods Hole which brought in catches of 10,000 to 25,000 pounds of dabs, fishing in 75 to 95 fathoms, through- out the period August 1951 to January 1952. Many of these fish were large, ranging from 4 to about 9 pounds in weight. And in this same region, in the spring, they appear to be plentiful in water much shoaler, for Capt. Arthur Nelson of Woods Hole reports a catch of 18,000 pounds taken in 25-30 fathoms in four days' fishing early in May 1952. Also, we have the definite evidence of commercial catches, as well as of newly spawned eggs taken in our tow net, that dabs are plentiful on Browns Bank also. Huntsman has calculated from fishing experi- ments that they are about one-tenth as numerous as cod in the Gulf of St. Lawrence. No general estimate of this sort is yet possible for the Gulf of Maine. But catches in gill nets (gear not very well adapted for flounder fishing) of 76 dabs to 1,055 haddock, 51 cod, 20 pollock, and 39 rose- fish near Boon Island on March 30; of 125 dabs to 40 other flounders, 89 cod, and 113 haddock in part of the net at the same locality on April 20; and of many dabs, but more cod and haddock, on May 3, 1913, 94 are pertinent here. This flatfish is often found in veiy shoal water in colder seas. They are often seen under wharves around Newfoundland, for example, according to Frost.95 And some are seined right on the beach 96 on the West Greenland coast. But we have never seen or heard of an adult specimen caught in less than 10 fathoms of water in the Gulf of Maine, probably because of the high summer temperatures of the shoaler waters, and they are the most plenti- ful in 15 to 60 fathoms there (in our experience). At the other extreme, 120 fathoms is the deepest definite record for the Gulf of Maine with which we are acquainted; hence this may be set as the lower limit to their occurrence there in any num- bers, which, by report, applies to the whole Ameri- can coastline, including the Scotian banks and the Grand Banks region. This preference of the dab for moderately deep water in the southern part of its range bars it from most of the Gulf of Maine harbors and river •' Recorded by Welsh. •s Research Bull. 4, Newfoundland Dept. Nat. Resources, 1939, p. 8. "Jensen, Meddel. Dansk. Komm. H avunderso'gelser, Ser. Fiskeri, vol. 7, No. 7, 1925, p. 24. FISHES OF THE GULF OF MAINE 265 mouths, which are such favored haunts for the winter flounder. But it enters the deeper estuaries and passages between the islands in the north- eastern part of the Gulf, those near Mount Desert, for example, Passamaquoddy Bay, and St. Mary Bay. "We hesitate to draw any definite conclusions from published statistics of the landings of "dab" 97 as to the regional abundance of this particular flat- fish in our Gulf, partly because of the likelihood that other flatfish may appear under this name, or dabs under some other name, and partly be- cause only a few of the otter trawlers fish in the deeper basins of the Gulf where dabs are known to be plentiful. The returns for 1945, if taken at face value, show about 48,000 pounds landed from off east- ern Maine; about 586,000 pounds from off central Maine; about 311,000 pounds from off western Maine; about 43,000 pounds from small grounds in the west central part of the Gulf; about 897,000 pounds from off eastern Massachusetts; about 8,000 pounds from Nantucket Shoals; about 910,000 pounds from the South Channel and Georges Bank combined; about 48,000 pounds from Browns Bank; and about 40,000 pounds from off western Nova Scotia (by United States fisher- men) ; or a total of some 2,890,000 pounds. It w as not until 1946 that the dab was listed (as "Cana- dian plaice") in the Canadian fisheries statistics for Nova Scotia; in that year landings for western Nova Scotia (Yarmouth County) were about 140,000 pounds, and about 41,000 pounds for the Nova Scotian side of the Bay of Fundy (Digby County). The presence of dabs or Canadian plaice of catchable sizes in the Bay of Fundy in general, and in Passamaquoddy Bay in particular, is interesting as evidence that this is not so stationary a fish there as it seems to be elsewhere, for none are reared there so far as is known (p. 266), so that the main- tenance of the local stock appears to depend on immigration from outside. Huntsman's observa- tion is interesting, too, that large ones form a much smaller proportion of the population in Passama- quoddy Bay and in the Bay of Fundy than they do in the Gulf of St. Lawrence. And it seems, simi- larly, that large ones are less plentiful relatively in •' It is only during the past few years that the landings of this particular flatfish have been reported separately, as "dab." Passamaquoddy Bay than they are in the western side of the open Gulf of Maine. The death rate may be higher in Passamaquoddy waters, as Huntsman has suggested, or it may prove that the fish tend to work out from there into the open Gulf as they advance in age. The dab is a spring spawner on both sides of the Atlantic, as is well known. The earliest date at which we have taken its eggs in our tow net in the Gulf of Maine has been March 4 (in 1920), off Casco Bay. We have also found the eggs on Browns Bank on the 13th, while Welsh records large female fish, half spent and with eggs exuding, as well as males with running milt, on the 14th of March, near Cape Ann, in 1913. But other fish of both sexes taken with them were unripe still, evidence that spawning is not general until the last of March or first part of April. Dab eggs have appeared regularly in our towings in April (twice in great numbers, namely off Seguin Island on the 10th and off Mount Desert Island on the 12th in 1920). Spawning continues unabated throughout May, when eggs were taken at nearly all our towing stations in 1915. And April and May similarly cover the height of the spawning season in the Bay of Fund}7, according to Hunts- man.98 Our latest seasonal record has been for a single egg, on the 14th of June in 1915. The dab spawns chiefly during May and June on the banks off Cape Breton and in the southern part of the Gulf of St. Lawrence: until the end of July on the southern part of the Newfoundland Banks (a few eggs were found by the Canadian Fisheries Expedition); until fall around the south- eastern and eastern coasts of Newfoundland, and along the outer coast of Labrador, according to Frost. And the eggs are reported from May into July off West Greenland, by Jensen. Tt spawns somewhat earlier in the North Sea than in American waters; i. e., from mid-January till May with the climax in March and April. Huntsman also remarks that there is a difference in the breeding season according to the depth of water, those living shoalest commencing to spawn the earliest, as the vernal warming of the water makes itself felt from above. But we have no clear evidence on this point to offer for the Gulf of Maine. ' Bull. 1. Biol. Board Canada, 1918, p. 14. 266 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Our egg records, added to Huntsman's observa- tions, show that the dab spawns all around the Gulf of Maine, from Cape Cod on the west to Cape Sable on the east, including the Bay of Fundy, and from close inshore out to the 50-fathom contour. It also spawns on Browns Bank (p. 265), and, while we found no eggs on Georges Bank either in February, March, April, or May, of 1920, the fish is so common there and so stationary in general that it is likely that we simply missed its eggs, either by a failure to tow over the precise spawning localities or by timing our visits between the waves of production. Dabs also spawn abundantly on Sable Island Bank (no doubt on all the other Nova Scotian Banks); off Cape Breton; in the shoaler parts of the Gulf of St. Lawrence;99 throughout the general region of the Grand Banks; off the east coast of Newfoundland; along the outer coast of Labrador to Hamilton Inlet at least;1 and as far north along the west coast of Greenland as the species is known to exist, as is proven by the presence of its eggs in the water there in quantities.2 Although the dab is rather a deep-water fish compared to most of the other flatfishes that are common in the Gulf of Maine, it is doubtful whether it ever spawns at depths much greater than 50 fathoms in the Gulf, for we have few egg records from more than a mile or two outside the 50-fathom curve, while these few have been based on only one or two eggs each. And we have trawled spawning females off Mount Desert, in 20 fathoms. This concentration of our egg catches inside the 50-fathom contour implies that the dabs that live deeper in our Gulf tend to work up into shoaler grounds to spawn. Beyond this, there is no reason to suppose that the}7 gather in any definite localities for the purpose. The temperatures and salinities in which the eggs are produced can be stated rather definitely for the Gulf of Maine because the dab lies close to the bottom, if not actually on it. The earliest spawning takes place at nearly the minimum temperature for the year, averaging about 37° for all the March and April stations where eggs were '• Dannevig, Canadian Fisheries Expedition (1914-15) 1919, p. 18, figs. 11, 12, and 13. ' See Frost, Res. Bull. No. 4, Newfoundland Dept. Nat. Eesources, 193S, chart 2, for the regional and seasonal distribution of dab eggs in Newfoundland and Labrador waters. s Jensen, Meddel. Dansk. Komm. Havunders0gelser, Ser. Fiskeri, vol. 7, no. 7, 1925, p. 24. taken. And while the water warms to 41°-43° F. by late May and early June at the depths known to be inhabited by the ripe fish, we have not found its eggs where the bottom temperature was higher than about 40°. Thus the optimum for breeding may be set at 37°-40° for the Gulf of Maine as a whole. Dabs spawn freely in 31°-32° off Cape Breton, and even in water as cold as 29.3°-32° in the Gulf of St. Lawrence, in New- foundland waters and northward, as well as along the West Greenland coast, proving that the lowest polar temperatures are no bar to the ripening of its sexual prwlucts. Neither does the distribution of the bottom stages suggest that warmer water is needed for the survival of the resultant larvae. In the Gulf of Maine the dab spawns in rela- tively low salinities, the range there being only from about 31.8 per mille to about 32.8 per mille at the bottom at the stations where eggs were taken in any number. But it does so in con- siderably more saline waters in the other side of the Atlantic, generally speaking. Although this flatfish spawns so generally throughout the whole area that it inhabits, there is evidence that different regions differ in their suitability as nurseries, either for its eggs or for the larvae. The southwestern part of the Gulf of Maine must be favorable in this respect, for we have taken larval dabs at 14 stations there, most of these off the Massachusetts Bay region. And they have also been taken at various localities off the southeast coast of Nova Scotia; on the Newfoundland Banks; in the Gulf of St. Lawrence; along the east coast of Newfoundland: in the Strait of Belle Isle; and northward for some dis- tance along the outer coast of Labrador. But it seems that reproduction does not succeed in the Bay of Fundy, for neither the larvae nor the young fry have ever been found there, although dabs spawn there and the eggs develop, at least partially. Failure to find any dab larvae off the coast of Maine east of Penobscot Bay, though eggs are produced there in abundance may be due to the prevailing drift from northeast to southwest along this part of the coast, because of which buoyant eggs produced there are likely to hatch a considerable distance to the west of where they were spawned. The influence that this drift may have on the distribution of larval fish in the Gulf of Maine offers a fertile field for future study. FISHES OF THE GULF OF MAINE 267 Commercial Importance. — This is an excellent pan fish, but there is no special demand for it in New England markets, as distinguished from other flat fishes of about the same size. If the landings reported as "dab" do not include any significant proportion of other flounders, and if most of the dabs that are taken are reported under that name, the yearly catch in the Gulf by United States fishermen ranged between about 2,700,000 pounds and about 4,400,000 pounds for the period 1942 to 1947, averaging about 3,600,000 pounds. In 1946 Canadian fishermen brought in an additional 181,200 pounds from the eastern side of the Gulf and from the Bay of Fundy, plus an indeterminate amount landed in Shelburne County from Cape Sable to the Yarmouth County line.3 We have no doubt that the catch could be increased greatly in our Gulf if any special demand were to develop for dabs. The dab lives too far out from the land, on the whole, and too deep, and it does not bite eagerly enough for it to be of any interest to anglers along our shores. Summer flounder Paralichthys denlatus (Linnaeus) 1766 Flounder; Fluke; Plaicefish Jordan and Evermann, 1896-1900, p. 2629. Description. — The summer flounder is left- handed; that is, it lies on the bottom on its right 1 The landings for that year were 60,100 pounds /or Shelburne County as a whole. side, with its eyes on its left-hand side, and its abdomen is on its left edge as it rests on the bot- tom, which differentiates it at a glance from the American dab (p. 259). It is large-mouthed, like the sand flounder, which is similarly left-handed (p. 290) ; but its two ventral fins are alike and each of them is separated from the long anal fin by a considerable space, whereas the upper left-hand ventral fin of the sand flounder is continuous with the anal fin. The only Gulf of Maine flatfish with which the summer flounder shares its left-handed- ness, large mouth, and symmetrical ventral fins, is its close relative, the four-spotted flounder (p. 270) , but the color pattern of the latter is distinctive (p. 270) and it has fewer fin rays. The summer flounder is one of our narrower flounders. Its dorsal fin (85 to 94 rays) originates opposite the forward margin of the eye; its anal fin has from 60 to 73 rays; the margin of its caudal is rounded, and its pectoral fins and ventral fins are smaller than those of the dab, relatively. Color. — It has long been known that flatfishes are generally dark on a dark bottom and pale on a pale one. Perhaps the summer flounder is the most variable in color of all our local species and the one which adapts its pattern the most closely to that of the ground on which it lies. It is white below and of some shade of brown, gray, or drab above, like most flatfishes. But it can assume a wide range of tints, from nearly white on white sand through various hues of gray, blue, green, . ,,<{ ■ ■ Figure 134. — Summer flounder (Paralichthys denlatus), Maryland. From Jordan and Evermann. Drawing by A. H. Baldwin. 268 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE orange, pink, and brown to almost black.4 Its upper surface is variegated witb pale and dark, as a rule, with the pattern fine or coarse accord- ing to the bottom, and it may or may not be marked with small eyespots of a darker tint of the general ground color. Mast's experiments show that it is slower in adapting its coloration to the actual colors of the bottom than to the general pattern, and also that it responds more rapidly to yellows and browns than to reds, greens, or blues, on which the adaptation may not reach its maximum for two or three months. He also observed that the skin simulates the pattern of the background, and does not reproduce the latter. Size. — Summer flounders ordinarily grow to a maximum weight of 15 pounds or so, and to a length of 3 feet, or a little more, though one of about 30 pounds has been reported as taken off Fishers Island about 1915. 6 The largest of which we find definite record weighed 26 pounds. The largest on record, taken in sport fishing, was 37 inches long, weighing 20 pounds, caught at Oak Beach, N. Y., September 7, 1948, by F. H. Kessel, but the average size of the fish caught is only 2 to 5 pounds. The relation of length to weight is about as follows : 6 Average weight. Length P™nds 15-16 inches 1 to 1% 17-18 inches 2 to 2% 20 inches 3 to 3H 22 inches 4 27 inches 8 30 inches 10 37 inches 20 Habits.- — Many fluke come close inshore during the warm half of the year, when they are caught regularly both along open coasts and in bays and harbors, the smaller sizes often from docks and bridges, and some even run up into fresh wTater rivers. But the great majority of the population, especially of the larger ones, lie farther offshore even at that season, in depths of 8 to 10 fathoms and deeper, at least in the northern part of the fluke's geographic range, as illustrated by the fact that nearly 40 times as many (by weight) are landed in New Jersey and in New York by otter trawlers as from the many pound nets operating there.7 And all of those that do come close inshore from Chesapeake Bay northward move offshore again at some time during the autumn, presum- ably to escape winter chilling. The earliest landings from offshore of which we have heard for southern New England have been on October 6th, when some were brought in to Woods Hole from northwest of Nantucket Light- ship, from 25 fathoms, and on the 16th of that same month, when the dragger Eugene H landed 6,000 pounds, taken west of Nantucket Lightship in about 25 fathoms. Corresponding to this, only a few are seen near Woods Hole after the middle of October, or after the last week of November near New York. And very few reappear near New York before the first week in May, or before about the 10th of May near Woods Hole. It has been learned since the first edition of this book appeared that the medium sized and larger ones, at any rate, pass the winter and early spring out on the continental shelf from the 25 to 30 fathom contour about to the 80 fathom contour. Otter trawlers now make paying catches there as far north and east as the offing of southern New England, and as far south as the offing of northern North Carolina, during the part of the year when there are only a few "fluke" inshore, or none at all. In 1950 and 1951 , for example, the Eugene H? fishing in the general offing of Marthas Vineyard, brought in many fares ranging from a few hundred poimds to more than 20,000 pounds, between the first week of October and the third week of May, with the most productive fishing between early January and mid-April, from 25 to 75 fathoms. But it is doubtful whether many of them work deeper than that, for the Albatross III did not take any at depths greater than 80 fathoms off southern New England or New York in mid-May, 1950. Fluke spend most of their lives on bottom, or close to it, as other flatfishes do. During their stay in shoal water they prefer sandy bottom, or mud, where they are often seen. And it takes one only an instant to bury itself to the eyes in the sand. Fluke often lurk in eel grass, or among the piling of docks; but they are swift swimmers when disturbed. < Mast, Bull. TJ. S. Bur. Fish., vol. 34, 1916, p. 177. ■ Nichols and Breder, Zoologiea, N. Y. Zool. Soc, vol. 9, 1927, p. 177. • From Goode, Fish. Ind., U. S., Sect. 1, 1884, p. 179; Hildebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 43, Pt. 1, 1928, p. 167; and World Record Marine Game Fishes. Amer. Mus. Nat. Hist., New York, to 1950. 7 1947 landings, New York and New Jersey, about 2,300,000 pounds by otter trawlers; only about 80,000 pounds from pound nets. 'Information contributed by Capt. Henry Klimm. FISHES OF THE GULF OF MAINE 269 This is a predaceous fish, like the halibut, feeding largely on smaller fish of various sorts, on squids, crabs, shrimps, and other crustaceans; on small shelled mollusks; on worms, and on sand dollars. It is very fierce and active in pursuit of prey, often following schools of small fish right up to the sur- face, to jump clear of the water in its dashes, actions very different from those of the sluggish dab and winter flounder. Little is known of its breeding habits. The fact that nearly ripe females have been taken in Octo- ber in Chesapeake Bay, in November and Febru- ary at Beaufort, N. C, and as late as April 15, at 75 fathoms off Nantucket,9 whereas Beaufort fish taken in March and April appeared to be spent, show that it is a late autumn, whiter, and early spring spawner.10 This implies that the flukes that spawn in the northern part of their range do so well offshore, and this may also be true of them in the southern part of their range, for fluke that were kept in aquaria at Beaufort through the winter failed to spawn. The eggs of the summer flounder laid naturally have not been described yet. But it is likely that they are buoyant like those of the four-spotted flounder (p. 271). And their future "lefthanded- ness" and large mouths are foreshadowed at an early stage in the development of the larvae. Larvae either of the fluke, or of a form (P. albigvMus Jordan and Gilbert, 1882), so closely allied that it may prove a race of that species, resemble cor- responding stages of the four-spotted flounder in their deep outlines and large heads, hut the pig- mentation on the rear part of their body is less dense. At a length of 10 mm. the right eye has nearly completed its migration, and the outlines of young fry 26 mm. long approach those of the adult." Young fry taken in Chesapeake Bay, had in- creased in length from about 0.9-2.4 inches long in May and June, to 3-5 inches in the last week of July; were 4.7-7.1 inches by December and January when one year old or a little less; about 8-10 inches long in the following October, when they were a little short of two years old ; and they ' Trawled by the Eugene II. Gapt. Henry Klimm, in 1951. " We dare not draw any conclusions as to spawning season from Hilde- brand and Cable's table (Dull. IT. S. Bur. of Fisheries, vol. 46, p. 470, table 12) of the seasonal distribution of young fry of different sizes because two species of flounders are included there. 11 Our account of the young stages is based chiefly on Uildebrand and Cable's description (Bull. U. S. Bur. of Fisheries, vol. 46, pp. 469-475), from Beaufort, N. C. measured 10% to 11 inches by their second May; i. e., when a little more than 2 years of age. The subsequent rate of growth has not been traced, so far as we know. General range. — Continental waters of the east- ern United States, from Maine to South Carolina, possibly to Florida,1- chiefly south of Cape Cod. Occurrence in the Gulf of Maine. — This is the most important flatfish commercially to the west and south of Rhode Island, and the one most sought after by sportsmen there. It is also plentiful offshore eastward to Nantucket Shoals and to the western part of the so-called South Channel, whence about 531,000 pounds were landed in 1947 (most recent year for which infor- mation is at hand). Trawlers also pick up a few on the southwest part of Georges Bank (about 6,000 pounds in 1947), as well as a fish here and there on other parts of the bank.13 But there is no reason to suppose that fluke ever stray eastward and northward as far as Brown's Bank, or to outer Nova Scotian waters. Coastwise, the angle of Cape Cod is the northern boundary to the regular range of the fluke in any great abundance. A number are caught each summer in Pleasant Bay, Chatham, Mass.,14 where we read of one of 1 1 % pounds taken as early as the last week of May, in 1951, 16 a few in Town Cove, Orleans, some miles farther north, and a fluke is picked up occasionally by someone casting into the surf on the outer Cape Cod beach.16 And they were so common near Provincetown and along the inner shore of Cape Cod as far as Well- fleet during the period from 1840 to 1850 that Captain At wood carried them regularly thence to Boston, recording a catch of 2,000 pounds in a single afternoon inside Provincetown Harbor. But this is the most northerly region where fluke have ever been known to occur in commercial quantities. Even there its numbers were so reduced by a few years of hard fishing that they were described bv Goode 17 in 1884 as "onlv 13 Florida is usually given as the southern limit for this flounder, but it is possible that the early records from that State (there are no recent ones) actually referred to the southern flounder (P.lethostigmus),& common Florid- Ian fish. » 645 pounds reported from the northwest part of Oeorges in 1947, 100 pounds from the northeastern edge, and 157 pounds from the central and sutitheas tern part. i* This opens on the outer coast of Cape Cod. 11 Reported in Salt Water Sportsman, June 1. 1981. 18 There is a record of this, by Kendall, in 1896, and we have known of other cases, of late years. » Fisb. Ind. U. S., Sec. 1, 1884, p. 178. 270 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE occasionally taken" there. And they have never reappeared in any abundance, so far as we can learn, a fact suggesting that the local body of fish concerned was not very numerous, and that it received but few recruits from the more abundant stock to the southward. The fluke is so raTe a straggler north of Cape Cod Bay that there is only one definite record — for Casco Bay (specimens collected in 1873). We may add that we have never seen or heard of one caught in the inner part of Massachusetts Bay, and that it is unknown in the Bay of Fundy. Importance. — This is one of the best of our flatfishes on the table, usually bringing a higher price than any other except the halibut; in 1947 it sold for 15 cents on the average in New Bedford, the halibut about 21 cents. And the landings of fluke from within the limits of the Gulf of Maine, totaling about 543,000 pounds (mostly from near Nantucket Shoals) were worth about $90,000 to fishermen that year. This is also the gamest of our flatfishes, biting freely on almost any bait, even taking artificial lures at times, while large ones put up a strong resistance when hooked. It is too bad that the fluke is not so common north of Cape Cod as it is to the south. Four-spotted flounder Paralichthys oblongus 1815 (Mitchill) Jordan and Evermann, 1896-1900, p. 2632. Description. — This flatfish resembles the summer flounder (p. 267) so closely in its general make-up that we need mention only the points of differ- ence. Most apparent of these are that it has fewer dorsal fin rays (72 to 81 dorsal and 60 to 67 anal rays, contrasted with 85 to 94 and 60 to 73, respectively, in the summer flounder), and that its mottled gray back is invariably marked with four large, oblong, and very con- spicuous black eye spots edged with pale pinkish, two of them situated at each margin of the body, as the illustration shows (fig. 135). Incidentally, we have seen two of them on which the lower side, rearward from the gill openings was as dark as the upper side, and marked, similarly, with four eye spots; also others that were more or less dark below.18 This is also a much smaller fish than the sum- mer flounder, for the adults average only about 12 inches long with 16 inches as about the maximum. Habits. — Although this is a rather common fish about Woods Hole in May and June, and is still more numerous along the coast of New York, very little is known of its habits. It does not usually come into as shoal water as the sum- mer flounder often does, being caught most often in 7 to 17 fathoms in Vineyard Sound, for example, near Woods Hole. And the many that have been trawled by the Albatross II and Albatross III between Georges Bank and northern North Car- olina, have been generally distributed from about 23 fathoms down to at least 150 fathoms. w Fish trawled by the Eugene HofT Marthas Vineyard. Jan. 27 to Feb. 3, 1950, at 47 to 07 fathoms. . Figure 135. — Four-spotted flounder (Paralichthys oblongus), Woods Hole. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 271 Its diet is much the same as that of the summer flounder; chiefly small fish and squid, with crabs, shrimps, shellfish, and worms. It spawns from May until mid-July. The eggs are buoyant, 0.95 to 1.05 mm. in diameter, with a single oil globule of 0.16 to 0.19 mm.19 The early stages have not been described previously, but certain large mouthed and lefthanded M flat- fish larvae of 8 to 11 mm. that have been taken in tow nets off New Jersey, by the Grampus in 1913, and from Nantucket Shoals southward by the Albatross II subsequently, seem likely to have been young four-spots, not summer flounders, because they were taken in June and July (p. 269). If this identification is correct, an aggregation of pigment over the rear part of the trunk, combined with deep outline and a large head are distinctive for this species. Small fry of 2 to 3 inches have been taken at Woods Hole in autumn, evidence that the fry of this flounder complete their meta- morphosis and take to bottom about 3 months after they are hatched. General range.- — This flounder has been taken between the eastern part of Georges Bank and the coast of South Carolina.21 Its center of abundance appears to lie between southern New England and Delaware Bay. Occurrence in the Gulf of Maine. — The four- spotted flounder is so plentiful along the conti- nental shelf as far eastward as the general ofling of Nantucket, and to the neighboring part of Georges Bank that we counted about 1,800 of them trawled there, by the Eugene II in 56 hauls at 47 to 67 fathoms, January 27 to February 3, 1950, also 968 of them on the southwestern part of Georges, in 26 to 75 fathoms, in late June 1951. And a few were trawled in 1931 by the Albatross II also along the southern and southeastern parts of Georges Bank. But this last is their most easterly known outpost. The only records for the four-spot in the coastal waters of our Gulf are from Monomoy at the southern angle of Cape Cod; from the vicinity of Provincetown (where Storer saw a number of '• Eggs artificially fertilized by O. E. Sette of the Bureau of Fisheries. » Lefthandness foreshadowed In the larger ones by the fact that it Is the right eye that had begun to migrate. » The most southerly record for it is fur five specimens trawled in the general ofling of Charleston, S. C, lat. 33°00' N., long. 77°44' W., at 92 fathoms, by Albatross III, January 30, 1950. Another flounder, AncylopsMa quadrocellala QUI, similar In appoarance, for it is strikingly marked with four large spots, is found along the South Atlantic and Gulf coasts. them in June 1847) ; and from somewhere on the northern shore of Massachusetts Bay where one was taken by the United States Fish Commission in 1878.22 This is a fair table fish but there is no market for it at present. Yellowtail Limanda ferruginea (Storer) 1839 Rusty flouxder Jordan and Evermann, 1896-1900, p. 2644. Description. — The yellowtail is right-handed (that is, its eyes are on the right side and its vis- cera are at the right-hand edge as the fish lies on the bottom), and small-mouthed like the winter flounder, the smooth flounder and the witch. But it is easily distinguished from the first of these by its more pointed snout, thin body, arched lateral line, and more numerous fin rays; from the smooth flounder by the last two characters as well as by the concave dorsal (left hand) profile of its head and by being scaly between the eyes; and from the witch by its arched lateral line, its less numerous fin rays, concave dorsal (left) profile of the head, and especially by lacking the mucous pits on the left (white) side of its head that are conspicuous on the witch (p. 285). The yellowtail is a comparatively wide flounder, nearly one-half as broad as it is long, with an oval body. The dorsal (left hand) outline of its head is more deeply concave than in any other Gulf of Maine flounder; its head is narrower; its snout is more pointed, and its eyes are set so close together that their rounded orbits almost touch each other. The fact that its mouth reaches scarcely as far back as the eyes, with its small teeth and thick fleshy lips, marks it off at a glance from all the large-mouthed flounders. The dorsal fin (76 to 85 rays) originates over the eyes, its middle rays are the longest. Its anal fin is similar in outline to the dorsal, but is much shorter (56 to 63 rays), and it is preceded by a short, sharp spine pointing forward. The two ventral fins are alike, and each of them is separated by a consid- erable space from the anal fin. But the pectoral fin on the blind side is slightly shorter than its mate on the eyed side. The scales are rough on the eyed side, but smooth on the blind side. " In one paper (Am. Jour. Set, Ser. 3, vol. 17, 1879, p. 40) Qoodo and Bean state that this specimen was trawled in Oloucester Harbor; in another paper (Bull. Essex Inst., vol. 11, 1879, p. 7) they credit It to the mouth of Salem Harbor. 272 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE ,: '..I'Mii'H.'i- .- ■'■'- - ■ ./ ■ '•■ ; 'I.I..'- .■ V. | . A- A. SSA ^"'1 v^'~ Figure 136. — Yellowtail (Limanda ferruginea) , Gloucester, Mass. From Jordan and Evermann. Drawing by H. L. Todd. Color. — The yellowtail is more constant in color than most of the other Gulf of Maine flatfishes. Its eyed side, including the fins, is brownish or slaty olive, tinged with reddish and marked with large irregular rusty red spots. The caudal fin and the margins of the two long fins are yellow, the yellow tail in particular being a very diag- nostic character. The blind side is white, except for the caudal peduncle which is yellowish. Size. — This is a medium-sized flatfish. Several hundred adults caught in gill nets between Cape Ann and Cape Elizabeth (measured by Welsh) ran as follows: Males, average length 15% inches, extremes 11% inches to 18% inches; females, aver- age length 18 inches, extremes \b}{ inches to 21% inches. This series includes the largest specimens that have ever been reported. A yellowtail 12 inches long weighs about one-half pound; one 15 inches long, about 1 pound; and one 18 inches long about 2 pounds. Habits. — A yellowtail is caught in very shoal water now and then: We heard, for example, of several taken in Pleasant Bay, Cape Cod, in 1950. But 5 to 7 fathoms may be set as its upper limit, generally speaking. Thus it keeps to rather deeper water than either the winter flounder or the smooth flounder. On the other hand, most of those caught are at least from no deeper than 50-60 fathoms,23 and the bulk of the catch is made 23 One was taken at 50 fathoms by Albatross II, September 5, 1926, on the northwestern part of Georges Bank, and two of about 10 inches at 90-95 fathoms on the northern edge of Georges Bank, by Cap'n Bill II, August 22,1952. shoaler than 40 fathoms. We saw many yellow- tails trawled by the Albatross III off Marthas Vineyard and Nantucket in 20 to 40 fathoms, in May, 1950, but only 6 in 41 to 50 fathoms, and none in deeper water. Again, in late June 1951, Eugene H averaged about 240 yellowtails per trawl bawl, at 26 to 45 fathoms on the western part of Georges Bank, but took only three of them in deeper hauls. Almost any sandy bottom or mixture of sand and mud suits them, and most of those that Welsh saw taken in gill nets on the Isles of Shoals-Boon e Island grounds (p. 274) were over fine black sand between the hard, rocky patches. Rocks, stony ground, and very soft mud are shunned by yellow- tails, as they are by most of the other flatfishes. The yellowtail feeds chiefly on the smaller crustaceans such as amphipods, shrimps, mysids, and on the smaller shellfish, both univalves and bivalves, and on worms. It is also known to eat small fish, but it is not likely that it can catch these often. Its European relative also feeds on sea urchins, starfish, and on algae at times. And it is probable that our yellowtail would be found equally omnivorous were then stomachs examined from various localities. Fish in breeding condi- tion are empty as a rule. The diet of the yellowtail suggests that it is one of the more sluggish of our flatfishes, and there is no reason to suppose that it ever travels about much after it once takes to the bottom except that it has been described, in Massachusetts Bay, as "in- FISHES OF THE GULF OF MAINE 273 habiting the deep water ... in summer, and approaching the shores in whiter," 24 as do various other ground fishes that tend to avoid high temperatures. If the yellowtails are as stationary as they seem to be, they must be subject to considerable range of temperature from season to season at different depths, in one part of the Gulf or another, from a maximum of about 52°-54° to a minimum of about 33°-36°. And some of them are exposed to still lower temperatures on the Grand Banks, and in the Gulf of St. Lawrence. The eggs of the yellowtail, artificially fertilized by Welsh in 1912, and hatched at the Gloucester hatchery, were buoyant, without oil globule, spherical, very transparent, and with a narrow perivitelline space. One hundred eggs measured by hini ranged from 0.87 mm. to 0.94 in diameter, averaging about 0.9 mm. The surface of the egg is covered with very minute striations, and the germinal disk is of a very pale buff color while alive. The embryonic pigment gathers in three groups shortly before hatching (which takes place in 5 days at a temperature of 50° to 52°); one group on the head, a second group in the region of the vent, and a third group half way between the vent and the tip of the tail. Unfortunately the fish which Welsh hatched were destroyed accident - ally, so we cannot describe the early larval stages. Larvae of 11 mm. are still symmetrical. But the left eye is already visible above the profile of the head at 14 mm. (fig. 139, Grampus specimen), all the fins are outlined, with their rays present in the final number (76 dorsal and 59 anal hi the specimen illustrated). Thus, they show enough of the distinctive characters of the adult for posi- tive identification. The early larval stages of yellowtails and of whiter flounders resemble one another closely; in fact, it is probable that some of the young flat- fishes pictured by A. Agassiz 26 as winter floiuiders were yellowtails in reality. But the number of fin rays usually places the larvae in one species or the other after these appear. And the yellow- tail does not take to bottom until upward of 14 mm. long, whereas the whiter flounder completes its metamorphosis when it is only 8 to 9 mm. long. Figure 137. — Egg. Figure 138. — Larva, 10.3 mm. " Qoode and Bean, Bull. Essex Inst., vol. 11, 1879, p. 6. '• Agassiz, Proc. American Acad. Arts, Sci., N. Ser., vol. 6, 1879, pi. 4. Figure 139. — Larva, 14 mm. Yellowtail (Limanda ferruginea) Captures of young fish 2 to 4 inches long in February; 2}i to 4% inches long hi April; 2% to 5% inches long in May; 3 to 5 inches long in June; and 3 to 6% inches in July indicate that the yellow- tail grows to an average length of about ;") inches by the time it is one year old. Its subsequent rate of growth has not been traced. General range. — North American continental waters, from the north shore of the Gulf of St. Lawrence, the Labrador side of the Strait of Belle Isle,26 northern Newfoundland (there are specimens from St. Anthony's in the Museum of Comparative Zoology), and the Newfoundland Banks, southward to the lower part of Chesa- peake Bay.27 It is most plentiful on the western » Recent records from the Labrador side of the Strait are of one from Barge Bay, July 29, 1910 (Jeffers, Contrib. Canad. Biol., N. Ser., vol. 7, No. 16, 1932, p. 210) ; and of another taken at Forteau Bay, June 29, 1949, by the Blue Dolphin Expedition, reported to us by Richard H. Backus. » We have records of one taken off Hog Island, Va., In lat. 37° 41' S. (Big- elow and Schroeder. Bull. U. S. Bur. Fisheries, vol. 48, 1939, p. 340). And it was reported from the southern part of Chesapeake Bay by Uhler and Lugger (Rept. Comm. Fish., Maryland, 1876, p. 95; 2d Ed., 1876, p. 79.) 274 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE half of Georges Bank; in the western side of the inner parts of the Gulf of Maine; on the Nan- tucket grounds; and off southern New England. This flatfish is represented in north European waters by the European dab, Limanda limanda, a close ally, from which it is distinguishable by its smaller scales, more pointed snout, more nu- merous fin rays, and shorter pectoral fins. We should also mention the deep-water dab (Limanda beanii Goode), for while it has not been taken within the limits of the Gulf of Maine it would not be astonishing to find it on the seaward slope of Georges Bank, for it has been taken westward and southward from Marthas Vineyard in depths of 120 to 896 fathoms.2* It differs from the rusty dab in a shorter head (occu- pying only two-elevenths of the total length instead of one-fourth) ; in the fact that the dorsal (left hand) profile of its snout is convex, not con- cave; in having only about 64 dorsal fin rays instead of 76 or more; in having only 88 rows of scales along its lateral line instead of 90 to 100; and in the fact that its tail fin is marked with a conspicuous black blotch on the outer rays on each side. Occurrence in the Gulj of Maine. — Little was known of the distribution of the yellowtail in our Gulf previous to the introduction of the otter trawl there, for it is seldom seen close inshore; while its mouth is so small that one is seldom caught on hooks as large as those that are used for cod, pollock, or for haddock. But it has proved so abundant since then, in the general region of Nantucket Shoals and in the neighboring side of the so-called South Channel, that about 4,400,000 pounds were landed thence in the most recent year (1947) for which we have information. The western half of Georges Bank as a whole is good yellowtail ground also. But yellowtails seem to be less numerous on the eastern half of the Bank (though generally distributed there), and less so on Browns Bank, as is illustrated by the landings (in pounds) for 1947,29 as follows: northwest Georges Bank, 930,000; southwest Georges Bank 1,740,000; northeast Georges Bank 210,000; central and southeast Georges Bank 540,000; and Browns Bank 40,310. >■ Localities are listed by Ooode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, p. 430). » To nearest 10,000 pounds. Yellowtails are so plentiful on the sandy bottoms in the eastern side of Cape Cod Bay, also, and on Stellwagen Bank, that (with winter flounders) they have long been the mainstay of the draggers that fish there; no doubt the greater part of the 1,150,000 pounds of yellowtails that were reported as taken off eastern Massachusetts in 1947 were trawled on these particular grounds. There are yellowtails in the deeper parts of Massachusetts Bay, too, as Goode and Bean30 remarked long ago. And since Welsh saw many hundred of them taken in gill nets (not very effective gear for flatfishes) between the Isles of Shoals and Great Boars Head, during March and April of 1913, the yellowtail must be one of the most numerous of its tribe in the western side of the Gulf in general, in suitable depths. Apparently they are less plentiful, however, around tbe Gulf to the north and northeast, for the reported catches for 1945 were only about 44,500 pounds for Cumberland County, Maine,31 which covers Casco Bay and the grounds in its offing; only about 9,000 pounds for Knox and Han- cock Counties combined, and none for Washington County in that particular year, though a few hundred pounds have been reported from "eastern Maine" in some other years. We have taken no yellowtails in the deep basins of our Gulf nor have we heard of any there, probably because of the depth, for the bottom would seem hard enough for them in the eastern trough, at least, even if it is not in the western, or in the bowl west of Jeffreys Ledge. They cer- tainly are uncommon in the Bay of Fundy, too, if not altogether lacking there. And though Huntsman did find a few in St. Mary's Bay, Nova Scotia, United States fishermen bring in only a few hundred pounds from off western Nova Scotia in some years, and none at all in others, though considerable amounts are brought in from the outer Nova Scotian grounds, as mentioned below (p. 275). Most of the yellowtails that are caught in the inner part of the Gulf of Maine are in 10 to 30 fathoms of water, though they are reported in Shoal water at the mouth of Penobscot Bay; those caught on Georges Bank are in 20 to 45 or 50 fathoms (see p. 272). " Bull. Essex Inst., vol. 11, 1879, p. 6. 11 Apart from those that were brought in to Portland by the large trawlers from more distant grounds. FISHES OF THE GULF OF MAINE 275 Trawlers find yellowtails in even greater num- bers off the southern New England coast than on Nantucket Shoals, at the proper depths, as illus- trated by reported landings thence of about 17% million pounds in 1947. 32 And they are moder- ately plentiful offshore, as far as the offing of New York.33 But southern New Jersey is about the southern limit to their regular ocurrence.34 Turning our attention eastward, we find the yellowtail plentiful all along the outer Nova Scotian banks, where about 2,700,000 pounds were taken in 1947 by vessels from Massachusetts,36 besides about 2% million pounds by Canadian vessels. They are also reported as numerous on the southern part of the Grand Banks from experi- mental trawling by the Newfoundland Fishery Research Commission, but are "not in any num- bers along the Newfoundland coast," 36 so far as is known, though they are recorded from as far north as the Strait of Belle Isle, as already noted (p. 273). They are also distributed generally in the Gulf of St. Lawrence, but no information is available as to their numbers there, for none are reported from the Gulf in the Canadian Fisheries statistics. The neighborhood of the Isles of Shoals and of Boars Head, at 20 to 30 fathoms, certainly is an important spawning ground for the yellowtails;87 so, too, the edges of Stellwagen Bank where we have caught spawning specimens. In fact, it is likely that yellowtail eggs are produced in abun- dance all around the western and northwestern periphery of the Gulf, between the 20 fathom and 50 fathom contours; few, however, in the eastern side, and none in the Bay of Fundy; nor have we found any of its eggs anywhere over depths greater than 50 fathoms. No doubt the yellowtail spawns as actively on the offshore Banks as it does inshore, for though we have not actually found its eggs there we have taken larvae only 7 to 11 mm. long over the western and eastern parts of Georges » Landings in 1947 in Massachusetts ports, from grounds westward from Nantucket Shoals, about 12 million pounds; landings in Rhode Island, about 2H million pounds; landings in Connecticut, about 3 million pounds. " About 3H million pounds were landed in New York in 1947. " Albatross II trawled many yellowtails as far southward as the offing of Delaware Bay (lat. 38° 32' N., long. 74° 24' W.) in 12 to 28 fathoms during February, April, and June, of 1929 and 1930. 31 The Newfoundland Fishery Research Commission (Rept., vol. 1, No. 4, 1932, p. 1 10) reports 680 yellowtails taken per 10 hours trawling on Banquercau. '• Frost, Research Bull. 14, Newfoundland Dept. Nat. Resources, 1940, p. IS. 17 Welsh obtained many ripe fish there. Bank in July,38 as well as near Gloucester and near the tip of Cape Cod in July and August. To the eastward and northward, yellowtail spawn on Sable Island Bank, Banquereau Bank, and the Newfoundland Banks, eggs (no doubt of this species) having been collected on these grounds by the Canadian Fisheries Expedition in 191 5.S9 In the opposite direction, it certainly breeds as far westward as New Jersey, for our tow net yielded 88 of its larvae (6.5 to 19 mm. long) 11 miles off Sandy Hook on August 1, 1913; adult fish approaching ripeness have been trawled as far southward as Little Egg Inlet, N. J., in April 1930. Spawning, Welsh found, begins on the Isles of Shoals-Boone Island ground by the middle of March; and many ripe fish were taken there during the last half of April, but the majority were still green as late in the season as May 8, though others were already spawned out. And spawning must last all summer, for we have trawled many ripe males and females in depths of 17 to 25 fathoms on the edge of Stellwagen Bank at the end of July; have taken eggs indistinguishable from those of the yellowtail in our tow nets in June, July, and August, with one even on Sep- tember 11: and have taken its newly hatched larvae (6 mm. long) off Race Point as late as August 31. And the individual females evidently spawn over a considerable period of time, for Welsh found that only a small part of the eggs ripened simultaneously. Importance.- — The yellowtail is one of the most valuable of the flatfishes caught within the Gulf of Maine. It compares favorably in quality with the summer flounder and the winter flounder, but because its body is thinner it brings a lower price to the fishermen. Thus in 1947 the average price, as landed in New Bedford, was about 8 to 9 cents a pound for yellowtails; winter flounders, about 9 to 10 cents a pound; and summer flound- ers, about 17 to 18 cents a pound. All the yellow- tails that are brought in find a ready sale and they make up a large part of the fillet of sole sold to consumers. In 1947 our Gulf yielded between 15 and 16 million pounds of them. But yellow- tails live rather too deep to be of any interest to anglers. >' Station 10059, July 9, 1913; and station 10224, July 23, 1914. '• Dannevlg (Canadian Fisheries Expedition [1914-15], 1919, p. 17) refers these provisionally to the European dab, which does not occur on our side of the Atlantic. Its egg is indistinguishable from that of the American species. 276 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Winter flounder Pseudopleuronectes americanus (Walbaum) 1792 Blackback; Georges Bank flounder; Lemon sole; Flounder; Sole; Flatfish; Rough flounder; Mud dab; Black flounder Jordan and Evermann, 1896-1900, p. 2647. Description. — This is a small-mouthed, right- handed species (eyes on the right side and viscera on the right). But it is easily separable from the yellowtail, which is similarly characterized, by the fact that its lateral line is nearly straight (at most only slightly bowed abreast the pectoral fin) ; that the dorsal profile of its head is less concave; that its nose is blunter; that its eyes are farther apart; that it has fewer fin rays; and that its fins are less tapering in outline. The most obvious differences between the winter flounder and the smooth flounder (p. 283) is that the former is rough scaled between the eyes, the latter smooth there, and that the winter flounder has the larger number of anal fin rays. On the other hand, it has only about two-thirds as many dorsal rays as the witch (p. 285) ; it lacks the mucous pits that are conspicuous on the left (lower) side of the head of the witch, and its tail is much larger proportionately than that of the witch. It is oval in outline, about two and one-fourth times as long to the base of the caudal fin as it i? wide, thick-bodied, and with proportionately broader caudal peduncle and tail than any of our other small flatfishes. Its dorsal fin (60 to 76 rays) originates opposite the forward edge of the eye, and is of nearly equal height throughout its length. Its anal fin (45 to 58 rays) *° is highest about midway, and it is preceded by a short, sharp spine. Its ventral fins are alike on the two sides of the body, and both of them are separated from the long anal fin by a considerable gap. The mouth is small, not gaping back to the eye, and the lips are thick and fleshy like those of the yellowtail. The left (under) half of each jaw is armed with one series of close-set incisor-like teeth, but the right (upper) side has only a few teeth, or it may even be tooth- less. The scales are rough on the eyed side, including the space between the eyes, but they are smooth to the touch on the blind (white) side. Color. — The winter flounder, like other flat- fishes, varies in hue according to the bottom on which it lies, but it is the darkest of Gulf of Maine flatfishes as a rule. Large ones are usually of some shade of muddy or slightly reddish brown, olive green, or dark slate above, sometimes almost black. And they vary from plain or more or less mottled to definitely marked with smaller or larger spots of a darker shade of the general ground tone. There usually is a wide variation in this respect, among any lot of flounders. And fish caught on Georges Bank average more reddish in «Perlmutter (Bull. Bingbam Oceanogr. Coll., vol. 11, Art. 2, 1947, pp. 19, 20) gives a detailed tabulation, and graph of the number of dorsal and anal fin rays from upwards of 1,100 specimens including both the smaller inshore form and the larger Georges Bank form. Figure 140. — Winter flounder (Pseudopleuronectes americanus). From Goode. Drawing by 11. L. Todd. FISHES OF THE GULF OF MAINE 277 hue than those caught inshore. But this rule does not always hold (see p. 277). The blind side is white, more or less translucent toward the edge, where it is often faintly tinged with bluish, and the lower side of the caudal peduncle is yellowish on some specimens, but is pure white on others. The long fins usually are tinged with pinkish, reddish, or are yellowish on the eyed side; the ventrals and pectorals of the eyed side are of the general ground tone, but their mates on the blind side are pure white. Small fish average paler and more blotched or mottled than large ones. Various color abnormalities have been recorded (fish, for example, that are partially white on the eyed as well as on the blind side, or with the blind side yellow-edged) and it is not uncommon to see specimens with dark blotches on the blind side. In fact, one-third of the fish caught near Provi- dence, R. I., during the winter of 1897-98 were these "black bellies," as fishermen call them, but the commissioners of fisheries of that State esti- mated them as forming only 4 percent of the catch in 1900. And none (or at most only an occasional fish) has been seen since. In 1898, some fry that had been hatched artificially from eggs of black-bellied flounders were released in Waquoit Bay, southern Massachusetts, where this race had been unknown previously, and several "black bellies" 7 to 8 inches long (hence probably two years old) were taken there in 1900, probably the offspring of this planted stock.41 Winter flounders change color to some ex- tent to suit their surroundings, usually being very dark on mud, and pale on bright sand bottoms. But field experience suggests that they have less control over shade and pattern than the summer flounder has. Size. — The largest winter flounder on record caught inshore was one 22% inches long mentioned by Scattergood ; 42 Nichols and Breder 43 report one 20 inches long, weighing 5 pounds ; and Welsh saw three of about 19^ inches, weighing 3%, 3%, and 4 pounds, respectively, that were caught near Boon Island in April 1913. But fish longer than 18 inches or heavier than 3 pounds are unusual inshore, the general run of adults caught there being from 12 to 15 inches in length and 1% to 2 '■ Bull.. U. S. Fish Comm., vol. 19, 1901, pp. 305-306. « Copeia 1952, p. 206. « Zoologies, New York Zool. Soc, vol. 9, 1927, p. ISO. pounds in weight. Flounders grow larger on Georges Bank, where many of 4-6 pounds are taken, and where they often are caught up to 7-8 pounds ; we have handled one Georges Bank fish of 25 inches, weighing 8 pounds. Remarks. — The winter flounder shows some tendency to break up into local races in the number of its fin rays,44 in the size to which they grow, and perhaps in other characteristics. The most interesting of these races, from the fisheries standpoint, is the population on Georges Bank, for the flounders tend to grow larger there than they do anywhere inshore. This fact was first brought to scientific attention in 1912, when some of these large flounders from Georges were received by the Bureau of Fisheries, to be made the basis of a new species, Pseudopleuronectus dignabUis, by Kendall.45 Since that time this Georges Bank flounder has been accepted pro- visionally as a separate species, supposedly charac- terized by rather more numerous fin rays, by reddish color, and by a caudal peduncle yellow on the under side, as well as by large size. But our own comparison of specimens of the winter flounder group of various sizes, from Georges Bank, with others from the No Mans Land ground, from Nantucket Shoals, and from many localities, inshore, from Labrador to New York, leads us to conclude that it is simply a large, more rusty-brownish, local race of the winter flounder, for we find no definite regional discontinuity in the number of fin rays or of gill rakers, in the teeth, or in color (p. 277). The names "black- back" and "lemon sole," as used by fishermen, have no bearing on the case, for their choice of the one or of the other is based solely on the size of the fish in question (p. 282 ).46 Habits. — Tide mark, high or low according to the stage of the tide, is the upper limit for this flounder. It runs up into brackish water in river mouths, and we have even caught them in the Susquehanna River, tributary to Chesapeake Bay, where the water was fresh enough to drink.47 Its lower limit cannot be stated definitely. It is plentiful certainly at 10 to 20 fathoms in Cape « See Bumpus (American Naturalist, vol. 32, 1898, pp. 407-412) and espe- cially Perlmutter (Bull. Bingham Oceanographic Coll., vol. 11, Art. 2, 1947, pp. IS- 23) in this connection. « Bull. U. S. Bur. Fish., vol. 30, 1912, p. 391. pi. 57. *8 Perlmutter has already emphasized this point in his detailed study of the blackback (Bull. Bingham Oceanogr. Coll., vol. 11, Art. 2, 1947, p. 18). <> HUdebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 43, Pt. 1, 1928, p. 170. 278 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Cod Bay and on Stellwagen Bank, while the gill- netters sometimes take very large ones at about this same depth about Boon Island. According to general report, however, few, if any, are caught deeper than this in the inner parts of the Gulf except in the Bay of Fundy, where they are to be taken in winter on soft bottoms down to 30 to 50 fathoms. On Georges Bank they are taken mostly between 25 fathoms and 45 fathoms; 70 fathoms is the deepest definite record for them there of which we know. Usually the smaller fish live the shoalest and the larger ones deeper. But we have seen large flounders caught so often in only a few feet of water that no general rule can be laid down. The young fry are found chiefly in the shallows. Most of those that are caught inshore are on muddy sand, especially where this is broken by patches of eelgrass. But winter flounders are common enough there on cleaner sand, on clay, and even on pebbly and gravelly ground. And the populations on the offshore banks are on hard bottom of one type or another. "When they are on soft bottom they usually lie buried, all but the eyes, working themselves down into the mud almost instantly when they settle from swimming. And flounders that live on the flats usually lie motionless over the low tide to become more active on the flood, when they scatter in search of food. They keep near the bottom, and we have never heard of them coming up to the surface as the summer flounder so often does (p. 269). But though they spend most of their time lying motionless, they can dash for a few yards with astonishing rapidity, to snap up any luckless shrimp or other victim that comes within reach, or to snatch a bait, as any one may see, who will take the trouble to watch them on the flats on a calm day. It is in this manner that they usually feed, not by rooting in the sand. But flounders can sometimes be attracted by stirring the bottom with an oar when they are not biting, or by dragging anchor to bring up small animals from the mud, an old trick. How close inshore they may come (how shoal) in any particular locality at any particular time depends largely on local conditions of temperature. Generally speaking, the summer temperature is low enough for their comfort close in to shore and up to within a few feet of the surface all around the open coast line of the Gulf, and among the island passages, but the winter temperatures may be uncomfortably low for them in enclosed situa- tions locally. Tn Passamaquoddy Bay, for in- stance, where the temperature of the water falls close to the freezing point in winter, those that are closest inshore in summer work out in winter unless the year is a very mild one. Others, how- ever, that are living at 15 fathoms or so remain there the year around, while it is only in winter that they are known to descend as deep as 30 to 50 fathoms in the Bay of Fundy.49 In shallow enclosed bays, however, or harbors, where extensive flats are heated by the sun at low tide in summer but are exposed to very severe chilling in winter, the flounders tend to desert the flats for the deeper channels during the heat of summer, work back again into shoal water in autumn, desert the ice-bound flats once more in winter, and then work up again in spring. Duxbury Bay is a case in point, also Barnstable Harbor, where we have speared many of them in spring, while wading on the flats. A migration of flounders out into deeper water in the summer and back to shoal for the winter is generally characteristic south of New York, where the coastal waters are warmer, hence the common name "winter flounder." They are very scarce, for instance, in the bays of southern New Jersey in summer, but very plentiful there in winter. And many are caught in Chesapeake Bay from November to the first of June, but none are taken in shoal water there in summer or early autumn. It has long been believed that the winter flounder is one of the most stationary of our fishes, apart from seasonal movements of the sorts just men- tioned, and apart from a general tendency (re- cently emphasized by Perlmutter)60 for the fry that are produced in bays and estuaries to work offshore as they grow older. This essentially stationary nature has been demonstrated recently by extensive marking experiments that have been carried out in Long Island Sound, along southern New England, and on the coast of Maine, for about 94 percent of the recaptures were made in the general areas where the fish had been tagged. Thus the popidation consists "of many independ- ent localized stocks inhabiting the bays and estuaries along the coast" as Perlmutter words it, «• As proved by captures in shrimp trawls, as reported by Huntsman. '• Bull. Bingham Oeeanogr. Coll., vol. 11, Art. ?, 1947, p. 17 FISHES OF THE GULF OF MAINE 279 with the fish merely tending to scatter "from population centers, a characteristic phenomenon with nonmigratory animals." S1 But some of them may stray for considerable distances. Thus winter flounders tagged at Waquoit Bay, near Woods Hole, in 1931 were recovered off Chatham, on the outer coast of Cape Cod, and on Nantucket Shoals. The case of one that was tagged near Block Island on April 17, 1941, and was recaught on the central part of Georges Bank (lat. 41°45' N., long. 67°06' W.) on August 27, 1945, is especially interesting,62 as showing that some interchange does take place between the inshore and offshore populations of adult fish. The normal distribution of the winter flounder covers a wide range of temperature at one season or another, from a minimum close to the freezing point of salt water around Newfoundland, in Nova Scotian waters, in the Gulf of St. Lawrence, and in the shoaler parts of the Gulf of Maine in late winter, to a maximum of about 64°-66° F. in shallow water in the southwestern part of the Gulf in summer, and of perhaps about 68°-70° in the southern part of its range. They sometimes perish by the thousands in very hot spells of summer weather, if they are trapped in shallow enclosed bays, as happened in Moriches Bay, Long Island, N. Y., in 1917, be- tween July 29 and August 4, when the ah tem- perature rose to 82°-89°, and the temperature of the water on the very shallow flats nearly as high, probably.63 But wTe have never heard of this happening in the Gulf of Maine where cooler water is always close to hand. On the other hand, they may succumb to anchor ice in winter if they are overtaken in very shoal water in a severe freeze, for dead "flounders" of one sort or another are sometimes reported in such locations after unusually severe weather. And observa- tions at Woods Hole have shown that freezing temperatures (say 30° to 29°) drive them down into slightly warmer water. Experience at the Boothbay and Woods Hole hatcheries, combined with the results of the trawl fishery (p. 283), proves that those living a few fathoms down are as active in winter as they are in summer, both north and south of Cape Cod. " Perlmutter, Bull. Blngbam Oceanopr. Coll., vol. 1 1 , Art. 2, 1947, pp. 26, 27, " This specimen Is on display at the Laboratory of the U. S. Fish and Wild- life Service at Woods Hole. « This occurrence Is described by Nichols (Copeia, No. 55, 1918, pp. 37-39), also by Nichols and Breder, Zoologica, N. Y. Zool. Soc, vol. 9, 1927, p. 79. Bean, it is true, has described the winter flounder as going into "partial hibernation in the mud in winter,54 but (as Breder55 has pointed out) the reason the hook-and-line fishermen cannot take them in late winter or early spring may simply be that they will not bite then, this being the spawning period when winter flounders fast, as so many other fishes do. According to Sullivan56 diatoms are the first food taken after the yolk of the larval flounder is absorbed. A little later they begin preying on the smaller Crustacea, and Sullivan invariably found isopods in the stomachs of fry that had just passed their metamorphosis. A series of young flounders 1 to 4)j inches long from Casco Bay were found by Welsh to have fed chiefly on isopod crustaceans, with lesser amounts of copepods, amphipods, crabs, and shrimps, which together formed 36 percent of the stomach contents; worms (39 percent); mollusks (2 percent); and various unidentifiable material (22 percent). Linton57 who examined about 398 young flounders of various sizes at Woods Hole, likewise found them feeding chiefly on amphipods and on other small Crustacea, together with annelid worms. And his tables of stomach contents show an increase in the ratio of mollusks to Crustacea as the fish grow. The adult winter flounder, like the yellow- tail (p. 271), is limited by its small mouth to a diet of the smaller invertebrates and of fish fry. Sometimes they are full of shrimps, amphipods, small crabs, or other crustaceans; sometimes of ascidians, seaworms (Nereis), or other annelids; or of bivalve or univalve mollusks. Three hun- dred "seed" clams, for example, were found in an 11-inch flounder at St. Andrews, New Brunswick.58 And it seems that they often bite off clam siphons that protude from the sand. They also eat squid, holothurians, and hydroids; occasionally they capture small fish; and they sometimes take bits of seaweed. Examination of the stomachs of adults taken at Woods Hole in February 1921 by Breder showed that they cease feeding when they are about to spawn. In spite of its small mouth the winter flounder bites very readily on clams, pieces of seaworm. or « Bull. r,0, New York State Mus., Zool., 9, 1903, p. 778. » Bull. U. S. Bur. Fish., vol. 38, 1923, p. 311. - Trans. Amer. Fisheries Soc, vol. 44, 1914-15, No. 1, p. 138. " App. 4, Report U. S. Comm. Fish. (1921) 1922, pp. 3-14. » Fisheries Research Board of Canada, Progress Reports of the Atlantic Coast Stations, No. 52. January 1052. p. 3. 280 FISHERY BULLETIN OF THE FISH AXD WILDLIFE SERVICE almost any other bait for that matter, provided the hook is small enough. Breeding habits. — The winter flounder is a winter and early spring breeder, spawning from January to May (inclusive) in New England. The season is at its height during February and March south of Cape Cod and in the Massachusetts Bay region,59 but it is somewhat later along the coast of Maine; near Boothbay spawning commences about March 1 and continues until about May 10 or 15 with the chief production of eggs usually taking place from March 30 to April 20, according to information supplied by Capt. E. E. Hahn, former superintendent of the Boothbay hatchery. Local differences of this sort in the spawning season are probably due to variations in the tem- perature of the water. After the severe winter of 1922-23, for example, when the vernal warming of the coastwise waters was slower than usual, Captain Hahn wrote us from Boothbay that "the fish were 10 to 15 days later in spawning than in any previous year, the first eggs being taken on March 24." On Georges Bank spawning fish have been reported in April and into May. Thus spawning is well under way inshore while the water is still near its coldest for the year; i. e., about 32° to 35° F. in the Woods Hole region, about 32° to 37° near Gloucester, and about 31° to 35° near Boothbay, according to precise locality and depth. And the major production of eggs takes place there before the water has warmed above about 38°, with about 40° to 42° as perhaps the maximum for any extensive spawning in the inner parts of our Gulf. The picture is not so clear for Georges Bank, for we do not yet know how early in the season flounders commence spawning there. Those that spawn on the Bank in April may do so in temperatures ranging from about 38° to perhaps 42°, depending on the year, on the precise date, and on the locality. Winter flounders spawn on sandy bottom, often in water as shoal as 1 to 3 fathoms, but as deep as 25 to 40 fathoms on George Bank, and they do so throughout the range of the fish, including the Bay of Fundy, where Huntsman found its larvae common near the mouths of estuaries. Most of the eggs are produced in salinities from about 31 to 32.3 per mille in the inner parts of the Gulf, to somewhere between 32.7 and 33 per mille on Nan- M This species was propagated artificially at the Woods Hole, Gloucester, and Boothbay hatcheries in large numbers. tucket Shoals and on Georges Bank. But those that spawn in estuaries are known to do so in brackish water, in salinities as low as 11.4 per mille near Woods Hole, for instance. Individual females produce an average of about 500,000 eggs annually, and nearly 1,500,000 have been taken from a large one of 3% pounds. They spawn at night, at least those did that were kept in the tanks at Woods Hole, where they seemed indifferent to the electric lights overhead. And Breder 60 describes the fish of both sexes as swimmmg in a circle, about one foot in diameter, clockwise so that the vent is outward, with the eggs from the females flowing back along the upper side of the anal fin and along the tail. After about 10 seconds of activity, they sink motionless to the bottom. This species is peculiar among our local flatfishes in that its eggs are not buoyant but sink to the bottom, where they stick together in clusters, usually so closely massed that the individual eggs are forced into irregular outlines. They are 0.74 to 0.85 mm. in diameter, and newly shed eggs have no oil globule, but some of them (if not all) develop one as incubation proceeds.61 Incubation occupies 15 to 18 days at a temperature of 37° to 38° F., which is about what they encounter in nature. The young larvae, which are 3 to 3.5 nun. long at hatching, are marked by a broad vertical band of pigment cells that subdivides the post anal part of the body, a characteristic feature; and the end of the gut also is heavily pigmented. In water of about 39° the larva grows to 5 nun. in length, and the yolk is absorbed (fig. 142) in 12 to 14 days. The vertical fin rays begin to appear in 5 to 6 weeks after hatching, at a length of about 7 nun., and the left eye has moved upward by then untd about half of it is visible above the dorsal outline of the head, while the whole left eye shows from the right side and the fins are fully formed in larvae of 8 mm. Metamorphosis continues rapidly.62 The left eye moves from this position to the right side of the head; the pigment fades from the blind side; the eyed side becomes uniformly pig- mented; and the little fish now lies and swims with the blind side down, its metamorphosis complete when it is only 8 to 9 mm. long. * Copeia, No. 102, 1922, pp. 3-4. •' Breder, Bull. U. S. Bur. Fish., vol. 38, 1923, fig. 274g. »2 Williams, Bull. Mus. Comp. Zool., vol. 40, 1902, No. 1, pp. 1-58, pis. 1-5. See also Sullivan (Trans. Amer. Fish. Soc, vol. 44, 1914-15, pp. 125-136, figs. 1-4) and Breder (Bull. U. s. Bur. Fish. vol. 3S, 1923, p. 311). FISHES OF THE GULF OF MAINE 281 Figure 143. — Larva, 5 mm. Figure 141. — Egg. Figure 142. — Larva, 4.5 mm. Figure 144. — Larva, 8 mm. Winter flounder (Pseudopleuronedes americanus.) The youngest larval stages are made indentifi- able as winter flounders by the pigment bar just mentioned. After the fin rays appear their small mouth separates them from any of the large- mouthed flounders; their short, deep body, com- bined with the small number of fin rays, separates them from the witch; and the number of fin rays marks them off from the yellow tail (p. 273). The winter flounder also completes its metamorphosis at a smaller size than either of these other small- mouthed flatfishes (pp. 287 and 273). The rate of development of the larvae is gov- erned by temperature, occupying from about 2% to about 3^ months, according to the data available, and the larvae that are hatched later may catch up with the earlier hatched ones before their meta- morphosis takes place. Larvae in their later stages have been taken in abundance in the tow nets at Woods Hole. But their habits in aquaria suggest that they are less at the mercy of the tide and current than our other flatfishes are, for they have been described as alternately swimming upward and then sinking, to lie for a time on the bottom, instead of remaining constantly adrift near the surface, as the larvae of most of the flatfishes do at a corresponding stage in their development. At any rate, we have not taken any in our tow- ings in the open Gulf 63 that were certainly iden- tifiable as winter flounder. Judging from a large series from Casco Bay, measured by Welsh, and from others seen by us off near Boothbay Harbor and at Mount Desert, the fry of the previous winter grow to an average length of 1% to 3% inches by August, with an occasional specimen as long as 4 inches; they are 2 to 4 inches long by the end of September; and 4 to 6 inches long off southern New England in January and February, when nearing 1 year old, which probably applies north of Cape Cod as well. They may grow somewhat faster in more southern (warmer) waters, as in Chesapeake Bay, where fish of the year are 4% to 7 inches long in January and February.9* Welsh also concluded, from measurements gathered from various sources, that the winter flounders are 5 to 7% inches in length at 2 years of age, 7% to 9% inches at 3 years, and 9% to 10 inches long when 4 years old, which accords with 8 to 10 inches at 2 to 3 years in New York waters as reported by Lobell 65 and by Perlmutter.68 Prob- ably they mature sexually at 3 years, for most of the spawners are upwards of 8 inches long. Our only information as to the rate of growth of older lisli is that one tagged near Block Island, April 17, 1941, when it was 10% inches long, was 17 inches long when it was recaptured on Georges Bank, 4 years and 4 months later. •J Three larvae taken in the Oulf in July 1912, were provisionally identified by Welsh as this species. 210941—53 19 « Hildebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 13, Pt. 1, 1928, p. 169. « 28th Kept., New York Conserv. Dept. 1939, Sup., Pt. 1, p. 86. « Bull. Bingham Oceanogr. Coll., vol. 11, Art. 2, 1947, p. 17. 282 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE General range. — Atlantic coast of North America from the coast line out to the offshore fishing banks; common from the Strait of Belle Isle,67 the north shore of the Gulf of St. Lawrence where it has been characterized as "all along the coast," 68 and southern and southeastern Newfoundland to Ches- apeake Bay; recorded from the southern part of the Grand Banks,69 and as far north as Ungava Bay, northern Labrador;70 and from as far south as North Carolina and Georgia.71 Occurrence in the Gulf of Maine. — This is the commonest shoal water flounder, and perhaps the most familiar of all the ground fishes of the Gulf of Maine. There is no bay or harbor from Cape Cod to Cape Sable, no inter-island passage, and no stretch of open coast where it is not to be caught, unless the bottom be too smooth and hard, except, perhaps in the very turbid waters at the head of the Bay of Fundy. As one looks down at low tide from some pier where the water is clear enough, or from a boat, drifting over the flats, one is almost sure to see a flounder here and there, lying partly buried in the sand or mud. And they often come into water so shallow that it is easy to spear them. A flounder spear used to be almost as familiar an instrument along our coasts as an eel spear. With most of the flounder population of the in- ner parts of the Gulf living shoaler than 30 fathoms (20 fathoms is the deepest we have caught one there, close in to Little Duck Island, off Mount Desert), the zone occupied by them around the coast north of the elbow of Cape Cod is hardly as much as 8 to 10 miles wide, measured from the outer headlands or islands, except for Stellwagen Bank which lies a few miles farther out, and off Cape Sable, where their outer-depth limit lies something like 15 miles offshore. But their range extends out along the offshore rim of the Gulf, in somewhat deeper water, to include the Nantucket Shoals region as a whole (they must be plentiful " Jeffers (Contrlb. Canadian Biol., N. ser., vol. 7, No. 16, scr. A, General, No. 13, 1932, p. 210) reports it as not uncommon at Raleigh, on the New foundland side of the Strait. » Stearns, Proc. U. S. Nat. Mus., vol. 6, 1883, p. 125. •» At 2 stations, see Eept. Newfoundland Fish. Res. Lab., vol. 2, No. 3, 1938, p. 79. ™ Reported from Fort Chimo, Labrador by Kendall (Froc. Portland Soc. Nat. Hist., vol. 2, Pt. 8, 1909, pp. 225, 233); specimen in U. S. Nat. Museum, collected in 1882 or 1883 by L. M. Turner and identified by T. H. Bean. 71 Reported from Beaufort, N. C. (by Yarrow, Proc. Acad. Nat. Scl., Phila- delphia, vol. 29, 1877, p. 205); from the Neuse River, near New Bern, N. C. (by Smith, North Carolina Geo!, and Econ. Surv., vol. 2, 1907, p. 390); and from Georgia (by Hildebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 43, Pt. 1,1928, p. 170). to account for the 2 to 4 million pounds of black- backs and lemon soles that are brought in from there yearly) and from the shoaler parts of Georges Bank. The flounders on Georges run so much larger than they ordinarily do in-shore that they have been described as a separate species (p. 277). Dur- ing the summer of 1913 these soles (as they are called now, if they weigh more than 3 pounds) con- stituted about 4 percent by number of all the fish of all kinds that were caught on Georges by the several otter trawlers that carried investigators from the Bureau of Fisheries. Nowadays most every otter trawling trip brings in anywhere from a few hundred to several thousand of them accord- ing to depth and precise location on the bank. About 4 million pounds of lemon soles (larger than 3 pounds) and blackbacks (smaller than 3 pounds) were brought in from Georges Bank as a whole in 1947. They seem not to be so plentiful on Browns Bank, to judge from a catch of about 23,000 pounds of large sole and smaller blackbacks there by United States vessels in that same year. But much larger numbers are landed in the fishing ports along the outer coasts of Nova Scotia; about 420,000 pounds of flounders and soles combined, in 1946, the most recent year for which we have seen the Canadian Fisheries statistics. Fluctuations in abundance.- — Declining catches in the fyke nets that were used to take brood fish for the Booth Bay (Maine) hatchery leave no doubt that winter flounders were decidedly less abundant in that vicinity from 1934 to 1940 than they had been from 1925 to 1933. And some decrease in their abundance during the same period is indi- cated for the southern Cape Cod shore by the catch records of the Woods Hole hatchery; also along Connecticut and near New York, by the evidence of fishermen's logbooks.72 Importance. — The winter flounder, whether blackbacks or lemon soles, is the thickest and meatiest of all the flatfishes smaller than the halibut that are common on our coasts eastward and northward from the elbow of Cape Cod. In 1946 (most recent year when statistics are available for the Canadian catch as well as for the United States catch), the inner parts of the Gulf, from the tip of Cape Cod around^to Cape Sable, " For details, see Perlmutter, Bull. Bingham Oceanogr. Coll., vol. 11, Art. 2, 1947, pp. 6-13, who has made a special study of the blackback. FISHES OF THE GULF OF MAINE 283 yielded not far from 4 million pounds of flounders smaller than 3 pounds (blackbacks) to New- England fishermen,73 and about 49,000 pounds of fish heavier than 3 pounds (lemon sole). Nan- tucket Shoals, and the neighboring side of the so-called South Channel yielded about 5 million pounds of blackbacks and 1 million of soles; Georges Bank n about 3 million pounds of the larger soles and about 600,000 pounds of the smaller blackbacks. In addition to all this, Canadian fishermen caught some 4,400 pounds of flounders at the mouth of the Bay of Fundy on the New Bruns- wick side, 16,200 pounds of "flounders and soles" on the Nova Scotian side of the Bay, and 82,000 pounds off the west coast of Nova Scotia. Our Gulf as a whole thus yielded something like 14 million pounds of winter flounders, large and small, in the year in question, which seems to have been a representative one. Most of the commercial catch is made today by the otter trawlers, a small part on hook and line, or in nets of one sort or another. Years ago numbers were speared on the flats; as lately as 1919, about 7,000 pounds were reported as taken w The amount cannot be stated any more precisely because of uncertainty as to bow much of the 2H million pounds of black backs reported that year from Cape Cod was caught off the Oulf of Maine coasts of the Cape, and how much off the south shore of Massachusetts. N Including the statistical area classed as Eastern Side South Channel. in this way on Cape Cod. But flounder spearing has gone out of fashion so completely of late that no flounders, only eels, are listed under the heading "spears" in the Massachusetts landings by gear for 1945 or for 1946.75 Flounder fishing, too, for amusement and for home use goes on in harbors, estuaries, and other sheltered situations all around the shores of the Gulf, from bridges, piers, and small boats. And the number taken in this way must be very large in the aggregate for flounders are easy to catch (as well as very toothsome) provided the hook is not too large (Nos. 4 to 8 are best) and the bait is on bottom. Pieces of clam, of large snails, of sea worms (Nereis) or of squid, shrimp, and mussels, all are good. And they will take angle worms. Smooth flounder Liopsetta putnami (Gill) 1864 Smoothback flounder; Eelback; Foolfish; Christmas flounder; Plaice Jordan and Evermann, 1896-1900, p. 2650. Description. — -This flatfish is right-handed (eyes on the right side) and small-mouthed like the winter flounder, yellow tail, and the witch. It resembles the whiter flounder (with which it is often caught) closely in its general outline and in '• "Spears" are not Included for 1947; only "harpoons," for larger game. ■'i'.\NKvt;£$f4; "■ ,v -■-. ',*' > Figure 145. — Smooth flounder (Liopsetta putnami) , Salem, Mass. From Goode. Drawing by H. L. Todd. 284 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE the considerable thickness of its body. But it is distinguishable from the winter flounder by the fact that the skin of its head between the eyes is smooth and scaleless. Females are more easily recognized than males, their bodies also being smooth to the touch on both sides; males are nearly as rough skinned on the eyed side (except between the eyes) as the winter flounder, but they have much longer pectoral fins than the latter. Both sexes have fewer anal fin rays (only 35 to 40) and dorsal fin rays (about 56), too, while the caudal fin of the smooth flounder is narrower and more rounded than that of the winter flounder. The smooth flounder can always be separated from the yellowtail by the facts that its very prominent lateral line is straight, not arched, that the dorsal (left) profile of its head is straight, not concave; and that it has fewer fin rays. It has little more than half as many dorsal and anal rays as the witch, and its long fins are highest midway of the body and tapering toward the head and tail, whereas they are nearly uni- form in height from end to end in the witch. It lacks the mucous pits that are so characteristic of the blind side of the head of the latter, a convenient field mark for separating these two species. The smooth flounder is peculiar among our local flatfishes for its sexual dimorphism. Besides the difference in the scales of the two sexes noted above, the pectorals on the eyed side are longer (about four-fifths as long as the head) and more pointed on the males than they are on the females. Color. — The smoothback varies from grayish to dark muddy or slaty brown above, or to al- most black, either uniform or variously mottled with a darker shade of the same tint; the dorsal, anal, and caudal fins are of the general ground color. These fins were mottled darker or paler, in specimens we have examined, but Storer de- scribed them as black spotted. The blind side is white. Size. — This is the smallest flatfish that is com- mon in the Gulf of Maine, for it grows to a max- imum length of only about a foot, and to a weight of about a pound and a half. Habits. — This flatfish is confined to the close vicinity of the coast throughout its geographic range, occurring chiefly in estuaries or river mouths, and in sheltered bays and harbors; mostly on soft mud bottom. Correspondingly, it is found from tide line down to a maximum depth of per- haps 15 fathoms, with 2 to 5 fathoms as its zone of greatest abundance in our Gulf. It prefers soft bottom to hard; so much so that a seine haul on soft mud yielded 23 smooth flounders to 4 winter flounders in St. Mary Bay, whereas another haul, only 100 yards or so dis- tant, but on harder bottom, brought in only 3 smooth flounders to 189 winter flounders, as we learn from Dr. Huntsman's notes. The shoal water habit of the smooth flounder exposes it to temperatures close to the freezing point of salt water in winter, and as high as 60° in summer, and perhaps higher temperatures still in some places. Little more is known of its life. But its small mouth suggests a diet similar to that of the winter flounder, and Kendall found that yoimg fry 3 to 4 inches long from Casco Bay has been feeding chiefly on small crabs, shrimps, unidentified crustaceans, and polychaete worms. Winter is its breeding season, females nearly ripe having been taken in Salem Harbor in December and spent fish at Bucksport, Maine, the first week in March, which corroborates fishermen's reports of more than half a century ago that it comes into Salem Harbor to breed at about Christmas time. It is not known whether the eggs sink or are buoyant, nor have its larvae been seen. General range. — The smooth flounder is Arctic- boreal. It is definitely recorded from as far north as Ungava Bay, hence no doubt occurs along the Atlantic coast of Labrador; it is de- scribed as the most plentiful flatfish along the coasts of the Strait of Belle Isle at all seasons; 76 its young are common in Pistolet Bay on the New- foundland side of the Strait in shallow sun- warmed pools,77 and there are two specimens from the north shore of the Gulf of St. Lawrence in the Museum of Comparative Zoology (col- lected mary years ago, labeled "Labrador"). Evidently it is widespread on the southern side of the Gulf of St. Lawrence, for it is the next most plentiful flatfish after the winter flounder on the Cape Breton shore and at the Magdalens, accord- ing to Cox;78 it is reported from Prince Edward '• Jeffers (Contrib. Canadian Biol., N. Ser., vol. 7, No. 16 (Ser. A, No. 13 1922, p. 210). There are specimens from St. Anthonys, northern Newfound- land, in the Museum of Comparative Zoology. " Eept. Newfoundland Fishery Res. Comm., vol. 1, No. 4. 1932, p. 110. '• Contrib. Canadian Biol. (1918-1920). 1921 p. 113. FISHES OF THE GULF OF MAINE 285 Island,79 also from Trois Pistoles; 80 and it has been classed by Huntsman 81 as characteristic of the estuarial transition from fresh to salt waters in the southern side of the Gulf of St. Lawrence generally. We find no record of it on the outer coast of Nova Scotia between Cape Breton and Cape Sable; but we suspect that it has been over- looked there, for it is widespread in the Gulf of Maine to Massachusetts Bay, as detailed below, and has been reported as a stray as far south as Providence, R. I. Its range probably is continuous in the north with that of its polar relative (L. glacialis) of the Arctic coasts of North America and Siberia. In- deed, it is a question whether any valid distinction can be drawn between the two species. Occurrence in the Gulf of Maine. — The smooth flounder is to be found in estuaries, river mouths and harbors, all along the shores of the Gulf, from the Bay of Fundy to the northern side of Massa- chusetts Bay. Localities whence it has been re- corded in print, or has been definitely reported otherwise, are Annapolis basin, Minas Channel and St. Mary Bay; Grand Manan; Bucksport at the mouth of the Penobscot River; Belfast in Penobscot Bay; Casco Bay; Portland; Salem Harbor; and Boston Harbor. Apparently the latter is the southern limit to its regular occur- ence for while there is a specimen in the Museum of Comparative Zoology, from Provincetown at the tip of Cape Cod, it seems to be unknown in Cape Cod Bay, along the outer shore of Cape Cod, or in the Woods Hole region, though a stray individual has been caught at Providence, R. I.82 This flatfish (often confounded with the winter flounder) has been found so often in various mar- kets among the winter flounders as to suggest that it is more plentiful along the coasts of northern New England, than is realized, general!)'. In Casco Bay and in estuaries of the Bay of Fundy such as the mouths of the St. CroLx and Annapolis Rivers it is abundant in summer, which no doubt applies equally to the intervening coast line. But it is said to run up into harbors in Massachusetts Bay in autumn and winter only;83 '» Cornish, Contrib. Canadian Biol. (1906-1910) 1912, p. 81. «° Vladykov and Tremblay, Nat. Canad., vol. 62 (Ser. 3, vol. 6), 1935. p. 82; (many specimens reported). »' Trans. Roy. Soc. Canada, Ser. 3, vol. 12, Sect. 4, 1918, p. 63. •' This specimen, formerly in the Museum of Comparative Zoology, is no longer to be found. 8* Our experience corroborates this to the extent that we have never seen il there in summer. nor would such a local difference be astonishing in the case of a cold-water fish, which might well be driven out into slightly deeper water by summer heat in the southern and western parts of the Gulf, but not in the northern and eastern parts. Commercial importance. — This is an excellent table fish for its size, sweet-meated and thick- bodied like the winter flounder. But it is neither large enough, plentiful, nor widely enough dis- tributed in the open Gulf to be of any commercial importance. Witch flounder Glyptocephalus cynoglossus (Lin- naeus) 1758 Gray sole; Craig fluke; Pole flounder Jordan and Evermann, 1896-1900, p. 2657. Description. — The witch or "gray sole" as it is now named in the United States fishery statistics, is right-handed (viscera on the right hand as the fish lies) and small-mouthed like the winter flounder, the smooth flounder and the yellow tail. But there is little danger of confusing it with any of these for its fin rays are much more numerous, its body narrower relatively, its head much smaller, and the open mucous pits on the blind side of its head large and conspicuous. It is two and one- half to three times as long as it is broad (deep, in reality), elliptical in outline, very thin but with its head occupying only about one-fifth of the total body length, and it has a very small mouth. The dorsal (left-hand) profile of its head is convex. It has 100 to 115 dorsal fin rays and 87 to 100 anal rays, and the anal fin is preceded by a short, sharp spine pointing forward, which is a prolongation of the post-abdominal bone. The two long fins are of about uniform width throughout most of their lengths, except that they narrow gradually toward head and tail. The pectoral fins and the ventral fins are alike on the two sides, or nearly so, while the caudal fin is much smaller, relatively, than that of the yellowtail, of the winter flounder, or of the smooth flounder, though similarly rounded in rear outline. The lateral line is straight, as a rule, but it is somewhat arched abreast the pectoral fin in some specimens. The teeth are small, incisorlike, and in a single series. There are about 12 open mucous pits or depressions on the blind side of the head, and less obvious ones on the eyed side also. The whole body and head (except for the tip of the 286 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 146. — Witch flounder (Glyptocephalus cynoglossus) . From Goode. Drawing by H. L. Todd. gnout and the lower jaw) are scaly, but the scales are smooth to the touch, which make the witch as slippery to hold as a female smooth flounder (p. 284). Color. — By all accounts (and the fish we have seen are in line with this) the witch is less variable in color than most of the flatfishes. Most of them are brownish or russet gray on the eyed side, either uniform or with darker transverse bars, with the vertical fins of the general body hue, tinted or tinged with violet, and either plain or spotted. The pectoral fin membrane on the eyed side is dusky or even black, a feature distinctive of this particular flatfish. The lower (blind) side is white, and more or less dotted with minute dark points. An occasional fish is colored on the under side as well as on the upper side; one of this sort, 19 inches long, was landed at the Boston Fish Pier early in March 1931. Size. — The maximum length is about 25 inches, and fish of 23 or 24 inches, weighing about 4 pounds, are not uncommon. But the general run of those caught are only about 12 to 20 inches long. Habits. — The witch flounder is rather a deep- water fish, seldom caught shoal er than 10 or 15 fathoms once it has taken to bottom, though taken occasionally close inshore (see footnote, p. 288). Off the American coast the best catches are made between about 60 fathoms and about 150 fathoms. Thus the Albatross III caught an average of about 57 witch flounders per trawl haul at 100 to 150 fathoms on the southwestern part of Georges Bank in mid-May f 950. but an average of only about one fish per haul between 31 fathoms and SO fathoms. And they have been trawled widespread down the continental slope as deep as 858 fathoms off southern Nova Scotia; to 732 fathoms off Marthas Vineyard; to 788 fathoms off Chesapeake Bay; and to 602 fathoms off North Carolina.84 In Swedish waters, according to Melander,85 the best catches are made between 80 and 140 fathoms. They are caught most abundantly on fine muddy sand, on clay, or even on mud. They are said to frequent hard reefs in Scandinavian waters, but this does not seem to be the case in the Gulf of Maine, though they are common there on the smooth ground between rocky patches. When the witch has once taken to the bottom it seems to be even more stationary in our gulf than some other flounders, for it is caught the year round, with no evidence that it moves in or off shore with the change of the seasons. In Swedish waters, however, it is said to work up into shoaler water in autumn, and deeper again in late winter and spring.86 It occurs in the Gulf of Maine in temperatures ranging from about 35°-38° F. (late winter and early spring), to 45°-48° (late summer and early autumn), according to precise locality and depth. In the Gulf of St. Lawrence it occurs in the icy « Goode and Dean (Smithsonian Contr. Knowl., vol. 30, 1895. p. 433) give a long list of deep-water stations for the witch off southern New England. » Pub. de Circonstance No. 85, Cons. Internat. Esplor. Mer, 1955, p. 3. « Melander, Pub. de Circonstance, Cons. Internat. Explor. Mer. No. 95, 1925, p. 3. FISHES OF THE GULF OF MAINE 287 cold waters (30°-32°) on the banks as well as in the higher temperatures (40°-42°) of the deep channels.87 Apparently it is never found in any numbers in water warmer than 50°, but we hesitate to propose high temperature as the factor barring it from shoal water because there is no evidence that it works inshore in our gulf in winter when this bar would not operate. Food. — It feeds on invertebrates, like other small-mouthed flatfishes; European experience points to small crustaceans, starfish, small mol- lusks, and worms, as its chief diet.88 It is not known to eat fish and does not take a bait often. Breeding as it does through a long season, over many degrees of latitude, and in both sides of the Atlantic, the witch spawns in temperatures ranging from close to the freezing point of salt water up to 48°-50° F. (p. 288). And experiments, added to captures of eggs naturally spawned, and of newly hatched larvae, have shown that incubation pro- ceeds normally in water at least as cold as 45°-46° F., and as warm as 50°-55° F. The eggs are buoyant, spherical, transparent, with narrow perivitelline space (the perivitelline space is broad in the eggs of the dab or Canadian plaice, which overlap them in dimensions), without oil globule, and 1.07 to 1.25 mm. in diameter. As noted (pp. 288 and 203), there is danger of confusing •' According to Huntsman, Trans. Roy. Soc. Canada, Ser. 3, vol. 12, Sect. 4, 1918, p. 63. •• No witch-flounder stomachs have been examined In the Oulf of Maine, so far as we know. newly spawned witch eggs with those of the cod and haddock, for they overlap these in size and in season. But identity is easily recognizable after a few days' incubation, for black pigment is to be seen in the gadoid eggs soon after the embryo is visible as such, but does not appear in the witch- flounder eggs until after hatching. Incubation occupies 7 to 8 days at temperatures varying from 46° to 49° F., and the newly hatched larvae are about 4.9 mm. long, with a larger yolk sac than those of our other flatfishes. The yellow and black pigment becomes aggregated into five transverse bands on body, yolk (now much reduced in size), and fin folds within a few days after hatching, when the larva is 5 to 6 mm. long. One of these bands is at the region of the pectoral fin, one at the vent, and three of them on the trunk rearward from the vent. The yolk is entirely absorbed in about 10 days after hatching, the caudal rays have begun to appear at a length of 15 mm., the rays of the vertical fins are well advanced at 21 mm. and they are complete in their final number at about 30 mm. The eyes are still symmetrical, or nearly so, up to this stage. But the left eye has moved to the dorsal surface of the head in larvae of about 40 mm. And the migration of the eye is complete at a length of 40 to 50 mm., when the young fish takes to the bottom. The witch is perhaps the most easily recogniz- able of Gulf of Maine flatfishes throughout its Figure 147. — Egg (European). After Cunningham. TV Figure 149. — Larva (European), 16 mm. After Kyle. Figure 148.— Larva (European), 10 days old, 5.6 mm. Figure 150.— Smallest bottom stage (European), 42 mm. After Holt. After Petersen. Witch flounder (Glyptocephalus cynoglossus) 288 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE larval stage. The transverse pigment bars are diagnostic prior to the appearance of the caudal rays, while the curiously concave ventral profile of the throat region with the comparatively long slender trunk are equally so, thereafter. And the great number of dorsal and anal fin rays, coupled with the small mouth, make identification easy after the fins are formed. The witch also grows to a larger size before it completes its metamorphosis than does any other of the right- handed, small-mouthed flatfishes that are found in the Gulf of Maine. Measurements of the young (American as well as European) , suggest that the free-drifting stage may last as long as 4 to 6 months for the witch, which is much longer than for any of our other flatfishes. Fry of 2K to 4% inches, and of 3K to 4% inches, such as we have trawled in July and August, re- spectively, probably are in their second summer, their sizes depending on how early in the season they were hatched the year before. The sub- sequent rate of growth has not been traced for American fish. If Molander's 89 estimate for European fish is correct, the size group centering at 6% to 8 inches that was prominent in our August catches of 19.36 were in their third summer.90 And subsequent growth is very slow. General range. — Moderately deep water in both sides of the North Atlantic. Its European range is from northern Norway and Iceland south to the west coast of France. In American waters its free-drifting larvae are reported from as far north as the Strait of Belle Isle, around the coasts of Newfoundland, and over the Grand Banks region in general.91 The adult is known from the Gulf of St. Lawrence; the south coast of Newfoundland; the southern part of the Grand Banks; in Cabot Strait; along outer Nova Scotia and the Scotian Banks; throughout the Gulf of Maine; and thence westward and southward along the continental shelf and slope as far as the offing of northern Virginia (lat, 37° 50' N.) in moderate depths, to the offing of Cape Hatteras in deep water.92 Occurrence in the Gulf of Maine.- — The distribu- tion of this flatfish in our Gulf is governed by the " Based on the structure of the otoliths; Pub. de Circonstance, No. 85, Cons. Perm. Intemat. Explor. Mer. 1925, pp. 12-14. « Bigelow and Schroeder, Biol. Bull., vol. 76, 1939, pp. 318-319. •i See Frost, Res. Bull. No. 4, Newfoundland Dept. Nat. Resources, 1938, Chart 6, for Newfoundland localities. m Goode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, p. 433) list it from lat. 34° 39' N., 603 fathoms. fact that it is a fish of at least moderately deep water, seldom caught as shoal as 10 fathoms.93 In fact, its very existence remained unsuspected by Massachusetts fishermen until 1877, when the United States Fish Commission caught numbers of them while trawling in the deeper parts of Massachusetts Bay. Since that time it has been reported (or we have trawled it, or both) from St. Mary Bay on the Scotian side of the Gulf; in the Bay of Fundy and its tributaries (where Hunts- man describes it as taken very generally below 15 fathoms, if not in any great numbers) ; at East- port; off Mount Desert, where we have trawled it as shallow as 10 fathoms; near Monhegan Island; off Seguin Island ; off Cape Porpoise ; near the Isles of Shoals (where Welsh saw a few taken from the gill nets set in about 25 fathoms in April 1913) ; in the deep trough to the westward of Jeffreys Ledge; in Ipswich Bay; near Gloucester; off Boston Har- bor; at various localities in the deeper parts of Massachusetts Bay; and in both branches of the deep trough of the Gulf west and east down to a depth of 140 fathoms; in the deep channel, between Browns Bank and Georges Bank, and on the slope to the southeast. Trawlers bring them in regularly from Browns Bank, also from Georges, where Welsh found them widespread, and from Nantucket Shoals. This is enough to show that the witch is to be expected anywhere in our Gulf where the water is deeper than 15 to 20 fathoms, if the bottom is suitable. The largest catches are made on the so-called South Channel grounds which include the slopes that lead down from the offing of Cape Cod on the one side and from Georges Bank on the other, into the southwestern part of the basin; farther north off eastern Massachusetts; and off western Maine. And the published statistics suggest that gray soles are about as plentiful as the American dabs are on the various grounds where the trawlers work regularly. Reported landings of gray soles by New England vessels in 1947 were as follows for the several statistical areas:94 Browns Bank, 44,000 pounds; off western Nova Scotia, 2,000 pounds; off eastern Maine, 17,000 pounds; off central " A stray specimen, picked up in a pound net at Eastport, Maine, many years ago, was reported by Gill (Proc. Acad. Nat. Sci., Philadelphia, 1873, p. 360) as a new species, Glyptocephalus acadianus. « To the nearest 1,000 pounds. FISHES OF THE GULF OF MAINE 289 Maine, 12,000 pounds; off western Maine, 630,000 pounds; small grounds in west central part of Gulf, 77,000 pounds; off eastern Massachusetts, 582,000 pounds; South Channel grounds, east and west, 629,000 pounds; other parts of Georges Bank, 94,000 pounds; Nantucket Shoals region, 16,000 pounds.96 More precise evidence as to their local numbers on suitable bottoms in the appropriate depths is that as much as 500 pounds have been taken in a 15- to 20-minute haul with a small beam trawl in Massachusetts Bay, and that we caught 48 of them in Ipswich Bay in 22 fathoms, in a short haul with an 8-foot beam trawl on July 16, 1912. We also saw 519 of them, 10 to 22 inches long, trawled on the southwestern part of Georges Banks by the Eugene H, in 41 hauls at 26 to 65 fathoms in late June 1951, and learned that this dragger caught 9,000 pounds on the northeastern edge of Georges Bank, in 85 to 95 fathoms, October 12-18, 1951. Neither the witch flounder nor the American dab is as plentiful as the yellowtail on good flounder bottoms, or the flatfishes of the winter flounder group (blackbacks plus lemon soles). And At- lantis took only 156 witch to 279 dabs on soft bottom at 90 to 103 fathoms during experimental trawling in the deeper parts of the Gidf in August 1936. Gray soles are at least moderately plentiful off southern New England. The Albatross III, for example, took 90 there in one trawl haul at 101 to 150 fathoms in mid-May 1950, a few as shoal as 31 to 40 fathoms. And a few thousand pounds are landed yearly in New York and in New Jersey ports.98 But records of the witch from farther south than New Jersey are of an occasional fish only. Reported landings suggest that gray soles are about as plentiful all along the Nova Scotian banks as they are in the Gulf of Maine region. In 1947, for example, New England vessels landed about 555,000 pounds of them from the various grounds from the eastern part of Browns Bank to Banquereau, about half of which came from the Horseshoe ground between Halifax and Sable Island.97 And they seem to be moderately plenti- »• An additional 182,000 pounds were landed in Cape Cod ashing ports, source not stated. •• About 19,000 pounds in New York In 1947, about 28,000 pounds in New Jersey. •» Only a lew thousand pounds are reported yearly from Nova Scotia, in the Canadian fishery statistics. 210941—53 20 ful in the southern part of the Gulf of St. Lawrence, for Cox 9S wrote of many (large and small) as taken off the Cape Breton shore, and in Cabot Strait off Cape North. But no information is available as to then numbers elsewhere in the Gulf of St. Lawrence, or on the Grand Banks. It seems that the witch does not breed success- fully in the Bay of Fundy; at least its eggs have never been found there, nor have its larvae. But probably it does so in other parts of the Gulf in general, including the offshore Banks, though our only positive egg records for it have been off Pe- nobscot Bay, and at the mouth of Massachusetts Bay. And there is no reason to doubt that the more northerly populations are equally self sup- porting, for the pelagic larvae have been taken at many localities on the more easterly of the Nova Scotian Banks; in the Gulf of St. Lawrence; over the Grand Banks; and along the south and east coasts of Newfoundland, by the Canadian Fisheries Expedition of 1915, and during the cruises of the Newfoundland Fishery Research Commis- sion more recently.99 But there is no evidence that the witch spawns to any extent to the west of Cape Cod. Captures of eggs, certainly of this species, in our tow nets in July and August, with larvae up to 20 to 23 mm. long as early as the first week of July, but others as small as 9 to 10 mm. as late as mid-October, show that the witch is a late spring and summer spawner in the Gulf of Maine as it is in European waters also, with the peak of produc- tion probably falling in July and August. Thus its spawning season overlaps that of the haddock, (p. 207). Its eggs are shed in the Gulf of Maine in tem- peratures ranging from 39° to 41° F. at the begin- ning of the season, to 43° to 48° in midsummer. But (being buoyant) the temperature may be con- siderably higher at the level where their develop- ment takes place than deeper down where the spawning fish lie. In fact, it is doubtful if any eggs develop in our Gulf in water as cold as 42° to 43°. Neither is there any reason to suppose that witch eggs develop in water any colder than this in the Gulf of St. Lawrence, or off Newfoundland, for the surface stratum to which they rise after « Contrib. Canad. Biol. (1918-1920) 1921, p. 113. •• See Frost, Research Bull. 4. Newfoundland Dept. Nat. Resources, 1938 Chart 6, for Newfoundland localities. 290 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE they are shed is comparatively warm (upward of 45°) in these seas also, during the spawning season. At the other extreme, our captures of eggs and of newly hatched larvae near the surface in July prove that the latter may be hatched in the Gulf in water at least as warm as 50° to 55°. But the upper limit to normal development cannot be stated from the evidence yet in hand, for with a temperature gradient as steep as it is over most of the Gulf of Maine in summer a difference of only a few fathoms in the depth at which the eggs or young larvae are suspended may mean a difference of several degrees of temperature. One result of the protracted spawning season, combined with the long period occupied by larval development, is that witch larvae of various sizes are to be taken in tow nets throughout the summer and early autumn, as appears from the following table of our catches on the Grampus. Date Num- ber of larvae Length in millimeters Date Num- ber of larvae Length in millimeters July 7, 1915 July 8, 1913 July 9, 1913 July 19, 1916 July 22, 1912.... July 24, 1912.... Aug. 5, 1913 Aug. 9, 1913 Aug. 14,1912... 109 19 1 100+ 1 2 27 7 1 8 to 23.5. 8.5 to 21.6. 14. 5 to 19. 9.5. 8.5 and 16.5 5.5 to 12.5. 10 to 23. 18.5. Aug. 15,1912... Aug. 24, 1912... Aug. 25, 1914 Aug. 26, 1913... Aug. 29, 1916... Aug. 31, 1912... Sept. 29, 1915... Oct. 18, 1915-.. Nov. 1, 1916 3 6 19 2 100+ 20+ 22 1 20+ 18.5 to 37.5. 10 to 18. 10 to 19. 8 and 14. 5 to 19. 9 to 16.5. 10 to 14. 9 5 29.5 to 50. All of these catches, like those for other larval flatfishes, and for larval gadoids, have been con- centrated in the southwestern part of the Gulf, which must be an important nursery for the witch also. And we may note in passing that the presence of young fry at all stages from immedi- ately after their metamorphosis (that is, 4 to 6 months old) in the Bay of Fundy, where few or none are hatched, points to an immigration of the late larvae, or of the youngest fry, into the Bay, either just before they take to the bottom or soon after they have done so. Importance. — The witch was of no commercial importance in our Gulf a quarter of a century ago; few fishermen distinguished it from other flounders then, and no record was kept of the catch. It is an excellent table fish; and the bases of its fins are provided with astonishingly large amounts of ge- latinous fat for so thin a flounder, of the sort for which the European turbot is famed. It is now in such demand that it brings about as high a price as either the yellowtail or the American dab. In 1947, for instance, the average price at Massachusetts ports was about 7 cents for gray sole, about 8 cents for yellowtad, about 7 cents for dab, about 9 to 12 cents, according to size, for flatfish of the winter-flounder type (blackbacks and lemon sole), about 17 cents for summer flounders (fluke), and about 25 cents for halibut. The Gulf yielded between 2 million and 2% million pounds of gray sole both in 1946 and in 1947, corresponding to something like 1 to 1% million individual fish. The otter trawl is the only gear now in use in our waters that is adapted to the capture of witch flounders on a commercial scale. They live too deep, and their mouths are too small for them to be of any concern to small-boat fishermen. Sand flounder Lophopsetta, maculata (Mitchill) 1814 Windowpane; Spotted flounder; New York plaice; Sand dab; Spotted turbot Jordan and Evermann, 1896-1900, p. 2660. Description. — This is the closest North American relative of the European turbot and brill. It is left-handed (eyes and viscera at the left-hand side) and large-mouthed, like the summer and four- spotted flounders, but it is readily separable from both of these by the outlines of its ventral fins. In all other Gulf of Maine flatfish (except for the hogchoker, p. 296) these are narrow at the base and widen toward the tip, but the ventrals of the sand flounder are as wide at the base as they are at the tip, each simulating a detached segment of the anal fin. Furthermore the two ventral fins are not alike either in location or in size, the left-hand (upper) fin, which is the longer of the pair, being practically a continuation of the anal fin so far as its appearance goes, whereas the right-hand (lower) ventral fin is situated a short distance up the right-hand side of the throat. The general appearance of the dorsal fin is no less diagnostic, for its first 10 or 12 rays are not only free from the fin membrane along the outer half of their lengths, but they are branched toward their tips, so that they form a conspicuous fringe which is without parallel among Gulf of Maine flatfishes. Further- more, the sand flounder is more nearly round in outline than any of our other local flatfishes (it is only about one and one-half times as long as it is FISHES OF THE GULF OF MAINE 291 broad), and so thin through that its body is trans- lucent when it is held up against the light. Its pectoral fins, too, are longer than in our other left- handed flatfishes; its caudal fin is more rounded; and its teeth smaller although the gape of the mouth is wide. The dorsal (right) fin (63 to 69 rays) tapers toward the tail; the anal (left) fin (46 to 52 rays) tapers toward head and tail, while both of these fins are noticeably thick and fleshy at the base; and there is no free anal spine. The pectoral fin on the eyed side is longer and more pointed than its mate on the blind side; the scales are smooth to the touch ; and the lateral line is bowed abreast of the pectoral fin. Color. — The sand flounder varies less in color than most shoal-water flatfishes do, the general ground tint of its eyed side (both as described by previous authors and in those we have seen) being of a pale and rather translucent greenish olive or slightly reddish or light slaty brown more or less mottled with darker and paler, and usually (if not always) dotted with many small brown spots of irregular shapes. Some fish are also marked on the body and on the bases of the dorsal, anal, and caudal fins with while spots that vary in number and in size from fish to fish. But others lack these spots. The dorsal, anal, and caudal fins are of the general body tint, more or less mottled with darker, while the pectoral of the eyed side is dark crossbarred or speckled. The blind side is white in most of them, but specimens have been seen on which it was irregularly dark-blotched.1 Size. — The sand flounder is said to grow to a maximum length of 18 inches and to a weight of 2 pounds. But the largest we have seen (from Waquoit on the southern shore of Massachusetts) , were about 15 inches long. And adult fish run only about 10 or 11 to 12 inches in length. Sand flounders from southern New England measured by Moore 2 averaged about ){ pound at 8 inches : about ){ pound at 10 inches; about % pound at 12 inches; and a little more than 1 pound at 14 inches. Habits.- — The sand flounder is a shoal-water fish. Its upper limit is close below the tide mark, and the 20 to 25 fathom line probably marks its lower limit, in general, in the coastal zone north of Cape Cod. But Moore reports it as occurring regularly down to 27 fathoms off Connecticut, and > Moore, Bull. Bingham Occanogr. Coll., vol. 11, art. 3, 1947, p. 20. » Bull. Bingham Owanogr. Coll., vol. 11, art. 3, 1947, p. 53, fig. 12. Figure 151. — Sand flounder (Lophopsetta maculala). From Jordan and Evermann. Drawing by H. L. Todd. 292 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Welsh saw it taken on Georges Bank down to 30 or 40 fathoms, while the Albatross III trawled a few on the southwest part of the Bank along this same depth zone in May 1950. It is caught chiefly on sand bottom off southern New England and southward, as its name implies, but its comparative abundance in Casco Bay and in Minas Channel shows that it also frequents softer and muddier grounds in the Gulf of Maine. The sand flounder is a year-round resident off the southern New England coast, and probably this applies to it in the Gulf of Maine also, there being no evidence that the adults carry out any migrations inshore or offshore, with the change of the seasons. But such of the young fry as settle to bottom in shallow water inshore tend to work offshore as they grow, and deeper, while tagging experiments off southern New England have shown that individual sand flounders may wander along the coast for considerable distances, or across open water, much as winter flounders do (p. 279) . Some of them went as far as 80 miles in 3 months.3 And it is probable that the wanderings of the adults play an important part in the intermingling of local populations. The adult sand flounder is necessarily attuned to a wide temperature, occurring as it does over many degrees of latitude, and in shallow waters where it is exposed to the extremes of winter chilling and of summer warming. Such of them as winter in shoal bays experience winter tempera- tures close to the freezing point of salt water in winter, not only in the northern part of their range, but even as far south as the Connecticut shore.4 And it is probable that the entire popula- tion hi the Gulf of Maine winter in water colder than 36° F. But these same fish summer in temperatures of 50° to 70°, according to locality and depth. And some sand flounders summer in still higher temperatures farther south. Never- theless, it seems that temperature is the factor that governs the northerly range of the species and its local abundance, for it is only where the surface waters warm to 55° or higher in summer, as happens in Massachusetts Bay, in Casco Bay, in Minas Channel, and over the southern shallows • For details, see Moore's (Bull. Bingham Oceanogr. Coll., vol. 11, art.3 1947, pp. 58-63) detailed study of the sand flounder in southern New England waters. * Warfel and Merriman (Bull. Bingham Oceanogr. Coll., vol. 9, art. 2, 1944. pp. 61-62) give details of temperature and salinity for Connecticut waters, with references. of the Gulf of St. Lawrence, that the sand flounder is able to maintain itself in any numbers. Ap- parently either its eggs or its young larvae, or both, fail to develop in lower temperatures (p. 293) . And these isolated breeding centers are not pro- ductive enough to stock the intervening stretches of shoreline in the case of a fish as stationary as the sand flounder. Thus its distribution is some- what analagous to that of the oyster. The large mouth of the sand flounder suggests that it feeds on active prey. Welsh, in his field notes, remarked, in fact, that sand flounders caught off Atlantic City, N. J., were full of "schizo- pod shrimps" (mysids) and of these alone, and mysid shrimps (Neomysis americana) had similarly been the predominant item in all months of the year, for 654 Long Island Sound fish examined by Moore,5 with shrimps of other kinds ranking second. Moore also concluded that the few fishes included in their diet were not enough to class the sand flounder as a fish eater. But hake, herring, launce, and silversides have been found in their stomachs at Woods Hole, while North Carolina specimens had eaten fish, also crabs and shrimps.8 And we suspect that they seize small fish whenever they can, for we once hooked a sand flounder only about 12 inches long on a 2%-ounce metal jig, while we were casting for striped bass in the surf on Orleans Beach, Cape Cod. A variety of small invertebrates other than shrimp have also been found in their stomachs; Vinal Edwards noted annelid worms, crabs, squid, small mollusks, ascidians and even seaweed, to which Moore adds gammarids and other small Crustacea, worm tubes, sea cucumbers (holo- thurians), glass worms (Sagitta), and sand. A larval sand flounder 11.5 mm. long examined by Moore7 contained minute copepods (Temora and Centropages) and amphipods (Unciola and Lepto- cheirus). The sand flounder is a late spring and summer spawner in the northern part of its range; thus Welsh found them spawning late in June at Gloucester, and ripe fish are taken at Woods Hole in May and June, while Moore reports sand flounders ripe in Long Island Sound from early May to August, with some still incompletely spawned out there in September. And it seems ' Bull. Bingham Oceanogr. Coll., vol. 11, art. 3, 1947, pp. 54-58. • Smith, North Carolina Econ. Geol. Survey, vol. 2, 1897, p. 392. 'Bull. Bingham Oceanogr. Coll., vol. 11, art. 3, 1947, pp. 26-27 FISHES OF THE GULF OF MAINE 293 that they commence spawning still earlier in the season to the westward and southward, for Nichols and Breder8 report young fry 20 mm. long in Sandy Hook Bay by May, while the sizes of the young fry taken in winter in Chesapeake Bay suggest that they are hatched there as early as March or April.9 It is not yet possible to state the extremes of temperature within which the sand flounder spawns. But 50° to 60° F. has proved favorable for hatching artificially fertilized eggs at Woods Hole, with even 70° not too warm for successful incubation. The eggs are spherical, transparent, buoyant, 1 to 2 mm. in diameter (measurements taken at Gloucester by Welsh), with a single color- less or pale-lemon oil globule of 0.15 to 0.28 mm. And the surface of the egg shows faint irregular markings. Incubation occupies about 8 days at 51°-56°; its duration has not been recorded for higher temperatures. The sand flounder, like the winter flounder, completes its metamorphosis while it is smaller than either the yellow tail (p. 273) or the witch (p. 287). Thus the dorsal and anal fin rays were complete and the ventral fins had formed in one only 8)4 mm. long (fig. 153), and its right-hand eye had already moved around to the back-line of the head, while the migration of the eye is com- pleted, and they are ready to take to bottom by the time they have grown to 10 mm. long.10 Rate of Growth. — It seems that the sand flounder passes through its larval stage more rapidly than most flatfishes do, for many of its fry with the migration of the eye completed have been taken at Woods Hole only ] to 2 months after spawning commences there. One that was kept in an aquarium there by Williams lI grew from 10 mm. to 22 mm. in length in 11 days; and Nichols and Breder's I2 observation that fry of the year in Sandy Hook Bay grew from an average length of about %-inch (to the base of the tail fin) in May, to about 2 to 2/8 inches by late September, is in line withTra,cy's statem ent 13 that the fry are 2 to 3 inches long in July in Rhode Island water's, growing to ' Zoologies, New York Zool. Soc., vol. 9, 1927, pp. 181-182. • Hildebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 43, Pt. 1, 1928, p. 172. '• Williams (Bull. Mus. Comp. Zool., vol. 40, 1902, No. 2) has given a brief account of the anatomical changes that take place during the passage of the eye In the sand flounder, and a more detailed account for the winter Sounder. For photographs of larvae and small fry, see Moore (Bull. Bingham Oceanogr. Coll., vol. 11, art. 3, 1947, fig. 3). ii Bull. Mus. Comp. Zool., vol. 40, 1902, p. 3. » Zoologies, New Ycrk Zool. Soc, vol. 9, 1927, pp. 181-182. H Kept. 40, Comm. Inland Fish. Rhode Island, 1910, p. 106. 4 inches and upwards in December. Fry only 1 to 2 inches long reported by Nichols and Breder at Orient, N. Y., in December, seem to have been from a late-hatched brood. Figure 152. — Larva, 5.5 mm. Figure 153. — Larva, 8 mm. Sand flounder (Lophopsetta maculata). Moore concludes, from her very detailed study of the growth zones on scales and otoliths, that sand flounders in Long Island Sound average about 4}£ inches long when they are 2 years old (i. e., at the beginning of their third summer); about 7K inches at 3 years; about 9 to 10 inches at 4 years; about 11 inches at 5 years; about 11 ){ inches at 6 years; and about 12 inches at 7 years.14 And Gulf of Maine fish probably grow at about this same rate. They mature at 9 to 10 inches; i. e., in the third or fourth year, according to the fore- going schedule. General range. — Coastal waters of eastern North America, from the Gulf of St. Lawrence to South Carolina; most abundant west and south of Cape Cod, north and east of which it is confined to favorable localities. Occurrence in the Gulf of Maine. — This flounder is not common in the Gulf of Maine, except locally. Dr. W. C. Kendall found it at Monomoy; we have caught one (p. 292) on the outer shore of Cape Cod; Storer found it at Province town, where he saw a considerable number in shoal water; it is reported from North Truro; from Gloucester » Bull. Bingham Oceanogr. Coll., vol. 11, art. 3, 1917, pp. 47-51. 294 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Harbor, where a considerable number were col- lected in 1878 (Welsh found it there in 1916), and at Milk Island nearby. But we have not learned of it anywhere else in the Massachusetts Bay region, and it has never been recorded between Cape Ann and Cape Elizabeth, nor did Welsh see it taken there by the gill-netters during the spring of 1913. It has been reported repeatedly at several localities in Casco Bay, which seems to be a local center of abundance. But it cannot be common along the eastern Maine coast or on the New Brunswick side of the Bay of Fundy, for the only records from this stretch of coastline are from Bucksport, from Eastport, and from Passama- quoddy Bay where one was taken in 1880 and another in 1912. Minas Channel on the Scotian side seems to be a second center of abundance, like Casco Bay, for Leim found it common there.15 Huntsman reports it in St. Mary Bay also. But we have found no other record of it along the western coast of Nova Scotia. Welsh saw it taken by the otter trawlers on Georges Bank in June 1913, and we have seen it there on four recent trawling trips, including about a dozen specimens trawled by the Albatross III on the southwest part of the bank and off Nan- tucket in 22 to 39 fathoms, in mid-May 1950, and 132 taken by the Eugene H in that same general region, in 36 hauls at 25 to 45 fathoms, in late June 1951. Beyond this, nothing is known of it on the offshore fishing grounds. The evidence of the Gloucester specimens men- tioned above proves that it breeds in the Massa- chusetts Bay region to some extent, while its local abundance suggests the same for Casco Bay, as does the capture of its larvae for Minas Channel. It may also breed at the heads of the warmer and shoaler bays between Casco Bay and Grand Manan. Seemingly it does not do so in any of the estuaries on the New Brunswick side of the Bay of Fundy for no larvae have ever been found in Passamaquoddy Bay, a fairly representative situation, probably because of low temperature. But we have no doubt that the local stocks in the Gulf of St. Lawrence (p. 294) are self-sustaining. The sand flounder is much more plentiful west of Cape Cod than it is anywhere in the Gulf of Maine, southward at least to Chesapeake Bay, where it is very generally distributed in depths down to 25 fathoms, especially in the southern '« Huntsman. Contrlb. Canadian Biol. (1921), No. 2. 1922, p. 70. part. And it is reported as common at Beaufort, N. C.16 The sand flounder is known only here and there to the eastward and northward of our Gulf. Its pelagic larvae have been reported on Middle Ground off Halifax and near Sable Island ; 17 a few adults have been taken in Chedabucto Bay, eastern Nova Scotia; IS Cox 19 states that it is "by no means uncommon" around the Magdalen Islands, in the southern side of the Gulf of St. Lawrence, where Huntsman ^ classes it as charac- teristic of the warm surface stratum inshore; and it has been taken off Port-au-Port on the west coast of Newfoundland.21 Importance. — Sand flounders are so small and so thin bodied, and so few of them are caught in the Gulf of Maine that they are of no commercial importance there, nor likely to be. However, a market developed for them during the war years in New York, where a much larger supply was near at hand, culminating in landings of about 340,000 pounds in 1944, and about 360,000 pounds in 1945. But as Moore has pointed out,22 the demand fell off during 1945, as the war drew to its close. And now the sand flounder is a neglected fish again. Gulf Stream flounder Citharichthys arctifrons Goode 1880 Jordan and Evermann, 1896-1900, p. 2683. Description. — This little flatfish is left-handed (eyes on the left-hand side and viscera at the left- hand edge as the fish lies), with a wide mouth gaping back as far as the forward edge of the eye; with a nearly straight lateral line; and with both of its pectoral fins well developed, though the one on the eyed side is considerably larger than its mate on the blind side. Its left-hand ventral fin stands on the midline of the body, but the right- hand ventral fin is a short distance above it on the blind side, and while the two ventral fins are alike in females, the one on the blind side is much the >• Smith, North Carolina Geol. and Econ. Survey, vol. 2, 1897. p. 392. » Report 1, No. 4, Newfoundland Fishery Res. Comm., 1632, p. 110. » Cornish, Contrib. Canadian Biol., (1902-1905) H07, p. 90. » Contrib. Canadian Biol., (1918-1920) 1921, p. 113. » Trans. Roy. Soc. Canada, Ser. 3, vol. 12, Sect. 4, 1918, p. 63. " Rept., Newfoundland Fish. Res. Comm., vol. 2, No. 1, 1933, p. 127. >s See Moore (Bull. Bingham Oceanogr. Coll., vol. 11, art. 3, 1947, p. 71) for detailed tabulation of the New York landings, 1943-1945, from the Dally Market News Service, Division of Fishery Industries, U. S. Fish and Wild- life Service. The sand flounder is not included in the general fisheries sta- tistics published yearly by the Fish and Wlldlifo Service. FISHES OF THE GULF OF MAINE 295 YY-' mm •■■ ■.?!;,:■ 'SSr-ia* Figure 154. — Gulf Stream flounder (Citharichthys arctifrons), off Rhode Island. From Goode and Bean. Drawing by H. L. Todd. longer of the pair in males. The body is ovate in outline and very thin. The long (ventral and dorsal) fins are of moderate breadth, with the dorsal fin (78 to S3 rays) originating over the forward margin of the eye, the anal (loft-hand edge) fin (61 to 67 rays) originating a little in advance of the pectorals; and the caudal fin rounded. The scales are so large that there are only about 40 rows of them along the lateral line. Fish living at different depths vary so widely in the number of fin rays that future studies may reveal the existence of distinct races, if not of species.23 Color. — Light brown above, with the scales usually more or less outlined with darker brown; brownish white below. Size. — Maximum length about 7 inches. Remarks. — This little flatfish parallels the sum- mer, four-spotted, and sand flounders (the latter « A second species of this genus (C unicornis Goode 1880) may be expected on the outer slope of Georges Bank in depths of 100 fathoms and more, since it has been taken ofl Marthas Vineyard In 115 to 150 fathoms. The male is separable from C. arctifrons by the fact that there are several short spines on the eyed side of the head above the upper lip (the head of arctifrons is spine- less although old fish may have a bony protuberance on tho snout). Further points of distinction are that unicornis has fewer fin rays (only about 74 to 77 dorsal rays and 60 anal rays) and that its body is broader (actually higher) Parr (Bulletin of the Bingham Oceanographlc Collection, vol. 4, art. 1, 1931/ has published a revision of the genus Cuharkhtfiys of the western Atlantic. We have towed the pelagic larvae of still a third small deep-water flounder {MonoUne sessilkauda Goode 1880) off the seaward slope of Georges Bank (Bull. Mus. Comp. Zool., vol. 41, No. 8, 1917, p. 277), while the adults have been trawled in depths of 100 fathoms and more off Marthas Vineyard and thence westward and southward along the continental slope. It Is left- handed like the summer, four-spotted, and sand flounders, with arched lateral line, but it has no pectoral fin on the blind side. For a detailed description of it. see Goode and Bean, Smithsonian Contrib. Knowl.,'(vol. 30, 1895, p. 452). its closest Gulf of Maine ally) in its lef t-handedness. But it is distinguishable from all of these by its nearly straight lateral line; by the great disparity in size between its two pectoral fins; and by its very large scales. Its narrow shape and the fact that none of its dorsal fin rays are branched are further points of distinction between it and the sand flounder; also it is much smaller at maturity than any of the flatfishes that are common in the inner parts of the Gulf of Maine. Habits.- — Little is known of its habits. It is found chiefly in water deeper than 40 fathoms but it has been trawled as shoal as 12 fathoms. Apparently it spawns from spring through sum- mer, for we have found females with well-developed ovaries in February, while Goode had ripe ones in September. It is not large enough to be of commercial value, but we can witness that it is excellent on the table. General range.- — Eastern coast of America, along the outer part of the continental shelf from the southwestern part of Georges Bank to the offing of Charleston, S. C., where the Blake took it many 3rears ago,24 usually at depths of 40 to 200 fathoms, but occasionally as shoal as 12 to 18 fathoms. A fish occupying this geographic province is misnamed when it is called "Gulf Stream," but this is the only English name by which it has been known. Occurrence in the Gulf of Maine.- — This little flatfish has never been reported from the inner » Goode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, pp. 443- 444) give a long list of localities where it has been trawled, along the conti- nental shelf from the offing of Nantucket to the offing of Charleston, S. O. 296 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE parts of the Gulf, nor is it to be expected there, to judge from its general distribution. But the Albatross I took one in a tow net over the south- western part of Georges Bank at about the 82 fathom (150 meters) contour line.25 And subse- quent captures of scattered specimens in that general neighborhood in 1931 by the Albatross II; at 8 stations (30 specimens) between the offing of Nantucket and longitude about 67° 10' W., in 41 to 150 fathoms, by the Albatross III in May 1950; and on those same general grounds in 39 to 65 fathoms by the Eugene H in late June 1951, show that its regular range extends eastward far enough to include not only the slope of Nantucket Shoals, but the southwestern sector of Georges Bank arc as well, at the appropriate depth. And it must be considerably more plentiful on the outer part of the shelf off southern New England, for the Albatross III has trawled a considerable number of them there, including one catch of 100 off Montauk Point, in February 1950, and another of as many more off Rhode Island on May 13, 1950, at 41 to 50 fathoms. » Station 20045, lat. 40° 18' N., long. 68° 09' W., February 22, 1920. Hogchoker Achirus fasciatus Lacepede, 1803 American sole Jordan and Evermann, 1896-1900, p. 2700. Description. — This fish is the closest relative, in northeastern American waters, of the famous sole of Europe. It is right-handed and small-mouthed, and it can be told at a glance from all other Gulf of Maine flatfishes by the fact that it has no pectoral fin on either side. Its mouth gapes along the general fore-and-aft line as the fish lies, with the upper jaw projecting beyond the lower, whereas the gape is oblique in all other local flatfishes, and it is their lower jaw that projects. Furthermore, the rounded outline of the head of the hogchoker, and the lack of a definite snout, gives it an aspect very different from that of any other Gulf of Maine flatfish. Equally diagnostic among right-handed species is that its right-hand ventral fin is continuous with the anal fin; its long fins are highest toward their rear ends; its dorsal (left-hand) fin originates at the very tip of the nose (thus, further forward than in our commoner flounders); and its small eyes are set flat instead of in prominent orbits. Other characters worth mentioning are that the Figure 155. — Hogchoker (Achirus fasciatus), Woods Hole. After Jordan and Evermann. Original drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 297 gape of its mouth is shorter and much more crooked on the blind side than it is on the eyed side (an asymmetry that has been emphasized in most of the descriptions of this species); that it is evenly oval in outline without a definite caudal peduncle; and that there are 50 to 56 dorsal-fin rays and 36 to 42 anal-fin rays, but no pre-anal spine. The scales are very rough on both sides, those of the upper part of the head and chin on the eyed side and on the wbole head on the blind side are larger than the body scales, and its skin is slimy with mucus. Color. — Dusky or slaty olive to dark brown on the eyed side, barred transversely with a varying number (usually 7 or 8) of indistinct darker stripes, with a dark longitudinal stripe along the lateral line, and sometimes with pale mottling. The dorsal, caudal and anal fins are of the general body tint, variously dark clouded. The blind side is dirty white, usually marked with dark round spots which vary in size and number from fish to fish. But some specimens lack these spots. Size. — Eight inches is about the maximum length. Habits. — The hogchoker is confined to the im- mediate vicinity of the coast, is most common in bays and estuaries where the water is more or less brackish, and sometimes runs up into fresh water. It is a late spring and summer spawner. At Woods Hole fish apparently ripe have been taken in May, while in Chesapeake Bay ripe or nearly ripe fish have been collected in June, July, and August. One female, 6% inches long, con- tained about 54,000 eggs about 0.3 mm. in diameter, whether buoyant or not is not known. It reaches a length of 2 to 3 inches at one year of age, and matures when about 4% inches long. It feeds chiefly on annelid worms and on small crustaceans.26 Fragments of algae also have been found in hogchoker stomachs, but these probably were swallowed with its animal prey. General Range. — Off the Atlantic and Gulf coasts of North America, from Massachusetts Bay to the Atlantic coast of Panama. The hogchoker is abundant in Chesapeake Bay and to the south- ward, and moderately common as far north as southern New England, but it is rare north of Cape Cod. Occurrence in the Gulf of Maine. — This little flatfish has been reported from Provincetown (where Captain Atwood spoke of it as plentiful); from Boston Harbor, whence the Museum of Comparative Zoology has several, all caught long ago; from the mouth of the Charles River (two specimens reported in 1847); and from Nahant (one taken in 1840). But it is more than three- quarters of a century since it has been brought to scientific attention anywhere to the north of Cape Cod; if it is caught there from time to time, as it doubtless is, it has not been recognized. It is not known north or east of Cape Ann, nor on the offshore banks. Importance. — The hogchoker is said to be deli- cious eating. But it is so small that it is of no commercial value even in Chesapeake Bay where it is plentiful. Incidentally, the rumored origin of the name "hogchoker" is that hogs that "feed on fish discarded on the beaches, have great diffi- culty in swallowing this sole, because of the extremely hard, rough scales." " THE JOHN DORIES. FAMILY ZEIDAE American John Dory 28 Zenopsis ocellata (Storer) 1858 Jordan and Evermann, 1896-1900, p. 1660. Description. — The John Dory is easily dis- tinguishable from all other Gulf of Maine fishes of similar body form by its long dorsal fin spines, bony armor, tiny tail fin, and the curious profile of » Hildebrand and Scbrocder, Bull. U. S. Bur. Fish., vol. 43, pt. 1, 1928, p. 176. » Hildebrand and Schroeder, Bull. D. S. Bur. Fish., vol. 43, Pt. 1, 192S, p. 177. 89 Separable from the common John Dory of Europe by having three anal spines instead of four, and by a greater development of the bony plates. its head. Like the butterfish it is very deep (only about one and three-fourths to twice as long as it is deep) and very much flattened sidewise. Its body is rounded in side view, with the dorsal profile of its head noticeably concave, its large mouth is set very obliquely, and its caudal peduncle is very slender. Its dorsal fin is in two parts, spiny and soft ra3red ; the former, originating over the upper corner of the gill covers, has 9 to 10 spines; the first, second, and third spines very long, the others graduated. And all the spines are filamentous toward the tip. The soft dorsal fin 298 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 156. — American John Dory (Zenopsis ocellata), Provincetown. From Goode. Drawing by H. L. Todd. (25 to 27 rays) is somewhat longer than the spiny dorsal fin, but less than half as high, and its anterior rays are only about half as high as the posterior ones. The two dorsal fins, together, occupy the entire length of the back of the fish from nape of neck to caudal peduncle. The anal fin (24 to 26 rays preceded by 3 short stout spines) corresponds to the soft dorsal in location, height, and outline. The very small caudal fin is brush shaped, the ventral fins are very long, with the rays free at their tips, and they are situated in front of the pectorals. The pectorals are short and rounded. The skin is naked except for a series of bony bucklers, each with a hooked thorn or double thorn; two or three of them along the base of the spiny dorsal fin and four along the base of the soft dorsal; two in front of the ventral fins; one in the midline behind the ventrals, followed by six pairs along the belly to the anal fin; and five along the base of the anal fin. Color. — Silvery all over. Specimens that we have seen up to about 10 inches long are marked on either side with about 12-24 vaguely outlined dark spots, irregularly arranged, and fish up to about 15 inches long retain some of the spots.29 But it seems that the spots tend to fade out with growth, for larger specimens that we have at hand, 16-20 inches long, have only one vague blotch on each side, a short distance behind the gill opening. Size. — The largest four specimens yet seen measured 18% and 18% inches; 30 19 inches, weigh- " This agrees with the original account of the species (Storer, Proc. Boston Soc. Nat. Hist., vol. 6, 1858, p. 386) and with a photograph of one about 3H inches long, from Campobello, New Brunswick, sent us by Dr. A. H. Leim. » Taken off Long Island, N. Y., by the Albatross III, lat. ZWP N., long. 72°08' W., May 12, 1960. FISHES OF THE GULF OF MAINE 299 ing 3 pounds;31 and 20 inches, weighing A){ pounds and 24 inches, weighing 7 pounds.32 Remarks. — The presence of plates along the base of its first (spiny) dorsal fin, as well as along the bases of its second (soft) dorsal and anal fins, and of only three anal spines marks our fish off from its close counterpart, the European John Dory {Zeus faber), which has four stout anal spines and lacks plates along the first dorsal fin. Other structural differences are that the plates are much larger in our species than in the European, but the thorns smaller and less conspicuous ; 33 that the base of each of the dorsal fin spines (except for the first and last one or two) is armed in the European species with a stout thorn (not in the American); and that the upper profile of the head is much the more deeply concave in the American species. Habits. — All that is known of the habits of our John Dory is that we found two butterfish 6 to 7 inches long and one squid in the stomach of a large one (of about 18)2 in.) trawled by the Albatross III about 74 miles off Long Island, N. Y., May 12, 1950, at 72 fathoms; and that the ovaries were well developed with orange colored eggs 1.2 to 1.4 mm. in diameter, in a 20-inch female that we saw trawled between January 27 and February 2 on the outer part of the shelf off Marthas Vineyard.34 General range. — Outer part of the continental shelf from the latitude of Chesapeake Bay to the vicinity of Sable Island, Nova Scotia, and perhaps to the Laurentian Channel that separa tes the Nova Scotian Banks from the Newfoundland Banks. It reaches the inner parts of the Gulf of Maine now and then as a stray. Occurrence in the Gulf of Maine. — Only four specimens are known to have been taken in the inner parts of the Gulf of Maine. One (the speci- men from which the species was described) was found at Provincetown, at the tip of Cape Cod many years ago; one found in a herring weir at Campobello Island, New Brunswick, at the mouth of the Bay of Fundy in 1942; 3S one trawled 25 miles off Cape Ann in 75 fathoms, January 1948.38 One also was trawled on the northeastern edge of Georges Bank in the summer of 1941,37 and one taken in Cape Cod Bay, July 7, 1952, by the dragger Santina. It is to be expected anywhere along the seaward slope of the offshore rim of the Gulf, for the dragger Eugene H took them in nearly every trawl haul on the southwest slope of Georges Bank, near Veatch Canyon, at about the 75-fathom contour line, in late March 1951, some hauls bringing in several hundred (estimated) specimens. Other speci- mens M have been trawled recently on the outer part of the continental shelf southeast of Cape Henry, Va., from between 28 and 50 fathoms;39 off Long Island, New York, in 72 fathoms and from between 145 and 200 fathoms; off Marthas Vineyard in 55 to 68 fathoms; off Nantucket in 60 to 75 fathoms; on Emerald Bank off Halifax, Nova Scotia, in 70 fathoms; and west of Sable Island, Nova Scotia, at 02 fathoms. GRAMMICOLEPID FISHES. FAMILY GRAMMICOLEPIDAE 40 Grammicolepid Xenolepidichthys americanus Nichols and Firth 1939 Nichols and Firth, Proc, Biol. Soc, Washington, vol. 52, 1939, pp. 85-88. Description. — This curious little oceanic fish resembles its near relative the John Dory (p. 297) in the arrangement of its fins, and in general shape, with body so strongly flattened sidewise as to be as thin as a pancake, and with a slender caudal peduncle. But it has a much smaller mouth " Caught on the northeast edge of Georges Bank In the summer of 1941 and reported In the Boston Traveler for September 9 of that year. " Taken 85 miles off Marthas Vineyard by the dragger Eugene II, May IS, 1950. :' Double and sometimes triple In the European 7. faber. » Trawled by the dragger Eugene H from between 55-68 fathoms. than the John Dory, its scales are linear in shape with their long axis dorso-ventral, so that the sides of the trunk are cross marked with a large number of narrow lines, closely crowded together, and the series of bony plates that arm the dorsal and ventral edge of the body of the John Dory are replaced in the Grammicolepids by a double series of short thorns that embrace the bases of the dorsal and ventral fins. Each side of the trunk of the only species known from our waters is » Reported to us, with a photograph, by Dr. A. n. Leim. « This specimen, trawled by the Agatha and Patricia, is in the Museum of Comparative Zoology. " Reported in the Boston Traveler for September 9, 1941. » Specimens seeu by us or reliably reported. » Reported by Firth, Copeia, 1931, p. 162. " For a recent account of this family see Myers, Proc, V. 9. National Museum, vol. 84, 1937, pp. 145-156. 300 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE /; ' . Figure 157. — Grammicolepid (Xenolepidichthys americanus), Georges Bank. Drawing by H. B. Bigelow. after the original illustration by Firth and Nichols. Tail fin armed with about 11 or 12 conspicuous, horizon- tally flattened spines, pointing rearward. Size. — The only specimen yet seen is about 4 inches (100 mm.) long, to the base of its tail fin. We need only add, further, of our species, that the forward division of the dorsal fin consists of 5 spines, the forward edge of the first saw-edged, and all of them filamentous toward the tip; that the second dorsal fin, of 33 soft rays (separated from the first by a considerable gap), is about as high as two-thirds the diameter of the eye; is of about equal height from end to end, and reaches back to the caudal peduncle; that the tail fin is deeply forked, its tips pointed, and its upper lobe longer than the lower (unless this is the result of mutila- tion) ; that the soft-rayed anal fin, corresponding to the second (soft) dorsal fin, is preceded, after a considerable gap, first by a short, smooth spine, then by another very long spine, saw-toothed most of its length, but filamentous toward its tip; and that the ventral fins, of 1 stout, saw-edged spine followed by 6 soft rays, stand a little in advance of the brush-shaped pectorals. Color. — After preservation in alcohol, the color is "pale, with a series of dark marks on the midline of the back, and about 10 narrow dark bands extending downward from these to the level of the top of the eye . . . the flattened spines, scattered over the body are blackish. Base of anal with a series of dusky blotches, and posterior part of caudal dusky".41 Range and occurrence in the Chilf oj Maine. — So far known only from Georges Bank, where the only specimen yet seen was picked up, in a bucket, from the Sword Fisherman America. A closely related species, X. dalgleishi Gilchrist 1922, is known from the Caribbean, South Africa, and the Philippines. along both its front margin and its rear margin for 41 Quoted from the original account by Nichols and Firth. FISHES OF THE GULF OF MAINE 301 SNIPE FISHES. FAMILY MACRORHAMPHOSIDAE Snipe fish Macrorhamphosus scolopax (Linnaeus) 1758 Jordan and Evermann, 1896-1900, p. 759. Description. — The snipe fish is given so unusual an appearance by a long tubular snout with small toothless mouth at the tip, combined with a very long, stout dorsal fin spine that is saw-toothed along the rear edge that it could hardly be mis- taken for any other Gulf of Maine fish. Its body is about two-fifths as deep as long, measured from front of eye to base of caudal fin, so flattened sidewise that it is only about one- third to three-eighths as thick through as it is deep; the snout, measured from the front of the eye is about 1% times as long as the depth of the body. The eye is noticeably large. The two dorsal fins stand far behind the mid-length of the trunk. The first dorsal is of 5 to 7 spines and very short, the second, far the longer, is pointed, with about 11 to 13 soft rays; and the two dorsal fins are separated by an interspace nearly as long as the base of the first dorsal. The anal, with 19 to 20 rays, is much longer than the second dorsal, but lower; the caudal is square- tipped, of moderate size. The very small ventrals are located considerably behind the pectorals. The snout, head, and sides are clothed with small rough scales. And the body is further stiffened with bony plates, of which there are 2 longitudinal rows of 4 each, high up on each side behind the gill opening; also 3 longitudinal series of 6 each along the lower breast and belly in front of the ventral fins, followed by 3 pairs behind the latter and finally by a single plate close in front of the anal fin, these last forming a sharp keel. Color. — Pinkish or reddish on sides above, fading to silvery white below. Described as sometimes golden above. Size. — Maximum reported size about 6% inches (16 cm.);42 the few we have seen were about 4 inches long. General range. — Widespread in warm seas. Eastern Atlantic, from the coast and Banks of Morocco, where it is sometimes taken in numbers, and the Mediterranean, northward to southern England (Cornwall, Devonshire) ; so far known in the western Atlantic only from the offing of Nan- tucket and from Massachusetts Bay. Occurrence in the Gulf of Maine.— Oddly enough, the few records of this eastern Atlantic fish in our side of the Atlantic have all been within the limits of the Gulf of Maine; namely, one reported from Massachusetts Bay; *■ a second trawled south of Nantucket, at the 130-fathom contour line,44 both many years ago; and eight specimens trawled in that same general vicinity (lat. 39°59' N., long. 69°47' W.) at 80 fathoms, by the Albatross III on May 14, 1950. Evidently it reaches the inner parts of the Gulf only as a stray, and at long intervals although it is taken from time to time by otter trawlers along the southwestern edge of Georges Bank in 75 to 85 fathoms. « One of this sUe is pictured by Murray and n)ort (Depths of the Ocean, 1912, p. 397, fig. 268). « Qoode and Bean (Smithsonian Contrib. Knowl., vol. 30, 1895, p. 483), without further details. « Oricinal of Ooode and Dean's illustration (Smithsonian Contrib. Knowl., vol. 31, 1895, pi. 127, flg. 396). Figure 158.— Snipefish (Macrorhamphosus scolopar), off Nantucket. 302 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE THE SILVERSIDES. FAMILY ATHERINIDAE These are small fishes, smelt-like in appearance, except that they have a spiny dorsal fin as well as a soft dorsal fin; do not have the adipose fin, and have much smaller mouths than the smelt. Two species are known from the Gulf of Maine. KEY TO GULF OF MAINE SILVERSIDES 1 . About 24 rays in the anal fin Common silverside, p. 302. Only about 15 or 16 rays in the anal fin Waxen silverside, p. 304. Silverside Menidia menidia (Linnaeus) 1766 Green smelt; Sand smelt; White-Bait; Capelin; Sperling; Shiner Jordan and Evermann, 1896-1900, pp. 800, 2840. Description.- — This silvery little fish is often confused with the young smelt, but it does not require very close examination to tell them apart for the adipose fin characteristic of the smelt is lacking in the silverside, while the latter has a spiny dorsal fin as well as a soft dorsal fin instead of the one soft dorsal only, as the smelt does; this last character distinguishes it equally from young herrings; its anal fin too is much longer than that of the smelt. It is a slender fish, about one-sixth as deep as long, not counting caudal fin ; thin-bodied but with rounded (not sharp-edged) belly; with short head; large eye; and small mouth, gaping hardly as far back as the front of the eye, and set very obliquely. Both head and body are clothed with large scales. The first dorsal fin (3 to 7 spines) is smaller than the second and originates about midway between the tip of the snout and the base of the caudal fin; the second dorsal has 7 to 10 soft rays and origi- nates over the middle of the anal. The anal (of 23 to 26 rays, the first stiff and the others soft) is falcate in outline. The caudal peduncle is slender, the tail moderately forked.45 Color. — Translucent bottle green above, with top of head, nose, and chin dusky. The upper parts of the sides are thickly speckled with dark brown, and there is a silver band outlined above by a narrow black streak, running along each side from close behind the pectoral fin to the base of the caudal fin. The belly is white. Size. — The silverside grows to a length of about h){ inches, adults usually running 4 to 4}i inches long. Habits. — Silversides tend to congregate in schools usually made up of even-sized individuals. They frequent sandy or gravelly shores chiefly, and there is no reason to suppose that they ever ven- ture out to sea. At high tide they are often seen among the sedge grass (Spartina), where it grows sparsely between tide marks, particularly about the inner bays and in river mouths where they follow the tide up and down the beach within a few yards of the water's edge. They also run up into brackish water; near St. Andrews, in fact, they are chiefly found in brackish situations though more generally distributed on the New Brunswick shore further up the Bay of Fundy and on the Nova Scotian side as a whole. They do not or- dinarily descend deeper than a fathom or so in summer. But some of them, at any rate, sink *8 The common silverside is represented on the coasts of the eastern United States by two races, a southern and a northern, not, however, very distinct and connected by such various intergradations that they hardly deserve two names, subspecies menidia for the southern and subspecies notata for the northern. The southern form has fewer scales than the northern, only 4 instead of 5 spines in the first dorsal, and is rather a stouter-bodied fish. Kendall has given an account of the genus (Report, U. S. Comm. Fish (1901) 1902, p. 241). For a recent discussion see Bayliffe (Publ. 90, Chesapeake Biol. Lab., Maryland Dept. Nat. Res., 1950. p. 5). -•F??VP?'-r.-'*7? 7i'^'- Figure 159. — Silverside (Menidia menidia), Connecticut. From Goode. Drawing from H. L. Todd. FISHES OF THE GULF OF MAINE 303 deeper in winter, probably to avoid low tempera- ture, for Hildebrand and Schroeder 46 found them at depths of 5 to 27 fathoms at that season in Chesapeake Bay. But this is not their universal habit, for they are taken in winter through the ice, as well as in summer, in the southern side of the Gulf of St. Lawrence. And they are resident throughout the year wherever found, generally speaking. Such, at least, is the case along south- ern New England. They are omnivorous, feeding chiefly on cope- pods, mysids, shrimps, small decapod shrimps, amphipods, Cladocera, fish eggs (including their own), young squid, annelid worms, and molluscan larvae. Insects, too, that fall into the water have been found in their stomachs, as have algae and diatoms mixed with sand and mud. On tbe other hand, they are harried by every predaceous fish that comes close inshore, especially by bluefish and by striped bass. It was interesting in this connec- tion to find that a dolphin (rare in the Gulf) , taken at Sandwich, in Cape Cod Bay, in July 1951 (p. 361) was packed full of silversides. They spawn in May, June, and early July " on the southern New England coast. Spawning may begin a little later in the Gulf of Maine, corre- sponding to more tardy vernal warming, while Leim writes 48 that they do so in June at Prince Edward Island. The gap in the presence of silversides in abun- dance that seems to exist along the cool-water stretch from the western side of the Gulf of Maine to the southern side of the Gulf of St. Lawrence (p. 304) suggests that they need summer tempera- tures as high as 68° or so for successful reproduc- tion. But young fry and adults alike are indiffer- ent to temperatures down to a degree or two above the freezing point of salt water, witness their presence in winter in Cape Cod Bay (p. 304) and below the ice in the bays on the northern side of Prince Edward Island (p. 304). When the sdversides are spawning they gather in schools to deposit their eggs on sandy bottom, often among the sedge grass at high tide, or above low-water mark. Capt. John B. Smith has described them spawning in the sedge at the head '• Bull. U. S. Bur. Fish., vol. 43, Pt. 1, 1928, p. 189. « Kuntz and Hadcllffe (Bull. U. S. Bur. Fish., vol. 35, 1918, p. 127) de- scribe Its development, and Hildebrand (Bull. U. S. Bur. Fish., vol. 38'. 1921- 22] 1923 that of the southern race. For a recent account of Its life history see Bayltffe, Pub. 90, Chesapeake Biol. Lab., Maryland Dept. Nat. Res. ,1950. •' Proc. Nova Scotlan Inst. Scl., vol. 20, Pt. 2, 1940, p. 38. of Buzzards Bay, June 13, 1872, rolling from side to side, some jumping clear of the water, and in such multitudes that the water was "whitened with the milt, and the grass was so full of eggs that they could be taken up by the handfull," while small fishes of various kinds were "helping themselves to the dainty repast." 49 Figure 160.— Egg. Figure 161. — Larva, 8 mm. Figure 162.— Fry, 13 mm. Silverside (Menidia menidia) . After Kuntz and Radcliffe. The eggs, 1.1 to 1.2 mm. in diameter and each bearing a bunch of sticky filaments, sink and stick fast in ropy clusters or sheets. Incubation occu- pied 8 or 9 days in the laboratory at Woods Hole. The yolk is absorbed before hatching, at which time the larvae are about 3.85 to 5 mm. long, and the dorsal, anal, and caudal fins are formed in larvae of 12 to 15 mm. in length. The young grew to a length of 9.3 to 11.7 mm. during the first 20 days in the aquaria. Probably they grow more rapidly at liberty, for all sizes from fry of an inch or less to adults are to be found throughout the summer. Probably the silverside attains maturity at 1 year of age. General range. — The northern variety of this silverside is common locally from the southern side of the Gulf of St. Lawrence and the outer «Goode, Fish. Ind. U. S. Sect. 1, 1884. p. 457. 304 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Nova Scotian coast to Massachusetts Bay, and very abundant thence southward to Chesapeake Bay, south of which it gives place to the southern form or intergrades with it ; the southern form has been detected as far north as Woods Hole, but never east of Cape Cod. Occurrence in the Gulf of Maine. — -The silverside is to be found all around the shores of the Gulf from Nova Scotia to Cape Cod, always, however, closely confined to the coastline. They are ex- ceedingly plentiful around the sandy shores of Cape Cod Bay. And while we have seen them from Chelsea Beach in Boston Harbor, from Beverly and from Gloucester, many summers spent on the coast leave us with the impression that the silverside is neither as omnipresent nor as abundant from Massachusetts Bay northward, although large schools of them are often to be seen here and there along the sandy beaches on the Maine coast. Bushels, in fact, have been caught in a single haul of the seine in Casco Bay and very likely could be elsewhere. Silversides are seldom seen along the stretches of rocky coast exposed to the open sea, which make up a large part of the northern shore line of the Gulf of Maine. In Passamaquoddy Bay Huntsman tells us60 "they are largely restricted to brackish water and hence not very common," but they must be rather generally present in suitable situations around the shore line of the Bay of Fundy, being reported from St. John and Kennebecasis Bay, from Annapolis basin and from St. Mary Bay. Nothing is known as to their status along the Nova Scotian coast of the open Gulf of Maine, or even whether there are any silversides there at all. Halifax is the most northerly locality where they are recorded on the outer coast of Nova Scotia.51 But Leim 52 reports them so plentiful in the shallows of the southern side of the Gulf of St. Lawrence that "hardly a seine haul has been made without catching several, and as many as 3,500 have been taken at once" in Malpeque Bay on the north shore of Prince Edward Island, where they are taken in winter through the ice, as well as in summer. Enough of them, in fact, are sometimes caught there to be worth canning.63 Their abundance there con- trasted with their evident scarcity along outer Nova Scotia suggests the presence of an isolated population (or populations) in suitable situations in the southern side of the Gulf of St. Lawrence, able to maintain itself because summer tempera- tures in the shallows there are high enough for its successful propagation. Importance. — The chief function of the silverside in the economy of the sea is to feed predaceous fishes such as bluefish, mackerel, and striped bass. The silverside is of no commercial value north of Cape Cod, being too small and too soft to answer the never satisfied demand for bait for the offshore fisheries, but they are very generally used to bait eelpots on the Rhode Island coast, and they are excellent on the table, fried, as whitebait. Waxen silverside Menidia beryllina (Cope) 1866 Jordan and Evermann 1896-1900, p. 797 (Menidia gracilis Gunther). Description. — This species resembles the com- mon silverside so closely in general appearance that it would be apt to be overlooked among the schools of the latter were it not paler in color, » Contrib. Canadian Biol. (192!) 1922, p. 61. •i Cornish (Contr. Canadian Biol. [1902-1905], 1907, No. 9) does not include it in his list of the fishes of Canso. '= Proc. Nova Scotian Inst. Sci., vol. 20, Pt. 2, 1940, p. 38. « Needier, Rept. Fish. Res. Board Canada (1941) 1942, p. 11. Figure 163. — Waxen silverside (Menidia beryllina). From Kendall FISHES OF THE GULF OF MAINE 305 and stouter bodied as a rule. A more dependable difference, which will always serve to separate the two (for which neither color nor form can be relied upon) is that the anal fin is much shorter (only 15 or 16 rays) in the waxen silverside than in the common silverside. Color. — Pale greenish on the back, silvery below; the sides with a well-defined silvery band bounded above by a dark line; scales on the back with numerous brown dots; fins without markings. Size. — Smaller than its relative menidia, the maximum length being about 3 inches. General range. — Cape Cod to South Carolina. Occurrence in the Gulf of Maine. — Specimens reported by Kendall 54 (1902) from Truro, and from Sandwich in Cape Cod Bay, with one taken in Cohasset, on the southern shore of Massachu- setts Bay in the autumn of 1939, 55 are the only records for this fish within the Gulf of Maine, where it appears only a stray from warmer waters to the west and south. At Woods Hole, where it is abundant, its habits are the same as those of the common silverside, though it spawns some- what later (in June and July). THE MULLETS. FAMILY MUGILIDAE Mullets have two separate dorsal fins, the first spiny and the second soft-rayed. Their ventral fins are on the abdomen behind the point of in- sertion of the pectorals; their tails are forked and they have large scales. Their closest affinity among Gulf of Maine fishes is with the silversides, which they resemble somewhat in the relative size and locations of the fins; but they differ from the silversides in their short, broad heads, small eyes, and relatively deeper and thicker bodies, while they have only 24 vertebrae instead of 35 or more. Furthermore, they are vegetable amd mud eaters instead of carnivorous, their stomachs are thick walled and gizzard-like, the intestines long, corresponding to their food. The lining of the belly of the mullet is black while that of the silverside is pale. There are many species of mullets. Most of them, however, are tropical, and only one has ever « Occ. Pap. Boston Soc. Nat. Hist., vol. 7. No. S. 1908, p. 66, as Menidia beryttina subspecies area Kendall (Kept. U. S. Comm. Fish. (1901) 1902, p. 261.) " This specimen is in the Museum of Comparative Zoology. been known to strav within the confines of the Gulf of Maine.58 Mullet Mugil cephalm Linnaeus 1758 Common mullet; Striped mullet; Jumping mullet Jordan and Evermann, 1896-1900, p. 811. Description. — The common mullet, the only one of its numerous tribe (there are more than 100 species of mullets) that has been known to stray north of Cape Cod, has a spiny first dorsal and soft second dorsal fin, the two well separated as in the silverside, and its ventrals are located on the abdomen. It is a much larger fish than the silver- side, however, and even very young mullets of the »• The so-called red mullet or goat fish (Mullut auratiu) of more southern waters, which is not a true mullet but belongs to a different family (Mullidae), is taken from time to time near Woods Hole, and it has been reported from HaUfu Harbor, Nova Scotia (by Leim, Proc. Nova Scotian Inst. Scl., vol. 17, No. 4. 1930, p. XLVI), hence it may be eipected as a stray in our Gulf, though it has not actually been found there as yet. There is no danger of mistaking it for a mullet, for it is bright crimson, with a fleshy barbel on its chin, and with its ventral flns far forward, below its pectorals. Figure 164. — Mullet (Mugil cephalus), Woods Hole. From Goode. Drawing by H. L. Todd. 306 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE size of the latter (4 to 5 inches long) are easily separable from silversides by the fact that their anal fin is only about half as loog in relation to the length of the body, while the second dorsal orig- inates over the origin of the anal instead of well behind it. Furthermore, the head of the mullet is shorter; its nose blunter; its profile quite different (compare fig. 164 with fig. 159); its eye smaller; its body stouter (about one-fourth as deep as long) ; and it lacks the silvery side stripes so char- acteristic of the common silverside. There are four spines in its first dorsal fin, 1 spine and 8 soft rays in the second dorsal, 3 spines and (usually) 8 rays in the anal. Young fish, 2 inches long or less, have only 2 spines in the anal, the first soft ray later developing into a spine.67 The first dorsal stands over the tips of the pectorals or close behind them; and the tail is forked moderately deep. The soft dorsal fin and anal fin are almost naked (they are scaled in most of the other American mullets), but the body and head are clothed with large rounded scales. Color. — Adults are bluish gv&y or greenish above, silvery on the lower part of the sides and below; the scales on the sides have dark centers which form longitudinal lines; the fins are sometimes partly dusky. Young fry are bright silvery. Size. — The common mullet grows to a length of 2% feet in warmer waters, but small specimens alone have been found along our northern coasts. General range. — Both sides of the temperate Atlantic; from Brazil to Cape Cod on the Ameri- can coast, and as a stray to outer Nova Scotia; also along the west coast of America from Mon- terey (Calif.) to Chile, and in other parts of the Pacific. Occurrence in the Gulf of Maine. — Mullets are common as far north as New York, less so to Woods Hole, but so rarely do they stray past Cape Cod that there are only a half dozen records of them in the Gulf of Maine, viz, at Provincetown, at Essex M in northern Massachusetts, at Freeport, Harraseeket River, Clapboard Island, and Casco Bay in Maine, each based on an odd fish. And one has also been taken in Bedford Basin near Halifax, Nova Scotia.69 Mullet are more likely to visit the cool waters of the Gulf in late summer or early autumn than at any other season. They have been known to winter as far north as New York, in the mud, but it is not likely that the few strays that round Cape Cod survive the cold sea- son, nor is there any reason to suppose they ever breed in the Gulf, for immature fish alone are found at Woods Hole. THE BARRACUDAS. FAMILY SPHYRAENIDAE The slim bodied barracudas, with their long, pointed heads, somewhat resemble the pikes in general appearance. But they are distinguishable from the latter at a glance by having two dorsal fins. The lower jaw projects beyond the upper, and both jaws are studded with large pointed teeth of unequal sizes. The gill covers are scaly, and there is a well-developed lateral line. The first dorsal is spiny, the second soft-rayed. The anal is roughly opposite the second dorsal, the 17 See Jacot (Trans. Amer. Microscopical Soc, vol. 39, 1920, pp. 204-214) or a study of the growth of the mullet. ventrals opposite the first dorsal, the pectorals short, the caudal forked. Northern barracuda Sphyraena borealis DeKay 1842 Jordan and Evermann, 1896-1900, p. 825. Description. — The combination of slender shape with long head, projecting lower jaw, a first dorsal ■ There is (or was) a specimen so labeled in the collection of the Boston Society of Natural History. M Reported by Vladykov, Proc. Nova Scotian Inst. Sci., vol. 19, 1935, p. 6. Figure 165. — Northern barracuda (Sphyraena borealis), Woods Hole. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 307 situated opposite the ventrals, a second dorsal opposite the anal, and a forked tail, separates the barracuda from any other Gulf of Maine fish. Color. — The adult is olivaceous above, silvery below. The young have dusky blotches along the back and along the lateral line. Size. — This is the smallest of the barracudas, few growing longer than one foot. General range. — -Atlantic and Gulf coasts of America from Cape Cod to Panama. Occurrence in the Gulf of Maine. — A specimen, about 2 inches long, found alive in the surf at Nauset Beach, Cape Cod, September 26, 1930, by the late Dr. Edward P. Richardson, is the only record for the Gulf of Maine. Young fry, a few inches long, have been taken from time to time in Vineyard Sound, however, and in Buzzards Bay on the southern coast of Massachusetts between July and December. THE STICKLEBACKS. FAMILY GASTEROSTEIDAE Sticklebacks are small fish, made easily recog- nizable by the presence of two, three or more stout free spines on the back in front of the dorsal fin (spines that they can erect or depress at will) and by the fact that each ventral fin is represented by an even larger spine with only one or two rudimen- tary rays. Some of them have bony plates in the scaleless skin, but others do not. KEY TO GULF OF MAINE STICKLEBACKS 1. Seven dorsal spines or more Nine-spined stickleback, p. 307 Not more than five large dorsal spines 2 2. No bony plates on the upper part of the sides, but there is a bony ridge on either side of the abdomen Four-spined stickleback, p. 311 The upper part of the sides are armed with bony plates, and there is a plate in the midline of the belly, but there are no ridges on the sides of the abdomen 3 3 Many (28 or more) plates on each side Three-spined stickleback, p. 308 Only 5 or 6 plates on a side Two-spined stickleback, p. 310 Nine-spined stickleback Pungitius pungitius (Linnaeus) 1758 Jordan and Evermann, 1896-1900, p. 745. Description.- — The nine-spined stickleback is a slender little fish five to 6 times as long (not counting the caudal fin) as it is deep, with a very slim caudal peduncle. The latter usually has a well-developed longitudinal keel on either side; but this keel may be very low or even wanting. There are no bony plates along the sides of the body, but only along the bases of the anal and dorsal fins and on the caudal keels. There are no true scales. The most distinctive character is that there usually are 9 spines on the midline of the back (from 7 to 12 have been counted) in a con- tinuous row from close in front of the pectorals to the dorsal fin, set in a slightly zigzag line and leaning alternately to one side and to the other. The spines are weakly curved rearward; wider at the base than at the tip; fairly uniform in size; about one-half to one-third as long as the height of the dorsal fin; each has a small triangular fin membrane at its base; and there is a shallow groove along the back, into which the spines can be de- pressed. Each ventral fin is represented by a stout curved spine thicker and longer than the dorsal spines. The dorsal and anal fins (the former stands above the latter) are alike in form, tapering from front to rear, the anal preceded by a single stout recurved spine. The tail fin is weakly rounded. Size. — Large adults are seldom more than 3 inches long, more commonly 2 to 2%. Figure 166. — Nine-spined stickleback (Pungitius pungitius). From Bigelow and Welsh. 308 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Color. — Usually dull olive brown above, the upper part of the sides faintly barred or blotched darker; the belly silvery; the pubic and thoracic regions often black. The color varies, however, with the season of the year, with the state of sexual maturity, and with the color of the bottom on which the fish is living, those on dark mud being darker and those on bright sand paler. All become more brilliant during the breeding season when reddish tints appear under the head, the belly turns greenish, and black dots develop here and there over the entire body. The male has also been described as assuming a rosy tint beneath. Habits. — Since the range of the nine-spined stickleback hardly touches the open waters of our Gulf, we need only note that its mode of life is much the same as those of its three-spined relative next to be considered (p. 308); that it is similarly destructive to the spawn and young of other fish, and similarly pugnacious. Probably it spawns in summer w on the shores of the Gulf, for its breeding season in northern Europe covers June and July. The male often (but not always) builds a nest attached to grass or weeds which the female spawns, and he guards nest and eggs until the latter hatch, which occurs in about 12 days. General range. — This is one of the most widely ranging of northern fishes, occurring both in fresh water and in salt in the northern parts of both hemispheres; from northern Scandinavia to France, the western Mediterranean and the Black Sea on the European coast; from Arctic seas south to New York along the American, and westward to Saskatchewan and Alaska. Occurrence in the Gulf of Maine. — This stickle- back is to be found all around the shores of the «° At Woods Hole it spawns in April and May. Gulf of Maine from Nova Scotia and the Bay of Fundy to Cape Cod, but it is chiefly restricted there to harbors and the creeks in salt marshes, where large numbers may often be taken in com- pany with the mummichogs that swarm in such locations, and where it is to be found throughout the year. It is also found in fresh water. In fact, the most exposed situations around the Gulf, where we have heard of it, are Biddeford Pool, Maine,61 Passamaquoddy Bay,62 and St. Mary's Bay on the west coast of Nova Scotia. Commercial importance. — This stickleback is of no commercial importance in America, but it is sometimes tried out for oil in northern Europe when enough can be caught. Three-spined stickleback Gasterosteus aculeatus Linnaeus 1758 Two-spined stickleback; Stickleback; Thorn- fish; Thornback Jordan and Evermann, 1896-1900, p. 747. Description. — The three-spined stickleback has a very slender caudal peduncle, and squarish tail fin, like its nine-spined relative, but it is a stouter fish, being about one-fourth as deep as long, and it is more flattened sidewise. Its most diagnostic characters are the number of dorsal spines, of which there are three (occasionally four and rarely five), with the first two usually much the larger, and each with a small triangular fin membrane; the small size of the anal spine (this is free in the three-spined stickleback but attached to the fin by the fin membrane in the four-spined) ; and espe- cially the presence of a series of 28 to 33 bony plates on each side, besides a single ventral plate « MaeCoy, Bull. 74, Boston Soe. Nat. Hist. 1935, p. 16. " Huntsman, Contr. Canadian Biol., (1921) 1922, p. 61. "I! Figure 167. — Three-spined stickleback (Gasterosteus aculeal-us), Woods Hole. From Jordan and Evermann. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 309 on the lower surface between and behind the ventral fins. The fact that the dorsal fin (1 spine, 10 to 14 rays) originates some distance in front of the anal (1 spine, 8 to 10 rays) is diagnostic also, while its ventral spines are longer and stouter than those of the nine-spined stickleback. This is one of the most variable of fishes; Smitt,63 lists no less than 32 named species or races based on its varieties. Thus its dorsal spines may be long or short, and they vary in number as noted above; its bony plates range from none at all to very well developed; and its caudal peduncle may be keeled or it may not. Most American authors have recognized an American species, at the least as contrasted with a European, the former supposedly with longer dorsal spines; the latter with shorter. But the long-spined, as well as the short-spined form is known to occur on the other side of the Atlantic with every possible gradation between the two. Seeing that we have found both in the Gulf of Maine among fish indistinguishable otherwise, we believe that the various forms are environ- mental races of the one species. And this is well established for the relative strength of the dermal armature, which is weak in fresh water, but strong in salt. Color. — This stickleback is extremely variable in color, a fact hardly mentioned in most American accounts. Thej7 are deep grayish, or olive, or greenish-brown above, or sometimes blue; paler and often with silvery reflections on the sides; silvery on the belly. The fins are pale, except that the fin membranes often are red. In breeding season the males are described as turning reddish below from nose to vent and often up the sides. Tn females, the whole body except the top of the back may then be reddish; at the same time the back turns brownish with trans- verse bands, and the sides develop brassy reflections. Size. — Maximum length about 4 inches, but few of them are more than 3 inches long. Tt matures sexually at a length of about 2 inches. Habits.- — This is distinctively a shore fish like all the sticklebacks, the great majority of them living their whole lives in estuarine situations. But it is equally at home in sea water of full salinity as in fresh water. And enough stray » Scandinavian fishes, vol. 2, 1895, p. 648. out to sea for it to be rather a common experience to pick up a few here and there in the tow net, far from land. On such occasions they usually hide in clumps of floating eelgrass (Zostera) or of rockweed (Fucus, Ascophyllum) ; indeed we have learned to expect a stickleback or two whenever we dip up bunches of weed of any size. These wanderers keep to the surface except, perhaps, in very rough weather. Tt is a permanent all-the-year resident where- ever it is found alongshore, entering creeks and the mouths of streams in the spring to spawn, and dropping down into slightly deeper water for the winter. Tn such situations it probably lies in schools in a more or less sluggish condition while the temperature is lowest.64 It is a pro- verbially pugnacious fish, using its spines with good effect as weapons of offense and defense, even on other fishes much larger than itself. Tt feeds indiscriminately on the smaller inverte- brates, on small fish fry, and on fish eggs, to which it is exceedingly destructive in fresh water. The diet list of specimens examined by Vinal Edwards at "Woods Hole included copepods, of which they are often full, isopods, schizapod shrimps, and young squid, while some had fed on diatoms only. And it is not only omnivorous but very voracious. This stickleback affords the classic instance of nest building and of the care of eggs among fishes, and its nesting has been described so often in popular natural histories that a bare outline will suffice here.66 Recent studies in Europe make it likely that this stickleback spawns chiefly in brackish or fresh water, if not exclusively there, for which purpose it enters the estuaries and the mouths of streams. The spawning time is pro- bably the same in the Gulf of Maine (May to June) as in north European waters,66 when the fish assume the nuptial dress described above, and the males fight fiercely. It is the male that builds the nest, selecting some sheltered spot in shoal water for his purpose, or some rock pool. Here he makes a barrel-shaped mass of bits of grass, weed, and other vegetation an inch or so in diameter, cementing it together with mucous threads, which he spins from his kidneys, and « Large numbers are sometime? seined ia winter in Scandinavian waters. « Smitt (Scandinavian Fishes, vol. 2, 1895, pp. 653-656) and Regan (The freshwater fishes of the British Isles, 1911, pp. 247-249) give accounts of the nest building on which the following is based. •» About Woods Hole it spawns from May until the last week. In July. 310 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE weighting it down with pebbles. He then escorts one or a succession of females to this nest, and each of them deposits about 100 to 150 eggs in the central cavity. The male then enters the nest to fertilize the eggs, which stick in clumps to each other and to the nest. Incubation occupies 6 to 10 days, during which period the male guards the nest, driving away intruders large or small. He tears down the nest when hatching-time ap- proaches, but he continues to guard the fry until these can shift for themselves. Many males die after spawning. Those that survive go back to sea in summer; the females, too. Figure 168.— Egg. Figure 169. — Larva, newly hatched, 4.3 mm. Figure 170. — Larva, 6.3 mm. Three-spined stickleback (Gasterosteus aculeatus). After Kuntz and Radcliffe. The young fish are 4.25 to 4.5 mm. long when hatched. The yolk sac is absorbed in three or four days; when a week old they are almost 8 mm. long; and the fry are of adult form with fins and spines fully formed when 6 weeks old, and 14 to 16 mm. long.67 They are 1% to 2 inches (40-50 mm.) long when 2 years old, 2 to 2% (50-55 mm.) at 3 years, according to European studies. General range. — Coasts and fresh waters of the northern hemisphere; from Labrador, the Strait of Belle Isle and northern Newfoundland to lower Chesapeake Bay on the eastern coast of America, and represented on the northwestern coast by a form (Gasterosteus cataphractus Pallas 1811) that probably is identical with the Atlantic species. Its European range is from northern Norway and Iceland to Spain, the Mediterranean, and the Black Sea. Occurrence in the Gulf of Maine. — This stickle- back is very plentiful all around the shores of the Gulf from Nova Scotia to Cape Cod, living indifferently in brackish water and in salt. The ditches and creeks of the tidal marshes, brackish ponds and lagoons, rock pools, and weedy shores in shallow water are its favorite habitats. It may be found practically anywhere in such places, often in great numbers and in company with other sticklebacks, for it is the commonest of its tribe in the Gulf, as it is about Woods Hole. And so many of them drift out to sea around the shores of the open Gulf that we have taken them on the eastern part of Georges Bank; over German Bank; in the western basin off Cape Cod; near the Isles of Shoals; off Seguin Island; and off Matinicus Island. In the Bay of Fundy, however, they are known only close to land and in the mouths of estuaries. Importance. — This little fish is of no commercial value in America. In Scandinavia, however, it is sometimes seined in such quantities that it is worth boiling down for oil. Two-spined stickleback Gasterosteus wheatlandi Putnam 1867 68 Jordan and Evermann, 1896-1900, as Gasterosteus gladiun- cul-us Kendall, p. 2836. Description.- — This stickleback is said to differ from the three-spined stickleback in having a deeper body, fewer fin rays (9 or 10 dorsal and 7 or 8 anal) ; fewer dermal plates (5 or 6 as against 28 to 33); a caudal peduncle without keels; and a strong cusp both above and below at the base of " Figures of stages in development of this fish are given by Kuntz and Radcliffe (Bull. U. S. Bur. Fish., vol. 35, 1918, p. 131); A. Agassiz (Proe. Amer. Acad. Arts Soi., vol. 17, 1882, p. 288, plate 9), and by Ehrenbaum (Nor- disches Plankton, vol. 1, 1905-1909, p. 319). M This is the Gasterosleus blaculeatus of MitchUl 1815 and Storer 1867; bispinosus of Walbaum 1792; gladiunculus of Kendall 1896, but not the O. bispinosus of Jordan and Evermann 189G, which is a variety of Q. aculeatus. For the reason for using the specific name wheatlandi, see Hubbs, Occasional Papers, Museum of Zoology, University of Michigan, No. 200, 1925. FISHES OF THE GULF OF MAINE 311 the ventral spine. Dr. Kendall writes 69 that care- ful examination of large series has convinced him that this is actually a distinct species, not a race of the extremely variable three-spined stickle- back, although he saw one specimen apparently intermediate between the two. Color.— Grass-green above in life, mottled and finely speckled with black on the top of the head and back; sides of head and body golden with dark blotches; breast silvery; ventral fins scarlet. Habits. — Its mode of life is the same as that of the three-spined species so far as known, and sticklebacks of this type have been described as building nests with bits of straw on sandy bottom in New York waters, 70 but the two species or races have been confused so often that nothing more definite can be said of its habits. General range.- — Newfoundland to New York. Occurrence in the Gulf of Maine.- — Sticklebacks of this type are common in company with the three-spined sticklebacks in Passamaquoddy and St. Mary Bays 71 and in the Bay of Fundy. They may be expected anywhere on the Maine coast, being recorded at Winter Harbor; off Monhegan Island; off Seguin Island; from Casco Bay and its tributaries in both salt and brackish water; and from Kittery. They have also been taken at Swampscott, in Massachusetts Bay, and they are fairly common in summer at Woods Hole. We have taken them in our tow-nets, also, off Cape Porpoise; on Platts Bank; in the Western Basin of the Gulf of Maine; and on German Bank. « Proc. U. S. Nat. Mus., vol. 18, 18%, p. 624. '• Bean, Bull. 60, New York State Mus., Zool. 9, 1903, p. 341. » Huntsman, Contrib. Canadian Biol. (1921) 1922. p. 61. Four-spined stickleback Apeltes quadracus (Mitchill) 1815 Bloody Stickleback Jordan and Evermann, 1896-1900, p. 752. Description.- — The four-spined stickleback has no bony plates in its scaleless skin, but it does have a bony ridge on each side of the abdomen, making it triangular in cross section, with flat belly and sharp back; this gives it an aspect very different from the other sticklebacks. It is fusiform in side view, tapering to the rather pointed nose and to the slim caudal peduncle. There are two to four free dorsal spines standing close one behind the other, inclining alternately to one side or the other, and another spine is attached to the dorsal fin by the fin membrane. The anal fin is similarly pre- ceded by an attached spine, and each ventral fin is represented by a stouter curved spine, strongly saw-edged, followed by about two slender rays. The dorsal fin stands over the anal as in the nine- spined species, but both these fins are more rounded in outline, wliile the caudal fin is rela- tively longer and narrower than in any of our other sticklebacks. Color. — Brownish olive or greenish brown above, with dark mottlings that alternate below the lat- eral line with the silvery white of the belly. The fin membrane of the ventrals is red. Males are much darker than females. Size. — One and one-half to two and one-half inches long. habits.- — This is a common little fish in the salt marshes, where it consorts with other stick- lebacks and with mummichogs. Like tbe three- Figure 171. — Four-spined stickleback (Apeltes quadracus), Woods Hole. From Jordan and Evermann. Drawing by A. H. Baldwin. 312 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE spined stickleback it often runs up into fresh- water, though it is primarily a salt and brackish water fish. And it is never found far in from the coast or out at sea. In the Woods Hole region this stickleback spawns as early as May and as late as the last week of July, after which spent females are found; but the onset of spawning may be somewhat later in the cooler waters of the Gulf. The male builds a nest of plant fragments which it brings in its mouth, cemented with mucous threads that he spins out of a pore near his vent; a small rudi- mentary affair, however, compared with that of the three-spined stickleback, described by Ryder72 as less than 1 inch in diameter, conical, with an opening at the top. Finally, the male stickleback picks up the eggs that have been laid by the female and deposits them in the hollow at the top of the nest, guarding them, presumably, during incuba- tion. The eggs are yellow, approximately 1.66 mm. in diameter; they sink like those of the other sticklebacks and stick together in clumps. Incu- bation occupies six days or thereabouts at labora- tory temperature (about 70°). Newly hatched larvae are about 4.5 mm. long and similar in appearance to those of the three-spined species but more densely pigmented.73 General range.- — This is an American fish, known along the coast from the southern side of the Gulf of St. Lawrence and Nova Scotia to Virginia; at home both in salt water and in brackish, and running up into fresh water. Occurrence in the Gulf of Maine.- — This stickle- back is common all around the shores of the Gulf on the Nova Scotian side as well as the New Eng- land side. We have taken it at Yarmouth; Huntsman 7i records it from St. Mary Bay and along the New Brunswick shore well within the Bay of Fundy (Maine has usually been given as its northern limit), and there are many locality records for the coasts of Maine and Massachusetts. But it is so much more closely restricted to estuar- ine situations than is its three-spined relative (p. 310) that we have never taken it in our tow nets nor do we find a single record of it in the open sea. On the south shore of New England it is a year-round resident. Probably this is equally true in the Gulf, where it may be expected to gather in the bottoms of the deeper creeks in winter, as it is known to do in Chesapeake Bay. It resembles the three-spined stickleback in its feeding habits so far as known (copepods and other small crustaceans being its chief diet) and in its general mode of life. THE PIPEFISHES. FAMILY SYNGNATHIDAE The forward portion of the head has the form of a long tubular snout in the pipefishes, with the small mouth situated at its tip; the skin is armed with rings of bony plates; there is only one dorsal fin (soft-rayed) , the body is very slender, and there are no ventrals. The snout recalls that of the trumpetfishes (p. 316), but pipefishes differ from them and from most other bony fishes in the struc- ture of their gills, which form tufts of small rounded lobes, instead of the familiar filaments. Their general affinity in this respect is with the group of which the sticklebacks are the most familiar ex- ponents. There are many species of pipefishes in warm seas, but only one inhabits the Gulf of Maine regularly, while a second has been re- corded there — a stray from the south. '» Bull. U. S. Fish. Comm., vol. I, 1882, p. 24. " The early development is described by Ryder (Bull. U. S. Fish Comm., vol. 1, 1882, p. 24) and by Kuntz and Radclifle (Bull. TJ. S. Bur. Fish., vol. 35, 1918, p. 132). " Conlrib. Canadian Biol. (1921) 1922, p. 61. KEY TO GULF OF MAINE PIPEFISHES 1. Dorsal fin with 35 to 41 rays; 18 to 20 bony plates in front of the vent and 36 to 42 behind it Common pipefish, p. 312 Dorsal fin with 29 to 31 rays; 16 or 17 bony plates in front of the vent and 31 to 35 behind it Pelagic pipefish, p. 314 Common pipefish Syngnathus fuscus Storer75 1839 Jordan and Evermann, 1896-1900, p. 770 as Siphostoma fuscum (Storer). Description. — This is a very slender little fish, particularly so behind the vent, males being about 35 times as long as they are deep and females about 30 times. The head occupies one-eighth to one- '* This is the only pipefish that occurs regularly on our northern coasts. Jordan, Evermann, and Clark (Rept. U. S. Comm. Fish. [1928], 1930, Pt. 2, p. 242) refer it to the genus St/rides Jordan and Evermann, 1927. For a syn- opsis of the various other species of the genus see Jordan and Evermann, Bull. 47, U. S. Nat. Mus., Pt. 1, 1896. p. 961. FISHES OF THE GULF OF MAINE 313 Figure 172. — Pipefish (Syngnalhus fuscus) . After Bigelow and Welsh. ninth of the total length (in the trumpetfish it is nearly one-third); the snont is tubelike, blunt ended, with the small toothless mouth at its tip. The gill openings are very small. The body is hexagonal in cross section in front of the vent but is four-sided behind the dorsal fin and it is clothed in an armor of bony plates connected in rings, of which there are 18 to 20 in front of the vent and 36 to 42 behind the latter. The abdomen of the male is wider just back of the vent than elsewhere, with two lateral flaps that meet along the midline to form the so-called "marsupial" or brood pouch. The female lacks these. The dorsal fin (35 to 41 rays and 5 or 6 times as long as it is high) covers 4 or 5 of the bony rings in front of the vent and as many behind it. The caudal fin is rounded, its middle rays the longest. The anal is very small, close behind the vent; the pectorals are of moderate size; there are no ventral fins. Color. — Greenish, brownish, or olive above, cross-barred and mottled with darker. The lower parts of the gill covers are silvery. The lower parts of the sides are sprinkled with many tiny white dots, and the longitudinal angles separating sides from abdomen are marked by longitudinal brown bars. The lower surface is colorless on the snout, but pale to golden yellow thence bark to the vent, with the marsupial flaps flesh-colored. The dorsal and pectoral fins are pale, but the caudal is brown.78 Pipefishes change color according to the color of their surroundings. We have seen them of various shades of olive and brown; and red ones have been described. Size. — Usually 4 to 8 inches long; occasionally up to 12 inches. Habits. — The chief home of this pipefish is among eelgrass or seaweeds, both in salt marshes, " Colors based on Storer's (Fishes of Massachusetts, 1867, p. 412) account and on the specimens we have examined. 210941—63 r-21 harbors, and river mouths, where it often goes up into brackish water, and on more open shores as well. In such locations it is caught as often today by boys dipping up mummichogs for bait as it was when Storer wrote of it, nearly a century ago. The pipefish, like the three-spined stickleback, sometimes strays out to sea on the surface, and while we have never taken it in our tow nets, Kendall n has often found it under floating rock- weed along the Maine coast. But they are so seldom taken at any distance out from the land that the capture of four specimens at a depth of 19 fathoms south of No Mans Land, February 5, 1930, is of present interest, though outside the limits of our Gulf. There is no reason to suppose pipefish are at all migratory, for they are resident in the eelgrass {Zostera) at Woods Hole throughout the year. They usually propel themselves by the dorsal fin, but they can travel swiftly when alarmed, with eel-like strokes of the tail from side to side. And they are able to roll their eyeballs separately, an interesting habit described many years ago by Lyman.78 They feed chiefly on minute Crustacea (copepods especially and amphipods), also to some extent on fish eggs, on very small fish fry, and no doubt indiscriminately for that matter on any small marine animals. And their snouts are so disten- sible that they can swallow larger prey than one might expect. In capturing its prey, the pipefish has been described as expelling the water from the snout and pharynx by muscular action, depending on the return rush to sweep in its victims. Pipe- fishes have few enemies so far as known. « Bull. U. S. Fish Comm., vol. 18, 1896, p. 623. '• Proc. Boston Soc. Nat. Hist., vol. 7, 1861, pp. 75-76. 314 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Breeding.79 — On the southern shores of New England pipefish breed from March to August; probably through this same period on the shores of the Gulf of Maine. Male pipefishes nurse the eggs in the brood pouch (p. 313), the flaps of which ordinarily lie flat but are swollen and have their edges cemented together during the breeding season. The protruding oviduct of the female is inserted into the opening of the pouch of the male and a dozen or more eggs are passed over. This occurs several times in succession, with intervals of rest, until the pouch is filled, the male working the eggs down toward the rear end of his pouch by body contortions. Fertilization is supposed to take place during the transference of the eggs from one parent to the other. The eggs become embedded in the lining of the brood pouch, and it has been established for the European pipefish (probably this applies equally to our North American species) that the embryos within the eggs are nourished by the epithelial lining layer of the pouch, so that the latter functions as a placenta.80 Incubation occupies about 10 days, according to Gudger, and the young are retained in the brood pouch until they are 8 or 9 mm. long, when the yolk sac has been absorbed. The young pipefish are then ready for independent existence, and once they leave the pouch they never return to it, as young sea horses (Hippocampus) are said to do (p. 315). Several observers agree on this, among them Miss Marie Poland (now Mrs. C. J. Fish), who kept pipefish under observation at the laboratory of the United States Bureau of Fisheries at Woods Hole during the summer of 1922. Pipefish fry kept in aquaria have been found to grow from about %-inch (10 mm.) to about 2%- inches (70 mm.) in length within about 2 months after hatching.81 It is probable that they mature when about 1 year old. General range. — Coast of eastern North America, in salt and brackish water, from the southern side '» For a historical survey and a general account, of the breeding of the closely allied Siphnstoma floridae see Gudger (Proc. U. S. Nat. Mus., vol. 29, 1900. pp. 447-500, pis: 5-11). 1 For detailed (if somewhat divergent) accounts of this interesting phe- nomenon see Huot (Annales des Sciences Naturelles, Ser. 8, Zoologie, vol. 14, 1902, pp. 197-288) and Cohn (Anatomischer Anzeiger, Centralblatt fur die gesamte wissenschaftliche Anatomie, vol. 24, 1904, pp. 192-199). M Tracy, 40th Kept., Rhode Island Comm. Inland Fish., 1910, p. 93. of the Gulf of St. Lawrence 82 and outer Nova Scotia at Halifax, to South Carolina.83 Occurrence in the Gulf of Maine. — The pipefish has been recorded from so many localities along Maine and Massachusetts that it is evidently to be expected anywhere there, in suitable situations; it is not uncommon in the Bay of Fundy; it has been reported from outer Nova Scotian waters and is common locally in the southern side of the Gulf of St. Lawrence, as noted in the preceding paragraph. It is probable also that they breed in every favorable locality all around the shores of the Gulf, but there are local differences in this respect, for while St. Mary Bay, Annapolis Basin, and Cobequid Bay, on the Nova Scotian shore of the Bay of Fundy, are breeding centers according to Huntsman, large specimens alone are known about Passamaquoddy Bay on the New Bruns- wick side. No doubt the estuarine waters from the Massachusetts Bay region to Penobscot Bay are favorable nurseries. Importance. — The pipefish is of no commercial importance. Pelagic pipefish Syngnathus pelagicus Linnaeus 1758 Jordan and Evermann, Siphostoma pelagicum (Osbeck), 1895-1900, p. 767. Description. — Most of the species of pipefishes resemble one another so closely that they can be named only by critical examination. The pelagic pipefish differs from its common shore relative of New England (5". fuscus) by having fewer dorsal rays (29 to 31) and fewer rings of bony plates, of which only 16 or 17 are in front of the vent, and 31 to 35 behind the vent. General range. — Tropical Atlantic, northward with the Gulf Stream; also the Mediterranean, and the Southern Pacific and Tropical Indian Oceans. Occurrence in the Gulf of Maine. — A single specimen, 3% inches (89 mm.) long, taken on Georges Bank (Lat. 42° 09' N.; Long. 66° 41' W.), September 20, 1927, by the Albratross His, the only Gulf of Maine record. This specimen was dipped up with a mass of gulf weed (Sargassum), and was the only one found in a large amount of weed that was examined. 11 Leim (Proc. Nova Scotian Inst. Sci., vol. 20, 1940, p. 38) found them com- mon at Prince Edward Island. " There is a specimen from Charleston, S. C, in the Museum of Com para- i ive Zoology. FISHES OF THE GULF OF MAINE 315 THE SEA HORSES. FAMILY HIPPOCAMPIDAE Sea horse Hippocampus hudsonius DeKay 1842 Jordan and Evermann, 1896-1900, p. 777. Figure 173. — Sea-horse (//ippoca mp us Audsonius), Virginia. From Goode. Drawing by H. L. Todd. Description. — The sea horse grotesquely re- sembles the "knight" in an ordinary set of wooden chessmen in its sidewise flattened body, in its deep convex belly, in its curved neck and in its curious horse-like head carried at right angles to the general axis of the body. The head is surmounted by a pentagonal star-shaped "coro- net," and the snout is tubular with the small oblique mouth at its tip, like that of its relative the pipefish. It has a sharp spine on each side above the eye and one behind it, a third over the gill cover, and a fourth on the side of the throat, which sometimes terminate in short fleshy fila- ments; also a blunt horn between the nostrils. Its neck, body, and tail are covered with rings of bony plates, 12 rings on the trunk, 32 to 35 on the tail, and each body ring is armed with four blunt spines. The body tapers suddenly behind the anal fin to a long tail, which is four-cornered in cross section, curled inward, and strongly pre- hensile. In the male the lower surface of the fore part of the tail bears the brood pouch, opening by a slit in front. The dorsal fin (about 19 rays) originates about midway of the length of the fish, opposite the vent, and runs backward over three and one-half rings to within half a ring of the commencement of the tail sector of the trunk. The very small anal fin stands opposite the rear part of the dorsal fin. The pectorals are of moderate size, broad based and round tipped; it has no ventral fins and no caudal fin. Color. — Light brown or dusky to ashen gray or yellow, variously mottled and blotched with paler and darker, sometimes spangled with silver dots, sometimes plain colored. European sea horses change color according to their surroundings, tints of red, yellow, brown, and white all being within their capabilities, and it is probable that the Ameri- can species is equally adaptable. Size. — Adults usually are 3 to 6 inches long; one of 7% inches is the largest on record.84 Habits. — Sea horses dwell chiefly among eel- grass and seaweed,85 where they cling with their prehensile tails, monkeylike, to some stalk. They usually swim in a vertical position by undulations of the dorsal fin, not with the tail, the trunk being too stiff for much sidewise motion. Sea horses feed on minute Crustacea and on various larvae, in fact on any animal small enough, sucking in their prey as the pipefish does (p. 313.) They breed in summer 86 and the breeding habits resemble those of the pipefish, the male nursing the eggs in his brood pouch where they are deposited a few at a tune by the female in repeated pairings. The young, of which there may be as many as 150, are about 10 to 12 mm. long at hatching. When the yolk sac is absorbed the father squeezes them out of the brood sac, and they already resemble the adult in general appearance within a few days after they are set free. According to some students they swim out and in at will, but this calls for verification. " Bull. New York Zool. Soc, vol. 16, 1913, p. 972. »•" QUI (Proc. U. S. Nat. Mus., vol. 28, 1905, pp. 805-814) has given an ex- cellent account of the habits and life history of the sea horse. »« Ryder. (Bull. U. S. Fish Comrn., vol. I, 1882, pp. 191-199) describes its development. 316 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE General range. — Atlantic Coast of North America, occurring regularly from South Carolina to Cape Cod, and to Nova Scotia as a stray. Occurrence in the Ouij of Maine. — The sea horse is not common much beyond New York. Only a few are foimd each year about Woods Hole, chiefly in July, August, and September, and they so rarely stray past the elbow of Cape Cod that we have found only one definite (Provincetown) and one dubious (Massachusetts Bay) record of its capture in the inner parts of the Gulf of Maine, dead or alive; and one record for Georges Bank. Three specimens of the sea horse were also reported from Nova Scotia more than % of a century ago;87 and Vladykov and McKenzie have reported one, picked up in Terrance Bay, on the outer Nova Scotian coast, Sept. 18, 1934, by V. Slaunhite.88 Commercial importance. — The sea horse is of no commercial value, but it is an object of constant interest to visitors to marine aquaria. THE TRUMPETFISHES. FAMILY FISTULARIIDAE The trumpetfishes are characterized by their slender bodies and tremendously long heads, as well as by the fact that the anterior bones of the skull are prolonged in a very long tube with the small mouth at its tip. The only other Gulf of Maine species with which they could possibly be confused is the pipefish (p. 312). In the latter, however, the tubular snout occupies only about one-eighteenth of the length of the fish whereas in trumpetfishes it is nearly one-fourth. Further- more, the pipefishes lack ventral fins which the trumpetfishes have, while the caudal fin of the trumpetfishes is forked, but that of the pipefishes is rounded. Trumpetfish Fistularia tabacaria Linnaeus 1758 CORNETFISH Jordan and Evermann, 1896-1900, p. 757. Description. — The slender body and very long tubular snout of this fish are mentioned above. The body (to base of caudal fin) is about 30 to 35 times as long as it is deep and only about two- thirds as deep as it is thick. The head occupies almost one-third and the snout about one-fourth of the body length. The bones of the snout are so loosely united that the snout is very distensible. The mouth is small, situated somewhat obliquely at the tip of the snout, and the lower jaw projects a little beyond the upper. The caudal fin is deeply forked and its middle rays are prolonged in a filament about as long as the snout, but which is likely to be broken off. Both the dorsal and the anal fins are triangular, higher than long, the former stand- ing exactly above the latter, about three-fourths of the distance back from eye toward base of caudal fin. The ventrals are very small, and are consid- erably nearer to the eye than to the rear end of the body (about one-third of the way from eye toward the base of caudal fin). The skin is without scales but with a row of embedded bony plates or shields along either side, conspicuous rearward. Color. — Greenish brown above, the back and sides marked with many large, oblong, pale blue spots and with about 10 dark cross bars; the lower surface is pale and silvery; the caudal filament deep blue. Size. — Said to reach a length of 6 feet, but the few specimens that stray northward are much smaller. General range. — Tropical, southward to middle Brazil, and common among the West Indies; rarely " By Knight (Catal. Fishes Nova Scotia, 1866, p. 9), as //. brairostris Storer (1839); later by Jones, (Proc. Nova Scotian Inst. Sci., vol. 5, Part 1, 1882, p. 95) as H. antiguorum Leach 1814. " Proc. Nova Scotian Inst. Sci., vol. 19, 1935, p. 5. Figure 174. — Trumpetfish (Fistularia tabacaria), from near Woods Hole. After Storer FISHES OF THE GULF OF MAINE 317 wandering northward as far as the Massachusetts Bay region, and straying to Nova Scotia.89 Occurrence in the Gulf of Maine. — There are only two records of the trumpetfish from the Gulf of Maine: a specimen taken at Rockport, Mass. (north side of Cape Ann) in September 1865, preserved in the collection of the Essex Institute, where it was examined and identified by Goode and Bean 90 and a second taken on the northern edge of Georges Bank by the trawler Flying Cloud on October 6, 1947, in a haul at 70 fathoms.91 Like other tropical fishes, however, it is not so rare west of Cape Cod, and a few small ones are taken at Woods Hole almost every year. THE MACKERELS. FAMILY SCOMBRIDAE The mackerels are a very homogeneous group, all of them agreeing in having a spiny dorsal as well as a soft dorsal fin, several small Unlets behind the latter and behind the anal, a very slender caudal peduncle, a deeply forked or lunate caudal fin, a very shapely form tapering both to snout and to tail, and velvety skin with very small scales. All, too, are predaceous, swift swimmers, and powerfully muscled, while all are fish of the open sea and more or less migratory. In the following key we mention all species so far actually recorded from within the limits of the Gulf of Maine, but it would not be astonishing if still others were to stray in from the open Atlantic from time to time.92 KEY TO GULF OF MAINE MACKERELS M 1 . The two dorsal fins are separated by a space at least as long as the length of the first dorsal 2 The two dorsal fins adjoin each other or are separated by a space much shorter than the length of the first dorsal.. 3 2. The sides below the mid line are silvery, not spotted Mackerel, p. 188, 317 The sides below the mid line are mottled with dusky blotches Chub Mackerel, p. 209, 333 3. Body scaleless, except along the lateral line and in the region of the shoulders (the so-called "corselet")-- 4 Entire body covered with scales •> 4. The lower part of the sides, below the lateral line, is marked with dark longitudinal bands, but there are no definite dark markings on the back Striped Bonito, p. 335 There are no dark markings on the lower side below the lateral line, but the back has dark markings False Albacore, p. 336 5. The anal fin is about twice as high as long; the corselet of large scales is obvious Tuna, p. 338 The anal fin only is about as high as long; there is no corselet of large scales 6 6. Second dorsal fin noticeably lower than the first dorsal Common Bonito, p. 337 Second dorsal fin at least as high as first dorsal 7 7. Lateral line with an abrupt downward curve under second dorsal fin Ca valla p. 349 Lateral line descending gradually 8 8. Sides with a few rows of oval bronze or yellowish spots and with one or two longitudinal dark streaks King Mackerel p. 348 Sides with bronze spots but without longitudinal dark streaks Spanish Mackerel p. 347 Mackerel Scomber scombrus Linnaeus 1758 Jordan and Evermann, 1896-1900, p. 866 Description. — The mackerel is fusiform in out- line, tapering rearward to a very slim caudal peduncle and forward to a pointed nose. Its body is about four and one-half to five and one-half times as long as it is deep, oval in section, thick and firm-muscled as are all its tribe. Its head is long (one-fourth of length to caudal) and its mouth large, gaping back to the middle of the eye (the premaxillaries are not protractile), while the jaws, " Dr. A. H. Leim reports the capture of a specimen at Port Mouton, Nova Scotia, on September 10, 1931; the specimen was recorded later by Vladykov (Proc. Nova Scotian Inst. Sci., vol. 19, 1935, p. 5) as Fistularia icrrata. which are of equal length, are armed with small, sharp, slender teeth. The eye is large, and the hollows in front of and behind it are filled with the so-called "adipose eyelid," a transparent, >» Bull. Esses Inst., vol. XI, 1879, p. 4. •i This specimen is in the Museum of Comparative Zoology. •i Fraser-Brunner, Ann. Mag. Nat. Hist. Ser. 12, vol. 3, No. 26, 1950, pp 131-163, has recently given a synopsis of the mackerels, with useful keys and excellent illustrations for all known species. We follow him in uniting them all in the old and inclusive family Scombrldae rather than Jordan, Evermann and Clark (Rept. U. S. Comm. Fish. (1928) Pt. 2, 1930) who have distributed them among four families, Cybiidae, Katsuwonidae, Scombridae, and Thunnidae. •> The long finned Albacore (Thunnus alalunga Bonnaterre, 1788) has been taken at Woods Hole, also on Banquereau Bank, off eastern Nova Scotia (Ooode and Bean, Bull. Essex Inst., vol. XI, 1S79, p. 15), so is likely to showup in the Gulf of Maine sooner or later. It is made easily recognizable among North Atlantic mackerel fishes by its very long pectoral fins which reach back past its second dorsal fin. 318 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 175. — Mackerel (Scomber scombrus). A, egg (European), after Holt; B, larva, 3.5 mm.; C, larva, 4.6 mm.; D, larva, 7.8 mm.; E, larva, 15 mm.; B-E, drawings by Luella E. Cable. gelatinous mass in the form of two scales, a for- ward and a hinder, which cover the eye except for a perpendicular slit over the pupil. There are two large dorsal fins: the first originat- ing over the middle of the pectoral fins when the latter are laid back is triangular, of 10 to 14 (usually 11, 12, or 13) rather weak spines that can be laid down along the midline of the back in a deep groove; the second dorsal, separated from the first by an interspace longer than the length of the latter, is smaller (9 to 15 rays, usually 12) and is followed by several small finlets, of which there are usually 5, but sometimes 4 or 6. The anal fin is similar to the second dorsal in shape and size, originates slightly behind it, and is similarly suc- ceeded by 5 small finlets that correspond to the dorsal finlets in size and shape. The caudal fin is broad, but short and deeply forked. The caudal peduncle bears two small longitudinal keels on either side but no median lateral keel, the absence of the latter being; a distinctive character. The ventral fins stand below the origin of the first dorsal and are small, as are the pectorals. The scales of the mackerel are so small that its skin feels velvety to the touch; indeed they are hardly to be seen on the belly with the naked eye, but those about the pectoral fins and on the shoulders are somewhat larger. Color.— The upper surface is dark steely to greenish blue, often almost blue-black on the head. The body is barred with 23 to 33 (usually 27 to 30) dark transverse bands 94 that run down in an ir- regular wavy course nearly to the mid-level of the body, below which there is a narrow dark streak running along each side from pectoral to tail fin. The pectorals are black or dusky at the base, the dorsals and caudal are gray or dusky. The jaws and gill covers are silvery. The lower parts of the sides are white with silvery, coppery, or brassy reflections and iridescence; the belly silvery white. « Hunt (Copeia, No. 117, pp. 53-59, April, 1923) describes the variations In these stripes among young mackerel. FISHES OF THE GULF OF MAINE 319 But the iridescent colors fade so rapidly after death that a dead fish gives little idea of the bril- liance of a living one. Size. — Most of the grown fish are between 14 and 18 inches long; a few reach a length close to 22 inches. Fourteen-inch fish weigh about 1 pound in the spring and about 1 ){ pounds in the fall when they are fat; 18-inch fish weigh about 2 to 2% pounds; a 22-inch mackerel will likely weigh 4 pounds. An unusually large mackerel is taken occasionally; in 1925, for example, the schooner Henrietta brought in one weighing 7}i pounds. Habits. — Mackerel are a swift-moving fish, swimming with very short sidewise movements of the rear part of the body and of the powerful caudal fin. When caught they beat a rapid tattoo with their tails on the bottom of the boat until exhausted. And they require so much oxygen for their vital processes that when the water is warm (hence its oxygen content low) they must keep swimming constantly, to bring sufficient flow of water to their gill filaments, or else they die.95 Despite their great activity, they do not leap above the surface, as various others of their tribe do, unless perhaps to escape some larger fish. The mackerel, like the herring, has the habit of gathering in dense schools of many thousands. It is not known how long these schools hold together; it would be especially interesting to know whether they do so through the winter when our mackerel are in deep water, but the general opinion of fish- ermen is that they do so throughout the migra- tions at least. Although the mackerel may scatter and the schools mix more or less, especially when they are feeding on the larger and more active members of the free-floating fauna as is said to happen in British waters, the members of any given school usually are all of about the same size, i. e., of the same age. Fish of the year almost always school separately from the others as Sette " has pointed out; he has also pointed out that this tendency of the fish to separate according to size is probably due to the fact that the larger ones swim faster than the smaller ones. Mackerel school by themselves, as a rule. But sometimes they are found mingled with herring, alewives, or shad, as Kendall 97 described. We m This interesting fact seoms first to have been reported by Hall (Amer. Jour. Physiol., vol. 93, 1930, pp. 417-421), and we have observed the same thing in the aquaria at the Woods Hole Oceanographic Institution. » Fish. Bull. V. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, p. 264. •' Bull. U. S. Bur. Fish., vol. 28, 1910, Pt. I, p. 287. have yet to learn how mackerel schools hold to- gether, whether by sight or by some other sense. And various explanations have been proposed to account for the schooling habit, such as that it is advantageous for feeding, that it is a concomitant of spawning (this would not explain its persistence out of the spawning season, however, or the fact that any given school is apt to contain green, and spent as well as ripe fish even at spawning time), or that it affords protection from enemies. But when all is said, the instinct prompting it remains a mystery. At any rate, schooling is not a neces- sity, though usual. When mackerel are at all plentiful, and even when they are not, numbers of single wandering fish are often hooked by persons trolling for them, and by flounder and cunner fishermen. Schools of mackerel are often seen at the surface. In the daytime they can be recognized by the appearance of the ripple they make, for this is less compact than that made either by herring or by menhaden. Neither do mackerel ordinarily "fin" or raise their noses above the surface, as is the common habit of the menhaden (p. 114). An observer at masthead height can per- haps see a school of mackerel as deep as 8 to 10 fathoms by day, if the water is calm, and the sun behind him. On dark nights the schools are likely to be betrayed by the "firing" of the water, caused bjT the luminescence of the tiny organisms that they disturb in their progress. Sette w reports one case of a school recognized by its firing as deep as 25 fathoms; but the water is seldom (if ever) clear enough in the Gulf of Maine for a submerged light to be visible from above, more than 15 fathoms down.99 The trail of bluish light left behind by individual fish as they dart to one side or the other, while one rows or sails through a school on a moonless, overcast night when the water is firing, is the most beautiful spectacle that our coastal waters afford, and one with which every mackerel fisherman is familiar. No one knows how greatly the movements of the mackerel, from day to day, result from invol- untary drifting with the circulatory movements of the water, which are different at different depths, and how greatly they depend on the directive swimming of the mackerel themselves. Our only •> Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, p. 267. » For observations on the visual transparency of the water of our Quit, see Bigelow, Bull. U. S. Bur. Fish., vol. 40, Pt. 2, 1927, p. 822. 320 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE contribution in this regard is that we once were able to follow on foot beside a school that was advancing along the Scituate shore at a rate of about 3 to 4 miles an hour, against a tidal current of about one-half knot, until the fish swung offshore and out of our sight. The speed at which a school travels when it is not disturbed depends, it seems, on the size of the fish of which it is composed. It has been observed by Sette that mackerel less than one year old swim at about 6 sea miles per hour (10 ft. a second) while circling inside a live car; yearlings at a rate of about 11% sea miles per hour (19 ft. a second), or nearly twice as fast. We find no definite observations on the normal speed of the larger fish, and no one knows how rapidly a mackerel may swim for a short distance, if it is disturbed. Mackerel seen during the warmer months of the year are always swimming, but this rule may not apply in winter, when the water holds more dis- solved oxygen because colder, and when it is probable that their demand upon it is lower. The mackerel is a fish of the open sea; while numbers of them, small ones especially, often enter estuaries and harbors in search of food, they never run up into fresh water. Neither are they directly dependent either on the coastline or on the bottom in any way at any stage in their lives. They are often encountered far out over the outer part of the shelf of the continent. But they are most numerous within the inner half of the continental shelf during the fishing season, and their normal range seems not to extend oceanward beyond the upper part of the continental slope, in which they contrast with their relatives the tunas, the bonitos, and the albacores. The depth-range of the mackerel is from the surface down to perhaps 100 fathoms at one season or another. (We recur to this in discussing the occurrence of mackerel in the Gulf of Maine, page 325.) From spring through summer and well into the autumn, the mackerel are in the upper water layers; shoaler, mostly, than some 25 to 30 fath- oms, and schools of all sizes come to the surface more or less frequently then. But they frequently disappear from the surface, often for considerable periods. And it seems, from fishermen's reports, that the larger sizes tend to swim deeper than the smaller ones, on the whole, especially in mid and late summer.1 It is probable, also, that their vertical movements during the warmer part of the year, when they are feeding actively, are governed chiefly by the level at which food is most abun- dant, which for the most part is shoaler than about 50 fathoms, at least on our side of the Atlantic. The highest temperature in which mackerel are commonly seen is about 68° F. (20° C). At the opposite extreme they are sometimes found in abundance in water of 46°-47° (8° C.) ; and com- mercial catches are sometimes made in water as cold as 44°-45° (7° C), but odd mackerel only have been taken in temperatures lower than that 2 in American waters. Large catches of mackerel are made, however, by trawlers in the North Sea in whiter in water as cold as 43°-45° (6°-7° C). But as Sette has emphasized, the European mackerel differs racially from the American, and may differ in its temperature rela- tions as well. Food. — We may assume that the diet of the young mackerel is at first much the same in the Gulf of Maine as it is in the English Channel,3 namely, copepod larvae and eggs, the smaller adult copepods, various other minute pelagic Crustacea, and small fish larvae. But the young fish depend more and more upon larger prey as they grow. Our Gulf of Maine mackerel have repeatedly been seen packed full of Calanus, the "red feed" or "cayenne" of fishermen, as well as with other copepods (we have examined many in this condition). They also feed greedily, as do herring, on euphausiid shrimps (p. 89), especially in the northeastern part of the Gulf where these crustaceans come to the surface in abundance. Various other planktonic animals also enter regu- larly into the dietary of the mackerel. Thus, Doctor Kendall writes in his field notes that some of the fish caught on the northern part of Georges Bank hi August 1896, were packed with crab larvae, others were full of Sagittae, others, again, of Sagittae and amphipods (Euthernisto) , of small copepods (Temora), or of red feed {Calanus), so that even fish from the same school had selected the various members of the drifting community in varying proportion. ' See Sette, Fish. Bull. U. S. Fish and Wildlife Service, vol. 61, Bull. 49, 1950, p. 267, for further discussion of this point. » Sette (Fish. Bull, U. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, p. 257) mentions one winter record from about 40° (4.5° C.) on Georges Bank. ' Lebour (Jour. Mar. Biol. Assoc. United Kingdom, vol. 12, N. Ser., No. 2, 1920, p. 305) gives diet lists for 90 larval mackerel ranging from 5 mm. to 13.5 mm. in length, taken in the English Channel. FISHES OF THE GULF OF MAINE 321 Similarly, 1,000 mackerel caught near Woods Hole from June to August contained pelagic amphipods (Euthemisto) , copepods, squid, and launce;4 others taken off No Man's Land have been found full of shelled pteropods (Limacina). And a series of small fish examined by Vinal Edwards contained copepods, shrimps, crustacean and molluscan larvae, annelid worms, appendi- cularians, squid, fish eggs, and fish fry such as herring, silversides, and launce. In short, prac- tically all the floating animals, not too large or too small, regularly serve for the nourishment of mackerel except the Medusae and ctenophores, and a diet list for any given locality would include all the local pelagic Crustacea and their larvae. Mackerel have often been seen to bite the centers out of large Medusae, but, as Nilsson suggests, they probably do this for the amphipods (Hyperia) that live commensal within the cavities of the jellyfish, not for the sake of the latter. Mackerel also eat all kinds of small fish, to a greater or less extent according to circumstances. In the Gulf of Maine they devour large numbers of small herring, launce, and even smaller mack- erel. They likewise feed on pelagic fish eggs when available, oftenest on those of their own species. And they bite greedily on almost any bait, especially if it moves, such as a bit of mack- erel belly skin, a piece of clam, a piece of sea worm (Nereis), a shining jig, spoon or spinner of appro- priate size, or an artificial fly, white, red, or silver- bodied. Side by side with these comparatively large objects mackerel are also known to take various microscopic organisms, chiefly the com- moner peridinians and diatoms, but they never feed extensively on these as menhaden do (p. 114). And copepods are so plentiful in the Gulf of Maine, and the vegetable plankton that swarms in April has so largely disappeared over most of the Gulf before the mackerel appear later in the spring, that we doubt if they are ever reduced to a vegetable diet there or anywhere in American waters. Mackerel are also known to feed on bottom animals to a small extent. Nilsson, for example, reports various worms and hydroids and even small stones from their stomachs, but our expe- rience in the Gulf of Maine is to the effect that this would be exceptional there, if it happens at all. ' Nilsson (Publ. de Circ, Conseil Perm. Internat. Eiplor. Mer, No. 69. 1914) gives a similar list for Swedish waters 210941—03 22 Most authors describe the mackerel as feeding by two methods: either by filtering out the smaller pelagic organisms from the water by their gill rakers 5 or by selecting the individual animals by sight. A good deal of discussion has centered about the relative serviceability of these two methods of feeding. Probably the truth is that when forced to subsist on the smaller objects in its dietary it must do so by sifting them out of the water, but that it selects the more desirable when- ever opportunity offers to exercise its sight. It is not yet known how small objects the fish is able to pick out. It takes fish individually of course, and such large Crustacea as euphausiid shrimps and amphipods, just as the herring does, which evi- dently applies to the larger copepods, to judge from the fact that mackerel stomachs are often full of Calanus or of one or two other sorts in localities where indiscriminate feeding would yield them a variety. Whether they select the smaller copepods and crustacean larvae is not so clear. Captain Damant,6 whose experience in deep-sea diving has given him an exceptional opportunity to observe mackerel feeding under natural condi- tions, describes fish among which he was at work 20 to 40 feet deep in Lough Swilly (Ireland), as "feeding on plankton, not by steadily pumping the water through the gill filters but snatching gulps from different directions and making little jumps here and there." It has been a commonplace from the earliest days of the mackerel fishery that the fish are fat when last seen in the autumn, but that most of them are thin when they reappear in spring, obviously suggesting that they feed little during the winter. This is corroborated by the fact that the mackerel taken on bottom by British and French trawlers between December and March usually are empty, and that a few mackerel taken by the Albatross II along the continental edge off Chesapeake Bay in February 1931 were very emaciated. But mackerel taken in winter some- times have food in their stomachs; some of them even are fat.7 • The mackeral has long rakers with spines on the foremost gill arch only, and these are not fine enough to retain the smallest organisms. See Bigelow, Bull. U. S. Bur. Fish, vol. 40, Pt. 2, 1926, fig. 42 C, D for photographs of the gill rakers. • Nature, vol. 10S, 1921, pp. 12-13. ' Sette (Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Bui'. 49, 1950, pp. 259, 202) reports some fat mackerel in winter. 322 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE The immature fish feed and fatten from the time they appear in spring. And it also seems that the schools of older fish destined to spawn late in the season feed until the actual ripening of their sexual products commences, for large catches of the maturing fish were regularly made on hook and line in June in the Gulf of St. Lawrence where spawning takes place in July (until the eggs began to run, in fact). But these large mackerel would not bite after that until they had spawned out (last half of July or first part of August). And available evidence, American as well as European, is to the effect that fish destined to spawn soon after their vernal appearance inshore continue their winter fast until they have spawned, when they commence feeding greedily. Enemies. — The mackerel falls easy prey to all the larger predaceous sea animals. Whales, porpoises, mackerel sharks, threshers, dogfish, tuna, bonito, bluefish, and striped bass take heavy toll in particular. Cod often eat small mackerel; squid destroy great numbers of young fish less than 4 or 5 inches long, and sea birds of various kinds follow and prey upon the schools when these are at the surface. A considerable list of parasitic worms, both round and trematode, are known to infest the digestive tract of mackerel. But they seem more immune to danger from sudden un- favorable changes in their environment than the herring are, for they are never known to be killed by cold, and they seldom strand, except when small ones are driven ashore by larger fish. Breeding. — Mackerel spawn off the American coast from the latitude of Cape Hatteras to the southern side of the Gulf of St. Lawrence. The spawning area covers almost the entire breadth of the continental shelf southward from Cape Cod, but it is confined more closely to the vicinity of the coast thence northward. Available data point to the oceanic bight between Chesapeake Bay and southern New England as the most productive area, the Gulf of St. Lawrence as con- siderably less so, and the Gulf of Maine and coast of outer Nova Scotia as ranking third.8 Mackerel do not resort to any particular breeding grounds, but shed their eggs wherever their wandering habits have chanced to lead them when the sexual products ripen. It follows from this, and from the > See Sette (Fish. Bull. U. S. Fish and Wildlife Service, vol. 50, Bull. 38, 1943, pp. 158-164, and especially fig. 3) for discussion of spawning seasons and temperatures, and the relative importance of different spawning areas. fact that mackerel vary so widely in abundance over periods of years that the precise localities of greatest egg production may be expected to vary from year to year, depending on the local concen- trations of the fish. The mackerel spawns in spring and early summer. As it does not commence to do so until the water has warmed to about 46° F. (8° C), with the chief production of eggs taking place in temperatures of, say, 48° to 57°, the spawning season is progressively later, following the coast from south to north. Thus the chief production takes place as early as mid-April oft' Chesapeake Bay; during May off New Jersey; in June off southern Massachusetts and in the region of Massachusetts Bay; through June off outer Nova Scotia; and from late June through early July in the southern side of the Gulf of St. Lawrence, where eggs have been taken from early June to mid-August.9 Mackerel have never been found spawning in autumn, so far as we can learn, though a considerable number of eggs that we towed in Massachusetts Bay early in November of 1916 resembled mackerel eggs from the hatchery so closely that we would not have hesitated to identify them as such, bad they been taken in summer. They may have been the product of a belated fish, but more likely of some other Scom- broid. The mackerel is a moderately prolific fish; females of medium size may produce as many as 400,000 to 500,000 eggs in the aggregate, according to various estimates,10 with 546,000 reported for one weighing 1% pounds. But it is seldom that as many as 50,000 are set free at any one time, and often many fewer, for the members of a given school spawn over a considerable period. And recent observations u have shown that our earlier statement that they spawn chiefly at night was not correct. The eggs are 0.97 to 1.38 mm. in diameter, with one large oil globule,12 and drift suspended in the water, chiefly shoaler than the 5-fathom level. The rate of development is governed by the • See Sette (Fish. Bull. U. S. Fish and Wildlife Service, vol. 50, Bull. 38, 1943, pp. 158-163) for a more detailed statement. » Brice, Manual of Fish Culture, 1898, p. 212; Moore, Rept. U. S. Coram. Fish. (1898) 1S99, p. 5; Bigelow and Welsh, Bull. U. S. Bur. Fish., vol. 40, Pt. 1, 1925, p. 208. " Sette, Fish. Bull. U. S. Fish and Wildlife Service, vol. 50, Bull. 38, 1943, p. 165. " A series of Gulf of Maine eggs measured by Welsh were about 1.1-1 .2 mm. in diameter, with an oil globule of 0.3 mm. FISHES OF THE GULF OF MAINE 323 temperature of the water. Recent experiments by Worley 13 (which corroborate early hatchery experience) have shown that incubation takes about 150 hours at 54°; 115-95 hours at 57°-61°; about 70 hours at 64°-65°; and about 50 hours at 70°; with normal development limited to temperatures between about 52° (11° C.) and 70° (21° C). Newly hatched living larvae are 3.1 to 3.3 mm. long u with large yolk sac, and with numerous black pigment cells scattered over head, trunk, and od globule which give them a characteristic appearance. The yolk is absorbed and the mouth formed, the teeth are visible, and the first traces of the caudal fin rays have formed by the time the larva is about 6 mm. long. The rays of the second dorsal and anal fins and of the ventrals appear at about 9 mm. (to end of caudal fin); the first dorsal when the total length of the larva is about 14 to 15 mm. The dorsal and anal finlets are distinguishable as such in fry of 22 mm., and the tail fin has begun to assume its lunate shape, but the head and eyes still are much larger than in the adult, the nose blunter, and the teeth longer. At 50 mm. the little mackerel resemble their parents so closely that their identity is evident. Rate of growth. — The sizes of the. mackerel fry taken during the mackerel survey carried out by the U. S. Bureau of Fisheries in 1932,15 added to other available evidence show that our mackerel grow to a length of about 2 inches during the first 1 to 2 months alter they are hatched, a rate about the same as in British and Norwegian waters.16 This size is reached earlier or later in the season, depending on the date when any particular lot of fry was hatched. Thus mackerel fry of 1% to 2% inches obviously spawned that spring, have been taken at Woods Hole, both in the first half of June I7 and in the last 10 days of July,18 fry of 2% to 5 inches hi the first half of August, and fish of about 6K inches at the end of that month. Similarly, Captain Atwood found fry of 2 inches and shorter in July in the Massachusetts Bay " Jour. Oen. Physiol., vol. 16. 1933, pp. 841-857. 14 They shrink somewhat when preserved. " See Sette (Fish. Bull. U. S. Fish and Wildlife Service, vol. 50, Bull. 38. 1943, pp. 173-178) for detailed statistical analysis of these. " See Ehrenbaum (Rapp. et Proces Verb., Conseil Perm. Internat. Explor. Mer. vol. 30, 1923, pp. 21, 25) for a discussion of the early growth rate of the European mackerel. " Bigelow and Welsh, Bull. U. S. Bur. Fish. vol. 40, 1925, p. 204. '• Sette, Bull. U. S. Fish and Wildlife Service, vol. 50, Fish. Bull. 38, 1943, p. 178, fig. 8. region, i. e., about a month after the local mackerel schools had spawned out. Fry of 3% to 4% inches (obviously of the same season's crop because too small for yearlings) have been taken at Gloucester in August, and Captain Atwood reports them as 6% to 7 inches long, near Provincetown by October. Many of these little fish, up to 7 or 8 inches long (now large enough to be caught in the fish traps and known as tacks or spikes) are caught along the western shores of the Gulf of Maine and along southern New Eng- land during the fall. And measurements of thousands of young mackerel from the Gulf and from southern New England, compiled by the U. S. Bureau of Fisheries, have shown that the fry of the year average 8 to 9 inches, or longer, by the end of their first autumn, before they leave the coast for the winter. But broods produced in different years may grow at different rates, probably depending on feeding conditions, as well as on the dates when they are hatched. Thus fry spawned in the spring of 1927 averaged 8% inches in November, but those spawned in 1928 averaged 9% inches then. Our mackerel run about 10 to 11 inches long in spring and early summer of their second year of growth (they are known now as tinkers) , which agrees closely with Stevens' 19 estimate for mack- erel of the English Channel, based on studies of scales and otoliths. They grow to about 12 to 13 inches by that autumn, or to 14 inches in years of especially rapid growth,20 and the year- lings usually are a little longer in the Gulf of Maine than at Woods Hole, and longer at Woods Hole than off Long Island, N. Y. It remains to be seen whether these differences are due to temperature, to the varying richness of the food supply, or perhaps to crowding. It is also a question for the future whether the differences persist into later life. The brood of 1923, which may perhaps be taken as typical, averaged almost 14% inches in then third autumn, about 15% inches in their fourth, about 15% inches in their fifth, about 16 inches in their sixth, 16% inches in their seventh, and about 16% inches in their eighth years. Thus the American mackerel, like the European, grows very slowly after its third "Jour. Marine Biol. Assoc. United Kingdom, vol. 30, No. 3, 1952, pp. 549-56S. ■ Fry spawned in 1927 averaged about 1334 inches but those spawned in 1928 averaged only about 12^ inches in tbeir second November according to Sette. 324 FISHERY BULLETIN OP THE FISH AND WILDLIFE SERVICE summer, although it is long lived. The two sexes grow about equally fast. Nilsson's studies 2l point to a slightly slower rate of growth for the North European mackerel. But American mackerel have been found to vary so widely in this respect that the reported differ- ence may have been only an accident of obser- vation. A few fish of both sexes may mature sexually in their second year; about % of the males and % of the females spawn in the third year; and prac- tically all of them do so in their fourth year, i. e., when three full years old.22 This coincides with the transition from fast growth to slow, as might be expected, the ripening of the sexual products being so great a strain that the adult fish do little more than recover before winter. Once a mackerel has matured sexually, it no doubt spawns yearly throughout life, as most other sea fishes do. Proportions of the sexes. — In American waters males have been described as predominating largely over females.23 But more recent obser- vations have shown that there are about as many of the one sex as of the other, as there are in Sweden also.24 General range. — Both sides of the North Atlantic ; Norway to Spain off the European coast25; from the northern side of the Gulf of St. Lawrence and Strait of Belle Isle 26 to Cape Lookout, N. C.27 off the American coast. Migrations, and occurrence in the Gulf of Maine. — The occurrence of the mackerel in the Gulf of Maine is closely bound up with the sea- sonal movements of the species as a whole, for this is a migratory fish wherever it occurs, appear- ing at the surface and near our coasts in spring, to vanish thence late in the autumn. The di- rections and extent of the journeys which it carries out have been the subject of much discussion ever " Publ. de Circ, No. 69, Cons. Perm. Internat. Explor. Mer, 1914. « Sette, Fish. Bull. U. S. Fish and Wildlife Service, vol. 50, Bull. 38, 1943, p. 156. a Smith, Report U. S. Comm. Fish (1900) 1901, p. 128. •* Nilsson, Pub. de Circ. No. 69, Cons. Perm. Internat. Explor. Mer, 1914. " There is a fairly constant racial difference between American and British mackerel (Garstang, Jour. Mar. Biol. Assoc. United Kingdon, New Sor., vol. 5, No. 3, 1898, pp. 235-295), the latter showing a larger number of trans- verse bars, being more often spotted between them, and more often having 6 dorsal flnlets instead of 5. !« Jeflers (Contr. Canad. Biol., N. Ser., vol. 7, No. 16 [ser. A, General No. 13J, p. 207) reports that several mackerel were caught in 1929 at Raleigh, on the Newfoundland coast of the Strait of Belle Isle, where nono had been seen In recent years. " Coles, Copeia, No. 151 , February 1926, pp. 105-106 records a three-quarter pound mackerel taken at Cape Lookout in February 1925. since the fishery first assumed importance, because of their intrinsic interest, because of their bearing on the prosecution of the fishery, and because this fish has been the subject of much international dispute. The point chiefly at issue has been whether the main bodies of mackerel merely sink when they leave the coast in autumn and move directly out to the nearest deep water, or whether they combine their offshore and onshore journeys with the extensive north and south migrations in which most fishermen have long believed.28 The great majority of the mackerel have with- drawn from the coast by the end of December, not only from the Gulf of St. Lawrence, but from the entire inshore belt as a whole, not to be seen there again until the following spring or early summer, and it is not yet known definitely where the bulk of them go, though the subject has been widely discussed. Mackerel, it is true, have been caught, and have been found in the stomachs of cod and pollock in January, February, and early March at various localities on and around the outer Nova Scotian banks westward from Sable Island Bank; on the southern and northwestern parts of Georges Bank; in the deeper water be- tween the latter and Nantucket Shoals; on Nan- tucket Shoals; and along the middle and outer parts of the continental shelf off southern New England, off New York, off New Jersey, off Dela- ware Bay, off Virginia, and off northern North Carolina. Most of these winter records have been along the 30-70 fathom contour zone, but some- times as shoal as 4-5 fathoms off Nova Scotia, and as shoal as about 10-20 fathoms (near Am- brose Lightship) off New York,29 as deep as 90 fathoms off Chesapeake Bay.30 Most of these winter records have been based on odd fish only, i. e., not enough to suggest the presence of any great concentration of mackerel.31 But there were enough of them off New York in January, February, and March of 1949 for com- mercial fisheries to bring in what Gordon 32 has •» The literature dealing with this subject is very extensive. See especially Goode, Collins, Earll, and Clark (Rept. U. S. Comm. Fish. [18S1] 1884, p. 91); Tracy (37th Annual Report, Rhode Island Commissioners of Inland Fish- eries, 1907, p. 43); and Sette (Fishery Bull. U. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, pp. 268-313) for the American mackerel. » Gordon, Marine Life, Occ. Pap., vol. 1, No. 8, March, 1950, p. 39. » Sette (Fish. Bull. U. S. Fish and Wildlife Service, vol. 60, Bull. 49, 1950, pp. 260-261, table 1) lists several such instances besides those cited previously by Bigelow and Welsh (Bull. U. S. Bur. Fish. vol. 40, Pt. 1, 1925, p. 196). >> Three hundred pounds seems to be the largest winter catch definitely reported up to 1951. Ji Marine Life, vol. 1, No. 8, 1950, p. 39. FISHES OF THE GULF OF MAINE 325 characterized as "huge amounts." He also reports "a large body of fish" off Montauk in mid- February of 1950. Schools of "mackerel" have also been reported as sighted at the surface on several occasions in winter, but none of these seem to have been brought in. Direct evidence carries us only this far. But the indirect evidence of temperature is suggestive. Thus, the Gulf of St. Lawrence (where ice some- times forms), outer Nova Scotian waters, and the upper 50 fathoms or so within the Gulf of Maine which chill to 35°-39° F. (2°-4° C.) or colder, are all too cold by late winter for mackerel, which are never encountered in commercial quantities in temperatures lower than about 45° F. (7° C.). In most years this applies equally to the inner part of the continental shelf as a whole, southward as far as northern Virginia, for the water usually cools there to 37°-40° F. (3°-4° C.) at the time of the winter minimum. But the mackerel need only move out to the so-called warm zone at the outer edge of the shelf to find a more suitable environ- ment, for the bottom water there is warmer than 44°-46° F. (7°-8° C.) the year round as far north and east as the central part of Georges Bank, and about 41° F. (5° C.) along outer Nova Scotia. Available evidence thus supports Sette's 33 con- clusion that the bulk of the American mackerel winter on the outer edge of the continental shelf from the offing of northern North Carolina to the mid-length of Georges Bank, 30 to 100 miles off shore according to location, in depths of perhaps 50 to 100 fathoms. The few that are caught closer to land and in shoaler water in winter either represent the inshore fringe of the main population, or they are strays. Perhaps some winter off Nova Scotia as far east as Sable Island Bank. And it would not be astonishing should it prove that some winter in the deep eastern trough of the Gulf of Maine, where the temperature of the bottom water, at depths greater than 75 fathoms or so, does not fall below about 41° F. (5° C.). A few mackerel have, in fact, been caught on cod lines in deep water off Grand Manan in winter,34 while two were found among kelp near Yarmouth, Nova Scotia, on December 28, in 1878.35 » Fish. Bull. 49, U. S. Fish and Wildlife Service, vol. 51, Fish. Bull. 49, 1950, p. 261. « Collins, Kept. U. S. Comm. Fish (1882) 1883, p. 273. « Ooode, Collins, Earll, and Clark, Rept. U. S. Comm. Fish. (1881) 1884, p. 98; cited from the Yarmouth, Nova Scotia, Herald, January 2, 1879. Sette 38 has pointed out, however, that some other factor besides temperature must have to do with the wintering habits of the mackerel, for they disappear as completely from the surface and from inshore in the southern part of their range as they do in the northern even in very warm years such as 1932, when the water (surface to bottom) was warmer than 45°-46° F. (7°-8° C.), from New Jersey southward, even at the end of the winter. On the other hand, the event (probably ab- normally low temperature) that was so destructive to the tilefish in March, 1882 (p. 429), did not affect such of the mackerel as were wintering on the fcilefish grounds, for they reappeared that summer in normal numbers, a point to which Sette 37 has called attention already. Two additional facts which support the view that our mackerel do not travel very far in winter are (a) no mackerel, young or old, have ever been taken outside the edge of the continent, or any- where on the high seas far from land for that matter; (6) their reappearance in spring takes place so nearly simultaneously along some hundreds of miles of coastline that they can hardly have come from any great distance. Thus time and increased knowledge have corro- borated the view of Captain Atwood and of Perley, of more than half a century ago that mackerel winter offshore in deep water and northward from the latitude of Virginia, not in the far south nor out in the surface waters of the warm parts of the Atlantic. The winter home of the American mackerel appears to correspond rather closely to that of the mackerel of British seas, some of which winter on the deep northern slope of the North Sea, some in the deeper parts of the English Channel, and many on the outer edge of the continental shelf southwest of Ireland, mostly deeper than 60 fathoms.38 The failure of the otter trawlers to take com- mercial quantities of mackerel off Chesapeake Bay in winter when they fish there intensively, leads Sette39 to conclude that our mackerel » Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Fish. Bull. 49, 1950, p. 527. « Sette, Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, p. 257, Footnote 3. " Ehrenbaum (Rapp. et Proc.-Verb. Cons. Perm. Internat. Explor. Mer, vol. 18, 1914) summarizes what was known of the life history of the European mackerel up to that time. And Steven (Jour. Marine Biol. Assoc. United Kingdom, vol. 27, 1948, pp. 517-539) has recently outlined the chief winter- ing grounds. » Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, p. 261. 326 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE winter in the mid-depths, not concentrated on the bottom. This, however, would imply that the wintering mackerel manage to hold position for two or three months in some way without drifting far with the movements of the water. Another possibility is that they do keep on bottom, or near it, but somewhat deeper down the contin- ental slope than the trawlers ordinarily fish,40 perhaps concentrated in the many gullies, large and small, with which the upper part of the slope is seamed all along from the offing of Chesapeake Bay to Georges Bank, much as the mackerel of the Celtic Sea and English Channel winter "on the sea floor, densely packed in places where its level is interrupted by banks and gullies."41 Whichever of these alternatives is the correct one, the oft repeated assertion that the adipose eyelids of the mackerel become opaque in winter has no foundation. And they certainly do not hibernate in thousands along the coasts of Green- land and Hudson Bay,42 and of Newfoundland, with heads in the mud and tails protruding as a vice admiral, no less, has described them; a wholly imaginary tale, we need hardly add.43 They may winter in a more or less sluggish state. But the presence of food in the stomachs of some of the winter-caught fish, added to the fact that some of them are fat though others are thin, shows that they move about more or less even then, and feed more or less.44 Most American students have looked on the vernal warming of the surface water to about 45° F. as the stimulus causing the mackerel to quit their winter quarters. European studies, however, have shown that the date of their re- appearance in spring is not closely associated with any particular temperature. And if the mackerel winter on bottom along the edge of the continent, vernal changes in the temperature of the surface water nearer to land would be wholly outside their ken. The European mackerel usually keep to the bottom on their spring migration until close in to the land before rising to the surface. But this <° The southern trawl fishery is mostly shoaler than 70 fathoms. « Steven, Jour. Marine Biol. Assoc. United Kingdom, vol. 27, 1948, p. 537. « Mackerel do not range that far north. « Cited from Lacepede, Hist. Nat. Poissons, vol. 3, in Buffon, Hist. Nat- urelle, 1802, p. 32. ** Ehrenbaum (Rapp. et Proces Verbaux, Cons. Perm. Internat. Explor. Mer, vol. 18, 1914, p. 13), whose studies of the fish entitle his view to great weight, thinks that the mackerel of northern Europe probably are torpid during part of their stay on the bottom. generalization does not apply to the American fish, for while some may swim deep (so, only can we account for the fact that the first schools often show as early in Massachusetts Bay as on Georges Bank or off Nantucket) mackerel in great num- bers are first sighted 30 to 50 miles offshore, and this all the way from the latitude of Cape Hatteras to the mouth of the Gulf of Maine. The first mackerel "show" off the Cape Hatteras region at any time between about March 20 and April 25, usually early in April, and by the middle of April off Delaware Bay. As the water warms they spread northward and shoreward, being joined, it seems, by additional contingents from offshore. They reach the offing of southern New England some time in May, and they are plentiful on Nantucket Shoals by the first week of that month, as a rule. The date when they are first sighted off Cape Cod in the southwestern part of the Gulf of Maine varies from the last of April or first of May (April 29 in 1901, May 2, in 1898) to the first of June, with May 10 about the average. The earliest dates of commercial catches, for example, made in one particular set of traps near Provincetown have varied between May 14 and June 19. And the fish are plentiful in the western side of the Gulf of Maine as a whole by the end of the first week in June at the latest, if it is fated to be a good mackerel year. Mackerel (usually in smaller numbers) also appear on the Nova Scotian side of the Gulf about as early as they do in its western side; thus they were reported almost simultaneously off Yarmouth, Nova Scotia, and off Chatham on Cape Cod in 1898; in 1922 they were sighted off Yarmouth on May 7th, and off Cape Sable on the 11th. And they may appear even earlier in the season at Cape Breton, and as early well within the Gulf of St. Lawrence and in the eastern side of our Gulf. In 1894, for example, mackerel were first reported off Cape Breton on May 5 and at Gaspe on May 12, but not until May 16 45 at Yarmouth on the Gulf of Maine coast of Nova Scotia. But few of them show along the coast of Maine or in the Bay of Fundy until toward the end of June. Sette 4e has made the very interesting discovery that two distinct populations are represented among the American mackerel, a southern and a <» Huntsman, Canadian Fisherman, vol. 9, no. 5, 1922, pp. 88-89. « Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950. FISHES OF THE GULF OF MAINE 327 northern, with rather different migratory habits, and differing also in the relative success of repro- duction in different years.47 The nature of these two contingents is not known, whether genetic or environmental. It is probable (though not proved) that the southern contingent tend to winter in the southern part of the wintering zone. The main bodies of mackerel that appear in spring along the middle Atlantic coast belong to this contingent, also most of those taken off southern New England. They summer for the most part over Nantucket Shoals; on the western part of Georges Bank; and in the western and northwestern parts of the Gulf of Maine, which they enter in the western side around Cape Cod. And they do not journey farther east than the coast of Maine. On the other hand, it seems the mackerel that appear early in the season along the Nova Scotian shore of the Gulf, to spread later to Maine, belong to the northern contingent, and also a scattering of those that enter the western side of the Gulf. These appear to winter mostly eastward from the Hudson Gorge, and their vernal migration carries most of them past our Gulf, to pass the summer along outer Nova Scotia, and in the southern side of the Gulf of St. Lawrence.48 A few mackerel (mostly small) from the southern contingent remain all summer in the coastwise belt from Long Island to Nantucket. Apart from these, however, the whole body of American mackerel have deserted the southern grounds altogether by the early summer, to spend the later summer either in the region of our Gulf, off Nova Scotia, or in the Gulf of St. Lawrence.49 If the view now held is correct as to their migratory routes, some of the mackerel that sum- mer in our Gulf may come from as far as the offing of North Carolina; others from as nearby as the offing of New York or of southern New England. The vernal journey of the Gulf of St. Lawrence *7 This conclusion, seemingly conclusive, is based on analysis of the size (i.e. age) composition of the mackerel population at various times and places, with some evidence from tagging experiments. The data are too extensive for discussion here. *> For further information as to migrations of the northern contingent, see Sette, Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, pp. 269, 285. <• We have found no positive record of mackerel takon In late summer anywhere south of Delaware Bay, although they are plentiful off this part of the coast in spring. Bell and Nichols, it is true, speak of "mackerel" as found in tiger-shark stomachs off North Carolina (Copeia, No. 92, 1921, pp. 18-19), but Dr. Nichols writes us that these were "Just Scombroids and probably not Scomber scombrtis." mackerel may be anywhere between, say 300 to 350 miles, and 700 miles, depending on whether they have wintered off outer Nova Scotia or as far west as the western slope of Georges Bank. It seems certain that some of the mackerel that are first sighted on Nantucket Shoals and on Georges Bank in May remain on these offshore grounds all summer, both spawning and feeding there, for they provide good fishing there any time from June to September or October, in some years. The farther advance of such of them as continue northward into the Gulf of Maine covers a period of some weeks, with the first-comers followed by other schools later. And it seems certain (as just remarked) that fish resorting to our Gulf, do so summer after summer, never visiting the outer coast of Nova Scotia, much less a region as far afield as the Gulf of St. Lawrence. But it is an interesting question for the future, whether a given school returns to the same part of the Gulf, year after year. Many of the mackerel that summer in our Gulf have already spawned farther south (p. 322). Others, however, are still hard, but they are soon taken there with eggs or milt running. Spawning iti the Gulf of Maine is at its peak in June in most years, with the proportion of spent fish increasing through July, and only an occasional ripe fish as late as the first of August. But a year comes occa- sionally, such as 1882, when spawning is not at its height in the Gulf until July, with ripe fish con- tinuing plentiful until August. And our towings there have yielded a few mackerel eggs as early as May 6, as late as September l.60 The spawning season is at its height in the Gulf of St. Lawrence during the last half of June and the first half of July, continuing into August, a fact well known by the hook-and-line fishermen of half a century ago, because the ripe fish will not bite at that time, and more recently corroborated by the egg catches of the Canadian Fisheries Expedition.61 It seems from the relative numbers of eggs taken from place to place, that Cape Cod Bay is the only subdivision of our Gulf that has rivaled the more southern spawning grounds in egg production dur- ing the particular years when intensive studies » See Blgelow and Welsh (Bull. U. S. Bur. Fish., vol. 40, Pt. 1, 1925, p. 206), for details. •' Dannevig. Canadian Fish. Exped. (1914-1915); 1919, p. 8. 328 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE have been made.62 Mackerel also spawn to some extent thence northward, as far as Casco Bay, but we believe very few do so farther east than that along the coast of Maine. Neither is it likely that mackerel breed successfully in the northern side of the Bay of Fundy for neither eggs nor larvae have been taken there though some production may take place on the Nova Scotian side for Huntsman reports eggs at the mouth of the An- napolis River. And while a moderate amount of spawning takes place along the outer coast of Nova Scotia,63 it seems that the eggs do not hatch in the low temperatures prevailing there, for no larvae have been found. But the southern side of the Gulf of St. Lawrence, where the surface waters warm to a high temperature in summer, is an extremely productive spawning ground (p. 322). Since the large adult mackerel tend to keep farther offshore than the small ones (p. 328), such of them as spawn in our Gulf do so at least a few miles out. Very few eggs, for example, were found in 1897 (a year of plenty) in the inshore parts of Casco Bay,64 though this was formerly thought to be a productive spawning ground. Once the mackerel have entered our Gulf, schools are to be expected anywhere around its coastal belt, at any time during the summer; also on Nantucket Shoals, on the western part of Georges Bank, and on Browns Bank, as just noted (p. 327). And while adult fish seldom venture within the outer islands or headlands, good catches of them have been made well up Penobscot Bay, and young ones 6 to 10 inches long often swarm right up to the docks in various harbors in summers of plenty.88 Mackerel are proverbially unpredictable in their appearances and disappearances at any particular place, hence the common saying that "mackerel are where and when you find them." This is partly because the schools are constantly on the move, but partly because it is only while they are school- ing at the surface or near it that they are seen. " Subsequent information, and especially the result of tow nettings on the southern grounds in 1929, 1930, 1931, and 1932 (Sette, Fish. Bull. U. S. Fish and Wildlife Service, vol. 50, Bull. 38, 1943) have shown that the Gulf of Maine as a whole is much less productive than the more southern spawning grounds, not more so as Bigelow and Welsh (Bull. U. S. Bur. Fish., vol. 4, Ft. 1, 1925, p. 206) believed. " Sparks, Contrib. Canadian Biol, and Fish., N. Ser., vol. 4, No. 28, 1929. '< Moore, Rept. U. S. Comm. Fish. (1898) 1899, p. 16. « Sette (Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, p. 297) discusses this point further. When they sink to lower levels in the water, as they often do, they drop out of sight entirely, unless some of them chance to be picked up by drift netters. Large mackerel are more prone to disappear in this way than small ones, especially in late summer or early autumn. In 1906, for example, the schools of large fish vanished from the Massachusetts Bay region in June, to reappear the 27th of July, on which date 28 seiners made catches ranging from 18 to 250 barrels each. And in 1S92, a year of abundance, they disappeared (that is, sank) in August, not to appear again in any abundance anywhere in the Gulf of Maine until October. The view has grown that when this happens the mackerel have deserted the Gulf for the time being. But it was common knowledge in the days before the introduction of the purse seine, when it was the regular practice to lure the fish to the surface by throwing out ground bait, that large mackerel summer as regularly in the Gulf as small, and that good hook-and-line catches of large fish could be made in one or another part of the Gulf through the season from June to October, even when none showed at the surface. Their disappearances in summer merely mean that the fish have sought lower levels in the water; that they have wandered to some other part of the Gulf; or perhaps that the schools have dispersed more or less. When they sink in summer in our Gulf, it is not likely that they descend very deep. In the first place the water deeper than about 40 to 50 fathoms is colder than 46°-47° F. (8° C), i. e., than they seem to prefer; in the second place the planktonic animals on which they feed are more concentrated above the 50-fathom level than deeper. And a year comes, now and then, when mackerel of all sizes school at the surface all summer long.66 Sette's 67 painstaking analysis of the relative frequency with which schools are seined in different localities has shown that mackerel are seen far the most often in the southwestern part of the Gulf and out along the western part of Georges Bank, with the chief concentrations in one part or another of Massachusetts Bay and off the outer shore of Cape Cod to Nantucket Shoals, though great numbers are also caught along the Maine coast, close inshore. '• 1882 was an example of this. " Fish. Bull. U .S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, p. 297, fig. 17. FISHES OF THE GULF OF MAINE 329 Mackerel contrast in an interesting way with herring in this respect, the latter being caught in by far the greatest numbers in the northeastern corner of the Gulf, i. e., just where there usually are fewest mackerel. But there is much variation from year to year in their relative abundance from place to place as appears from the following table of catches, made in two successive years when the total landings from the Gulf, as a whole, did not differ greatly (landings at Boston, Gloucester, and Portland by the vessel fishery, stated in pounds). Fishing grounds 1916 1917 Georges Bank 3,701,597 624,086 South Channel 77,157 13,600 Nantucket Shoals 2,516,414 6,277,830 Off Chatham 2,017,753 3,938,452 Off Race Point 99,250 621,751 Stellwagen Bank 1,559,972 519,550 In some years few mackerel are seen at the sur- face in the Gulf eastward of the Isles of Shoals, 1926, 1927, 1933, 1934, and 1935 were examples. In other years, however (e. g., in 1928, 1929, 1930, 1931, and 1932) many schools are sighted and seined along the coast of Alaine as far eastward as the vicinity of Mount Desert Rock (see fig. 176, based on Sette's painstaking analysis). But the experiences of the old time hook-and-line fishermen suggest that the mackerel tend to move north- ward and eastward in general from the Massachu- setts Bay region, for they made their best late- summer and early-fall catches between Cape Eliz- abeth and Mount Desert Rock in most years, notably about Monhegan Island. And the results of hook-and-line fishing are a far better clue to the presence or absence of mackerel than the seine catches are, since they draw from the fish that are deep down, as well as from those that may chance to be at the surface. The Nova Scotian side of the Bay of Fundy has been a profitable mackerel ground, occasionally, but only for short periods and at long intervals. Thus good catches were made there for some years previous to 1876, but this fishery was abandoned a few years later for want of mackerel. There were enough fish there again in the early 1900's to yield about 7 million pounds in the 6-year pe- riod 1901 to 1906.68 But we have not heard of any large catches made anywhere in the Bay of Fundy since that time, so events of the sort must be out of the ordinary. And very few mackerel are ever « Sctte and Needier, Inv. Rept. 19, U. S. Bur. Fish., 1934, pp. 1-48. reported along the New Brunswick side of the Bay. In most years, mackerel are few over the central deeps of the Gulf (fig. 176), but a year comes now and then when they are plentiful there, as hap- pened in 1882 (a year of great abundance), when great numbers were caught between Georges Bank, Browns Bank, and Cashes Ledge, and thence northward to within 40 miles or so of the Maine coast. Most of the early season catch, in fact, was made in this deep water region that year, and in the weirs along the west coast of Nova Scotia. But the fish disappeared thence later in the season. And large catches have never been reported from the eastern part of Georges Bank to our knowledge. Figure 176. — Average distribution of mackerel in the Gulf of Maine, July through September, based on relative frequencies of catches recorded for each 10- mile rectangle, 1926 through 1935. After Sette. As a rule, the schools tend to stay nearer the coast in years when small (i. e., young) fish dom- inate the population. The entire Gulf of Maine catch, for example, was taken within 45 miles of 330 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE land in 1926, when the stock was dominated by fish hatched in 1923, i. e., were in their third year. In years of this sort, anglers fishing in harbors, or going out in charter boats for the day, do well, catching the smaller sizes chiefly. But in 1928, when the same year class dominated as had in 1926 (i. e., fish now in their fifth year), only about two-thirds of the catch was made that close in, with about one-third of the catch taken more than 45 miles out at sea. Nineteen twenty-nine may serve as another example, with more than one- half (57 percent) of the large fish caught more than 45 miles out, but less than 1 percent of the small ones, and a few large ones, taken as far out as 80 miles. But even the fully grown fish do sometimes come close inshore; we have ourselves caught mackerel within a few yards of the beach in the southern side of Massachusetts Bay, as large as any that we have seen taken anywhere. Fishermen have long realized that mackerel are most likely to be found where there is a good supply of "red feed" (copepods) or other small animal life in the water. A relationship has, in fact, been found to hold in the English Channel between the catches of mackerel and the numbers of copepods present.69 And while no attempt has been made yet to relate the local abundance of mackerel in our Gulf, or the depths at which they swim with the supply of food on a statistical basis, the mere fact that they do fatten in our waters is evidence enough that they manage in some way to congregate where food is plentiful. But it appears that their vernal journey, from their wintering grounds to the Gulf and to Nova Scotian waters, is directed by some impulse to migration more definite than the mere search for food. Thus while a large proportion of the mack- erel did travel along the zone of abundant plank- ton in the only year (1930) when their advance along the coast has been compared with the quantitative distribution of the animals on which they prey,60 they deserted the waters south of New England that year whde the food still was abund- ant there, for regions (Gulf of Maine and eastward) where there is no reason to suppose that feeding conditions were any better at the time.61 M Bullen, Jour. Marine Biol. Assoc. United Kingdom, vol. 8, 1908, pp. 269, 302. « Bigelow and Sears, Mem. Mus. Comp. Zool., vol. 54, No. 4, 1939. pp. 259-261. « See Sette (Fish. Bull. U. S. Fish and Wildlife Service, vol. 51, Bull. 49, 1950, p. 302) for a further discussion of the influence of feeding conditions on the movements of the American mackerel. As autumn draws on, the fish that summer along the Maine coast (chiefly belonging to the southern contingent) seem to work back southwestward toward Cape Cod, for catches were made suc- cessively off Portland, near Boon Island, and off Cape Ann, in the days when mackerel were caught on hook and line. It is probable, too, that such of the fish from the northern contingent as have entered the Gulf in the eastern side join in this general autumnal movement around the coast to the westward and southward, rather than that they leave by the route along which they enter, for schools have often been reported, and actually followed, swimming southward at the surface across Massachusetts Bay. And while reports of this sort are likely to be based on misconception,62 they are corroborated in this instance by the fact that the latest catches are always made either in or off Massachusetts Bay, along the outer shore of Cape Cod, or on the neigh- boring parts of Nantucket Shoals, never either on Georges Bank, which would be on the direct route of any fish swimming westward from Nova Scotia, or in the inner parts of the Gulf of Maine. Sette's studies indicate that the bulk, at least, of the mackerel of the southern contingent have moved out of the Gulf around Cape Cod and past Nantucket Shoals by late September or October in most years. But many of the fish of the northern contingent coming from Nova Scotia, and perhaps even from the Gulf of St. Lawrence, usually provide good fishing off Cape Ann and southward through October and late into Novem- ber,63 with good commercial catches until mid- December in some years. In 1913, for example, 1,200 mackerel were caught off Gloucester on December 10; 3,000 off Chatham, Cape Cod, a day or two earlier; and nearly 1,000 barrels (200,000 pounds) were seined off the Massachusetts coast during the early part of that month in 1922. In mild winters schools of mackerel are sometimes reported and even caught off the outer coast of Nova Scotia as late as Christmas time; i. e., somewhat later than off Cape Cod. But the M The successive approach of one school after another to the coast often sug- gests a long-shore movement of the fish. Thus Kendall (Bull. U. S. Bur. Fish., vol. 28, Pt. 1, 1910, p. 287) tells of an instance when seiners reported "following" the schools continuously eastward along outer Nova Scotia, although the fish taken ofl Liverpool proved to be of quite different sizes from the catch made later about Cape Breton. - In 1922, for example (Gloucester Times of April 26, 1923), mackerel netters fishing near Cape Ann did well right through November, with a catch of about 1,200,000 pounds (6,000 barrels) for the month. FISHES OF THE GULF OF MAINE 331 whole body of Gulf of Maine, Nova Scotian, and Gulf of St. Lawrence mackerel have withdrawn thence by the end of December at the latest, except for odd stragglers. And when they do de- part, they must sink at once to lower levels in the water, for schools are never sighted on their jour- ney offshore and southward; they simply drop out of sight. Abundance. — It has been common knowledge since early colonial days that mackerel fluctuate widely in abundance in our Gulf from year to year, perhaps more widely than any of our other im- portant food fishes, with periods of great abun- dance alternating with terms of scarcity, or of almost total absence. In good years the fish may appear in almost unbelievable numbers; schools or associations of schools, miles in length, are re- ported; and it is common to see 50 or more sep- arate bodies of fish from the masthead at one time. Mackerel, in short, seem to be everywhere, and a tremendous catch is made. But perhaps only an odd school will be seen here and there the next year, and the fishery will be a flat failure. The period from 1825 to 1835 was one of abun- dance. In 1831, for example, more than 380 thousand barrels (76 million pounds) of salt mack- erel (in those days most of them were salted) were landed in Massachusetts ports. But mackerel were scarce for the next 8 years (1837-^5), only 50,000 barrels being landed in Massachusetts in 1840. The Massachusetts catch then fluctuated violently from 1851, when the landings rose once more to 348,000 barrels, down to 1879. The fleet brought in something like 294 million fish from Nova Scotian and United States waters combined in 1880. And this introduced a period of extraor- dinary abundance, culminating in 1885 when the catch reached the enormous total of 500,000 barrels (100,000,000 pounds). But this was followed in its turn by a decline so extreme, so widespread, so calamitous to the fishing interests, and so long continued, that the catch was only about 3,400 barrels (equivalent to 582,800 pounds of fresh fish) for the entire coast of the United States in 1910 (when the stock of mackerel fell to its lowest ebb) with almost none reported in Massachusetts Bay or along the Maine coast. Mackerel then increased again in numbers; slowly at first, then more rapidly, as appears from the fact that the catch for the Gulf of Maine and for the banks at its mouth was about four times as great in 1911 (about 2% million pounds) as it had been the year before, rising to about 4% million pounds in 1912, 5 million in 1913, 7% million in 1914, to something more than 11 million in 1915, and 16 to 16% million each for 1916 and 1917. But this period of multiplication fell far short of equaling the banner years of the 1880's. And the catches fell off again so rapidly after 1917 that the Gulf of Maine yield for 1919 64 was only about one-quarter as great as it had been in 1917. Although 1920 saw some slight recovery, 1921 (with a local catch of only about 1 million pounds) proved the worst mackerel season for our Gulf since 1910. The stock then built up enough (following the familiar seesaw pattern) for the Gulf to yield about 25 million pounds of mackerel in 1925. Since that time down to 1946, the Gulf of Maine catch has ranged between a low of about 20 million pounds (1937) and a high of about 59 million (1932). Thus the catch of mackerel in our Gulf may be 50 to 100 times as great in a good year as in a poor. The average Gulf of Maine catch for the period 1933-1946 was about 37 million pounds, yearly. Various far-fetched explanations have been proposed for these astounding ups and downs in the catches from year to year, such as that the fish have gone across to Europe; have sunk; or have been driven away or killed off by the use "! the purse seine. Actually, these changes re- lied the ups and downs in the numbers of the fish that arc in existence from year to year. Mack- erel, in short, were extremely plentiful in 1885, very scarce in 1910, moderately plentiful in 1916 and l'.i 1 7. very scarce again in 1921, and they have been moderately plentiful since about 1925, but probably not so plentiful as they were in the 1880's.05 It has long been known for the herring and for some other species that the prime factor in deter- mining the abundance of the fish is the comparative success of reproduction from year to year, years favorable to the production and survival of larvae presaging several seasons of abundance, or vice-versa. And comparison of the relative pro- portions of mackerel of different sizes (that is, of different ages) in the total catches from year to year has shown that this is equally true of the »' In 1919, 4,091,345 pounds. •' See especially, Sette, U. 8. Bur. Fish., Fishery Circular Xo. 4, 19.11. 332 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE mackerel.66 When there is an abundant crop of young mackerel, the fishing is good during the next several years, but the catches then fall off, if another good brood does not soon appear upon the scene. The course of events since the low point in 1910 may then be reconstructed about as follows: In 1910, when the stock of mackerel was at its lowest, most of the fish caught were large, sug- gesting that few young had survived for several years past. Unfortunately, no information is available as to the composition of the population from the point of view of size for the next three years, when the catch was progressively somewhat larger, but great numbers of small fish, (apparently yearlings), were reported in 1912, pointing to a good breeding season in 1910, in 1911, or in both. In 1914 fish smaller than 1% pounds again formed nearly 60 percent, by weight, of the catch in and off the Gulf of Maine, and approximately 80 per- cent in 1915, with an even greater preponderance in actual numbers between small (young) fish and large (old). These little fish, hatched dur- ing the period 1910 to 1912 or 1913, were respon- sible, as they grew, for the fairly good catches made in the Gulf in 1916 and 1917.67 But the produc- tion of fry must have been very poor in 1916 and 1917, for the Gulf of Maine catch was only about one-seventh as great in 1919 as it had been in 1916. And reproduction must have practically failed in 1918 or in 1919, for the mackerel caught in 1920 ran very large, both south of New York that spring, and in our Gulf during that summer and autumn. The population was now back again in about the same state as it had been in 1910, the cycle having run through a period of 10 years. The parallel goes further, too, for 1921 must have seen a wave of production to account for the swarms of small fish that appeared along the New England coast from Woods Hole to Mount Desert during the summer of 1922. This again presaged a great in- crease in the catches of mackerel for the next few years to come (more than 1 1 million pounds were taken in the Gulf of Maine in 1923). And Sette's studies show that 1923 was another productive year, resulting in a catch more than twice as great in 1925 as it had been in 1923, and about 3 times as great in 1926.68 The very large catch of about 59 million pounds in the Gulf in 1932 was pre- ceded similarly by the presence of great numbers of yearlings hi 1929, evidence of successful repro- duction in 1928. Thus, it seems that the proportion of fish of different ages in the catch in any one year may be used as a basis for predicting the success or failure of the run of mackerel for the next year; such predictions have in fact been attempted by Sette 69 with fair success. No record has been kept, so far as we know, of the relative numbers of mackerel of different ages, of late years. But a failure of reproduction, fol- lowed by a slump in the catch, may come at any time, for history has a way of repeating itself, especially where fishes are concerned. Nothing definite is known as to what determines the success or failure of reproduction of the mack- erel in any given year. Towings by the U. S. Bureau of Fisheries make it likely that the actual production of eggs is usually sufficient. But the vitality of the eggs spawned in any given year goes back to the physiological condition of the parents. And studies of the composition of the stock of fish in periods of high production and of low suggest that there is some correlation between the number of adult mackerel existing in the sea at any time, and the success with which they breed, for it seems that years when great numbers of fry survive always fall when the parent fish are scarce, average large, and also average very fat (by general report). One hypothesis is that the mackerel tend to grow fast when there are only a few of them and go into the winter in excellent condition, hence are able to produce eggs of high vitality and in abundance; but they do not fare so well indi- vidually when plentiful, hence, do not emerge from their winter quarters in as good physio- logical condition in spring, so that fertilization and incubation may be less successful, and such larvae as hatch may be less strong. On the other hand, all this may be insignificant as compared with the success or failure of the larvae in sur- viving the dangers and difficulties of subsistence that confront them. Onslaughts by enemies, M Sec especially Sette, U. S. Bur. Fish., Fishery Circular No. 4. 1931. " Gulf of Maine catch, 16,391 ,095 pounds in 1916; 16.021 ,619 pounds in 1917. •■ Gulf of Maine catch, 11,007,676 pounds in 1923; 25,475,S76 pounds in 1925; 33,152.766 pounds in 1926. « U. S. Bur. Fish., Fish. Circ, No. 4, 1931; No. 10, 1932; No. 14, 1933; No. 17, 1934; Fishing Gazette, vol. 50, No. 6, 1933, pp. 9 and 21. FISHES OF THE GULF OF MAINE 333 abundance and ready availability of food, tem- perature, and salinity of the sea water all act upon the young fish to make their existence pre- carious; a favorable environment depends on a happy combination of all these. Importance. — The mackerel is a delicious fish, but it does not keep so well as some other fishes that have less oil in their tissues. When mackerel are rather plentiful they are one of the four most valuable fishes of our Gulf commercially, sur- passed in dollar value only by the haddock, cod, and rosefish, as appears from the following table of landings in New England for the years 1943- 1947. When the fishery fails, as it does periodically tlirough periods of several years (p. 331), the value of the catch decreases accordingly, and to a point where it is only a negligible fraction of the total yield and value of the Gulf of Maine fishery. Species 1943 1944 $8,650,000 (1.1) 4,000,000 (1.2) 4,350,000 (1.1) 3,180,000 (1.1) $7,550,000 (1.2) Cod 3,500.000 (1.5) 4,300,000 (1.1) 2,400,000 (1.3) Species 1945 1946 1947 Haddock Cod $7,000,000 (1.4) 4,280.000 (2.3) $8,800,000 (1.2) 3.940.000 (1.4) $8,900,000 (1.3) 2,780,000 (1.4) 4.200,000 (l 4) 3,840,000 (1.3) 4,750,000 (1.7) 3,100,000 (1.3) 2.340.00(1 (1.1) 2,000,000 (1.2) Note.— The total value of the catch landed in Maine, New Hampshire, and Massachusetts, including fish from grounds outside the Gulf of Maine, as well as from within the Gulf, may be determined by multiplying the values by the figures in parentheses. Most of the mackerel were caught formerly with hook and line, ground bait being thrown out to lure the fish close enough to the vessel.70 But this way of fishing was gradually given up about 1870, when the use of the purse seine became general. And practically the entire catch of mackerel of the past 70 years has been made with purse seines, with pound nets, weirs and floating traps coming second, and gill nets a poor third. In 1943, for example, when the total Gulf of Maine catch was between 53 and 54 million pounds, about 80 percent was taken in purse seines; between 12 and 13 percent in pound nets, weirs, and floating traps; and between 3 and 4 percent (between 1 and 2 million pounds) in gill nets (anchored or drifting), but only 1,700 pounds on hand lines. Otter trawlers, too, '» See Qoode and Collins, Fish. Ind. U. S„ Sect. 5, vol. 1, 1887, pp. 275-294, for an excellent account of the hook and lino fishery. bring in scattering mackerel from the offshore banks: 2,400 pounds, for example, in the year in question. Many anglers, also, troll or bait-fish for mackerel all along the coast from Cape Cod to Penobscot Bay; as far as Mount Desert if mackerel are on the coast that far east. In good years it is not unusual for 3 or 4 anglers fishing from a party boat to bring in one or two hundred fish. And in summers when young tinkers are plentiful inshore many of them are caught from the wharves in various harbors. If one chooses to troll, an ordinary pickerel spinner, No. 3, serves well, especially if tipped with a small piece of pork rind or with mackerel skin; a small metal jig similarly adorned, or any small bright spoon. Mackerel will also take a bright artificial fly, and bite greedily on a white piece of clam, a piece of mackerel belly, or on a sea worm (Nereis) , especially if attracted by ground bait. Chub mackerel Pneumatophorus colias (Gmelin) 1789 71 Hardhead; Bullseye Jordan and Evermann, 189&-1900, p. 866, Scomber colias Gmelin. Description. — The hardhead (by which name it is commonly known to fishermen) resembles the common mackerel so closely that we need mention only the points of difference. Most important of these, anatomically, is the fact that the hardhead has a well-developed swim bladder connected with the esophagus, which the mackerel lacks. But it is not necessary to open the fish to identify it for there is a characteristic color difference between the two, the mackerel being silvery-sided below the mid line whereas the lower part of the sides of the hardhead (otherwise colored somewhat like the mackerel) are mottled with small dusky blotches, and the chub has a larger eye than the mackerel. Less obvious differences are that the dorsal fins are closer together in the chub and that there are only 9 or 10 spines in its first dorsal fin instead of 11 or more, which is the usual count in the mackerel. " This genus is separated from Scomber by having a well developed swim bladder which the true mackerel lacks (see Starks, Science, N. Ser., vol. 54 1921, p. 223). 334 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE : ■' --'-V ::'j.Ci-:'S : l'<-:.'M'<^^ Figure 177. — Chub mackerel (Pneumalophorus colias), Provincetown, Mass. From Gootle. Drawing by H. L. Todd. Size.- — This is a smaller fish than its better known relative, growing to a length of about 8 to 14 inches only. Habits. — Hardheads school like mackerel, and their feeding habits are much the same, for Doctor Kendall found fish on Georges Bank in August 1896, full of the same species of pelagic Crustacea and Sagittae that the mackerel had taken at the same time and place, while specimens taken at Woods Hole had dieted chiefly on cope- pods, to a less extent on amphipods, Salpae, appendicularians, and young herring. They fol- low thrown bait as readily and bite quite as greedily as mackerel do. Their breeding habits have not been studied. General range.- — Temperate Atlantic Ocean, north to outer Nova Scotia and to the Gulf of St. Lawrence in the west,72 to England in the east. It is represented in the Pacific by a close ally, Pneumatophorus japonicus. It is a more southerly fish than the mackerel. Occurrence in the Gulf of Maine. — Goode,73 long ago summarized the early history of the chub mackerel in our waters, which briefly was as follows: It was tremendously abundant during the last of the eighteenth century and early years of the nineteenth, down to 1820-1830. Thus Capt. E. E. Merchant, an experienced and observant fisherman, described them as so plentiful off Provincetown from 1812 to 1820 that three men and a boy coidd catch 3,000 in a day on hook and T; It is reported from St. Margaret Bay and Halifax by Vladykov (Troc. Nova Seotian Inst. Sci., vol. 19, 1935, p. 7), and Schmitt (Monographic de l'lfle de Anticosti, 1904, p. 285, Paris) credits it with "apparitions irregu- lieres" at Anticosti. » Fish. Ind., U. S., Sect. 1, 1884, p. 303. line. But it practically disappeared from the United States coast some time between 1840 and 1850. It is interesting to note, as Captain At- wood pointed out, that destructive methods of fishing had nothing to do with the case, for its disappearance antedated the introduction of traps, pounds, or purse seines; it also antedated the re- appearance of the bluefish (p. 386) ; hence cannot be blamed on these sea pirates. So completely did the hardheads vanish that the Smithsonian Institution tried in vain for 10 years prior to 1879 to obtain a single specimen. But a school was taken in the summer of 1879 in a trap at Province- town (where representatives of the U. S. Fish Commission were stationed at the time), and though none were seen in 18S0 there w_ere some off the coast of New York in 1886. We find no definite record of the status of the hardhead during the next decade. But Bean 7* describes them as abounding off New York in 1896, swimming up little creeks in such numbers that they could be dipped in boat loads. And hardheads were taken singly and in schools by the mackerel fleet on Georges Bank during that same August,76 while many were caught on hook and line from the Grampus in Block Island Sound during the first week of that September. Kendall found them at Monomoy, the southerly elbow of Cape Cod in 1898, and they were suffi- ciently restablished by then for Smith 76 to de- scribe them as uncommon to abundant at Woods Hole. They then dropped out of the published record again (they are not separated from the '< Bull. f.O, New York State Mus., Zool. 9, 1903, p. 383. »• Field Holes supplied by Dr. W. C. Kendall. '• Bull. U. S. Fish. Comm., vol. 17, 1898, p. 95. FISHES OF THE GULF OF MAINE 335 common mackerel in the fishery returns) until 1900, when they were found in the Casco Bay region. There is no reason to suppose that they appeared in any numbers anywhere on our coasts during the period 1900 to 1906, but in the latter year many were taken in the traps near Woods Hole, also in 1908. And the mackerel fleet found great schools of hardheads on Georges Bank in 1909, when vessels brought in fares of 50,000 to 100,000 of them during the first week of July," their small size (500 to 700 to the barrel) suggest- ing that there had been a great production of hard- heads a year or two previous. Fishermen speak of catching a few from time to time since then, but no great numbers. We caught one at Cohasset on the south shore of Massachusetts Bay in September 1942. In its years of plenty, which fall at long in- tervals, however, the chub mackerel is likely to appear wherever mackerel do off the Massa- chusetts coast, especially about Provincetown. Thus 13,420 pounds were taken in traps at North Truro, in 1952, between August 11 and October 5. Other definite Gulf of Maine records are mostly78 about Casco Bay and one from Johns Bay, Maine. We found no record of it farther east along the coast of Maine ; it is unknown in the Bay of Fundy, nor does it seem to reach the west Nova Scotian coast. But in good "hardhead" years, it is to be expected all along Georges Bank and on Browns as well, to judge from its occasional visits to the outer coast of Nova Scotia. " Boston Herald, July 9, 1919. " Scattergood, Trefethen, and Coffin (Copeia, 1951, No. 4, p. 298). report one caught in August 1949. Importance.- — The chub mackerel is as choice a table fish as the mackerel, and no distinction is made between them in the market, other than the size of the individual fish. Striped bonito Euthynnus pelamis (Linnaeus) 1758 Oceanic bonito Jordan and Evermann, 1896-1900, p. 868, Gymnosarda pelamis. Description. — The various fishes commonly called bonitos, albacores, and tuna, are fusiform in shape like all their family, tapering to a pointed nose and to an extremely slender caudal peduncle. But they are much stouter-bodied than mackerel or chub mackerel, and their second dorsal fin originates close to the rear end of the first dorsal, instead of being separated from the latter by a long interspace. The present species is about one-fourth as deep as it is long; its caudal peduncle has one prominent median longitudinal keel on either side, with a smaller keel above it, and another below at the base of the tail. The very deeply concave contour of its first dorsal fin (fig. 178) is enough to separate it at a glance from the common bonito (fig. 180), or from a young tuna (fig. 181), and from the Spanish and king mackerels (figs. 182, 183). The fact that its sides have dark markings below the lateral line, but not above the latter, is the readiest field mark by which to distinguish it from its close relative the false albacore (p. 336), Ln which the reverse is true. Also, its anal fin originates Figure 178. — Striped bonito (Euthynnus pelamis). After Smitt. 336 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE farther forward than in the false albacore; i. e., under the middle of the second dorsal fin, instead of under the first dorsal flnlet. Another distinctive character (shared, however, by the false albacore, p. 335) is that it has no body scales except along the lateral line, and covering a very prominent corselet on the forward and upper part of the trunk, which is outlined in the illustration (fig. 178) . Its lateral line curves down- ward suddenly below the second dorsal which is not the case in its relative alleteratus (p. 336). The first dorsal fin (about 15 spines) is not only much longer, relatively, than that of the mackerel, but its upper edge is abruptly concave behind the second spine, with the last 9 or 10 spines much shorter. The second dorsal is tri- angular, with concave rear edge; almost the whole of it stands in front of the anal; the anal is as large as the second dorsal and of about the same shape. There are about 8 little finlets behind the second dorsal, and about 7 finlets behind the anal. The pectorals are of moderate size, reach- ing back only about midway of the first dorsal. The tail fin is very short but broad and lunate in outline. Color. — Deep steel blue above, with the lower part of the sides, the throat and the belly shining white. Each side is barred behind the corselet with 4 to 6 longitudinal blue or brown stripes, the upper ones terminating at their intersection with the lateral line, the lower 3 or 4 fading out as they near the caudal peduncle.79 " The number of stripes is different in different geographic regions; Ameri- can fish usually show only 4; 7 have been described for Japanese specimens; there usually are 4, and sometimes 5 or 6, on each side in the European bonito. Size. — This bonito grows to a length of about 30 inches. General range. — -Warmer parts of all the great oceans, the Atlantic, Pacific, and Indian. Occurrence in the Gulf of Maine. — A specimen obtained at Provincetown in 1880 by J. Henry Blake is the only record for this oceanic fish in the Gulf, but it sometimes appears in numbers about Woods Hole, where 2,000 to 3,000 were taken in 1878, but where it did not show again until October 1905. False albacore Euthynnus alleteratus (Rafinesque) 1810 Little Tunny; Bonito Jordan and Evermann, 1896-1900, p. 258. This bonito resembles the striped bonito (p. 335) very closely in body form, in size and arrangement of its fins, and in the fact that its body has no scales except on the forward and upper part of the trunk, the corselet, and along the lateral line. But it is distinguishable from the striped bonito by its color pattern, for it is above its lateral line that its sides bear dark markings, not below. And its lateral line does not bend downward appreciably below the second dorsal fin.80 Also, its anal fin originates relatively farther back than in the striped bonito, i. e., under the first dorsal finlet instead of under the middle of the second dorsal fin. M For further differences between the species of Emhimnus, see Fraser Brunner, Ann. Mag. Nat. Hist., Ser. 12, vol. 3, 1950, p. 150. Figure 179. — False albacore {Euthynnus alleteratus), Woods Hole. From Jordan and Evermann. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 337 Color. — Steel blue above, glistening white lower down on the sides and on the belly. The sides are without markings below the lateral line, except for a few dark spots below the pectoral fin, but are marked above the lateral line with dark wavy bands, in various patterns. Size. — -About the same as E. pelamis, i. e. growing to about 2)i feet. General range. — This, like its relative pelamis is a tropical-oceanic fish, widespread on the high seas, in all the great oceans. Occurrence in the Gulj oj Maine.- — False albacores are picked up from time to time near Woods Hole, in July or August. But the only records of them within our Gulf are of 200 to 300 taken in a trap at Barnstable, in the autumn of 1948,81 and of 28 taken in another trap in Cape Cod Bay, near Sand- wich, on September 1 1 , 1949.82 Like various other tropical fishes they come our way only as strays from warmer seas; they are likely to be in schools whenever they reach our Gulf. Common bonito Sarda sarda (Bloch) 1793 Bonito; Skipjack; Horse mackerel Jordan and Evermann, 1896-1900, p. 872. Description.- — This bonito is shaped much like a small tuna, being thick and stout bodied, about one-fourth as deep as it is long (not counting the caudal fin), and similarly tapering to a pointed snout and very slender caudal peduncle. It is tuna-like also, in that its body is scaled all over, that its caudal peduncle has median longitudinal keels, and that its two dorsal fins are so close to- gether that they are practically confluent. But !' Reported to us by Frank Mather of the Woods Hole Oceangraphic Institution. All of these, weighing 2,498 pounds, were caught on September 16 in the trap of John Vetorino. « Schuck, Copeia, 1951, p. 98. the shape of its fins distinguishes it at a glance from a small tuna, the only regular member of the Gulf of Maine fish fauna, with which it is apt to be con- fused,83 its first dorsal being relatively much longer than that of the tuna (about one-third as long as the body, not counting the caudal, and with about 21 spines), and its second dorsal considerably longer than high, whereas the second dorsal is at least as high as it is long in the tuna. The mouth, too, of the common bonito is rela- tively larger than that of the tuna, gaping back as far as the hind margin of the eye, and its jaw teeth are larger, with the two to four in the front of the lower jaw noticeably larger than the others. The shape of its first dorsal, with nearly straight upper margin marks it off from the oceanic bonito (p. 335), also from the false albacore (p. 336), in both of which this fin is very deeply concave in outline; the uniform scaliness of its body, also, is diagnostic, as contrasted with them. We need only note further that its first dorsal fin is triangular, tapering regularly backward, with only slightly concave upper edge; that the margins of the second dorsal and anal fins are deeply concave; that it has 7 or 8 dorsal finlets and 7 anal finlets; that its tail fin is lunate, much broader than long; and that its lateral line is not deeply bowed below the second dorsal, but is only wavy. Color. — The color of this bonito is so distinctive as to be a ready field mark to its identity, for while it is steely blue above with silvery lower part of the sides and abdomen, like most of the mack- erel tribe, the upper part of the sides are barred with 7 to 20 narrow dark bluish bands running obliquely downward and forward across the lateral line. While young its back is transversely barred 13 No one should take a bonito for a large mackerel, its dorsal'flns'being close together, while those of the mackerel are far apart. Figure 180. — Common bonito (Sarda sarda). After Smitt. 338 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE with 10 to 12 dark-blue stripes, but these dark cross-bars usually disappear before maturity. Size.- — This bonito grows to a length of about 3 feet and to a weight of 10 to 12 pounds. Habits. — The bonito is a strong, swift, predace- ous inhabitant of the open sea and like all its tribe travels in schools. When they visit our northern waters they prey upon mackerel, ale- wives, menhaden, and other smaller fish such as launce and silversides; also upon squid. They are very likely to be noticed, for they jump a great deal when in pursuit of their prey. Further to the southward the bonito spawns in June; but it is not likely to spawn in the Gulf of Maine, nor does it do so in the northern part of its European range. Presumably its eggs are buoyant like those of other scombroids. Young 5 to 6 inches long have been reported as common off Orient, N. Y., early in September.86 But nothing is known of its rate of growth. General range. — Warmer parts of the Atlantic, including the Mediterranean; north to outer Nova Scotia,86 on the American coast and to Scandinavia on the European coast. Occurrence in the Gulf of Maine. — Cape Ann is the northern limit to the usual occurrence of the bonito within our Gulf. It has been taken oc- casionally in Casco Bay, while one was recorded from the mouth of the Kennebec River in Septem- ber 1930 and two more in July 1932.87 But we find no definite record of it east of this on the coast of Maine, or in the Bay of Fundy, although the young have been reported from Halifax on tbe outer coast of Nova Scotia. Its usual limitation to the southern half of the Gulf appears clearly in the location of the commercial catches. In 1919 8S for example, pound nets, traps, and other gear, accounted for almost 34,000 pounds in Cape Cod Bay, but only 90 pounds about Cape Ann, while the entire catch landed in the fishing ports of Maine during that year was only half a dozen fish (44 pounds). And there have been so 81 Xichols and Breder (Zoologies, New York Zool. Soc, vol. 9, 1927, p. 123). •'"Fair numbers'* have been taken in St. Margarets Bay, also some in mackerel traps near Lunenberg, and one was taken at Cape Breton, Nova Scotia, in October 1937 (McKenzie, Proc. Nova Scotian Inst. Sci., vol. 20, 1939, p. 16) . It is also reported from the mouth of Halifax harbor (Jones, Proc. and Trans. Nova Scotian Inst. Sci., vol. 5, pt. 1, 1882, p. 88). One specimen, 276 mm. long, was taken off Centre East Pubnico, September 12, 1951 (reported to us by A. H. Seim). " Reported by Walter H. Rich. « Nineteen nineteen is the most recent year, the published statistics for which mention bonito in the regional breakdown of the total Massachusetts catch. And there is nothing in the published fishery statistics to suggest that the status of the bonito has changed since then. few of them in Maine waters of late that none at all were mentioned in the fisheries statistics for that State of late years. Bonito have been known to reach Cape Ann in larger numbers in the past, as happened in 1876, when 73 were taken in one August day in a weir near Gloucester. And probably they are far more plentiful every year out at sea in the southern part of the Gulf than these meager returns would suggest, for fishermen often mention schools of them. Capt. Solomon Jacobs reported them as very plentiful, in August 1896, for instance, in the deep water to the northward of Georges Bank. And we have seen schools of large scombroids, (probably bonito) splashing and jumping off Cape Cod more than once in August. Apparently bonito visit New England shores only in the summer and fall. Thus the earliest catch made by a certain set of pound nets at Provincetown over a period of about 10 years was in July (1915), the latest on October 4 (1919). The bonito is more regular in its occurrence west and south of the Cape, being common in some years at Woods Hole and especially off Marthas Vineyard, whence about 57,000 pounds were mar- keted in 1945. And party-boat captains have described Buzzards Bay and the waters around the Vineyard and Nantucket as full of them in some recent summers. Importance. — The bonito is a good food fish. It readily bites a bait trolled from a moving boat, once one has the lure that it will strike on the particular occasion. A good many are caught in this way off southern New England, and we can assure the reader that a bonito is one of the strong- est fish that swims, weight for weight, and one of the swiftest. Bonito are picked up now and then in Cape Cod Bay by anglers trolling for other fish; we heard of two taken in this way off Well- fleet, on August 29, 1950. But they are never abundant enough in the Gulf of Maine to be worth fishing for there with hook and line. Tuna Thunnus thynnus (Linnaeus) 1758 Blue fin tuna; Horse mackerel; Great albacore; Tunny; Albacore 89 Jordan and Evermann, 1896-1900, p. 870. Description.- — The two dorsal fins of the tuna are practically continuous, a character (with the •' A comprehensive list of publications dealing with the tunas Is given by Corwin, Division Fish and Game of California, Fish Bull. No. 22, 1930. FISHES OF THE GULF OF MAINE 339 Figure 181. — Tuna (Thunnus thynnus). After Smitt. large number of finlets) sufficient in itself to sep- arate a very young one from either of our true mackerels. A small one is readily separable from the striped bonito and from the false albacore by the fact that the entire trunk of the tuna, includ- ing the belly, is scaly, the upper outline of its first dorsal fin only very slightly concave; and from the common bonito (p. 337) by a second dorsal that is considerably higher than it is long by the shape of its anal with only weakly con- cave margin, by the small size of its jaw teeth; and by the midline of the roof of its mouth armed with hairlike teeth. The plain coloration of the tuna, without dark markings, is still another con- venient field mark for separating small ones from any of the bonito tribe that have been reported from our Gulf. The tuna is shaped like a bonito rather than like a mackerel, with robust body, about one- fourth to one-sixth as deep as long, tapering to a pointed nose and to a very slender caudal pe- duncle which bears a strong median longitudinal keel on either side. The first dorsal fin (13 or 14 spines), originating close behind the axil of the pectoral, is triangular, its upper edge weakly con- cave, tapering backward from its first spine, and with the last spine very short indeed. And it can be laid down, flush, in a groove along the back. The second dorsal (about 13 rays, not depressible) is almost confluent with the first (a little lower than the latter in young fish and a little higher in old) is at least as high as it is long or higher, deeply concave behind, and with sharp-pointed apex. The anal fin originates under the rear end of the second dorsal to which it is similar in out- line and size (about 12 rays). Usually there are 9 or 10 dorsal finlets and 8 or 9 anal finlets, behind the second dorsal fin and the anal fin, respectively. The tail fin is much broader than long, its margin evenly lunate, its two lobes sharp pointed, much as it is in the bonitos. The pec- toral and ventral fins are of moderate size, the former scimitar-shaped and much longer than broad.90 Color.- — The back is dark lustrous steel blue or nearly black, with gray or green reflections; the checks silvery; the sides and belly silvery gray, often with large silvery spots and bands, and iridescent with pink. The first dorsal is dusky to blackish; the second dusky to reddish brown; the dorsal finlcts yellow with dark edgings. The anal fin is silvery gray; the anal finlets the same, or yellow; the caudal dusky but more or less silvery; the ventrals and pectorals blackish above and silvery gray below.91 Size. — This is the largest Gulf of Maine fish, except for some sharks; a length of 14 feet or more, and a weight of 1,600 pounds being rumored, with fish of 1,000 pounds not rare. The heaviest Rhode Island fish on record, taken about 1913, weighed 1,225 pounds, while 4 or 5 fish have been brought into Boston that weighed approximately » The tunas and their allies are discussed by Jordan and Evermann (Occas. Papers, Cal. Acad. Sci. vol. 12, 1926); Fraser-Brunner (Annals and Maga- zine Nat. Hist., Ser. 12, vol. 3, 1950, pp. 142-146) has recently given a con- venient key to all known species of tunas, with eicellent illustrations; and Godsil and Holmberg have recently discussed the relationships of the blue- fin tunas of New England, Australia, and California (Fish. Bull. 77, Cali- fornia Dept. Nat. Resources, 1950). 11 The foregoing description of the color is based on accounts of freshly caught tuna by Storer (Fishes of Massachusetts, 1867, p. 65) and by Nichols (Copeia, No. Ill, 1922, pp. 73-74); and on fish we have seen. 340 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE 1,200 pounds each, and one in 1924 that is said to have reached 1,300 pounds; and Sella92 men- tions a "fairly well authenticated instance" of one caught 60 to 70 years ago off Narragansett Pier, R. I., that weighed in the neighborhood of 1,500 pounds, was divided among the various hotels, and fed 1,000 people. The largest caught so far on rod and reel weighed 977 pounds and was 9 feet 9 inches long.93 One of 932 pounds, taken at Wedgeport, Nova Scotia, by H. E. Teller, in September 1951, is the largest that has been caught on rod and reel in the Gulf of Maine.94 Another of 864 pounds °5 was 9 feet 4 inches long and 88 inches in girth. Large tuna of the same length and caught the same day may vary as much as 100 pounds or more in weight, depending on their condition, as pointed out by Crane.96 Lengths and weights of tuna, before being dressed, caught in Massachu- setts Bay and off Ipswich in July and August 1951 were as follows: 28 inches, 17 pounds; 34 inches, 30 pounds; 42 inches, 56 pounds; 60 inches, 144 pounds; 63 inches, 172 pounds; 66 inches, 188 pounds; 68 inches, 200 pounds; 88 inches, 516 pounds; 93 inches, 587 pounds. Off Bimini, in May and June, 1950, two 88-inch tuna averaged 415 pounds and three 93-inch fish averaged 450 pounds indicating that they are much thinner in the spring in their more southern habitat than they are in summer to the northward.97 In the western side of the Mediterranean, where tuna run smaller than in our Gulf, a 500- pound fish is very large and this is equally true off the California coast. But tuna weighing as much as 1,595 pounds (725 kilograms), if the stated weights are reliable, have been reported from the eastern parts of the Mediterranean and from the Bosphorus near Constantinople..98 Habits. — The tuna is a strong, swift fish and an oceanic wanderer like all its tribe. Probably its chief reason for holding to continental waters along our coasts during the warm seasons is that " Intemat. Rev. Gesamten Hydroblol. Hydrogr., vol. 25, Ft. 1-2, 1931. p. 60. " Caught by Coram. D. W. Hodson at Cape Breton, Nova Scotia, Sep- tember 4, 1950. « Reported In Salt Water Sportsman, for Oct. 1, 1951. " Caught near Jordan Ferry, Nova Scolia, by Alfred Kenny In 1950. » Zoologica, New York Zool. Soc, vol. 21, No. 16, 1936, p. 207. " These records are from unpublished data furnished by Frank Mather of the Woods Hole Oceanographic Institution and Howard Schuck of the U. S. Fish and Wildlife Service, who have given us much first-hand information on the habits of the tuna. M Heldt, 10 Rapp. Comm. Internat. Explor. Medit., vol. 11, 1938, p. 343 its prey are more concentrated there and hence more easily caught than over the ocean basin. The small, medium, and fairly large-sized fish, up to 350-500 pounds or so, commonly travel in small schools of half a dozen to 30 or 40 fish, but sometimes in much larger schools, and each school is usually composed of fish of about the same size: we have never heard of large and small tuna schooling together. And it seems that the very large fish usually are solitary.99 When tuna are at the surface, as they often are, they are proverbial for their habit of jumping, either singly or in schools ; they may do this when swimming about, or harrying smaller fishes, or less often, when traveling in a definite direction, in which case all that are jumping do so in the same direction. Frank Mather, for instance, reports seeing a school of 200-pounders, jumping in unison, 2 or 3 feet clear of the water. When large tuna jump, they sometimes fall flat, making a great splash, but they reenter the water a little head-first as a rule, though they do not make as complete and graceful an arc in the air as the various oceanic kinds of porpoises usually do. When schools, at the surface, are not jumping, they often splash a good deal and they are conspicuous then. We remember, for instance, sighting a large school so employed, off the Cohasset shore at a distance of about 3 miles, on one occasion. Even if they are neither jumping nor splashing, as is more com- monly the case, the wakes that large ones leave behind them betray their presence, if the sea is smooth. They sometimes cut the surface with the sickle- shaped second dorsal fin and with the tip of the caudal fin, on calm days, and they have been photographed while so doing.1 But we have not seen this and experienced tuna fishermen have told us that tima are not often seen finning. In any case, it seems that the first dorsal fin is laid back, when they do fin; at least we have never heard of a tuna as showing both of its dorsals above the surface, except after it had been hooked.2 •» Crane (Zoologica, New York Zool. Soc, vol. 21, 1936, pp. 207-211) has given a readable account of the tuna off Casco Bay, which we cannot better, and with which our own sightings of tuna agree. i See Farrington (Fishing the Atlantic, 1950 [approximate date], upper photo facing p. 421), for an excellent photograph of a tuna finning. s See Farrington (Fishing the Atlantic, 1960 [approximate date], lower photo facing p. 421), for an excellent photograph of a hooked tuna showing the first dorsal fin as well as the second dorsal. FISHES OF THE GULF OF MAINE 341 Tuna often break the surface when striking a bait, or they may even leap clear then. But for some reason they do not jump ordinarily after they are hooked, but first make one or more swift shallow runs and then tend to bore deep unless in very shallow water. Tuna prey on smaller fishes, especially those of the schooling kinds, the particular species depend- ing on the local supply. In the Gulf of Maine they destroy great numbers of herring, large and small; also mackerel of which they are often full. They have been reported as pursuing silver hake; 26 tuna contained these, out of 30, that were examined by Crane at Portland, Maine, in July 1936. She also reports a rosefish (Sebastes) in one. No doubt they take whatever small fishes are available locally, and a tuna has been known to swallow a whole dogfish as large as 8 pounds. Southward from Cape Cod they prey on men- haden, as predaceous fishes do in general. They also eat squid: Crane found squid, in two, at Portland, and quantities of euphausiid shrimps (Meganyctiphanes) in two others. It is not unu- sual for tuna to strand in pursuit of prey. But this is a timid fish and easily frightened though so voracious. Tuna have no serious enemies in the Gulf of Maine, but killer whales take toll of them in Newfoundland waters where, writes Wulff 3 "one or more times annually, usually in September, orcas will ravage the tuna schools in the bays they frequent most." The tuna is a fish of at least moderately warm seas. The smaller sizes seem rather closely restricted to regions where the surface layer is warmer than 60°-62°, and while large ones are regular visitors in summer to the eastern side of our Gulf where the water warms only to about 50°-54°, this, seemingly, is about the lower limit to the thermal range they favor.4 Few tuna, for ex- ample, whether large or small, are seen in the Passamaquoddy region in most summers (p. 343) though the multitudes of small herring there would seem to offer ideal feeding conditions, but where the temperature rises only to about 52°-54° even by August, when it is highest. And seasonal chilling is generally accepted as the factor that I Interaat. Game Fish Assoc. Yearbook, 1945, p. 65. * The tuna that visit the west coast of Newfoundland find summer temper- tures as high as 59°-G0° along the south coast of Newfoundland, and 55°-57° In Trinity and Conception Bays on the southeastern part of the Newfound- land coast. drives them from our northern coasts in the autumn. Tuna tolerate a wide range of salinity, and they run well up into bays, and even into harbors in pursuit of herring; the bays on the outer Nova Scotian coast for example; Bras D'or "lake," Cape Breton; Bonne Bay on the west coast of Newfoundland; and Trinity and Conception Bays on the southeastern coast of Newfoundland. But we have never heard of one entering brackish- water. Tuna are as definitely migratory as the mackerel is, those that visit our coasts working northward in spring, to drop out of sight again late in the autumn.5 They are said to be around Jamaica throughout the year, but most plentiful there in March and April 6 Ordinarily they appear ear- liest on the Bahaman side of the Straits of Florida in the first or second week in May; next off New Jersey, off Long Island, off southern New Eng- land, and in Cape Cod Bay in June. But they have been reported well within the Gulf of Maine by the last week of May (p. 342), or nearly as early as in Bahaman waters. This, with the added fact that they are not known to approach the American coast anywhere between the Bahama Channel and North Carolina or Virginia 7 sug- gests that we may have two separate populations, a southern and a northern. They usually arrive in Bonne Bay, on the Gulf of St. Lawrence coast of Newfoundland in late June or in early July, and a week or two later in Trinity and Conception Bays, on the south- eastern part of the Newfoundland coast.8 Finally, we should point out that it is not known yet whether the tuna populations of the two sides of the Atlantic are entirely separate, one from the other, or whether more or less inter- change takes place between them. The vertical range of the tuna is from the surface down to an indeterminate depth; the only barriers likely to limit their descent are the • See Heldt (Bull. No. 5, Station Oceanographiquc de Salambo, 1926), and Sella (Int. Rev. Hydrobiol.. Uydrogr., vol. 24, 1930, p. 446) (or accounts of the migration and food of tuna in the Mediterranean and eastern Atlantic. • Information contributed by Capts. Eddie Wall and Walter Whiteman, for which we are indebted to Frank Mather of the Woods Hole Oceano- graphic Institution. ' Frank Mather of the Woods Holo Occanographic Institution informs us that a 600-pound tuna has been taken in a trap 200 miles south o( Chinco- teague, Md., and that small ones are taken off Chiucoteague. "Tuna" are reported from time to time off North Carolina, also. But it is not yet certain whether these actually are "blueflns." • Wulff, Internat. Game Fish Assoc. Yearbook, 1943, p. 65. 342 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE low temperatures they would encounter in regions where there is a strong thermal gradient (the Gulf of Maine is an example, p. 344), the increasing scarcity of prey, and, perhaps, darkness. The breeding habits of the tuna remained a mystery until recently. And while it is now known that those that visit the Mediterranean spawn in June and July, both the spawning grounds of our American tuna and their spawn- ing season are yet to be learned. The eggs (Mediterranean) are buoyant, small for so large a fish (1.05-1.12 mm. in diameter) with one oil globule of about 0.27 mm. The larval stages have also been recorded in the Mediterranean in abundance; and the characters determined by which they may be distinguished from allied species.9 Tuna fry of 3% inches (81 mm.) and about 6 inches (152 mm.) have also been pictured and are described from the Gulf of Mexico by Fowler.10 Rate of growth. — The rate of growth of so large a fish is naturally a matter of much interest. Young fry grow so rapidly that fish hatched in June in the Mediterranean reach a weight of a little less than % pound to a little more than 1 pound (300-500 grams) by September. Accord- ing to studies by Sella, based on the number of concentric rings in the vertebrae" for 1,500 in- dividuals, Mediterranean tuna average about 10 pounds at 1 year of age, about 21 pounds at 2 years, about 35 to 36 pounds at 3 years, about 56 pounds at 4 years, about 88 pounds at 5 years, about 128 pounds at 6 years, about 170 pounds at 7 years, about 214 pounds at 8 years, about 265 pounds at 9 years, about 320 pounds at 10 years, about 375 pounds at 11 years, about 440 pounds at 12 years, about 517 pounds at 13 years, and 616 to 660 pounds at 14 years of age. Average lengths of 20 to 24 inches in their second summer of growth, 27 to 34 inches in the third, 35 to 40 inches in the fourth, and 42 to 46 inches in the fifth, reported by Westman and Gilbert 12 suggest about the same growth rate for • See Sella (Atti Realo accad. Lincei, Roma, Ser. 5, vol. 33, Fasc. 7-8, semestr. 1, 1924, p. 300) and Sanzo (R. Comit. Talass. Ital. Mem., No. 189, 1932) for description of the larvae; Hcldt (Bulls. 5 and 18, Station Oceano- graphique Salambo, 1926 and 1930) for summaries of all previous observations on the breeding habits and larval stages. io Monogr. 6, Acad. Nat. Sci. Philadelphia, 1944, pp. 261, 373. u Memoria No. 166, R. Comitato Thalassografico Italiano. 1929, p. 10. 13 Copeia, 1941, pp. 70-72, based on' length frequencies for those up -to 3 years of age and on scale studies for the older ones. the American tuna. Thus the giants of 800 pounds and heavier have reached a very respect- able age. According to Sella 13 Mediterranean tuna weighing only 35 pounds may already be sexually mature. But nothing definite is known about the American fish in this regard. General range. — Warmer parts of the Atlantic (including the Mediterranean) , Pacific and Indian Oceans; 14 north regularly to the western, southern and southeast coasts of Newfoundland,16 on the western side of the Atlantic; to Iceland and northern Norway (Lofoten Islands) on the European side. Occurrence in the Gulf of Maine. — The tuna is a yearly visitor to our Gulf. Every fisherman knows the tuna or horse mackerel, as it used to be called, and this great fish visits all parts of the Gulf of Maine, but we do not understand its comings and goings much better now than when Storer called attention to its abundance about Provincetown nearly a century ago. Scarcity is not to blame for this (it is common enough) but the fact that little attention was paid to it until recently for want of market value. And while a demand for tuna has developed of late, as is reflected in the catches (p. 346), and while many anglers now fish for them (p. 347), most of the resulting information is confined to the few inshore localities where they either seem to be the most plentiful, or where they are caught most easily from small craft, or incidentally in the fish traps. It is now known that tuna are to be found all around the shores of the Gulf from Cape Cod to eastern Maine; in the Bay of Fundy; also along the west coast of Nova Scotia. And fishermen often report them on Nantucket Shoals, Georges Bank, and Browns. In ordinary years the first of them are likely to be seen as early in the season between Cape Ann and the Maine State line as they are off Cape Cod. In 1950, for example, the earliest report of them was off Hampton, N. H., May 26; the next off Plum Island, Mass., on June 9; and it was not until about June 16 that word came of one hooked in Cape Cod Bay, and of the first fish (one of 462 pounds) harpooned off Plum " Memoria No. 156, R. Comitato Thalassographico Italiano, 1929, p. 6. '* Sella's recent studies (Internat. Rev. Ges. Hydrobiol., Hydrogr., vol. 25, 1931, pp. 48-50) showed no characteristic differences between the bluefln tuna of tho two sides of the Atlantic, and those of different oceans appear, at most, to represent races of a single wide-ranging species. " Vesey- Fitzgerald and Lamonte (Game Fishes of the World, 1949, p. 183) report tuna from Hamilton Inlet. FISHES OF THE GULF OF MAINE 343 Island. This may have been an early year. But pounds, caught in a herring weir at Grand Manan tuna are to be expected throughout the western during that October. side of the Gulf generally by the middle or end of The regional contrasts in local abundance June, which is about as early as they ordinarily within our Gulf may be illustrated for a repre- appear in any numbers off southern New England; sentative year by the reported catches of tuna by and they appear on the Nova Scotian side of the counties around the coast from southwest to Gulf by the first of July if not earlier. In 1950, northeast, for 1945. for example, upwards of 450 had been landed from Massachusetts: Poonds Ipswich Bay by July 31, the largest weighing 734 Barnstable (chiefly Cape Cod Bay) 301, 900 pounds.16 The peak season usually is from about Plymouth 600 the middle or end of July to the middle of Sep- M-Efex— " 50> 300 tember off Massachusetts; July and August off ' Y' Casco Bay; through August and September along Cumberland] western Nova Scotia. Sagadahoc |vicinity of Casco Bay " - 815,300 The vicinity of Provincetown, with Cape Cod Lincoln 900 Bay, has long been known as a center of abund- HD°X k™ S°° ance for tuna. Other well known centers are from Washington" " 0 Cape Ann north to Boon Island and from the Nova Scotia: Ipswich Bay-Plum Island shore out to Jeffreys Annapolis ' 0 Ledge some 30 miles off shore; off the mouth of Yarmouth 35,800 Casco Bay and for some distance thence eastward; ape and the vicinity of Wedgeport, on the west coast In most 3*ears the tuna that are seen and caught of Nova Scotia, where the International tuna near Provincetown at the tip of Cape Cod, and matches are held. Fewer are seen along the in Cape Cod Bay, are small (so-called "school eastern coast of Maine, though we are told that fish" weighing less than 200 pounds with many a fishery for tuna has developed during the current as small as 30 to 70 pounds; and few of those summer off Southwest Harbor, Mount Desert caught there in most years are large. The smallest Island,17 and in the New Brunswick side of the reported in the inner part of the Gulf of Maine Bay of Fundy. was a run of 20- to 26-pound fish (2-year-olds) It is expecially interesting that there are so taken in Cape Cod Bay in October 1950.20 And few tuna in the Passamaquoddy region in most good catches of "school" fish of 30-70 pounds, years that the capture of even an occasional fish but few larger, if any, are being made again off in the local weirs causes comment, for the astound- the tip of Cape Cod around the shores of Cape ing abundance of small herring there would seem to Cod Bay at this writing (August 5, 1951), and offer them an inexhaustible supply of food. But have been for several weeks past. Large numbers a summer comes now and then when they are far of even smaller tuna, averaging about 11 pounds, more plentiful there than usual; thus Passama- have been encountered on the southwestern part quoddy waters are said to have "teemed with of Georges Bank (p. 344), and many of these little tuna" in the summer of 1937 18 when as many as ones (from 8 pounds or so upwards) are caught 7 were taken at Campobello in a single seining; off southern New England every summer and and several were reported again and a few caught autumn, especially near Block Island.21 On the in Passamaquoddy Bay in the summer of 1945. I9 other hand, most of those found northward from Dr. Huntsman writes us that "schools" were Cape Ann, and in the Nova Scotian side of the reported there in the summer of 1951, when the GuU are large, few of them as small as 100 pounds, water was warmer than usual. And Leslie Thus, the average live weights of 1,641 tuna that Scattergood reports 22, ranging from 113 to 161 were landed at Portland, Maine, during the period 1926 to 1935, varied between 495 pounds " Reported by Henry Moore In tbe Boston Herald, July 31, 19.10. *> Reported by Frank Mather of-tbe Woods Hole Oceanographlc Instltu- >; Information supplied by Frank Mather, of the Woods Hole Oceano- tlon. graphic Institution. si Frank Mather, of the Woods Hole Oceanographlc Institution, reports >• Atlantic Fisherman, vol. 18, No. 9, October 1937, p. 28. a catch of 110 of them, weighing about 10 pounds, off No Mans Land, on » Atlantic Fisherman, vol. 26, No. 8, September 1945, p. 52. September IB. 1951. «S 344 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE and about 630 pounds yearly, as appears from the following table.22 Year Number Largest Smallest Average 1926 90 176 152 172 107 91 162 268 423 945 858 905 967 840 800 909 955 913 416 74 447 423 420 409 93 68 33 515 1928 1929 ._.. 1930.. 1931 1932 1933 1934 1935 525 Thirty-two fish caught at the mouth of Casco Bay in 1950 averaged 46S pounds, the heaviest 643 pounds; 23 the smallest among 34 measured by Crane,24 at Portland, Maine, weighed 65 pounds, the heaviest 860 pounds. And many fish are taken of 700 pounds and heavier. Similarly, 23 tuna caught during the international match at Wedge- port, Nova Scotia, in the second week of August 1950, weighed from 362 pounds to 744 pounds, and 72 taken there during the match of the previous year averaged about 360 pounds, the largest weigh- ing 857 pounds. Also, most of the tuna caught in the Gulf of St. Lawrence are rather large. The reason for this regional segregation of tuna of different sizes is not known, or for the variation therein from year to year. We suspect that tem- perature is chiefly responsible; i. e., that the larger fish are more tolerant than the small of the lower temperatures prevailing in the northern and north- eastern parts of the Gulf, and in more northerly regions. Especially suggestive in this connection is the fact that the tuna run so large off Wedgeport, western Nova Scotia, where the abundant herring offer excellent feeding conditions, but where the water does not ordinarily warm above about 54° F. along the open coast, though to a somewhat higher figure locally, in enclosed situations. So many tuna come so very close inshore in Cape Cod Bay that nearly all of the commercial catch made there is taken in the traps; large schools have even been sighted within Provincetown Harbor (on October 11, 1950, for example 25) and occasion- ally a tuna comes into the surf either to strip the reel of some surf fisherman or to be landed (p. 347) . The tuna that are taken north of Cape Ann are farther out; all of them, however, are caught » Data gathered by the late W. H. Rich of the U. S. Bureau of Fisheries. *• Caught by Capt. Earl Larrabee; reported in Saltwater Sportsman for Aug. 25, 1950. * Zoologica, New York Zool. Soc., vol. 21, No. 16, 1936, p. 207. «* Reported In Cape Cod Standard Times, October 11, 1950. within 30 miles or so of the land, at farthest. And while a great concentration of tuna was en- countered by the Albatross III on the southwest- ern part of Georges Bank, on September 18, 1950, when 25 were hooked and landed, all very small, about 11 pounds apiece, it is unusual to see any large number on the offshore banks. The tuna that are seen or caught in our Gulf all are near the surface, or at least where the water is not more than 35 to 40 fathoms deep. How deep down they might be found is not known. But it is likely that they tend to keep within 50 fathoms or so of the surface, for the deeper water in the Gulf is colder than tuna appear to like (p. 341). In some years the tuna appear to remain fairly stationary in whatever part of the Gulf they visit, for weeks at a time, as is indicated in the consist- ency of catches, or the sightings reported, which is equally true of them in Newfoundland waters, according to Wulff.26 In other years they may disappear suddenly from one locality or another, after a brief stay, and without any apparent reason. In 1926, for example, when about 70 fish were taken in July off Casco Bay, only 17 were caught there in August, 3 in September, and only 1 in October (the 4th).27 In 1950 they deserted the Ipswich Bay-Plum Island region during the last week of August, not to reappear there in any numbers that season, though they continued plentiful enough off the Maine coast farther north to be worth fishing for until the end of September, with some in the Cape Cod Bay-Provincetown region until early October. There are tuna in good numbers along the outer Nova Scotia coast, off Shelburne, the vicinity of Liverpool at the mouth of the Mersey Kiver, the mouth of the La Have Kiver, Mahone Bay, and St. Margaret Bay being centers of abundance as appear from landings of 258,000 pounds in Lunen- burg County and 201,000 pounds in Halifax County in 1950. A few, also, are seen and caught around Cape Breton. It was here that the record size fish was taken with rod and reel (p. 344). Catch records suggest that only a few visit the southern side of the Gulf of St. Lawrence; 400 pounds were reported from the Gulf shore of Cape Breton in 1944, none in 1946, and it was only in one year (1925) that any were reported (975 a International Game Fish Assoc., Yearbook, 1943, p. 65. « Data from Waller H. Rich of Portland, Maine. FISHES OF THE GULF OF MAINE 345 pounds) M from Prince Edward Island during the period 1917 to 1928. They may visit the west coast of Newfoundland more regularly; for Wulff speaks of them as common and gives a photo- graph of tuna finning at the surface in Bonne Bay, M but we have not heard anything to suggest that they are anywhere near so plentiful there as they are in Nova Scotian waters or southward. Wulff writes of them as "few" on the southern Newfoundland coast, at present; but they appear to be regular visitors to Conception and Trinity Bays on the southeast coast; Mr. Tibbetts in- formed us that he once saw an abundance of tuna in Notre Dame Bay, midway of the east coast of Newfoundland; and they are reported from Ham- ilton Inlet, Labrador, then most northerly known outpost on the American Coast of the Atlantic. Most of the tuna disappear from the coasts of Maine and of northern Massachusetts by the end of September, or by the first part of October at the latest, depending on whether the season is an early one or a late.30 But considerable numbers remain in Cape Cod Bay and around the tip of Cape Cod until well into October, or even into November in some years. Thus in 1950 large schools were seen in Provincetown Harbor, and more than 5,000 pounds of small fish, averaging about 75 pounds, were landed there on October ll,31 while in 1949 about 2,000 pounds were caught nearby between November 1 and 14. The dates of the earliest and latest catches, made by a set of 8 traps, at North Truro, Cape Cod Bay, during the period 1943 to 1952 32 are illustrative. Year Earliest catch 1943 July 8 1944 June 29 1945 June 25 1946 June 15 1947 June 21 1948 June 11 1949 June7 1951 July7 1952 June 24 The monthly catches, by these same traps, mark July and August as the most productive » Sella, Internat. Rev. Oes. Hydrobiol., Hydrogr., vol. 25, 1931, p. 50. » Internat. Game Fish Assoc. Yearbook, 1943, p. 66. 80 In 1950 seven tuna of200-300 pounds were caught off Boars Hoad, Maine, during the first week of October. " Reported In Cape Cod Standard Times, October 11, 1950. •» Information contributed by the Pond Village Cold Storage Co. 210041—53 23 months. The number of pounds of tuna (dressed weight) follows: Month Largest catch Lates catch Oct. 6 Oct. 9 Oct. 26 Oct. 28 Oct. 28 Nov . 14 Oct 15 Oct. 24 Smallest catch 0 12, 255 5,029 33 2, 365 330 0 June 17, 520 July 148, 139 August 185, 305 September 70, 125 October 43, 603 November31 2, 197 Catches have also been reported along western Nova Scotia as late as the third week in October, and Wulff writes of tuna lingering through the month in the bays of Newfoundland, which is as late as they remain in any part of our Gulf. Tuna are never reported as seen moving south- ward on their way out of the Gulf to their winter quarters; they drop just out of sight. The wintering grounds of the particular bodies of tuna that summer in the Gulf of Maine, and of those that go farther east and north, are not known. Small (20-50 lb.) fish, it is true, have been caught occasionally in coastal waters off southern New England from January to March; off Block Island, for example, in 1928.3S But the bulk of the northern contingents certainly travel farther. It is probable that they winter in deep water as the Mediterranean tuna do, perhaps along the continental slope off our Middle Atlantic coast, perhaps so much farther south that some of the tuna seen (and caught) in spring in the Straits of Florida are our Gulf of Maine and Nova Scotian fish, on their way north again. We are equally in the dark as to the spawning grounds of the American tuna, for although the Gulf of Maine fish are of breeding age, no ripe ones have ever been seen off the New England or Canadian coasts, or even fish approaching ripeness. Abundance. — -We dare not guess how many tuna are in our Gulf in any summer, there being no way to estimate how large a proportion of them the yearly catch represents. We suspect that they are fewer than reports would suggest, for being so large, a few hundred of them make a great show if they are at the surface, whereas an equal number of mackerel, for instance, would never be noticed. Neither is any definite informa- tion available as to their annual fluctuations in M The year 1944 is omitted from the calculation for September-November, because the traps were not fished after September 14tb that year. u The only catch recorded for November was 2,197 pounds In 1949. « Sella, Internat. Rev. Qesamten Hydrobiol., Hydrogr.. vol. 25, 1931, p. 62. 346 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE abundance, though fishermen are well aware that The largest Gulf of Maine catches of which we their numbers in any part of the Gulf do vary have heard were of 336 fish, weighing about 75,000 widely from year to year. Thus it is on record pounds taken at one lift of 3 traps set for mackerel that they were scarce in the Massachusetts Bay on the Barnstable shore of Cape Cod Bay, Aug. 5, region for two or three years prior to 1904, but 1948; 37 and of 120,000 pounds of fish ranging were plentiful that summer. Commercial land- from 25 to 30 pounds seined some 50 miles east of ings suggest that they were scarce again in 1943, Cape Cod on September 18, 1951, by the Western when the landings came to only about 380,000 Explorer, chartered by the U. S. Fish and Wildlife pounds for Maine and Massachusetts combined.36 Service for experimental fishing for tuna. But they appear to have been much more Importance. — Horse mackerel were formerly plentiful again off the Maine coast in 1945 (catch regarded as a nuisance on the Atlantic coast, for there about 850,000 lb.) ; more plentiful than they bands of them made trouble for fishermen by have been since, if the commercial landings are a following herring or mackerel into the traps and reliable index to the ups and downs of the tuna pounds, to tear their way out again through the population, which they may not be. The follow- net unless harpooned. Many years ago, when fish ing catch statistics of tuna landings (in pounds) 0il was more valuable than now, a few were some- suggest that the stock built up more slowly, from times harpooned for oil, which was tried out of the 1943 low in Massachusetts waters, to a peak the heads and bellies, but there was no sale for in 1948, which was a big year on the Ipswich Bay their meat. The tuna, however, has been highly grounds (p. 343), as well as in Cape Cod Bay. valued as a food fish for many years, not only in Year Maine Massachusetts the Mediterranean, but on the west coast of the 1943 25,000 129,500 United States. And a local demand has developed 1944 "«SiS 5?S! on our coast> supplied chiefly by local fisheries 1945 859,500 356,400 ' . ., -, » t> tij 1946 421 800 57!, xoo off Casco Bay, m the Cape Ann-Boone Island 1947 186, 600 726, 400 region, and in the Cape Cod Bay region. 1948 -- 229,100 1,627,000 With this increasing demand, the reported In 1949, the catch by traps in Cape Cod Bay landings on the Maine and Massachusetts coasts alone was 811,160 pounds, suggesting a total of have risen from about 94,000 pounds in 1919, to more than a million pounds from the Gulf of around 250,000 pounds yearly in the early Maine coast of Massachusetts. 1930'a, and to about 1 to nearly 2 million pounds During that banner season 2,164 large tuna for the years 1945 to 1948, this last representing were taken on hand lines where the draggers work, around 3,000-6,000 fish, if they averaged 300-400 off Ipswich Bay; while 806 fish (305,300 lb.) were pounds in weight (see table, p. 346). The average taken off Wedgeport, Nova Scotia, in 1948, 1,760 value to the fisherman in 1946 was about 7-9 fish (449 362 lb ) in 1949 cents per pound and all that are caught now sell But the/ were scarce' in 1950, to judge from readily. The annual catches off the entire coast reports coming in from all along our coast: a of Nova Scotia ranged from 152,000 pounds to week's fishing, for instance, by the same number of about 1,550,000 pounds dunng the period 1917 to rods and at about the same date, yielded only 1928; from 402,000 to 1,820,000 pounds for the about half as many on the famous Soldiers Keef 5 years !942-46. off Wedgeport during the International Match The commercial catch off the coasts of Maine is that summer as it had the year before. Many made mostly by harpoon; that off northern fewer were caught by anglers in Cape Cod Bay Massachusetts by hook and line and by harpoon; in 1950 than in 1949, and the tuna disappeared that off the Cape Cod Bay region mostly in the from the Ipswich Bay region unusually early that traps.38 year, as noted above (p. 344). It is too early (August 5) to forecast how the 1951 Season may ,7 A photograph of part of the catch was published In the Boston Herald, develop August 6, 1948. r * s» In 1945 about 60 percent of the catch reported for Maine was by harpoon . almost all the remainder on hand lines; in 1946 about 98 percent was bar- s' The weights given in the Fisheries statistics are for the dressed fish, and pooned. About 86 percent of the Massachusetts catch was taken in traps of represent about 80 percent of the live weight. one sort or another in 1945. about 90 percent In 1946. FISHES OF THE GULF OF MAINE 347 But experiments are in progress, by the U. S. Fish and Wildlife Service, to find whether a prof- itable seine fishery or long line fishery can be developed for these great fish in our Gulf, with the hope of providing a more dependable supply, and through a longer season. The sporting qualities of the tuna in our Gulf deserve a word, though an extended account would be out of place here. Encouraged by the famous tuna fishing off the coast of southern California, and by the knowledge that tuna run much larger in the Gulf of Maine than they do on the Pacific coast, several anglers had caught these huge fish with light tackle at various points in New England and Nova Scotia by 1925, when the first edition of this book appeared. Small tuna will often take artificial lures especially if trolled at high speed and close to the vessel's stern, while large ones will take a hook baited with herring, mackerel, or other fish. And tuna fishing has now grown to be so popular and successful a sport that many party boats go out regularly off Provincetown, in Cape Cod Bay, to the Ipswich Bay-Isles of Shoals-Boone Island region, off Casco Bay, and off Wedgeport on the Nova Scotian side.39 To date, the largest tuna that has been landed on rod and reel in the Gulf of Maine was one of 932 pounds, caught by H. E. Teller at Wedgeport, NovaScotia, September 11, 1951 (p. 340). Another of 927 pounds was caught in Ipswich Bay, August 4, 1940, by Dr. J. B. Vernaglia. We have heard of one of 180 pounds landed with ordinary surf- casting tackle on the beach at Plum Island, a " Farrington (Field and Stream magazine for August 1950, p. 84) has recently given an interesting account of the methods employed by rod and reel anglers, in these localities. Crane (Zoologica, N. Y. Zool. Soc., vol. 21, No. 16, 1936, p. 210) describes In a readable way the small boat harpoon fishing for tuna off Casco Bay, Maine. notable feat.40 Even a small tuna, such as a thirty-pounder that was caught on a black plug by Wm. Lakaitis, surf casting at North Truro on the night of July 28, 1951, is a far more stronger adversary in the surf than a striped bass of equal size. Spanish mackerel Scomberomorus maculatus (Mitchill) 1815 Jordan and Evermann, 1896-1900, p. 874. Description. — The Spanish mackerel has the outline of the slender mackerel rather than of the stout bonito, its body being nearly 4% to 5 times as long as it is deep. But there is no danger of confusing it with either of the true mackerels, first, because its two dorsal fins (like those of the bonitos) are hardly separated, and second, because of its color pattern, Its high second dorsal, slender form, and spotted sides mark it off at first glance from our bonitos, while its color, slender form, long first-dorsal fin, and the outline of its second dorsal distinguish it from a small tuna. The most clear-cut distinc- tion between the Spanish and its close relatives the king mackerel and the cavalla, is that the pectoral fins of the Spanish arc naked but those of the last two are mostly covered with scales. The ventral fins, also, originate definitely behind the origin of the first dorsal in the Spanish, under it or only a very little rearward in the king; and the color differs. The most distinctive anatomic character of the Spanish, king, and cavalla among our local scom- broids is the large conical jaw teeth. The caudal peduncle of the Spanish mackerel is keeled; its lateral line wavy; its first dorsal fin (17 to 18 <• Landed on August*12, 1950,[by[M. L. Insleyn. #fc^s. Figure 182. — Spanish mackerel (Scomberomorus maculatus) . From Goode. Drawing by H. L. Todd. 348 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE spined) is triangular; its second dorsal (14 to 18 rays) is concave and originates a short distance in front of the anal, which is similar to it in form and size. It has 8 or 9 dorsal and as many- anal finlets. Its pectorals are naked; its caudal is deeply lunate, with the outer rays decidedly longer than those of the common mackerel. It has 32 teeth, or fewer, in each jaw. Color.- — The Spanish mackerel is dark bluish or blue green above, pale below, like all scombroids, and silvery, its sides marked with many small, oblong-oval, dull orange or yellowish, spots, both above the lateral line and below, these spots being a very diagnostic character. The fact that the membrane of the front one-third of its first dorsal fin is black (blue in the king, p. 348), whereas its rear part is greenish white, is an equally useful field mark. The second dorsal and pectoral fins are pale yellowish with dusky edges; the anal and the ventrals are white. Size. — The maximum weight is about 9 or 10 pounds,41 maximum length about 36 inches, but the fish average less than 3 pounds as caught. General range. — Both coasts of North America, north commonly as far as Chesapeake Bay in the Atlantic, and to Maine as a stray; south to Brazil. Occurrence in the Gulf of Maine. — The Spanish mackerel is a summer visitor all along the Atlantic coast of the United States, as far north as New York; less regularly along the southern coasts of New England, though a few are taken during most summers at Woods Hole. But it is only a stray in the colder waters of the Gulf of Maine, where occasional fish are taken in Cape Cod Bay every «> A weight of 25 pounds is recorded by Smith (North Carolina Oeol. Econ. Surv., vol. 2, 1907, p. 191) for a specimen observed in a Washington, D. C, fish market. If the identity was correct and this was not the closely related king mackerel Scomberomorus regalis it must be considered a case of giantism. year or two. In 1896 the local catch rose to 37 fish (Provincetown and Truro traps), and there is record of it at Lynn, Mass. But Spanish mackerel are so rare north of this point that Monhegan Island is the only locality-record for Maine, and the most northerly known outpost for the species. King mackerel Scomberomorus regalis (Bloch) 1793 KlNGFISH Jordan and Evermann, 1896-1900, p. 875. Description. — The king mackerel resembles the Spanish mackerel closely in general appearance, but its pectoral fins are mostly covered with scales ; its ventrals are below the first dorsal, instead of definitely behind the origin of the latter; its head is relatively longer, its nose more pointed, its teeth more numerous (about 40 in each jaw), triangular and very sharp pointed; and the upper half of its first dorsal is deep blue. Furthermore, the king mackerel is marked by a narrow brownish stripe running from close behind each pectoral fin to the base of the caudal, crossing the lateral fine as the latter bows downward below the second dorsal fin. Its side spots, too, are mostly below the lateral line and arranged in rows, whereas the spots of the Spanish mackerel are irregularly scattered, with about as many above the lateral fine as below it. Size. — Said to reach 35 pounds, but the average weight is between 5 and 10 pounds. General range. — Atlantic Coast of North Amer- ica, Cape Cod to Brazil, abundant among the West Inches and around southern Florida. Occurrence in the Gulf of Maine. — This southern fish is recorded by Dr. W. C. Kendall at Monomoy, at the southern elbow of Cape Cod, but it has not been taken elsewhere in the Gulf of Maine. NSs Figure 183. — King mackerel {Scomberomorus regalis), Key West, Florida. From Goode. Drawing by H. L. Todd FISHES OF THE GULF OF MAINE 349 Ca valla Scomberomorus cavalla (Cuvier) 1829 42 Cero Jordan and Evermann, 1896-1900, p. 875. Description. — The pectorals of the cavalla are scaly, and its anal fin, like that of the king mackerel originates about under the origin of its second dor- sal, in which it differs from the Spanish mackerel (p. 347). In fact, it resembles the king mackerel so closely in general appearance that the one might easily be taken for the other by anybody not used to handling the two fish as southern fisherman are. But the lateral line (very conspicuous in both) is a sure clue to identity, for this dips downward abruptly in the cavalla under the forepart of the second dorsal fin, but slopes down only gradually there in the lung mackerel. Other points of differ- ence are that the outline of the first dorsal fin is concave in the cavalla (nearly straight in the king) ; that the cavalla has a large number of teeth (about 40 in each jaw, as against about 30); that its body is more slender (about one-sixth as deep as it is long); and that the upper forepart of its first dorsal fin is not noticeably darker than the remain- der of the fin. Color. — Iron gray above, silvery lower down on the sides and on the belly; the sides marked with darker gray or yellowish spots, which tend to disappear in large fish. Size. — Said to reach a length of a little more than 5 feet, and a weight of about 100 pounds. The rod and reel record is 73% pounds, for one taken off Bimini, Bahamas, February 1935, by L. B. Harrison. General range. — Warm parts of the Atlantic; south to Brazil in the western side ; north regularly to North Carolina (June-November);43 occasion- ally to southern Massachusetts; and as a stray to the southern part of the Gulf of Maine. Occurrence in the Gulf of Maine. — The only Gulf of Maine record of which we know is of one 20% inches long (to base of caudal fin) taken in a trap at North Truro, Cape Cod, in August 1949.44 Figure 184. — Cavalla (Scomberomorus cavalla). Woods Hole. From Goode. Drawing by H. L. Todd. THE ESCOLARS. FAMILY GEMPYLIDAE These fishes are closely allied to the true mack- erels, the most obvious differences being that they lack the keels on the sides of the caudal peduncle so characteristic of the mackerels. Escolar Huvettus pretiosus Cocco 1829 Oilfish; Scourfish; Plaintail Jordan and Evermann, 189G-1900, p. 879. Description. — This fish suggests the mackerel family in its slender fusiform shape and in the general arrangement of its fins. And its first (spiny) dorsal (13 to 15 spines), like that of the Spanish mackerel, is much longer than the second dorsal (18 soft rays). But it is separable at a « Fowler (Proc. Acad. Nat. Sci. Philadelphia, vol. 66, 1905, p. 766) refers this species to a new subgeDus Sierra. glance from all Gulf of Maine mackerels by the facts that it has only 2 dorsal finlcts and 2 anal finlets, and that its skin is set with bony plates armed with short spines instead of being velvety with small scales, as it is in the case of the mackerel tribe. The caudal fin is deeply forked. The first dorsal is much lower than the second, and the anal is situated below the second dorsal, which it parallels in its outlines. Color. — Described as purplish brown, darkest above, with blackish patches, and with the in- side of the mouth dusky. *3 Taylor (Survey Marine Fisheries North Carolina: University of North Carolina Press, 1951, pp. 261-265) has given an interesting survey of the seasonal presence of various southern game fishes off the North Carolina coast. ** This specimen was received through the kindness of John Worthington and the Pond Village Cold Storage Co. 350 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE «Sk V> Figure 185. — Escolar (Ruveltus pretiosus), Georges Bank. From Goode and Bean. Drawing by J. C. van Hook. Size. — The escolar grows to a weight of at least 100 pounds. General range. — Tropical parts of the Atlantic, and the Mediterranean, in moderately deep water (usually 100 to 400 fathoms); also widespread in warm latitudes in the North Pacific 46 and in East Indian waters. It is plentiful around Cuba though not reported from Puerto Pico; is known from Bermuda; and it has been taken as a stray as far north as the Bay of Biscay in the east and to the Grand Banks of Newfoundland in the west. There are regular fisheries for it off Cuba and about the Canaries ; also in the Pacific.46 Occurrence in the Gulf of Maine. — Two escolars, respectively, 49 inches long and 6 feet long, were brought in to the United States Fish Commission from Georges Bank during the autumn of 1891.46 It has not been seen in the Gulf of Maine region since then. The nearest record of it to the south- ward, with which we are acquainted, is of two trawled about 92 miles off Cape May, N. J., in January 1950.47 THE CUTLASSFISHES. FAMILY TRICHIURIDAE The cutlassfishes are characterized by a scale- less, band-shaped body tapering to a slim pointed tail, with one dorsal fin extending the whole length of the body; the anal is also long but is very low. The ventrals are absent or rudimentary, and there is no distinct caudal fin. Their large mouth is armed with strong teeth of various sizes. They inhabit the surface waters of tropical seas. Cutlassfish Trichiurus lepturus Linnaeus 1758 Hairtail; Scabbardfish; Silver eel; Ribband- fish Jordan and Evermann, 1896-1900, p. 889. Description. — The most striking characteristics of the cutlassfish are its band-like form tapering to a pointed whiplike tail without caudal fin; its single long dorsal fin (about 135 rays) originating close behind the eyes, about two-thirds as high at its midlength as the body is deep, and diminishing to nothing some distance in front of the tip of the tail; its long anal composed of very low detached spines pointing backward; and its long barbed fangs in the front of the mouth, four in the upper and two in the lower jaw. The depth of the body equals about one-thirteenth to one-fifteenth of its total length, about one-seventh to one eighth of which is occupied by the head. The snout is pointed, the mouth gapes back to below the eye and the lower jaw projects beyond the upper. Each of the jaws is armed with 7 to 10 smaller teeth behind the fangs. The anal fin is reduced to a series of short inconspicuous spines, about 100 to 110 in number, without connecting fin membrane, running back from the vent nearly to the tip of the tail. The small pectorals are situated a little in advance of the rear corners of the gill covers. There are no ventral fins and the skin is scaleless. Color. — Plain silvery all over. The dorsal fin is plain yellowish or dusky green in life, dark edged or speckled along the margin with black; the tips of the jaws dusky. Size. — Maximum length about 5 feet. General range. — All warm seas; abundant in the West Indies and Gulf of Mexico; not rare along the '• Gudger (American Naturalist, vol. 62, 1928, p. 467) and Nordhoff (Natural History, vol. 28, 1928, p. 40) give accounts of the geographic distribution of the escolar, and of the fisheries for it In tropical waters. « Approximate location 41° 40' N., 67° 44' W. See Goode and Bean, Smith- sonian Contrib. Knowl., vol. 30, 1895, p. 197. " LaMonte, Marine Life, vol. 1, No. 8, 1950, p. 40. FISHES OF THE GULF OF MAINE 351 Figure 186. — Cutlassfish (Trichiurus Upturns), Florida. From Goode. Drawing by H. L. Todd. South Atlantic coast of the United States, occa- sionally straying north as far as Massachusetts Bay. Occurrence in the Gulf of Maine. — The cutlass- fish is only an accidental straggler north of Cape Cod. One was taken at Wellfleet in the summer of 1845, and one in Salem Harbor also many years ago, and it is recorded from Lynn by Kendall.48 There is no report of it farther north in the Gulf of Maine, or for Canadian waters. THE SWORDFISHES. FAMILY XIPHIIDAE The upper jaw and snout of the swordfish (there is only one species) is greatly prolonged, forming a flat, sharp-edged sword. It has a very high first- dorsal fin and a very small second dorsal, both of them soft rayed ; a broad lunate tail ; two separate anal fins, the second very small ; and a strong longi- tudinal keel on either side of the caudal peduncle. It has no ventral fins, and the adults have neither teeth nor scales. The spearfish family (p. 357) is the only other group represented in the Gulf of Maine fauna which at all resembles the swordfish, but spearfish have ventral fins and minute teeth; their swords are round edged, and either there is one long continuous dorsal fin or, if there are two, the first is several times as long, relatively, as it is in the swordfish. Swordfish Xiphias gladius Linnaeus 1758 Broadbill Jordan and Evermann, 1896-1900, p. 894. Description. — The salient feature of the sword- fish is the prolongation of its upper jaw into a long, flattened, sharp-edged 40 and pointed "sword" occupying nearly one-third the total length of the fish. This sword is of itself enough to identify <> The Massachusetts Bay and Provincetown records listed by Kendal (Occ. Pap. Boston Soc. Nat. Hist., vol. 7, No. 8, 190S, p. 76) are based on the Wellfleet specimen. He also credits it to Monhegan I.f Maine, quoting Storer as his authority, but Storer stated in his latest mention of the species that only two had come to his notico; the Wellfleet specimen just mentioned, and one taken at the head of Buzzards Bay. « In Its tropical relatives, the sailflsh and spearfish, the sword is round edged, spearlike, and relatively shorter. Figure 187. — Swordfish (Xiphias gladius). After California Fish and Game Commission. 352 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE the fish at a glance among all our northern fishes. On a fish 10 feet 10 inches long, which we har- pooned on Georges Bank on the Grampus in July 1916, the sword was 42 inches long from its tip to the eyes. The swordfish is moderately stout of body, only slightly flattened sidewise, deepest just behind the gill openings, and tapering rearward to a slender caudal peduncle, which bears a single strong longitudinal keel on either side. Apart from the sword the head is short; the lower jaw is pointed, and the mouth so wide that it gapes far back of the very large eyes, which are set close to the base of the sword. Swordfish (except young fry) are both toothless and scaleless. The first dorsal fin originates over the upper angle of the gill openings and is much higher than long (about 39 to 40 rays), with deeply concave rear margin. The second dorsal is very small and set far back on the caudal peduncle. There are two anals likewise. The second is as small as the second dorsal and located below the latter, while the first is similar to the first dorsal in outline but shorter, and located well behind it, close to the second anal. The pectorals are narrow, very long, scythe- shaped, and set very low down on the sides below the first dorsal. The caudal fin is short, but as broad as half the length of the fish from top of lower jaw to base of caudal fin, with deeply lunate margin and pointed tips. There are no ventral fins.60 Color. — While all swordfish are dark above and whitish with silvery sheen below, the upper surface varies from purplish to a dull leaden blue or even to black. The eye has been described as blue. Very young swordfish, like very young tuna, are transversely barred, but none small enough to show this pattern has ever been found within the limits of the Gulf. The colors fade soon after death. Size.- — Swordfish grow to a great size. The heaviest definitely recorded from the Gulf of Maine was one caught on Georges Bank in the summer of 1921 by Capt. Irving King and landed at the Boston Fish Pier, that weighed 915 pounds dressed, hence, upwards of 1,100 pounds alive.61 This specimen was not measured, but the sword was more than 5 feet long, so that the total length of the fish must have approximated 15 feet, and 16 feet seems to be about the maximum length, though fish as long as this are very unusual. The heaviest landed in Massachusetts during 1922 weighed 637 pounds dressed; that is, upward of 750 pounds live weight,62 while the largest taken in 1931 weighed 644 pounds dressed and was 13 feet long including its sword, which measured 44 inches. One that weighed 925 pounds before it was dressed was landed in 1932; also one weighing 650 pounds dressed, which must have weighed 800 pounds alive; while one of 850 pounds (dressed?), brought in to Halifax, Nova Scotia, was said to have been the largest ever landed in that port. And several, weighing more than 500 pounds, dressed, are reported almost every year.53 But the general run are much smaller. Thus the average dressed weights of sundry fares of fish landed in Portland, Boston, and Gloucester in the years 1883-1884, and 1893-1895 were between 200 pounds and 310 pounds, falling to 114-186 pounds for the years 1917, 1919, 1926, and 1929-1930. And general report has it that Block Island fish run smaller than Georges Bank and Cape Breton fish. A 7-foot fish weighs about 120 pounds; 10- to 11-foot fish about 250 pounds; fish of 13 to 13K feet, about 600 to 700 pounds, as taken from the water. The rod and reel record is 860 pounds, for one 13 feet 9 inches long caught off Tocapila, Chile, April 28, 1940, by W. E. S. Tuker. Swordfish fry are quite different in appearance from their parents, having only one long dorsal fin and one long anal fin, a rounded tail, both jaws equally prolonged and toothed, and the skin covered with rough spiny plates and scales. But fish of half a pound weight such as are caught in abundance in the Mediterranean already resemble the adults, except that they have minute scales until 30 inches long. Habits.** — The swordfish is oceanic, not de- pendent in any way either on the coast (except as this offers a supply of food), or on the bottom; and it is a warm-water fish, most plentiful in localities and at depths where the temperature is higher than about 60°. But occasional captures M In the sailfisbes and spearfishes the body is scaly, the jaws are toothed, ventral fins are present, and the first dorsal fin is much longer than that of the swordfish. " Fishing Gazette for September 1921, p. 13. " Gloucester Times, April 26, 1923. « See Rich (Proc. Portland Soc. Nat. Hist., vol. 4, Pt. 2, 1947, pp. 34-37) for these and other large fish landed from year to year. » Rich (Proc. Portland Soc. Nat. Hist., vol. 4, pt. 2, 1947) has recently given an extended account of the occurrence and habits of our swordfish. FISHES OF THE GULF OF MAINE 353 of swordfish on halibut lines set near bottom as deep as 200 fathoms, together with the fact that swordfish are by no means rare on the Newfound- land Banks, whence several fish were brought back by the American cod fleet in 1920, proves that temperatures as low as 50° to 55° do not bar it, at least for a brief stay. Although swordfish may gather in certain localities they do not school, but are always seen scattered, either singly or at most two fish swim- ming together. Earlier published accounts, state- ments by fishermen, and our own rather limited experience all agree on this point. On calm days they often lie quiet on the surface, or loaf along with both the high first dorsal fin and the tip of the caudal fin above water, and they are easily harpooned while so employed, often allowing a vessel to approach until the pulpit projecting from her bow comes directly above the fish. When a swordfish is swimming at the surface, its first dorsal fin and the upper part of its tail fin both show above the water whereas a marlin shows its caudal only. One can tell a surfacing swordfish from a shark by its sharp-pointed dorsal (that of a shark is more broadly triangular) and by the fact that its tail fin seems to cut the water in a direct line, not wobbling from side to side as the tips of the tails of most sharks do (other than the mackerel shark tribe), if they show above the water at all. When swordfish are at the surface, they jump a good deal, perhaps in vain attempts to shake off the remoras that so often cling to them. We saw one leap clear of the water four or five times in rapid succession close to the Grampus, off Sbel- burne, Nova Scotia, on July 28, 1914. Reports by fishermen, and our own experience, are to the effect that they surface only during the hours of daylight. The swordfish is a fish-eater. During its stay in American waters it feeds on mackerel, men- haden, bluefish, silver hake, butterfish, herring, argentines, rattails (Macrourus bairdii), and indeed on any smaller fish, buckets of which have been taken from swordfish stomachs. Squid, too, are often found in them and may be their chief diet at times. And the jaws of one of the giant squids (genus perhaps Architeuthis) , taken from the stomach of a swordfish harpooned on the northern edge of Georges Bank,65 was an especially interest- ing find. One that we examined on Georges Bank, » Rich, Proc. Portland Soc. Nat. Hist., vol. 4, pt. 2, 1947, p. 23. 210941—53 24 July 24, 1916, was full of silver hake, one taken there in August 1929 contained 5 large haddock (p. 199), while another harpooned off Halifax con- tained a squid (Ilex) and fragments of silver hake. They have often been described as rising through schools of mackerel, menhaden, and other fishes, striking right and left with their swords, then turning to gobble the dead or mangled fish. And we have seen them so employed on more than one occasion, to judge from the commotion. It is not unusual for swordfish taken on the offshore banks to contain deep-sea fishes of one kind or another in their stomachs; many such instances have been recorded,68 sometimes swal- lowed so recently that they are still in good condi- tion when the swordfish is opened.67 And since these so-called "black fishes" live outside the edge of the continent, mostly below 150 fathoms, this is good evidence that the swordfish found on the banks that front our Gulf do some of their foraging at considerable depths farther out at sea. It also seems that they sometimes strip lines set for hali- but and tilefish of the fish already caught, for they are sometimes brought up tangled in the line. It was not out of the ordinary for swordfish to be hooked on long lines set for halibut in the days when this fishery flourished (p. 255). Goode M cites a number of cases, including one when 13 swordfish were caught in this way on one halibut trip. And fishermen have told us of more recent instances. Swordfish have often been hooked and landed on hand lines, also. A case is on record of 7 taken in this way on one trip, in the South Channel, in 15 to 25 fathoms of water, the bait being whole mackerel; evidence that swordfish seemingly do not insist on live food. We also read that of old, fishermen from Marthas Vineyard and Nantucket sometimes took them while trolling with some sort of silvery fish as bait, forecasting the big game anglers of today. Many tales are current of swordfish attacking slow moving vessels without any provocation, and driving their swords through the planking, either »■ The late Wal'er H. Rich of the U. S. Bureau or Fisheries reports the following genera as taken from swordfish on Oeorges and Browns Banks: A'epismrus, Chaulioi'j,), Chiasmodon, Lampadcna, Macrotlomi, Mijctophum, Xotoscopelut, and Stomlas. « Kingsley (Science, N. Ser., vol. 55, 1922, pp. 225-226) reports two freshly ! i ill nved stomlatids (Echiostomi barbitum) being taken'from the stomach of a swordfish harpooned on the offshore slope of Georges Bank. » Rept. U. S. Comm. Fish., (1880) 18S3, pp. 353-354. 354 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE in "fits of temporary insanity," as Goode 69 expressed it, or more likely, while pursuing dolphins or other fish. Most of the attacks of this sort reported from tropical seas seem actually to have been by spearfishes (p. 357) but some in northern waters seem almost certainly to have been by the broadbill. A case in point is that of the schooner Volunteer, out of Gloucester, which received a strong blow near Block Island, August 7, 1887, apparently from a 300-pound swordfish that was seen swimming alongside, and which proved to have lost its entire sword when it was harpooned and brought on board.60 We can only add that we have never heard of a swordfish making an unprovoked attack on any of the fishing vessels that pursue them every summer, or on any of the other craft, large or small, that cruise off our coasts. But fish that have been harpooned often turn on their pursuers, and it is a common event for one to pierce the thin bottom of a dory. We have, indeed, known several fishermen of our acquaintance to be wounded in the leg in this way, but always after the fish had been struck with the harpoon. Under these circumstances swordfish have been known to drive their swords right through the planking of a fishing vessel.61 Stories of swordfish attacking whales are time- honored traditions of the sea, mostly with no more foundation than the myth that they ally them- selves with the harmless thresher shark for the purpose. As a matter of fact swordfish are easily frightened, and they will not often allow a small boat to come within striking range, which made harpooning from dories difficult in the old days.62 But for some occult reason they will allow them- selves to be almost run down by a larger vessel without paying the least attention to its approach until aroused by its shadow, or by the swirl of water under its forefoot. But we have never heard of a swordfish actually being struck by a vessel; they always sound or turn aside in time. Swordfish fight gamely on the surface or below when harpooned. Storer long ago wrote that they sometimes sound with such speed and force as to » Fish. Ind. V. S., Sect. 1, 1884, p. 345. «° Related by Rich (Proc. Portland Soc. Nat. Hist., vol. 4, Pt. 2, 1947, pp. 48-49). «' Many cases of this sort are mentioned In the rather extensive literature dealing with the swordfish. « Rich (Proc. Portland Soc. Nat. Hist., vol. 4, Pt. 2, 1947, p. 71) so informs as. drive the sword into the bottom, which fishermen say is by no means uncommon; and we saw this off Halifax in August 1914, when a fish more than 10 feet long, which we had harpooned from the Grampus, plunged with such force that it buried itself in the mud beyond its eyes in 56 fathoms of water. When finally hauled alongside it brought up enough mud plastered to its head to yield a good sample of the bottom. Full-grown swordfish are so active, so powerful, and so well armed that they have few enemies. Sperm and killer whales and the larger sharks alone could menace them. And while we can find no evidence that swordfish ever fall prey to the first two, Captain Atwood found a good- sized swordfish in the stomach of a Mako shark. A swordfish erman described seeing two large sharks bite or tear off the tail of a 350-pound swordfish, which he afterwards harpooned. A 120-pound swordfish, nearly intact with sword still attached, was found in the stomach of a 730- pound Mako taken near Bimini, Bahamas, while another Mako of about 800 pounds, harpooned off Montauk, Long Island, was seen attacking a swordfish, and was found to have about 150 pounds of the flesh of the latter in its stomach when it was landed (p. 24). .And Rich 63 mentions that other like cases have been reported. Young swordfish would, of course, be preyed upon by any of the larger predaceous fishes. Swordfish are infested with many parasites besides the remoras, several of which are often found clinging to one fish. No less than 12 species of worms and 6 of copepods have been identified from fish taken off Woods Hole. The eggs of the swordfish have not been seen, or have not been identified if seen; probably they are buoyant. Neither is anything definitely known of the rate of growth of the swordfish. It has been supposed that the young fish of half a pound to 12 pounds that are taken in winter in the Mediterranean are the product of the past spring's spawning, which would call for unusually rapid growth. But the very large size to which swordfish grow may equally be the result of long life, as it is in the case of the tuna (p. 342). General range. — Both sides of the Atlantic Ocean; north to northern Norway, southern and « See Farrlngton (Field and Stream magazine, vol. 47, February 1943) and Rich (Proc. Portland Soc. Nat. Hist., vol. 4, Pt. 2, 1947, p. 44) for more detailed accounts. FISHES OF THE GULF OF MAINE 355 western parts of the Gulf of St. Lawrence, south coast of Newfoundland and Grand Banks, south to latitude about 35° south. Also in the Medi- terranean and Red Seas; about the Cape of Good Hope; and widespread in the Indian Ocean and in the Pacific Ocean, both north and south of the equator. Occurrence in the Gulf of Maine. — The swordfish seems to have attracted little attention in the Gulf in colonial days, and though it has long supported a lucrative fishery off New England, we know little more of its life there today than in 1883 when Goode M published his Materials for the History of the Swordfish. The outer half of the continen- tal shelf off Block Island and southern Massachu- setts; the offshore parts of the Nantucket Shoals region; Georges Bank; the deep channel between Georges and Browns; Browns Bank and La Have; and the banks off the outer coast of Cape Breton are its chief centers of abundance off our coasts. On these grounds 25 or more are often sighted in a day, sometimes that many are in view at one time; in fact, "one skipper reports counting 47 fish in sight at one time, after a week-long breeze had died out to a flat calm," 65 and some 10 to 20 thou- sand of them are harpooned every summer off the New England coast, with as many more off eastern Nova Scotia.66 An occasional swordfish is seen off Massachusetts Bay also, and along the Maine coast nearly every year. During some summers, of which 1884 was one, numbers of them appear there, and on such occasions some are taken in the Gulf from Cape Cod to Browns Bank, with Jeffreys Ledge and a zone about 10 to 12 miles off the coast from Boon Island to Cape Elizabeth perhaps their favorite inshore resort. But the great majority keep strictly to the offshore banks during most years, and they are seldom seen in the Bay of Fundy. Thus we find only 2,500 pounds (10 or 12 fish) brought in by the shore fishermen of Cum- berland County, and 3 or 4 fish (800 lbs.) landed in York County in 1919, while none was reported as caught off the coast of Maine in 1945, though 193,000 pounds were landed on the Nova Scotian side of the Gulf (Yarmouth Co.) in that year and the offshore catch was considerable. Swordfish seem to be less plentiful along the outer Nova Scotian coast from Cape Sable to the Gut of Canso than on Georges Bank or on Browns, though a few are brought in from the various fishing banks every summer (p. 357). But the amounts reported from the outer (Atlantic) coasts of Cape Breton are so large as to show that they are likely to be as numerous there as they are anywhere abreast of the Gulf of Maine, or off southern New England, and perhaps more con- centrated. These regional variations may be il- lustrated by the landings for 1945, which were as follows for United States and Canadian vessels combined: offing of southern New England, west- ward from Nantucket Shoals, about 242,000 pounds ; 67 near coast of eastern Massachusetts, probably one fish; 68 coast of Maine, 400 pounds; Bay of Fundy (including both shores), 0; Nan- tucket grounds and Georges Bank region (includ- ing South Channel grounds), about 800,000 pounds; off west coast of Nova Scotia and on west- ern part of Browns Bank, about 671,000 pounds; Nova Scotian coast and banks from eastern part of Browns to offing of Cape Canso, at the entrance to the Gut of Canso, about 219,000 pounds; outer (Atlantic) coast of Cape Breton,69 about 2,059,000 pounds. A few are harpooned on the Gulf of St. Lawrence shore of Cape Breton also; 600 pounds were re- ported there in 1936, 200 pounds (one fish?) in 1943, and 1,000 pounds (4 or 5 fish) in 1946. The only other definite report of swordfish in the Gulf of St. Lawrence that has come to our notice is from Bonne Bay, on the west coast of Newfound- land, where Wulff n saw one, and had a strike from it (he did not hook the fish). But some few are seen and harpooned on the Grand Banks, and also along the south coast of Newfoundland, most often along the stretch between Port au Basque, on Cabot Strait, and Hermitage Bay. Here, writes Wulff, they sometimes come so close inshore that they "have been harpooned from the small wharfs, from shore, and from dories in the almost landlocked harbors," which we have never known to happen in the Gulf of Maine. " Rept. U. S. Comm. Fish. (1880) 1883, pp. 298-394, pis. 1-24. •! Rich, Proc. Portland Soc. Nat. Hist., vol. 4, Pt. 2, 1947, p. 71. » See Rich (Proc. Portland Soc. Nat. Hist., vol. 4, Pt. 2, 1947, pp. 55-82) for a more detailed survey of the distribution of swordfish on Georges, Browns, and the Nova Scotian Banks. WThe weights given are dressed; live-weights would be about 1M times as great. 88 Forty pounds reported, but this probably is an error, for it is not likely that a swordfish that small was taken there. M Victoria, Cape Breton and Richmond Counties, Nova Scotia. '• fnternat. Game Fish Assoc. Yearbook, 1943, p.6ti. 356 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Swordfish are summer fish on the North Ameri- can coast like the tuna, and their presence (often reported) in the blue water72 between the outer edge of the continent and the inner edge of the Gulf Stream proper, ofT southern New England and the Gulf of Maine, added to the fact that few are seen along the coast south of New York, makes it likely that they come in from offshore, direct. They appear about simultaneously off New York, off Block Island, on Nantucket Shoals, and on Georges Bank, sometime between the 25th of May and the 20th of June, but seldom on the Scotian Banks until somewhat later, or in the inner parts of the Gulf of Maine before July. They are most numerous in July and August, and they van- ish at the approach of cold weather. None have ever been reported east of Cape Cod after the first half of November, so far as we can learn (in 1875 one was taken on Georges in November in a snowstorm)73 and most of them are gone by the last week in October, though some fish have been taken off New York and New Jersey in December and even in January. A case in point is that 13 were entangled in long lines set for tilefish in 95 to 125 fathoms off Long Island between De- cember 20, 1921, and January 1, 1922.74 General report has it that the fish caught early in the season average not only thinner but con- siderably smaller than those caught later, a phe- nomenon still awaiting satisfactory explanation.76 Nearly all the swordfish that visit us weigh upwards of 50 to 60 pounds. One of 6 pounds 7 ounces, taken by the schooner Anna on Georges Bank, August 9, 1922 (now or formerly to be seen at the Boston Fish Bureau); a second of 7 pounds, taken by the schooner Courtney on a long line, on Browns Bank in 1931; a third of 1){ pounds, taken on a long line by the Dacia on Western Bank, early in September 1931; and a fourth 28K inches long to tip of lower jaw (its sword was broken off short), weighing 5% pounds, caught by the trawler Winchester, August 15, 1951, on the south- east part of Georges Bank in a haul which fished at 46 fathoms76 are the smallest Gulf of Maine and Nova Scotian specimens of which we have heard. Goode77 also reported a sword, only 3% inches long, found sticking in the nostril of a mackerel shark caught at Gloucester, probably picked up somewhere off southern New England for this particular shark does not ordinarily range farther south than that.78 In the Mediterranean, however, young fry as small as half a pound are often brought to market. It is generally believed that swordfish come di- rectly in from the open seas when they appear on the offshore banks in spring; a few to enter the Gulf of Maine, but the majority to work slowly eastward along the outer part of the continental shelf. When they depart in autumn it is to return to the open Atlantic, but they are never seen on then journey offshore, or southward; they simply drop out of sight as the tuna do. No ripe fish, male or female, have ever been seen off our coast. The ovaries and spermaries of most of those examined79 have shown no signs of ap- proaching maturity; most of the fishermen, too, of whom we have inquired have assured us that they have never seen "spawn" in swordfish, though they had dressed hundreds. And while the captures of 3 fish with ovaries containing eggs in early stages of development have been reported, one brought into Provincetown in September 1909, a second with ovaries weighing 15 pounds, brought to New Bedford on June 25, 1922,80 and a third of about 150 pounds killed off Marthas Vineyard in July 1924, events of this sort are so unusual that they cause wide comment. Evidently the swordfish that summer off our coasts spawn during the part of the year when they are elsewhere; probably in the subtropical parts of the Atlantic basin, for Lutken 81 found swordfish fry as small as 10 mm. (evidently hatched only a short time previous) between the latitudes of 20° and 39° N. The fact that the fish are thin when they return to us in spring, but fatten during the summer stay, is further evidence that they are spent before they appear off our coasts. Abundance. — Our only clue to the numbers of swordfish that visit our waters is the poundage '2 This Is often spoken of as the "Gulf Stream"; Its more accurate name Is the "slope water." « Rich, Proe. Portland Soc. Nat. Hist., vol. 4, Part 2, 1947, p. 58. »< Townsend, Science, N. Ser., vol. 56, 1922, pp. 18-19. » Rich (Proc. Portland Soc. Nat. Hist., vol. 4, Pt. 2, 1947, p. 43) Is "Inclined to think" that there are two or more "distinct year-schools" In our waters. " Reported to us by George Kelley of the U. S. Fish and Wildlife Service. The specimen is in Its collection In Woods Hole. " Ftsh. lnd. U. S., Sect. 1, 1884, p. 348. " See Rich (Proc. Portland Soc. Nat. Hist., vol. 4, Pt. 2, 1947, pp. 37-39) for additional records of small Gulf of Maine swordfish. 76 Many have been opened with this point In mind; some by us. «o Townsend, Science, N. Ser., vol. 60, 1922, pp. 18-19. » Spolla Atlantica, In Kong. Danske Vidensk. Selsk. Skrlft, Ser. 5, Nat. Math. Sect., vol. 12, No. 6, 1880, pp. 444-445. FISHES OF THE GULF OF MAINE 357 landed yearly. The smallest year's catch reported as landed at Portland, Gloucester, and Boston, within the period 1904 to 1929 was 883,000 pounds (in 1919), the largest 4,593,000 pounds (in 1929), the average about 2,000,000 pounds, or anywhere between 4,000 and 18,000 fish per year. And the landings in New England ports ran from 1,715,000 to 5,070,000 pounds during the decade 1930 to 1939 for southern New England and the Gulf of Maine. The interruption of swordfishing by Ger- man submarines and by the diversion of manpower was reflected in much lower landings during the first two years of the war, as was to be expected.82 But swordfishing picked up again after the war, to landings of about 1,250,000 pounds for southern New England and the Gulf of Maine, including western Browns Bank, in 1944 (New England and Canadian landings combined), to about 2,850,000 pounds in 1945, to about 2,500,000 pounds in 1946,83 and to something like 2,000,000 pounds in 1947.84 A catch of somewhere between 2 million and 3 million pounds would be a reasonable expectation for southern New England and the Gulf of Maine combined in average years. The catch off Cape Breton, eastern Nova Scotia, has run between 1% and 3 million pounds of late years (1939-1946), averaging a little more than 2 million until in 1947, when it fell to about 770,000 pounds.86 The Nova Scotian catches were not lessened by the submarine menace during the war years. It is not known what percentage of the total number of swordfish off our coasts is represented by the catches. But, at least, they do not suggest that any extreme ups and downs took place prior to 1947. Importance. — Appreciation of the swordfish as a food fish is of rather recent growth. Down to the middle of the past century it was unsalable in Boston and brought a very low price in New York, but of late years the demand woidd have taken care of a much greater supply than has been brought in. In 1919, the price to the fishermen averaged about 24 cents; in 1928, 22 cents; and 18 cents per pound in 1929 when a large catch was made. In 1945 it brought between 40 and 42 cents; and it rose to about 60 cents in 1946, but fell again to about 40 cents in 1947. Practically all the swordfish brought in to market are harpooned; we have never heard of one caught in net or seine, nor is it likely that any net now in use would hold a large one. Sword- fish have also been taken from time to time on hand lines and on long lines baited for cod or halibut with mackerel or other fish (p. 353). But the numbers caught in these ways have never been large enough to figure to any extent in the total catches, and are not likely to be.86 Occa- sional swordfish have been caught by anglers of late years, on rod and reel, and sport fishermen would agree that a good-sized broadbill is the premier prize of the sea. THE SPEARFISHES OR MARLINS AND THE SAILFISHES. FAMILY ISTIOPHORIDAE The spearfishes and sailfishes, like the sword- fish, have a sword formed by the prolongation of the snout and upper jaw. But their sword is rounded toward the tip, not flattened, and narrower than that of the swordfish. Their bodies, too, are closely clothed with narrow lanceolate scales, pointing rearward in general and embedded in the skin, either wholly or with their sharp tips pro- jecting slightly (fig. 188), and their first dorsal fin is much longer, occupying the greater part of the back behind the nape, and it can be depressed ,a Landings were only about 545,000 pounds In Massachusetts and 7,000 pounds In Yarmouth County. Nova Scotia in 1942; about 479,000 pounds in Massachusetts and about 17,300 pounds in Yarmouth County In 1943. •' Most recent year for which the landings have been published for the Canadian coast of the Gulf of Maine and for the ports in New England. 84 The Canadian catch statistics for 1947 have not reached us yet. 11 Information from Dr. A. H. Leim of the Fisheries Research Board of Canada. into a groove along the back. They fall in two groups, sailfishes with very large, sail-like dorsal fin, and spearfishes or marlins with lower dorsal. The sailfish (Istiophorus americanus Cuvier and Valenciennes 1831), so common in the wanner parts of the Atlantic, is included in the following Key because it has been taken at Woods Hole on several occasions, though not yet recorded from the Gulf of Maine. It is readily recognizable by the fact that the first dorsal fin is much higher than that of the marlins while the ventral fins of the sailfish are 2- or 3-rayed instead of being reduced to a single spine, as in the marlins. The two dorsal fins of the sailfish have usually been described as connected even in the adult. This, » Rich (rroc. Portland Soc. Nat. Hist., vol. 4, Pt, 2, 1947, pp. 67-82) gives an Interesting account of the methods of the New England swordfishery. 358 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE in fact, is given as the chief distinction between it and the marlins by Goode87 by Jordan and Evermann,88 and by Boulenger.89 But there is actually a considerable gap between the two fins in large specimens as Bean90 remarks and as appears on Goode's own illustrations of a sailfish taken at Newport, and of a skeleton. Two species of marlins, the blue and the white are known off the middle and north Atlantic Coasts of the United States. But it is not yet clear whether the enormous marlins, with violet cross-stripes on the sides, that are caught off the North Coast of Cuba91 are simply very large blue marlin, a separate subspecies, or even a species. And the marlins of more southern waters still await critical study. KEY TO SPEARFISHES OR MARLINS, AND SAILFISHES Middle Atlantic and North Atlantic Coast of United States 1. First dorsal fin much higher than the body is deep and sail-like; ventrals of 3 rays each.. Sailfish, p. 357 The first dorsal fin is not higher than the body is deep; ventrals reduced to one spine each 2 2. Apex of first dorsal fin and tips of pectorals pointed. Blue marlin, p. 358 Apex of first dorsal and tips of pectorals rounded White marlin, p. 360 Blue marlin Makaira ampla (Poey) 1860. Skilligalee Jordan and Evermann, 1896-1900, p. 892, Telrapterus amplus." Description. — The marlin parallels the swordfish in the prolongation of the bones of the upper jaw to form a sword. But that of the marlin is slender and rounded above, not broad and flattened as in the swordfish, also only about half as long relatively. The two fish differ widely, too, in the '» Rept.U.S.Comm. Fish., 1880, p. 296. « Bull. 47, U. S. Nat. Mus., Pt. 1, 1896, p. 890. " Cambridge Natural History, vol. 7, 1904, p. 680. » Bull. New York State Mus., 60, Zool. 9, 1903, p. 404. 81 Hemmlngway (In Vesey-Fitzgerald and Lamonte, Game Fishes of the World, 1949, p. 158) reports these striped marlins weighing up to 1,250 pounds off northern Cuba. M Jordan and Evermann in their general review of the giant mackerel-like fishes, tunnies, spearfishes, and swordfishes (Occ. Papers, Calif. Acad, of Scl., XII, p. 28, 1926) separate the spearfishes into two genera Telrapterus with the front of the first dorsal fin little if any higher than the median part of the fin and Makaira, with the front part of the first dorsal higher than the median part of the dorsal. relative sizes of the first dorsal fin,93 which extends along fully two-thirds of the length of the trunk from the nape backward in the marlin and is, futhermore, of characteristic falcate outline. But more important systematically, if less ap- parent, is the fact that the adult marlin has ventral fins which the swordfish lacks, though they are reduced, it is true, to one long spine each (actually 5 spines fused together). Futhermore, the second dorsal fin and the second anal fin of the marlin are relatively larger, and the pectorals smaller than those of the sword- fish, while there are two small longitudinal keels on either side of its caudal peduncle instead of one broad one; its body is more slender; and its head is relatively shorter. Careful examination would show that the spearfish is not naked but has small scales imbedded in the skin and that there are small teeth in its jaws and on the roof of its mouth. The blue marlin differs from its close relative the white marlin in the shape of the apex of its first dorsal fin and of the tips of its pectorals, both of which are pointed; also in the much darker color of its sides and belly; and in the fact that it grows much larger. The blue marlin is deepest abreast the pectorals, (about 6/4" times as long, not counting the caudal fin, as it is deep), tapering evenly to the caudal peduncle, and its upper jaw in front of the eye (including the sword) is about twice as long as the length of its head behind the eye.94 The first dorsal fin, with 47 to 48 stiff rays is separated from the second dorsal by a space about as long as the latter in one fish seen by us; by a shorter space in another. The first anal fin (2 spines and about 12 or 13 rays), situated below the rear part of the first dorsal, is triangular, its long first rays forming a sharp angle. The short second anal is similar to the second dorsal fin and originates a little in front of the latter. The ventrals stand below the pectorals; the caudal resembles that of the swordfish in its lunate outline. Color. — Dark dull blue on the back and on the sides down about to the level of the eyes, washed with coppery reflections, also on the bill, with 93 Very young marlins have only one continuous dorsal fin, but this separates later into two. M See Shapiro (Amer. Mus. Novitat., No. 995, 1938) for a study of the changes in proportional dimensions that take place with growth, in the blue marlin. FISHES OF THE GULF OF MAINE 359 Figure 188. — Blue marlin (Makaira ampla). Drawing by Jessie Sawyer, based on Bahama and Marthas Vineyard specimens. Left, a piece of skin from the upper part of the side of the Marthas Vineyard speci- men, with epidermis scraped off to show scales, about twice natural size. rather abrupt transition to much paler gray-blue lower down the sides and on the lower surface, the belly being as dark as the lower part of the sides; the sides cross-marked with about 13 indistinct violet-blue stripes, about 1 to 1% inches wide on a fish 8 feet long, showing pale against the dark blue of the upper parts of the body, but dark against the paler blue of the lower part of the sides. First and second dorsal fins, pectoral and ventral fins, and first anal fin dark rather vivid blue. Caudal fin of about the same color as upper part of trunk; second anal fin of same pale gray-blue as the belly.85 Size. — Blue marlins run fully as large as sword- fish. Reports are current of fish of 1,000 pounds being harpooned; the rod and reel record is 742 pounds.96 Many weighing more than 500 pounds are caught off the north coast of Cuba and on the Bahamas side of the Straits of Florida every year,97 and one taken on the southern part of Browns Bank, weighed 575 pounds dressed, when landed, 11 Description based on a "blue" about 8 feet long from tip of bill to fork of tail, and weighing 169 pounds, fish taken near Bimini, Bahamas, June 1941, by R. W. Foster, mounted by the well-known fish taxidermist, H. Pfleuger of Miami, Fla., and now in the Museum of Comparative Zoology. " Caught at Bimini, Bahamas, June 19, 1949, by Aksel Wichfeld. " See Farrington (in Vesey-Fitzgerald and Lamonte, Game Fish of the World, 1949, p. 154) for a readable account of the blue marlin of Bahaman waters as a game fish. or about 700 pounds alive. A very large one may measure as much as 15 feet,98 but the rod and reel record fish, mentioned above, was only 12 feet 10% inches long. Another fish caught in the Ba- hamas weighed 650 pounds (not dressed), and measured 12 feet 1 inch; a third, of 621 pounds was 12 feet 3 inches long.99 General range — Warm parts of the northwestern Atlantic, straying northward to the Gulf of Maine. It has been reported near Sable Island, but the very small specimen in question may have been a white marlin (p. 360). Occurrence in the Gulf of Maine. — This southern warm-water fish was reported from the South Channel, between Georges Bank and Nantucket Shoals, between 1877 and 1880, by the fishing schooner Phoenix. No other marlins that we can be sure were blues were reported within the limits of the Gulf of Maine until September 5, 1930, when a small one 6 feet 10 inches long,1 was har- pooned on the southern part of Browns Bank. And a very largo one was caught in that same vicinity by the Col. Lindbergh the following July, " The blue marlin Is said to reach 26 feet, but we think this much exagger- ated. » Reported to us by Frank Mather, of the Woods Hole Oceanographic Institution. 1 This specimen is in the Museum of Comparative Zoology. 360 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE and brought into the Boston Fish Pier. A marlin about 5 feet long was taken on Georges Bank by the schooner Ethel Merriam, on August 5, 1925, but this may have been a white (p. 359). Blue marlins are sighted at long intervals off Marthas Vineyard. And fishermen report them now and then along the southern edge of Georges (aDy very large marlin is a blue) but do not har- poon them, for they have no market value. They are game fish par excellence, and much sought after off Cuba and in the Bahaman side of the Straite of Florida. They also support a consider- able commercial fishery off the north coast of Cuba.2 White marlin Makaira albida (Poey) 1860 Jordan and Evermann, 1896-1900, p. 892. Telrapierus imperator (Bloch and Schneider) 1801, in part. Description. — The white marlin differs from its larger relative the blue marlin in its rounded first dorsal and pectoral fins, in the pale color of the lower part of its sides, and in its white belly (p. 359) ; and in its smaller size. Few grow larger than 125 pounds; the rod and reel record stands at 161 pounds.3 This fish was 8 feet 8 inches long. General range. — Western North Atlantic; com- mon in Cuban and Bahaman waters, and off south- ern Florida; north regularly in summer to the offing of Delaware Bay in abundance, and to southern New England waters in lesser numbers. Occurrence in the Gulf of Maine.- — So many white marlin come northward, as far as New York waters that about 500 were taken off Montauk, Long Island, on rod and reel during the 11 years 1925- 1936, and more than 150 in 1935 alone.4 And a few are caught off the southern Massachusetts Islands in most summers. But their usual turning point is west of Nan- tucket. True, Farrington6 speaks of "great quantities" of them as seen on Georges Bank; but we cannot find that any marlin caught there has been identified positively as a white, though one about 5 feet long taken on August 5, 1925 (p. 359) may perhaps have been one. The meager record suggests that they may stray oftener to outer Nova Scotian waters, for a 5-foot fish weigh- ing 21 pounds, caught on Sable Island Bank, August 18, 1931, probably was a white marlin, while Farrington reports one harpooned off Glace Bay near Sidney, in 1945, and others sighted off Halifax that same year. Figure 189. — White marlin (Makaira albida). From Goode. THE DOLPHINS. FAMILY CORYPHAENIDAE The dolphins (two species are known) are moderately slender and flattened sidewise, with slightly projecting lower jaw, a massive blunt head, a long, rather high dorsal fin without spines, extending from close behind the head to near the base of the caudal fin, an anal similar to the dorsal in shape but shorter, and a widely forked tail. They have small comb-like teeth in the jaws and on the roof of the mouth. • Farrington (in Vescy-Fltzgerald and Lamonte, Game Fishes of the World, 1949, p. 153) gives an interesting account of this fishery. » One caught off Miami, Fla., Mar. 20, 1938, by L. F. Hooper. Common dolphin Coryphaena hippurus Linnaeus 1758 Jordan and Evermann, 1896-1900, p. 952. Description. — The dolphin differs from related fishes in that its long tapering body is most massive and deepest close behind the head, and that its dorsal fin, originating over the gill cover, extends back nearly to the base of its deeply * Information supplied by Frank Mather of the Woods Hole Oceanographic Institution. ' In Vesey-Fitzgerald and Lamonte, Game Fish of the World. 1949, p. 155. FISHES OF THE GULF OF MAINE 361 Figuee 190. — Dolphin (Coryphaena hippurus). From Goode. Drawing by H. L. Todd. forked tail fin. These characters, with its side- wise flattened form, notably steep forehead, deeply forked tail, and large ventral fins, separate it at a glance from the few other Gulf of Maine fishes which have long dorsal fins with bodies ,/hat are deepest forward. Its anal fin, 26 to 30 iays, originating about midway of its body, is about half as long and half as high as the dorsal which has 55 to 65 rays. The lobes of its deeply forked tail are long and slender. Its moderately long ventrals and pectorals are situated the one below the other. Color. — The dolphin is famous for its brilliant hues and for the vivid waves of color that flash across it when first taken from the water. Alive, in the sea, its sides are largely vivid blue, variously mottled and washed with gold; its tail largely golden yellow. Size. — Maximum length about 6 feet. Habits. — The dolphin, despite its blunt snout, is one of the swiftest of fishes. Voyagers on tropic seas often see them leaping in pursuit of small fry, or when pursued themselves by larger fishes. In sailing ship days dolphins were often caught by trolling from the stern. Offshore, they feed largely on flying fish; the Sandwich specimen mentioned later had some silversides in its stomach. General range.- — Cosmopolitan in warm seas; northward along our Atlantic Coast to southern New England, where it is rare inshore, occasionally straying as far as the outer coast of Nova Scotia. Occurrence in the Gulf oj Maine. — A dolphin about 3'^ feet long (now in the collection of the Boston Society of Natural History) and weighing 23 pounds, taken 60 miles south-southwest of Cape Sable, in the deep gully between Browns and Georges Banks by the trawler Natalie Ham- mond, August 15, 1930, was the first record for the Gulf of Maine; a second was taken in a trap at North Truro on Cape Cod Bay, in August 1949 6 (a season when many were taken off Marthas Vineyard) ; a third at Sandwich, on the southern shore of Cape Cod Bay in mid-July 1951.7 THE SEA BREAMS OR POMFRETS. FAMILY^BRAMIDAE The sea breams are usually considered the most nearly related to the dolphins. But they rather suggest the butterfishes (Family Stomateidae, p. 363) in general appearance, with single, long, falcate soft-rayed dorsal fin; anal similar to the dorsal; lunate tails, very small ventrals; and deep, sidewise flattened bodies. They are to be expect- ed only as strays in the Gulf of Maine. Johnson's Sea Bream Taractes princeps (Johnson) 1863 Johnson, Proc. Zool. Soc. London, 1863, p. 36. Description. — This sea bream is unique among Gulf of Maine fishes in its general appearance. • Reported by Schuck, Copeia 1951. p. 171. ' We saw this specimen in the collection of the U. S. Fish and Wildlife Service at Woods Hole. 362 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Figure 191. — Johnsons sea bream (Taractes princeps), Browns Bank. After Bigelow and Schroeder. In the adult the body is massive, flattened side- wise, about half as deep as it is long to the base of the tail fin. The dorsal and anal fins are long, scythe-shaped in front, each followed by a row of low rays that are detached along their outer parts, but are joined by scaly skin along their bases, giving the fins the outline shown in figure 191. The tail fin is deeply lunate; the long pectorals originate a little in advance of the dorsal, and the very small ventral fins a little in front of the pectorals. The eyes are large, oval, with vertical di- ameter longer than the horizontal diameter. There is no lateral line. A striking character is that the scales winch clothe the vertical fins and the body (about 43 scales along the median longitudinal row) vary greatly in size, being largest along the sides, smallest on the back, breast, and fins. They vary also in shape, their exposed margins being either concave, convex, notched, or straight. This species is separated from Brama raii Bloch 1781,8 the only one of its relatives yet recorded from our North Atlantic coast, by its larger scales {Brama raii has 80 or more in the median longitudinal row) and by the fact that its ventral fins originate slightly, but distinctly, in front of the pectorals. Color. — The body and head of a specimen, three days after death, were blackish, tinged with salmon on the gill covers and along the sides; the dorsal and anal fins were dusky, with the free ends of the short rays pale; the caudal was black but with its concave margin white; the pectorals were gray. Size. — Maximum length about 3 feet. General range. — Known only from Madeira in the eastern Atlantic, and from Browns Bank in the western. Occurrence in the Gulf of Maine. — This fish is mentioned here on the basis of one specimen, about 33 inches long, caught on a long line on Browns Bank, off Cape Sable, in January 1928, by the schooner Wanderer. A detailed account and comparison with allied species is given by Bigelow and Schroeder.9 It seems certain that Taractes is very rare in American waters, at least in the depths in which commercial fishermen operate, for so conspicuous a fish would almost certainly be reported, if caught. Nothing is known of its habits except that it seems to be common around Madeira in deep water. 8 Brama rail has been taken at Woods Hole and on the Grand Banks. • Bull. Mus. Comp. Zool., vol. 69, 1929, pp. 39-50, 1 pi. FISHES OF THE GXJLF OF MAINE 363 THE BUTTERFISHES. FAMILY STROMATEIDAE The members of this family are deep bodied and very much flattened sidewise, with one long dorsal fin that is soft rayed except for a few short weak spines at its forward end, an anal fin of corre- sponding size and shape, a deeply forked caudal fin, a blunt nose, and a small mouth. The two species that occur on the east coast of North America lack ventral fins, but the extremity of the pelvic bone projects through the skin as a spine but this is so short that it is likely to be over- looked unless felt for. Two species occur in the Gulf of Maine: one (the butterfish) a common summer visitor, the other (the harvestfish) a rare stray from the south. KEY TO GULF OF MAINE BUTTERFISHES 1 . The forward one-fourth of the anal fin is only about 2 or 3 times as high as the rear portion of the fin. The margins of the anal and dorsal fins are only slightly concave in outline Butterfish, p. 363 The forward one-fourth of the anal fin is at least seven times as high as the rear portion of the fin. The anal and dorsal fins are both very deeply concave in outline Harvestfish, p. 368 Butterfish Poronotus triacanthus (Peck) 1800 Dollarfish; Shiner; Skipjack; Sheepshead; Harvestfish Jordan and Evermann, 1896-1900, p. 967, as Rhombus triacanthus. Description. — The most distinctive characters of the butterfish are its very thin deep body, like a flounder on edge; the fish is only about twice as long as it is deep to the base of its tail fin (the only common Gulf of Maine species of this shape), combined with a single, long, soft-rayed dorsal fin, an anal fin almost equally long, and a deeply forked tail, but no ventral fins. The absence of ventral fins separates it from the pompanos; the dorsal without obvious spines from the scup (p. 411) and John Dory (p. 297); the lack of detached dor- sal spines from the triggerfishes, which are, fur- thermore, very different in general aspect (p. 520). And it is easily distinguishable from its relative, the harvestfish (p. 194), which is rare in northern waters, by its much lower dorsal and anal fin (compare fig. 192 with fig. 194). The dorsal fin (about 45 rays) originates close behind the axils of the pectorals and tapers at first abruptly and then gradually backward, while the anal (about 40 rays) narrows evenly from front to rear. There is a forward-pointing spine close in front of the dorsal fin, so short as hardly to be visible though it can be felt; also 3 very short spines in front of the anal, almost wholly embedded in the skin, the first of which points forward. Both the dorsal fin and the anal extend rearward almost to the base of the caudal fin. Distinctive, also, are the long pointed pectoral fins, short head, blunt snout, small mouth, weak teeth, and the short and slender caudal peduncle, which does not have longitudinal keels. The scales are very small, and are easily detached when the fish is handled, and there is a row of very conspicuous mucous pores below the forward half of the dorsal fin. Color. — Leaden bluish above, pale on the sides, with numerous irregular dark spots which fade after death. The belly is silvery. Size.' — The largest are about 12 inches long; the general run are about 6 to 9 inches long. The weight runs about 1% ounces at 6 inches, 4 to 4% ounces at 8 inches; about 1 pound at 11 inches (if fat). The largest weigh about lVt pounds. Habits. — Astonishingly little is known of the manner of life of the butterfish considering how familiar and valuable it is. As a rule they travel in small bands or loose schools; and draggers re- port catching several times as many by night as by day, suggesting that they are active enough to dodge a trawl, except during the hours of darkness. They often come close inshore, into sheltered bays and estuaries, hence their frequent capture in pound nets. And it shows so decided a preference for sandy bottoms rather than for rocky or muddy, that few are taken in traps on muddy ground while other traps along the sandy beach nearby may yield considerable numbers. General experience is that the butterfish keeps chiefly near the sur- face during its stay near the coast, and schools are often to be seen. At Cohasset (on the south side of Massachusetts Bay), for instance, schools of butterfish, fifty to a few hundred, are often to be seen where the flats are covered by only 4 or 5 feet 364 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE 4t& : Figure 192. — Butterfish {Poronolus triacanthus) , New Jersey. From Goode. Drawing by H. L. Todd. of water. Though definite evidence is lacking, we believe butterfish seldom descend deeper than 15 to 30 fathoms during the summer, and that most of the fish caught by the otter trawlers on the Nantucket grounds and on Georges Bank in sum- mer are picked up by the trawl on its way up or down, not while dragging on bottom. In fact, mackerel fishermen often take a few butterfish on Georges in their purse seines. But such evidence as is at hand is to the effect that they spend the winter and early spring near bottom, and in depths down to 100-115 fathoms (p. 367). Food. — The butterfish feeds on small fish, squid, Crustacea such as amphipods and shrimp, and annelid worms. And ctenophores have been found in butterfish stomachs at Woods Hole, though these watery objects are not a regular item in its diet. Breeding habits. — Butterfish begin spawning in the Gulf of Maine in June, soon after their arrival. The height of the reproductive season is in July and their eggs have been taken throughout August. Observations at Woods Hole suggest that butter- fish spawn some few miles out at sea, returning to the coastwise waters when they are spent.10 We have taken its eggs in our tow nets at several stations in Massachusetts Bay, and it would not be astonishing to find them anywhere off the New England and western Nova Scotian coasts or on the Scotian side of the Bay of Fundy, Huntsman having found large spawning individuals in St. Mary Bay in July. But despite the considerable number of butterfish eggs that are produced in the Gulf of Maine, we doubt whether the latter is a favorable nursery for this fish, for we have taken its larvae only twice there (off Cape Cod on August 16 and on Georges Bank on July 23, 1916) a total of only 3 specimens, 5 to 30 mm. long, although we have made hundreds of hauls widely distributed inshore as well as offshore at the season when they might be expected. Neither have young butter- fish been reported from the Bay of Fundy. But- terfish fry are very plentiful, however, along the shores of southern New England. The eggs are buoyant, transparent, spherical, 0.7 to 0.8 mm. in diameter, usually with a single oil globule of about 0.17 to 0.2 mm. In newly spawned eggs, however, there may be two globules, which coalesce as development advances." At a temperature of 65° F. (about the summer state of the surface of Massachusetts Bay) incubation occupies less than 48 hours. And it is probable that development can only proceed in compara- tively warm water, though the lower temperature limit to successful reproduction is not known. The larvae are about 2 mm. long at hatching and they are characterized shortly after by their short deep form, by their 30 muscle-segments, and by the row of black spots along the ventral edge in the ■• Kuntz and Radcliffe, Bull. V. S. Bur. Fish., vol. 35, 1918, p. 112. II A large series of butterfish eggs artificially fertilized at the Gloucester hatchery have been available for comparison with the pelagic eggs taken in the tow nets. FISHES OF THE GULF OF MAINE 365 post anal region.12 The dorsal, anal, and caudal fin rays are visible in larvae of 6 mm., when the body has already begun to assume the deep thin form so characteristic of the adult butterfish. At a length of 15 mm. the caudal fin is deeply forked, the dorsal and anal fins are formed, and the little fish resembles the adults sufficiently for ready identification. Figure 193. — Butterfish (Poronotus triacanthus) . A, egg; B, larva, 6 mm.; C, fry, 15 mm. After Kuntz and Radcliffe. During the first summer young butterfish often live in the shelter of the large jellyfishes as young haddock do, and Goode 13 graphically described the fry of 2 to 2% inches as swimming among the tentacles of the red jellyfish (Cyanea), sometimes 10 or 15 little fish under one jellyfish, where they find protection from larger fish, but to which they sometimes fall prey. This association, however, u Information furnished by O. E. Sette. The illustrations of larvae 2.1 mm. and 3.4 mm. long credited by Kuntz and Radcliffe (Bull. U. S. Bur. Fish., vol. 35, 1918, figs. 63 and 64) to the butterfish and reproduced in the previous edition of this book (Blgelow and Welsh, Bull. U. S. Bur. Fish. vol. 40, Pt. 1, 1925, fig. 116, c and d) have since been pioved to belong to one of the hakes (Urophycis). " American Fishes, 1888, p. 222. is not essential to their welfare, for fry are often seen living independently at the surface, particu- larly in sheltered bays west and south of Cape Cod. On one occasion in late August 1925, on Nantucket Shoals, we observed numbers of .young butterfish 1-1% inches (2G to 39 mm.) long swim- ming free in the upper stratum of water. And we have seldom found young butterfish with the many Cyanea that we have captured in the Gulf of Maine. It seems that the fry hatched earliest in the season grow to a length of 3 to 4 inches by autumn, great numbers of that size having been taken in Rhode Island waters in October. But late- hatched fish probably are not more than 2 to 3 inches long at the beginning of winter, and they can grow little during the cold season, for little fish of 3 to 5 inches are seen again in the spring. A series of measurements made by Welsh at Atlantic City, N. J., in August 1921, throws some light on the subsequent rate of growth. The fish fell into two groups: one ranging from 4 to 5% inches (averaging about 4%) and the other from 7% to 10% inches. Very likely those of the first group (which were much the more numerous) were in their second summer, for Hildebrand and Schroeder I4 record a growth of from 4 inches to 5% inches from May to October in Chesapeake Bay; those of the second size group were in their third summer, some perhaps in their fourth. It is probable that the butterfish matures when 2 years old, and upward of 7 inches long. General range.- — -Atlantic coast of North America from the offing of South Carolina and from coastal North Carolina waters to the outer coast of Nova Scotia and Cape Breton; northward as a stray to the Gulf of St. Lawrence 15 and to the south and east coasts of Newfoundland;16 southward to Florida in deep water. Occurrence in the Gulj of Maine.— This is a reg- ular summer visitor to the Gulf of Maine, locally abundant along the shores of Massachusetts, less common along the coast of Maine. Butterfish are common also in some years along the Nova Scotian coast of the Gulf; great numbers were caught in " Bull. U. S. Bur. Fish. vol. 43, Pt. 1, 1928, p. 214. " Hoar (Copeia, 1937, p. 238) records two large ones from Margaree Harbor on the Gulf of St. Lawrence shore of Cape Breton, and cites an earlier record for the coast of Quebec. '• It is reported from Rose Blanche on the south coast of Newfoundland, and from Bulls Bay and Ferryland on the east coast of the Avalon Peninsula (Rep. Newfoundland Fish. Res. Commission, vol. 1, No. 4, 1932, p. 108, and vol. 2. No. 1, 1933, p. 125. 366 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE St. Mary's Bay, for example, in 1910-1913 and again in 1938, though few were taken during the intervening years.17 But they appear only irregu- larly and in small numbers on the New Brunswick shore of the Bay of Fundy, though they have been taken repeatedly in Passamaquoddy Bay. The diminution in the numbers of butterfish, following from south and west to east and north around the coast line of the Gulf may be illus- trated by catches for 1938 a fairly representative year 18 when catches in pound nets and floating traps around the shores of Barnstable, Plymouth, and Essex Counties, plus those landed in Boston and Gloucester by seiners and trawlers fishing off- shore, amounted to 943,500 pounds, whereas only about 18,000 pounds were reported from the entire coast from the Massachusetts line to and including the region of Casco Bay, and none at all from farther east than that along the coast of Maine. Butterfish also appear in the Nantucket Shoals region and on Georges Bank in summer, often in good numbers. About 1,000 fish, for example, were caught on Georges during one trawling trip in 1913; and otter trawlers accounted for nearly two-thirds of the total landings for Massachusetts in 1938, about one-half of those for 1945, most of which probably came from these offshore grounds. We have heard no rumor of them on Browns Bank but doubtless they occur there, for "fair quantities" usually visit Halifax Harbor in summer and autumn, according to McKenzie,19 in fact, he cites one instance when about 1,500 of them were taken from two traps there in one day. And they are said to be common eastward as far as Canso.20 But this appears to be the normal limit to their range, for strays, only, have been taken in the Gulf of St. Lawrence (p. 365), or on the Newfound- land coast (p. 365). Season. — Butterfish are warm season fish along our coasts; we refer of course to the temperature of the water, not to that of the air. They may appear off Rhode Island by the last half of April and about Woods Hole by the middle of May, though they are not plentiful in the Woods Hole region until in June. And it is likely that these early comers move in across the shelf from off- shore, rather than that they have followed along " McKenzie, Proc. Nova Scotian Inst. Sci., vol. 20. 1939, p. 14. 18 This is the most recent year for which butterfish have been mentioned in the statistical breakdown by counties for Maine. " Proc. Nova Scotian Inst. Sci., vol. 20, 1939, p. 17. " Cornish, Contributions to Canadian Biology (1902-5) 1907, p. 85. the coast, for from April 8 to 12, 1953, the Eugene H trawled 22,000 pounds of butterfish, close to bottom, in 85 fathoms south to Martha's Vine- yard, and in 1950 the Albatross III trawled 10 to 723 butterfish per haul, May 11 to 18, along the 40-80 fathom zone off southern New England, where small commercial catches were also being made at the time. During the season of 1913 21 the first butterfish were reported on Georges Bank June 5 to 8. But it is not until the end of that month or early in July that they are plentiful anywhere north of the elbow of Cape Cod. The earliest catches, for example, in one set of traps off North Truro, on Cape Cod Bay, were not made until June 26-28th in 1947, or until July 29th in 1948, but on May 29, 1951. From that time on there are butterfish in the inner parts of the Gulf throughout the late summer and autumn, also on Georges Bank. The following tabulation of the catches made in one set of 8 traps at North Truro, on the east- ern shore of Cape Cod Bay,22 suggests that butter- fish are likely to be the most numerous there in August, at least in good years, and rather more numerous in September and in October than in July. But they are exceedingly irregular and un- predictable in then appearances and their disap- pearances. Thus the traps just mentioned yielded butterfish on only one day in July, 2 days in August, 3 days in September, and 3 days in October in the years 1948 and 1949 combined, though catches as great as 2,856 to 7,490 pounds were made on three of these occasions. The ap- proximate catches, in pounds, for the years 1946 through 1950 follow: Maximum Minimum Average Total July 5,900 0 1,760 8,810 August 53,101 0 11,450 57,260 September 15,100 90 5,850 29,250 October 26,440 120 8,425 42,130 In some years the peak for this locality may not come until October, as in 1947, when the catch by this set of traps was between five times and six times as great during that month (about 14,500 pounds) as during the next most productive month (July, about 2,300 pounds; August, about 2,500 pounds). Similarly, in 1950 the October catch 21 This is the only year for which lists are available of the number of fish of all species taken on Georges Bank by certain trawlers. " Information supplied by the Pond Village Cold Storage Co. of North Truro, Mass. FISHES OF THE GULF OF MAINE 367 of these traps was about 26,400 pounds following a peak in August (about 53,000 pounds). And they linger in numbers until well into November in the Cape Cod Bay region in some years; also on Georges Bank. Thus four or five traps at Provincetown yielded some 30,000 pounds during that month in 1915, while 2 traps at Barnstable, on the southern shore of Cape Cod Bay took 4,275 pounds of butterfish on November 17, in 1950.23 They may linger equally late into the season along the outer Nova Scotian coast in some years, as in 1938, when two traps at Halifax yielded about 1,500 fish on November 12th.24 They have been caught on Georges Bank until the end of that month; and in 1928 several hundred pounds were reported from Nantucket Shoals as late as the last week in December.25 But they all vanish from the coast by the end of December at latest, and usually earlier than that, not only from our Gulf but along the more southerly part of their range as well. It seems that the southern contingents simply move out to the outer edge of the continent into deeper and warmer water to winter, as the mackerel do also, for they are often caught by otter trawlers working out on the shelf between the latitudes of Chesapeake Bay and of Cape Hatteras in winter. The Albatross III trawled from 1 to 202 butterfish at a number of localities at depths of about 20 fathoms to at least 115 fathoms, between the offings of Charleston, S. C, and of Cape Hatteras in January and February of 1950. The case is not so clear for those that summer off southern New England and farther north and east. Butterfish, it is true, have been trawled in February near the 90-fathom line abreast of the eastern part of Long Island, N. Y. ;26 also late in March on the southwestern slope of Georges Bank (where the dragger Eugene H had the unusually large catch of about 15,000 pounds in 1951 in the last week of that month) and in April and in May off southern New England (p. 366). These, how- ever, may not have wintered in the vicinity, but may have been following along the outer part of the shelf northward, before turning shoreward toward their summer homes. " Information from John E. Vettorino, who operates these traps. « McKenzie, Proc. Nova Scotian Inst. Sci.. vol. 20, 1939, p. 17. » See Hildebrand and Schroeder, Bull. U. S. Bur. Fish., vol. 43, 1928, p. 215. for details as to their seasonal occurence in Chesapeake Bay. » Three fish taken by Albatross II, February 27, 1929. Abundance. — During the period 1928 to 1947 27 the reported catch of butterfish for Massachusetts ranged between 279,000 pounds and 2,250,000 pounds. Low points were in 1928 (about 580,000 pounds) and in 1946 (about 279,000 pounds); high, in 1932 (about 1,479,000 pounds), and during the period 1937-1940 (from about 1,226,000 pounds to about 2,250,000 pounds). And while this includes landings for the southern shore of the State as well as for the Gulf of Maine shore, the fluctuations that are indicated from year to year probably were paralleled north of Cape Cod. But the catch may be poor at any particular locality even in a good year, or vice versa. Thus the North Truro traps mentioned (p. 366) took only 1,230 pounds of butterfish in 1948, though this was a better-than average year for the Massachu- setts coast as a whole.28 If the fish caught average about one-half pound each, the Massachusetts fishery may thus be ex- pected to take somewhere between 560,000 and 4K million individual fish. But it is not known what proportion this may be of the total popula- tion of butterfish in the Gulf of Maine. Importance. — This is one of our best table fish, fat, oily, and of delicious flavor. Experience with many fresh from the net as well as on the table proves the old tale to be a myth that butterfish have a peculiar odor. However, they were often used to enrich land in planting during the first half of the past century, and appreciation of the fact that they are too good for this use is of recent growth. Even today the demand for butterfish in Boston is uncertain and the price widely vari- able. As late as 1938, 1,500 fish taken in traps at Halifax, Nova Scotia, were dumped for want of a market.29 The commercial catch is made mostly in pound nets, floating traps, purse seines, and otter trawls, and it was thought of old that they would never take a hook. But anglers have recently dis- covered that butterfish will sometimes bite a very small hook greedily, if baited with a bit of clam or with a small piece of a sea worm (Nereis). And 1,100 pounds were reported in 1945 as caught along the Massachusetts coast on hand lines. » Statistics are not available for 1929, 1934, 1936, or 1941. »■ Massachusetts catch, about 676,000 pounds. » McKenzie, Proc. Nova Scotian Inst. Sci., vol. 20, 1939, p. 17. 368 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Harvestfish Peprilus alepidotus (Linnaeus) 1766 Starfish Jordan and Evermann, 1896-1900, p. 996, as Rhombus paru (Linnaeus) in part. Description.- — The body of the harvestfish (not including the caudal peduncle) is almost as deep as it is long, and ovate in outline; its nose is rounded, mouth very small, and head very short. The outlines of its dorsal and anal fins afford the readiest field mark to separate it from its relative, the butterfish; both of these being very high and falcate in front, and continuing nearly straight, thence rearward (compare fig. 194 with fig. 192). The mucous pores, conspicuous in the butterfish, are lacking in the harvestfish. There is also a color difference between the two, the harvestfish being greenish silvery above, silvery sometimes tinged with yellow on its sides and belly, while the fins of some specimens are slightly dusky or yellow- ish. In all other respects (including size) it closely resembles the butterfish. General range. — From Florida northward along the middle Atlantic Coast of North America; rarely straying north to Cape Cod, Mass., and to Cape Elizabeth, Maine; represented by a close ally (Peprilus paru Linnaeus 1758) in West Indian- Brazilian waters.30 Occurrence in the Gulf of Maine. — This fish, al- though abundant to the southward, rarely strays as far north as the outer Cape Cod coast. A speci- men taken at Monomoy Point by Dr. W. C. Ken- dall in 1896; 5 or 6 caught in floating traps at Richmond Island, off Cape Elizabeth, Maine, in July 1929 ;31 one from the Damariscotta River, Maine, in August 1933 ;32 and one taken at Race Point at the tip of Cape Cod, in October 1949, 33 are the only Gulf of Maine records of which we know. a° See Meek and Hildebrand (Field Mus. Nat. Hist., Zool. series, vol. 15, Pt. 2, p. 411, 1925) for discussion. «i Reported to us by the late Walter H. Rich of the V. S. Bur. Fish. » MacCoy, Bull. 69, Boston Soc. Nat. Hist., 1933, p. 9. » Reported to us by Edgar Arnold. -^<\. fi' \- - f if -*tmZ '^:- Figure 194. — Harvestfish [Peprilus alepidotus), New York. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 369 THE RUDDERFISHES. FAMILY CENTROLOPHIDAE The closest affinities of the rudderfishes are with the mackerel-like fishes. They have moder- ately stout bodies, short blunt snouts with convex profiles, and a moderately deep caudal peduncle without longitudinal keels. The single dorsal fin extends from over the pectorals to the caudal peduncle; the front part of the dorsal is spiny, either reduced to a few flexible spines covered over by the skin so that it is hard to find them, or represented by several detached spines so short that they might be overlooked, and preceding the much longer soft-rayed part of the dorsal. The tail fin is only slightly emarginate; the anal fin is similar to the dorsal in shape but much shorter; the ventrals are below the pectorals, and are smaller than the latter. The mouth is small, with small teeth in the jaws. Only two species are known off the Atlantic coast of the United States. KEY TO GULF OF MAINE SPECIES 1. The single dorsal fin is preceded by 6-8 short detached spines; the sides of the head are scaly Barrelfish, p. 369 2. The dorsal fin is not preceded by any detached spines; there are no scales on the sides of the head Black ruff, p. 370 Barrelfish Palinurichthys perciformis (Mitchill) 1818 Logfish; Rudderfish; Black pilot Jordan and Evermann, 1896-1900, p. 964. Description. — The reduction of the spiny portion of the dorsal fin of the barrelfish to 6 to 8 short detached spines, with very small triangular fin membranes, closely followed by a long soft-rayed dorsal fin, marks the barrelfish from all other Gulf of Maine fishes, except for certain of the pompano tribe. The caudal fin of the barrelfish is only slightly emarginate instead of deeply forked and its caudal peduncle moderately stout and without keels instead of very slender. It suggest a tautog remotely in general appearance, especially in its rather stout body (about two-fifths as deep as long, not including the caudal fin), very bluntly rounded nose, convex forehead, and small mouth. But its rudimentary spiny dorsal fin and forked caudal fin are ready field marks to distinguish it. The soft dorsal fin (20 to 22 rays) arises about mid-way from tip of snout toward base of caudal fin; the anal (16 or 17 rays) somewhat farther back. Both these fins are moderately high and they taper slightly from front to rear. The anal is preceded by three short spines so nearly imbedded in the skin as to be hardly visible. Both the ventrals and the pectorals are large with rounded tips. The top of the head is scaleless but the sides of its head and the body are clothed with small rounded scales. The presence of the dorsal fin-spines and the scaliness of the sides of its head distinguish it from its close relative the black ruff (fig. 196). Figure 195. — Barrelfish (Palinurichthys perciformis). After DeKay. 370 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Color. — Described as varying from blackish to green in life, and as either as dark below as above, or paling to bluish white on the belly, the latter variously mottled with darker dots and bars. It is said to change color to accord with its surroundings. Size.- — Maximum length 12 to 14 inches and about 1 )i pounds in weight, but most of those seen are smaller. Habits.- — The barrelfish owes its common name to its habit of congregating about floating spars and planks or any drifting wreckage, or inside of barrels or boxes, where it is easy to catch one in a dip net. Off southern New England they are often found under gulf weed, or under any other raft of drifting seaweed or eel grass (Zostera) . And they sometimes gather about slow-moving vessels. Merriman 34 thinks its proper home is in the mid- depths offshore, but this is a question for the future. It feeds on the sundry small crustaceans, bar- nacles, hydroids, young squids, small mollusks, and salpae, which it finds near or attached to its floating homes; on ctenophores; likewise on fish fry, the diet lists of specimens taken at Woods Hole including herring, mackerel, menhaden, launce, scup, and silversides.35 Sometimes they contain seaweed, but we suspect that this is eaten for the animals attached to it, and not from a vegetarian taste. Nothing is known of its breeding habits. General range. — Atlantic Coast of North Amer- ica, Cape Hatteras to outer Nova Scotia;36 most plentiful south of Cape Cod. Probably it is "Trans. Connecticut Acad. Arts. Sci., vol. 36, 1945, pp. 842-S43. 31 Notes by Vinal Edwards. ** According to Vladykov and McKenzie (Proc Nova Scotian Inst. Sci., vol. 19, 1935, p. 87) occasional specimens are caught oS outer Nova Scotia in most summers. Recent records there are of one at Halifax, October 1924, and of another there September 1927 (Vladykov, Proc. Nova Scotian Inst. Sci., vol. 19, 1935, p. 6). oceanic, as Merriman 37 suggests, and more widely distributed than the foregoing would suggest, for one was found in a drifting packing case off Penzance Harbor, Cornwall. Occurrence in the Gulf of Maine. — The barrelfish is caught in some numbers in the traps near Woods Hole and to the westward, or is found drifting under mats of seaweed. They were unusually plentiful in Vineyard Sound, for exam- ple, in 1920.38 But it is so rare a fish within the Gulf of Maine that we have never seen it there,3' nor did Doctor Kendall find it on his various collecting trips along the Maine coast. In fact, the only published Gulf of Maine records for it that we have been able to find are one from Bos- ton Harbor; one from Salem; one from Annisquam; one from Gloucester;40 and one vaguely described as brought in from the fishing banks off the coast of Maine. We can now add one taken on the northern edge of Georges Bank by the trawler Squall on September 10, 1947." Black Ruff Centrolophus niger (Gmelin) 1789 Jordan and Evermann, 1896-1900, p. 963. Description. — The black ruff resembles the pilot fish (p. 372) in its general body form more than it does its closer relative the barrel fish (p. 369), being moderately slender (a little more than % as deep as it is long to base of tail fin) , with very blunt snout, strongly convex forehead, and small mouth. But its body (about 2% times as « Trans. Connecticut Acad. Arts Sci., vol. 36, 1945, pp. 842-S43. » Smith, Copeia, 1921, No. 91, pp. 9-10. 59 Our own experience with this fish is limited to a single occasion, south of Nantucket, when several were seen about a drifting box. <° Reported by MacCoy, Bull. 67, Boston Soc. Nat. Hist., 1933, p. 9. 41 This specimen now in the Museum of Comparative Zoology, and was received through the kindness of J. Miggins of the U. S. Fish and Wildlife Service. ^SEr* ~:^m Figure 196. — Black ruff (Cenlrolophus niger), Dennis, Mass. From Goode and Bean. Drawing by S. F. Denton. FISHES OF THE GULF OF MAINE 371 high as it is thick) is more flattened sidewise than that of the pilot fish; its caudal peduncle is much deeper, and has no lateral keels; its dorsal fin is considerably longer than that of the pilot fish, and there are no detached spines in front of its dorsal fin. The single dorsal fin of 3 flexible spines entirely covered over by skin and 35-38 soft rays reaches from over the pectoral fin to the caudal peduncle; the anal (3 spines concealed by skin and 20-22 soft rays) originates about under the midpoint of the dorsal and runs equally far back; both dorsal and anal fins are evenly gradu- ated in outline from front to rear; and both are fleshy and scaly along their bases. Its ventral fins are about under the pectorals; pectorals and anals are both small; and the caudal is moderately forked. Color. — Those we have seen (after a few weeks preservation in alcohol) are dark leaden-brown on back and sides, with the margins of the scales darkest, in so fine a pattern (because of the small size of the scales) that the general effect is sooty; the fins are darker, even, than the back; and the belly only a little paler. Other specimens have been described a as brownish pink all over, or brown, darkest above, some with irregular and obscure markings, either yellowish or dark blue. Size. — Grows to about 2 feet in length. General range. — Oceanic, and widespread in low and mid latitudes in the eastern North Atlantic; Madeira, the Azores, and the coasts of Spain north to the entrance to the English Channel, the Celtic Sea and southern Norway; also in the Mediter- ranean; and reported as a stray from Massachu- setts and from Georges Bank. Occurrence in the Gulf of Maine. — One specimen of this wanderer from Europe, about 12% inches long, was taken in a trap at North Truro, on Cape Cod Bay, September 6, 1890; u a second of 21 H inches was brought in from the northern edge of Georges Bank by the trawler Thomas Whalen in September 1936;" and a third of about 13 inches (330 mm.) to the fork of the tail was taken in a trap at North Truro June 23, 1951. 4S Another about 9 inches long was taken in 18884* at Dennis, Mass. But it is not known whether this record should be credited to our Gulf or to the southern coast of Massachusetts, since that town- ship fronts both on Cape Cod Bay and on Nan- tucket Sound. THE POMPANOS AND JACKS. FAMILY CARANGIDAE The pompanos are allied to the mackerels; both have deeply forked tails, very slender caudal ped- uncles, and ventrals situated below the pectorals. And, mackerel-like, most of them have two dorsal fins, the first hard-spined, the second soft-rayed. But they are readily separable from the mackerels by the fact that their first (spiny) dorsal, if they have one, is much shorter than the second (soft rayed) while it may be reduced to a series of very short spines, or even lost altogether in old age. And, except for the leather jacket, they either lack the dorsal and anal finlets so characteristic of the mackerel tribe, or have only one of each at most. They differ further from the mackerels in the number of vertebrae (only 24 as against up- ward of 30), and in that their premaxillary bones (fixed in the mackerels) are protractile (except in adult Oligoplites) , while their anal fin is preceded by two free spines that may either take the form of a permanent finlet or may be lost in old age. Warm seas support a host of species, but none of them is more than an accidental stray to the Gulf of Maine. KEY TO GULF OF MAINE POMPANOS 1. Rear parts of soft dorsal fin, from 7th ray backward, and of anal fin from 6th ray backward are so deeply indented between every two rays as to form a series of 12 to 14 nearly separate low finlets Leather jacket, p. 380 Rear parts of soft dorsal and of anal fins even-edged, not as series of finlets 2 2. Body very much flattened, sidewise; nearly or quite half as deep as it is long to base of caudal fin 3 Body moderately stout, less than two-fifths as deep as it is long to base of caudal fin 6 3. Back and belly rounded; pectoral fins reach not more than one-third the way back toward the base of the caudal fin True pompanos (genus Trachinotus) i7 Back and belly sharp-edged; pectoral fins reach at least half-way back toward the base of the caudal fin. 4 « Day, Fishes Great Britain, 1880-1884, vol. 1, p. 110. « Reported by Bean, Proc. U. S. Nat. Mus., vol. 21, 1898, p. 639 and now in the U. S. National Museum. « Reported by Bigelow and Schroeder, Copeia, 1937, p. 61. « Received through the kindness of John Worthington of the Pond Village Cold Storage Co. « Ooode and Bean, Smithsonian Contrib. Knowl., vol. 30, 1895, p. 214. " None of these southern fish have yet been reported within our Gulf. 372 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE 4. Second dorsal and anal fins are conspicuously falcate in shape, very high in front, tapering abruptly toward the rear; there are no enlarged bony plates along the lateral line on the caudal peduncle Lookdown, p. 379 Second dorsal and anal fins only moderately high in front, tapering rearward gradually; caudal peduncle with weak bony plates along the lateral line 5 5. Upper anterior profile of head concave; ventral fins very small; anterior rays of soft dorsal and of anal not elon- gate Moonfish, p. 378 Upper anterior profile of head convex; ventral fins as long as head or longer; anterior rays of soft dorsal and anal fins elongate, threadlike Thread fin (probably the young of the Cuban jack) , p. 381 6. There is only one well-developed dorsal fin (the soft rayed), the first (spiny) dorsal being reduced to a few short spines, without separate fin membranes Pilot fish, p. 372 There are two well-developed dorsal fins though the first (spiny) is smaller than the second 7 7. There is a detached finlet behind the dorsal fin and one behind the anal fin Mackerel scad, p. 374 There are no finlets behind the dorsal and anal fins 8 8. There is no finlet in front of the anal fin; and the anal is only about one-half as long as the soft dorsal Rudderfish, p. 373 There is a finlet of 2 short spines in front of the anal fin, and the anal fin is nearly or quite as long as the soft dorsal 9 9. The forward part of the lateral line is scarcely arched Goggle-eyed scad, p. 377 The forward part of the lateral line is strongly arched 10 10. The breast is naked, except for a small patch of scales in front of the ventral fins Crevalle, p. 375 The breast is covered with scales 11 11. The body (to base of tail) is not more than 3 times as long as it is deep; the soft dorsal fin has only 23 to 25 rays Hardtail, p. 376 The body to base of tail is more than 3 times as long as it is deep; the soft dorsal fin has 30 to 35 rays __ Saurel, p. 377 Pilotfish Naucrates ductor (Linnaeus) 1758 Rudderfish; Shark pilot Jordan and Evermann, 1896-1900, p. 900. Description. — The pilotfish is one of the more slender carangids (body about one-fourth as deep as it is long), round-sided, about two-thirds as thick as it is deep, and somewhat mackerel-like in appearance though with a blunter, more rounded nose and smaller mouth, while its caudal peduncle is conspicuously keeled on either side like that of a bonito. But its long second dorsal fin separates it from all the mackerel tribe. The first dorsal fin is reduced to three or four short inconspicuous spines, which are connected by a membrane in young fish but this membrane is lost with growth. The second dorsal (26 or 27 soft rays) is weakly concave in outline and originates midway between tip of snout and base of caudal fin. The anal fin is similar to the second dorsal in form, but is oidy about half as long (16 or 17 raj's), and is preceded by two very short spines. It resembles the rud- derfish in this but the first dorsal of the latter is well developed and has 7 spines instead of only 3 or 4. The ventrals, situated far forward under the pectorals, are about as large as the latter. The caudal is large and deeply forked. The edge of ^mm *S?r? :- '•$ Figure 197. — Pilotfish (Naucrales ductor), about 13 inches long, New Bedford, Mass. After Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 373 the gill cover is rounded in the adult but it bears a spine in young fry. Color. — Bluish, cross-barred with 5 to 7 dark bands, 2 or 3 of which run up on the dorsal fin and down on the anal. The outer margins of caudal, ventral, and pectoral fins are nearly black The caudal is white-tipped. Size. — Maximum length about 2 feet. General range. — A tropical fish of the high seas, rarely straying as far north as outer Nova Scotia.48 Occurrence in the Gulf oj Maine.- — -The only records of this species from within the Gulf are of one taken in a mackerel net in Provincetown Harbor in October 1858, the fish probably having followed a whale ship that arrived a few days pre- vious; one caught near Seguin Island in 1906; one off Portland in September 1921; one taken from a mackerel net at Provincetown in August 1924; three in 1929; one of them from the northern edge of Georges Bank in October, the other two from the South Channel to the southeast of Cape Cod (one in August, one in November) ; one off Portland, July 1931 ; and one picked up in a trawl on the northern slope of Georges Bank Oat. 42°10' N., long. 66°32' W.) October 10, 1933.49 We need only add that this is the fish that so commonly attends sharks in tropic seas, either picking up a living from the scraps left by the latter, or feeding on the parasites with which their protectors are infested. They often follow sailing vessels, also. *' Vladykov (Proc. Nova Scotian Inst. Scl., vol. 19, 1935, p. 0), reports two specimens taken on Sable Island Bank, and one from Sambro near Halifax, during the period 1932-34. "Reported to us by W. C. Neville of the U. S. Bur. Fish. Rudderfish Seriola zonata (Mitchill) 181 5. M Amberjack; Pilotfish Jordan and Evermann, 1896-1900, p. 902. Description. — The rudderfish is deeper bodied, relatively, than the pilotfish (body about three and one-half times as long as deep), so much flattened sidewise that it is almost as thin as a butterfish (p. 363), and with a pointed nose. Its first (spiny) dorsal fin is well developed, with 7 spines. There are 36 to 38 rays in the second dorsal fin (only 26 or 27 in the pilotfish) and the ventrals are relatively much longer than in the pilot. In young fry of 2 to 3 inches the second dorsal originates a little in front of the tips of the pectorals, but it origi- nates slightly behind the tips of the pectorals by the time the fish has grown to 8 or 9 inches, and still farther back in larger specimens.61 The anal fin (20 or 21 rays) is a little more than half as long as the second dorsal in the rudderfish, as it is in the pilotfish also. And in young fish it is preceded by one or two short spines which adults lack. The ventrals are a little longer than the pec- torals, and more pointed in large fish than in small; the caudal is deeply forked, its slender peduncle with a longitudinal keel on each side; the mouth gapes back to the forward margin of the eye and is armed with broad bands of hairlike teeth. The body is clad with small scales. " The interrelationships of the several Seriola! that have been described from our South Atlantic coast still remain in doubt. " We have examined specimens ranging from 3 to 9 inches In length taken in Cape Cod Bay, at Woods Hole, Now Bedford, and other localities. Storer's illustration, reproduced here, was of a 2-inch fish. **■ /.'.-■-:■"■'. i u '• ■ ,1 .,, i -/■-.Vi'i- ■■— Figure 198.— Rudderfish (Seriola zonata), young, in striped stage, Wellfleet, Mass. After Storer. 374 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE Color. — Bluish or silvery brown above, paler on the sides, and white below. In young fish (no large ones have been reported from within our limits) the sides are conspicuously crossbarred with 5 or 6 broad dark blue or brown bands, the last 4 run up on the dorsal fin and the last 2 or 3 down on the anal fin. There also is a dark band running obliquely from the first dorsal to the eye in some cases. All of these bands fade with growth, however, to disappear in large fish. The first dorsal is black, the anal white at the base, the ventrals black above, pale below, and the caudal dusky green, with white tips.62 Size. — Maximum length about 3 feet. General range. — Atlantic Coast of America, Halifax, Nova Scotia,63 to Gulf of Mexico. Occurrence in the Gulf of Maine.- — The rudder- fish is ordinarily a rare visitor to the Gulf of Maine, and most of those that have been seen there have been small, made conspicuous by their crossbarred pattern. Two were taken at Well- fleet in 1844 and 1849 (mentioned by Storer); another at Beverly in May 1866; one five inches long at Provincetown in 1870; and one at Salem sometime prior to 1879. A gap then follows in the record until September 1921, when one was caught by an angler fishing for smelt at a wharf in Portland Harbor.64 Another, of 5?2 inches was caught on September 22, 1929, also by an angler fishing for smelt; one of 6% inches was taken on Nantucket Shoals August 1, 1930 ;66 several were reported in 1949 at Boothbay Harbor, the Sheep- scot River, and at Gloucester.66 However, in the a We have do color notes from life. •' Reported by Leim, Proc. Nova Scotian Inst. Sci., vol. 17, 1930, No. IV, p. xlvi, as £?. dumerili. " Reported to us by Walter H. Rich. " Reported by Firth, Bull. 61, Boston Soe. Nat. Hist.; 1931, p. 12. M Reported by Scattergood, Trefethen, and Coffin, Copeia, 1951, p. 298. summer and fall of the years 1949-51 large num- bers of them were caught or observed in and around the traps at Barnstable, Cape Cod Bay, and one day's record catch by one set of pound nets, within this period, amounted to two barrels 67 indicating that, in some years, large schools of rudderfish are sometimes present in the latter region. Small fry 1 % to 7 inches long are regular summer visitors at Woods Hole. Mackerel scad Decapterus macarellus (Cuvier and Valenciennes) 1833 Jordan and Evermann, 1896-1900, p. 909. Description. — This scad is easily recognized among such of its tribe as are known from our Gulf by the presence of a small detached finlet between the second dorsal and the base of the caudal fin with another similar to it behind the anal.6S Furthermore, it is more slender than most of the other pompanos; its body is only about one-fifth as deep as it is long, and fusiform like the mackerel. But the great length of the second dorsal fin and the fact that there is only one dorsal finlet and one anal finlet would separate a mackerel scad from a mackerel at a glance. The mouth of the scad is smaller, and its premaxillary bones are protractile. Its triangular first dorsal fin (8 spines) originates over the middle of the pectorals. Its second dorsal (about 34 rays) is w Information supplied by Frank Mather who was informed of the 1949- 1951 catches at Barnstable by Capt. John Vetorino in whose traps many of these rudderfish were caught. 48 A second scad, the round robin (Decapterus punctatus), similarly characterized, is known as far north as the Woods Hole region. It has 40 or more scutes or shieldlike scales along the lateral line, instead of only about 30 or 31; its jaws are toothed, and it is spotted along the lateral line, char- acters that separate it from the mackerel scad. Figure 199. — Mackerel scad (Decapterus macarellus), Woods Hole. After Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 375 separated from the first dorsal only by a very short space and extends back nearly to the base of the caudal. Its anal fin is similar to its second dorsal in shape but is shorter (about 28 rays), originates about under the seventh or eighth ray of the second dorsal, and is preceded by 2 short stout spines. The ventrals are shorter than the pectorals and situated under them. The tail of the scad is less deeply forked than in most of the pompanos. In place of fleshy keels on the caudal peduncle, the rear half of its lateral fine is armed with a series of 31 keeled shields, largest on the peduncle, and all of them much larger than the ordinary scales, a very noticeable character. Color. — Described as slate blue or leaden above, silvery below, with a small black spot on the margin of the gill cover and with the axil of the pectoral black. We have not seen it alive. Size. — Maximum length about 1 foot. General range. — Warm parts of the Atlantic, rarely straying northward to the Gulf of Maine and to Nova Scotia. Occurrence in the Gulf of Maine. — A specimen caught with smelt in Casco Bay, Maine, in Oc- tober 1920, and another, 7 inches long, taken in a trap at Richmond Island, off Cape Elizabeth in September 1931, are the only Gulf of Maine records, though it has been taken at Canso and at Port Mouton Bay, Nova Scotia.63 But being common in the autumn about Woods Hole, where as many as 10 barrels have been taken from one •» This last flsh, a 2H-inch specimen, caught October 10, 1928, was recorded by Leim (Proc. Nova Scotian Inst. Sci., vol. 17, No. 4, 1930, p. ilvi). trap haul, it would not be surprising to find it north of Cape Cod any summer. Crevalle Caranx hippos (Linnaeus) 1766. Jack Jordan and Evermann, 1896-1900, p. 920 Description. — The presence of a well-developed first dorsal fin (8 spines) combined with an anal (about 17 rays, preceded by 2 short detached spines) nearly as long as the second dorsal (about 20 rays), but no detached finlets, separates this particular jack from all other pompanos known from the Gulf, except the goggle-eyed scad (p. 377), hard tail (p. 376), and the saurel (p. 377). Its arched lateral line and the presence of (usually) two pairs of small but plainly visible canine teeth in the lower jaw distinguish it from the goggle eye; its naked breast and its canine teeth from the hard tail and saurel. The dorsal profile, too, of the head of the crevalle (fig. 200) is characteristic, and the long scimitar-shaped pectoral fins are a convenient field mark to separate it and other members of its immediate tribe,64 from the pilot- fish, rudderfish, and mackerel scad, in which the pectorals are short and blunter. We need only call attention further to its deeply forked tail; to the row of keeled shields along cither side of its caudal peduncle; to its flattened oblong form (body « The yellow tail (Chtoroscombrut chrysurui), another species In this group straggles northward at times and, sometime, may be taken within the Gulf of Maine. It may bodistinguished from the crevalle, hard tall, saurel, and big- eyed scad by the fact that Its lateral line Is wholly unarmed, whereas In these species it is armed with bony plates, along part of its length at least. Figure 200.— Crevalle, or Jack (Coram hippos), Woods Hole, Mass. From Goode. Drawing by H. L. Todd. 376 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE only about two and one-half times as long as deep, but with caudal peduncle as slender as that of a mackerel), and to its blunt head. Color. — Greenish or greenish bronze above with golden sides; silvery below, sometimes with yellow blotches. There is a large black blotch on the gill cover, a fainter dark spot on the lower rays of the pectorals (in adults), and a black blotch in their axils. The fins are more or less yellowish; the edge of the dorsals is black. Very young fish have 5 or 6 dark cross-bars. Size. — Maximum recorded weight 36 pounds. General range. — Warm seas; abundant on both coasts of America; northward as a stray to the outer coast of Nova Scotia; 65 also among the East Indies. Occurrence in the Gulf of Maine. — We know of only two records of this southern fish from our Gulf, one specimen picked up on Lynn Beach on the shore of Massachusetts Bay during the sum- mer of 1847, and a second taken at Provincetown in 1933.66 But it is a regular summer visitor at Woods Hole though it is not common there. Commercial importance. — A famous game fish, but of minor commercial importance. Hard tail Caranx crysos (Mitchill) 1815 Yellow jack; Eunner; Yellow mackerel Jordan and Evermann, 1896-1900, p. 921. Description. — The hard tail resembles the cre- valle, saurel, and goggle-eyed scad in the rel- « Reported near Halifax, Nova Scotia, by Vladykov (V-roc. Nova Scotian Inst. Sci., vol. 19, 1935, p. 4). " Reported by MacCoy, Bull. 70, Boston Soc. Nat. History, 1934, p. 6. ative sizes and arrangement of its fins, in its deeply forked tail, in its slender caudal peduncle and in the presence of a row of bony shields along at least the rear part of its lateral line. But its scaly breast, the lack of canine teeth in its lower jaw, and the lack of a black spot on the pectoral fin separates it from the first of these; the fact that the bony plates increase in size, passing rearward along the lateral line, marks it off from the saurel, and its strongly arched lateral line from the goggle eye. Its hrst dorsal fin has 8 spines, its second, one spine followed by 23 to 25 rays, while its anal consists of a finlet of 2 short spines followed, after a distinct gap, by the soft portion with 19 to 21 rays. Color. — Greenish bronze above, golden or silvery below. The fins may show dusky cloudings, and there usually is a dark spot on the gill cover, near the margin, but none on the pectoral fin. Young fry are more or less distinctly cross-barred on the sides, but these bars disappear with growth. Size. — Maximum weight about 4 pounds and length about 22 inches. Northern examples are seldom more than a foot long. General range. — -Atlantic coast of America, Brazil to Rhode Island and to Nantucket Sound regularly, and as far northward as outer Nova Scotia as a stray; represented by a closely allied species in the Pacific. Occurrence in the Gulf of Maine. — The fact that this fish has been reported at Chatham on Cape Cod in 1933,67 at Provincetown, in Boston Harbor, « MacCoy, Bull. 70, Boston Soc. Nat. History, 1934, p. 6. BsS^** ; Figdre 201. — Hardtail (Caranx crysos). Woods Hole. From Goode. Drawing by H. L. Todd. FISHES OF THE GULF OF MAINE 377 off Gloucester,08 and in Ipswich Bay,69 and that 1 1 specimens, about 6 to 8 inches long, were taken in a fish trap at Barnstable on the shore of Cape Cod Bay on September 6, 1950, shows that it is more likely to round Cape Cod than is the crevalle. It is also reported from outer Nova Scotia.70 Young fish are not rare about Woods Hole and thence westward from July to November. Saurel Trachurus trachurus (Linnaeus) 1758 Rough scad Jordan and Evermann, 1899-1900, p. 910.71 Description. — The. saurel is distinguishable from all allied species yet known from New England waters by having about 75 bony plates along its lateral lines, as contrasted with about 30 or fewer in other Gulf of Maine carangids. It is a some- what deeper fish than the mackerel scad but more slender than the hardtail or the crevalle, its body (to the base of tail) being about 3% times as long as it is deep. Its first dorsal fin, of 8 spines, is closely followed by the long second dorsal of 25 to 30 soft rays. Its soft anal, opposite the second dorsal, has 24 to 26 rays, and is preceded by two small detached spines. The tail is deeply forked " One netted September 18, 1878. M Specimen now in the collection of the Boston Society of Natural History. '° A 5-inch specimen is reported from Port Mouton by Leim (I'roc. Nova Scotian Inst. Sci., vol. 17, 1930, No. 4, p. xlvi), ibnico, and near Halifax, by Vladykov (Froc Nova Scotian Inst. Sci., vol. 19, 1935, p. 4). "Nichols (Bull. Amer. Mils. Nat. nisi., vol. 12, 1920, p. 179) considers the western Atlantic saurel distinct from tho eastern Atlantic saurel and has proposed the name lathamt for it. But this separation bas not been adopted cenerally. Color. — Described as bluish green above, silvery below, with a black spot on the edge of the gill cover above its rear angle. Size. — Length about one foot. General range. — Known from nearly all warm and temperate seas, sometimes common off the Florida Keys. A few have been recorded from the vicinity of New York, one from Newport, R. I., and three from the Gulf of Maine. Occurrence in the Gulf of Maine. — One specimen of this rare fish was taken in Casco Bay on August 12, a second at Castine, Maine, on October 15, 1930,72 and a third at Sandwich, Mass., on Cape Cod Bay in the summer of 1950.73 Goggle-eyed scad Trachurops crumenophthalmus (Bloch) 17